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Dyce, 46 Sedley Rise, Loughton, Essex, 1G10 7LT. Vice Presidents R.E. Holttum, F. Jackson, R. Kaye, Irene Manton, S. Walker. General Secretary A.R. Busby, 42 Lewisham Road, Smethwick, Warley, West Midlands. Treasurer Dr. B.A. Thomas, Biological Sciences Department, University of London, Goldsmiths’ College, New Cross, London SE14 6NW. Membership Secretary Lt. Col. P.G. Coke, Robin Hill, Stinchcombe, Dursley, Gloucestershire. Meetings Secretary A.J. Worland, 702 Queens Close, Harston, Cambridgeshire CB2 5QN. Editors: Fern Gazette Dr. C.N. Page, Royal Botanic Garden, Edinburgh EH3 5LR. assisted by J.A. Crabbe, J.W. Grimes, A.C. Jermy. Bulletin M.H. Rickard, 9 Hammond Close, Pampisford, Cambridgeshire CB2 4EP. assisted by J.W. Dyce. British Fern Distribution Recorder A.R. Busby, Dept. of Biological Sciences, University of Aston, Gosta Green, Birmingham B4 7ET. Spore Exchange Organiser R.F. Cartwright, 73 Perry Mill Road, Peopleton, Pershore, Worcestershire. Committee P.G. Coke, J.A. Crabbe, Barbara S. Croxall, Mary Gibby, J.W. Grimes, N.A. Hall, A.C. Jermy, H.L. Schollick, F.J. Tingey, S.L. Williams. The Fern Gazette and the Bulletin are the journals of the British Pteridological Society, published annually. The Gazette publishes matter chiefly of specialist interest on international pteridology, the Bulletin topics of more general appeal. [See inside back cover for notes to contributors.] The BRITISH PTERIDOLOGICAL SOCIETY was founded in 1891 and today continues as a focus for fern enthusiasts. It provides a wide range of information about ferns through the medium of its publications and available literature. It also organises formal talks, informal. discussions, field meetings, garden visits, searches for rare fern books, plant exchanges and a spore-exchange scheme. The Society has a wide membership which includes gardeners, nurserymen and botanists, both amateur and professional, in Britain and overseas. Articles for the Society's publications are received from, and reflect the interests of, Society members. Membership is open to all interested in ferns and fern-allies, worldwide. Subscription rates (due on 1st January each year) are: Full Personal Members £4; Personal members not receiving the Fern Gazette £3; Student Members (full-time students under the age of 25) £3; Subscribing Institutions £5. Applications for membership should be sent to the Membership Secretary (address above), from whom further details can be obtained. (Remittances made in currencies other than Sterling are £0.50 extra, to cover bank conversion charges.) Back numbers of the Gazette and Bulletin are available for purchase from the General Secretary (address above), from whom further details can be obtained. FERN GAZ. 12(1) 1979 THE KILLARNEY FERN, TRICHOMANES SPECIOSUM, IN WALES R-H. ROBERTS 51 Belmont Road, Bangor, Gwynedd. ABSTRACT The history of botanical knowledge of the Killarney Fern (Trichomanes speciosum Willd.) in Wales is discussed. The first record of the Killarney Fern (7richomanes speciosum Willd.) in Wales appeared in 1863 when A.M. Gibson drew attention to a note in a Norwich newspaper that the fern had been found in North Wales by J.F. Rowbotham of Manchester (Gibson, 1863). Rowbotham also reported his discovery to Thomas Moore and his description of the habitat is worth repeating. “| found it’, he wrote, “in a large hole formed by fallen rocks alongside a cascade of water; and admission to this hole, which is about five feet high by four feet wide, is obstructed after a depth of about three feet by this Fern falling from the rocks at the top and growing out of the sides in the form of a beautiful curtain, down which the water is constantly trickling’ (Moore, 1863). He also added that the fronds were remarkably fine and abundant. The frond he sent to Moore had an overall length of nearly eighteen inches and was seven inches across at the widest part. A second frond in Rowbotham’s possession had a total length of about twenty-two inches. Moore does not disclose the locality beyond stating that it was in a part of the Snowdon range. The following year James Backhouse Jnr. sent specimens of this fern to the Botanical Society of Edinburgh. In the covering letter he stated that they had been found by him and his father, in 1863, “in a truly wild state, in Carnarvonshire,” where, he added, the plant was remarkably luxuriant (Backhouse, 1864). His specimens both at Edinburgh and the British Museum (Natural History) are equal to the finest Irish specimens and it is probable that they were from the same locality as Rowbotham’s, although Backhouse did not refer to Rowbotham’s discovery either then or subsequently. Five years later Backhouse found the Killarney fern in Merioneth, v.-c. 48. His specimen in the British Museum is vaguely localised ‘Mountains of Merionethshire”’ and dated 1869. Rather uncharacteristically, this find was not published by Backhouse, but in his next letter to the Botanical Society of Edinburgh, in 1875, he announced that he had “collected 7Trichomanes radicans in a new station in North Wales in 1873’ (Backhouse, 1876). Again he gives no locality — not even the name of the county — but it is probable that this was also in Merioneth, for Backhouse is stated to have seen the Killarney Fern in two widely separated localities in the county in 1874: one near Harlech and the other on Cader Idris, and it was flourishing at both of them twenty years later (Percival, 1894). It is almost certain that it was at one of these localities (near Harlech) that Backhouse showed the Killarney Fern to D.A. Jones, the well-known bryologist, who lived in Harlech (Jones, 1898), and to Mrs. Mary Richards of Dolgellau (Benoit & Richards, 1963). Although frequently visited, and no doubt too frequently collected, the fern survived in this site until 1968, but there is now no trace of it left (Dyce, 1975). These Merioneth localities became known to a number of fern enthusiasts. Specimens were collected from one of them in 1889 and sent to the British Museum by William Robinson, of Weston-super-mare, who claimed to have known the Killarney FERN GAZETTE: VOLUME 12 PART 1 (1979) Fern there and in “other localities” (all unspecified) for about twenty years (Britten, 1894). D.A. Jones, who knew the county well, gives only two localities for the Killarney Fern in his manuscript ‘Flora’ and these are the same as those mentioned by Percival (1894). The statement that the Killarney Fern had ‘been found growing luxuriantly in some abundance in several places extending over several miles in Wales” (Lowe, 1876) seems to be somewhat exaggerated. In 1887 the Killarney Fern was found independently in Caernarvonshire, v.-c. 49, by J. Lloyd Williams (1887), who was at the time a schoolmaster at the village of Garn Dolbenmaen, near Cricieth. However, he could not be certain that this was not the same locality as that in which the fern had been found previously, because that record had not been localised. Apart from describing the habitat as ‘‘’a damp hole near the top of a range of mountains” and that it was not on any part of Snowdon, Williams gave no indication of the locality. In his autobiography written many years later he gives a fascinating account of how he found the fern and indicates that the locality was Moel Hebog, near Beddgelert (Williams, 1945). There are, however some curious inconsistencies in Williams's account. According to this the fern was completely removed about a month after he first found it, and even though he made several visits to the site from 1887 until 1893 (when he left the district), he could find no trace of it. Twenty-four years later he visited the place again and was surprised to find the Killarney Fern once more flourishing in its rocky recess. Williams ascribes its recovery to the spores left after the fern itself had been completely removed. However, J.E. Griffith, of Bangor, author of the ‘Flora of Anglesey and Carnarvonshire’ (1895), gathered a specimen of the Killarney Fern from this locality in 1891 (Hyde, Wade & Harrison, 1969). In his ‘Flora’, published a few years later, Griffith wrote, | have seen this fern growing undoubtedly wild, in one place only. This was first found by Mr. J. LI. Williams. | refrain from giving the locality as it is so rare.”’ One can only surmise that Williams showed the locality to Griffith, although the latter does not say so. But it seems that he passed through Garn Dolbenmaen, where Williams lived, on his way to Moel Hebog, for he published records of plants which he saw in both these places in August 1891 (Bennett, 1892). It seems that Williams was mistaken when he wrote that the Killarney Fern had been destroyed in his locality before the end of the year 1887. But he was in his late eighties when he wrote this account and perhaps relied too much on his recollection of the events of over fifty years before. However, there is nothing to indicate whether Williams’s locality on Moel Hebog was the same as those found by Rowbotham and Backhouse. On the contrary, Rowbotham’s description can hardly refer to the Moel Hebog habitat and strongly suggests a different place, possibly one of the two others in which the Killarney Fern was stated to occur in Caernarvonshire. Professor J. Bretland Farmer (1948) claimed that he had known the Killarney Fern in the Cwm Glas area of Snowdon for over twenty years. Neither this locality nor another, which according to rumour is vaguely localised on the Carneddau, has ever been confirmed. Unfortunately, Farmer did not leave a voucher specimen, but it seems hardly credible that he could have mistaken any other fern for it. Nevertheless his record has remained unconfirmed for fifty years and is now regarded with a good deal of scepticism. Even when it was first recorded in 1863 there were rumours that the Killarney Fern had been deliberately introduced into Caernarvonshire by a local guide, but Moore (1863) thought that the luxuriant and well-established appearance of the fern found by Rowbotham was not consistent with such a view. Trimen (1871), on the other hand, was of the opinion that there was ‘reason to suspect an intentional ROBERTS: KILLARNEY FERN IN WALES 3 introduction.” Britten (1879-81) also refers to these rumours, but goes on to make the intriguing statement that the Killarney Fern was already known in two localities in Caernarvonshire about thirty years before its discovery by Rowbotham. The discoverers, he adds, were satisfied that the fern was truly native, but had kept their knowledge to themselves. Unfortunately there is more direct evidence that some planting did take place. In a tribute to the recently deceased Herbert Stansfield, E.H. Hawkins wrote: “| recall that some years ago he spent some time in North Wales, taking with him some plants of the Killarney Fern, which he planted in some wild and congenial places, with the hope that they may be found subsequently by fern hunters. This thoughtfulness was ever typical of him’’ (Hawkins, 1928). Herbert Stansfield was a nurseryman of the firm of F.W. and H. Stansfield of Sale, near Manchester, who specialised in British ferns and no doubt had access to Irish material of the Killarney Fern. Stansfield was not the only one who tried to introduce the Killarney Fern into suitable places. Its occurrence in Westmorland (where it was gathered in the Rydal district about 1863), in the Isle of Arran, and in Argyll, were all suspected to be due to deliberate introductions (Moore, 1863), as it was in the only Cornish locality (Davey, 1909). Even in Ireland this fern was planted in many places in Killarney, around Glengarriff and on Valentia Island (Newman, 1844), though the purpose there was to try to preserve the fern from the depredations of collectors, which even then threatened to exterminate it in its original localities. We shall probably never know where Stansfield made his introductions into North Wales, or indeed whether any of them were successful. His activities were too late to affect the early records, for he was only seven or eight years old in 1863, but they cast a shadow of doubt over all the Welsh localities of the Killarney Fern. On the other hand Backhouse’s remarkably fine specimens from both Caernarvonshire and Merioneth give the impression of being from old, long-established colonies of the fern — an opinion already expressed by Moore (1863). Rowbotham’s description of the habitat, quoted above, is also very convincing and does not suggest an introduction. This impression is supported, as far as Merioneth is concerned, by William Robinson’s specimen (BM) gathered in 1889, and A.J. Crosfield’s two specimens (BM) gathered in 1902 and 1904, although all these Merioneth gatherings may well be from the same locality, possibly that in the Cader Idris area, now lost, or, if known, wisely kept secret. The much smaller size of specimens (BM) gathered by T.J. Foggitt from ‘‘wet rocks near Harlech, Merioneth, 27 Sept. 1929,” suggests that they are from a different site. The largest of his two fronds is just under four inches long, including the stipe, and just over an inch wide. They are similar in size and shape to D.A. Jones’s specimens (NMW) gathered in 1905 “near Harlech,’’ and to a frond | was once shown which had been gathered from the well-known Merioneth locality where the fern now seems to be extinct. There is still a good deal of mystery attached to the Killarney Fern in Wales. Its discovery as recently as 1961 in Cardiganshire, v.-c. 46, by A. Neville Jones was certainly remarkable, for the colony he found is a luxuriant one, and it is difficult to understood why it remained undiscovered for so longin a comparatively well-botanised area. Perhaps, after all, we can only echo the words of Scully (1916) when referring to the Valentia Island records: that the fern is both indigenous and introduced in different localities in Wales. However, this should not affect our attitude to its conservation. Its disappearance from its only known Merioneth locality within the last ten years is most 4 FERN GAZETTE: VOLUME 12 PART 1 (1979) regrettable. Already the Caernarvonshire locality, rediscovered by G.M. Hughes in 1967, has become too well known, as indeed has the Cardiganshire one. Fortunately the Killarney Fern is now protected by the Conservation of Wild Creatures and Wild Plants Act 1975, but it is also to be hoped that a new attitude will prevail among botanists who visit the sites of this most attractive fern and that they will be content to look at and admire it without removing a single frond. ACKNOWLEDGEMENTS | am grateful to Dr. C.N. Page for his assistance in many ways. Thanks are also due to the Keeper of Botany, National Museum of Wales, the Regius Keeper, Royal Botanic Gardens, Edinburgh, the Keeper of Botany, British Museum (Natural History), Mr. J.W. Dyce, Mr. A.C. Jermy and Mr. M. Morris for their help with many of the references. REFERENCES BACKHOUSE, J. 1864. (Note on Trichomanes radicans) Trans. Bot. Soc. Edin. 8: 111. BACKHOUSE, J. 1876. Miscellaneous notices. Trans. Bot. Soc. Edin. 12: Appendix XX. BENNETT, A. 1892. Rep. Bot. Exch. Cl. 7: 326, 340. BENOIT, P.M. & RICHARDS, M. 1963. A contribution to a Flora of Merioneth. Ed. 2. Haverfordwest. BRITTEN, J. 1879-81. European Ferns. London. BRITTEN, J. 1894. Editorial note. J. Bot. Lond. 32: 372. DAVY, F.H. 1909, Flora of Cornwall. Penryn. DYCE, J.W. 1975. Meetings 1975: Barmouth, Gwynedd. Brit. Pterid. Soc. Bulletin, 7 (3). FARMER, J.B. 1948. Notes on the flora of Snowdonia, in CARR, H.R.C. & LISTER, G.A. (eds.) The Mountains of Snowdonia. Ed. 2: 144-155. London. GIBSON, A.M. 1863. Trichomanes radicans in Wales? The Phytologist. 