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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 <P 28-31, 33-40. 

5 L. alpin inum aS fa complanatum auct.; Alpine Clubmoss 


Native. fore) ountain g rassland, usually above 
BL, §,.F2. 34, 37, 42), 4 43-46, (47), 48-50, [51], ‘sh, ote (57), [58, 59], 
60, (62-66), 67-70, (1), 32,73; FIAT, (73; we 7 7). 791, (20), “ (83), 


86-92, Ae 94), 9, (100), 101, [102, 103], AHi01, 11. U2: 

H (1, 2, 16), 20, Gi, oy, [28], 29, (34), 35, 36, $640. 
SELAGINELLACEAE 

2 SELAGINELLA Beauv. 

7 S. selaginoides (L.) Link Lesser Clubmoss 


Native. Amongst moss or in short turf a ined in hilly districts. 
[17], 48-52, 54, 57-83, 85-112; HB, 9-40 
5 


2 S. kraussiana -oeastag A. — 


Introduced. E gre 1S itivation and has become established 
in a few place: 
6. t, 3. 24. 104: H3. 
ISOETACEAE 
3 ISOETES L. 
] I. lacustris L. Quillwort 


agri lakes and tarns with varied substrate, usually poor in dissolved 


[3], 40, (41), [42], 44, 46, [47], 48, 49, [S0, he 52, (61), 67, 69, 70, (72-74), 
[75], 79, [85], 86-90, 92, ee 95-112; Hi (2, 3 0, [9], 16, 20, (25), 27, 29, 
30, 32-36, (37), 38, 39, (40 

2 I. echinospora Durieu Quillwort 
Native. In lakes and tarns, usually on peaty subs 

2, 3, 9, 41, 46, 48, 49, [5S1, 87], 88, Pt, 92, 93, 96, 197198, 99, 103-105, [107], 
108, 110, 112: Hi, 29, 16,21,3 

3 I. histrix Bory 


Native. In peaty and sandy places with a high winter bred 
On Guernsey and Alderney, v.c.0, and on the Lizard, v 


EQUISETACEAE 

4 EQUISETUM L. 
I E. hyemale L. Rough Horsetail 
Native. Shady stream banks, damp woods. 
[4], (6), 11, (a), TiS, (17), Dil (26, 27, 29, 30, 36-39), [40], 41, [50, 51]. 
54, (55), (56}, (58), [59], 60-70, 72, 73, 75, 71, 81, 83, 84, [87]. 88 8-97, 104 

, 109; H1, (2, 4, 9), [11], 12, (13-15, 18, 19), "20, 1, 28, (29, 31, 33, 35), 
36, (38), 29, (40). 
E. x trachyodon A. Braun (E. —— x E. variegatum) 
Native. Shady marshy banks by 
59, 64, 88, 90-92, [103]; H1, 2, ‘A 0. “14-16, 21, 26-28, 33-36, 38-40. 
2 E. moorei Newm. (E. occidentale (Hy) Coste) 
Native. Dunes and banks by the sea, rarely inland. 
17, 88, 92; H2, 12, 20. 
3 E. ramosissimum Desf. 
Probably native; first found in 1947. 
Only one station by the R. Witham, near Boston, S. Lincoln, v.c.53. 


4 E. variegatum Schleich. ex Weber & Mohr Variegated Horsetail 
Native. Dune-slacks, wet banks and flushes on mountains. 
3, 4, (6), [20], 22, 41, [42], 4446, 51, 52. 58-60, 68-70, 


>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]. 
<x ASPLENOPHYLLITIS Alston (ASPLENIUM x PH YLLITIS) 
x Asplenophyllitis jacksonii Alston (A. adiantum-nigrum x P. scolopendrium) 
Found with apr in Jersey and Guernsey, v.c.0, and at Hartland, 
E. Cornwall, v.c 
9 


x Asplenophyllitis microdon (T. Moore) Alston 
(A. obovatum x P. scolopendrium 
Found in Guernsey, v.c.0, and Penzance, W. Cornwall, v.c.1. 
* oe confluens (Lowe) Alston 
(A. trichomanes * P. scolopendrium) 
Found eae tomes with parents. 
62, 70; H2. 
16 CETERACH DC. 


1 C. officinarum DC. (Asplenium ceterach L.) Rusty-back Fern 


Native. On walls and calcareous rocks and soil. 
0, 1-24, 26, 27, 30, 32-52, 56-60, [61], 62-65, 68-77, 81, [82, 86], 88, 89, 
98-101, [102], 103: HI-40. 


ATHYRIACEAE 
17 MATTEUCCIA Tod. 
I M. struthiopteris (L.) Tod. Ostrich Fern 
Naturalised in damp ground and in woods in a few places. 
13, 88, 89; H39. 
18 ATHYRIUM Roth 
] A. filix-femina (L.) Roth Lady-fern 
Native. In woods, on heaths, damp gullies, etc. 
In all vice-counties. 
2 A. alpestre Clairv. sensu la Alpine Lady-fern 
(Aspidium alpestre (Hope): Polypodium alpestre (Hoppe) Spenn.) 


1 A. alpestre Clairv. (Polypodium molle All., non Schreb.) 
are In shady places amongst alpine rocks and by mountain stream 


186, 87], 88, [89], 90, = [94], 96-98, 105, 106, [107], 108; [H34, 35]. 
2 A. flexile (Newm.) D 
ee flexile Peri ) T. Moore, non Fée; Athyrium alpestre vat. 
exile 
Native. High pitta corries. 
88, 90, 94, 96-98, [108]. 
19 CYSTOPTERIS Bernh. 


J C. fragilis (L.) Bernh. ttle Bladder-fern 
(Cc. dentate asm Bry os — ee ar ee CG. ee ‘CRoth) Desv.) 

Native. ocks yon man-made walls. 

0, 1-9, iT sg [13- cL 1, is. 19], ( OO), say [25], 26, 27*, 30, 32-44, 

46-52 , 54-60, 62-112; H1-3, 6-11, 13-17, 19, 20, 22-30, 32-40. 

2 C. dickieana Sim 


Native. In dark and wet rock clefts. First found in Kincardine. 

{71], 88, 90, 91 

a eT (Lam.) Desv. Mountain Bladder-fera 
n damp situations usually on bas 

a9 (69), "01, (86), 88, 90, (92), ak 97, 98, ian [104]. 


20 WOODSIA R.Br. 


ss a _— a R. Br. Oblong Woodsia 
n rock faces and rock ledges above 1,200ft. 
ash "49, rer 66, 69, 70, 72, [88], 90, [103]. 


2 W. alpina (Bolton) G Alpine or Round-leaved Woodsia 
(W. 


.) R.Br. 
Native. Rocky crevices on mountains from 1,900ft. to 3,000 ft. 
49, 88, "[89], 90, ri04 ]. 
ASPIDIACEAE 

21 DRYOPTERIS Adans. (VEPHRODIUM Michx.) 

/ D. filix-mas (L.) Scho Male Fern 

(Aspidium ype Lae Pa Nephrodium filix-mas (L.) Strempel; 
strea filix-mas (L.) C. Presl) 
Native. Woodlands, a ee mountain slopes, etc. 
Widespread throughout the British Isles 
D. filix-mas x D. lanceolatocristata re Sixipend Druce pro parte; 
Fogg remotum auct.; Lastrea remota auct.) 
Occurs i with the parents. 

ey 69, [99]; H15 
2 D. borreri Newm. 

Native. In similar situations to D. filix 
0, 1-11, 13-19. 25, 27, 30, 31, 33-36, 38. “40-42, 44-50, 53, 54, 56, 57, 60, 
62-75, 77, 78, 80, 82, 83, 85-92, 94-101, 104-106, 108, Il: Hi-4, i, a 9, 12, 
5S, 16, 20, aa 27-29, 33, 35, 36, 38, 40. 
D. x tavelii Rothm. (D. borreri x D. tien: 
Native. In similar situations as parents 
2, 3, 8, 9, 35, 36, 69, 85, 94. 
D X woynarii Rothm. (D. borreri x D. dilatata) 
Presumably native. Has been found by the side of Loch Lomond, Dumbarton, 
v.c.99, 


a? iggy bees Lats ) peak (Lastrea propinqua btinnirent non the aay ) 
Native. On screes, etc., in drier situations than previous tw 

9, 14, 34, 33, 40, “49, 54, 64, [65], 66, 69, 70, 72, 77, 87.90. 92, 07, 98, T01- 105, 
108, 110: Hil, 38. 


4 D. villarii (Bellardi) pane Rigid Buckler-fern 
A 


5 D. cristata L). A. Gra ested Buckler-fern 
(Aspidium cristatum (Ls Serta A se (L.) C "Pre 1) 

Native. On wet heaths and in and marshes 

es 8, 9], 10, IS, 17, 23, 23 07. “8. [38], 31. B71, 39, 56, 58, 61-64, 74-76, 

oh 

D. x (Newm “door ex Druce (D. cristata x D. lanceolatocristata; 
ioe wiainoss New 

Native. On wet heaths a: 2 atiailiel marshes 

15, [18, 24, 25] 27-29, 36. S840, an 62, 85, [91]; H2. 


6 D. lanceolatocristata (Hoffm.) Alston arrow Buckler-fern 
(Aspidium spinulosum Sw.; Lastrea_ spinulosa a “pial: Dryopteris 
spinulosa Watt) 

Native. See ahs oods, marshes, wet hea 

1-22, [23], 2 442, 44, 46, 48, 49, se V7). 78-81, [83], 85-92, 94-106, 

108, ‘110; HI-11, 13-20, 21-40. 


7 D. dilatata (Hoffm.) A. G Broad sop ees 
(Aspidium dilatata (Hoffin,) Sm.; Erotik 8 dilatata (Hoffm.) C. Prest; 
Dryopteris aristata Druce, non Kuntze austriaca auct.) 

Native. Woods, hedgebanks, in rock song 

Common throughout the British Isles. A send mountain form has been re- 

corded (as var. alpinum T. Moore) from: 70, 86, 88-90, 92, 88, 104, 109, 111. 

Some of these have proved to be the diploid form and ae collecting of 

fertile material is needed. 


D. < subaustriaca Rothm. (D. dilatata x D. filix-mas) 
Recorded from S. Devon, v.c.3, and from Dalystown, S.E. Galway, v.c.H15. 


D. x deweveri (Jansen) Jansen & Wachter a dilatata < D. lanceolatocristata) 
Native. Occurs frequently with the pare 
Recorded from 2, 6, 14, 17, 18, 27, 34, 36, 40, 61, 63, 69; H15, [22, 23], 34. 


8 D. aemula (Ait.) Kuntze Hay-scented Buckler-fern 
(Aspidium it (Ait.) Sw.; Lastrea aemula (Ait.) Brackenr.) 

Native. Woods, shady banks and rocks. 

1-5. [6], (8, 9), 14-17, 34, 36, 39, (40, 41), 44-46, (48), 49, [51, 52], 62, (63), 

66, 67, (68), 69-71, 74, (75), 76, (80, 81, 89, he aah 196], 97, 98, [99], 

100-105, [109], 110, 111; H1-18, 20, (21), 23, 24, 2 

22 POLYSTICHUM Roth 

1 P. setiferum (Forsk.) Woyna Soft Shield-fern 
(Aspidium poet oe Kit. ex Willd. : ; Polystichum angulare (Willd.) C. Presl; 
P. braunii auct.) 

Native. Woods, banks, hedgerows, etc. on light soils. 

0, 1-27, 31-52, 54-71, 73-75, [76, 77], 78, 80-83, 88, 95, 97, 98, [99], 100, 

101; Hi-40. 

2 P. aculeatum (L.) Roth ard Shield-fern 
(Aspidium aculeatum (L.) Sw.; A. lobatum (Huds.) oe Polystichum 
lobatum (Huds.) lire 

Native. Woods, hedgebanks, 

1-28, (29), 30-52, 54-83, [84], $5111: Hi-3, 5—40. 


P. x bicknellii (Christ) Hahne (P. aculeatum « P. setiferum) 

Native. Occurs occasionally in mixed populations of the parent species. 
Recorded for [1], 4, 12, 22, 39, 59. 

3 P. lonchitis (L.) Roth (Aspidium lonchitis on a ies ) Holly Fern 
Native. On screes, and in crevices of basic 

23*, (29, 32, 36, 41, 48), 49, 64-66, [68], 69, 70, 2 86-90, 92, 94, [95], 96-99, 
{103}, 104-108, (109, diy i, Se 16 (27), 28, 29, 34, (35). 

23 CYRTOMIUM C. Pres! 

i C. faleatum (L.f.) C. Presi 


An eastern asiatic species often cultivated i in the cool house 
Escaped in the Scilly Isles, v.c.1, in ma! v.c.17 and ‘pomibiy elsewhere. 


THELYPTERIDACEAE 


24 THELYPTERIS Schmidel (LASTREA Bory; GYMNOCARPIUM 
Newm.; PHEGOPTERIS (C. Pres!) Fée) 


I T. limbosperma (All.) H. P. Fuchs n Fern 
(T. oreopteris (E hrh.) Slosson; Aspidium oreopteris ‘een Sw.; 
Dryopteris oreopteris (Ehrh.) Maxon) 

Native. In woods and mountain Semen nd screes in siliceous areas. 

1-27, 33-81, 83, 85-112; H1-3, 6-14, 18, 20, 21, 24, 25, 27-31, 33-40. 


2 T. palustris Sch Marsh Fern 
AAspidium sal (L.) Sw.; Dryopteris ai eee (L.) A. Gray). 
n wet w 


31, 34, 36, (39), 40. 


3 T. phegopteris (L.) Slosson Beech Fern 
(Polypodium ty sar L.; Phegopteris polypodioides Fée; Dryopteris 
phegopteris (L. Chr. 

Native. In shady Chet i in siliceous rocks. 

1-6, (8), 9, 11, 13, 14,47, 21, 22, 33-36, 39-44, 46-51, 57-112; H1-3, 9, 14, 

16, 20, 21, 27-31, 33-40. 


4 T. dryopteris (L.) Slosson Oak Fi 
Polypodium dryopteris L.; Gymnocarpium dryopteris (L.) Newm 
Nola sh ie apc (L.) Fees gitar g ny sonferipsc Christ), 


N; a ons 00 
2-5, 9 11, a id I 18, 19], "33, “Al, 26, B31, 3437, (38), 39-44 [45], 


54. [56], 57-70, (71), eps 73, [74], 75-81, 83-101, [102], 
103-109, (i10}, 112; H(9, 20, 28, 29), 3 


5 T. robertiana (Hoffm.) Slosson Limestone Fern 
(Polypodium robertianum Ho ffm.; Gymnocarpium robertianum (Hoffm,) 
Newm.; Phegopteris vet (Hoffm.) A. Braun; Dryopteris 
robertiana (Hoffm.) C. Chr.) 


Native. Limestone screes and rocks. 
6, 7, 13, 17*, 23, 24, 28, 32-37, 39, 45, 47, [49], 50, 57, 59, 60, 62-66, 69, 70, 
88, [93]: HIIsS, 17], * [27]. 


POLY PODIACEAE 
25 POLYPODIUM L. 
1 P. vulgare L. sensu lato Polypody 
Native. Banks, walls, rocks, on mossy tree trunks, etc. Widesprea 


read. 
This aggregate apparently includes three species and s specimens from any 
locality are needed in order that their distribution may be elucidated. 


MARSILEACEAE 
26 PILULARIA L. 
/ P. globulifera L. Pe were Pillwort 
Pig On = eos ¢ es edges of pools, often submer 
0, » 5, 6, 8-1 20-22, 24-28, 31, (32, 33], ree 55, 58, 59. 61, 


6, 6s 67, 69, 70, eH, am, 83-85, 87-92, 95, 96, 98, 99, 104, 107, [108], 
110; Hi, 2, 16, 35, 36, 39, 40. 
13 


AZOLLACEAE 
27 AZOLLA Lam. 
s = oe ty Lam. 
Ditches, canals and other stretches of open wate 


(0. en eS: 11- ~ U6, 17, 18, 20-22, (23, 24), 25, 27, (29, 33-35), 37, 38, 
59, 25, 0; 2D, 3 


OPHIOGLOSSACEAE 
28 BOTRYCHIUM Sw. 
] B. lunaria (L.) Sw. Moonwort 
ative. Dry grassland, well drained banks, ledges etc. 
1-17, (19, 20), 29-29, (30), 32-112; H1-24, 27-40. 
2 B. lanceolatum (S. G. Gmel.) Angstr. 


Doubtfully native; found originally on Sands of Barry, Angus, v.c.90 and 
has also been recorded from v.c.57, 61, 69 and 79. All records need 
confirming 


3 B. matricariifolium A. Braun ex Koch (2. multifidum auct. 


) 
Native? Found on sandy sea shore at Stevenston, “Ayr, v.c.75 and needs 
confirming in v.c.69 and 90. 


4 B. multifidum (S. G. Gmel.) Rupr. (B. matricariae (Schrank) Spreng.) 
Native? Found on a grass topped wall in Strachan, Kincardine, v.c.91. 
29 OPHIOGLOSSUM L. 


I O. vulgatum L. Adder’s Tongue 
a subsp. vulgatum 
Native. Fens, damp meadows, woodsides, e 
0, 1-77, 79-89, [90], 91, 93-95, 98, 100-103, 7104], 105, 108, 110-112; H1-40. 
6 subsp. ambiguum (Coss. & Germ.) E. F. Warb. 
(O vulgatum subsp. potter nex E. F. Warb. pro parte) 
Native. Sandy ground, short turf near the sea 
0, 1, 4, 9, 41, 48, 110-112; Til, 25 Sy 30 


2 O. lusitanicum L. 


Native. Grassy tops of cliffs. 
Channel Isles, v.c.0, and Scilly Isles, v.c.1. 


omgdbrae REFERENCES 
1 Alston, A. H. G. 1930-58, MS. Ni 
: Botanical Hew and Exchange Club ‘of Br a Isles, Plant Records, in R ts for 1933-47. 
3 Botanica | Socie of the British Isles, Records, in Watsonia, 19095, and in Proceedings. 
4 Clapham, ae R., Tutin, T. G. & Warburg, E. F. 1952, Flora 54 the British Isles, Cambridge. 
re 3 Watso org) —— ingyen _ S.B.I. 1950 Conf. Rep.: 23-29. 
6 Dandy, J. E. 1958, * List of British Vi lar Plan 
7 Druce, G. C. 1933, Comital Flore of psy Brush ‘hes. pucker 
§ Hirde, 3. A. & Wade, A.B. 1954, fone lg Ed.3, Cardiff. 
> 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. <i Jermy, B.SC., F.L.S. 
nt of Botany, 
ae 3 Museum (Natural crete 
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, 
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 
upon paying an annual subscription of TEN SHILLINGS. Full details will be 
sent upon application to the SECRETARY. 


THE 


BRITISH FERN GAZETTE 


NEW SERIES 


VOL. 9 APRIL 1961 PART 2 


Editorial 


Over the past year the Society has expanded and brought into play new 
ideas in the way of more and varied meetings in the field. This issue of the 
nae puts forward yet another aspect, namely the inclusion of a paper on 

non-British, yet European, genus of ferns — Cheilanthes; this has been 
ticki for several reasons but it should suffice to mention two very valid 
ones: first, it is a genus often grown in the greenhouse by British fern lovers 
and to those who grow these plants, although often North American species, 
it will not be new; second, and most important if we are to appreciate every 
aspect of ferns, we must look at genera other than those which are represented 
in our own restricted flora. There is in preparation at the moment a Flora 
of the vascular plants of Europe; Volume 1 of this Flora Europaea will 
contain 46 genera of ferns and fern-allies and its use to both professional and 
amateur botanist will be immense. The paper on Cheilanthes is therefore 
welcomed as a useful contribution towards the aims of this large project. 

Fern Societies are not as numerous throughout the world as might be 
expected. I understand there is a recent one formed in Wellington, New 


Tomitaro Namekata, No. 481, Narita, Narita-Shi Chiba-Ken, Toyko, Japan) 
is very keen to exchange spores of any kind for growing. 

In 1960 the American Fern Society published the 50th anniversary 
volume of the American Fern Journal. The quality and scope of this journai 
over the past 50 years can well be taken as a standard to be kej, in the minds 
of Societies such as our own, an even older body in fact than the A.F.S. 
The papers are of interest to all our members and it is hoped to abstract 
several for the Gazette but this is a periodical that might well be taken with 
good effect by many of our provincial libraries. Indeed, the American Fern 
Society (Membership Secretary: Dr. W. S. Phillips, Department of Botany, 
University of Arizona, Tucson) would welcome members from this country 
and the $2.50 (eighteen shillings) annual subscription would be well spent. 


The BRITISH FERN GAZETTE Vol. 9 Part 1 (pp. 1-32) was published 3rd March 1960. 


33 


THE FERNS OF THE CHANNEL ISLANDS 


DAVID McCLINTOCK 
Bracken Hill, Sevenoaks, Kent 


Considering especially that they total only 75 square miles, i.e., less than 
two 10 km. squares, the Channel Islands have a particularly interesting 
variety of plants. Hundreds of thousands of visitors go there each year, but 
only very very few know what plants to look out for. 


Check lists for each of the main groups of Channel Islands, Jersey, 
Guernsey, hist Sark and Herm, are in hand, the first, for Herm, due 
to appear in 1961. Marquand’s and Lester Garland’s Floras ignored in 
effect any peso not available in the Islands. Consequently any specimens 
in mainland herbaria want reporting as well as present localities. I hope 
therefore these notes on some of the ferns may result in fresh records, 
which are always welcome. 


Lycopodium clavatum L. Stag’s Horn Clubmoss was listed by Gosselin in 
1788 


Selaginella kraussiana (Kunze) A. Braun. The only record for this Clubmoss 
is from Mr. P. J. Girard in July 1948 from under trees in a garden at Le 
Vallon, St. Martins, Guernsey. This is a plant which tends to be ignored for 
two reasons — that it is unobtrusive, and that people call it cultivated. But 
it can be a persistent weed of beds and turf, and the climate of the islands is 
right for it. 
Isoetes hystrix Bory. This small and inconspicuous Quillwort has been well 
own in moist spots, mostly on L’Ancresse Common, in Guernsey since 
1860. In 1957 we found it on several of the off-islets, often in good quantity. 
It is still to be seen in S.E. Alderney, but Marquand’s more northerly locality 
** facing the coast of France” was rediscovered, by Mrs. F. le Sueur, only 
in 1960. Here is yet another plant which so far no-one has definitely found in 
the other islands. There is an undated specimen at Oxford labelled curtly 
** Jersey. H. E. Fox,” but this is probably an example of the confusion often 
made between the various Channel Islands. It is usually at its best only up to 
about May, after which its fronds wither, before most visitors arrive. But 
I have seen them long and fresh and green in a wet August. 


Equisetum sylvaticum. L. There are old unsupported records for Guernsey 
and Sark for this small graceful Horsetail. 


E. telmateia Ehrh. This fine and equally graceful Horsetail has been long 
known in a restricted area north of the airport (although not originally so 
described!) in Alderney, and nowhere else in the archipelago, although 
there was a vague record of it about 1860 in Guernsey. In 1957, however, 
Mr. N. Jee showed it me in quantity as an exuberant weed in a friend’s 
garden in St. Peter Port. The present owners know nothing about its origin 
but I have since learnt that it is occasionally offered by nurserymen. 


34 


Osmunda regalis L. Royal Fern is still to be seen wild in Jersey and Sark. 
In Herm it was planted in two places, one of them very prominent, between 
1920 and 1923 during Sir Compton Mackenzie’s reign. 


Anogramma leptophylia (L.) Link. Many people — e.g., Mr. P. Taylor in his 
recent book — look on this small annual as only a Jersey fern, where it was 
found in 1852. But it was discovered in Guernsey, where there was a legend 
it was planted, as long ago as 1877. (Marquand calls it “ Native?’’). 
Admittedly it is rare there, and it seems that only a very few people ever 

new where to see it. During the last war the late Mr. W. A. Warry was 
warned off by the Germans when he went to revisit it, and returned only in 
1957 when he kindly led a party in search of it. The closest scrutiny of the 
very short stretch of road banks he indicated had failed, when one of us, 
Mrs. B. H. S. Russell, who had wandered 150 yards on, came on a fine 
colony, which is still there. It is not certain if this is Mr. Derrick’s precise 
original locality. It might well be in Sark or other islands but dries up 
before most visitors arrive. 


Adiantum capillus-veneris L. aia is clearly native in Jersey, although 
rare. It was recorded from Herm by Mr. G. Templeman in 1923 with no 
details and, so far as I know, no ies Close search has failed to refind 
it. In Guernsey its only place is on a tomb i in the Cemetery, where it seems 
to have seeded itself: it does the same in old stonework elsewhere in the 
British Isles. There is an old and unsubstantiated record from Alderney. 


Onoclea sensibilis L. There are records of the naturalised Sensitive Fern from 
Jersey by Mr. T. W. Attenborough from Plémont in August 1915, and by 
Mr. J. Piquet from Creux du Vis, St. Mary’s. 


Blechnum spicant (L.) Roth. Hard Fern is surprisingly scarce except in 


Asplenium obovatum Viv. Lanceolate Spleenwort is locally frequent, but it 
takes time to get one’s eye in to know it from the more common A. adiantum- 
nigrum. 1 find the shorter stipe often tells more readily than the less triangular 
outline and the sori being nearer the margin help later as a check. 


x Asplenophyllitis jacksoni Alston (Asplenium adiantum-nigrum L. X 
Phyllitis assoc um (L.) Newm). This rare hybrid was found by Mr. C. 
Jackson, in 1856, 57 and 58 at least, in Guernsey; and at Gréve de Lecq in 
Jersey in Noveliiiée 1863 by “ E.H.N.” (who was this?). It looks like an 
untidy broad-pinnuled Black Spleenwort. 


x A. microdon (T. Moore) Alston (Asplenium obovatum Viv. x Phyllitis 
scolopendrium (L.) Newm.) an even rarer hybrid, was found by Miss Wilkinson 
in 1855 in company with A. obovatum on a bank of rough masonry without 
mortar in St. Peter in the Wood, Guernsey; there is also a Guernsey specimen 
from ‘“* Mr. Griffiths ” in June 1863. It has a similar untidy look to the last, 
and may also be being overlooked. 


35 


Ceterach officinarum DC. Except in Jersey the Rusty-back is also curiously 
rare by comparison with its abundance in the often otherwise similar S.W 
England. One reason may be that walls are fewer. 


Cystopteris fragilis (L.) Bernh. Mrs. le Sueur writes to me ‘‘ There is some- 
thing very odd about Bladder Fern in Jersey — the only island from which 
there are any reports of it. It has been recorded on several occasions yet 
there is only one specimen. It is inconceivable that Stanley Guiton whose 
principal botanical interest was ferns should not have taken a specimen of 
it when it was found in the Mourier Valley in 1919. In spite of extensive 
‘searches in recent years it has not been found there. Later Father Burdo 
recorded it but also failed to provide a specimen — yet he collected 


says “ Lastraea Oreopteris Presl and Cystopteris fragilis Bernh. have been 
found here by the late Dr. Bull, and will have to be searched for.’”’ And 
now an undated unlocalised specimen of John Piquet’s has just turned up, 
together with one labelled ‘*‘ July 1898, very rare, JP,’ but if he had this, 
why is it not mentioned in Lester-Garland’s Flora?” This fern is said to have 
been planted out in Guernsey 80 years ago, presumably unsuccessfully. 


Dryopteris borreri Newm. Observations on the comparative frequency of 
this segretate of Male Fern and of D. filix-mas (L.) Schott would be welcome. 
It has not yet been proved for Jersey. 