2nd ser. 6: 608. GRIFFITH, J.E. 1895. Flora of Anglesey and Carnarvonshire. Bangor. HAWKINS, E.H. 1928. Tribute to H. Stansfield. Br. Fern Gaz. 5: 220—221. HYDE,H.A., WADE, A.E. & HARRISON, S.G. 1969. Welsh Ferns, Ed. 5. Cardiff. JONES, D.A. 1898. A Handbook to the Botany of Merionethshire: ms. in National Museum of Wales, Cardiff. LOWE, E.J. 1876. Our Native Ferns, 2. London. MOORE, T. 1863. Trichomanes radicans indigenous to Yorkshire and Wales. J. Bot. Lond. 7: 238—239. NEWMAN, E. 1844. History of British Ferns. London. NEWMAN, E. 1864. History of British Ferns. Ed. 4. London. PERCIVAL, J. 1894: Trichomanes radicans in Wales. J. Bot. Lond. 32: 372. SCULLY, R.W. 1916. Flora of County Kerry. Dublin. TRIMEN, H. 1871. Trichomanes radicans in England. J. Bot. Lond. 9: 174—175. WILLIAMS, J. LL. 1877. Trichomanes radicans in Caernarvonshire. J. Bot. Lond. 25: 215. WILLIAMS, J. LL. 1945. Atgofion Tri Chwarter Canrif, 4. London. FERN GAZ. 12(1) 1979 a PRELIMINARY INVESTIGATION OF TWO SOUTH-WEST ENGLAND POPULATIONS OF THE ASPLENIUM ADIANTUM-NIGRUM AGGREGATE AND THE ADDITION OF A. CUNEIFOLIUM TO THE ENGLISH FLORA C.N. PAGE & FRANCES M. BENNELL Royal Botanic Garden, Edinburgh. ABSTRACT Examination of the plants of the Asp/enium adiantum-nigrum aggregate from south Devon and west Cornwall questions the status of records of A. onopteris, but confirms A. cune/folium as new to the English flora. Asplenium onopteris, A. adiantum-nigrum and A. cuneffolium form a trio of inter- related species in the British Isles. All three are morphologically variable. Because A. adiantum-nigrum is an allotetraploid containing the chromosomes of the other two species (Shivas, 1969), its variation may be particularly wide, approaching and perhaps overlapping that of both its parents. Clearly, without the benefit of a chromosome count, there can be difficulty in separating all specimens in areas where more than one may be suspected to be present. The discovery that A. cune/folium can also exist as a tetraploid species in Britain (Roberts & Stirling, 1974; Sleep et al, 1978) and elsewhere (Deschatres et al, 1978) further complicates the problem of its inter-relationships and taxonomic separation. The finding of populations of this plant group in the Bovey Tracey area of South Devon during the British Pteridological Society's field week in the summer of 1978, which appeared to contain plants of clear A. adiantum-nigrum and ones of extremely attenuate form suggesting the presence of A. onopteris stimulated the present investigation to try to distinguish and identify them. The problem was further highlighted by also finding plants in the Lizard area of Cornwall which resembled A. cuneifolium, apparently linking through plants of rather intermediate appearance with plants of A. adiantum-nigrum. The following investigation was initiated to try to resolve the taxonomic problem arising. Bovey Tracey Population These plants occurred as widely-scattered specimens on lightly-shaded earth and rock hedgebanks in the neighbourhood of the village of Bovey Tracey in South Devon. They are very common locally in such habitats, but extraordinarily variable in appearance. Examination showed them to be mostly large-fronded plants. The cutting of the blade of many suggested they were typical A. adiantum-nigrum, but many more showed varying degrees of attenuation, more finely cut and upswept pinna form, through to at least one individual which had such extremely finely cut and attenuate fronds that it appeared very like A. onopteris (Page 11370). Study of the spores showed, however, a surprising uniformity. No specimens showed any significant degree of spore abortion. Compared with those of typical A. adiantum-nigrum spores were very slightly paler in colour, and showed an overall greater size range, with those of the most A. onopteris-like morphology proving only slightly smaller (31-57 wu) than those of plants of more A. adiantum-nigrum-like morphology (37-60 u). By contrast, plants of known A. onopteris from Ireland and the Canary Islands, proved to have distinctly smaller spores (25-43 1) than those of any of the Bovey Tracey plants. The spores thus present a slightly inter-grading picture FERN GAZETTE VOLUME 12 PART 1 (1979) *projdesja} Ajqeqoid si wnijojiauna ‘y ysiujoD ay} }eyi Guljsebbns ‘(p/6L) Buljans 7 siuaqoy Aq puno} jela}ewW Yysi1O9S Ul UOITENPIS 9Ui YIM Ajge4snoAey BIEAWIOD Sy} Ul pue ‘sajjeuus Ajzybijs Asan a4 WiNsjOssaUuNnd “VV jo esoul (4 ‘GOLLL 26eq — aul| peii0p) xOeUUaIS ye jeligqey yueqoue; |jeMuUIOD 3saM = pueju! jeoidA} e@ WOs} Wniblu-tunjuelpe “YY uMoU> yO uoljeyndod e 40 asoui YyIIM (B‘ZHZLL 26d — QuUl| PIJOS) PyezZi7 ay} ul JeIIGeY aa19S-y49O|Q ay WO4, WNIJOJIaUND “VV $O SjUue|d 410} JeY} aIeEdWOD 01 (rl) azis-aiods yo wesboIs!IH “€ FYNOIS O09 OS OV O€ ‘wnsbju -uinjueipe ‘yy pooh Bulsnogqubiau uey} 4ajjews Aj, ybiys AjjuaisisuOd Wwaas spuds} aul} ay} WOd} asOu} 1ng ‘| “Bly ul Si4a7dOUO “vy }O asOoU} Wod4 Ajseajd Way} saysinbuljsip yoea yo abues azis asods sabe) ays “(4 ‘OLELL a6egd — aut pijos) W4OJ-Puos} |a}2eNU|a}}e PuUue PapiAIp-Ajauly jsOWw YIM aUO LWO4J ABSOY? YUM (3 ‘ZOHLL 26ed ‘aul| pai10p) Aboyoydsiouw ayI|-Wwnsbiu-winjueipe ‘Vv SOW $O 9UO WOJ} BsOYU} asedwod oO} ‘UO!}e;ndod Aasei| AsAog ayi woijy sj}uejd jo sajdwes ul (qd) abues azis asods yo wesboisi “f FYNOIS 09 OS OV Of& — ew ee oe ee — — —S r--- *(3noyBnosyy pue ‘auisadA\i6b %OOL ul palunowW ‘uawiIoads yoea WO1} Painseaw saiods OG = azis ajdues) ‘aHuei azis sejiwis pue azis |jews Ajaniejas diayy Buimoys ‘(3 ‘Ely abled — aul| pijos) spueyjs| Aseued ‘esaWOd ee pue (FJ “u's sdijjiud — dul] pa110P) Puejas| ‘4409 ‘OD ‘auAH ybnoq WO4} Si4a7douo Winiuajdsy UMOUy $O sajdwes ul (qd) aBues azis aiods yo wesbolsiH “1, AHNOIS dq Ni azis O09 OS OV O€ proc occ ee eK - - - -- --------- S3YOdS 4O YAEIWNN Oc PAGE & BENNELL: ASPLENIUM ADIANTUM-NIGRUM AGGREGATE 7 corresponding with the overall morphology of the plants, but do not suggest, from their size, that A. onopteris is present in this south Devon population, at least as this species is recognised in Ireland and the Canary Islands. K ynance Population This consists of a large population of plants of the Asp/enium adiantum-nigrum group in the region of Kynance Cove on the west side of the Lizard Peninsula of West Cornwall. The area is of particular interest not only in being one of the most southerly and climatically oceanic points in the British Isles, but also in containing extensive outcrops of ultrabasic serpentine rock. A preliminary collection of fronds from the area was made by C. Ferreira during the winter of 1977-78, and a more extensive investigation of specimens from different adjacent habitats after examination of the Ferreira material made by one of us (C.N.P.) in the summer of 1978. Plants were examined in three distinct habitats: (a) in a large, natural, block-scree slope descending steeply from the cliff tops directly into the sea to the west of Kynance Cove (b) amongst small outcropping rock bluffs amongst the plateau-like downs running back from the tops of Kynance Cliffs to the north of Kynance Cove (c) in a recent man-made scree amongst disused soapstone quarry workings about one mile north-west of Kynance Cove. Plants from the whole area gave a strong impression of distinctness from those of inland stations for A. adiantum-nigrum in West Cornwall. Very many from habitats (b) and (c) differed from A. adiantum-nigrum in having fronds with fan-shaped segments, whilst in habitat (a), the plants appeared totally distinctive from A. adiantum-nigrum and to indeed be A. cune/folium. Examination of frond samples from habitat (a) showed them to differ significantly from A. adiantum-nigrum in the following features: 1. Overall fronds much less finely cut, with a broader triangular outline and coarser segments varying from slightly more to much more fan-shaped. 2. Ultimate serration of the frond margins less acute (but rarely actually obtuse). 3. Stipe much shorter, usually less than % to % the length of the blade, sometimes less. 4. The deep red-brown colouration more restricted to the lower part of the stipe, extending only rarely as high as the lower pinnae on the underside and rarely as much as half way up the stipe on the upper side, the remainder of the stipe and the whole of the rachis remaining green. 5. The lowermost pinnae pair much more broadly triangular and held more perpendicularly to the rachis, and not at all swept upwards on the frond. 6. A much stronger tendency for the veins on the frond undersides near to the margins to show as distinct whitish radiating lines (especially in the lowermost pinna pair). Plants from scattered rock outcrops (habitat ‘b’) on the top of the Lizard Downs proved much more variable in morphology, although many of them were stunted by varying degrees of exposure. Nevertheless, some approached A. cune/fo/lium in their morphology, whilst others appeared more close to A. adiantum-nigrum, with others apparently linking between these extremes. Examination of spores, however, showed none to be wholly abortive, as far as could be judged by light microscope examination, although many contained a proportion (estimated at up to 10%) which were sufficiently mis-shapen to regard as probably abortive. 8 FERN GAZETTE: VOLUME 12 PART 1 (1979) Plants from the disused soapstone quarry (habitat ‘c’) proved equally confusing. The habitat was a man-made rough loose boulder slope. Asp/enium plants were present mainly in sheltered pockets between the boulders, and consequently were rather larger than those on the tops of the downs. They were, however, of equally variable appearance, a few seeming more like A. adiantum-nigrum, but most more clearly A. cune/folium, but linked by apparent morphological intermediates. As with those on the Downs, spores appeared mostly good but with up to about 10% mis-shapen ones in many of the plants. One well grown individual which in morphology showed a particularly intermediate appearance, probably had a higher number of abortive spores than this. The morphology of the fronds in all of the serpentine rock habitats of the Lizard thus suggests that these are all far from typical populations of A. adiantum-nigrum, and that each of these habitats (‘a’-’c’) thus contains at least some plants of A. cuneifolium, thus adding this species as new to the English flora. These A. cune/folium plants match very closely indeed with those from the Scottish serpentine localities, and provide further evidence in support of the view (A. McG Stirling, personal communication) that these British plants are morphologically distinctive from at least those of central Europe. The close similarity of the spore size to that of the Scottish ones, suggests that these Cornwall plants are likely to prove tetraploid (although this needs cytological confirmation). In these Lizard habitats, A. cune/folium appears to be the dominant fern in the natural block scree slope, from where it spreads on to the outcropping boulders of the Downs and into the man-made serpentine scree of the soapstone quarry. Some questions remain about the identity of the plants of most A. adiantum- nigrum-like morphology in the latter two habitats, but the lack of obvious hybrids (at least as indicated through lack of total spore abortion) and the complete range of forms intermediate in overall morphology, suggests too that these are probably extreme forms of A. cune/folium, underlining the possibility that these can approach A. adiantum-nigrum very closely indeed. ACKNOWLEDGEMENTS It is a pleasure to acknowledge the guidance and stimulating discussion in the field of Messrs. J.W. Dyce and M.H. Rickard in south Devon and Miss Rosaline Murphy and Major E.W.M. Magor in West Cornwall. REFERENCES DESCHATRES, R., SCHNELLER, J.J., & REICHSTEIN, T. 1978. A tetraploid cytotype of Asplenium cuneifolium Viv. in Corsica. Fern Gaz. 11: 343—344. ROBERTS, R.H., & STIRLING, A. McG. 1974. Asplenium cuneifolium Viv. in Scotland. Fern Gaz. 11: 7—14. SHIVAS, M.G. 1969. A cytotaxonomic survey of the Asplenium adiantum-nigrum complex. Br. Fern Gaz. 10: 68—80. . SLEEP, A., ROBERTS, R.H., SOUTER, J.l., & STIRLING, A. McG. 1978. Further investigations on Asplenium cuneifolium in the British Isles. Fern Gaz. 17: 345—248. FERN GAZ. 12(1) 1979 9 AN ECOLOGICAL SURVEY OF THE FERNS OF THE BURREN, CO. CLARE, EIRE A. WILLMOT Derby Lonsdale College of Higher Education, Derby. ABSTRACT A systematic list of the ferns of the Burren (area includes parts of Co. Clare and South East Galway v.-c. H9 and H15) is given based mainly on records made in 1978 by the author. This includes notes on the distribution, abundance and habitats of the taxa. The list includes 35 species and 5 hybrids as recently recorded. The records for Polypodium x mantoniae and Dryopteris x tavelii are first county records for Co. Clare. The distribution of the commoner 24 species is shown by maps. Species lists are given for the more interesting or characteristic fern habitats. The phytogeograph- ical affinity of the flora is discussed on a European basis and is shown to be basically southern with a lesser western (atlantic) affinity. INTRODUCTION The Burren is the most impressive area of karst scenery in the British Isles. It is situated in the middle of the western coast of Eire, on the southern side of Galway Bay. The bed-rock over most of the area is Carboniferous Limestone, which has been stripped bare of soil by past glaciations. From a distance the general aspect of the area therefore is of a bare, grey, uninviting rock but closer examination reveals that this area is botanically one of the most interesting and beautiful of western Europe. Burren Hills Black Head OG + bboy i/ Burren Lowlands =) - rate > Ce —_ A H = Sandstone -shale Uplands ,eFanore*,*,* XK Recently glaciated shale lands FIGURE 1. The area referred to as the Burren, showing the four topographic regions. The dashed line in the north east corner separates Co. Clare (v.c. H9) to the south west from South East Galway (v.c. H15) to the north east. 10 FERN GAZETTE: VOLUME 12 PART 1 (1979) Historically the name Burren, which means great rock, referred to the Barony of Burren. The Barony occupied the north of Co. Clare roughly down to a line from Carron through Kilfenora and Lisdoonvarna to Poulsallagh. Botanically the name has a wider sense and includes surrounding areas. It is used here for a large area of northern County Clare (v.-c. H9) and a smaller, adjacent area of South East Galway (v.-c. H15). This is an area approximately 28km. north/south and 40km. east/west (fig. 1). This is slightly less than the area of Ivimey-Cook & Proctor (1966a) in their paper on the plant communities of the Burren, as it excludes a small area to the east around Gort. A bibliography of the area is given in Malloch (1976). Geologically the area consists of pure Carboniferous Limestone with only a few bands of chert and shale, overlain in the south-east corner by shales and sandstones of the same age. There is little glacial drift and that which there is bears a close relationship to the composition of the rocks it overlies (Finch 1971). Sweeting (1955) describes the topography of the area, which consists of four distinct regions in County Clare (fig. 1). Three of these form a block of high ground to the west of the fourth lower-lying region (Finch 1971). The three upland regions are the Burren Hills, the Sandstone-shale Uplands and the Recently Glaciated Shale Lands, the fourth region is the Burren Lowlands. The Burren Hills occupy the north-west of the area. They are an elevated block (300m) of horizontally-bedded limestone, dissected by two, broad, dry valleys running south from Ballyvaghan and Bealaclugga. To the east it forms a bold escarpment to the Burren Lowlands from Abbey Hill to Killinaboy and to the south it merges into the Sandstone-shale Uplands. There is a great deal of bare limestone, with soil in the valleys and depressions known locally as dolines. The hill sides are steep, showing the vertical edges of the limestone strata, with flat areas of limestone pavement between. The water table is well below the surface except for the deeper dolines. The only permanent river is the Cahir, which rises north-east of Slieve Elva and runs north-west to the sea at Fanore. The river only exists because the floor of its valley is choked with boulder clay. The Sandstone-shale Uplands form an elevated block, which drops in altitude from that of the Burren Hills in the north to around 100m in the south. The sub-soil is covered in poorly-drained, acid soils and peat, and there is abundant surface drainage by means of small streams. The division between the Sandstone-shale Uplands and the Recently Glaciated Shale Lands is not as dramatic as that between the former and the Burren Hills. There is merely a gradual decrease in altitude and an increase in recent glacial deposits. There are ice-deepened valleys in the shale with ridges of sandstone between. The soils are wet and acid. The Burren Lowlands consists of a low-lying (30—60m) plateau of horizontally bedded Carboniferous Limestone. There is more glacial drift than on the Burren Hills but there are still extensive areas of limestone pavement. However, the water table is only just below the surface and there is a chain of large permanent lakes from'Corrofin north-east to Boston. The very similar low-lying area of limestone to the south and west of Kinvarra, in South East Galway, is included here in this region, although it lacks the large Lakes of the Burren Lowlands. This is referred to as the Ardrahan Limestones in Praeger (1934). The climate of the whole area is highly oceanic like other western areas of the British Isles, though details for aspects other than rainfall are sparse (Finch, 1971; lvimey-Cook & Proctor 1966a). Rainfall is high in the upland regions, in the range 1250—1500mm but less in the lowlands, around 1125mm. The precipation-to- evaporation ratio is well in favour of precipitation, which means that most soils, even WILLMOT: FERNS OF THE BURREN 1 shallow soils over limestone (Grime, 1963) are leached and, in areas where the sub-soil is non-calcareous, podzolised. Where drainage is impeded, there are extensive areas of gleyed soils. The great botanical interest of the area lies in its high floristic diversity due to a mixture of phyto-geographical elements which is unique in western Europe. This is described in Lousley (1950), Praeger (1934) and Webb (1962). Plants which in Europe have a southern distribution, e.g. Adiantum capillus-veneris™ , grow alongside plants of a northern distribution and with yet others of Mediterranean, continental and arctic- alpine distributions. Another remarkable feature is the growth of the arctic-alpine plants at sea-level, which at these latitudes normally grow on mountain tops. FIGURE 2. Hazel scrub in Glen of Clab showing Polystichum setiferum also present was Pteridium aquilinum, Athyrium filix-femina, Dryopteris filix-mas and D. austriaca. A general survey of the vegetation of the area is given in lvimey-Cook & Proctor (1966a); and, more particularly, of fissures in the limestone in Dickinson et a/. (1964); of salt-marshes, fens and woodlands in Ilvimey-Cook & Proctor (1966b); and of temporarily flooded depressions in the limestone in Praeger (1932). Woodlands are scarce in the area. Semi-natural woodlands of Fraxinus exce/sior occur on some of the steeper slopes of the Burren Hills. There are small, ornamental plantations throughout the area and recent conifer plantations on the shales around Lisdoonvarna. Cory/us avellana scrub is abundant on the limestone hills and lowlands (fig. 2) and there are scattered hedges of Crataegus monogyna on the shale regions. Tall herb communities are common throughout the area, as are grasslands of an acidic nature on the shales and of a calcareous nature on the limestone. These range from semi-natural communities to small areas of improved grassland on the best soils. These communities merge into marsh and fen communities, especially around the lakes in the Burren *Nomenclature follows Clapham et al. (1962) for angiosperms and Jermy et a/. (1978) for pteridophytes. 12 FERN GAZETTE: VOLUME 12 PART 1 (1979) Lowlands. Blanket bogs with small areas of heath are common on the shales and grit- stones, with some raised bogs around the lakes in the Burren Lowlands. Sandstone or shale outcrops are occasional in their areas, and outcrops of limestone and limestone pavement (fig. 3) dominate large tracts of their areas. Stone walls are common in the area, generally of stone local to the region. These are usually drystone walls but occasionally, and more often in the shale and sandstone areas, there is soil between the stones. Mortared walls of all stone types also occur throughout the area. FIGURE 3. Limestone pavement with grykes near the lighthouse Black Head, the habitat of Adiantum capillus-veneris, Asplenium scolopendrium and A. marinum. METHODS Field records on the distribution maps (figs. 4—7) refer to records made by the author in August 1978. These were gathered on the basis that two to three examples of each relevant habitat were visited, in each of the 10 km. grid squares of the Irish national grid in the area. A relevant habitat was defined as a major habitat type, that occurred commonly in the topographical region in which the grid square was situated. These major habitat types are outlined in the previous section. At each site visit a subjective estimate of the abundance of each fern present was made, for each habitat type in which it occurred (see under Hab/tat Lists for explanation of abbreviations used). No attempt was made to record where habitats contained no ferns. There is a:bias in the records to sites near roads, due to the time available to study the area, but it is impossible to assess the strength or significance of this. A search was made of the major works that give fern records for the Burren: Corry (1880), Foot (1860), Ilvimey-Cook & Proctor (1966a), More (1898), Praeger (1901, 1909 & 1934) and Scannel & Synnott (1972). Where these give significant extensions to the geographical or ecological ranges of species, above that recorded here, details are given in the systematic list. Records given in Ivimey-Cook & Proctor (1966a) are plotted on the distribution maps (figs. 4—7) as they are given on a1 km. grid square basis. WILLMOT: FERNS OF THE BURREN 13 SYSTEMATIC LIST OF TAXA Each taxon recorded in the field for the study area is given, with notes on (a) its distribution, abundance and habitats as recorded in the survey and (b) significant extensions to these from the literature. Taxa, recorded in the literature but not this survey, are given a (b) type entry. Reference to abundance in habitats in (a) refers only to abundance in examples of the habitat where the species occurred; no account is taken of how often a species was present in the particular type of habitat. Grid references prefixed with c/rca are my interpretations of localities originally given without a reference. oui Huperzia selago (b) Recorded for summit of Gleninagh Mountain (c.M/17.09) and Slieve Elva (c.M/15.04) records communicated by D.A. Webb (Pers. comm. 1978) and in Jermy et a/ (1978) for M/1.0, M/1.1 and R/0.8. The card for the M/1.0 record in Jermy et a/ (1978) gives the habitat as blanket bog. 5.1 Selaginella selaginoides (fig. 4) (a) Scattered in small amounts in damp, calcareous, herbaceous communities in flushes around coast near Black Head and lakes in the Burren Lowlands. (b) Recorded in Jermy et a/ (1978) for M/2.1 and R/2.9.. 7.1 x 2 Equisetum x trachyodon (b) Recorded in Jermy et a/. (1978) for R/3.9 and in Praeger (1901) near the outlet stream to Lough Inchicronan (c.R/38.86). These are specimens in BM for the outlet stream to Lough Inchicronan dated 1925 and for the east shore of Lough Bunney (c. R/38.97) dated 1966. 7.2. &. variegatum (a) Widely distributed but very local in the Burren Hills and Lowlands. Rare to locally abundant in tall, damp, herbaceous vegetation dominated by monocotyledons, associated with flowing water or loughs. Specimens from Carran (R/28.99) and Lough Bunny (R/38.97) determined as probably var. majus Syme by C.N. Page. Also recorded Cahir River (M/15.08). (b) Recorded in Jermy et a/. (1978) for M/1.2 & M/3.1. 7.3 E. fluviatile (fig. 4) (a) Widespread but local in the Burren Lowlands and shale regions. In the Burren Lowlands, it is locally frequent to occasional as an emergent along the edges of loughs, and frequent to occasional in marshes fringing them. Occasional to frequent in damp, tall grass and marshy patches in fields in the Burren Lowlands and shale regions. Recorded once growing in old peat workings on blanket bog, Slievebeg (R/16.89). (b) Recorded in Jermy et a/. (1978) for M/1.0, M/1.1, % M/2.0 in the Burren Hills and R/0.9. 7.4 E. arvense (fig. 4) (a) Widespread throughout the Burren but usually only occasional to frequent, and it is less common in the limestone regions. Occurs usually in tall, often damp, grass and rarely in woods. (b) Recorded in Jermy et a/. (1978) for M/1.1, M/3.0 and R/3.8. 7.4x3 E. x litorale (a) Recorded as occasional in tall grass at northern end of Lough Bunny (R/38.97); and frequent on Cahir River (M/15.08) in tall, marshy grass by river and amongst stones in bed of river. (b) Recorded in Jermy et a/. (1978) for M/1.1. 15 E. sylvaticum (a) Restricted to the shale regions away from the coast but only very local even there. Occasional in damp, tall herbaceous vegetation and locally frequent in the recent, conifer plantation of Cragan West Wood (M/18.01). Recorded for R/13.90 and R/15.97. (b) Recorded in Jermy et a/. (1978) for R/1.8. FERN GAZETTE: VOLUME 12 PART 1 (1979) 14 18 LY UT bea | ANAT OBa Ga P| J88R M-F- 10? JEREEER Tso Tart | { far TT TTT BEESEGRGE000 GSSSSeG2B8b0nb005 > -on@ Boras AEeeSeeSoce SSSSOS00eC50a0 BESSEERESR0CE5 BSEEGH ESG2:GG0GC808200 0000000050 SenCSeEGs DUSSE Pe PEAS . bt c, E. arvense; , FIGURE 4. The distribution of the following in the Burren, circles Willmot 1978 and triangles d, E. palustre; e, E. telmateia and f, Ophioglossum vulgatum. lvimey-Cook & Proctor (1966a); a, Se/aginella selaginoides; b, Equisetum fluviatile; WILLMOT: FERNS OF THE BURREN 15 7.7 E. palustre (fig. 4) (a) Widespread but local in the shale regions away from the coast, although where it occurs it is occasional to frequent. Seen only once growing in a limestone region (M/15.08) and there on boulder clay. Habitats were mostly damp, tall grass and occurred once in old, peat workings on blanket bog, Slievebeg (R/16.89). (b) Ivimey-Cook & Proctor (1966a) record it frequently in the Burren Lowlands and Jermy et a/. (1978) for the same region in R/3.8. 7.8 E. telmateia (fig. 4) (a) Widespread but local in the shale regions, and occasional to frequent where it grows. It is very local in the Burren Hills on limestone, and in at least one of its two localities there, it grows on boulder clay (M/15.08). It usually occurs in tall grass and is especially abundant along roads round Lisdoonvarna. (b) Recorded in Jermy et a/. (1978) for R/0.9, and in the Burren Lowlands for R/3.9. 8.1 Botrychium l/unaria (b) Recorded in Jermy et a/ (1978) solely for the coastal squares M/1.0, M/1.1, M/2.1 & R/0.9. Older records give inland records for pasture near Lisdoonvarna (Corry 1880) and pasture near Inchicronan Lough c. R/38.86 (specimen in Herb BM dated 1905). Original cards for records in Jermy et a/. (1978) for M/1.0, M/2.1 & R/0.9 give grassland as habitat. 9.1 Ophioglossum vulgatum (fig. 4) (a) Widespread but local throughout the region on acidic and basic sub-soils, in tall grass, where it is rare in abundance. (ob) Recorded in Jermy et a/. (1978) for M/1.0, M/1.1, M/2.1, R/0.8, R/0.9, R/2.9 & R/3.9 Ivimey-Cook & Proctor (1966a) recorded it for damper, tall grass (R/17.93) and Corylus avellana scrub (R/26.93). Recorded in Praeger (1909) for Plantago sward near the sea. 10.1 Osmunda regalis (fig. 5) (a) Widely distributed but local on the shale regions and Burren Lowlands. It is occasional to frequent in areas of blanket bog and heath on the shale and at the edge of raised bogs around lakes in the Burren Lowlands. (b) Recorded in Jermy et a/. (1978) for M/1.0, R/0.8, R/1.8, and R/3.9. 13.1. Adiantum capillus-veneris (fig. 5) (a) Widespread but local in the Burren Hills near the coast. It is occasional in grykes and damp crevices in the limestone. (b) Recorded for a wider area of the Burren Hills and Lowlands in Jermy et a/. (1978): M/2.1, M/3.0, M/3.1, R/1.9 and R/2.9; and Praeger (1901) records it for two areas of crags north of Corofin (c. R/28.89). The present records and those in Jermy et a/. (1978) when viewed on a 10 km. basis indicate a distribution throughout the limestone areas. However, when these, along with the older records in Foot (1860) and More (1898), are viewed on a 1 km. basis there is a tendency for a coastal distribution. 14.2 Hymenophyllum wilsonii (b) Recorded in Jermy et a/ (1978) for M/2.0 and M/3.0, habitats on original record cards are “under moss-covered boulder” and ‘‘tree trunk’ respectively. D.A. Webb (Pers. Comm. 1978) records the species as an epiphyte on a tree trunk and as growing on a sandstone erratic in Fraxinus excelsior woodland come Corylus avellana scrub in the Poulavallan doline (c. M/28.02). 16. 1—3 Polypodium vulgare agg. (fig. 5) (a) Widespread and common throughout the area on walls where it is occasional to frequent. Widespread but less common on outcrops of limestone and shale, where it is occasional in abundance. It is also widespread, but much less common, as an epiphyte on Corylus avellana in C. avellana scrub, on Acer pseudoplatanus trees and rarely on Alnus glutinosa trees and Crataegus monogyna bushes. It is locally frequent on moss-covered rocks on the floor of C. ave//ana scrub on limestone, and on the floor of woods on limestone and shale. It was also recorded once, as occasional, on the banks of hedges in the shale regions (R/09.91). All segregated records below have been determined by J.M. Mullin of the British Museum (Nat. Hist). As only a few of the FERN GAZETTE: VOLUME 12 PART 1 (1979) 16 He EHH a @r A a> 5} bs S A 4 veneris; c, Polypodium -nigrum. circles Willmot 1978 and triangles Osmunda regalis; b, Adiantum capillus Asplenium scolopendrium and f, A. adiantum e, , d, Pteridium aquilinum; FIGURE 5. The distribution of the following in the Burren, vulgare agg.; Ivimey-Cook & Proctor (1966a): a, WILLMOT: FERNS OF THE BURREN 17 records for the P. vu/gare aggregate were critically determined, it is not possible to be completely confident of the differences in the ecologies of the species as recorded here. However, the preference of P. interjectum and P. australe for lime-rich substrates and the avoidance of such by P. vulgare support findings elsewhere Benoit (1966) and Jermy et a/. (1978). 