D. aemula (Ait.) Kunze. This is a surprising apparent absentee in view of 
its plentifulness in the S.W. of Britain and frequency in the Weald, and in 
Normandy. There is a century-old unverified record for it from Guernsey. 
Polystichum aculeatum (L.) Roth. The Hard Shield Fern has only two very 
old records, one each from Jersey and Guernsey, almost certainly erroneous, 
but it could presumably occur. P. setiferum (Forsk.) Woynar occurs but is 
rare. 

Thelypteris limbosperma (All.) H. P. Fuchs (T. Oreopteris (Ehrh.) Slossons). 

e sole mention is by J. Piquet in 1896, see under Cystopteris. 


Pilularia globulifera L. The only station for Pillwort was St. Peters Marsh, 
ersey, whence it went when this was drained a century ago. 


Azolla filiculoides Lam. There are now four records for this in Guernsey, 
all since 1952, but it is hardly wild yet and grows in no great quantity. So 
far, it is not known from the other islands. 
Botrychium lunaria (L.) Sw. There are vague records for Guernsey, but 
only one specimen of this Moonwort, collected there a century ago. 
Ophioglossum vulgatum L. ssp. vulgatum. Present records for the Common 
Adder’s Tongue are two places in Guernsey, two in Jersey (close together) 
and one — among dense bluebells — on Brecghou. Yet is is probably fairly 
widespread and overlooked — I know I easily overlook it! 

36 


O. vulgatum L. ssp. ambiguum (Coss. & Germ.) E. F. Warburg. This Adder’s. 
Tongue too could very well grow other than in Guernsey, where however, 
search in the S.W. of L’Ancresse Common has failed to turn it up since 1930. 
O. lusitanicum L. Few people see this partly because of its very small size — 
1 in.— and partly because it is over long before the holiday season. To see it 
at its best, go in December or January. It is still on the S. cliffs of Guernsey, 
where it was first detected in the British Isles in 1851 — the original letter is 
in the Brit.Mus.(N.H.). By an error it was recorded for Alderney in 1906. 
Yet the cliffs there, and in the other islands, seem not unsuitable. Both in 
Guernsey and St. Agnes, its only other British locality, its patches are near 
a large rock. There are specimens labelled “‘ Jersey,” by Sir Rawson W. 
Rawson, an experienced pteridologist, in 1862, but no details are known; 
and it is suspected that here is another example of the sort of geographical 
confusion made with the Quillwort and other records. — however 
Says several attempts were made to introduce it into Jerse 

In all 33 species have been known definitely from the ‘eas Islands, 
with five others unconfirmed. 

I am most grateful to Frances le Sueur for many helpful comments. 


IS IT A HYBRID OR A SPECIES?* 
FERN WARD CRANE 
Summit, New Jersey, U.S.A. 
There has been a good deal of confusion, especially in the Dryopteris 
crscenniag Sips St aha series, regarding the interpretation of a given 
specim Is it a hybrid or a species? The problem may be resolved witha 
few een of sulci and some diligent study. 

In order to recognize the characteristic abnormal spores of hybrid ferns, 
it is necessary first to become acquainted with normal spores from sori of 
accurately identified specimens of the species involved. The sculptured 
spores are distinct for each species, and hence they may be compared with data 
already published.! 2 

It is wise to prepare permanent slides, using the medium Permount, and 
to keep a file for reference and for comparative study. An excellent space- 
saver for housing the slides is a metal box containing plastic drawers com- 
monly used to store assorted oddments. The resolution of a simple compound 
microscope is sufficient for spore examination. 

After the student has become familiar with the spores of several related 
species, he may undertake the study of their suspected hybrids. Spores 
produced by a hybrid fern can be quickly recognized, for a majority of them 
are tiny and aborted. Normally in any large sampling, however, there are 
also to be found some unusually large mis-shapen spores, presumably fertile. 
These abnormal spores exhibit a combination of the characters of both 
parents. 

* Reprinted with permission from AMERICAN FERN JOURNAL, Vol. 48:164 (1958 
37 


It is sufficient for the beginner merely to separate the hybrids from the 
taxa meriting species rank on the basis of spore features. After one has 
mastered that phase of research, he may debate the parentage of these 
puzzling hybrids. Data to be considered specifically are gross morphologic 
details of the fronds and the microscopic characters of the indusia and 
abnormal spores 

Study of the specimens in several large herbaria shows that errors are 
often made in identifying D. intermedia, D. spinulosa, and their hybrid 
“var. fructuosa.” In these —— too, D. clintoniana, D. cristata, and 
their hybrid have been misinterpreted, yet the spores of each of these entities 
are distinctly characteristic. Curators of herbaria might well set aside un- 
needed extra sheets from Dryopteris collections for student work. Amateurs, 
too, will find this type of research interesting as well as useful in annotating 
their own specimens. 


REFERENCES 
1 CRANE, FERN WARD. Spore Studies in Dryopteris. I. II, 111. Amer. Fern J. 43: 159-169. 1953; 45: 
14-16. 1955; 46: 127-130. 1956. 
2 ERDTMAN, GUNNAR. Pollen and ae Morphology and ey Taxonomy. Vol. U. Gymnospermae 
Pteridophyta, Bryophyta. The Ronald Press Company, New York. 1957. 


THE GENUS CHEILANTHES SWARTZ AND ITS 
EUROPEAN SPECIES 
HANS PETER FUCHS 

The Hague, Netherlands 
The genus Cheilanthes is represented in Europe by five different species, 
including the two taxa most European taxonomists consider to belong to the 
genus Notholaena. This latter discrepancy in generic interpretation together 
with the fact that the scientific names of two out of the five European species 
do not agree with the provisions of the Int. Code Bot. Nomencl. 1956 may 

justify the following discussions. 

daries between the two entities known under the names 

n 


ys been rath 
the two genera are often considered to be distinct. FEE (1852) Gen. Fil. :155 
therefore called Cheilanthes a ‘‘ genus arduum,” keeping the two genera 
Cheilanthes and Notholaena still separated. More than a century ago 
METTENIUS (1859) in Abh.Senckenb.natf.Ges.3:47—58 drew attention to 
the fact that Notholaena was an artificial genus and included in Cheilanthes 
most of the species formerly considered to belong to that genus. However, 
practically nobody followed METTENIUS in this view until DOMIN (1915) 
in Bibl.Bot.20(85):8 who pointed out that the form of the pseudoindusium 
is a rather uncertain distinctive character. This view became more and more 
general even though most subsequent pteridologists hesitated to follow 


38 


METTENIUS and DOMIN as e.g. C.A.WEATHERBY (1943) in Amer. 
Fern J.33:69 who ia a short note on the typification of the genus Cheilanthes 
wrote about Notholaena: “(and nobody has ever known the boundary between 
the so-called genus and Cheilanthes)”, zs oat frente, ator in a posthumous 
publication, Not. ll R.TRYON (1956) in Contr. 
Gray Herb.(179):1—106]. Also other modern authors, e.g. HORVAT (1927) 
in Act.Bot.Inst.Bot.Univ.Zagreb.2:92, did not agree with the view of 
METTENIUS and DOMIN. On the other hand COPELAND (1947) Genera 
Filicum: 65-66 included Notholaena within the genus Cheilanthes together 
with some other smaller genera kept separate by some taxonomists. This 
systematic question, however, can only be solved by a thorough study of all 
species pertinent to Cheilanthes and Notholaena, especially all the tropic 
species,! and has no or little relevance to the generic interpretation of the 
European species, as they all belong to the genus Cheilanthes in its strict 
sense. 

The generic difference between Cheilanthes and Notholaena as generally 


Notholaena the marginal sori are naked or at most covered by protrusions 
of the epidermis, whilst the sporangia of the species belonging to the genus 
Cheilanthes in its strict sense are covered by the reflexed margin of the 
pinnules. In this respect the two European species so far generally attributed 
to the genus Notholaena seem by no means to belong to this generic group, 
but are clearly representatives of the genus Cheilanthes in its strict sense. 
In Cheilanthes marantae (L.)Domin as well as in the species generally known 
under the binomial Notholaena vellea (W.T.Aiton)Desv.[non R.Br.] the mar- 
ginal sori are covered by a very thin and narrow hymen originating from the 
edge of the pinnule. However, this feature can be seen only in a very young 
Stage. In adult plants quite the contrary effect is observed, as the edges of 
the pinnule are bent outwards and upwards respectively. Therefore, the 
former upper surface of the pinnule is then on the inner side of the folded 
pinnule, the former underside being covered with the sporangia, which now 
form the outer side of the pinnule. This effect is most probably caused by a 
change of turgescence due to strong sunlight, as it is not present in plants 
growing in shady places 

The scientific name generally accepted for the genus under discussion is 
Cheilanthes which was first published by SWARTZ (1806) in his Synopsis 
Filicum on page 126 with the following description: “* Capsulae in punctis 
discretis marginalibus, tectae (nec innatae); Indusiis e squamis membranaceis 
distinctis, interius dehiscentibus; 1. e crenulis marginis ipsius replicatis. 
SWARTZ ascribed eighteen species to his new genus, however, two of 
them as “ Inquirendae” but did not indicate a generic type. Thus, the 
typification of the genus was somewhat questionable, [cf. CHRISTENSEN 
(1906) Index Fil.:XLI and CHING (1941) in Hongkong Natur.10:194). 
" NZ:Dep Sa indwutes No.6, 5145 the spore morpholony, atleast ofthe only New Zealand 


aoe formerly put in the genus Notholaena R.Br., strongly suggests its paar fy ne ‘as om 
heilanthes Sw. am 


JOHN SMITH (1875) Hist.Fil.:282 seems to be the first in designating a type: 
Cheilanthes tenuifolia (Burmann [1768] Fl.Ind.: 237, sub Trichomane) 
Sw. (1806) Syn.Fil.:129. His choice seems to be as reasonable as any other 
and, therefore, there was no need to choose another species as lectotype, as 
has been done by MAXON (1913) in IIl.Fl. NE. U.S. & Can.: 25 and (1925) 
Pter. Porto Rico: 428, who, however, just designated the first species men- 
tioned by SWARTZ in the enumeration of the species, Cheilanthes micro- 
pteris Sw. (1806) l.c.:126, following the general attitude in typification of the 
U.S. botanists of those days. The fact that BRITTON and MAXON have 
been followed later on by WEATHERBY (1943) in Amer.Fern J.33:67-69 
would be irrelevant according to the provisions of the Int.Code Bot. Nomencl. 
1956:295, Appendix IV, para 4e, if PICHI-SERMOLLI in 1950 at the Seventh 
Int.Bot.Congr. in Stockholm [cf.PICHI-SERMOLLI (1952) in Int.Code 
Bot.Nomencl.:83; and (1953) in Webbia 9:393] had not followed 
WEATHERBY, BRITTON and MAXON in typifying the genus Cheilanthes 
Sw. by Cheilanthes micropteris Sw. As the generic name Cheilanthes Sw. with 
the latter as type-species is included in the list of nomina generica conservanda 
[cf.PICHI-SERMOLLI (1954) in Taxon 3:68] it seems that Cheilanthes 
micropteris Sw. has to be accepted as lectotype, there not being any taxonomic 
obstacle against this choice. However, this decision holds only as long as 
Cheilanthes Sw. is included in the list of nomina generica conservanda, and 
this seems not to be necessary as the following facts may explain. 


Cheilanthes Sw. was proposed as a nomen genericum conservandum and 
included later on in the list accordingly because of the existence of the 
generic name A/losorus Bernh.(1806) in Schrader, N.J.Bot.1(2): 5,36 which 
was considered to be congeneric with Cheilanthes Sw. by several pteridologists. 
BERNHARDI described the new genus as having “‘ Sporangia cathetogyrata, 
sessilia, subaggregata. Hyposporangia subcommunia, margine libero sub- 
pellucida’ and further stated “‘ Die Gattung Allosorus begreift vielleicht 
alle Adianta spuria Sw.: ausserdem Adiantum pusillum Willd.?  (Pteris 
acrosticha Balb.), Pteris crispa und (wahrscheinlich) heterophylla Sw. Ich 
habe ihr den Namen wegen der verschiedenen Zusammenhdufung der 
Saamenbehialter gegeben. Bei manchen stehen sie grésstentheils einzeln 
(wie bei Allosorus pusillus), bei andern in Haufen. Bei manchen auf jedem 
Hyposporangium nur eines, bei andern mehrere.” Thus, it becomes evident 
that the genus Allosorus Bernh. was at the time of its publication a “* mixtum 
compositum ”’ the morphologic character not being of generic value. This is 
also the reason why the generic name Allosorus Bernh. was used and typified 
by other authors in quite different ways. 


The first in interpreting this generic name was SWARTZ (1806) Syn.Fil.: 


425 who wrote in the “ Index”: “ Allosorus Bernh.— est Cheilanthes, 
and thus restricted Allosorus Bernh. to those species which are real Cheilanthes 


” 


ERNHARD! 
A liioni oe ee ath a2 yar. ONt wy Soe 
3s au or, £ t ao pores neither WILLDENOW m sone LI ever 
publinned the binomial in F tot ion, it is more likely that the latter really used the binomial on 
barium labels. 
40 


in the original sense of SWARTZ. However, this choice of SWARTZ 
hardly agrees with the original concept of BERNHARDI, as the latter in- 
cludes with some doubt the species of Cheilanthes Sw. in the ‘“‘Adianta 
spuria”’ whilst Adiantum pusillum [Allioni, in hb.?, ex] Sw. and Preris 
crispa (L.)All. are regarded as certainly belonging to the genus Allosorus 
by its author. Moreover the identification and restriction of the generic name 
Allosorus by SWARTZ was done only in the above-mentioned short note in 
the “Index” to his Synopsis and it seems quite probable that SWARTZ 
did not actually see BERNHARDI’s publication, but cited BERNHARDI’s 
new names only according to an oral or written communication which he 
most likely received from one of his two most important German corres- 
pondents, DANIEL MATTHIAS HEINRICH MOHR or FRIEDRICH 
WEBER. 


The typification, and at the same time restriction, of the generic name 
Allosorus by Osmunda crispa L. as first proposed by ROEHLING (1813) 
Deutschl.Fl.,ed.2,3:31 corresponds much better to the original concept. 
Moreover, ROEHLING was the first to typify the genus Allosorus and there 
is no reason whatsoever for not accepting this typification and emendation. 
The most critical pteriodologists such as H.WOYNAR (1913) in Mitt. 
nhatw. Ver.Steiermk.49:147 and CHING (1940) in Sunyatsenia 5:225 followed 
ROEHLING in typifying Allosorus Bernh. by Allosorus crispus (L.)Roehl. 
Thus, it seems to be irrelevant that in more recent times Allosorus Bernh. 
has been typified on quite different species by some American authors. 
UNDERWOOD (1899) in Mem.Torrey Bot.Club 6:276 followed the manner 
of typification generally accepted by the older American taxonomists and 
chose Preris viridis Forsk. (1775)Fl.aeg.-arab. :186, the first species listed by 
SWARTZ (1801) among his ‘‘ Adianta spuria.” That this choice hardly 
agrees with the original generic concept of A//osorus Bernh. has been 
mentioned before. 


Much closer to BERNHARDI’s concept of the genus comes the lecto- 
type chosen by FARWELL (1931) in Amer.Midl.Nat.12:243, i.e. Allosorus 
pusillus ({All. ex] Sw.) Bernh. (1806) in Schrader N.J.Bot.1(2):36. This choice 
would probably match even better the original idea of the generic type of 
Allosorus, being the only combination made by BERNHARDI under 
Allosorus, and also the only illustrated species of the genus [cf. BERNHARDI 
(1806) L.c. t.2.f.6]. However, as mentioned before, there is no reason to 
disregard the first choice of the lectotype and that made by ROEHLING 
therefore has to be accepted. Thus, it seems that the generic name A/losorus 
Bernh. cannot stand as a synonym of Cheilanthes Sw. in the strict sense, and 
the latter therefore does not need to be conserved against the former. 


KEY TO THE EUROPEAN SPECIES OF CHEILANTHES 


Underside of pinnules, at least of adult plants, glabrous, or rarely with some 


Scattered glanduliferous hairs; plants highly odoriferous (of cumarin) 
2 fragrans 


41 


Underside of pinnules + densely covered with hairs or paleae: 
Pinnules convex, with margins inrolled and thus + covering the 
sporangia: 
Upper surface of pinnules glabrous, lower surface with long hairs 
and no glandules; lamina oblong. persica 
Upper surface of pinnules with oncnsineal ‘pianidules along the 
midrib, lower surface with long hairs and glandules; lamina 
deltoi oe ... 5 hispanica 
Pinnules flat, or, sere i in adult poe in very open habitats, folded 
upwards along the midrib, with margins only turned slightly downwards: 
Upper surface of pinnules glabrous with scattered ae pi the 
midrib 1 marantae 


Upper sia of ee sieeccduieied ie ... 3 catanensis 


The first of the five known European species which was published with 
a binary name appears in the first edition of LINNAEUS’ Species Plantarum 
as Acrostichum Marantae L. The transfer of this species to Cheilanthes Sw. 
was made only in 1915 by DOMIN in a footnote in his publication Beitrage 
zur Flora und Pflanzengeographie Australiens: 


1. Cheilanthes marantae (L.)Domin (1915) in Bibl.Bot., Stuttgart 20:133, 
adnot.1. 
Acrostichum Marantae L. (1753) Spec.Plant., 2:1071 (‘* Maranta”’). 


Polypodium Marantae (L.)Hoffm. Shi! pcre co Taschenb.1795(2):5, 
partim, quoad syn.Linnaei et forsan oc.Styria 


gh ponies baie igr fod (i: poe il ree Fi. Germ a0 ):92 job quoad 
ei, C.Bauhini et Camerarii (“ Polystichum? Mar *)s 
Pinar Cterah van. var. io Ber jae in ae ; Bot.1799(1): me eit syn. 
ni pro 
Acrostichum peiad indy Cav. (1801) in Anales cienc.nat., Madrid 4:97. 
Ceterach Marantae (L.)DC.apud Lam. & DC. (1805) Fl.franc., ed. 3. 2-567. 
Acrostichum canariensis Willd. in L. (1810) Spec. Plant., ed.4, 5(1):121. 
Cincinalis Marantae (L.)Desv. (1811) in Mag.natf.Frde., Berlin 5:312. 
Cincinalis subcordata (Cav.)Desv. (1811) in Mag.natf.Frde., Berlin 5:312. 
Notholaena Marantae (L.)Desv. (1813) in J.Bot.appl., Paris 1:92. 
Notholaena subcordata (Cav.)Desv. (1813) in J.Bot.appl., Paris 1:92. 
Notholaena canariensis (Willd.)Desv. (1827) in Ann.Soc.linn., Paris 4:220. 
Pteris canariensis (Willd.)K.B.Presl (1836) Tent.Pteridogr. :145. 
Gymnogramme Marantae (L.)Mett. (1856) Fil.horti bot.Lips. :43. 


GEOGRAPHICAL DISTRIBUTION. Mediterranean region (Albania, Yugoslavia 
[Serbia], Italy, Sicily, Corsica, France, Spain, Cyprus, Syria) north to Austria 
and Switzerland, westwards to the Atlantic islands (Canaries, Cape Verdes 
and Madeira) and eastwards to the Crimean Peninsula [Chatyrdagh] and the 
Caucasus region. 


42 


Whilst the nomenclature of the first mentioned species does not imply 
any difficulty, the binomial -of the second, and also of two further European 


2:307, he was not aware of the fact that he had used this binomial for a quite 
different taxon in 1753; the homonymous binomial published in 1771 was, 
therefore, illegitimate at the time of its publication. The fact that the two 
homonymous species later on were placed in two entirely different genera, 
the taxon from 1753 in Dryopteris Adans. as D. fragrans (L.)Schott and the 
taxon from 1771 in the genus Cheilanthes Sw., is irrelevant. 
REICHARDT when editing the entirely new edition of the Systema 
Plantarum of LINNE was the “ee to realize the fact that LINNAEUS in 
1753 and in 1771 chose the same name for two taxonomically different 
species and proposed therefore, er correctly, for the taxon described in 
1771, a new name, Polypodium pteridioides Reich. Unfortunately, however, 
REICHARDT did not pay attention to the fact that the new proposed bino- 
mial was an illegitimate name because of the existence of the earlier homonym 
Polypodium pterioides Lam. (1778) Fl.frang.,ed.1,1:18.4 One year after 
T had published the new name for the taxon in discussion 
LINNE fil. transferred Polypodium fragrans L.(1771) to the genus 
Adiantum L. This first transfer, Adiantum fragrans L.fil. has to be considered 
as a nomen novum according to Art.72 of the International Code. In 1806 
SWARTZ published Cheilanthes fragrans Sw., which was considered by 
Practically all modern pteriodologists as being a different taxon from that 
Published by LINNAEUS in 1771 as Polypodium fragrans L. This inter- 
pretation of SWARTZ’ name, however, is only partly true. In fact SWARTZ 
was describing a quite different species from the East Indies which is known 
today as Cheilanthes opposita Kaulf., and excluded, though with some doubt, 
the taxon, called in modern publications C. pteridioides (Reich.) C.Chr. 
from the Island of Madeira. This latter taxon was confused with that from 
the East Indies for the first time by LINNE fil. when he published Adiantum 
fragrans L.fil. in 1781 and appears in SWARTZ’ Synopsis as two separate 
species, namely Cheilanthes odora Sw. and Cheilanthes suaveolens Sw. All 
this, however, is irrelevant, as SWARTZ based his Cheilanthes fragrans Sw. 
definitely on Polypodium fragrans L. (1771) and therefore automatically on 
Adiantum fragrans L. fil. Thus, all these names bear the same nomenclatural 
type, and SWARTZ had no right to exclude the type of the taxon. Though 
description and locality ascribed by SWARTZ to his Cheilanthes fragrans 
match a different taxon, the name has to stand with the nomenclatural type, 
and thus SWARTZ’ description and locality have to be excluded, and his 
combination must be accepted as the earliest legitimate name for the species 
in discussion: 
" Acording tothe Staples peace, neroidan) ced eae Bot Nomen 


1936 as “epithets treated as orthographic v: fern, 
adjecti tive, ie., a Medieval form composed of the latin words ee Hh casesep and the 


43 


2. Cheilanthes fragrans (L.fil.) Sw. (1806) Syn.Fil.: 127/325, excl.descr.et loc. 
Adiantum fragrans L.fil. (1781) Suppl.Plant., Syst. Veg., ed.13:447; nom.nov. 
Polypodium fragrans L. (1771) Mant.Plant.2:307; non L. (175 
ee <i! A alesamagy Reich. in L. (1780) Syst.plant., ed.noviss. si :424; non 
rp 778). 


Pteris ere Balb. (1801) Elenco piante cresc.cont.Torino :98 
Pteris fhiebges a fil.) Lagasca, Garcia & Clemente (1802) in Anal.Sci.nat., 


— pitiened Bory (1803) Essai Isles Fort., ant.Atlant.:311; sphalm. 
Allosorus pusillus (All.)Bernh. (1806) in Schrad.N.J.Bot.1(2) :36. 

Cheilanthes suaveolens Sw. (1806) Syn.Fil.:127. 

Cheilanthes odora Sw.(1806) Syn.Fil.:127/32 

woe suaveolens (Sw.)Poir.apud Lam. (1810) Encycl.méth., Bot., Suppl. 


ee odoratum Poir.apud Lam. (1810) Encycl.méth.,Bot.,Suppl.1:142; 


Adiantum eet (Sw. )DC.apud Lam. & DC. (1815) Fl.fran., ed. 3,5:237 


"bey A dia m”’). 
masgser ie ‘neropola of a (1819) in Abh.Preuss.Akad. Wiss. 1816-17: 
nud.; non Sw. (1806). 


pone “pail eee Poll. (1824) Fl. Veron.Prov.3:293 

—— pumilum [Bell., in hb., ex] Colla (1836) Herb.pedem.6:195, in 
observ. 

Cheilanthes maderensis 2 ages (1834), nomen nudum, ex] Lowe (1838) in Trans. 
Camb.phil.Soc.6 

pst es squamosa ra & Regel (1851) in Linnaea 24:174; non [Gillies, in 

ex] Hook.et Grey. (1829). 

Chet Avista ie (Balb.)Tod. (1866) Enum.Fl.sic.1:9; et in G.Sci.nat. 
Eco 

Cheianes Ti on P oe (1866) Enum.plant.sic.1:12; et in G.Sci.nat.Econ. 

o1:2 

Chelate ues sensu Nicotra (1891) in Malpighia 5:435; non Mett. 

Cheilanthes pteridioides (Reich. in L.) C.Chr. (1905) Index Fil.:178; non 
C. pteroi . (1806). 

Allosorus fragrans (L.fil.)Farw. (1931) in Amer. Midl.Natur.12:284. 


GEOGRAPHICAL DISTRIBUTION. Mediterranean region (Crete, Greece, Albania, 


icily, , France, Spain, Portugal, Balearic Is., Morocco, Algeria, 
Tonisia, Libya, eastwards through Asia Minor to Daghestan and the opposite 
mountain range near Dzhebel in Turkmenistan and to Afghanistan, the 
Iranian mountains and W. Himalaya (Kashmir). Also on the Canaries and 
adeira. 


A third European species was described by W.T.AITON (1789) Hort. 
Kew., ed.1,3:457 under the binomial Acrostichum velleum. This species was 
attributed to the genus Notholaena R.Br. by DESVAUX (1813). This com- 
bination, however, was a later homonym of Notholaena vellea R.Br. (1810) 


ae 


which is typified by an entirely different taxon. Also within the genus Cheil- 
anthes Sw. the specific epithet ‘‘ vellea’’ cannot be used, as there exists a 
transfer of R.LBROWN’s species to this genus by F. VON MUELLER (1866) 
Fragm.phytogr.Austral.5:123. 


Ten years after W. T. AITON had described his Acrostichum velleum, 
DESFONTAINES published [July 1799] the same taxon again under the 
binomial Acrostichum lanuginosum Desf., assuming it to be a new species, 
POIRET (1816) in LAMARCK, Encycl.méth.,Bot.,Suppl.4: 110 transferred 
DESFONTAINES’ species to the genus Notholaena R.Br., quoting as 
author of the new combination DESVAUX who, however, seems never to 
have published the combination Notholaena lanuginosa (Desf.)Desv. Whilst 
the specific epithet /anuginosa agrees with the provisions of the Code when 
the taxon is considered to belong to the genus Notholaena R.Br., it cannot be 
taken up under Cheilanthes Sw. as there exists a taxonomically different 
species C. lanuginosa Mart. & Gal. (1842) Mém.Acad.Brux.15:75/t.20.f.2. 


The earliest valid specific name for the species in question under 
Cheilanthes Sw. seems to be catanensis, first published as a supposed new 
species as Acrostichum catanense Cosentini in 1827. Thus, the following 
new combination is needed: 


3. Cheilanthes catanensis (Cos.)H.P.Fuchs comb.nov. 


Acrostichum catanense Cos. (1827) in Atti Accad.gioenia, ser.i,2:207/218. 
Acrostichum velleam W.T.Ait. (1789) Hort.Kew.3:457. 

Acrostichum lanuginosum Desf. (1799) Fl.Atlant.1:400/t.256 [July 1799]. 
Cincinalis vellea (Ait.)Desv. (1811) in Mag.natf.Frde, Berlin 5:312. 
ORR vellea (Ait.)Desv. (1813) in J.Bot.appl., Paris 1:92; non R.Br. 


Notholaena lanuginosa (Desf. [‘* Desv.’’, sec.]Poir.apud Lam. (1816) Encycl. 
méth.,Bot.,Suppl.4:110. 
Notholaena Plukenetii Fée (1852) Gen.Fil.:159. 
Gymnogramme lanuginosa (Desf.)A.Br. (1854) Ind.sem.horti reg.Bot.Berol. 
1854:17. 


spipstcrnata — (Cos.)Tod. (1866) ue Pi: —. Vasc.sic.:14; et in 
nat., econ., Palermo 1(3/4):221, in obse: 
Costin vel he )Tod. (1866) a Pl. ee Vasc.sic.:14; et in Giorn. 
., Palermo 1(3/4):2 
Siuccacab ne (Ait.)Kuhn a Fil. Afr. :61. 


GEOGRAPHICAL DISTRIBUTION. Mediterranean region (Crete, Greece, Italy 
as far north as Salerno, Sicily, Corsica, Spain, Portugal, Morocco, Algeria, 
Cyprus, Jordan, Palestine) expanding south to Ethiopia and east to Afghan- 
istan. Also on the Canaries and Madeira. 


45 


The fourth species of Cheilanthes occurring in Europe is C. persica 
(Bory apud Bélanger)[Mett.in litt.ex] Kuhn and it is not at all clear when 
the basionym, Notholaena persica Bory apud Bélanger, was published. In 
accordance with FOMIN apud KOMAROV 1934, FI.SSSR 1:76 all modern 
Russian floras cite 1833 as the date of publication of the second part of 

LANGER’s Voyage aux Indes-Orientales without any remarks, whilst 
FIORI 1943, Fl.ital.crypt.5:272, also adopting this date of publication for 
Notholaena persica states in a note that the date of publication of 
BELANGER’s work is not at all certain. PAMPANINI 1906, in Nuovo 
G.bot.ital.N.S.13:139-157 used the binomial Cheilanthes Szovitsii® Fisch.v. 
Waldh. & C.A. Meyer apud Hohen. throughout his paper and discussed at 
length the date of publication of BELANGER’s undated publication. 
KUHN, in Bot.Ztg.26:233 adopted 1833 and in Fil.afr. :73, 1831 (in error ?), 
RUPRECHT 1845, Beitr.Pfzenkde.Russ.Reiches 3:48 cited 1833 as the date 
of publication, however with doubt, whilst MILDE 1867, Fil.eur.atlant.:33, 
as well as NYMAN 1883, Consp.Fl.eur., Suppl.1:861 cited as the date of 
publication of Notholaena persica Bory apud Bélanger the year 1838, and 
cB CLA 1880, in Trans.Linn.Soc., Lond., ser.2,Bot.1:455 pointed 
out, that the binomial Notholaena persica was published four years after the 
publication of Cheilanthes Szovitsii, i.e. in 1842. PRITZEL 1872, Thes.lit. 
bot., ed.2:20, and SHERBORN & WOODWARD 1901, in Ann.Mag. 
nat.Hist., ser.7,7(40):390, and 1901, in J.Bot.,Lond.39:205 accordingly 
accepted as the date of publication for the cryptogamic part of BELANGER’S 
Voyage the year 1846, the latter authors pointing out, that according to a 
note in the Bibliographie de France in 1834 parts 2 and 3 of the work in 
discussion were published, but contained only plates and wrappers. Recently, 
a note appeared in the Fl.Males.Bull.15:730—-731 [August 1960] pointed out, 
that according to a review by BRANDES 1835, in Lit.Ztg.1835(4) :69 [January 
21, 1835] two printing sheets (i.e. pp.[1]}-32) and eight plates were already 
published before this date. Therefore, it is evident, that the binomial 
Notholaena persica Bory appearing in the second, and only published volume 
of BELANGER’S Voyage aux Indes-Orientales on page 23 has priority overt 
the name Cheilanthes Szovitsii Fisch. vy. Wald. & C. A. Meyer which indeed 
appeared as early as in December 1833 in a paper by HOHENACKER, 
entitled “* Enumeratio plantarum in territorio Elisbethopolensi et in provincia 
Karabach sponte nascentium,’ however only as a nomen nudum, but was 
validly published with a Latin description only in June or July 1838 in another 
paper by HOHENA , ‘‘ Enumeratio plantarum, quas in itinere per 
provinciam Talysch collegit Fr. R.Hohenacker”’: 


5 Dedicated to JOHANN JOSEF coat AL (-x made Coarmenioge: in Odessa, who collect ted 


plants in tinea — cia, Southern coe 5 as w een 1828 and os in ea yong Persia 
and the Caucasus, where in the region of wi ae he poe found d the species him 
6 That et —_ of publication of goa we $s work as accepted b ae as W cad as ce 
modern ssian floras and FIORI is correct, has been proved recently. «4 psi ox Seg (arith 
Museum (Natural History)] who very Kindly communicated the resul will 
be published shortly [cf. ROSS 1961, in J.Soc.Bibl.nat.Hist. 4:(in press tg 


46 


4. Cheilanthes persica (Bory apud Bélanger) [Mett., in litt., ex] Kuhn 
1868) in Bot.Ztg.26:233 [April 3, 1868]; Fil.Afr.:73. 
tate toe persica Bory apud Bélanger (1833) Voy.Indes. Orient., bot.2:23 
ochlaena”” {before March 25th, 1833]. 
pate Szovitsii iets v.Wald. & C.A.Meyer apud Hohen. (1833) in 
Bull.Soc. Nat. Moscou 1833(6):260, nomen nudum,, ex}Fisch.v. bbe d 
CUA: Meyer apud Hata (1838) in Bull Soc Nat.Moscou 1838(3):24 
cum des 
Cheilanthes Aas Vis. (1842) Fl.Dalm.1:42/t.1.f.1. 
Acrostichum microphyllum A.Bertoloni apud G.Bertoloni (1856) Propag. 
ric.6:343. 


Oevosporangium fimbriatum (Vis.) Vis. (1867) in Atti Ist.veneto, ser.3,12:656. 
Ocvosporangium Szovitsii SE ers v.Wald. & C.A.Meyer apud Hohen., 

nudum, ex]Fisch.v d. & C.A.Meyer apud Hohen.) Vis. (1867) i in Atti 
Ist. veneto, ser.5, 12: weS6 10-11. 


Ocvosporangium persicum (Bory apud Bélanger) Vis. (1872) in Atti Ist.veneto, 
ser.3,16:44, 


Notholaena_ Szovitsii oe v.Wald. & C.A.Meyer apud Hohen 
lum, ex]Fisch.v.Wald. & C. A. es apud Hohen. )Keyserling (1873) 
Polypod.et Cyath. Herb. Bungeani 
Mpyriopteris Szovitsii ({Fisch.v.Wald. & C.A.Meyer apud Hohen., nomen 
ee ex]Fisch.v.Wald. & C.A.Meyer apud Hohen.)J.Sm. (1875) Hist. 
Fil. :281. 


GEOGRAPHICAL DISTRIBUTION. Mediterranean region bondatahiie ti 
Yugoslavia [Dalmatia, Montenegro, Hercegovina], Aegean Is.) southwar 

Igeria eastwards through Turkey, Crimea, Iraq, Transcaucasia and Periia 
to Central Asia (Turkmenistan, Tien-shan, Pamir-Alai) and to the Himalayas 
(Kashmir and Punjab). 


5. Cheilanthes hispanica Mett. (1859) Abh. senckenb. natf.Ges.3:74. 


Cheilanthes fragrans var. hispanica (Mett.) Fiori (1943) in FLital.crypt. 
:270, in adnot [February 20th, 1943]. 


GEOGRAPHICAL DISTRIBUTION. Ranging from the south-western part of the 
Iberian peninsula [Portugal and $.W. Spain (Estremadura Mts.)] to Spanish 
Morocco and Tunisia. 
SUMMARY 
The generic value of Notholaena R.Br. is somewhat questionable and it 
seems that probably all species of this genus should be included in the genus 
Cheilanthes Sw. As to the two European species so far included in the genus 
Notholaena R.Br. the general taxonomic problem is irrelevant as both are true 
Cheilanthes in the strict sense. It is argued that the latter generic name does not 
need to be conserved against A//osorus Bernh., the latter by reason of typifica- 
tion not being congeneric with Cheilanthes Sw. Two out of the five European 
species bear generally used names which do not agree with the provisions of 
the International Code of Botanical Nomenclature. The following chinges 


47 


in nomenclature are therefore necessary: the species generally known as 
Cheilanthes pteridioides (Reich. in L.) C.Chr. has to be called Chelate 
fragrans (L. fil.)Sw.; the species generally called Notholaena vellea (W.T 
Desv.,non R.Br. needs a new combined name, Cheilanthes catanensis a 
H.P.Fuchs. General distribution of, and a key to, the five species is included. 


THE SOUTHPORT SPRING FLOWER SHOW, 1960 


Mr. Tom Bolton adjudicated Classes 8-20 (the fern classes) as in previous 
years. The following is a list of the prize-winners submitted by Mr. Bolton 
and, where it has been possible to obtain the information, names of the plants 
exhibited; the comments in quotation marks are those of Mr. Bolton. 


CLASS 8_ A group of hardy British ferns. 
Ist prize and the Challenge Cup: Mr. John Brookfield — “ the plants were 
very clean and well grown. I noted one plant of Victoriae that was 
age 


CLASS 9 Six ee ferns. 
Ist: rin —_ ookfield —‘‘... group included a very good plant of 
2nd: Mr. 8 B. Hayhurst, 
3rd: Mr. J. Pye. 
CLASS: 10 —— Scolopendriu 
Ist Mr; Hayhurst — Ss. “wlgare sana scolopendrium) crispum, S.v. 


_erspum "Drammen S.v 
2nd: Mr. J. Pye. 


CLASS: 11 Three Polypodium 
Ist: Mr. J. Bro obfield these were in large pots and an outstanding 
exhibit. 
2n . Ha ii igh sare vulgare cambricum, P.v. cristatum 
"Forestil, P.v. cong 


abt a Three Polystichum 
Hayhurst — Potpstictiaal setiferum divisilobum densum, P.s.d. 
a es oe gracile. 
2nd: Mr. J. Brookfield. 
3rd: Mr. J. Pye. 
CLASS 13 Three Athyriums. 
Ist: Mr. J. Brookfield. 
2nd: Mr. N. H. Almond. 
3rd: reo : . Pye Rg es evonly g vulgare crispum, S.v. crispum Drummondii, 
Athyrium f.-f. Victori A. plumosum Druery, Polystichum setiferum 
sulcherrbhae Bevis, Ps. Geri: densum. 
get 14 Three Lastreas. 
Mr. J. Brookfield. 
ad Mr. B. Hayhurst — Lastrea (Dryopteris) pseudomas ramossima, L.p. 
re —— aie ZS. Y eikecseled, filix-mas grandiceps. 
r 


CLASS: 15 thee oe apr 
Ist: Not giv “This was rather a poor class for exhibits.” 
2nd: Mr. J. Br rookfield. 


48 


ASS. 16 ees ebardy 4 British ferns. 
eh S 


one . Robinson — Cryptogramma crispa, Dryopteris aemula, Poly- 
‘stichum setiferum. 
3r Mr. B. Hayhurst — Adiantum capillus-veneris, Ceterach officinale, 


* Phyllitis picts ee (Scolopendrium vulgare). 


Soarag 17 ae Rab British fern 
Mr. ayhurst — Polystichum setiferum diversilobum plumosum. 
oa: Mr. 5 


3rd: Mrs. M. L. Bassnett — Scolopendrium vulgare crispum Campbellii. 
aes 18 Six greenhouse ferns. 
Mr. J. Brookfield. 


medi r. J. R. Eaves. 
3rd: wei B.H ayhurst — Adiantum “ Kensington Gem,” Onychium japonicum, 
vallia oes Pellaea rotundifolia, ‘Nephrolepis exaltata, Nephrolepis 
at ee elegantiss. 


CLASS 19 Three greenhouse ferns. 
1st; Mr. J. ap nese 
2nd: Mr. J 


CLASS 20 One greenhouse fern. 
Ist: Mr. J. Brookfield. 
an. Not give 
3rd: Mr. B. Hayburit — Adiantum ‘* Kensington Gem.” 


SHORTER NOTES 
THE REDISCOVERY OF ASPLENIUM x MURBECKII Dorfl. 
When botanising some years ago on Castle Crag in Borrowdale I 
noticed a very small stunted fern with fronds about an half an inch long 
growing on a ledge of the iron-hard volcanic rock almost out of my reach. 
However, I managed to get a couple of fronds and saw immediately that it 
was an Asplenium. Well grown specimens of A. trichomanes, A. septentrionale 
and A. ruta-muraria were in the vicinity and on first examination I took the 
fronds to be those of A. x alternifolium Wulf. (A. septentrionale x A. 
trichomanes). Later in the season I again visited the locality, this time with 
my son, who after a short rock-climb secured a good plant. I was now able 
to have a good look at the plant and I noticed that the apex of the pinnules. 
was very finely toothed and not rounded as in alternifolium and I began to 
have doubts as to its precise identity. However, I eventually planted it on 
a ledge in the fernery where it stayed for some time but it never increase 
Stature. Recently I decided to pot up the plant in porous volcanic scree a 
leaf-mould, both of which we have plenty at hand, and in this the fern grew 
Strongly and soon began to show its true character. 

It was then that I consulted my never-failing source of information the 
octavo edition of Thomas Moore’s Nature Printed British Ferns and found 
that A. ruta-muraria var. cuneatum T. Moore (now A.  murbeckii Dorfl.) 
fitted my plant very well both in description and in figure. I sent a specimen 
to Mr. Jermy who agreed that it could well be murbeckii and he sent some 
material on to Dr. D. A. Meyer (at the Botanic Garden and Museum, 


49 


Berlin-Dahlem) who has worked on Asplenium hybrids for some years and 
he confirmed the identification. Part of my original plant is now in cultivation 
at Chelsea Physic Garden. ; 
FRED JACKSON 
TRANSPLANTING FERNS 
Ferns are very accommodating plants and can be transplanted successfully 
at any time of the year, even during the hottest weather. Major alterations 
to my fern borders this year compelled me to move, during our warmest and 
driest spell of weather early in the summer, a very large growing plant of 
Dryopteris borreri with fronds almost four feet long. I did not hesitate to 
divide the several crowns although the root system suffered in the process. 
Given my usual treatment and well watered in, the fern settled down happily 
with no wilting of the fronds, and new ones were soon being thrown up. 
This habit of ferns is invaluable to the fern-hunter, who can confidently 
excavate a treasured find from the hedgerow at any season of the year and 
establish it successfully in the garden. 

Most growers will probably lift ferns carefully with as much root as 
possible and replant them with a minimum of disturbance, and some of my 
fern friends have looked rather doubtful when I have explained or demon- 
strated my method of dealing with ferns when transplanting or introducing 
them from the wild. As it has proved very successful with me, it may be 0 
interest to others. 

First, the plant is placed in a bucket of water and all the soil washed away 
from the roots so that I can see what I am doing. Buried among the roots of 
a large plant, particularly if it is an old one, will usually be found a length of 
dead stem which is broken off, and then beginning from the bottom all the 
old frond bases right up to the living fronds are pulled off. With big strong 
growing ferns I find a pair of large pliers very helpful for this task. Care 
should be taken to remove as few roots as possible; these grow from the 
bases of the fronds and the loss of some is unavoidable but has no adverse 
effect. 

It is remarkable how small a large fern rootstock becomes after this 
treatment, but replanted carefully and well watered in, it is soon growing 
vigorously again. With the removal of all the old stumps the new roots 
rom the bases of the living fronds are able to move easily and quickly into 
the surrounding soil and the growth of the plant benefits accordingly. 

J. W. DYCE 


POLYSTICHUM ACULEATUM PULCHERRIMUM BEVIS 

This beautiful fern was found in 1876 and for over 30 years was reputed to 
be barren, until a few spores were found on plants i in the possession of MS 
C. T. Druery and Dr. F. W. Stansfield. S 
able results, including Polystichum acilesituns. pulcherrimum Drueryi, P. 
aculeatum plumosum Greenii, and the P. aculeatum gracillimums. The first 


50 


two are strong-growing outdoor ferns, but the gracillimums are, to quote 

T. Druery, ‘“ of wonderful delicacy of cutting, the parental half-inch 
pinnules being extended sometimes to nearly three inches, and of almost 
hairlike tenuity, with splayed tassel-like terminals. 

No further recorded finds of spores on ‘* Bevis ’’ have been made since 
before the first World War, although there is no doubt that plants have 
borne spores since then. I have grown the fern since 1952, and each year 
have examined my plants hopefully but in vain. However, this summer 
when working in my fern garden, I lifted a frond of “ Bevis,” and to my 
utter amazement found myself looking at undoubted sori. On two adjacent 
pinnules on a pinna near the apex of the frond, I counted eight in all. Un- 
fortunately the discovery was made rather late, for most of the sporangia 
had dispersed. A large number still remained however, and the pinna was 
carefully broken off and placed in a folded paper. A few days later the spores 
were shed and were immediately sown in a prepared pan. 

At the time of writing I have a prolific crop of prothalli from which I 
hope to obtain many good ferns, and possibly something entirely new. 

YCE 


HERBARIUM SPECIMENS OF FERNS AND THEIR ABERRATIONS. II 
Following the policy outlined in the last number of the Gazette (pp. 26-27), 
r. T. A. Dyer has made a collection of pressed fronds from ferns growing 
in his garden at Alrewas. He pursued this self-imposed task with great 
enthusiasm and has made an excellent job of preservation and labelling and 
has provided a plan of his garden with specimen locations marked with their 
numbers ; 96 specimens handed over at the 1960 Annual General Meeting 
with a promise of more to come. Five specimens were sent by Mr. R. Kaye 
in November. 
It is hoped that this building up of the Society’s herbarium will continue 
with the sending of named and well-labelled specimens, especially fronds 
from prize-winning plants. 


J. A. CRABBE 


BOOK NOTES 
‘* BRITISH FERNS AND MOSSES ” 
By P. G. TAYLOR pp. 1-231 illustr. 
“* The Kew Series ” (Eyre and Spottiswode, 1960) Price 25s. 


‘* British Ferns and Mosses ” is the fourth volume of “‘ The Kew Series ” to 
appear and it maintains the high standard of production as others in this 
series of which R. D. Meikle’s “‘ British Trees and Shrubs ” is perhaps most 
well-known. Perhaps it should be said at the outset that it was the publishers’ 

unfortunate decision to include the mosses and this presented the author 


51 


with the uneviable task of choosing one-twelfth of our British species. It 


can be of little help to the beginner; but it is for what Mr. Taylor has written 
on ferns and their allies that this book will be considered. 

As in the previous volumes there is an Introduction [pp. 9-12] which 
acquaints the reader with the botanical terms used and with the life-histories 
of the fern (and moss) plant. Since there is a Glossary [pp. 220-226] it seems 
a pity to have confused the text throughout with definitions which often 
lack uniformity and become ambiguous [e.g. p. 15, “‘ Sori . . . are of several 
distinct shapes . . . kidney-shaped or reniform (as in Dryopteris . .), oval or 
linear (as in Asplenium..)...’]; the use of italics is confusing [why ‘‘ modified 
one — ne : 1. 162]. And is “ Hybridisation in the vascular 

togams ... of rare occurrence”? In ail our larger genera (eg. Dryopteris, 
ae pik a hybrids are formed frequently between species 
living in the same habitats. Apogamy and apospory are not discussed. 

The greater part of this book (pp. 23-176) is devoted to the description 
of the majority of our species of ferns and related groups, together with 
data on habitat preferences useful to all who grow ferns. There is a single 
key to all species mentioned; it might have been more workable to have 
separate keys for the major natural groups. In that excellent part of the 
key dealing with Equisetum Mr. Taylor makes it apparent that he has studied 
the genus in some detail; on the whole it is a practical key but I think the 
inclusion of geographical distribution at any stage in a key is dangerous 
because if a reader sees that his specimen is outside the range given he will 
rarely check its morphology (this could arise over Ophioglossum lusitanicum 
and O. vulgatum ssp. ambiguum). It would have been worthwhile to include 
in the key the odd species of flowering plant that could be confused with 
Isoetes (eg. Littorella uniflora) — in the latter species the leaf section (easily 
seen with a hand lens) is made up of several tubes; in Jsoetes there are four 
tubes only. 

It is true to say that, for the most part, the taxonomy of our British 
species of Pteridophyta is well known but I find it disappointing that Mr. 
Taylor has virtually ignored the fundamental work done during the last 
15 years by Professor Manton and her colleagues at Leeds University. It is 
true that the Introduction would have had to include a short discourse on 
chromosomes and the importance of meiosis but so much more could then 
have been discussed. On p. 166, under Polypodium, “‘ a form” is described 
which is in fact the diploid (P. australe Fée) and Taylor says “‘ results of the 
[recent] investigation suggest that it should perhaps be treated as a distinct 
variety or even a distinct species, though there seems to be some doubt as 
to its correct name.” Here is an example that clearly shows how cytological 
evidence has helped the taxonomist and there is an interesting story to be 
told if only Mr. Taylor could have mentioned chromosomes and meiosis. 


52 


Again in the genus Dryopteris, the work of Walker in the dilatata-spinulosa 
complex has been forgotten and in the D. filix-mas complex D. abbreviat 

(DC.) Newm. is omitted; Lovis’ work on the synthesis of Asplenium x 
alternifolium (Breynii) has, | am glad to say, been briefly mentioned [p. 118]. 

As the author says in the introduction he has based his classification 
(and presumably his nomenclature) on Warburg in Clapham, Tutin and 
Warburg, 1952, but he deviates, I am pleased to see, in upholding Gymmo- 
carpium [p. 161] with two species, G. dryopteris (L.) Newm. and G. 
robertianum (Hoffm.) Newm. both of which were placed in Thelypteris in 
Warburg. There is a minor slip on p. 133 where Linneus is given as the 
authority for D. filix-mas instead of (L.) Schott. 

The book is beautifully illustrated in both line and colour by Miss Ann 
Webster. Many of our species could be identified from these alone but there 
are two places where I feel the artist has been misdirected. A good character, 
mentioned by Taylor and used to distinguish between Equisetum arvense and 
E. palustre is the length of the lowest internode of the branch in relation to 
the length of the main stem nodal sheath; this could have been made clear 
in the magnified inset drawing on Plate 2 [E. arvense] and similarly in Fig. 9 
[E. palustre] if the total length of the lowest internode had been figured. In 
Plate 6 the magnified ‘‘ base of pinna ” labelled D. borreri does not show at 
all well the truncate apex to the pinnule; in fact the magnified drawing of the 
single pinnule of D. filix-mas is more typical of borreri. The absence of a 
scale in all the drawings is something for which Miss Webster cannot be 
forgiven; Lycopodium annotinum [Fig. 2B] is surely bigger than Selaginella 
Selaginoides [Fig. 3]. 

One can appreciate Mr. Taylor’s difficulty in putting over a modern 
concept of fern taxonomy to the non-specialist and there are occasions 
(e.g. in the descriptions of variants within a complex) where he himself is 
perhaps unsure. In such circumstances he has been wise to approach the 
subject as he had and what he has done is commendable. In spite of its 
seemingly high price, it is a book worth owning. 

A. C. JERMY 


FARNE IM HERZEN EUROPAS 
By GEORG EBERLE pp. I-VIII, 1-116, illustr. 1959 


Obtainable from W. Kramer and Co., Frankfurt am Main, Germany 
ice 


This book on Central European ferns is published as one of a series of 
Natural History Handbooks by the Seckenberg Natural History Society. 
The first part of the book contains sections on the structure and life-history 
of the ferns in general and this is followed by a section on re. taking 
into account the cytological work of Professor Manton and D. E. Meyer. 
The descriptive pages (21-40) show that the author is an obmervant field 
botanist and the 88 first-class photographs of these ferns in their natural 


a3 


habitats strengthen this view. Forty-four of our own native species are 
illustrated in 67 photographs and this alone makes this book a useful posses- 
sion of the British fern-lover. 

It should be noted that in a recent review Dr. D. E. Meyer pointed out 
that the lower photograph on p. 102 is not Polystichum x illyricum as 
labelled but is P. lonchitis. The shape of the frond and the inflexed lower 
pinnae of the Polypodium on p. 109 suggests that this is the hexaploid species. 

A. C. JERMY 


‘*THE FERNS AND FERN-ALLIES OF BRITISH COLUMBIA ” 
By T. M. C. TAYLOR pp. 1-154, illustr. 1956. 


Obtainable from the Provincial Museum of Natural History 
and Anthropology, Victoria, B.C., Canada 
Price 75 cents (incl. postage). 


This is No. 12 in the series of Handbooks — by the British Columbia 
Provincial Museum Department of Education. It is a neatly produced 
pocket-size book which covers a region not too dissimilar in climate and 
flora from our own and about 40 of our species are described and figured 
(by clear line drawings) together with ecological notes. One of the assets of 
this book to the British pteridologist is that it gives him a wider concept of 
our smaller genera. 
A. C. JERMY 


THE BRITISH PTERIDOLOGICAL SOCIETY 
THE ANNUAL EXCURSION 1960 


The 1960 Annual Excursion was held in North Somerset from the 3rd to 
11th September with headquarters at the Egremont Hotel, Williton. Ten 


Saturday, and we were joined on the Wednesday by Mr. and Mrs. T. A. Dyer 
and Mr. L. Burnett, and on the Friday by Mr. P. Temple. Fine weather 
favoured us and on only one day, Tuesday, did it rain. It so happened we 
planned to spend this day visiting some habitats of bog ferns, and they 
were seen under conditions very much to their liking, if not to ours. 

Two previous excursions in recent years have been held in this area 
comprising the Brendon and Quantock Hills, but not much ground can be 
covered in a week, and after this third visit there still remain many good 
places awaiting our future attention. As usual in the West Country, the 
hunting was chiefly among Polystichum setiferum, with the hope ever constant 
in our minds and undimmed by past failure, that we would come upon @ 
good variety of this fern which has produced so many of the finest known 
varieties. 

54 


There seems to be some magnetic attraction for us in a certain deep valley 
in the Brendon Hills. We commenced our last Excursion to the area in 1957 
with a visit to this spot which in 1956 had given us some good hunting. Again 
this year the decision to start here was unanimous. Nothing new was found, 
but some of the party were exploring the West Country for the first time, and 
it was pleasing to introduce them to this wild fern garden which has so much 
to show. We have recorded 14 species from this valley, some of them but 
not Polystichum setiferum, growing in great abundance. Dryopteris aemula 
is found here, a beautiful fern which never fails to arouse admiration in the 
beholder, and contributes greatly to the attraction of the place. 


During our previous excursions to North Somerset, most attention has 
been given to the Brendon Hills area, but this year we concentrated more on 
the Quantocks. A network of lanes run up and along the west side of these 
hills, most of them unexplored by us, and we decided it was time to remedy 
this omission. They are rich in ferns but in very few did we find more than 
nine or ten species. P. setiferum was common everywhere and provided 
interesting hunting. In one lane near Crowcombe the only plants of 
P. aculeatum seen during the week were found. There were only a few, but 
it was observed that most of the P. setiferum in the lane seemed to exhibit 
some P. aculeatum characters. Several of the plants were of vigorous growth 
with very wide crispate and foliose tripinnate fronds, and some good 
specimens were collected. It was near this lane that the find of the week was 
made by Mr. Greenfield — an incised form of Asplenium trichomanes. 


The one wet day was spent very appropriately further afield in the flat 
country beyond Bridgwater, in search of bog and water ferns. This low 
lying district is drained by a network of rhynes or ditches, in some of which 
the water fern Azolla filiculoides is to be found. One good colony was dis- 
covered, and near it, the green dust covering the surface of the water was 
found to be an extensive colony of the rare Wolffia arrhiza, the smallest known 
flowering plant. Near Shapwick, Thelypteris palustris is literally a weed 
infesting the moor, but we noted that in the open it was not so prolific as 
On our previous visit, and it was suggested that the exceptionally dry summer 
last year may have dried up the moor sufficiently to kill it. In the woodland 
nearby this fern grows rampantly with fronds three feet high, along with 
Dryopteris lanceolatocristata (spinulosa) and Osmunda regalis. Floundering 
in a boggy wood, and forcing a way through the undergrowth in a downpour 
of rain, could not be regarded as pleasant even by the most enthusiastic 
hunter of Royal Ferns however, and after finding two magnificent plants 
we felt we had achieved sufficient to excuse a quick retreat. 