16.1 P. vulgare (a) Widespread in the shale regions. Recorded as frequent on shale walls near Ballybreen (R/15.92 & 15.93) and occasional as an epiphyte on base of tree in mixed, deciduous woodland near Lisdoonvarna (R/12.98). (b) Recorded by Jermy et a/. (1978) for R/2.8 and there is an herbarium specimen in BM for a wall (most probably limestone) near sea at Blackhead (M/1.1) dated 1962. 16.2 P. interjectum (a) Widespread throughout the area. Occasional to frequent on limestone walls both mortared and drystone (R/05.96, 18.93, 18.94, 23.93 & 26.89). Rare as an epiphyte on an Acer pseudoplatanus tree (R/14.94) and a Corylus avellana bush in hazel scrub (M/29.02). Occasional on a hedge bank near Moymore House (R/09.91). (b) Recorded by Jermy et al. (1978) for M/1.1, M/2.1 & M/3.0, 16.2 x 1 P. x mantoniae (a) Recorded once for a mortared limestone wall at Ennistimon (R/13.90) as occasional. First county record for Co. Clare. 16.3 P. australe (a) Scattered throughout the area. Occasional to frequent on mortared and drystone limestone walls (M/10.00, M/17.10 and R/25.92); and rare on grykes at Burrin (M/28.11) and on moss- covered boulders in hazel scrub at Dromore Woods (R/34.86). (b) Recorded by Jermy et a/. (1978) for M/3.0 with habitat given as ‘‘limestone karst” on original card, and R/3.9. 17.1. Pteridium aquilinum (fig. 5) (a) Common throughout both the limestone, and sandstone and shale regions. It is occasional to frequent in tall, acidic grassland; widespread and frequent in abundance in short grassland over limestone; and occasional to frequent in Cory/us avel/lana scrub and scrub in general. It is locally abundant in acidic, tall grass and herb communities in a few places. It is also rare to frequent in grykes in the limestone pavements. 18.1 Thelypteris thelypteroides (a) Recorded once as locally abundant in marsh next to Lough Inchicronan (R/39.87) where it was recorded by Praeger (1901). (b) Jermy et a/. (1978) also give M/2.0. There are old records for Ballycullinan Lough (c. R/28.86) in Praeger (1901) and a specimen in herb. BM for marsh in Inchiquin Lough dated 1905 (c. R/26.89). 20.1 T. /imbosperma (b) Jermy et a/. (1978) give M/1.1 and an old record for R/1.9. 21.1 Asplenium scolopendrium (fig. 5) (a) Widespread and common throughout the area in both areas of limsetone and of sandstone and shale, more abundantly in damper and shaded habitats. It is rare to frequent on walls and outcrops of all rock types; occasional in grykes in the limestone; rare to occasional on the floor of Cory/us avellana scrub and woods; and occasional on hedge banks in shale regions. 21.2 A. adiantum-nigrum (fig. 5) (a) Widespread but local in the limestone regions and very local in the sandstone and shale regions. Rare on outcrops of limestone and shale. (b) Recorded over a wide range of both the limestone and shale regions in Jermy et a/. (1978): M/1.0, M/1.1, M/2.1, R/1.9, R/3.8 and R/3.9. lvimey-Cook and Proctor (1966a) record it for lime- stone pavement at Carran (R/28.99). 21.5 A. billotii (b) Recorded in Jermy et a/. (1978) for M/1.0. 18 FERN GAZETTE: VOLUME 12 PART 1 (1979) =25 === Spee rsces: Piet lel lel Spall | BY IV moe =e va NEBZEBNG “= a SCSOREERERDATEH af 258 TT | 1 vacliaal bail He ae ages SEEZEREr== Bab Dp a an FIGURE 6. The distribution of the following in the Burren, circles Willmot 1978 and triangles lvimey-Cook & Proctor (1966a): a, Asp/enium marinum; b, A. trichomanes agg.; c, A. ruta-muraria; d, A. ceterach; e, Athyrium filix-femina and f, Cystopteris fragilis. WILLMOT: FERNS OF THE BURREN iS) 21.6 A. marinum (fig. 6) (a) Widespread but local along the coast particularly the western coast. Occasional on walls and outcrops of both limestone and shale and frequent in grykes in the limestone (fig. 3). (b) Recorded in Jermy et a/. (1978) for M/1.0 and in Praeger (1901) for Ballyvaghan (c. M/22.08). 21.7. A. trichomanes agg. (fig. 6) (a) Widespread and common throughout the area. Occasional to frequent on limestone outcrops; and on walls of limestone and of shale. Rare in grykes in the limestone. Recorded once, as rare, in Corylus avellana scrub. (b) Ivimey-Cook & Proctor (1966a) record it more commonly in C. avellana scrub. All specimens collected would appear to be referable to subsp. quadriva/lens, which is the common subspecies on lime-rich substrates, but no specimens were critically determined. 21.9 A. ruta-muraria (fig. 6) (a) Widespread and common throughout the area. Frequent to occasional on outcrops of limestone; and on walls of limestone or shale. Recorded once as occasional in grykes in limestone. (b) Ivimey-Cook & Proctor (1966a) record it more commonly in grykes. 21.11 A. ceterach (fig. 6) (a) Widespread and common throughout the area except near the coast between Liscannor and Black Head. Widespread and occasional to rare on outcrops of limestone. Widespread and frequent to rare on walls of limestone, but much more local on shale walls. (b) Ivimey-Cook & Proctor (1966a) record it more commonly in grykes, and Jermy et a/. (1978) record it for R/0.9. 22.1 Athyrium filix-femina (fig. 6) (a) Widespread and common in the shale regions, but less widespread and less frequently encountered in the limestone regions. Frequent to occasional in damp, tall grass in the shale regions. Occasional in hedges; in Cory/us avellana scrub on limestone; and in woods throughout the area. (b) Jermy et a/. (1978) record it for M/1.1, R/0.8 and R/2.9. Plants with red rachises and with green rachises occur in the area. 23.1 Gymnocarpium dryopteris (b) Praeger (1901) records this species on outcrop by road between Roadford and Cliffs of Moher. 24.1 Cystopteris fragilis (fig. 6) (a) Widespread but local throughout the Burren Hills and the Sandstone-shale Uplands. Rare to occasional on damp outcrops of shale and limestone. One favoured situation in this type of habitat is around swallow-holes on the edge of the limestone, where streams passing off the shales go underground. Only occurs on limestone walls at their base amongst other vegetation. Recorded once in grykes as occasional. (b) Jermy et a/. (1978) record it for a wider range: M/2.1, M/3.0, M/3.1, R/0.8, R/1.8, & R/3.9. Prof. Webb considers it rarer in the Burren Lowlands than elsewhere in the area (D.A. Webb Pers. Comm. 1978). 26.2 Polystichum aculeatum (fig. 7) (a) Widespread but local throughout the Burren Hills and very local in the shale regions. Rare to occasional on limestone outcrops, in grykes and tall grass. Occasional in Cory/us avellana scrub; and rare in shale hedgebanks. (b) Recorded for the Burren Lowlands and for a wider range of the Burren Hills in Jermy et. a/. (1978) M/3.0 and R/3.9. Much less common and abundant than P. setiferum. 26.2 x 3 P. x bicknellii (a) Recorded once as rare on rock outcrops in quarry by road north east of Cross Bridge (R/25.91). (b) Recorded for R/2.9 in Jermy et a/. (1978), with habitat given as walls of old mineshaft on Original card. FERN GAZETTE: VOLUME 12 PART 1 (1979) 20 KG N KD a eae = NStal mci I] Ps * Ta BA fe a [ tt ae The distribution of the following in the Burren, circles Willmot 1978 and triangles FIGURE 7. , c, Dryopteris filix-mas; _P. setiferu Blechnum spicant. 2 ge 22 So o = & > Oo So i Eo SS =e Aya 7S = as o & BAS 3 S O » Oo 4 oO 2 es = 18) en aes ot oO fe) = oO (oa) a fe) fe) QO > ® & 2 —_——, WILLMOT: FERNS OF THE BURREN 21 26.3 P. setiferum (fig. 7) (a) Widespread but rather local throughout the area. Occasional! to frequent in woods. Rare to occasional in Cory/us avellana scrub on limestone, though sometimes locally abundant in the latter habitat (fig. 2). Occasional in hedges in a few places in the shale regions. Rare on walls and rare to occasional in tall grass. (b) Jermy et a/. (1978) give records for M/1.1, M/2.1, & M/3.1. 27.2 Dryopteris filix-mas (fig. 7) (a) Widespread and common throughout the area but never in large amounts. Occasional in deciduous woods and Cory/lus avellana scrub on limestone. Rare to frequent in tall herb and grass communities. Occasional on limestone walls. Occasional to rare, on limestone outcrops and walls. 27.2x3 D.x tavelii (a) Recorded once as rare, with parents, in tall grass by roadside near Ennistimon (R/13.90). First county record for Co. Clare. 27.3. D. pseudomas (fig. 7) (a) Widespread but rather local throughout the area. Occasional in woods and Corylus avellana scrub on limestone. Rare in grykes. Occasional in damp, tall grass communities in shale regions. (b) Recorded for M/3.0 in Jermy et a/. (1978). 27.5 D.aemula (a) Widespread but very rare throughout the area. Recorded three times: frequent in mixed deciduous wood Lisdoonvarna (R/15.97); occasional in Cory/lus avellana scrub Poulavallan (M/29.02); and rare in damp, tall, herbaceous vegetation Slievebeg (R/16.89). 27.9 D. austriaca (fig. 7) (a) Widespread and common in the shale regions, while less widespread and rare in the limestone regions. Rare to occasional in mixed deciduous woods; occasional but much less common in Corylus avellana scrub on limestone and rare to occasional in damp, tall grass and blanket bog in the shale regions. (b) Recorded more commonly in C. avellana scrub on limestone by Ilvimey-Cook & Proctor (1966a). 28.1 Blechnum spicant (fig. 7) (a) Rare but widespread in the shale regions and recorded once for the Burren Hills, as rare in Corylus avellana scrub. Occasional to frequent in heath and short, acid, grass communities in the shale regions. Occasional in woods on shale and very local but frequent where it does occur on shale outcrops. (b) Recorded over a wider part of the western and central portions of the area in Jermy et al. (1978): M/0.0, M/1.1, R/0.8, R/2.8 & R/2.9. HABITAT LISTS Species lists are given for some of the more interesting and/or characteristic fern habi- tats of the Burren. As in the systematic list, abundance only refers to abundance in examples of the habitat in which the species in question occurred. The abbreviations ‘D', ‘A’, 'F’, ‘0’, &'R’ stand for Dominant, Abundant, Frequent, Occasional and Rare respectively and ‘( )’ means exceptionally. ‘L’ stands for locally. How often the species occurred in the habitat is given as a percentage frequency, rounded to the nearest ten percent. Woodland No examples of woodlands consisting of ‘’"an open tree layer over-topping the hazel (scrub)’’ were examined (Ivimey-Cook & Proctor, 1966a). However, as these merge into hazel scrub, its species list can be taken as a good approximation for this habitat. The list here refers to mixed deciduous woodlands of an obviously planted nature, and to eight examples of the vegetation. 22 FERN GAZETTE: VOLUME 12 PART 1 (1979) Species Abundance Occurrence (%) Equisetum arvense R 30 Polypodium vulgare agg. O 40 Pteridium aquilinum R—O 40 Asplenium scolopendrium O — F(LA) 80 Athyrium filix-femina O 50 Polystichum setiferum O — F(LA) 100 Dryopteris filix-mas O 80 D. psuedomas O 50 D. aemula = 10 D. austriaca ~ R—-O 90 Blechnum spicant O 30 The paucicity of the flora for deciduous woods on limestone probably reflects the planted nature of the woodlands and lack of damp areas. Hazel Scrub This is one of the most characteristic types of vegetation in the Burren (fig. 2). It is species-rich and has a typical, woodland-ground flora. Although often restricted to more-sheltered situations and often wind-cut in more exposed areas, it is considered that hazel scrub could cover much more of the Burren (Ivimey-Cook & Proctor, 1966a), presumably if grazing pressure was reduced. Pollen analysis (Watts, unpubl.), however, indicates that it is not the climax vegetation at least in the east of the area. The list refers to eleven examples of the vegetation. Species Abundance Occurrence (%) Polypodium vulgare agg. R 10 Pteridium aquilinum O-—F 80 Asplenium scolopendrium R—O 70 A. adiantum-nigrum R 10 A. trichomanes agg. R 10 Athyrium filix-femina One} 30 Polystichum aculeatum O 10 P. setiferum R — O(LA) 80 Dryopteris filix-mas R—F 90 D. pseudomas O 30 D. aemula O 10 D. austriaca O 30 Blechnum spicant R 10 Allowing that my samples are much larger than those of Ivimey-Cook and Proctor (1966a), there is a close similarity between their values and mine for the occurrence of ferns in hazel scrub. The only large differences are that they record Asplenium trichomanes agg. and Dryopteris austriaca more commonly. Assuming these are real differences, there are no obvious explanations. Generally the flora has a wood- land aspect apart from Asp/enium adiantum-nigrum and A. trichomanes agg. These are rock crevice plants and grow on limestone outcrops in the scrub. Grykes Grykes (fig. 3), which are vertical fissures in limestone pavement, have long held a special fascination for botanists in the Burren (Dickinson et a/., 1964; Heslop-Harrison, 1960), probably because they contain a rich flora, in stark contrast to the bare —_———iit, FO. —_—— WILLMOT: FERNS OF THE BURREN 23 pavement around them. They do not constitute a single habitat, as pointed out by lvimey-Cook & Proctor (1966a), but a mixture of several distinct types, including gryke floors and rock crevices. Nevertheless grykes are considered a single habitat here, as this was considered sufficient for the survey. The list refers to fourteen examples of the vegetation. Species Abundance Occurrence (%) Adiantum capillus-veneris (M) O 10 Polypodium vulgare agg. Pteridium aquilinum D | D mn ON oi =) Asplenium scolopendrium O 90 A. marinum (M) F 20 A. trichomanes agg. (R) R 20 A. ruta-muraria (R) O 10 Cystopteris fragilis (R) O 10 Polystichum aculeatum O—R 20 P. setiferum R 10 Dryopteris filix-mas (W) O—R 20 D. pseudomas (W) R 10 (W) = Woodland-floor elements (R) = Rock-crevice elements and (M) = Maritime element The flora illustrates the multiple habitat nature of grykes, containing as it does woodland-floor and rock-crevice elements. The presence of the former is explained by the micro-climate of the gryke floors being like that of woodland floors in the vicinity (Dickinson et a/., 1964). A third element is the two species that only occur commonly in grykes near the sea, however, these species occur separately. lvimey-Cook & Proctor (1966a: Table XL) give a species list with higher frequencies of the rock-crevice element and also include Asp/enium ceterach. This is probably because their list refers to limestone pavement in general and not just grykes. However, Webb (1962) also records the Asp/enium spp. as common on the pavement. Limestone outcrops The list refers to natural outcrops and not man-made ones. However, this does not mean that it describes or even resembles the flora of outcrops before the advent of agriculture. No doubt originally outcrops would have been generally more shaded and have harboured a different flora at least in quantitative terms. This is the only commonly-occurring terrestrial habitat apart from walls, which are artifical rock outcrops, where ferns are the dominant element in the vegetation. The list refers to twenty-eight examples of the vegetation. The two maritime-element species only occurred in crevices near the sea. They occurred separately with practically no other vascular plants. lvimey-Cook & Proctor (1966a) note similar differences between crevices near and away from the sea. Asplenium scolopendrium and Cystopteris fragilis occurred more commonly in damper areas. This list only differs significantly from the Ilvimey-Cook & Proctor (1966a) list by the greater frequency of A. sco/opendrium. This may be due to the fact that A. scolopendrium prefers damper habitats and thus more often grows at the base of outcrops. | may have been more likely to record it as on the outcrop in such situations than lvimey-Cook & Proctor. 