Two more days were spent outside our immediate hunting grounds — 
on the northern edge of Exmoor. This was new territory which we wanted 
to learn something about for future visits; we followed up a record for 
Hymenophyllum tunbrigense which led us into a fine wooded combe south 
of Porlock and what we saw of the lanes between Dunster and Porlock 


55 


augered well for an excursion in this district. We did not find the Hymeno- 
phyllum but had an enjoyable day’s hunting with a bag of 14 species which 
included a large flourishing colony of D. aemula. 

During the week we recorded 20 fern species. 


“Osmunda regalis L. — Shapwick Heath. 
Pteridium aquilinum (L.) Kuhn — common everywhere. 


Blechnum spicant (L.) Roth — Druid’s Combe, Hawkcombe, Kingsbridge, Nettlecombe, Riscombe 
Exford. 


Sea scolopendrium (L.) N — Crowcombe, Hawkcombe, Holford, Huish Barton, — oa 
gsbridge, Se aicaifcen,: Metlacsebs, Riscombe Exford, Stogumber Station, Westwood. 
LPs adiantum-nigrum L.— Crowcombe, Hawkcombe, Holford, Huish Barton, rishi 
Kingsbridge, Monksilver, Stogumber Station, Westwood. 


A. {ieee L. — Crowcombe, Hawkcombe, Holford, Huntscott, Kingsbridge, Monksilver, Ris- 
mbe ae psbabie es Station. 


A. ruta-muraria L. — Crowcombe, Hawkcombe, Holford, Huntscott, Kingsbridge, Stogumber Station, 
Ceterach era DC. — Crowcombe, Holford, Shapwi 


Aen | Mere -femina (L.) Roth — Crowcombe, se ond! Combe, Hawkcombe, Holford, Huntscott. 
ridge, pape Pre: AVE REI Riscom 
Dorie i «mas (L.) Sc — Crowcombe, engine guns Hawkcombe, 
Huish Barton, ‘Sun tscott, Kingebridgs, Monksilver, Nettlecombe, *Riscombe Exford, 
Bes tation, Westwood. 


D. oO Newm., — Cua Druid’s Combe, Hawkcombe, Holford, Huntscott, Monksilver, 
Nettlecombe, Riscombe Exford, Stogumber Station. 


D. lanceolatocristata (Hoffm.) Alston — Shapwick Heath. 


D. dilatata (Hoffm.)A.Gray — Crowcombe, Druid’s Combe, Hawkcombe, Holford, Kingsbridge, 
Monksilver, Nettlecombe, Riscombe Exford 


D. aemula (Ait.) Kuntze — Druid’s Combe, Hawkcombe. 
Polystichum_setiferum (Forsk.) Woynar —Crowcombe, Druid’s Combe, Hawkcombe, imate’ 
Huish Barton, ee Kingsbridge, sicaksibver, Nettlecombe, Stogumber Station, Westwoo 
culeatum — Crowcombe. 
Thelypteris into (All.) H.P.Fuchs — Druid’s Combe, Hawkcombe, Riscombe Exford. 
T. palustris tt — Shapwick Heath. 
nha ames satioe L. — Crowcombe, Druid’s ee Biscdelg we Holford, Huish Barton, Hunts- 
t, Kingsbridge, iMooksilver, Nettlecombe, Riscombe Exford, Stogumber Station, Westwood. 
pe piace Lam. — Burrough Bridge. 
The vice-county numbers and grid-references of the localities quoted 
are as follows; Burrow Bridge v.c. 5, 31/3530; Crowcombe v.c. 5, 31/1436; 
Druid’s Combe v.c. 5, 31/0037; Hawkcombe v.c. 5, 21/8745; Holford v.c. 
5, 31/1541; Huish Barton v.c. 5, 31/0538; Huntscott v.c. 5, 21/9243; 


1/07 
v.c. 5, 31/0537; Riscombe Exford v.c. 5, 21/8339; Shapwick v.c. 5, 31/4137; 
Shapwick Heath v.c. 5, 31/4137; Stogumber Station v.c. 5, 31/1137; West- 
wood v.c. 5, 31/1534 


Very few varieties were collected, but this year we can record one first- 
rate find — Asplenium trichomanes “ incisum’ found in the Crowcombe 
area by Mr. Greenfield, growing on a roadside wall in a large colony of 
normal forms of the fern. Some very fine incised forms of A. trichomanes 
have been found in the past, but as far as is known are no longer in existence, 
and Mr. Greenfield’s find is probably superior to any incisum in cultivation 
today. It is intended to put a frond into the Society’s Herbarium. Other 
finds included a colony of A. trichomanes with terminal forking of which 
several were collected by the party, and some plants of crispate and foliose 
P. setiferum. 

J. W. DYCE 
56 


REPORTS OF MEETINGS 1960 


THURSLEY AREA, SURREY, SUNDAY MAy 22Np. Eight members attended this 
meeting which met at Godalming B.R. Station at 10 a.m.; from there 
transport was by means of private cars. We made our way S.W. on the A3 
road and explored first some wet woods close to the Thursley road turning. 
Several ferns were in abundance and of particular note were Dryopteris 
lanceolatocristata, D. dilatata and, by all appearances, the hybrid between 
the two. The first-mentioned species was definitely less developed, whereas 
the dilatata exhibited full crowns of open fronds. We continued to Thursley 
Bog to search for Lycopodium inundatum, L. clavatum and L. selago, all of 
which were recorded for Thursley by C. E. Salmon early this century. Only 
L. inundatum was found however, and this was frequent in the wetter hollows 
which had escaped burning earlier in the year. The afternoon was spent 
at Waggoner’s Wells, just over the Hampshire border, and here a number of 
Dryopteris were observed in an attractive setting of Montia sibirica in full 
bloom. Throughout the day a number of stone-brick bridges were investi- 
gated with the hopes of finding Ceterach officinale, but in vain. 


TUNBRIDGE WELLS, SUNDAY JUNE 26TH. Five members attended this outing 
to the wooded district south of Tunbridge Wells. At Eridge Green we 
recorded ten fern species, including Hymenophyllum tunbrigense on the 
sandstone rocks. The common fern was Dryopteris dilatata, and a fine 
colony of D. borreri was found. The other species were Polypodium vulgare 
growing on the rocks, Dryopteris filix-mas, Athyrium filix-femina, Pteridium 
aquilinum and a few plants of Thelypteris limbosperma, Asplenium adiantum- 
nigrum and Blechnum spicant. We hoped to find Dryopteris aemula at 
Saxonbury Hill but were unlucky. The six species found here were P. 
_ A. filix-femina, D. dilatata, D. lanceolatocristata, B. spicant and 
T. limbosperma. 


We finished the day at Mayfield in search of Polystichum aculeatum but 
again were unlucky 


BRENTWoop, Essex, SATURDAY OCTOBER Ist. This outing was held jointly 
with the London Natural History Society in Thorndon Park Woods. It 
rained most of the time but our party of ten, which included six of our 
members, had an enjoyable day. Ferns were found only in small areas and 
we recorded only seven species. Athyrium filix-femina was abundant and 
the other common species were Dryopteris filix-mas, D. borreri and D. 
dilatata. One colony of D. lanceolatocristata was found and a visit was 
paid to Essex’s only recorded colony of Polystichum setiferum which consists 
of about three plants. 


NORTHERN MEETINGS. Two meetings were held in the North, the first at 
Bakewell on August 13/14th, but only one member turned up to support the 
Meetings Secretary. The second held in Borrowdale on October 8/9th was 


57 


more successful and the five members participating agreed that in spite of 
wet weather it was well worth while. A total of 22 species was found, and 
included Osmunda regalis, Hymenophyllum wilsonii, and the rare Asplenium 
septentrionale. Quite a few plants of the spleenwort were seen, and the 
filmy fern covered the rocks in great sheets. 


EXHIBITS AT THE A.G.M. Some well-grown potted ferns were shown by the 
President, including Thelypteris limbosperma, T. phegopteris and Polypodium 
vulgare collected durning the Excursion to Ambleside in 1959. 

Mr. Greenfield exhibited his Asplenium trichomanes “ incisum” found 
during the 1960 Excursion to Somerset and the Secretary also showed 
fronds from the same fern, as well as the pinna from his plant of Polystichum 
aculeatum pulcherrimum Bevis showing the rare phenomenon of sori on this 
fern. 

Mr. J. F. Jackson from Borrowdale had two very interesting exhibits, 
fronds from an Asplenium septentrionale hybrid with A. ruta-muraria (A. X 


murbeckii Dorfl.) and a narrow form of Dryopteris dilatata. 
J.W.D. 


OBITUARIES 
EDWARD ARTHUR ELLIOT 

The death of the Rev. E. A. Elliot on the 19th February, 1960, was briefly 
announced in a notice sent with the last issue of the British Fern Gazette. 

Mr. Elliot had been Editor of the Gazette since 1949 and Secretary of 
the Society since 1950, holding both Offices until 1959, when failing health 
forced him to resign. He had in fact needed medical treatment intermittently 
for some considerable time, but made light of it and carried on his various 
duties with great determination. He was able to continue local walks for 
botanical purposes and recorded a large number of plants for the Mapping 
Scheme of the Botanical Society of the British Isles. 

Elliot was born in 1890 at Kingsbridge in South Devon. His 
ancestors had been connected with that part of Devon for several hundred 
years, and he himself owned property near the village of South Milton until 
a few years ago. He was educated at Sherborne School and Christ Church, 
Oxford. He was ordained in 1915 and appointed to a curacy at Portsea 
(Portsmouth), one of several large parishes in poor districts in England 
considered to be beneficially tough training grounds for the Clergy. For 4 
time in the 1914-1918 War he served in a non-combatant capacity with the 
British Army in France and Belgium. After a temporary break-down in 
health he left Portsea in 1921. Subsequently, in 1924, he became Vicar of 
Dunstall, near Burton-on-Trent, doubtless glad to resume life in rural sut- 
roundings; and it was in Staffordshire that he married. While at Dunstall 


58 


he joined the British Pteridological Society, and developed a keen interest 
in ferns. The fruit of some painstaking work on scales under the microscope 
was printed in Volume VI of the Gazette, and he was from that time in 
constant touch with the Editor, Dr. F. W. Stansfield at Reading. He left 
Dunstall in 1929 and took charge for three years of the parish of Longstone 
on the eastern fringe of the Derbyshire Dales. In 1932 he became Rector of 
Cubley — some ten miles west of Derby, and remained there until 1936. 
During this period he was made President of the North Staffordshire Field 
Club, with which he had been associated for some years. His final move was 
to South Stoke, near the southern end of the Chiltern Hills. Among other 
advantages was the proximity of Oxford. He frequently visited the City and 
seldom missed an opportunity of going into the Botanic Garden. Reading 
was equally accessible, and he was able to pay occasional visits to Dr. Stans- 
field, thus extending his knowledge of fern varieties and learning a good deal 
about the back history of the British Pteridological Society. This experience 
stood him in good stead on his appointment as Secretary in 1950 at a time 
when the Society had only partially succeeded in reviving itself after the 
suspension of its functions during the War. 

Mr. Elliot might almost be said to have a passion for making friends, 
more particularly among people interested in botany and gardening. He 
maintained good relations with our members at home and abroad and was 
Pscangaial — to receive personal visits from members of the American 
Fern Soc 

In ane, of his work for the Society over a long period of years 
carried on until illness enforced his resignation, he was awarded the Stansfield 
Memorial Medal at the Annual General Meeting on the 26th September, 
1959 — the fourth award since the medal was instituted in 1938. This tribute 
gave him great satisfaction. 

Mr. Elliot was buried at South Milton, where mural tablets to the 
memory of several of his forebears can be seen in the village church. 

P. GREENFIELD 


J. DARGUE DIXON 
It was with great regret that members who had met Mr. Dixon in the North 
of England heard that he had died, at the age of 83, on the 22nd of February, 
1960. 


. Dixon was elected a member of the Society at the Annual General 
Meeting held at Southport on the 27th August, 1931. At the time he was 
living at Rochell Park, Dumfries. It was not until for business reasons he 
moved to Hest Bank on Morecambe Bay that he became well known to 
several of our members as a skilful and enthusiastic cultivator of ferns. He 
was elected to the Committee of the Society in 1936. 

A fernery Mr. Dixon built inside a large greenhouse with sandstone 
brought from Scotland was a source of admiration to those who saw it 
both for the pleasant effect of the rock, and the splendid development of the 


59 


ferns. Some of us invited to his house at Hest Bank and hospitably received 
there, saw ferns growing on a terraced bank, the species in this case being 
those for which shade was not essential. In later years, after the loss of his 
wife, Mr. Dixon was fortunately able to live with a married sic at 
Morecambe where he was not far away from our veteran member 
Whiteside. The writer has pleasant recollections of joining these two ccinesdl 
on more than one occasion for a talk on old times. After the Meeting at 
Ambleside in 1959, Mr. T. Dyer on behalf of members present conveyed their 
greetings to Mr. Dixon and Mr. Whiteside, an attention which they much 
appreciated. 
P. GREENFIELD 


THE SECRETARY’S AND TREASURER’S REPORT 


We can look back with some satisfaction on a year of progress, marred only 
by one sad event — the death of the Rev. E. A. Elliot; tribute to him is 
paid elsewhere in this issue. We have widened our scope to deal with world 
ferns and their allies, but the British ferns still continue to be our major 
interest, catered for not only by our Annual Excursions but by day meetings 
and weekend meetings held in different parts of the country. The large 
majority of our British members should now be able to attend at least one 
meeting annually without having to journey long distances, and we hope that 
these activities will develop accordingly 

The last issue of the British Fern Gazette has been accepted as a great 
success, and there has been a healthy increase in our membership. We hope 
this will continue, for the stronger we are the better our services will be. 

At the A.G.M. on 17th September we had to accept with regret the 
resignation of Mr. T. H. Bolton from the Presidency of the Society. He has 
held the office for two years and we thank him for his services to us during 
this period. In his place, Professor R. E. Holttum was elected President and 
we welcome him to this office; Mr. Bolton becomes a Vice-President. Dr. 
Davidson was elected a Vice-President, and Mr. R. F. Cartwright was 
elected to the Committee. A complete list of the Officers and Committee 
appears on the front inside cover. 

We are re-writing our Constitution, but were unable to get it completed 
- before the A.G.M. Copies will be sent to members well before the next 
meeting when it will be submitted for approval. 

The state of our finances formed an important part of the discussions. 
The appended financial statement shows a great increase in expenditure, 
which is inevitable if we want to develop the usefulness of the Society. But 
our income must also increase, and there are two courses open to us — 
increase our membership or raise the subscription rate. We do not feel the 
time is propitious for the latter course, although in the next year or so when 
our annual programme is more fully developed, we shall have to give it 


60 


serious consideration. In the meantime everyone can help to increase our 
membership by bringing the Society to the notice of friends interested in 
ferns and recruiting them to our ranks. 

It was pointed out that members could always consider leaving a small 
legacy to the Society in their wills. A painless method and one worth thinking 
about! 


Finally, prompt payment of subscriptions helps us greatly, and those 
who have not already done so, are asked to give this their attention now. 


FINANCIAL STATEMENT at 30th JUNE 1960 
as presented and accepted at the A.G.M. on 17th September 1960 
eo so 


59 £4. d. 
30th June 
To Balanc 87.3.3 tte Vol. 9, No. 1 8215 0 
Subscriptions 57 10 0 | Subscriptions to: 
le of: o> 
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 
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BRITISH FERNS & THEIR VARIETIES 
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Dwarf Trees, Flowering Shrubs 
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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 <! 
ca; 
2 


WK) bh 


THE SECRETARY’S AND TREASURER’S REPORT 


Progress has been maintained during the past year. The second issue‘of the 
new Gazette has upheld the standard set by Part 1 and has attracted wide 
attention both at home and abroad. It is satisfactory to know that our 
overseas subscribers are increasing in number and that we have something 
of interest to offer them. At home our meetings have increased — our 
widely scattered membership does not allow many to attend each meeting, 
but at some we have had a dozen and more members and this can be con- 
sidered very satisfactory. Meetings in the North however have not been so 
well supported. We would like to see these a strong feature in our annual 

rogramme. The Society was born in the North, we have a very large 
membership there and it is fitting that we should be active in this area where 
so much field work is awaiting our attention. 

The Annual General Meeting was held on 27th September, during the 
Annual Excursion, at Prenteg near Portmadoc in North Wales. Professor 
R. E. Holttum was again elected President and all the other Officers were 
re-elected. The resignation of Mr. H. Wainwright from the Committee left 
eight members who were re-elected and two new members, Messrs. R. M- 
Payne and R. S. W. Pollard were elected to bring the number up to full 
strength. 

The new Constitution of the Society was approved and appears in this 
ssue. 

Our satisfactory progress unfortunately does not extend to our financial 
affairs — in fact progress has been at the expense of finance. The cost of the 
Gazette was again heavy but was justified to ensure a good production which 


92 


would consolidate the favourable impression created by the previous issue. 
This is long term policy which we are confident will eventually pay dividends, 
but in the meantime we are faced with the fact that an annual subscription of 
10s. which was considered necessary as far back as 1920 is inadequate for 
our needs today, and we have had to increase the amount to 17s. 6d. The 
appended Financial Statement shows the position. 


e hope that all our members will take a realistic view of the i increase 
leads me to my usual final remarks — many subscriptions still remaining 


unpaid do not help our bank balance, and I would urgently request that these 
be forwarded to me at your earliest convenience — or earlier! 


FINANCIAL STATEMENT AT 30TH JUNE, 1961 


1960 
30th £0 4.2 6 2. To Balance... ... £10 5 5 
To Balance . aea 52 eee Gea eo ae 
Subscriptions 48 10 0 | Cost of Census List... 20:7: 31 
Donations 7 0 O | Subscriptions to: 
Sale of Census Lists 15 0 R.HLS. . pan laa Heme 2 
Advertisements 110 0 Council for Nature ... 1 0 0 
Sundry Debt 3; 3.8 
Advert 110 0 Postages and _ Incidental 
Sale of Gazettes t to Expenses: 
Leyden University 19 18 0 + pagl Sak nA i s 4 
—_—_ Editor ... ey ue 
4 a Se or 8 19 10 
Sundry Credito 
Due to ar ‘Caicos ie 
Gazette, V _ 9 No. ei 4 : 
Index to Vol. 8 anf 8 11 6 
Balance : 10°53 
*431°°6"° 3 £131 6 8 
1961 
June 
30th 


(signed) J. W. Dyce, Hon. Treasurer 
(signed) PETER TEMPLE, Hon. Auditor 


CONSTITUTION OF THE BRITISH PTERIDOLOGICAL SOCIETY 
as adopted at the A.G.M., 27th September, 1961 
1 NAME The Society shall be called “‘ The British Pteridological Society.” 


2 OBJECTS The objects of the Society are to promote and encourage the 
study and conservation of Ferns and other Pteridophytes and to 
do all such acts as shall further the active and corporate life of the Society 
and may lawfully be done by a body established only for purposes reco recognized 
by the law as charitable. 
93 


3 MANAGEMENT The Officers of the Society shall consist of a President, up 
to six Vice-Presidents, a Secretary, Treasurer, and Editor 
of Publications. 

The Management of the Society shall be in the hands of a Committee 
consisting of not more than ten elected members with the addition of the 
Officers. Three members shall constitute a quorum. The Committee shall 
have the power to co-opt additional members for special purposes. 

The Officers and members of the Committee shall be elected at an 
Annual General Meeting, and shall remain in office until the next Annual 
General Meeting when they shall retire but be eligible for re-election. In the 
event of a vacancy occurring during the year, such vacancy may be filled by 
the Committee. 

A resolution in writing signed and agreed to by all the members of the 
Committee for the time being in the United Kingdom shall be as valid and 
effectual as if it had been passed at a meeting of the Committee duly called 
and held, and may consist of several documents in the like form each signed 
by one or more members of the Committee. 


4 MEETINGS The Annual General Meeting of the Society shall be held at 
such time and place as the Committee may decide for the 
purpose of passing accounts, electing members, officers and the Committee 
and transacting the general business of the Society. General and Field 
Meetings shall be held at the discretion of the Committee 
An Extra-ordinary General Meeting shall be caited by the Secretary 
within thirty days of receiving a request in writing of the Committee or of 
not less than twenty members stating the purpose for which such meeting is 
required. 
At least 21 days notice of any general meeting shall be given to all 
members. 


5 ORDINARY MEMBERSHIP AND SUBSCRIPTIONS A candidate for membership 

must be proposed and seconded 
by members of the Society in writing and may be elected at a Committee 
Meeting or at a General Meeting. 

The annual subscription payable by members shall be payable in 
advance, and due on the Ist of July in each year, or on the election of the 
me: In the event of the election of a member after 31st March his 
subscription shall cover the succeeding year from Ist July. 

The rates of subscription shall be fixed by a General Meeting from time 
to time. 

6 HONORARY MEMBERSHIP Committee may elect any person or persons 
Honorary Membership of the Society. 
Honorary members shall be aa to all the privileges of membership, but 
shall not be liable for payment of any subscription. 
94 


7 REMOVAL FROM MEMBERSHIP Any member failing to pay his subscription 

within six calendar months of its becoming 
due shall be liable to have his name removed from the List of Members of the 
Society and upon such removal shall cease to be a member but shall be liable 
for his subscription for the current year and for any previous years. 

Any member whose conduct in the opinion of the Committee is pre- 
judicial to the interests of the Society may be removed from membership by 
a two-thirds majority vote of those present at a meeting of the Committee 
on the Agenda paper of which the words ‘‘ Removal of a Member ”’ shall 
have appeared; provided no member may be so removed unless due notice 
has been sent to the member of the intention of the Committee to proceed 
under this Rule and of the nature of the charges made and an opportunity 
has been afforded of answering such charges to the satisfaction of the 
Committee. A member so removed shall forfeit any claim upon the Society. 
8 AWARDS The Committee may make awards in respect of plants of 

tstandi ientific or horticultural interest 


9 ALTERATIONS TO RULES This Constitution may be altered as follows: 

Any proposal for alteration or addition to this 
Constitution shall be in writing, and shall be signed by two members, and 
be deposited with the Secretary not less than thirty days before the Annual 
General Meeting. Such alteration or addition shall be included in the 
Agenda of that meeting and circulated to all members and shall not be 
carried unless the votes of two-thirds of the members present at the meeting 
are cast in its favour. 


10 DISSOLUTION The Society may be dissolved by a vote of at least three- 
quarters ery of those siigao- ne voting at an Extra 

ordinary General Meeti 

for dissolution is passed ‘the sities funds and satel of the Society shall 

be disposed of to an agreed charitable organisation of similar interest or in 

default to a similar charitable body decided by the Council for Nature. 


11 Notices The accidental omission to give notice of a meeting to, or the 
non-receipt of notice of a meeting by, any member shall not invalidate any 
Proceedings or resolutions at any meeting of the Society or any Committee 
thereof. 


OBITUARY 
JOHN COCHRAN 
It is with great regret that we have heard of the death on 29th November, 
1961, at the age of 94, of our old member John Cochran of Kilmarnock. 
He was a joiner by trade, but was devoted to his garden and his plants. He 
specialised in chrysanthemums and bulbs for which he won many awards at 
Flower Shows, but his great interest was in his ferns of which he had a large 


95 


collection. He joined our Society in 1928, and right up to the end of his long 
life continued to be one of our staunchest supporters. It was a great sorrow 
or him when at the age of 89, he felt he was unable to give his ferns proper 
praberrany and had to part with them. The collection was gifted to his native 
of Kilmarnock, and is still being well cared for. 
The evening of his life was peaceful and happy in the care of his son and 
daughter-in-law, to whom we extend our deepest sympathy. 


NOTICE TO MEMBERS 


The Society now subscribes to the AMERICAN FERN SOCIETY and 
receive their quarterly journal. This is available for circulation among 
m 


undertaken to look after this. For the time being the AMERICAN FERN 
JOU may be kept for up to two weeks only, please, and returned to 
Miss Tetley. Applications must Sneath be accompanied by a stamped 
(44d.) addressed envelope, 9in. 


ADDITIONS TO MEMBERSHIP LIST 


BANGOR, maiden oe ac of North Wales, ree Library, Bangor, Caerns. 

H. Vv. Y, Pucketty Farm, Faringdon, 

G. B. EVANS, "3. A., si = t., Memorial Buildings, University College of North 
Wales, Bangor, Caern 

BRIAN J. FORD, 44, Richmo 2s Road, Cardiff, Glam 

Dr. w. GATZI, " Dufourstr., 118, St. Gallen, Switzerlan d. 

Mrs. C. E. URSULA HOLTTUM, 80, Mortlake Road, Kew Gardens, Richmond, Surrey. 

Dr. FRANCES M. JARRETT, The Herbarium, Royal Botanic Gardens, Kew, Richmond, 


Surrey 
LEYDEN Rij iksherberium, Nonnensteeg 1, Leyden, Holland. 
LUCKNow, National Botanic Gardens, India. 
DENYS MORGAN, M.Sc., F.L.S., Dept. of Botany, University College, Roma, Via 


Miss H. D. M. NORMAN, 57, Oakington Avenue, ited Park, Middlesex. 

B. J. PERRY, B.Sc., 7, Church St., Exmout 

Miss ALISON M. RUTHERFORD, The Cottage, Camis Eskam Farm, Nr. Helensburgh, 
Dumbartons 

Dr. HOPE SHERMAN, Pleasant Valley, Connecticut, U.S.A. 

Miss ae SLEEP, B.Sc., Dept of Botany, University of Leeds, Leeds 2, Yorks. 

Mrs. S. B. SPENCER, York Gate, Adel, Leeds 16, Yorks. 

Mrs. M. R. STUART, Tigh-a-Chladaich, Moulin, Pitlochry, P erthshire. 

p. w. woop, 9, Slievemoyne Park, Newcastle, Co. Down, Northern Ireland. 


96 


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 


BRITISH FERN GAZETTE 


VOL. 9 PART 3 MAY, 1962 


Contents 


The Polypodium Vulgare Complex ... se M.G.SHIVAS 65 
(Mrs. Trevor Walker) 


A Revised List of Pteridophytes from St. Kitts G.R.PROCTOR 71 


Short Notes : 
“Lastrea Dilatata var. Alpina” ss H. V. CORLEY 80 


Hymenophylium Transplants in Kent and Sussex 
GARETH B. EVANS 84 


Cystopteris Alpina and C. Dickieana in the Rock Garden 
P, GREENFIELD 85 


Notes on the History of British Fern Species I : Confusion in the 
Filmy Ferns G. B. EVANS and A.C. JERMY = 81 
British Pteridological Society : 
Field Meetings for 1961 
The Secretary’s and Treasurer‘s Rea 
Constitution of the British Pteridological Society 
Obituary .. 
Notice to stactiiers 
Additions to Siecateiahtes Su 


RRESSS 


Courier Co. Ltd., Tunbridge Wells 


THE 
BRITISH FERN 
GAZE FE 


VOLUME NINE PART FOUR” 


1963 


EDITED BY <A.-C: SERMY 
ASSISTED BY J. A. CRABBE 


Mhssoun! Boranicee 
SEP 19 1966 
GARDEN LIBRARY 


iG THE BRITISH 
PTERIDOLOGICAL SOCIETY 


BRITISH PTERIDOLOGICAL SOCIETY 
Officers and Committee 1962-63 


PRESIDENT 
Professor R. E. Holttum M.A., SC.D., F.L.S. 


VICE-PRESIDENTS 
T. H. Bolton Esq. 
Dr J. Davidson M.B., CH.B., F.R.C.P. (Edin.) 
P. Greenfield Esq. 
Reginald Kaye Esq. 
Dr 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 
R. 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 A120 
of AND 


upon paying an 
SIXPENCE. Full details wal be seat npék anton 0 ie SECUTAR 


THE 


BRITISH FERN GAZETTE 


VOL. 9 1963 PART 4 


VARIATION IN POLYSTICHUM IN THE BRITISH ISLES 


J. W. DYCE 
Loughton, Essex 


About the middle of the nineteenth century ferns were becoming popular 
plants and it must have been about this time that interest in variation was 
fully aroused; up till then little reference to varieties appeared in the fern 
books beginning to pour out; soon after, we find E. J. Lowe, T. Moore and 
others listing varieties more fully and the many fine ones of Polystichum 
setiferum must have done a lot to awaken an interest which became almost 
hysterical enthusiasm. From 1850 a succession of enthusiasts hunted, 
recorded and grew varieties; E. F. Fox was one of the first, later collaborating 
with Colonel A. M. Jones in the breeding of many fine varieties, notably 
plumose divisilobes which mark the zenith of development in this form of 
P. setiferum; G. B. Wollaston followed and we find his name associated with 
numerous plants of quality, his proliferum being the forerunner of many 
fine acutilobes, multilobes and divisilobes; S Elworthy and the Rev. c. 
Padley t thers 
were found ni Moly and Wills whose names are coupled with divisilobum and 
pPulcherri 
The hak plants were grown and passed on in the magnificent 
Collections of a select cavalcade of collectors; the last of these was W. B. 
Cranfield who made seemingly adequate precautions in his will for their 
safety for posterity; at his death these plans did not work out as he intended 
and most of the cream of his collection was irretrievably lost. Fortunately, 
among our meagre legacy made up of the wreckage from Cranfield and of 
gems still existing in private and nursery collections, sufficient fine varieties 
remain to give pleasure to presentday growers. 
The classification and nomenclature of fern varieties has been much 
bedevilled. _Untoetanately many ig ae early enthusiasts Sermee to orn 
Serene . deficien ree and irregulare. 
so freely depicted in literature and nursery catalogues, were veritable 
monstrosities fitted only for the rubbish heap and these, together with many 


The BRITISH FERN GAZETTE Vol. 9 Part 3 (pp. 65-96) was published 5 July 1962. 


98 BRITISH FERN GAZETTE: VOLUME 9 PART 4 (1963) 


minor variations, undoubtedly did a great disservice to the cult by offending 

“many people and by creating a most elaborate confusion of names; today 
such varieties are found in the wild quite frequently but are hardly ever 
collected for cultivation. 

This endeavour to clear the confusion of varieties of our three species 
of Polystichum has not been dressed to conform to the International code of 
nomenclature for cultivated plants; it is felt that to have done so would have 
rendered many of the names unnecessarily unfamiliar in the British Isles, the 
epicentre of cultivation of fern varieties. 


1. Polystichum lonchitis, the Hollyfern, is a rare alpine found on 
mountains in northwest and central areas of Scotland, the north of England, 
Snowdonia and the west of Ireland. The pinnate fronds are narrow and rigid, 
up to 12 inches and sometimes more in length and the pinnae are close-set 
and very spinose. It is very difficult to cultivate away from its natural 
habitats. : 

Two types of variation are recorded, a crested and a dense imbricate; 
it is very unlikely that any now exist in cultivation. 


2. Polystichum aculeatum, the Hard Shieldfern, is widely but rather 
sparingly distributed, nowhere to be found in great abundance and perhaps 
commonest in Scotland and the north of England. It grows in woodlands, 
hedgerows and, particularly in the north, in rocky ravines. The bipinnate 
fronds are up to 3 feet in length, leathery in texture, with smooth shiny 
dark-green unstalked pinnules; unipinnate juvenile fronds tend to resemble 
the Hollyfern. It grows freely and happily in cultivation with little attention. 

P. aculeatum has given few varieties and, with one exception, none of 
these seems to have survived; the outstanding exception, pulcherrimum and 
its progeny, includes some of the finest ferns we possess. 


(a) Pulcherrimum 


P. aculeatum pulcherrimum Bevis (Fig. 7). This unique and remarkable 
wild find by J. Bevis in a Somerset hedgerow in 1876 was given to Dr Wills 
and is one of the most beautiful of British ferns; it is vigorous and graceful 
and is described by Druery as “‘ with long attenuate silken divisions delicately 
imbricate at frond tips and almost always barren ”; it is evergreen, hardy and 
strong-growing in the border, needs a minimum of attention, propagates 
freely from its offsets and altogether is one of the finest and easiest of ferns to 
grow. On rare occasions a very few spores have been found and the results 
from them have been startling: 

P. aculeatum pulcherrimum Druery (Fig. 6) (progeny of ‘ Bevis’). An 
improved form of the parent and equally hardy, strong and easy growing. 

P. aculeatum gracillimum (Fig. 5) (progeny of ‘ Bevis’). A most re- 
markable series of plants, none quite alike and a complete departure from all 


DYCE: POLYSTICHUM VARIATION 99 


ities 


Figs. 1 - 7. Pinnae from the middle sector of the frond of some — nf gto 

Varieties. 1 P. se tiferum aeetobany 2 P. s. divisilobum longipinnatum Carbonell 

3 P.s. Pulcherrimum *‘ Mo ly’s ’.4P. s. cristatum, 5 P. acu — gracillimum, 
6P.a ¥ pulttervtilaat Ditieey, Vie pulcherrimum Bevis. 


100 BRITISH FERN GAZETTE: VOLUME 9 PART 4 (1963 


the usual types of variation. Again, Druery’s description cannot be bettered 
—‘ several are of wonderful delicacy of cutting, the parental half-inch 
pinnules being extended sometimes to nearly 3 inches and of almost hairlike 
tenuity with splayed tassel-like terminals, imparting by their weight a 
peculiarly beautiful pendulous character to the fronds ”; such plants usually 
need the protection of a glasshouse but some of them, not 80 finely developed, 
retain the parental hardiness and can be grown 

Two plants of this variety exhibited by Druery received First-class 
Certificates from re Royal Horticultural Society. 

Be leat. G (progeny of ‘ Bevis ’). This plant appeared 
in C. B. Green’s batch of seedlings. ‘Iti isa distinct and plumose plant of great 
beauty with finely-cut pinnules, and fronds about 18 inches long. The 
parental robustness has been inherited in full, making it an easy i. 
plant. It obtained an Award of Merit from the Royal Horticultural Soci 


(b) Plumosum 


Lowe records plumose and foliose varieties, but it is doubtful whether 
they still exist. 


(c) Densum 


Dense and imbricate varieties were recorded by Druery and Lowe but do 
not appear to be still in existence 
(d) Acutilobum 

A few finds with acutely lobed pinnules, and proliferous like the setiferum 


acutilobes, have been recorded; one at least is still grown and can 
purchased. 
(e) Cristatum 
This section includes all forms with frond apices and/or pinnae divided. 
Cresting is the commonest variation in ferns and most of the few 
recorded varieties of P. aculeatum come under this heading; it is doubtful 
whether many still exist. P. aculeatum grandiceps Jones is still listed; it is 
very distinct, with large narrow erect fronds and branched terminal crests. 


Polystichum setiferum, the Soft Shieldfern, is common in the south 
and oe of England, South Wales and Ireland. It needs more warmth, 
shelter and drier conditions and these it finds in Devon, Dorset and Somerset, 
its chief centre in England where it grows abundantly and almost exclusively 
in sheltered lanes and hedgerows, being often the dominant plant. It becomes 
infrequent towards the east and north and in Scotland, apart from a few 
places in the west, is practically non-existent. The normal form of this fern 
is not so strong-growing as aculeatum and the fronds are rarely over 2 feet 
in length, 18 inches being a good average; an exception is a very common 
variation with tripinnate and sub-tripinnate fronds which is always very 


DYCE: POLYSTICHUM VARIATION 101 


conspicuous by its upright vigorous growth, up to 4 feet and more. The 
habit of the plant is spreading with more or less arched and drooping fronds 
which are soft and paler green, in marked contrast to the dark hard glossiness 
of aculeatum. The pinnules are spinose and somewhat crescent-shaped, 
broad at the base with a distinct slender stalk. Except in very sheltered 
conditions the fronds are not so persistent and die down much more quickly 
than those of the other two species. The normal form of setiferum makes an 
acceptable garden plant and is found in gardens throughout the country. 
Except in sun-baked positions, it is an easy plant to grow and its graceful 
soft green fronds will adorn shady corners and borders with a minimum of 
attention. An annual mulch of rotted leaves, a dusting of bonemeal very 
occasionally and, particularly in the <a" ~~ water during hot and dry 
spells will keep this fern happy and in dition for many years. 

The Soft Shieldfern is ee oh ret in any colony of the fern 
there is a wide range of minor variation in shape and habit; more extreme 
variation produces a vast number of forms. Lowe in British ferns and where 
found (1890) published a table of divisions, groups and sections covering 366 
named varieties; the British Pteridological Society’s selected list in Druery, 
The book of British ferns (1902) reduced the number to 173; today, after 
minor variations and undesirable monstrous forms are removed from the 
Survivors, we are left with a greatly reduced list which, by still using Lowe’s 
classification, appears as follows: 


VARIATION IN POLYSTICHUM SETIFERUM 
DIVISION A CHANGE IN FROND SHAPE 
Group 1 Cristatum 
(a) capitatum (crested apex) 
(b) cristatum (crested apex and/or pinnae) 
(c) percristatum (crested apex, pinnae and pinnules) 


Group 2 All other changes in frond shape 
(a) brachiatum (rhachis branching into three near the base) 


) 
(d) cruciatum (pinnae or pinnules twin-branched) 
(€) congestum (pinnae dense and/or overlapping) 


DIVISION B CHANGE IN PINNULE SHAPE 


Group 1 Plumosum 
(a) topes (increased development, thin texture, fructification scanty 
or absent) 


(b) foliosum rea texture and fructification normal) 
(c) pulcherrimum (lower pinnules, and sometimes upper, sickle-shaped, 
deeply incised and brite ped into thread-like Se 
Character more 


Fructficati tion generally a 
intermittent) 


102 BRITISH FERN GAZETTE: VOLUME 9 PART 4 (1963) 


Group 2 Divisum 
(a) acutilobum ni pinnules, all penny acutely lobed, upper and 
lower pinnules of approximately eq gth) 
(b) dvslobum m: (pinnate Pinos, all divisions acutely lobed, low 
onger than upper ones aivisions hoe 
highly iy doveliiied, approaching sub-quadripi innate or even 
a 


q 
(c) decompositum (subdivision on normal lines) 


Group 3 All other pinnule changes 
(a)  lineare (pinnules narrow, acutely lobed and undivided). 
(b) rotundatum (pinnules broad or roun 
(c) crispatum (pinnules crisped) 
(d) setosum (pinnules spiny) 


DIVISION A CHANGE IN FROND SHAPE 


Group | Cristatum 


As already stated, cresting is the commonest distinctive type of variation 
in ferns, and P. setiferum has contributed in full measure a multiplicity of 


named which can all be gathered under this one heading; these are: 
peigaoe (with branching te capitatum (having a head), coronatum 
(crowned), corymbiferum (bearing a cluster or corymb), cristulatum (having 


small aa grandiceps nia a a arge head), multifidum (very much split), 
percristatum (crested all through), polydactylum (many-fingered), ramosum 
(branched), ramo-cristatum (crested and branched), ramulosissimum (having 
many small heads); this does not exhaust the list but they all mean more OT 
less the same thing and can be sorted into three sections under the group 
heading. 

Cresting, considered generally, does not enhance the beauty of P. 
setiferum and cannot be regarded as very desirable except for the collector 
interested in diversity of form. Capitatum in all its shapes is clumsy and top 
heavy and the names given to its variations (acrocladon, coronatum, 
corymbiferum, grandiceps, multifidum, etc.) are hardly less so. Cristatum 
(Fig. 4) can be more acceptable if the cresting is small and neat, but even then 
one has the feeling that the pinnae are overloaded and lacking in the grace of 
the normal form. Percristatum is a stage further with minute cresting at the 
tips of the pinnules and this does add something which helps to soften the 
general effect. On the whole, this group is more interesting than beautiful 
and most of the many named varieties seem to have sunk into oblivion. 
There has been very little mention of them in fern literature for many years. 
A few are still listed in fern catalogues and some possibly linger on un- 
recognised in fern borders known only as crested polystichums. 

In the wild these crested forms still turn up occasionally and they seem 
to be one of the few surviving kinds of wild variation in P. setiferum. 
Intensive hunting by members of the Society in the West Country since the 
war has produced very few noteworthy finds, but these include three examples 


DYCE: POLYSTICHUM VARIATION 103 


of cristatum and one percristatum, all found by myself, two of them during 
the autumn of 1962. 

Lowe (1890) contains 71 ferns which can be included in this group; 
they are reduced to about 24 in Druery (1902); today’s catalogues contain 
only a few 

P. setiferum cristatum gracile Moly, a percristatum with dark green erect 

crested fronds, small crests on the pinnae, and the pinnules regularly 
fanned at the tips. My percristatum found last year is very similar, 

P. s. cuneatum cristatum, fronds erect with terminal crests and wide 

wedge-shaped pinnules 

P. s. ramo-cristatum, dwarf dark-green broad-pinnuled form, repeatedly 

anched 


P. s. Thompsoniae Lowe seedling. fronds terminating in compactly 
branched heads. 


Group 2 All other changes in frond shape 

The five types of variation included in this group cover all other changes 
likely to be found. In brachiatum the branching of the fronds does nothing 
to enhance their appearance and only destroys their grace; its chief value is 
Curiosity. Deltoideum and angustatum are more acceptable, particularly if 
accompanied by some pinnule change. Under angustatum can be placed the 
variety revolvens in which the pinnae are curved so as to give the fronds a 
narrow tubular shape; a good form can be very attractive. In cruciatum the 
Pinnae are doubled and grow out from the rhachis at wide angles from each 
Other, one upwards and one downwards; the overlapping with the other 
doubled pinnae gives a criss-cross effect up both sides of the rhachis and a 
good example of this variation is most effective. In congestum the axes are 
Shortened with the result that the normal sized pinnae and pinnules are very 
Close together and imbricate; provided it is regular and constant, this variation 
is well-worth growing but more often it is temporary, induced by adverse 
Conditions, and such plants removed to the comfortable conditions of 
garden life flourish and become normal. 

The sections in this group at one time contained a large number of 
named varieties, but most of them were worthless, being irregular, inconstant 
Or near-normal. It is not surprising that very few remain. The first 
Sections seem to have disappeared from cultivation. Deltoideum and 
angustatum can still be noted in the wild, but are regarded as within the 
normal range of variation and are left in peace. Revolvens is probably still 
grown in some gardens, but does not seem to be obtainable commercially. 
Cruciatum and congestum can both be purchased, the former in a few forms, 
the most noteworthy being: 

P. s. Wakeleyanum Moore, a narrow cruciate variety with crested apex. 

Found in South Devon in 1860. 


104 BRITISH FERN GAZETTE: VOLUME 9 PART 4 (1 963) 
DIVISION B CHANGE IN PINNULE SHAPE 


Group 1 Plumosum 
Division B contains the cream of the at se varieties and, probably 
because their great beauty has ensured a more widespread distribution, we 
find that a number of the old named forms still survive and form the bulk of 
Polystichum collections today. In Group 1 there have been many notable 
and remarkable plants but most of them have been lost. In the p/umosum 
section itself I can find no record that the following continue to exist: 


P. s. plumosum Elworthy, found in Somerset by Elworthy and stated to 
be very beautiful and distinct. 
P. s. plumosum Patey, found in Dorset, with thin papery finely-cut fronds. 
P. s. plumosum Wollaston, found in Devon, robust and fertile. 
P. s. plumosum grande Fox : 
. . : All raised, very robust and splendid 
P. s. p. perfectum Birkenhead 1 
plants. 
P. s. p. splendens Col. Jones 
s. plumosissimum Birkenhead, finely divided and, under extremely 
ee avourable conditions, Druery says “all the ultimate divisions 
extend into long tongue-like tails, piling up a moss-like growth 
upon the frond. A type of apospory; unique 


Some of these ferns were strong growers and it is possible they still survive 
unrecognised. However, we do still have with us: 


P. s. plumosum grande Moly. This tall strong-growing barren variety is 
in my collection and produces annually, without fail, large heavy 
tripinnate fronds over four feet long and wide in proportion; the 
close-set, overlapping pinnae are like small fronds, with pinnules 
over one inch long, each resembling a small pinna with central 
rhachis and stalked pinnulets. It is truly a magnificent plant but the 
fronds are so dense and massive that, unless given some support, 
they are quickly broken at the base and complete the season lying 
on the ground. It is an easy fern to grow and the fronds remain 
fresh and green well into the following spring. 


The next section, foliosum, is described as sub-plumose but in all other 
respects normal. This type of variation is met with frequently in the wild but 
is very seldom striking enough to induce the finder to collect it. Lowe lists 
many named forms but today only one seems to be in nursery catalogues, 
it is called foliosum and described as having broad erect fronds with very 
deep-cut pinnules. I think it is probable that the named examples of foliosum 
depended chiefly on other characters for their differences and the oliose 
character as such showed little if any variation. It would follow frei this 
that in the strict sense there is only one foliosum. 


DYCE: POLYSTICHUM VARIATION en eg 105 


The pulcherrimums were an ephemeral race and throughout the history 
of fern hunting it has been the ambition of all hunters to find one of these 
varieties in which F, W. Stansfield (Gazette 1:275) says “* the species rises to 
its highest flight of genius, at all events so far as wild varieties are concerned.” 
They were too good to last and those that did last found the strain too much; 
most of them reverted to normal setiferum permanently, while some would 
throw up a frond in character occasionally, but more often only a few pinnae, 
In all, only sixteen examples of pulcherrimum were found in the sixties and 
seventies of last century, nine of them by Moly! None have ever been found 


or even iat seasons produce fronds exhibiting the pulcherrimum characteris- 

tics. Th e so distinctive that they couldn’t be missed. The peculiar 
oR neni of the pinnules gives an extremely graceful and feathery 
appearance to the fronds which, in addition, are aposporous and the pinnule 
tips and abortive sori can under moist conditions develop prothalloid 
growths, producing young plants when layered. Only one pulcherrimum has 
survived : 


P. s. pulcherrimum ‘* Moly’s Green’ (Fig. 3). This is one of Moly’s wild 
finds which, unlike all the others, is a vigorous grower; it develops, 
however, a multiplicity of crowns which, if allowed to remain, 

_ reduce the size of the plant and effectively prevent it from showing 
its true character. In common with the others it is intermittent and 
only a well-grown plant kept to a single crown is likely to be 
persuaded to show the pulcherrimum character, something well 
worth striving for. Although I have grown this fern for 10 years, I 
have not yet been fortunate enough to coax it into exhibiting ‘its 
full glory, but that is probably because my fern borders are too 
exposed and get too dry in summer and I am always too busy to 
give the individual attention required by such plants. Given a good 
deep root-run in shaded conditions and all offsets removed as soon 
as seen, it should not be too difficult to succeed with ‘Moly’s Green’. 


Group 2 Divisum 

This group embraces most Polystichum varieties grown today; in the 
wild, the decompositum section is commonly found as tall vigorous plants 
with large tripinnate and subtripinnate fronds. Subdivision is on normal 
lines but, since the normal form of setiferum is a handsome and acceptable 
garden plant, so even more is the wild decompositum. It is only the ease with 
which they can be found and the vision of the finer forms of gentle blood 
beyond which blinds us to their beauty and leads us to dismiss them casually. 
Some outstanding ones have been found and named, as have been many 
Which were not particularly outstanding. None of them seem to have 


106 BRITISH FERN GAZETTE: VOLUME 9 PART 4 (1963) 


survived, but it must not be overlooked that decompositum is the link between 
the normal and the more elaborate and desirable forms; it should not be 
dismissed lightly. The best known was P. s. decompositum splendens Moly, 
said to be only occasionally fertile and bulbiferous. The Jones Nature Print 
shows this to be a very good tripinnate form but with nothing about it to 
indicate any latent possibilities, for Moly claimed that it was the parent 
used by Jones and Fox to produce their new race of plumosum divisilobum, 
the finest ever raised. However, Col. Jones found many decompositums 
himself and from one of these raised a plant also named decompositum 
splendens which without doubt was the real parent. One wonders if this fern 
still lingers on in some corner unrecognised. Another thought intrudes: 
how often during hunting in the post-war years have we stopped to look at a 
decompositum with similar potentialities and discounted it at its face value. 


Although the best known, Lawes s ween was by no means the best 
m ollas 


plants do not appear to have been capable of emulating the achievement of 
Jones’ splendens. 


this section can also be placed all varieties classed as multilobum 
and conspicuilobum. As the names indicate, the lobes of their divided pinnules 
are many and conspicuous, but these characters in themselves are not out- 
standing enough to make them particularly sought after and I cannot find 
that any 


We now move on to the sections acutilobum and divisilobum. The name 
acutilobum is often given to varieties with acute undivided pinnules but 
these belong more appropriately to Jineare in Group 3. This leaves us with 
two very similar sections which will be considered together as it is difficult 
to draw an exact line between them. The definitions are Wollaston’s, and 
by almost general consent were accepted by all hunters and growers. 
Sues character of larger lower pinnules in divisilobum is rather a 

eak point, as borderline plants in certain years or in different cultural 
iediie: may have longer or shorter lower pinnules. Padley considered 
there should be two classes — decompositum and acutilobum, the latter to 
include divisilobum. However, even though it is difficult to allocate some 
varieties with certainty, it still cannot be denied that there are two definite 
sections, with divisilobum showing much more highly developed pinnules, 
and Wollaston’s arrangement cannot be bettered. 

Acutilobes are still to be found fairly frequently as wild _ and post- 

war hunting has produced a few. Usually they have been found on dryish 
banks as quite dwarf-growing plants and have not been very distinctive, their 
quality being more suggestive than actual; in fact, /ineare described them 


DYCE: POLYSTICHUM VARIATION 107 


better than acutilobum. Several have been collected during hunts, chiefly 
because there was nothing better to collect and we wished to have something 
to take home with us. Growing these plants has convinced me, particularly 
after last year’s cool and wet summer, that we have been wrong to regard 
them lightly. My first acutilobum found in 1953, developed some years later 
into a plant of quality almost rivalling Dr Stansfield’s ‘ Howley Acutilobe ’, 
a first-rate find of 1914. A few years later, other finds, dwarf-growing in dry 
banks, were transplanted in my borders and have grown in size year after year 
until the past summer when they excelled themselves. One has developed 
into a fine proliferous acutilobum with finely divided but not very narrow 
pinnules, another has extremely narrow fine pinnule segments giving it an 
appearance of grace and lightness, while another has retained the /ineare 
character, but has become a really good specimen of its kind. Close watch 
will be kept on all these plants to see whether they retain their high standard 
in future years. 

One of the best known of the many wild finds of acutilobum is that 
found by Wollaston in Devon in 1852, acutilobum proliferum Wollaston. At 
one time it was thought that only the acute-lobed varieties were proliferous 
and all were classed accordingly as proliferums. It was later discovered that 
this tendency to produce bulbils was shared by many other forms of the 
species, including some of the normal ones, and the name lost its appropriate- 
ness when applied to the acutilobes. However we still associate the proliferous 
habit with acutilobum more than any other type of variation and the well- 
known named varieties possessed the habit; these included P. s. acutilobum 
Gray and six examples of P. s. a. proliferum, of Allchin, Choule, Dadds, 
Hartley, Wills and Wollaston. That so few really good finds were made 
points to the fact that most of them are very much alike and it may be that, 
with few exceptions, this quality in setiferum is a rather inconstant one. 
My own finds mentioned above may prove this to be so if they fail to hold 
the standard achieved last year. 

Named acutilobums surviving today are few. I am told that the Allchin 
and Hartley varieties can be obtained, but it seems that, so far as the name Is 
concerned anyhow, Wollaston’s has disappeared. Many plants very similar 
can be found flourishing in fern collections named only acutilobum ot 
acutilobum proliferum, so it would seem reasonable to assume that this 
vigorous variety is still with us. However, as the members of the section are 
So very much alike, even to the proliferous habit, unless they can be definitely 
identified, or have some distinctive quality to lift them above the others, it 1S 
better that they remain unnamed. The few acutilobums in my collection are 
without names (e.g., Fig. 1) all bulbiferous and similar in appearance and it is 
quite possible that they are survivors of the named coterie. I am quite 
content to leave them incognito. 

When we come to divisilobum we find that the species has been much 
more active. Lowe's list of variations is longer than any of his other Poly- 


108 BRITISH FERN GAZETTE: VOLUME 9 PART 4 (1963) 


stichum lists and all following lists are the same, including those in nursery 
lo And still today divisilobum seems to have survived better, 
possibly because of its greater beauty. What I said in my opening remarks 
to Division B applies most of all to this section. In divisilobum pinnule 
division has run riot, splitting up into a multitude of fine segments, pinnae 
and pinnules all overlapping to give the fronds the appearance of great 
plumes. Many, too, are proliferous and the tiny fronds bursting from the 
bulbils along the rhachis add to the effect. One of the finest is divisilobum 
Bland, easily obtainable and propagating true from its innumerable bulbils. 
This fern was found in Ireland and is by far the best wild find in the section. 
To find a wild divisilobum has been an ambition of hunters since the cult 
began and the finds were many, chiefly in the West Country. Today the most 
intensive hunting in that area fails to produce a single specimen! Outstanding 
forms were found by all the great hunters of the last century, notably 
Elworthy, Jones, Moly, Padley and Wills, but none of them were as dense as 
* Bland ’ and none seem to be living today, at least not as named specimens. 
The failure to ensure the safe disposal of the last great collection was their 
note inen Plenty of unnamed divisilobums, mostly seedling plants, are to 
easy to obtain. 


Prov the béiginintes ng both nurserymen and amateurs were busy raising 
new plants from spores and bulbils. The outstanding work in this field was 
by Jones and Fox, who evolved a new race of plumose divisilobes by breeding 
from a wild find of decompositum. These surpass ‘ Bland ’ in their develop- 
ment and the finest of them, Baldwinii, although no longer available in the 
original, has passed on to us much of its refinement in its progeny which still 
exist under the same name. Other named divisilobes obtainable are: 


P. s. plumoso-divisilobum, Esplan type, rivalling Baldwinii when well 


P. s. divisilobum densum, dwarf-growing, plumose. 

P. s. divisilobum Iveryanum, crested. 

P. s. divisilobum longipinnatum Carbonell (Fig. 2), slender and graceful 
with very long lower pinnules 

P. s. divisilobum decorum, very ‘a cut, 

P. s. divisilobum grandiceps, with a heavy crested head. 


Group 3 All other pinnule changes 

This group gathers together all the remaining worthwhile types of 
setiferum variation and, apart from lineare section which contains plants 
with simple clean-cut graceful and symmetrical lines, forms rather an anti- 
climax. The old-time /ineare lists contained many irregular rogues wit 
ragged, rudimentary and missing pinnules, most of them fit only for the 
rubbish heap. The section is exemplified by ‘ Hirondelle’, one of Moly’s 


DYCE: POLYSTICHUM VARIATION 109 


finds, so named from the striking resemblance of the pairs of opposite 
pinnules to swallows in flight. ‘ Hirondelle ’ was far from robust and is now 
no more. Many /ineare plants are in cultivation however, strong-growing 
and approaching ‘ Hirondelle’ in beauty. Some are named gracile but the 
majority are wrongly included under acutilobum. Good specimens can be 
found in the wild. 