24 FERN GAZETTE: VOLUME 12 PART 1 (1979) Species Abundance Occurrence (%) Adiantum capillus-veneris (M) R 10 Polypodium vulgare agg. O 20 Asplenium scolopendrium O 60 A. adiantum-nigrum R 10 A. marinum (M) O 10 A. trichomanes agg. O) = IF 70 A. ruta-muraria O-—F 60 A. ceterach R—O 50 Cystopteris fragilis R—O 20 Polystichum aculeatum R—O 10 P. setiferum R 10 Dryopteris fillix-mas R—O 20 D. pseudomas O 10 (M) = Maritime element Limestone walls This is the most striking habitat for ferns because it is so common, especially near roads, and because ferns constitute a large part of its vascular flora. The distribution maps of Asplenium trichomanes agg, A. ruta-muraria and A. ceterach for my records show the pattern of major roads in the south of the area. The list refers to twenty-five examples of the vegetation on drystone walls and to twenty-two examples on mortared walls. Species Abundance Occurrence (%) Drystone Mortared Drystone Mortared Polypodium vulgare agg. O-F O-—F 60 60 Asplenium scolopendrium O R 70 40 A. adiantum-nigrum R—O _ 10 0 A. trichomanes agg. O-—F O-—F 60 80 A. ruta-muraria O O-—F 40 100 A. ceterach O-F R—F 50 80 Cystopteris fragilis O — 10 0 Polystichum setiferum R—F R 10 10 Dryopteris filix-mas O O—R 20 10 Cystopteris fragilis and Dryopteris filix-mas only occurred at the base of drystone walls, undoubtedly due to the preference of these species for damper habitats. The overall similarity of the floras of limestone walls and outcrops is obvious but there are important differences. Walls in the area often have soil cappings but out- crops rarely do and in at least one area Po/ypodium vulgare agg. shows a preference on limestone, for areas of thin soil (Willmot, 1977). The major differences in occurrence of species between drystone and mortared walls may be due to differences in size of crevices. Mortared walls have a predominance of smaller, presumably drier, crevices, whereas drystone walls have larger, presumably damper, ones. It is possible then that Asplenium ruta-muraria and A. ceterach, which amongst the outcrop-ferns occur in the driest situations, are commoner on mortared walls because they grow in the smaller, drier crevices. On the other hand A. sco/opendrium may occur more commonly on the drystone walls because it prefers damper crevices. WILLMOT: FERNS OF THE BURREN 25 Shale walls These are less numerous than limestone walls. The list refers to eight examples of the vegetation of drystone walls and to nine examples of that on mortared walls. Species Abundance Occurrence (%) Drystone Mortared Drystone Mortared Polypodium vulgare agg. O-F O 60 40 Asplenium scolopendrium = R 30 30 A. marinum — O 0 20 A. trichomanes agg. R O-—F 10 70 A. ruta-muraria O-—F O-—F 40 80 A. ceterach O O-—F 10 30 Athyrium filix-femina O F 10 10 Cystopteris fragilis _ E 0 10 Polystichum setiferum _ A 0 10 Dryopteris austriaca O — 10 0 The difference in abundance and occurrence between the two types of wall for A. scolopendrium, A. ruta-muraria and A. ceterach mirror the differences for these species between the two types of limestone wall. This is presumably for the same reason. In terms of occurrence the mortared, shale walls are more like limestone walls than the drystone, shale walls. This is possibly due to the mortar increasing the base status of the otherwise acid soil that collects in the crevices of the shale walls. The hypothesis that the soil in shale walls is acid if mortar is not present, is supported by the appearance of a calcifuge element on shale walls e.g. Dryopteris austriaca and Athyrium filix-femina. SUMMARY OF FLORA Thirty-five species are recorded from the area, which is 60% of the native fern flora recently recorded for Ireland (Jermy et a/. 1978). An idea of the richness of the flora of the area can be obtained from Jalas & Suominen (1972). They record only seven out of forty-eight 50 km squares with more species than the square containing this study area. Note that the study area is about half the area of one of their squares. Also five hybrids out of nine recently recorded for Ireland were found. Two, of these, Polypodium x mantoniae and Dryopteris x tavelii are first county records and a third Polystichum x bicknellii a second county record for Co. Clare. The record for D. x tavelii is only the second for Ireland, though the hybrid is doubtless under-recorded there. The most valuable contribution of the fern flora of the area to that of the British Isles is the abundance of Adiantum capillus-veneris. This has nine of its thirty-three recent localities (Jermy et a/. 1978) in the area. Efforts should therefore be made to conserve the best of these localities. Two species, Asp/enium viride and Dryopteris carthusiana, which are widespread elsewhere in Ireland in habitats that occur in this area are absent from the study area. The ease of recognition of the species and the extent to which the area has been studied suggest that these are probably genuine absences. However, they should be looked for in the area. The ferns illustrate the well-known commingling of northern and southern elements in the flora (Praeger, 1934). Examples of northern species are Huperzia selago and Selaginella selaginoides and of southern species Adiantum capillus-veneris and Polypodium australe. There is, however, a complete absence of any arctic-alpine element although there is one in the flowering plant flora (Praeger 1934). A more 26 FERN GAZETTE: VOLUME 12 PART 1 (1979) detailed analysis of the phytogeographical affinities of the flora is given below and in Table 1. This is based on the European distribution of species as recorded in Jalas & Suominen (1972) and Jermy et a/. (1978). The figures for the British Isles refer to recently-recorded, native species of the British Isles excluding the Channel Isles. Minor elements are excluded, as are a few species whose distribution is intermediate between the classes given or is indeterminate. On the same basis there are eleven widespread species in Europe which all occur in the Burren; seven arctic-alpine species none of which occur; and seven northern, montane species two of which occur. TABLE 1 : THE PHYTOGEOGRAPHICAL AFFINITIES OF THE FERN FLORA OF THE BURREN Atlantic Sub-atlantic Continental Total British British British British Isles Burren Isles Burren Isles Burren Isles Burren Northern Southern Widespread Total Overall Table 1 shows the atlantic/sub-atlantic and southern affinities of the flora. Most striking is the presence of all species of southern, atlantic and southern, sub-atlantic affinity in Europe which are in the British fern flora. This is particularly so since the area is so far north relative to the definition of southern Europe in Tutin et al. (1964). This is probably due to the relatively mild climate of the area for a region 53°N, which is caused by the warming effect in winter of air masses moving over the area from the Atlantic. This southern nature to the flora is not so marked in the bryophyte flora of the area. Ratcliffe (1968) records 18 out of his 25 northern atlantic bryophytes in Mid-West Ireland, and 16 out of his 27 southern atlantic. Note, however, that Ratcliffe’s Mid-West area is much larger than the Burren. The relatively high representation of atlantic/sub-atlantic species and low representation of continental species is to be expected in an area on the western seaboard of Europe. Although the relatively higher over-representation of sub-atlantic compared to atlantic species is worthy of comment. This may be due to the fact that although the Burren is on the western seaboard of Europe and thus has a damp, oceanic climate, this is partially ameliorated by the exposed and/or free-draining nature of many of the habitats in the area. Birks (1976) gives a more detailed analysis of the phytogeography of European pteridophytes, dividing them into twenty-one floristic elements, that is groups of species with similar distributions. Examination of the placing of the thirty-five species from the Burren amongst Birks’ groups confirms the southern and western (atlantic) affinities of the flora and lack of northern, eastern and alpine elements. However, the character of the flora that emerges from this examination is rather more southern and less western than the previous analysis (Table 1) suggested. This is probably a truer picture of the affinities of the flora as Birks’ methods were more objective. One ecological problem in the area has not been discussed: the occurrence of species in more acidic habitats than those in which they usually occur. The best example is Pteridium aquilinum, a calcifuge which occurs commonly in short calcareous grassland over limestone pavement. Webb (1962) records it in such habitats more especially in the east of the area. In such grassiand it is usually stunted (Webb ee ee a WILLMOT: FERNS OF THE BURREN 27 1962) being about 40 cm. high and it does not grow in dense masses but as separate fronds. It is temping to suggest this anomalous occurence is due to leaching of the top layers of the soil caused by the high rainfall of the area. However, Grime (1963) demonstrates that some calcifuges occur in calcareous soils in the area without surface leaching. Jermy et a/ (1978) suggest an alternative hypothesis that there might be a calcicole ecotype involved, similar to the diploid cytotype on limestone in Spain. As no calcicole ecotypes have been found in calcifuge species growing on calcareous soils (Grime 1963), the ecotype explanation appears unlikely. As a number of calcifuge flowering plants occur in similar habitats in the area (Grime 1963 & Webb 1962), a more general explanation appears probable. However, Grime (1963) has analysed the ecologies of a number of calcifuge species growing on calcareous substrata in various sites in the British Isles and could find no general explanation. Amongst the other species growing in more calcareous habitats than usual, Oreopteris limbosperma & Blechnum spicant have each been recorded once in a locality which suggests that they were growing in a habitat similar to P. aquilinum. | have recorded O. /imbosperma once growing in soil over Carboniferous Limestone in Derbyshire. Dryopteris pseudomas is recorded in Clapham et a/. (1962) as usually on acid soils but is recorded here as growing once on a limestone outcrop, once in a gryke and once amongst limestone blocks. Willmot (1977) records the species as growing rarely on Carboniferous Limestone in Derbyshire. ACKNOWLEDGEMENTS My thanks are due to Prof. D.A. Webb of Dublin Univ. for pointing out some of the more interesting pteridological sites and problems of the area. My thanks are due to the following authorities for determining critical material in the genera indicated: A.C. Jermy and J.M. Mullin of the British Museum (Nat. Hist.), Po/ypodium and Dryopteris; C.N. Page of the Royal Botanic Garden, Edinburgh, Equisetum; and A. Sleep of Leeds University, Po/ystichum. | also wish to thank F.H. Perring of the Biological Records Centre, Monk’s Wood for information on habitats of records in Jermy et al. (1978). REFERENCES BENOIT, P.M. 1966. Some recent work in Wales on the Po/ypodium vulgare aggregate. Fern Gaz. 9 : 277-282. BIRKS, H.J.B. 1976. The distribution of European pteridophytes : A numerical analysis. New Phytol. 77 : 257—287. CLAPHAM, A.R., TUTIN, T.G. & WARBURG, E.F. 1962. Flora of the British Isles. Cambridge. CORRY, T.H. 1880. Notes of a botanical ramble in the County of Clare, Ireland. Proc. Rep. Belt. Nat. Hist. Phil Soc., 167—207. DICKINSON, C.H. PEARSON, M.C. & WEBB, D.A. 1964. Some micro-habitats of the Burren, their micro-environments and vegetation. Proc. R. Ir. Acad. 63B : 291—302. FINCH, T.F. 1971. Soils of County Clare. Dublin. FOOT, F.J. 1860. On the Ferns of West Clare, being a list of those growing west of a line drawn north and south through the town of Tulla. Wat. Hist. Rev. 7 : 36—40. GRIME, J.P. 1963. Factors determining the occurrence of calcifuge species on shallow soils over calcareous substrata. J. Ecol. 57 : 375—390. HESLOP-HARRISON, J. 1960. A note on temperature and vapour deficit under drought conditions in some microhabitats of the Burren limestone, Co. Clare. Proc. R. Ir. Acad. 67B : 109—114. IVIMEY-COOK, R.B. & PROCTOR, M.C.F. 1966a. Plant Communities of the Burren, Co. Clare. Proc. R. Ir. Acad. 64B : 201—311. IVIMEY-COOK, R.B. & PROCTOR, M.C.F. 1966b. The application of association analysis to phytosociology. J. Eco/. 54: 179-192. JALAS, J. & SUOMINEN, J. 1972. Atlas Florae Europaeae, Vol. 1. Helsinki. 28 FERN GAZETTE: VOLUME 12 PART 1 (1979) JERMY, A.C., ARNOLD, H.R., FARRELL, L & PERRING, F.H. 1978. Atlas of ferns of the British Isles. London. LOUSLEY, J.E. 1950. Wild flowers of Chalk and Limestone. London. MALLOCH, A.J.C. 1976. An annotated bibliography of the Burren J. Ecol. 64: 1093—1105. MORE, A.G. 1898. Cybele Hibernica 2nd Edn. Dublin. PRAEGER, R.L. 1901. Irish Topographical Botany. Proc. R. Ir. Acad. 7B. PRAEGER, R.L. 1909. A tourist’s flora of the West of /reland. Dublin. PRAEGER, R.L. 1932. The flora of the turloughs : a preliminary note. Proc. R. Ir. Acad. 47B : 37—45. PRAEGER, R.L. 1934. The Botanist in Ireland. Dublin. RATCLIFFE, D.A. 1968. An ecological account of the atlantic bryophytes in the British Isles. New Phytol. 67 : 365—439. SCANNELL, M.J.P. & SYNNOTT, D.M. 1972. Census Catalogue of the Flora of lreland. Dublin. SWEETING, M.M. 1955. The landforms of north-west County Clare, Ireland. Trans. Inst. Br. Geogr. 