Rotundatum is the opposite of lineare, with broad rounded pinnules. 
Good forms are distinctive and should be strong-growing, so are probably 
fairly common in gardens. I cannot find that they have survived as named 
specimens. Crispatum has been noted at times in the wild in recent years 
but no extreme forms worthy of collection and naming have been seen and 
I can find no trace of any in soe agit Setosum can be regarded as a fuller 
development of the spininess of the normal species; named forms have not 
survived ; the character is quite a hehcsies one in wild colonies and, rarely, one 
is seen which is admired but is not extreme enough to be collected. In 
section can be included serratum and perserratum, the deep-cut saw-like 
edges of the pinnules giving a similar spiny look to the fronds; again, there 
were a few named forms but none are grown by name today. 

It is hoped that this brief survey will help to give a clearer picture of 
variation in Polystichum and remove some of the nomenclature confusion. 
It may enable some members to identify unnamed plants in their possession, 
plants which excite curiosity and are cherished for their rare beauty or 
unusual form. Perhaps some of the treasures which we mourn as lost will be 
brought to the light of day. If so, my labours will be well rewarded. 


REQUEST 


POLYSTICHUM VARIETIES 
As may be appreciated from the foregoing account, the tracking down of fern 
varieties is an arduous task. 

Anyone who can provide evidence of living specimens of decried 
varieties is asked to communicate with J. W. Dyce, Hilltop, 46 Sedley 
Loughton, Essex. - 

Dried fronds with data are particularly desirable; for some instructions 
on drying and labelling see Gazette 9: 27. 


110 BRITISH FERN GAZETTE: VOLUME 9 PART 4 (1963) 


MEIOSIS IN ASPLENIUM X MURBECKII FROM BORROWDALE 


J. D. LOVIS, 
Department of Botany, University of Leeds 
INTRODUCTION 
The hybrid between Asplenium ruta-muraria L. and Asplenium 
ee (L.) Hoffm., A. x murbeckii Dérfler, is certainly of very rare 
ce in Britain dnd there are indeed very few authenticated records. 
Undoubte d specimens of A. x murbeckii originating from Stenton Rock, 
near Dunkeld, Perthshire (1853 and 1858) and from Arthur’s Seat, Edinburgh 
(1861) are preserved in the Moore Herbarium at Kew but as far as I am 
aware, prior to Mr Fred Jackson’s discovery of a specimen on Castle Crag 
in Borrowdale, this hybrid had not been recorded in Britain during this 
century. ere is an earlier but until now unrecognised specimen of this 
hybrid from Borrowdale in the British Museum Herbarium, collected in 
1927 by H. W. Pugsley, but mistaken for A. x alternifolium Wulfen. Diminu- 
tive forms of A. x murbeckii (and all the known British specimens merit this 
description) have much the same aspect as small forms of A. X oe 
and these two hybrids can in fact readily be confused. However, A. 
murbeckii may be simply distinguished from A. x alternifolium by Pe 
minutely serrate tips of its pinnae, the several individual fine teeth being 
narrowly acute or even lanceolate. In A. x alternifolium the tips of the 
pinnae are more rounded, and where teeth are present these are obtuse or 
subacute. 

It is quite evident that Mr Jackson’s discovery (Jackson 1961) is 2 
significant event in the study of British ferns and he should indeed be con- 
gratulated. Very generously, Mr Jackson sent a portion of his plant to the 
Chelsea Physic Garden and subsequently another portion to the Botany 
Department Experimental Garden at Leeds. This plant has not yet been 
fertile since its arrival in Leeds, neither has the Chelsea plant produced any 
fertile fronds since 1961. It would seem that this hybrid does not thrive in 
an urban atmosphere, even within the protection of a greenhouse. The 
Leeds plant was certainly visibly damaged by the appalling week of con- 


EXPLANATION OF PLATE I 


Figs. Permanent mg iphone a Lo koe ae of diakinesis in Asplenium 
iarbeckdl from Castle Crag, Borrowdale. Magnification x 1000, photographed with 
z2mm N.A. 14a sorhanieie with x 10. rr ” Fig. 1, 60 bivalents and 24 


univalents. Fig. 2, 62 bivaleds and 20 univalents. Fueplarmsory. diagrams on text- 
figures 2 and 3. Figs. 3, 4, and 5. Glycerine jelly mounts of sporangia of A. X 


roportion of abortive spores are formed, 
but this proportion varies widely from Daieatainnn to sporangium. Fig. 3. Rupturee 
sporangium showing mis-shapen spores. Fig. 4. Spo nee ovo showing group 0 

apparently good spores. Fig. 5. Entirely abortive sporangium 


2 
> 
> 


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_<Smm, linearae-lanceolatae, fuscibus-fulvibus virgibus 
centralibus. Frondes 10-18(-25) cm longae, in forme fas stigate sen nsim erga apice, 
apices acutae, aris pinnis; stipes fuscae-fulvae: pinnae 1-2cm longae, 


ob inali i 

mediis 41-49). Sori (4-) 5-8(-10) ad pinnis mediis. Sporae in medio chlorale 
27-50p longae (ordo mediis 34-4312. Chromosomata gametophyti 72, sporophyti 
144. 


Holotype: GERMANY, Bavaria: Kienberg by Ruhpolding D. E. Meyer. 
Living in culture (Hort. Berol.), as sporophytes Nr. 42, 43, 47 and 82. 


1° The following quotation is taken from Heywood, V. H., 1960. Feddes Repertorium 63:183. se eee 
an example is — in oo eomage a trichomanes where the only certain character (and by no means an 
unambiguous a to sep — he bon pom the tetraploid is the form a: - ‘details of the agers 


A. pre Boll has vane pu lished oan pacific Rong s oheeae o is 

ua cmon of A. trichomanes subsp. bivalens D. E, Meyer psi Nr. 133), gave the following 
dee ms: in chloral medium, 23-5-35-5u long (mean 30-5u), 18-0-27-5u cog a ion 3928 
in air, 22: 70-33" 5u long (mean 27-9), 18-0-27-5u broad (mean 22-14). 100 spores 
medium. 


LOVIS: ASPLENIUM TRICHOMANES 153 


of upper pinnae square or som oblique, termi i inconspocuo 
narrow. Stomata 35-57, long (range of means 41-49). Sori (4-) 5-8 (-10) on 
middle pinn pO. n chi medium 27-50» long (range of means 


THE NEW DIPLOID TAXON WITHIN THE ASPLENIUM TRICHOMANES COMPLEX 
It has already been stated above that the diploid subspecies recognised 
by Meyer, subsp. bivalens, found only on non-calcareous rock, and particu- 
larly frequent in central and northern Europe, should be called subsp. 
trichomanes. Another diploid taxon is now known, which is apparently 
confined to limestone rock in central and south-eastern Europe. This plant 
is described below as subsp. inexpectans. In this section the discovery of this 
plant is briefly described, and the two diploid subspecies are compared. 

It was with very great surprise that I discovered that a single plant of 
Asplenium trichomanes collected by J. A. Armitage of the Leeds City Museum 
in 1961 from limestone near Ston on the Peljesac peninsula on the southern 
Adriatic coast of Jugoslavia was diploid. My surprise was occasioned both 
by the morphology and the ecological origin of this plant. The origin from 
limestone was most unexpected, since although diploid A. trichomanes was 
known from a great variety of rock, including serpentine and mica-schist as 
well as a iderabl ber of acidic types, no diploid plants had previously 
been found on calcareous rock. Moreover, this rule, the absence of diploid 
A. trichomanes from limestone, held good not only in Europe, but also in 

orth America}, and Australia’. 

Subsequently, another single plant of unusual morphology collected by 
Miss Anne Sleep of this Department in 1962 from limestone in the Tajgetos 
mountains of Greece also proved, against prediction, to be diploid. 


ore recently, some plants of Asplenium trichomanes collected by 
ia have proved to 


Professor Melzer and Professor T. Reichstein, to study this plant in situ in 
several localities in Austria. 

__ The familiar diploid form of Asplenium trichomanes, 
is notable not merely for its avoidance of calcareous rock, 


subsp. trichomanes, 
but also for its 


8 Type material of A. trichomanes subsp. quadrivalens D. E. Meyer (Meyer Nr. 47), gave the following 
i ions: in chloral medium, 33-5045-50 long (mean 39:3), 25-5-37°5u meen < eS 
air 29-5-43-5Sy long (mean 38:2), 25-5-33-5u broad (mean 29-6). 100 spo meas figures refer 
medium. In common with all the other spore measurements rage in this paper, th 
to the dimensions of the exospore, the perispore 
* BRITTON, D. M., 1953, Amer.J Bot. 40:577. 


BRITISH 


FERN GAZETTE: 


TEXT-FIGURE 2 


VOLUME 9 PART 


5 


(1964) 


LOVIS: ASPLENIUM TRICHOMANES 155 


remarkably constant morphology. As is demonstrated in Text Fic. 2: 5 and 6, 
like all other forms of A. trichomanes, subsp. trichomanes is very susceptible to 
modification in size and development by environmental conditions, but this 
nvironmental influence is predictable, and this subspecies remains a 
relatively uniform taxon with a distinctive facies recognisable wherever it is 
found 


The number of known samples of subsp. inexpectans is still very small, 
but it is nevertheless evident that in direct contrast to subsp. trichomanes, 
inexpectans as presently conceived is a very variable taxon. Whether or not 
the Peljesac plant properly belongs in the same taxon as the Austrian collec- 
tions is a problem which only more collections and much more study can 
resolve. Nevertheless, although variable, subsp. inexpectans in all its known 
forms is clearly distinct from subsp. trichomanes and merits at least sub- 
specific separation. It will obviously be of great interest to discover what 
degree of genetic relationship exists between these two taxa, but this can only 
be determined in due course by experiment. 

The most obvious morphological differences between subsp. trichomanes 
and subsp. inexpectans lie firstly in the shape of the pinnae, which in inex- 
pectans are comparatively square or oblong, not suborbicular, without the 
distinctive auricle seen in well-grown trichomanes, and secondly, in the 
insertion of the pinnae, which is not oblique in inexpectans, but more or less 
square, even near the tip of the frond. In subsp. inexpectans the terminal 
Pinna is usually conspicuous, and the frond ends abruptly, lacking the 
gradually tapering tip seen in subsp. trichomanes. 

It is certainly much easier to distinguish subsp. inexpectans from subsp. 
trichomanes than it is to distinguish inexpectans from some of the forms of 
subsp. quadrivalens found within its geographical range. More than 
cannot usefully be stated at the present time, but it is already clear to me that 
the Asplenium trichomanes complex is very much more complex and variable 
on the forested limestones of central Europe than it is in Great Britain, and 
a great deal more study is required. 


Asplenium trichomanes L. subsp. inexpectans Lovis subsp. nov. 


Pal 
Frondes 7:5-10(-12) cm longae, terminales abruptae, apices 
Pinnis; stipes fuscae rubrae-fulvae; pinnae 0-4-0-8(-1-0) cm longae, qu 8, : 
oblongae, aliquando angustatae oblongae, plerumque lateratae-parallelae, 
texturis delicatis, sub-distantae, insertio pinnis superis quadratus vel parum obliquus, 


TEXT-FIGURE 2 Photo-copy silhouettes (reproduced natural size) of fi “ ; 
Fic. | from plant in cultivation at Leeds, ai others pressed from wild. Fic. 1 su Se aie, 
pe } i ” Switzerland 


n, Pel ninsula, Jugoslavia. Fics. 2 and 3 subsp. inexpectans, 
iederésterreich pore oy Fie subsp. trichomanes, Val d@’Antabbia, Canton Ticino, Swi 
(selected as comparable in size to Fic. 3.) Fics. Sand 6 subsp. tric , Bosco-Gurin, 
Ticino, Switzerland, demonstrating effect of environment. $ from very exposed on serpen 
tine boulder. Fic. 6 from ly shaded base of other side of same boulder. Fic. bag quadri aod 
Draycott Rocks, Mendip’ til. Somerset, England, from very deep 5 ee g insertion 
tion i a thai of subsp. trichomanes. Fi. 8 subsp. quadrivalens 
on, Engiand, crevice in mortared wall. 


156 BRITISH FERN GAZETTE: VOLUME 9 PART 5 (1964) 


pinna terminalis ee pea lata. Stomata 31-48» longae (ordo mediis 

40-42). Sori rh 10) a mediis, aliquando condensa in media pinna. 

Se kot a 5h eas ey rondila 33-37). Ciinaadomand gametophyti 36, 
yti 


Holotype: AUSTRIA, Niederésterreich: at the Langenbrucke, Gutenstein, on 
shaded limestone rocks, H. Melzer, April. 1963, cult. in Basel & Leeds, 
JDL1963:150T in Herb. B.M. 

Paratypes (all cytologically determined) AUSTRIA,—Steiermark : Baren- 
schiitzklamm, by Mixnitz, Melzer, 3 Oct. 1962, cult. in Basel, TR. 738, 
JUGOSLAVIA, Hrvatska: ne Ston, Peljesac peninsula, J. Armitage, 17 May 
1961, cult. Leeds Univ. s.n.. GREECE, Peloponnisos: Taiygetos moun- 
tains, in limestone gully, A. Sleep AS62/29, 2 Aug. 1962, cult. Leeds Univ. s.n, 


Rhizome scales <3-5mm, lanceolate, with red-brown central stripe. 
Fronds 7: 5-10 (—12) cm. long, abruptly terminated, apex obtuse, 10-24 pairs of 
pinnae; petiole dark red-brown; pinnae 0-4-0-8 (-1-:0) cm long, square or 


oblong, some etimes narrowly oblong, usually parallel-sided, texture delicate, 

sub-distant, square or slightly oblique, terminal 
pinna usually conspicuous, broad. Stomata 31-48» long (range of means 
40-42). Sori 4-7 (-10) on middle pinnae, sometimes crowded in middle of 
pinna. Spores 29-5-41-5u long (range of means 33-374). Chromosome 
number n= 36, 2n=72. Caloareous om af in Central and SE Europe. (Austria, 
Jugoslavia and Greece. ) 
A DISCUSSION OF RECENT NOMENCLATURAL ACTIVITY WITH REGARD TO THE 
ASPLENIUM TRICHOMANES COMPLEX 

It is obvious that the production within one year of no less than three new 

specific names relating to a complex generally believed to consist of only two 
main taxa must inevitably result in confusion. It may assist clarification of 
this confusion if the publication of these names is discussed in more detail 
here: 


1 THE CHOICE OF RANK Not infrequently, cytotaxonomists are repri- 
manded for an alleged tendency for over-enthusiastic production of new 
“‘ cytological” species. For example, one may quote Davis & Heywood**, 
who write with evident disapproval, “‘ Thus, there is a school of cytotaxono- 
mists which insists that diploids and polyploids must always be kept apart as 
separate species, no matter what the apparent degree of morphological 
differentiation involv 

The most ironic tetas of the present confusion surrounding the 
nomenclature of the Asplenium trichomanes complex is that tkis confusion 
has manifestly been produced by classical taxonomists or nomenclaturalists 
reacting against the reluctance of the cytotaxonomists to give the segregates 
specific rank! It is very remarkable that although Meyer, who having first 
studied these plants for some ten years, produced the first taxonomic treat- 
ment of the cytotypes®, regards them as deserving only subspecific rank, 


RIES Ree ee CET 
4 DAVIS, P, H. and HEYWOOD, V. H., 1963. Principles of Angiosperm Taxonomy. Edinburgh, p.213. 


157 


LOVIS: ASPLENIUM TRICHOMANES 


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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 <n atum erendicen 
3rd Messrs Brookfield: Athyrium filix-femina Druery, A.f.-f. sec nae Dryopteris lineata cristata, 
Phyllitis scolo, ee variegata, Polypodium cambric: hadwinii, Polystichum 
pls tin grac "illim 
as 10 Three distinct varieties of Phyllitis scolopendrium 
N. H. Almond: bifida, Bolton’s ea ‘ondii, vad tict viel: 
aaa Messrs Brookfield: cristata (2), undui 
3rd. B. Hayhurst: crispa, ‘ Robinson’. ares speciosa. 
eure 11 Three distinct varieties of Polypodium vulga 
Messrs Brookfield: cornubiense, grandiceps bifido-mulifum, * Oxford ’. 
oe B. Hayhurst: bifidum, cambricum hadwinii, pulcherrimum. 


gage 12 Three distinct varieties of Polystich 
Z ema oe, ~ istlobems densum, iresyamtem. eel 
bs sium, pulcherrh eee e 
ves Renokfield< Leaded hi, pulcherrii Bevis, setiferum. 


CLASS 13 Three distinct varieties of Athyrium esrgiiene 
Ist F. Jackson: craigii, cristatum, nudicaule cristatum. 


Druery 
3rd N. H. Almond: ci capltatun, cristatum, frizelliae 


SOUTHPORT FLOWER SHOW 1963 193 


CLASS 14 Three distinct varieties of Dryopteris (including wali: and Thelypteris): 
Ist B. Hayhurst: angustata c evistata, line eata, reorient 


2nd __s“E. Jackson: Jineata cristata lydactyl 


3rd Messrs Brookfield: lineata cristata, crispa cristata Barnes, cristata, 


CLASS 15 Three distinct-varieties of Asplenium: bagh ceria species were submitted]. 
st F. Jackson: oe septentrionale, 
2nd __—sONN«’. Robinson: ruta-muraria, septentr ronal, pret: jomanes 
3rd Messrs Broo kfie Id: pei tum-nigrum, trichomanes, viride. 


CLASS 16 Three distinct normal species of hardy British ferns: 


Ist . Robinson: Cryptogramma crispa, Polypodium vulgare, Polystichum setiferum. 

2nd__—s*F. Jackson: Asplenium trichomanes, Dryopteris est ee ten Woodsia ilvensis. 

3rd Messrs Brookfield: Adiantum capillus-veneris, Cry, amma crispa, Polypodium vulgare. 
CLASS 17 One British f racy hwreniceDaoiegad! 

Ist Messrs Brookiield: 4 i 

2nd Mrs Wilkinso: ustin: Po ceniee rire es prereset 

3 F. Jackson: igre apres, -femina frizelliae x fieldi 


CLASS 18 Six dissimilar greenhouse ferns: 
B. Hayhurst: Adiantum cuneatum, Cyrtomium falcatum, Nephrolepis exaltata, Onychium 
japonicum, Platycerium recpanayh Pteris cristata. 
2nd = Messrs Brookfield: Adiantum ‘ Kensington Gem’, Asplenium bulbiferum (2), Davallia 
canariensis, Nephrolepis, ‘Platyeariam alcicorne. 


CLASS 19 Three dissimil 


use ferns 
Mere iin e grin ‘ epsiniacis Gem’, Davallia canariensis, Ne phrolepi 
lumosa. 
2nd ——s«B. Hayhurst: Adiantum cuneatum, Adiantum ‘ Kensington Gem’, Nephrolepis exaltata. 


CLASS 20 One greenhouse f 
Ist B. Hayhurst: Adian : n Gem’ 
2nd . McCorie: Adiantum cuneatum 


3rd M. Plowman: Adiantum cuneatum. 


R. KAYE, N. ROBINSON 


BOTTLE GARDENS AND FERN CASES by Anne Ashberry, with 
illustrations by N. Creina Glegg. 120 pp., 33 photographs, 41 drawings. 
London; Hodder & Stoughton, June 1964. Price 25 shillings. 


Following the theme of her earlier books on miniature gardening Miss 
Ashberry, whose Essex nursery specialises in dwarf conifers and alpine 
Plants, describes her experiences with a variety of adaptations of the principle 
of the Victorian Wardian Case, including cider flagons, decanters and an 
antique wasp catcher, some arrangements (bottle gardens) being completely 


yi 
removal of dead flowers, taking of herbs for cooking, etc. She hopes to Pe 
a modern vogue which will be of immense joy and help to those who fin 
More orthodox cultivation difficult or impossible. 


plant treasures, will realise that this book is a labour of love by real fern 
J. A. CRABBE 


194 BRITISH FERN GAZETTE: VOLUME 9 PART 5 (1964) 
NOTES FROM THE ROYAL BOTANIC GARDENS, KEW 


GLEICHENIA: this genus consists mainly of tropical species that grow on 
waste or barren ground at the edges of forests, roadsides, etc., where they 
get a great deal of light. Such conditions seem easy to emulate in the 
hothouse but I find these ferns fail to make much growth if potted in 
rich fern compost. Instead, if put in bracken peat only, and given plenty 
of water, they will soon spread and cover the staging. They apparently 
dislike having their foliage cut, so any withered or dead frond is best 
left on the plant. Gleichenia may be raised from spore if sown within a 
week; for this reason the number of species in cultivation is small 
(only G. linearis at Kew). 


TREE-FERNS: there is a tendency for some species which are easy to grow 
in the heated house, e.g., Cyathea hookeri, C. sinuata, to have weak 
trunks and fall over. If the trunk is covered with sphagnum moss held 
in place with wire netting, root production throughout the stem is 
increased and these bind together to help strengthen the trunk. Such 
plants do much better in open beds in the house rather than in confined 
tubs, and with a healthy root mat formed above ground they can always 
be cut off at ground level and lowered if the fronds are reaching the roof. 

OPHIOGLOSSUM PENDULUM: a plant from Ceylon, on the roots of a 
Birdsnest Fern (Asplenium nidus), is now fruiting. 

FERNERIES, KEW H. J. BRUTY 


WILLIAM CULLEN BUCHANAN 


On 13 September last year died Willie Buchanan, the last of the family to 
farm Garscadden Mains, Bearsden, Dumbartonshire, for almost 150 years. 
He was born at the farm on 27 January 1887, and while there built an 
entrancing landscape rockery. Shortly after the Second World War, when 
most of the farmland was taken over for Glasgow development, he retired 
to a house nearby where he constructed a new half-acre garden. 


Quite unique, a bachelor, an uncannily successful grower of very difficult 
plants, Member of the British Pteridological Society, the Alpine Garden 
Society, the Scottish Rock Garden Club and Fellow of the Royal Horti- 
cultural Society, he had been interested in plants all his life. Hundreds of 
thriving hardy plants, including rare ferns, were assembled in his beautiful 
new garden*, and one sincerely hopes his treasures have found good homes. 


EES Saar Taran 
aoe tributes and pictures of the Douglasbank garden, see J. Scottish Rock Garden Club 91) April 
1 = 


WILLIAM CULLEN BUCHANAN 195 


As a judge at the Shows of the Scottish Rock Garden Club his services 
‘were always in great demand. His unassuming generosity gave freely of his 
plants and advice to anyone interested and keen, although he found it 
difficult to suffer fools. He was most unwilling to put anything down in 
writing; it is believed that some of the early “‘ Plant Notes” in the Journal 
of the Scottish Rock Garden Club were by him. “‘ Some favourites ” in 1954 
{loc.cit. 14:65, 15:142) caused him considerable annoyance because the 
editor had inserted “‘ by W.C.B.”; the second part contains interesting short 
notes on a number of our rarer British ferns. 


Willie will long be remembered with great affection by all who knew him. 
In his memory the Scottish Rock Garden Club has inaugurated an annual 
“William Buchanan Lecture” devoted to the cultivation of rare and 
difficult plants. 


JAMES DAVIDSON, REGINALD KAYE 


FREDERICK GEORGE PRESTON 


My father, an Honorary Member of the Society, died at Cambridge on the 
8th of January this year aged eighty-one, leaving a widow, two sons and a 
daughter. 


Born at Warborough in Oxfordshire, his outstanding career in horti- 
culture began in 1904 with training as a student gardener at the Royal 
Botanic Gardens, Kew. In 1909 he was appointed to the staff of the 
Cambridge Botanic Garden as Foreman-in-charge of the Outdoor Collection. 
From 1919-1947 he was Superintendent of the Garden. After his retirement 
he lived at Shelford on the outskirts of Cambridge where, until he 
ill about two years ago, he continued his longstanding work with the Royal 
Horticultural Society, on committees and advisory panels, and as an examiner 
for the National Diploma of Horticulture. 


His published work on horticulture includes books, articles in journals 
and contributions to the R.H.S. Dictionary of gardening. 


In recognition of his many services to horticulture, Cambridge 
University honoured him with the degree of Master of Arts; the Royal 
Horticultural Society made him an Associate of Honour in 1936 and two 
years later he received their highest award, the Victoria Medal of Honour. 

G. H. PRESTON 


me 


Sexe 
a Pode nee tee) F 
ve aah | 


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THE 
BRITISH FERN GAZETTE 


VOL.9 PART 5 1964 


Contents 
Dryopteris assimilis S.Walker in Britain 
S. WALKER & A. C. JERMY 137 


A diploid form of Asplenium ruta-muraria 
J.D. LOVIS & T. REICHSTEIN 141 


The taxonomy of Asplenium trichomanes in Europe J. D. LOVIS 147 
Two species of Diplazium in Europe ... rep A. C. JERMY 160 
Asplenium majoricum Litardiére A. C. JERMY & J. D. LOVIS 163 


The Polypodium vulgare complex in North America 
R. M. LLOYD & F. A. LANG 168 


Cultivated ferns me ae ay ae J. A. CRABBE 178 
Short notes: 

Notes from the Royal Botanic Gardens, Kew H. J. BRUTY 194 

Certificates of Merit oe. ee J. W. DYCE 183 

Distribution of the British ferns = their allies J. W. DYCE 184 

The European Chainfern in Corsica... G. SCHULZE 162 
British Pteridological Society, reports of meetings ... ats er 186 
The Southport Flower Show 1963... me i sie tse 191 
Obituaries: 

William Cullen Buchanan... wis we me oe ae 194 

Frederick George Preston... ea ae oe ree ae 195 
Book review at Si hice aut ak beg cay ae 193 


Courier Co. Ltd., Tunbridge Wells, Kent 


TTie 
BRITISH FERN 
GAZE] TE 


VOLUME NINE PART SIX 


1965 


Mesoun: Borg Nice 
SEP 19 1966 


GARDEN LiBRaRy 


THE BRITISH 
bo PTERIDOLOGICAL SOCIETY 


BRITish FIERIDOLGGICAL ~SOCIETY 
Officers and Committee 1964-65 


PRESIDENT 
Reginaid 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 
Hilltop, 46 Sedley Rise ° 
Loughton, Essex 
EDITOR 


A. C. Jermy BSc FLS 
British Museum (Natural History) 
Lo SW7 


ASSISTANT EDITOR 
J. A. Crabbe 
British Museum (Natural History) 
London SW7 
COMMITTEE 
Cartwri T. A. Dyer FLS, 

Py, Hallizey. Se AMIEE Macias, Secretary) B. decile 
F. J. Healey, Jean R. Healey BSc, F. Jackso 
Frances M. Jarrett MA PhD FLS. R. M. Payic 

R. S. W. Pollard LAMTPI, N. Robinson 
AUDITOR 
Peter Temple LRIBA MInstRA 


The SOCIETY is open to all interested in any way in FERNS and FERN 
ALLIES u Se paying an annual subscription of SEVENTEEN SHILLINGS & 
SIXPENCE. Full details will be sent upon application to the SECRETARY. 