217: 33—49. TUTIN, T.G., HEYWOOD, V.H., BURGES, N.A., VALENTINE, D.H., WALTERS, S.M. & WEBB, D.A. 1964. Flora Europaea. Vol. 1. Cambridge. WATTS, W.A. Unpubl. Quoted in Ilvimey-Cook & Proctor (1966a). WEBB, D.A. 1962. Noteworthy plants of the Burren : a catalogue raisonné. Proc. R. Ir. Acad. 62B >: 117-34. WILLMOT, A. 1977. A pteridophyte flora of the Derbyshire Dales National Nature Reserve. Fern Gaz. 17: 279—284. REVIEW THE EIGHTIETH BIRTHDAY OF R.E. HOLTTUM, 1975. A special issue of the Gardeners’ Bulletin Singapore, Vol. 30, 312pp published 15th October 1977 edited by Chang Kiau Lan. Available from the Botanic Gardens, Singapore 9, price $49.40 (about £11.12) post free. This birthday issue for Richard Eric Holttum contains introductory biographical accounts and 22 papers of which 13 are about ferns. Descriptive taxonomical papers including B.J. Hoshizaki on Platycerium (13—15); G.J. de Joncheere on Humata (45— 58); K. lwatsuki on Meringium (63—74); M.G. Price on Dryopteris (239—250) and A.C. Jermy & T.G. Walker on Botrychium. Three papers describe spore morphology: E. Soepadmo & E.E. Khoo on Malayan Dennstaedtiaceae (85—95); F.S. Liew, on Oleandraceae (101—110) and K.U. Kramer on ‘Synaptospory : a hypothesis’ (79—83) in which the author elaborates an interesting theory in which spore ornamentation ts said to hold spores together thus allowing two prothalli to establish themselves, and thus help cross fertilization. A very interesting account and discussion by Herb and Florence Wagner (251—267) on fertile-sterile leaf dimorphy is stimulating and that by D.W. Lee (21—29) on iridescence in Selaginella is similarly so. Two floristic papers: A.G. Piggott, the ferns of Gunong Ulu Kali (31—43) and a larger paper on the lime- stone hill flora of Malaya I, by S.C. Chin (165—219) make interesting reading. The latter gives notes and a key to 97 species of ferns in 43 genera. Last, a paper listing all Holttum’s new taxa and name changes in ferns to July 1975 by J.A. Crabbe (221— 238) which lists over 700 items abstracted from 420 papers and shows the product- ivity of this Grand Old Man of Pteridology, whom this journal seeks to honour. A.C. JERMY FERN GAZ. 12(1) 1979 29 THE ASSOCIATIONS BETWEEN PTERIDOPHYTES AND ARTHROPODS URI GERSON The Hebrew University of Jerusalem, Faculty of Agriculture, Rehovot, Israel. ABSTRACT Insects belonging to 12 orders, as well as mites, millipedes, woodlice and tardigrades have been collected from Pteridophyta. Primitive and modern, as well as general and specialist arthropods feed on pteridophytes. Insects and mites may cause slight to severe damage, all plant parts being susceptible. Several arthropods are pests of commercial Pteridophyta, their control being difficult due to the plants’ sensitivity to pesticides. Efforts are currently underway to employ insects for the biological control of bracken and water ferns. Although Pteridophyta are believed to be relatively resistant to arthropods, the evidence is inconclusive; pteridophyte phytoecdysones do not appear to inhibit insect feeders. Other secondary compounds of pteridophytes, like prunasine, may have a more important role in protecting bracken from herbivores. Several chemicals capable of adversely affecting insects have been extracted from Pteridophyta. The litter of pteridophytes provides a humid habitat for various parasitic arthropods, like the sheep tick. Ants often abound on pteridophytes (especially in the tropics) and may help in protecting these plants while nesting therein. These and other associations are discussed. It is tenatively suggested that there might be a difference in the spectrum of arthropods attacking ancient as compared to modern Pteridophyta. The Osmundales, which, in contrast to other ancient pteridophytes, contain large amounts of phytoecdysones, are more similar to modern Pteridophyta in regard to their arthropod associates. The need for further comparative studies is advocated, with special emphasis on the tropics. INTRODUCTION This is the fourth and final installment in a series of review papers intended to explore the relationships between arthropods and the lower green plants. These reviews, while not intended to be comprehensive, are meant to draw attention to some hitherto- neglected areas of arthropod-plant inter-relationships. Former parts dealt with mosses, lichens and algae, respectively (Gerson, 1969; 1973; 1974-76). The associations between arthropods and pteridophytes have been of some recent interest to entomologists. The continuing weed problem of bracken (Pteridium aquilinum) in some parts of the world (Braid, 1959), and the relatively new problem of Sa/vinia as a nuisance in Asian and African waterways (Anders and Bennett, 1975) are being tackled by a biological control approach. Some Pteridophyta, of ornamental interest, are grown commercially; their pests have become of economic importance. Finally, the discovery of insect moulting hormones in many pteridophytes has brought forth a series of biochemical studies and some speculations concerning the role of these and other compounds in regard to insects. Balick, Furth and Cooper-Driver (1978) compiled a fairly comprehensive list of about 420 insects and mites believed to be herbivorous on pteridophytes. The presence of primitive as well as advanced insects among these arthropods suggested to Balick et a/. (1978) the possibility of co- evolution of arthropods and pteridophytes, both before and after the radiation of angiosperms. The main interest of Balick et a/. (1978) concerned arthropod feeding on Pteridophyta. This will be the first association to be discussed. 30 FERN GAZETTE: VOLUME 12 PART 1 (1979) ARTHROPODS FEEDING ON PTERIDOPHYTA Diverse arthropods, mainly insects and mites, feed on pteridophytes wherever these grow. The insects include representatives of several orders (Table 1). Among the sucking insects, the Hemiptera are dominant. Some examples are the mirid bugs Bryocoris pteridis and Monalocoris filicis, which feed mainly on sporangia (Southwood and Leston, 1959). The whiteflies A/eurotu/us nephrolepidis and Filicaleyrodes williamsi (Mound, 1966), the mealybugs Nesopedronia cibotii (Beardsley, 1971) and Spilococcus filicicola (Hussey, Read and Hesling, 1969) and the aphids /diopterus nephrelepidis and Sitobion ptericolens (Robinson, 1966) all settle on and suck from the fronds. Among the Thysanoptera (thrips), the fern thrips, Leucothrips nigripennis and the gall-making Pteridothrips pteridico/a will serve as examples. Many fly (Diptera) larvae are found on pteridophytes, the Anthomyiid genus Chirosia being restricted to them. Other representatives are the gall midges Dasineura filicina and D. pteridicicola, which form galls on bracken fronds, the Agromyziid Phytoliriomyza pteridii which mines in bracken (Spencer, 1973) and the Drosophilid Drosophila notha induces multiple galls on bracken in New Guinea (Kirk, 1977). Several beetles (Coleoptera) feed on pteridophytes. These include the notorious fern weevil, Syagrius fulvitarsis and its relatives (Marshall, 1922), the pteridophyte-specific Megacolabus (May, 1973), some Chrysomelids (Kirk, 1977) and Poecilips pteridophytae, a Scolytid from New Guinea (Gray, 1970). Many caterpillars of butterflies and moths (Lepidoptera) were collected from Pteridophyta. The Pyralid Samea mul/tiplicalis was tried for the biological control of Salvinia in Africa (Bennett, 1972). The leather®af fern borer, Undulambia polystichalis (Pyraustidae) is a pest of Rumohra adiantiformis in Florida (Short, Driggers, Kuitert and Roberts, 1971) as is the Florida fern caterpillar, Ca//opistria floridensis, a Noctuid (Pirone, 1970). Theichobia verhuellella induces galls on several pteridophytes (Hering, 1937). The Hymenoptera have many pteridophyte-feeders among members of the Selandriinae, including the Equisetum-associated Loderus and Dolerus (Benson, 1962), Blasticotoma filiceti and many Strongy/ogaster spp. (Smith, 1969). Mites reported from Pteridophyta comprise the gall-making Eriophyids TABLE 1 : THE ORDERS OF INSECTS (BASED ON RICHARDS AND DAVIES, 1977). ORDERS WITH WHICH ARTHROPODS ARE KNOWN TO BE ASSOCIATED ARE MARKED BY AN ASTERISK. APTERYGOTA Us 2. 3. 4. Thysanura (Bristle-tails) Diplura Protura Collembola* (Spring-tails) PTERYGOTA EXOPTERYGOTA . Ephemeroptera (May flies) . Odonata* (Dragonflies) . Plecoptera (Stoneflies) . Grylloblattodea . Orthoptera* (Grasshoppers and Crickets) . Phasmida (Stick Insects) . Dermaptera (Earwigs) . Embioptera . Dictyoptera* (Cockroaches and Mantids) . lsoptera (Termites) . Zoraptera . Psocoptera* (Booklice) . Mallophaga (Biting Lice) . Siphunculata (Sucking Lice) . Hemiptera* (Bugs) . Thysanoptera* (Thrips) ENDOPTERYGOTA . Neuroptera (Lacewings) . Coleoptera* (Beetles) . Strepsiptera (Stylopids) . Mecoptera* (Scorpion Flies) . Siphonaptera (Fleas) . Diptera* (Flies) . Lepidoptera* (Butterflies and Moths) . Trichoptera (Caddis Flies) . Hymenoptera* (Wasps, Ants) a GERSON: FERNS AND ARTHROPODS 31 Phytoptus pteridis (Lawton, 1976) and Nothopoda pauropus (Anthony, 1974), the fern mite Hemitarsonemus tepidariorum (Cameron, 1925) and several Tenuipalpids, among them Jenuipalpus lygodii (DeLeon, 1966). A detailed list of insects and mites reported from Pteridophyta was compiled by Balick et a/. (1978). Few other arthropods are known from pteridophytes. Hussey et a/. (1969) reported that the woodlouse Porcellio laevis damages Adiantum roots in greenhouses. Kuhnelt (1976) cited observations on the Diplopods Tauveriu/us and Pteridoiulus, both said to live in pteridophyte rhizomes. Unspecified millipedes (Diplopoda) and sowbugs and pillbugs (lsopoda, woodlice) were reported to feed on the tender new growth of pteridophytes (Hoshizaki, 1975). Horning, Schuster and Grigarick ((1978) collected Tardigrades from Pteridophyta in New Zealand. Feeding may be on any part of the plants. Roots, rhizomes, stems, fronds and spores are eaten. Members of pteridophyte-feeding genera may utilise different parts of the same plants. Thus Chirosia parvicornis mines in bracken frond-tips, C. crassiseta mines the stem, and C. a/b/tarsis mines in both the stem and leaf stems (Lawton, 1976). Most feeding insects (Coleoptera, Hymenoptera, Lepidoptera) have chewing mouth parts; many of them in fact skeletonise fern fronds (Beer, 1955; Swezey, 1921). Others, such as aphids, whiteflies and scale insects, suck out the contents of pteridophyte tissues. Feeding aphids induced circular, chlorotic areas on pteridophyte fronds, chlorosis extending along the midrib (Severin and Tompkins, 1950). As damage extended to newly developing fronds, it was concluded that the causative agent had a systemic nature. There was no evidence that plant viruses were involved (but see below). Pteridophyte feeders comprise species confined to one plant alone (monophagous), others which feed on several species of Pteridophyta but on no other plants (oligophagous) and arthropods which attack pteridophytes as well as higher plants (polyphagous). A special, minor group are some aphids which alternate between Pteridophyta and other angiosperm host plants. Examples are Shinjia pteridifoliae, alternating between bracken (Pteridium) and Viburnum (Miyazaki, 1968) and Aulacorthum pterinigrum, on Pteris and Vaccinium (Richards, 1972). Gall makers are usually monophagous. Several arthropods induce galls on pteridophytes, mites being said to cause more than half of the known galls (Mani, 1964). This author, however, included only mites and Diptera among arthropod gall makers (Mani, 1964; fig. 126), ignoring Hemiptera (Beardsley, 1971), Coleoptera and Thysanoptera (Docters van Leeuwen, 1938), as well as Hymenoptera and Lepidoptera (Buhr, 1964-65). The Pteridophyta appear to have fewer arthropod-incuded galls than other large plant groups (Mani, 1964). Several quite specific insects were found during projects aimed at the biological control of pteridophyte weeds (Bennett, 1966; Wieczorek, 1973). Such specificity is of paramount importance in these projects, the insects therefore undergoing vigorous starvation tests. In other cases, the reported specificity may reflect only lack of knowledge, as most pteridophyte feeders appear to be oligophagous. Feeding on several pteridophyte species has been reported in mites (Beer, 1954; DeLeon, 1966). Thysanoptera (Hussey et a/., 1969), Hemiptera (Mound, 1966; Gosh, 1974; McKenzie, 1967; Southwood and Leston, 1959), Diptera (Wieczorek, 1973), Coleoptera (Kirk, 1977; Swezey, 1921), Lepidoptera (Swezey, 1921; Hering, 1937) and Hymenoptera (Benson, 1962; Smith, 1969). 3 FERN GAZETTE: VOLUME 12 PART 1 (1979) Many of the polyphagous insects which feed on Pteridophyta are pests of agricultural crops (Table 2). While listing these species, it became evident that two superfamilies of plant-feeding Hemiptera, namely the Aphidoidea (aphids) and Coccoidea (scale insects) show a marked difference in specificity to pteridophytes. Among aphids, specificity is the rule, only very few species feeding on other plants also. Pteridophyte-associated scale insects, on the other hand, are by and large not restricted to these plants (see also Eastop, 1973). Even the fern scale, Pinnaspis aspidistrae, has many angiosperm host plants (Dekle, 1976). Primitive arthropods are believed to be more closely associated with Pteridophyta than recent ones. Cooper-Driver (1978) has suggested that the more ancient insect orders (except the Orthoptera) are better represented among pteridophyte feeders than would be expected. This was taken to indicate a prolonged association between pteridophytes and these insect orders. Gall makers in particular are said to have been recruited from the older arthropod groups. Mani (1964) who considered the mites to be ‘‘undoubtedly”’ the oldest group of cecidozoa (gall-forming animals), stated that they are responsible for more than half of all known pteridophyte galls. He further wrote that of the Thysanoptera (thrips), only the more general (and thus presumably older) Terebrantia induce pteridophyte galls. The millipedes (Diplopoda) TJaueriu/us and Pteridoiulus live in pteridophyte rhizomes; KUhnelt (1976) finds this interesting in view of “the great geological age of ferns and diplopods’’. Members of the sub-family Selandriinae (Hymenoptera: Tenthridinidae) are regarded as the most generalised in the family, and the genus Hem/taxonus as especially primitive (Smith, 1969). Members of this genus feed on Pteridophyta. On the other hand, Heptamel/us, a related but highly advanced genus, also lives on these plants (Smith, 1969). As the latter case shows, pteridophytes have also become hosts to arthropods in more recent geological periods. For instance, Docters van Leeuwen (1938) notes that despite the reported antiquity of the Pteridophyta, relatively few galls occur on their older members, most galls being found on what are nowadays called '’Polypodiaceous”’ ferns (Lovis, 1977). Feeding of the Scolytid beetle Poecilips pteridophytae on bracken in New Guinea was believed by Gray (1970) to be of recent origin, due to the highly specialised feeding and the relatively recent geological origin of New Guinea. Occurrence of the endemic Hawaiian mealybug /Nesopedronia on introduced pteridophytes is probably also quite recent (Beardsley, 1971). Hering (1951) concluded that several Dipterous leaf miners have only recently transferred to the Pteridophyta. A special form of feeding is on the ‘nectaries’’ or on sap exudates. Darwin (1877) noted that the secretion of bracken nectaries is attractive to ants. Bees, Elaterid beetles, numerous flies as well as ants were reported by Meikle (1937) to visit these nectaries. Adult sawflies, whose larvae feed on bracken, drink sap exuding from wounded fronds (Beer, 1955). Little is known concerning pteridophyte litter breakdown in the soil through arthropod activity. C. Overgaard Nielsen (in Elton, 1966) reported that the polyphagous millipede G/omeris marginata is an important consumer of bracken litter in Denmark. Harding and Stuttard (1974), reporting on former work, found large numbers of the Oribatid mite Platynothrus peltifer and the springtail Onychiurus procampatus in bracken litter. These authors concluded that .. . “much remains to be investigated concerning the role of microarthropods in the decomposition of litter of pteridophytes and other cryptogams’”’. GERSON: FERNS AND ARTHROPODS 33 ARTHROPODS AS PESTS OF PTERIDOPHYTA Several pteridophytes (Asp/enium bulbiferum, Rumohra adiantiformis and others) are commercially cultivated; arthropods which damage them are plant pests which require control measures. The fern mite, Hem/tarsonemus tepidariorum, lives in the innermost recesses of unopened frond and pinnae, and feeds there. Damage is manifested as minute brown depressions, leaf deformations and stunted and asymetric growth, resulting in dwarfed, weak plants (Cameron, 1930). The fern aphid, /diopterus nephrolepidis, is another destructive pest of glasshouse pteridophytes. Infested fronds curl and turn black (Hussey et a/. 1969). The leatherleaf fern borer, Undulambia polystichalis, has recently become a major pest of Rumohra adiantiformis in Florida (Short et a/., 1971) requiring weekly control measures. Several non-insect arthropods, like Isopoda (sowbugs and pillbugs) and Diplopoda (millipedes) feed on the tender new growth of commercial pteridophytes (Hoshizaki, 1975). Other pteridophyte pests are polyphagous insects which damage many agricultural plants. Some representative, non-specific pteridophyte pests are listed in Table 2. Many of these pests must be controlled by chemicals, a problem aggravated by pteridophyte sensitivity to certain insecticides. Pirone (1970), for instance, advocated using only pesticides of plant origin (like pyrethrum or nicotine), and warned against Organophosphorus compounds. Fluffy ruffle fern, Nephrolepis exaltata, was very sensitive to acaricides like Omite and Plictran, Acarol causing leaf deformations and burns (Short and McConnell, 1973). The chemical control of pteridophyte pests thus poses some special problems. The fern weevil, Syagrius fulvitarsis, invaded the Hawaiian Islands and became a destructive pest of the large Sad/eria cyatheoides there in 1919. Pemberton (1948) discovered that the weevil originated from Australia and found an efficient natural enemy (the Hymenopterous Braconid /schiogonus syagrii) in New South Wales. This parasite was introduced to Hawaii and controlled the pest there. Pemberton collected several other pteridophyte-infesting weevils, belonging to the genera Syagrius and Neosyagrius; these were described by Marshall (1922). The fern weevil also invaded England and Ireland, infesting many Pteridophyta in the Dublin Botanical Gardens (Lloyd, 1944). One way of controlling weevils, as narrated by Lloyd (1944), was by placing infested plants into water, the beetles then floating to the surface. Dozens of weevils and their larvae were thus found to infest single pteridophyte plants. BIOLOGICAL CONTROL OF PTERIDOPHYTA The tables are turned when pteridophytes become weeds and insects are brought in to control them by feeding on them. The prime example of a pteridophyte weed is bracken, which has long been an agricultural pest in various parts of the world (Rymer, 1976). Sa/vinia, on the other hand, has only become a nuisance in tropical and sub- tropical waterways in more recent times (Bennett, 1966). While chemical and agrotechnical measures were being taken, biological control by insects was not neglected. The resultant surveys (Bennett, 1966; Kirk, 1977; Simmonds, 1967; Wieczorek, 1973) have materially contributed to general knowledge concerning pteridophyte arthropods. Of the ten phytophagous insects found on Sa/vinia by Bennett (1966), three, namely the Pyralid Samea mu/tiplicalis, the weevil Cyrtobagous singularis and the Orthopteran Paulinia acuminata, were considered most promising. They were subsequently released at various sites in Central Africa, and P. acuminata became established there (Anders and Bennett, 1975). 34 FERN GAZETTE: VOLUME 12 PART 1 (1979) TABLE 2 : SOME POLYPHAGOUS PESTS WHICH ALSO FEED ON CULTIVATED PTERIDOPHYTA. Thysanoptera Heliothrips haemorrhoidalis Schneider, 1966 Thrips tabaci Pirone, 1970 Hemiptera Trialeurodes vaporariorum Schneider, 1966 Coccus hesperidum Pirone, 1970 Planococcus Citri Hussey et. a/., 1969 Chrysomphalus aonidum Dekle, 1976 Tachycines asynamorus Schneider, 1966 Otiorrhynchus sulcatus Schneider, 1966 Other cases of insect injury to bracken include reduced germination due to spore feeding by deep soil springtails (Collembola) (Conway, 1953), and conspicuous injury to isolated stands by sawflies (Beer, 1955). Balick et a/. (1978) reported that arthropod damage to wild pteridophytes in tropical Mexico may affect the reproductive capacity of these plants. Invertebrates thus have the potential to affect pteridophyte survival in the field. PTERIDOPHYTE RESISTANCE TO ARTHROPODS Notwithstanding all cases of arthropod feeding on Pteridophyta, a concept of intrinsic pteridophyte resistance to insects and mites has found its way into the literature. This concept was formulated in the following quotation: ‘In spite of the similarity of their foliage to that of the flowering plants, ferns do not commonly serve as food plants for insects. They are, in fact, strikingly immune from insect pests of all sorts. This is hardly what might be expected from the long presence of this group of plants, their enormous development in the past, and their persistence at the present time in quite considerable abundance. Why they should be so sparingly selected as food plants does not seem to have been adequately explained’’ (Brues, 1920). Although this generalization was already challenged by Swezey (1921), Brues later (1946) reinforced it. This concept was supported by studies in plant galls (Docters van Leeuwen, 1938: Mani, 1964), by Elton’s (1966) observations on bracken arthropods and more recently by Cooper-Driver (1978). Apparent pteridophyte resistance to grazers was subjected to experimental studies. Soo Hoo and Fraenkel (1964) incorporated Nephrolepis exaltata frond extracts into a diet for the polyphagous moth Prodenia (=Spodoptera) eridania, and reported that this extract contained a water-soluble feeding deterrent. Soon afterwards (Kaplanis, Thompson, Robbins and Bryce, 1967; Takemoto, Ogawa, Nishimoto, Arihara and Bue, 1967) it became known that many Pteridophyta contain ecdysones (insect moulting hormones). These were believed to play a role in pteridophyte defence against herbivorous arthropods (Slama, 1969). Carlisle and Ellis (1968), however, reported that a diet of air-dried bracken fronds did not interfere with the moulting cycle of the desert locust, Schistocerca gregaria. Furthermore, there were indications that these insects do not take up ecdysones from the food into their blood. Hikino, Ohizumi, and Takemoto (1975) reviewed and studied the effect of ingested GERSON: FERNS AND ARTHROPODS 35 phytoecdysones on insects. They concluded that the absorption of these compounds by insects is slow and limited, their excretion rapid, and absorbed ecdysterone is rapidly catabolised into compounds with little or no moulting hormone activity. Jones and Firn (1978) showed that amounts of phytoecdysteroids obtained from bracken were considerably below levels which affected several phytophagous insects belonging to three different orders. These authors believed that ecdysteroids might still act as nontoxic feeding deterrents. Hendrix (1977) offered dried bracken-leaf meal to larvae of the polyphagous moth 7richoplusia ni and reported some inhibition of feeding, which was not, however, considered to be due to phytoecdysones. The effect of prunasine, a cyanogenic glycoside present in bracken, on herbivorous insects was studied by Cooper-Driver and Swain (1976). In the south of England bracken was found to be polymorphic in regard to this glycoside, as well as to its hydrolase. Although 96% of individual plants of most populations contained prunasine and its hydrolase (and were thus cyanogenic), there were a few populations in which most individual plants were acyanogenic. Bracken disks from the various populations were offered to Schistocerca gregaria. Those originating from cyanogenic plants were hardly touched by the locusts, whereas the others, even though they contained the cyanogenic glycoside itself, were eaten to a large extent. The authors concluded that the production of hydrocyanic acid from prunasine probably has a positive role in protecting bracken against herbivores. In a later study (Cooper-driver, Finch, Swain and Bernays, 1977) they showed that when bracken was periodically collected and offered to the locusts, there were two peaks in inhibition, during late May and in late August. The first period coincided with a maximum of cyanogenesis, the second with a peak in tannin production. The importance of these feeding deterrents for general feeders like locusts was thus demonstrated. The level of several possible feeding deterrents produced by bracken during its growth cycle was studied by Lawton (1976). He did not exclude any bracken component, including phytoecdysones, from inhibiting arthropod herbivores. As noted, there is no clear evidence that such hormones actually protect ferns, but their occurrence in pteridophytes provides an unexpected link between these two groups. Lists of phytoecdysone-containing Pteridophyta were presented by Hikino, Okuyama, Sin and Takemoto (1973) and by Russell and Fenemore (1971). As recent studies on pteridophyte-associated arthropods show, these plants actually have a considerable number of herbivores. Lawton (1976) compared the bracken fauna to that of other English plants and showed that the Pteridium- supported fauna is not an improvished one. (See also Lawton and Schroder (1977) in regard to Dryopteris villari’). Further studies will likely provide similar results in regard to other Pteridophyta. Although, as noted, pteridophytes are supposed to deter the feeding of arthropods, Milne (1968) obtained complete development of several springtail species which were given bracken spores as their only nutrient. Balick et a/. (1978) reported that insects appeared to prefer pteridophytes over angiosperms as a food source in tropical Mexico. Another kind of defence mechanism was called the ‘set a thief to catch a thief” principle (Lawton, 1976). Ants which visit pteridophyte nectaries may repel other insects, coming for that or other purposes. Little is known about this mechanism at present, but Bentley (1977) believes that most plants gain some protection from herbivore damage by the ants which visit their nectaries. 36 FERN GAZETTE: VOLUME 12 PART 1 (1979) TABLE 3 : TOXICITY OF SOME PTERIDOPHYTE EXTRACTS TO THREE SPECIES OF INSECTS. (A: Effect of injecting pteridophyte extracts into American cockroaches, Periplaneta americana; 1 = 100% paralysis at 1 day, without recovery; 2 = 100% paralysis at 2 days, without recovery; 3 = 50% mortality at 3 days; 4 = less than 50% morality at 3 days. B and C: Effect of immersion of German cockroaches, Blatella germanica, and milkweed bugs, Oncopel/tus fasciatus, respectively, into aqueous fern extracts for one minute; N = less than 20% average mortality in 4 days). (From Table 1 in Heal et a/., 1950). Pteridophyte Part of Plant Adiantum sp. Entire Anemia mexicana Entire Cheilanthus microphyllus Entire Dryopteris filix-mas Leaves D. marginalis Rootstocks Equisetum arvense Entire (fresh) E. arvense Entire (dry) E. bogotense Entire E. hyemale var. californicum Stems E. robustum Stems lsoetes dodgei Entire Lycopodium clavatum Entire L. complanatum var. flabelliforme Fronds Marsilea vestita Entire Notholaena sinuata Entire Osmunda clay toniana Rhizomes Pellaea ornithopus Entire Polypodium angustifolium Entire P. neriifolium Entire Polypodium sp. Rhizomes Pteridium aquilinum Leaves P. aquilinum Stems P. caudatum Leaves P. caudatum Stalks Selaginella myosurus Entire psa | | sear aies e caeeoe,| ea Pe Parag res, Pe 3 2 3 3 3 1 2 3 4 3 2 4 4 1 3 3 4 2 2 4 2 3} 3 4 2 | | ZZ Zz || ANTI-ARTHROPOD SUBSTANCES FROM PTERIDOPHYTA Heal, Rogers, Wallace and Starnes (1950) assayed extracts from many plants for insecticidal properties. Some of their results, in regard to pteridophyte extracts, are presented in Table 3. It is noteworthy that bracken was not among the most toxic. In a further set of tests, extracts of fewer Pteridophyta were assayed against more insect species. Extracts of Lycopodium annotinum, L. clavatum, L. complanatum var. flabelliforme and L. quadrangulare, as well as Anemia mexicana, Dryopteris marginalis and Marsilea vestita were used. Of several household pests, only the black carpet beetle, Attagenus piceus, was consistently affected. Woollen fabric impregnated with extracts of all Lycopodium spp., Anemia mexicana, and Dryopteris marginalis reduced feeding injury of this pest by 90% or more. Filicin, a drug originating from Dryopteris, gave 90% kill of the aphid Aphis rumicis (Hartzell and Wilcoxon, 1941). Additional information on the insecticidal activity of Dryopteris and other Polypodiaceae was provided by Jacobson (1958). Seaward (1976) suggested that bracken layers placed in Roman dwellings at Vindolanda (north England) produced an insecticidal exudate which adversely affected stable flies. A liquid from boiled bracken fronds was reported to be very effective against rose aphids (Long and Fenton, 1938). Botanists at a European meeting were seen by Rymer (1976) to burn bracken in order to repel nuisance midges. Rhizomes of Pe//aea involuta (crushed in milk) are being applied by GERSON: FERNS AND ARTHROPODS 37 Southern Sotho (Africa) tribesmen to counteract spider bites. They also use a decoction from rhizomes of Po/ystichum pungens as an enema for horse bots (Watt and Breyer-Brandwijk, 1962). ARTHROPOD-TRANSMITTED VIRUSES IN PTERIDOPHYTA Feeding by sucking insects (especially Hemiptera) is often accompanied by transmission of plant viruses. Eastop (1977, appendix 1-11-2) maintains that no aphid- transmitted viruses occur in the Pteridophyta. Nienhaus, Mack and Schinzer (1974), however, suggested that a viral disease of Po/ypodium vulgare and Dryopteris filix-mas may be vectored by aphids. They innoculated the angiosperm test plant, Nicotiana glutinosa, with this virus. Aphids (Myzus persicae, a vector of many plant viruses) were allowed to feed on W. glutinosa for one minute, and then placed onto healthy test plants. These \V. g/utinosa later showed symptoms of the same fern virus. The disease may therefore be transmitted by aphids. PTERIDOPHYTES AND THEIR LITTER AS SHELTER FOR ARTHROPODS Old or dying pteridophytes are inhabited by various non-specific insects in Hawaii (Swezey, 1921). Many beetles, belonging to the families Nitidulidae, Staphylinidae and Curculionidae were reported from insect-damaged pteridophytes by Gray (1970, 1972), none of these beetles apparently feeding on Pteridophyta. Roaches deposit and live in older tuber ferns (Gomez, 1974; Yapp, 1902); the latter author also found a centipede in that habitat. Bracken litter maintains a considerable dampness, allowing arthropods to flourish there (Elton, 1966). Though none of these arthropods appear to be specific to bracken, their numbers may exceed those found in oak litter (Elton, 1966). Certain members of the bracken litter, like the pill millipede, G/omeris marginata, are important consumers of bracken litter (Elton, 1966). Frankland (1966), in her study of bracken breakdown in the soil, noted some mites and Collembola in decaying petioles, but found no evidence that they were actually feeding on bracken. She considered them to be mycophagous, and even to aid in fungal dispersal. Bracken litter arthropods may occur on the plant above ground; the mite Chamobates is one example (Lawton, 1976). The bracken litter fauna may have some value as prey for small vertebrates, like the bank vole, which uses bracken as cover (Elton, 1966). Other small vertebrates also shelter there, and their parasites and inquilines possibly find the litter suitable for host finding and resting. An example is the sheep tick, /xodes ricinus, which completes its life cycle in the humid bracken litter (Nicholson and Patterson, 1976). ANTS AND PTERIDOPHYTA Ants are often associated with pteridophytes. At least two species visit bracken nectaries, where they suck up the abundant secretion and possibly also gnaw shallow excavations (Darwin, 1877). In the tropics many ants nest fortuitously in pteridophytes (Yapp, 1902), but many more constant associations have also been reported. Camponotus sp., for instance, regularly visits the Neotropical potato fern, Solanopteris brunei, although it was never observed to breed there (Gomez, 1974). At a more advanced level, the ants consistently nest within a certain pteridophyte species; this relationship is regarded as symbiotic. Azteca sp. regularly built its nest on and in- touch with the potato fern (Gomez, 1974). The tubers served queen ants to start new colonies, and were also used as brooding chambers, in