BRIT. FERN’ GAZ. 9 (6): 1965 197 


VARIATION IN ATHYRIUM IN THE 
BRITISH ISLES 
REGINALD KAYE 


Waithman Nurseries, Silverdale, Lancashire 


AsstractT. About eighty varieties of Athyrium filix-femina remain today from 
a heritage of several hundreds; they are ae va discussed in a classification 


based on that used by E. J. Lowe in 1890. No extant varieties of A.distenti- 
folium or a ‘fel are known to the author. ‘ Var iety ’’ is used in preference 
to “* it is synonymous under the International Code of 


cultiv with which 
basen peat for Culivessd Plant 


Three native species of ladyfern grow in the British Isles: 

1 Athyrium distentifolium, the Alpine Ladyfern, is locally abundant above 
one thousand feet in the central Highlands of Scotland up to Sutherland, in 
shady places by streams and among rocks. It is very similar to 3, differing in 
having a rudimentary indusium which usually falls off or becomes invisible 
ong before sori nearer tl in of the lamina; 

a tighter’ more “ golden-green colour when growing in its 1 native mountain 

fastnesses. There seems to be no difficulty in cultivating it in lowland habitats. 
I know no living varieties. 

2 Athyrium flexile. This rare species is often included within 1, but it 
differs in having fronds Sule 9-12 inches long; stipe very short and deflexed: 
pinnae short, distant, reflexed; pinnules well-separated and deeply cut. 
It has been reported from high alpine corries in Glen Clova, Ben Lawers 
and Ben Almond, but has not been seen lately. In 1962 the British Pterido- 
logical Society spent some hours in the mountains at the head of Glen Clova 
and founds lots of distentifolium, but no one sighted flexile. No varieties are 


ra wn. 
Athyrium filix-femina, the Ladyfern, is very common and widely distri- 
a throughout the British Isles in damp shady places. The perennial root- 
stock bears dark-brown scales and becomes branched and woody and covered 
with old frond-bases. The deciduous fronds are lanceolate, from 12-60 
inches long, and usually bipinnate. The pinnae usually end in slender acute 
points. The pinnules are sometimes so deeply cut that the frond is almost 
tripinnate. The stipe and rhachis is usually green, darkening towards the 
base, but is sometimes purplish-red. It is quite difficult to find two plants 
ape: fronds exactly alike. 
nnate tendency of British ferns to produce variations is shown to a 
greater pny in the ladyfern than in any other species, and while there are 
certain well-defined varieties, these are combined and recombined to such a 
degree that there seems to be no easy way to classify them. The number of 
varieties in cultivation today (see list on page 203) which have come down to 
us from the collections of seventy years ago is difficult to estimate, but 


The BRITISH FERN GAZETIE Vol. 9 Part 5 was published 22 July 1964 


198 BRITISH FERN GAZETTE: VOLUME 9 PART 6 (1965) 


is probably in the region of 80, though there are no doubt in gardens quite 
number existing whose names and origins are unknown to the owners. 
On the other hand many plants exist which have been raised from spores, 
intentionally or by chance, which do not quite agree with any of the described 
varieties, but which may be quite good or even excellent varieties in their own 
ight. There are also very many irregular forms to be seen which are not worth 
keeping and should be destroyed to make room for more-worthwhile ones. 
E. J. Lowe in British Ferns and where found (1890) described and classified 

no less than 313 varieties of ladyfern (including 75 which had received First- 
Class Certificates from the Royal Horticultural Society or the Royal Botanic 
Society); in his Our native Ferns (1876) he had given 294 (including 58 he had 
not seen, raised or found by other fern enthusiasts). There is no doubt, how- 
ever, that in Lowe’s day there was a tendency to record and name every 
slight variation or irregularity with resounding, often hyphenated, latinised 
ioe om C. T. Druery in British Ferns and their Varieties (1910) de scribed 
50 ladyfern varieties which may or may not all have been in cultivation at 

the time. Very few new varieties have been described since 1910. In 1892 the 
catalogue of H. B. May & Sons (Upper Edmonton, London) listed 138 
adyfern varieties, while in 1908 W. & J. Birkenhead (Sale, Manchester) 
offered only 88; these collections ete long been dispersed and the nurseries 
have disappeared. In 1920 W. F. Askew listed 36 varieties of ladyfern in his 
Borrowdale Nurseries catalogue; this fine collection is now mostly oe 


varieties of Athyrium there. Soon after the war I acquired the entire collection 

Robert Whiteside in which there were many ladyferns which he had 
collected in a lifetime devoted to growing hardy ferns. In my own fern 
catalogue some 27 varieties of ladyfern are listed, though I have others. 
many of these are now transferred to reserve beds for propagation. 

The only way to propagate many varieties is by division, a slow process. 
Certain kinds can be reproduced true-to-name from spores, e.g., ‘Frizelliae’ 
comes at least 90% true; others are not so reliable and sporelings have to be 
grown-on for two or three years and rogued before they can be offered as 
reasonably true-to-name. 

Basically, variation is of two kinds: variation in frond-shape due to 
cresting, and variation in pinnule-shape. Both types can occur together, 
making classification somewhat difficult; I remember discussing the problem 
with W. B. Cranfield and he rather deprecated the tendency of British ferns 
trying to do too many things at once and recommended that some varieties 
be discarded on this account. It is said, however, that beauty lies in the eye 
of the beholder and, provided that a variety has beauty and symmetry OF 
interest of some sort, it should at least be recorded and grown by those who 
wish to cultivate it. 


cms 


inches 
FIGURES 1-9 Varieties of . bien Abate filix- femina 
I Psst’ Medusae’, 2 ‘Foliosum Grandiceps’, 3 idiceps’ (dwarf form), 
gree Bolton’ origi clone), 5 Setigerum “Corymbiferum Grubb’ 
(original clo 6 ‘Acrocladon’ (original clon zelliae’ (original clone), 
Frizelliae C istatum’ jews spore), 9 “Cc eorabiferans 


KAYE: ATHYRIUM VARIATION 199 


Lowe (1890) divided 313 varieties of ladyfern into two main divisions, 
five groups, and fifteen sections. As was done by Dyce for Polystichum 
(GAZETTE 9: 97-109), I have adapted Lowe’s classification to analyse the 
ladyfern situation today. 


DIVISION A CHANGE IN FROND SHAPE 
Group | CRISTATE 

(a) capitate (apex crested) 
This section is exemplified by ‘Coronatum’ which makes a neat plant with 
a large corymbose crest, but the pinnae are not crested; my plant is more 
robust than the one described by Moore. ‘Capitatum Wollaston’ I had from 
William Askew but lost it. 

(b) cristate (apex crested in one plane; pinnae crested) 
Very many forms are to be found of this common type of variation in which 
the crests are fan-shaped and in one plane, for example ‘Cristatum’ (Fig. 27). 
When the divisions of the crests are long and slender and well-separated 
they are known as polydactylous; a good example today is the very attractive 
*Multifidum 


(c) percristate (apex, pinnae and pinnules crested) 
I have fine large examples of ‘Percristatum’, and a particularly pretty plu- 
mose one mentioned below under B(a). 


(d) corymbose (apex crested in several planes to give a bunched 
cluster; pinnae crested) 
Fine forms, for example ‘Corymbiferum’ (Fig. 9), are in cultivation, though 
it is doubtful whether many of the varieties described in Druery still exist. 
‘Diadematum’ has a very large head and many of the pinnae are heavily 
crested. ‘Glomeratum’ is a particularly fine example, almost a section in 
spa defined by the pinna-crests being corymbose and wider than the pinnae. 
mmatum Barnes’ with a green stem and ‘Gemmatum Bolton’ (Fig. 4) 
sk a red stem are perfect examples of sare corymbose varieties, 
both with large almost spherical crests on all pinnae. ‘Depauperatum 
Edelsteinii’, which I acquired with the Whiteside parental might well be 
included here; apart from the fact that it is not depauperate, it has a pecu- 
liarly twisted terminal crest and the whole plant is a particularly vivid green. 
A corymbose crest, combined with other characters, occurs in other sections. 
(e) grandicipital (terminal crest wider than rest of frond; pinnae 
crested, reduced, or absent 
In this section are many forms ranging from tall strong-growing hnaagoa bee 


well-developed pinnae to varieties with reduced or depauperat 
“Nudicaule Cristatum’ has no pinnae at all except those reel in 


200 BRITISH FERN GAZETTE: VOLUME 9 PART 6 (1965) 


immense terminal crest, so that the frond is reduced to a ball of verdure on a 
long stalk. ‘Caput-Medusae’ (Fig. 1) might be regarded as an extreme 
grandicipital, or it might equally be included as a ramose under 2(b). The 
many fine grandicipitals raised by Ellworthy seventy years ago I have not 
seen since before the Second World War. There are dwarf, otherwise typical, 
forms of grandicipitals, one of which (Fig. 3) is about 12 inches high; 
‘Cristulatum Stansfield’ is about 9 inches high. 


Group 2 OTHER CHANGES IN FROND SHAPE 


(a) cruciate (a cross formed by opposite pinnae; each pinna usually 
reduced to a single pair of enlarged, + right-angled, opposite 
pinnules) 

This is a large section and is usually combined with cresting. I do not know 
whether the morphology of cruciation has been worked out; one theory is 
that the whole of the pinna, except the pinnules nearest the rhachis, is sup- 
pressed, the two remaining pinnules developing to the size of pinnae at an 
angle to one another, thus forming a cross with the opposite pair of pinnules. 
A perfect example is ‘Fieldii’ (Fig. 12) whose long arching fronds show a 
chain of crosses with no cresting. ‘Pritchardii’ (Fig. 10) is another charming 
example on more slender lines, with more than four pinnules involved in 
each cross so that the pinnae form little tufts. ‘Cruciatum Cristatum’ 
differs from ‘Fieldii’ in being crested, slenderer and about half the length. 
‘Angustocruciatum Cristatum’ (Fig. 11) has very narrow fronds, perfectly 
cruciate throughout, with a neat flat crest. The crested cruciate par excel- 
lence is ‘Victoriae’ (Fig. 13), a wild find discovered in a country lane in 
Stirlingshire over a hundred years ago; the original form is still the strongest, 
making fronds 2-3 feet long of perfect symmetry, narrowly lanceolate, with a 
good terminal crest and crests to all pinnae; divisions from the original are 
still with us and show no loss in vigour; how long these divisions of the 
original clone will continue to maintain their fine constitution remains to 
be seen; plants raised from spores show the original form but have never 
equalled its size; some very graceful fine forms have been raised; ‘Victoriae 
Foliosum’ (Fig. 14) is in cultivation; ‘Setigerum Victoriae’ is included under 


ic): : ruciatum Congestum’ is a stiff dark-green dwarf of 6-9 inches, not 
crested. 


(b) ramose (lower part of main axis divided once or more) 
‘Acrocladon’ (Fig. 6) the finest example of this section, produces a ball of 
verdure by repeated branching of the rhachis; as it produces spores only 
occasionally it has to be increased by division, a slow process; when spores 
have been produced and sown, even more-remarkable forms have been raised, 
but whether these are still in cultivation I do not know; it was originally 4 
wild find. ‘Ramocristatum Sim’ does not appear to be about, though 
unidentified ramocristates are in cultivation. The brachiate varieties, in 


cms 


inches 
FIGURES 10-19 Varieties of Athyrium igpattitionan 
10 ‘Pritchardii’ Lan clone), 11 ‘Angustocruciatum neh m’, 12 ‘Fieldii’ 
(Original clone), 13 ctoriae’ (original clone), | ctoriae Foliosum’, 15 
‘Flabellipinnulum’, 16 “Minutisimumn’, 17 yea Minus’, 18 ‘“Crispum 
Coronans’, 19 ‘Vernoniae’ (original clone). 


KAYE: ATHYRIUM VARIATION 201 


which the rhachis divides into three as though the basal pinnae had developed 
into fronds (which may be the cause of brachiation) have, I think, been lost. 


(c) congested (pinnae close together, often overlapping, usually 
dwarf) 


Several varieties exist, seldom exceeding 6 inches in height. ‘Congestum 
Minus’ (Fig. 17) is extremely congested with pinnae and pinnules all over- 
lapping; it comes true from spores. ‘Congestum Grandiceps’ is heavily 
crested and suitable for the rock garden. ‘Crispum’ is a ramose, crisped 
and crested dwarf, forming 4-inch-high mats. “Crispum Coronans’ (Fig. 18) 
is more heavily crested but seldom exceeds 5 inches. ‘Minutissimum’ (Fig. 
16), though not strictly congested, might be included here; it is a perfect 
miniature ladyfern no more than 2-3 inches high in the open and 5-6 inches 
in shade, a charming wee fern for the front of the fern border. 


(d) angustate (fronds narrow with short pinnae; pinnae twisted, 
reflexed, or reduced to bal 
The narrowest Jadyfern frond is shown by ‘Frizelliae’ (Fig. 7), found wild on 


grows about 12 inches high; sports from the original include several striking 
crested varieties such as ‘Frizelliae ners (Fig. 8), ‘Frizelliae Capitatum’, 
‘Frizelliae Ramocristatum’, and some of these attain 18 inches or more; 
there are ramose dwarf forcna® one very dwarf, otherwise normal form of 
Only 3 inches I had from Askew, but it is gone. ‘Curtum’, found in Lakeland, 

is a neat dwarfish variety with very short pinnae; the width of the frond seldom 
exceeds 3 inches, usually less; there is a neat crested form of this, 9-12 inches 
high. Unequal growth of the upper surface of pinnae can result in twisted 
and reflexed varieties, such as ‘Revolvens’; this is still in cultivation I believe, 
but to my way of thinking it is more curious than beautiful. The more 
desirable ‘Reflexum Fox’ was a neater and more interesting plant, but I fear 
it may be lost. 


DIVISION B CHANGE IN PINNULE SHAPE 


(a) plumose (pinnules pinnate or bipinnate; sori scanty or absent) 

This section contains some of the most beautiful ladyferns, their tripinnate 
and sometimes quadripinnate form giving the fronds an especially feathery 
and graceful appearance, coupled with a particularly lucent, sometimes golden- 
green colouring. Several forms have been found in the wild and recorded by 
their finders, though not many of the originals have persisted to the present 

y. ‘Plumosum Stansfield’ (Fig. 20) is one of the many forms found in 
the wild. The finest forms have been propagated from spores. In a well- 
8rown specimen of ‘Plumosum Druery’ (Fig. 21) the large quadripinnate 
fronts are a wonderful sight. I have a similar form, ‘Plumosum Penny’ 


202 BRITISH FERN GAZETTE: VOLUME 9 PART 6 (1965) 


(Fig. 23), raised by Penny of Lancaster. Charles Grubb had some out- 
Standing seedlings which still exist I believe. A further development, the 
examples of ‘Plumosum Superbum’ raised by Druery and others, are extra 
feathery and crested; ‘Plumosum Superbum Druery’ is still about and is 
usually prominent at Southport Show. There is a nice percristate plumose 
in cultivation, very delicate in appearance. ‘Plumosum Miultifidum’ (Fig. 
24) is a distinct broad-fronded variety with loosely-crested pinnae on 
multifid lines. ‘Plumosum Furcillans’ (Fig. 22) is very dainty and lightly 
crested, and there are several others which tend to lose their attractive 
plumose character as they become more heavily crested. The best varieties 
have to be increased by careful division and, in consequence, are scarce. 
Perhaps the most remarkable variation in this section is ‘Clarissima Jones’ 
(Fig. 26); this has extremely slender attenuated pinnules, and in a good 
environment can produce fronds 3-4 feet long and 2 feet wide; I had thought 
it extinct, but recently acquired what I believe to be the true form; as it 
takes about five years before a double crown is produced, it is not likely 
to appear in Woolworth’s for some time. ‘Clarissima Bolton’, found in 
Lancashire, is smaller than Jones’s and is even more finely cut; plants of it 
usually appear at Southport Show, but they are the jealously guarded trea- 
sures of one or two growers and pearls would not buy them. ‘Clarissima 
Cristatum Garnett’ (Fig. 28), also known as ‘Fimbriatum Cristatum Gar- 
nett’, is a most elegant variety with very wide light-golden-green fronds, 
chastely crested, and it grows quite tall; I have had fronds over 3 feet long 
and 2 feet across; it is slow to increase. These forms of ‘Clarissima’ all 
show the phenomenon of apospory, but very few youngsters so raised have 
been worth growing. 


(b) foliose (pinnules wider than normal, almost confluent, often 
overlapping) 
The pinnules are lushly foliose. The only living example I have seen, ‘Folio- 
sum Grandiceps’ (Fig, 2), I acquired with Whiteside’s collection. 
(c) dissect (pinnules finely divided into teeth or bristles) 
This is a large section. ‘Setigerum’, found in Lakeland by J. Garnett, is 


beautiful things such as ‘Setigerum Cristatum Garnett’ and ‘Setigerum 
Corymbiferum Garnett’; ‘Setigerum Corymbiferum Grubb’ (Fig. 5) is a 
superb red-stemmed perfectly symmetrical plant. The feathery ‘Pulcher- 
rimum’ comes into this section, but is seldom seen. Grubb raised ‘Setigerum 


my original plant has worked up to five crowns in the last fifteen years, sO 
in another fifteen years I may be able to list it under ‘stock limited’. 


cms 


inches 
oe FIGURES 20-28 Varieties of Athyrium filix-femina 
gh i oben Sencar Rit goal clone), 21 ‘Plumosum Dru atl ee clone), 
< urci umosum Penny’ (original l ‘Plum 
M ginal clon mosum 
pe stedaoi : 25 ‘Ho owardii’ (original clone), 26 6 “Clar rissim a Jones’ elie clone), 
atum’, 28 ‘Clarissima Cristatum Sadie oneal clone). 


KAYE: ATHYRIUM VARIATION 203 


(d) depauperate (pinnules reduced, irregular, or missing) 

Many of the depauperate varieties are asymmetrical and not worth garden 
room. Exceptions are the symmetrical varieties such as ‘Mediodeficiens 
Polydactylum’, a very pleasing plant with loose-fingered crests and with 
the segments much reduced towards the rhachis. Other good forms such as 
‘Defectofurcillans’ and ‘Paucidentatum Cristatum Mapplebeck’ are extremely 
rare if indeed they still exist. 

(e) heteromorphic (pinnules rounded, narrow, or jagged) 
‘Flabellipinnulum’ (Fig. 15) is an interesting variety with narrow fronds, 
Slender pinnae and tiny pinnules. ‘Vernoniae’ (Fig. 19) is very distinct 
with the pinnae crisped and of broadly triangular outline. I doubt whether 
any of the varieties with tiny lunulate pinnules (for example ‘Pulleri’) or 
narrow pinnules still remain; I would very much like to have them if anyone 
knows of their existence. 


VARIETIES OF ATHYRIUM FILIX-FEMINA KNOWN BY THE AUTHOR 
TO IN CULTIVATION TODAY 
Angustate: ‘Curtum’, ‘Curtum Cristatum’, ‘Frizelliae’, ‘Frizelliae ne 


tatum’, ‘Frizelliae Cristatum’, ‘Frizelliae Multifidum’, Friz elliae Nanum 
‘Frizellia ae Ramocristatum , ‘Re flexum Fox’, ‘Revolven 

Capitate: ‘Capitatum Wollaston’, Reon wer 

Congested: ‘Congestum’, ‘Congestum Cristatum’, Cong estum Shana ale 


Cotes estum Minus’, ‘Cri aa “Cris spum Coronans’, eMinutis 
ove ne Be sr ness ae forms), ‘Depauperatum Edelsteini’, 
aoe ‘Gemma Barnes’, ‘Gemmatum Bolton’, ‘Glome- 


Cristate: “Cristatum’ = several cristate forms in other sections), ‘Multi- 

fidum’, ‘Poly 

Cruciate: ‘An se ae Cristatum’, ‘Crucia um Congestu um’, ‘Cruci- 
atum pevineectettt Field, ‘Pritchardii’, svictoriae’, Victoriae Foliosum’ u 
“Victoriae Gra 

Depauperate: ariecaiasllta ‘Mediodeficiens Polyactylum’, ‘Pauciden- 
tatum Cristatum Mapplebeck’. 


Dissect: ‘Denticulatum’, ‘Howardii’, ‘Setigerum’, ‘Setigerum Corymbiferum 
Garnet Scomean m Corymbiferum Grubb’, ‘Setigerum Cristatum Garnett’, 
‘Setizerum Vict 

Foliose: ‘Folios Geandidiee (also ‘Victoriae Foliosum’ above 

Grandicipital: ‘Caput-Medusae’, ‘Cri stu latum Stansfield’, ‘Grandiceps’ 


Heteromorphic: ‘Flabellipinnulum’, ‘Pulleri’ , ‘Vernoniae’, ‘Vernoniae 
Percristate: ‘Percristatum’ (several forms, also ‘Plumosum Percristatum’ 
below). 


Plumose: ‘Clarissima Bolton’, ‘Clarissima Jones’, ‘Clarissima Cristatum 
Garnett’ (also known as ‘Fimbriatum Crista tum Garnett’), ‘Congestum 


Paul’, ‘Plumosum Druery’, ‘Plumosum Penny’, » ‘Plumosum Stansfield’, 
*Plumosum Cristatum’, ‘Plumos Furcillans’, Plumosum Multi 
ercristatum’, ‘Plumos per ruery’, 


Ramose: ‘Acrocladon’, ‘Ramocristatum’, ‘Uncoglomeratum 


204 BRITISH FERN GAZETTE: VOLUME 9 PART 6 (1965) 


over the years. It is hoped that anyone possessing unusual forms of ferns 
will practice good cultivation and propagation to ensure the health and 
perpetuity of our ferny wealth. In nature, athyriums receive a yearly top- 
dressing of leaves, which helps to maintain a moist compost around the 
rhizome. In gardens, the soil-level usually remains the same or falls because 
of weeding and tidying, and consequently the apex of the rhizome becomes 
more and more exposed as it grows upwards; for this reason replanting is 
advisable every three or four years. 
hope these remarks may have thrown some light on the intricacies of 
variation in the ladyfern and will help in sorting out some of the plants in 
readers’ hands. 
I am indebted to Mrs Joyce Parker of the British Museum (Natural 
History) for preparing the illustrations from plants growing in my nursery. 


REVIEW 


THE GARDENER’S FERN BOOK by F. Gordon Foster, with drawings and 

color plates by the author: j ] 

18 black-and-white Photographs: Princeton (New Jersey), New York, Toronto, 
ndon: Nostrand, 64 (U. 

1965 (U.K.); price $7.95; 62 shillings. 


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7 
Q. 
8 
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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 

<GRAMMITIS nu Pea (Maxon) Proctor, comb. noy., based on Polypodium 
nubigenum Maxon in Contr. U.S. Nat. Herb. 17: 599 (1916). P. Pree 
sensu Jenman in Bull. Bot. Dept. Jamaica II, 4: 120 (1897), non Desv. (1811). 
TYPE from Jamaica, Maxon U a 

“Gramm itis randallii (Maxon) Proc mb. noy., based on Polypodium 
Sy sini Maxon in Amer. Fern Journ. 16: “46 (1928). TYPE from Jamaica, 
Maxon 9280 (US. isotype IJ). 

vGRAMMITIS rigens (Maxon) Proctor, comb. nov., based on Polypodium 
rigens Maxon i he U.S. Nat. Mus. 27: 741 (1904). P. rigescens sensu 
Jenman in Bull. t. Dept. Jamaica IL 4: 117 (1897), non Bory ex Willd. 
(1810). TYPE pei Jamaica, Maxon 1346 (US). 


Neuropium Fée in Mém. Soc. Mus. Hist. Nat. Strasbourg 4: 201 (1850) (as 
Nevrodium). This generic name has priority over Paltonium Presl. Epim. Bot. 
156 (1851). The plant customarily called Paltonium lanceolatum (L.) Presl 
8 (i ‘aaa be known as Neurodium lanceolatum (L.) Fée, Mém. Foug. x 


“PITYROGRAMMA ebenea (L). Proctor, comb. nov., based on Acrostichum 

ebeneum L., Sp. Pl. 2: 1071 (1753). derostcium pn eit Cav . 242 

(1802); Pityrogramma tartarea (Cav.) Maxon in Contr. U.S. Nat. Herb. 17: 
173 (1913), Proctor in Bull. me eae "Sci. Ser. 5: 44 (1953), ete. 

In a communication W. T. Stearn opinion that 

the Linnaean epithet for this pow must reinstated and poirts out that 

“Article 8 of the International Code of Botanical ee val wage se 


220 BRITISH FERN GAZETTE: VOLUME 9 PART 6 (1965) 


ad before him a specimen which still exists in the Linnaean Herbarium 
(sheet 1245. 14, named and mg hh red in his pee hepa his dia fie not 


This practice must be discontinued, as a lectotype cannot properly be based 
ona a synonym, especially whsies a perfectly good type specimen 
is ava 


PITYROGRAMMA Williamsii Proctor, sp. 
Rhizoma curvato-ascendens ap‘ce sate. paucis occultis nitide brunneis 
subclathratis deltato-acuminatis ad 2 mm longis praeditum. Frondes 


purpureo-brunnei 5-10 cm longi. Laminae ambitu lineari-lanceolatae vel 
reed cst coiwcee ‘AS-27 cm longae, 3. dhe m latae, apice ee oe 


Sar apicem versus anguste nde a pect dense ces sede 
pinnae anguste deltatae, pinna longissima c. 3 cm cae 1.2 cm lata, brevipetio- 
latae, apice acutae; pinnulae e basi inaequilaterali cuneata ovato-oblongae, 


basi pinnatifidae omnino pin ; segmenta ultima flabellato-cuneata, 
| 1s; venae supra canaliculatae praeter apices leviter dilatatos qui 
desinunt versus apices lob a non-vascularis rigide 


membranacea, atroviridis, opaca, subtus pulvere eae dense induta. 
Sporangia numerosa in a ultimis penultimisque dispos 
izome nding, bearing at ae a few nebo, glossy brown, 
subclathrate, deltate acuminate scales up to 2 mm long. Fronds numerous, 
densely tufted, stiffly erect, 20-35 cm lon: na: views om purple-brown, 
] lade 27 cm 


3-pinnate; rhachis shining purple-brown, oe — win nged towar ard a apex, the 
wing densely yellow-powder beneath; pinnae narrowly deltate, the longest 
c. 3 cm long and 1.2 2cm broad, short-stalked, the apex acute; pinnules ovate- 


green, 
Sporangia numerous on the nites and penultimate vein 

TYPE: Parish of Westmoreland: coon Mountain n Wo ods, c. 1 mile 
W.S.W. of Rat Trap, elev. 1200-1300 ft., Proctor 21469, collected aces 23, 
1960. PARATYPEs: Westmoreland: Enfie Id, 3 miles so outh-eas t of Darliston, 
Proctor 4662; Leami ar paar Orcutt 7451 (Ki), Chocho Gully, along spe 


men c 
‘wi t Country c. 5 miles north of Quick Step, Proctor 4081. 
The latter differs chiefly i in having iecaaer, more membranous fronds bearing 
much less powder beneath. 


PROCTOR: TAXONOMIC NOTES ON JAMAICAN FERNS 221 


Pityrogramma williamsii occurs on partly- ee calcareous banks, 
at 


co p 
ultimate lobes in which the veinlets run towards the tips. The veins of P. 
renal always run to a small sinus. Named for Bernard Williams of Copse 
untain House, long a patron of Tuithican natural history, upon whose 
sath the type specimens were collected. 


rage oases LEUCOSTICTON Kunze in Linnaea 20: 380 (1847). 

r many years it has been assumed that the Jamaican fern named 
Yohinaiuns microchasmum Baker i n Journ. Bot. 16: 44 (1887), which was 
i j . & Cham. 


(aided by a grant from the Dodge Jones Foundation, made available by 
ndell) was able to observe P. plebejum ing abundantl 
Guatemala. It h ome nt that cage smum is n 


a eju. ring 

structure of the de ore and in the su aienowniac fronds with stouter 
and more or less winged s By these characters the Jamaican plant 
approximates the taxon nen Saest Te al leucostiction by yeaa big age Pa S 


s s. P. micro- 
chasmum should therefore be transferred to the synonymy of P. leicostiea. 
Maxon implied that the latter name might include pepo elements, but 
so far as I know this question has not eceved further st 


STIGMATOPTERIS jamaicensis (Desv.) Proctor, comb. , based on Cysto- 
Pteris jamaicensis Desv. in agers 7% Linn. Paris 6: 363 (1827). Dryopteris 
chlae ; in 


Con 
. Inst. Jamaica Sci. Ser. 7 36 (1933). “Desvaux says that his plant came 
from “Cuba et eee , but a specimen from his herbarium pater the 
type) at Paris simply says ‘in Antillis’. The species is now considered 
endemic to Jamaica. 


222 BRITISH FERN GAZETTE: VOLUME 9 PART 6 (1965) 


THE INFLUENCE OF SHADED CONDITIONS 
ON THE FERTILITY OF BRACKEN 


MATTHEW J. DRING 
Department of Botany, Westfield College, London, NW3 


ABSTRACT 
Qualitative observations in various parts of southern England, and a few 
northern localities, have shown that Bracken Preridium aquilinum 1s 
less fertile when growing in shaded situations. Measurements of 


tion with the other habitat factors measured. 


INTRODUCTION 


This investigation was originally suggested by the observation, in the 
summer of 1959, that large bracken fronds growing under fairly dense shade 
in a pine plantation on the North Downs, were apparently completely sterile. 
Apart from brief notes on light and frond morphology (Tansley 1939), 
research into the extensive literature on bracken (Braid 1959) revealed little 
beyond a few casual remarks that “sporophylls do not occur in nature in 
shaded situations” (Goebel 1905, Schwabe 1951), although Conway (1957) 
has concluded that ‘“‘fertile fronds develop in any habitat where the rhizome 
is well established, but that extreme conditions which retard vegetative 
growth, also restrict sporangial development.’ The only indication of a 
detailed investigation of the problem is a note by Conway (1957) that in 
“fairly thick shade” in a Stirlingshire oakwood, fronds were 21% fertile, 
but at the edge of the wood, 200 yards away, they were 40% fertile; no 
indication of the relative degree of vegetative development was given, nor 
any measurements of the physical factors of the habitat. 

Several detailed investigations of the effect of environment on the 
anatomy of bracken (Boodle 1904, Woodhead 1906) show that shading does 
have profound effects on the bracken plant, including the loss of the leaf 
hypoderm, and the reduction of the palisade layer, and of cuticle and leaf 
thickness. These investigators also suggested that in shaded situations the 
number of sterile leaves is very much greater than in exposed areas, and 
that the sori are small with reduced marginal indusia (Boodle 1904). 


attempt to measure these effects quantitatively in the field (Burke 1953), 
ignored this advice and gaily concluded that light was the only important 
factor influencing the size of shaded fronds. 


DRING: SHADE AND BRACKEN FERTILITY 223 


The present investigation aims to demonstrate that bracken is less 
fertile in shaded situations, in spite of satisfactory vegetative growth, and to 
examine its response to varying light intensities in the field. In addition, as 
many of the other physical factors as possible have been measured in an 
attempt to sort out the complex of factors involved. 


GENERAL OBSERVATIONS 


e more or less complete sterility of bracken fronds when growing in 
shaded situations has been observed in a large number of widely differing 
sites, mainly concentrated in southern England but also including mixed 
woodland in the Lake District and in southern Scotland. Such general 
observations have covered a wide range of British climatic types, and also 
a wide range of soils, including those derived from sandstones, limestones, 
clays and sand, of various ages between Cambrian and Quaternary. A few 
examples of areas in which bracken was observed to be fertile in the open 
and sterile in the shade are: 


(1) Michaelwood, Gloucestershire: mixed deciduous woodland on 
ambrian rocks 
(2) Brean Down, Somerset: exposed and maritime, on Carboniferous 
limestone. 
(3) Cheddar Wood, Somerset: oak-ash wood, on Carboniferous lime- 
stone. 


(4) Epping Forest, Essex: beechwood on Tertiary sands. 
(5) Esher Common, Surrey: oak-birch wood on Tertiary clays. 


In no locality have general observations contradicted the hypothesis that 
bracken tends to be less fertile in shaded situations. 


DETAILED OBSERVATIONS 
Localities studied 
Three areas were studied in detail and were deliberately chosen to 
contrast their geology, topography and climate. The bracken growing in a 
completely exposed situation on Braunton Burrows, North Devon, was 
examined as an example of a fairly xerophytic habitat, subject to a climate 
with considerable maritime influence. The edge of a mixed coniferous wood 
was studied in Ashton Court, Bristol, which was situated on a sandstone in 
the Carboniferous Limestone series. The area studied was on a 10—15° slope 
facing S.S.E., at an altitude of about 300 feet. The third area studied was 
the edge of a pine plantation on the North Downs, near Shere, Surrey, and 
was more or less level. The climate in this area is markedly drier, and with 
more extreme temperatures, than the other areas in south-west England. 
The geology was the clay-with-flints of the chalk plateaux, and the altitude 
was about 600 feet. 


224 BRITISH FERN GAZEI1TE: VOLUME 9 PART 6 (1965) 


Results 

The measurements of fertility in the three areas (Fic. 1) demonstrate 
the degree of response—a reduction to less than 5% of full fertility in dense 
shade—and also demonstrate that the effect is graded according to qualita- 
tive estimates of the degree of shading. 

The numbers 1 to 6 for the Ashton Court and Shere data represent 
strips about three yards wide and six yards apart, which were drawn at 
right—angles to the transect line, but parallel to the edge of the wood. All 
plant and habitat measurements were taken at random within each strip. 
As the population at Braunton was all fully exposed, samples were taken at 
random throughout the area. 

A reliable and convenient measure of the frond fertility was worked 
out with some difficulty. Initial observations soon established that fronds 
could not be simply classified as either fertile or sterile, but the degree of 
fertility of the basal pinna was found to be a reliable guide to the fertility 
of the whole frond. The development of a marginal indusium was taken to 
indicate potential fertility, and the number of the basal ten segments of a 
standard pinnule which showed such development was counted. This gave 
a value of 0 to 10, which could be rapidly transformed into percentage 
fertility. The standard pinnule used was the second lower pinnule of the 
bottom left-hand pinna. 

Having established that bracken shows a gradual decrease in fertility 
when growing in increasingly dense shade, an attempt was made to elucidate 
the factors controlling this effect, whether acting directly, or through the 
restriction of vegetative development. The factors considered in detail were 
the degree of vegetative development, edaphic factors, humidity and light 
intensity. 

Three measures of vegetative development were used, none of which 
were entirely satisfactory. The length of the standard pinnule was measured, 
but this often gave a poor estimation of the size of the frond as a whole, 
and certainly showed no correlation with the percentage fertility. The 
average height of the fronds and the number of fronds growing within a 
circle of one metre diameter, were also measured in each strip. The results 
are shown in Fic 2. It is clear that in both the Ashton Court and Shere areas 
the average frond height shows a positive correlation with fertility although 
individual fronds growing in deep shade occasionally reached heights of six 
feet or more. This reduction in the average height of the shaded fronds was 
associated with a change in morphology. In deeply shaded fronds the angle 
between the basal pinnae at the point where they join the rhachis was obtuse, 
usually between 135° and 180°, whereas in exposed fronds the angle was 
more acute, between 60° and 120°. Thus the pinnae of shaded fronds 
appeared to lie all in one plane, often at about 45° to the horizontal, whereas 
only the upper pinnae of exposed fronds lie in one plane, and the lower are 
borne on an erect rhachis. In addition the shaded fronds are a much paler 


idium —- in relation 


@ frond 


DRING: SHADE AND BRACKEN FERTILITY 
wl DEEP AK DEEP 
$ OPEN OPEN ——————? suape OPEN ae OPEN ————> 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 


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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 


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a 


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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: 


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‘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 


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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’, “<Crispa’, ’‘Ramo-cristata’. : : 

3rd N. H. Almond: ‘Drummondia ae’, ‘Ramo-cristata’, ‘Ramo-cristata 
( . 


a 
- 


oe 11 Three distinct varieties of Polypodium vulgare 
Messrs. Brookfield: ‘Cristatum’, ‘Oxford’, ‘Pulcher 
dad B. Hayhurst: ‘Grandiceps Cowan’, ‘Hadwinii’ gy ptr 
meee 12 fg distinct varieties of Polystichum: b 
. Hayhurst: P. setiferum ‘Divisilobum Densum’, P.s. ‘Divisilobum 
gf eine liso , P.s. ‘Imbricatum Cristatum 
2nd N. Robinson: P. aculeatum Sherr ny: mum Bevis’, P. setiferum 
‘Divisilobum ae te Jones’, Divi silobum pitied se 


eet 13 wis indices varieties of cea filix-fem 
B. Hayhurst: ‘Cruciatum Crist tatum’, picsasatene ‘Victoriae’. 
ad N. H. Almond: aa atum’, ‘Frizelliae Capitatum’, ‘Victoriae’. 
3rd_—sSON.. Robinson: ‘Cruciatum Angustatum’, ‘Gemmatum’, Vie 
CLASS 14 Three distinct varieties or species of Dryopteris (including Gym 
carpium and Neca abi all exhibits, except Brookfield’s eee 
fern, were Dryop ‘ soa? 
Ist B Hayhurst: D. Bie ‘Angustata Cristata’, D. dilatata Succisa , 
. filix-mas ‘Depauperata Cristata : ‘ndale’ 
2nd *,, Robinson: D. filix-mas ‘Lineata Cristata’, D.f. ‘Martindale, 


3rd —s Messrs Brookfield: D.borreri ‘The King’, D.filix-mas ‘Cristata’, 
Thelypteris palustris. 
—o 15 Three distinct varieties or species of Aspleniu ho- 
— — Brookfield: A. adiantum-nigrum, A. reniudesiebl A. trie 


pends 16 Thiee distinct normal species of hardy British ferns hice 
sshebin nson: Blechnum spicant, Poly podium salaare, Polystichut 
ah 
2nd Mess. “Brookfi field : aie aa capillus-veneris, Blechnum spicant. 
Gymnocarpium dryop 


FERNS AT SOUTHPORT FLOWER SHOW 1965 297 


seer 17 One se baal fern of any kind or variety 
. Hay hehe th tide baie abd ‘Gracillimum’, 
and Messrs Brookfe Id: Plumosum Superbum Druery’. 
3rd N. H. Almond: A af “Grist: 
4th J. Pye: Af. “Vict 
sa 18 mag Artem dy areenhouse ferns 
ayhurst: mtum ‘Kensington Gem’, Adiantum cuneatum, 
jen repair — atodes abeconeRe Nephrolepis 


2nd Messrs Brookfield: Adi antum “Kensin ington Gem’, Davallia cana- 
riensis, Nephrolepis ‘Cristata’, Nephrolepis cexaltata, Platycerium 
alcicorne, Polypodium ashes var. areolatu 
ce 19 Three dissimilar greenhouse ferns: 
Messrs. Brookfield: Adiantum ‘Kensington Gem’, Davallia 
canariensis, Nephrolepis exaltata. 
2nd B. Hayhurst : Adiantum ‘Kensington Gem’, Davallia bullata, 
Nephrolepis ‘Rosea’. 
eg = 20 fog greenhous m: 
ayhurst: Aight ‘Kensington Gem’. 
nd M. Plowman: Adiantum ‘Fritz Luthi’. 
3rd ~—sW.. W.. Howarth: Adiantae: ‘Pragtitis® 
27 Yewlands Drive, Fulwood, Lancs N. ROBINSON 


A VISIT TO FERNDENE, KENDAL 
During a Northern Weekend Meeting of the British Pteridological 
Society in October 1965 I suggested a visit to the former garden of 
Joseph Wiper* to see what remained of his ferns. The first meeting 
convened amongst the Lakeland fern enthusiasts to form our Society 
was held in Joseph Wiper’s shop. He had a magnificent collection of 
ferns in those days but sold many of them from time to time, and 
finally emigrated to Canada in 1911. his garden being taken over by 
Mr Monkhouse a bulldog breeder but also interested in the ferns. 
Joseph took specimens of his ferns to Canada and soon established 
a collection in Victoria B.C. After Joseph’s death his ferns were taken 
to Vancouver by his son Ernest*; later, his grandson Kenneth G. Wiper 
(son of George who died i in June 1964) formed another fern garden on 
Vissbnioet Island assisted by spores and plants from the old collection. 
The Wiper gardens created epicentres of the fern cult visited by fern 
growers from all parts, particularly Canada and U.S.A. 


* JOSEPH WIPER born Kendal 25 September 1848 died — B.C. 


Asis American Fern 

Sere sero fa member 1891 of the British Preridological Society. Fern 
finds: Athyrium filix-femina ‘Caudigerum’, Thelypteris limbosperma ‘Fili- 
fera’ and ‘Gracilis’. Obituary in Brit. Fern Gaz. 6:33 (1930). Portrait in two 
group photographs in Brit. Pterid. Papers Meeting 

* ERNEST WIPER born Kendal 26 September 1879 died Vancouver B.C. 
18 March 1964 leaving two sons (Joe and Theo). Member 1909 of the Satish 
Pteridological ety. 


298 BRITISH FERN GAZETTE: VOLUME 9 PART 7 (1966) 


_The Wiper family, who made Kendal Mint Cake a famous con- 
fection still in great demand by mountaineers as a concentrated food, 
started another wholesale and retail manufacturing confectionery 
business in Victoria and when Joseph died transferred to Vancouver 
where it is still carried on by two grandsons Joe and Theo (sons of 
Ernest).—I well remember meeting Ernest when he visited South- 
port Flower Show many years ago; he and his wife used to make many 
collecting trips into the Victoria mountains, and on three occasions 
lost their bearings and had to spend the night couched on cedar boughs 
among sheltering rocks till daylight made progress safe. 

The garden at Ferndene (formerly Fern Deane) has changed hands 
many times. I visited it ten years ago and saw many fine things still 
happy there, but in the meantime some landscape work has been done, 
a new garage introduced, and some of the old fern beds have gone. 
Mr Ingham the present owner was very interested in hearing about 
the history of the garden and although he knew little about the ferns 
is going to preserve what are there and inform me if he considers 
leaving, so that provision may be made to ensure the safety of the 
remaining specimens. The first plants just within the gateway were 
several Dryopteris filix-mas ‘Barnesii’ and we noticed several self-sown 
seedlings established in the wall. Several good forms of Polystichum 
setiferum ‘Divisilobum’ were the next finds, growing more or less 
happily below some bushes. Towards the house the sloping ground 
had been terraced with limestone dry walls and in these were sever 
varieties of crested athyriums, scme good, some obviously self-sown 
and of little account. Large clumps of Dryopteris filix-mas ‘Grandiceps 
Wills’ were noticed here. A large bush of Dryopteris filix-mas “Bol- 
landiae’ showing good autumn colour was fighting it out with ivy and 
tree roots in one corner. Dryopteris borreri ‘Crispa’ in large clumps 
was growing at the base of another wall. A stream trickling through 
an old rock-garden was lined with many ferns, a large clump of crest 
royal fern, more ‘Bollandiae’, and many polystichums were flourish- 
ing. A seedling on the lines of Polystichum setiferum ‘Cristatum Wol- 
laston’ but with ovate pinnules was rather striking and at the owner's 
Suggestion was taken by Fred Jackson for safe keeping. Polystichum 
aculeatum ‘Acutilobum’ was represented by two or three large clumps; 
and some rather good plumose athyriums also were noted. The most 
striking find was a large mass of Pteridium aquilinum ‘Cristatum . 
several square yards in extent, a very good form indeed with every 
pinna terminating in a fine crest, perfectly symmetrical. Plants and 
spores were given to Fred Jackson and myself. 

There are many gardens in the district where the old fern men 
grew their treasures, but on the deaths of their owners these collections 
were dispersed by gift or sale and most of the plants lost. It was 4 


KAYE: A VISIT TO FERNDENE, KENDAL 299 


great pleasure therefore to find just a part of Joseph Wiper’s original 
collection still surviving and even thriving, sixty years after he had 
left it in other hands. There must be many plants still to be found in 
other gardens, whose present owners have acquired them with the 
houses of former fern enthusiasts long passed away, plants which 
may be the last examples of varieties believed lost to cultivation. 

Waithmam Nurseries, Silverdale, Lancs. REGINALD KAYE 


REVIEW 


SPORES - FERNS, MICROSCOPIC ILLUSIONS ANALYZED 
Volume One by Clara §. Hires; xxiv + 548 pages, 1150 illustrations, 
8} x 11 inches; Millburn, New Jersey: Mistaire Laboratories, June 
1965; price $22.50. 


This volume, entitled VARIED PLANT GROUPS INT RODUCE 
TRUE FERNS THROUGH SPORE TETRAD STRUCTURE, is the 
first of “future volumes restricted to ferns” to prove the value of three- 
dimensional spore study, and designed to help a diversity of professions 
from artist to zoologist ranging through botanist, criminologist, doctor, 
evolutionist, food analyst, etc. 

he 1150 illustrations comprise about 700 drawings, 100 photo- 
graphs, 336 photomicrographs in black-and-white and 14 photomi- 
crographs in colour. The first quarter of the book is concerned with 
understanding the architecture of spores, emphasizing their solid geo- 
metry by means of such aids as three-dimensional models and 
microscopy. Scattered through the book are various reference guides 
(such as lists of persons, books, terms, etc.), various technical guides 
(concerning mounting media, mathematics, photomicrography, etc.), 
and historical information about the author and the 38-year-old Mis- 
taire Laboratories. The main part of the volume presents a collection 
of photomicrographs of spores which, apart from accompanying 
explanatory text, speak for themselves and form a valuable source of 
reference. 

However, there are three things which would have greatly 
enhanced the usefulness of this book ; (1) an alphabetical index to 
figures of plants, giving page references to each’ species and to parts 
(frond, indusium, spore, etc.), placed at the end (not as “Contents 
On page iv or “Mistaire ferns . . . used for research” on page 477, 
which I found elusive and exasperating), (2) a check on the authority 
of the plant names, by giving the source of each picture which should 
preferably have been taken from a herbarium specimen available for 
reference, (3) a better selection of pictures displayed uniformly for 
easier comparison, particularly as there is a varied amount of material 
presented, e.g. Dryopteris marginalis occupies thirty-two pages an 
Polypodium vulgare three. 

Nevertheless the author is to be congratulated for a valuable 


300 BRITISH FERN GAZETTE: VOLUME 9 PART 7 (1966) 


contribution beautifully set with very clear type and illustrations. Her 
enthusiasm was further shown when she issued with this first volume 
two Mistaire Awards of one-hundred dollars each, one “‘to the person 
contributing the most helpful, constructive criticism of my basic spore 
concepts’’, and one “‘to the first person who can honestly prove . . . 
that my basic spore concept . . . is incorrect’. In 1965 she published 
a Supplement consisting of a summary of her basic theories and a 
reprinting of ten plates “to provide better orientation and larger 
backgrounds to insure clearer outlines.’’ We look forward with great 
interest to the promised further volumes. 

British Museum (Natural History), London SW7 JAMES A. CRABBE 


REVIEW 


LEARN OF FERNS WE GROW by Sylvia B. Leatherman and 
Dorothy S. Behrends, [xii] + 166 pages, 15 x 22cm, 35 line-drawings 
by Mrs Behrends; Los Angeles, published by the authors, May 1965; 
price $3.85 ($4 postage paid) from B & L Books Dept. A, 2637 North 
Lee Avenue, South El Monte, California 91733. 


The authors of this attractively designed stencil-produced ring-bound 
book are well known in Californian and other h 


gia, not spores; the gametophyte is described ‘“‘simply and briefly” (pP- 
112, 129, etc.), but the result is confusing and sometimes misleading, 


swim” (p.130); a cultivar is any assemblage of cultivated individuals 
which propagates true-to-type and is not necessarily ‘“‘a hybrid” (p.132). 
‘ y review copy is dated ‘First printing, May 1965” f 
this first edition will soon be revised and the botanical inaccuracies 
corrected. Some system of quicker reference to the many useful an 
interesting points would be a blessing. The text in capital letters 1s not 
acceptable to everyone, but the authors presumably had a reason fOr 
using this unusual typography. 
British Museum (Natural History), London SW7 Auk. JERMY 


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 
REGINALD KAYE LTD 
SILVERDALE, LANCASHIRE 
CATALOGUE ON REQUEST 


BOOK NOW AVAILABLE 
VOLUME ONE 
SPORES - FERNS - MICROSCOPIC 
ILLUSIONS ANALYZED 
by CLARA S. HIRES 
580 pages, approx. 1150 illustrations, 14 in colour 
price $22.50 from 


MISTAIRE LABORATORIES, 152 Glen Ave, 
Millburn, New Jersey 07041, U.S.A. 


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 7 1966 
Conterits 
Psilotum nudum in Europe B. M. ALLEN _ 249 
Ferns in a wardian case G. GILBERT 251 


The identification of British Hymenophyllum species 
G. B. EVANS 256 
Dryopteris villarii in Britain O. L. GILBERT 263 
Origin of the fern flora of Tristan da Cunha 
A. F. TRYON 269 

Some recent work in Wales on the Polypodium vulgare 
aggregate P. M. BENOIT 

Some observations on Polypodium australe 

R. H. ROBERTS 283 

The British Pteridological Society’s Austrian Excursion 
J. W. DYCE 


277 


288 
Fern classes at Southport Flower Show 1965 : 
Judges’ report R. KAYE 297 
Prizewinners and their entries N. ROBINSON 296 
A visit to Ferndene, Kendal R. KAYE 295 
Book reviews : 
HAMLIN, Nature in New Zealand: Native Ferns i 250 


TRYON, The Ferns of Peru: Polypodiacae (Denn- 

staedtieae to Oleandreae) 287 

HIRES, Spores . Ferns, Microscopic Illusions analyzed 

LEATHERMAN & BEHRENDS, Learn of Ferns we grow 
Request for living material: Equisetum 


Bess 


E. C. Freeman Ltd., Glenny’s Acre, London, S.W.11 


toe 
BRITISH FERN 
GAZETTE 


VOLUME NINE PART EIGHT | 


1967 


Missour! BoTanic ~ 


SEP 2 1967 


ROEN WIBRARY 


THE BRITISH 
Od PTERIDOLOGICAL SOCIETY 


BRITISH PTERIDOLOGICAL SOCIETY 


Officers and Committee 1966-67 


PRESIDENT 
J. Davidson MB ChB FRCP (Edinburgh) 


VICE-PRESIDENTS 
Thomas H. Bolton 
Percy Greenfield 
R. E. Holttum MA ScD FLS 
Reginald Kaye 
Professor Irene Manton PhD FRS 
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 
(Joint Meetings Secretaries) 
R. F. Cartwright P. Temple LRIBA MInstRA 
143 Pittmansfield 42 Holly Park 
Harlow, Essex London N3 


T. A. Dyer FLS F. J. Healey F. Jackson 
Frances M. Jarrett MA PhD FLS J. D. Lovis BSc PhD 
R.S.W. Pollard LAMTPI _N. Robinson W. Wright 


AUDITOR 
Peter Temple LRIBA MiInstRA 


The SOCIETY is open to ll interested in any way in FERNS and FERN ALLIES 
pon paying i of TWENTY-FIVE SHILLINGS. Full details 
ae wll be sent upon aplication t the SECRETARY. 


BRIT. FERN GAZ. 9 (8) 1967 301 
FERN HYBRIDISTS AND FERN HYBRIDISING 
I. THE WORK OF EDWARD JOSEPH LOWE 

(1825-1900) 


JOHN D. LOVIS 
Department of Botany, Leeds University 


Both this paper and its companion owe their origin not to an inspiration 
of my own, but instead to a suggestion from Mr J. A. Crabbe, who 

informed me that as a result of discussion on the subject of fern hybrids 
at the Brantwood meeting of the Society, it was clear to him that an 
ee of the methods used by the Leeds atte in Lath ions ferns 

be welcomed by age of the Gaz rther 

assessment of the work “Of E. J. LOWE, as decctibed | in his book * Te m 
Growing ” (1895),* and in dectiodiny of his theory of “ multiple 
sitentnge™ , would also be of interest. It soon esas evident to me 
that it would be difficult to combine these two topics a 
within one article, with the consequence that in the end I have not 
attempted to do so, but intend to treat them separately. This article. 
a is solely concerned with the work of Edward Lowe, is presented 


I must confess at the outset that when this project was first suggested 
to me, I was totally ignorant of this particular book, * Fern Gro wing” 
by Lowe, and in writing this article I have a assumed that at least some 
of my readers will be as ill-informed in this respect as I was myself. 
I expect that given the opportunity most readers of the Gazette would 

owe’s account of his own work as interesting as I have done, but 
unfortunately this book seems to be rather rare, and indeed is described 
In the trade as ‘ scarce’. No copy was available in Leeds, and Ia 
very grateful indeed to Mr J. W. Dyce for the protracted and weneious 
loan of his own personal copy, without which I could not have made 
my analysis 

Toa very great extent, this book consists of a summary of Lowe’s 
own work as a fern hybridist. A considerable proportion of the first 
part of the book takes the form of direct quotations from Lowe’s 


deel will occ cssieealiy revea 
obviously 


*All page references given in the text of this article refer to this publication. 
The BRITISH FERN GAZETTE Volume 9 Part 7 was published on 24 June 1966 


302 BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967) 


Neto re of an idea (that ferns can cross and et reagnd bts. er 
e today accept without question as undoubtedly the 
cthee hand, latterly he persisted with equal saanlon in ye t of a 
concept, that of multiple parentage, which we today must regard as 
ill- founded, since basic to Lowe’s ideas was his belief that the union 
of one spermatozoid and one ovum is never alone sufficient to produce 
a sporophyte. 
One is surprised and intrigued to learn what difficulty Lowe had 
in ig ee the botanists of his day that indeed it was possible for 
to hybridise. Furthermore, one is astonished to discover that as 
fie as 1884, a paper submitted to Ly Linnean Society describing ee 


of his hybridisation ree nts was rejected, and returned to him 
“not proved ”’ (p. 63), a decision which clearly deeply wounded aa 
permanently offended poor Lowe. ome of us have reason to be 


thankful that the Fellows of the Linnean Society are to day no longer 
so sceptical concerning fern hybrids!). This is all the more ‘criartable 
in view of the relative sophistication of the experiment being r reported, 
dam is indeed of such interest as to merit description in some detail 
her 


"Wain may first note that like all of Lowe’s hybridisation experiments, 
the ovine was attempted by the simple device of making a 


close copies of pee idium aculeatum var. densum, spidium 
angulare var. Wakelyanum; these five hybrids had all the characters of 
Aspidium euleaders | (in the stronger consistency of the pinnae, and 
richer and deeper colour), ba not only this, they were a copy of the 
cruciate yond Wakelyanu 

he successful oeing of two species is clearly a more effective 
demonstration of hybridisation than a cross between two varieties, but 
in this rhe Lowe, as he pont out (p. 13), encountered the difficulty 


it to quote Lowe (p.1 De quite seecmebie for the male organs from one prothallus 
to sere another’’. It does however appear that Lowe did not entirely pre 
pport. ee 


bt Ferns... - 
equally between Lowe and four others (J. M. Barnes, J. E. Mapplebeck, J. Moly, 
aud Stansfield the elder), whom he states had formed “. . . independent conclusions 
... , Nearly about the same time, . . .”’, though no dates or references are quoted. 
I am indebted to Mr Crabbe for drawing my attention to the existence of this article, 
which provides a valuable perspective to the account given by Lowe of the activities 
of his contemporaries. 


LOVIS: FERN HYBRIDISTS L: > 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