which eggs were deposited and larvae tended by worker ants. 38 FERN GAZETTE: VOLUME 12 PART 1 (1979) Two Malaysian “myrmecophilous’” Pteridophyta and their ant associates were reported by Yapp (1902). Po/ypodium (=Lecanopteris) carnosa was associated with the ant Crematogaster yappi, and Polypodium (=Lecanopteris) sinuosum with Technomyrmex albipes. Ants entered both plants through openings excavated in large- celled, thin-walled tissue (with apparent water-storing functions), which, upon breaking down, gives rise to the galleries that run throughout the plant. These and other Malaysian Lecanopteris and their ant associates were also discussed by Holttum (1954). Lecanopteris was recently revised by Jermy and Walker (1975), who provided additional data on the accompanying ants, like a Camponotus sp. on L. spinosa. Daniels (1974) briefly referred to /ridomyrmex cordatus, an ant living in the rhizome galleries of Drynaria quercifolia in Australia. Advantages accruing to ants from this relationship are shelter, moisture and carbohydrates present in the plants’ ‘ripe’ parenchymatous cells (Gomez, 1974). Pteridophyta appear to have evolved towards mutualism with ants, as suggested by various anatomical modifications; thin-walled points of entry, ready-made gallleries and carbohydrate-containing tissues. Solanopteris brune/ produces an aldehyde-like substance which attracts the ant Azteca and may serve as an allomone, restricting these ants to the plant. What the pteridophytes gain from this association is not clear. Darwin (1877) rejected the possibility of defence against herbivores, a possibility which Lawton (1976) termed the ‘set a thief to catch a thief’ principle. Such protection is usually ruled out in regard to tropical ant ferns (Gomez, 1974; Jermy and Walker, 1975; but see Bentley, 1977 for a divergent view). Other postulated advantages include imported minerals and nitrogen from ants’ excreta (Holttum, 1954), and CO9 provided by actively respiring ants in the green rhizomes of Lecanopteris spinosa (Jermy and Walker, 1975). Janzen (1974) conducted a comparative study of the relationships between several epiphytes (including Phymatodes (=Lecanopteris) sinuosum) and_ their associated ant, /ridomyrmex myrmecodiae. He thought that ants may prefer epiphytes for nesting as they provide relatively dry cavities of long duration (in the tropics). He also pointed out that epiphyte-ant associations often occur when epiphytes develop on slow-growing vegetation with an insufficient supply of nutrients. Without the ants, which supply their remnants as well as CO9, the epiphytes, including Pteridophyta, may not have been able to survive in low productivity habitats. One such association appears to have implications for a Lepidopteran herbivore, the lLycaneid Hypochrysops theon medocus, whose larvae feed on Drynaria quercifolia. Ovipositing Lycaneid females alight on the undersurface of the fronds and walk about. Eggs are laid only if ants are encountered; otherwise females search other fronds. Larvae feed within rhizome galleries, where they coexist with the attendant ant, /ridomyrmex cordatus, which apparently never molests the larvae (Daniels, 1974). This appears to be another case of Lycaneid-ant symbiosis, a little understood series of associations (Owen, 1971). EPIZOIC SYMBIOSIS A different form of pteridophyte-arthropod mutualism was reported by Gressitt (1969) within the context of ‘’epizoic symbiosis’’. This involves weevils which support extensive plant growths on their backs. The beetles — mostly belonging to the subgenus Symbiopholus in the genus Gymnopholus — are structurally modified to accommodate the plants growing on them. These modifications include dorsal depressions surrounded by ridges, as well as various adapted setae and scales. A sticky secretion, which may promote propagule germination and subsequent growth, is also produced. Most plants found were fungi, algae, lichens and mosses; but a specimen of GERSON: FERNS AND ARTHROPODS 39 Gymnopholus (Symbiopholus) lichenifer had an _ unidentified pteridophyte gametophyte growing on it. The association was believed to be mutualistic; the weevils providing a favourable environment for the plants, the latter serving as protective covering for the beetles (Gressitt, Samuelson and Vitt, 1968). DISPERSAL AND AN ETYMOLOGICAL ASSOCIATION Some herbivores of Pteridophyta, especially those which attain pest status, were probably inadvertently brought to various parts of the world with their host plants by humans. The aforementioned fern weevil, of Australian origin, is one example. Arthropod dispersal of pteridophytes appears to be of minor importance. Ants were seen to Carry spores on their bodies, thus transferring them from tree to tree. Such spore transport was considered by Holttum (1954) to be “‘hardly necessary’’ as wind dispersal of spores was very effective. Thompson (1977) observed that Ap/rocal/us, a New Guinean weevil, sometimes has pteridophyte sporangia attached to it, although the beetles were never collected on cryptogamic plants. Many spore feeders possibly void some viable spores away from the feeding site and thus disseminate them. Finally, on a whimsical note, it might be mentioned that the scientific name for pteridophytes, and the names of the insect subclasses (as well as many orders) (Table 1) have a common root, the Greek word Pteron, a wing. Thus the names Pteridophyta, Apterygota, Pterygota, Diptera, Lepidoptera etc., have all originated from the same word. DISCUSSION The most interesting current problem in regard to pteridophyte-associated arthropods is the extent to which the plants’ defences deter or inhibit insect feeders. The opinions of Brues (1920, 1946), Elton (1966) and more recently Cooper-Driver (1978), were that Pteridophyta have far less than their share of herbivores. The discovery of phytoecdysones in pteridophytes (Kaplanis et a/., 1967), as well as other feeding deterrents (Lawton, 1976; Cooper-driver, 1978) tended to support the hypothesis that Pteridophyta are highly resistant to insect attack (Hendrix 1977; Slama, 1969). However, other, contradictory data were concurrently becoming available. The search for natural enemies of bracken and water pteridophytes disclosed that these plants were actually being attacked by a large and diverse arthropod fauna (Bennett, 1966; Kirk, 1977; Simmonds, 1967; Wieczorek, 1973). An analysis of the structure of an arthropod community on bracken (Lawton, 1976) showed that the size of this fauna fits well within the range of comparative angiosperm-associated faunas. Working in tropical Mexico, Balick et a/. (1978) reported that pteridophytes were apparently preferred over angiosperms as food for insects. On theoretical grounds, there is no reason why the pteridophyte fauna should be depauperated. Strong, McCoy and Rey (1977, and former papers) strongly argue that host-plant range is the most important factor determining the species-richness of its herbivores. Age of a host within a given region was considered to be of minor importance. Smith (1972) concluded that the number of endemic pteridophyte genera is less than half that of the flowering plants and that pteridophyte genera and species, on the whole, are more widely distributed than angiosperms. Upon applying the concept stated by Strong et a/., (1977) to Pteridophyta, it becomes reasonable to expect that these plants may actually have as many, or more, arthropods associated with them as the angiosperns. As to the role of pteridophyte chemical defences, the secondary substances, there is no doubt that they confer some protection upon the Pteridophyta (Cooper-Driver et a/., 1977). However, this by itself does not mean that FERN GAZETTE: VOLUME 12 PART 1 (1979) 40 ZLEL ‘Ae4spoy Pr6| ‘uosisse+-do|say (QepiuoljndsnD :e4aIdO0ajOD) suapnsjui sniubeAS expo | (aepinjoO,Y :esa1dOpiday) sasiwisse e1/og (QePInyIONY :esaidopiday) evedion; eixajdng LOGL “!ueINAO + (2eplulpasyzuay ‘e4aidouawApY) snpunsas 4sajsebo;AGu011g LOGL ‘!ueINAO + (aepiulpasyjzuay :e1a1douawWAY) aepunwiso 4a3se60/Ab6uU0115 (SEp!ulpesyiusy ‘B4aIGOUBWAH) SNII/AOpIgye snuoxewaf (Sepiuipasyiua| :e4a3dOUaW AH) snleZ/gnp snuOxeIWwal} (eepizAwoiby :esaidiq) eyjasejiy eEzAWOIsI/O1AYd (sepiAWwoulUYy :esaidiq) eulaisjzsAy e1sosiyD (sepipiudy :esaidiway) siaijiy CErOYdosOYduie]/) + (eepipiydy :esaidiway) sepunwso snzAwosap 696L “UUWS 9961 ‘uiWS LG6L ‘Gulsay LE6L ‘Guisay OBL ‘IezeAII\ S9OBL ‘HezeAIy epunwso (aepizAwoiby :e4aidiq) ‘ds ezAwoby enjeseyy 9961 aes (aepidjedinual :esy) /pobA/ sndjedinuas 6961 ‘buljseH 7@ peay ‘AassnyH (9eP19905 :esaidiwayH) aeassoo eljassies wnipobAT LZ6L ‘Aa|spseag (aepiss0c0pnasq :esajdiwiay) ‘dds esuospadosayy | sisa7doueszig LL6L ‘Auuadg (aepiai0g :e1aidooal\|) snzanpas snaiog LZ6L ‘Haq\i5 B YOI|AYR ‘Jabuis SE6L “uamnaa] UPA $19190q LOGBL ‘AlZUa>9/\ (aepijeyduAN) :esa1d0piday) spoomjsam e1ydAdNFZ (e4a1diq) sepliAwopisag (9eEp!19D0N0PNas, :esaldiwap) /uesOs snIDOIeUaY, (eapiopiudy :esaidiway) eyoaizasinba siydy (eapiopiudy :esaidiway) /zasinba uolgojs (aepizAwoiby :e1aidiq) ‘dds ezAwolsi7 (aepiuipasyzUa| :esaj}dOUusWAP) !uUl43ajOG ejjauibejas PIGL “UOS||!UUeISsSC VIGL “UOS||!UUeISsC ZL6L pegteds CIBL ‘UOSuag } gaunog_ | snofeudAjog snobeydobijo9 | snobeydouoyy +++ winjasinbZ uoxe} podoiyyy ajyAydopiieig ‘(s}ue\d Bulsamo|j UO pue susajy UO Bulpaay :snobeydAjod ‘susa} uUsapow se |jaM se }uaIdUe UO Hulpaay :snobeydobijo ‘snuab auo uo Ajuo Bulpaaj :snobeudouop\) “SAYOAIGYSH YISHL SO ALIDISIDSdS JHL GNV VLAHdOGIYdLld LNAISNV AWOS : 7 A1EVL GERSON: FERNS AND ARTHROPODS 41 the protection afforded to pteridophytes is much more efficient than that given to angiosperms. The latter are extremely diverse from the chemical point of view, and lumping all flowering plants together in order to compare them with pteridophytes, in regard to number of associated arthropods (Brues, 1920), is not convincing. What may be needed is a comparative study on one or two angiosperm groups with similar numbers of species, distribution and apparency (sensu Feeny, 1976). Plant chemistry alone possibly makes little difference to the total number of insect species which eventually evolve to exploit a given plant (Lawton, in press); or, as stated by Levin (1976), ‘no defence is sacrosanct’. The considerable insect and mite guilds which live on bracken and Sa/vinia suggest that arthropods have indeed evolved mechanisms to circumvent the plants’ protective chemicals. Other often ignored factors in host-plant selection are temperatures and specific habitats (especially in the tropics), as shown by Eastop (1973) in regard to aphids, and the non-chemical (i.e., plant architecture, abundance, seasonality) Components of apparency. There is no cause to exclude the Pteridophyta from such considerations. And yet, some of the available literature tentatively suggests that a distinction could perhaps be made between the ancient Pteridophyta (including Se/aginella and Equisetum) and the modern ones. Although records of ancient pteridophyte-associated arthropods are meagre, the pattern of these associations appears to differ from those of modern Pteridophyta. A difference between the number of animal-induced galls formed on ancient and modern Pteridophyta was noted by Docters van Leeuwen (1938). A listing of arthropods currently known to feed on and live on ancient pteridophytes suggests that most insects and mites which feed on them are either monophagous (restricted to one pteridophyte species or genus), or else polyphagous (subsisting on Pteridophyta as well as flowering plants). Aside from two exceptions (see below), ancient Pteridophyta are only very rarely attacked by arthropods which also feed on modern pteridophytes (Table 4). The exceptions are Osmunda and Todea, both members of the order Osmundales. They are attacked by aphids, sawflies, leaf-mining fly maggots and a weevil (Table 4), all of which also occur on one or more modern Pteridophyta. In so far as further collections confirm these observations, it could be argued that from the herbivore point of view, Osmunda and Todea are more similar to modern pteridophytes than to the ancient ones. According to current ecological concepts, the similarity is probably biochemical in nature. Hikino et a/. (1973), while assaying Japanese Pteridophyta for phytoecdysones, found no activity in most ancient pteridophytes. These included Equisetaceae, Lycopodiaceae, Selaginellaceae, Ophioglossaceae, Marattiaceae, Schizaeaceae and Wymenophyllaceae. Only in Hicriopteris glauca (Gleicheniaceae) and in six members of Osmunda (Osmundales) were phytoecdysones found. The related 7odea also showed insect moulting hormone activity (Russell and Fenemore, 1971). Nothing is known about arthropods of H. glauca, so it will not be further discussed. The Osmundales remain as almost the only group of ancient Pteridophyta with high phytoecdysone activity, and the only group which has herbivores in common with modern pteridophytes. it is tempting to postulate that these phenomena are related. As already noted, available experimental data do not confirm that phytoecdysones act as feeding deterrents in Pteridophyta. This does not imply that they did not affect herbivores in the past; the phytoecdysones may have played an important role in the co-evolution of Pteridophyta and their associated herbivores, but have now become a “redundant defence mechanism’’ (Jones and Firn, 1978). The associations between arthropods and the ancient Osmundales may well have initiated 42 FERN GAZETTE: VOLUME 12 PART 1 (1979) later co-evolutionary processes with modern pteridophytes. A relatively advanced state of Osmundales-arthropod co-evolution is also suggested by the disproportionately high number of insects found on the few extant species of Osmundales, as pointed out by G. Cooper-Driver (in lit.). The comparative studies of Lawton (1976) and Kirk (1977) on bracken arthropods in England and Papua New Guinea, respectively, emphasize the differences in these faunas between temperate and tropical regions. Britten (1881), and Cooper- Driver (1978), among others, noted dissimilarities in the extent of pteridophyte susceptibility to insect attack between temperate and tropical regions, but quantitative data are scarce. And yet over 90% of all extant Pteridophyta grow in the tropics (Manton, 1973) and evolution in tropical regions operates in fundamentally different ways than in temperate zones (Dobzhansky, 1950). Tropical Pteridophyta doubtless carry a multitude of unstudied arthropods (Balick et a/., 1978); relevant studies thus promise to enrich and modify our concepts about pteridophyte-arthropod associations. ACKNOWLEDGEMENTS The work invested in the preparation of this review was undertaken during a sabbatical at the School of Environmental Science at the University of Bradford. It is a pleasure to thank my host there, Dr. Mark R.D. Seaward, as well as Professor M.J. Delany, for their generous hospitality. Professor |. Manton, The University of Leeds, Professor G. Cooper-Driver, Boston University, and Dr. J.H. Lawton, University of York, commented on the manuscript and made some literature sources available to me. Dr. V.E. Eastop, of the British Museum (Natural History) supplied records of fern-associated aphids, and A.C. _ Jermy, of the same institute, offered helpful advice. To all of them | wish to extend my sincere thanks. REFERENCES ANDERS, L.A. and F.D. BENNETT. 1975. Biological control of aquatic weeds. Annu. Rev. Entomol. 20: 31—46. ANTHONY, M. 1974. Contribution to the knowledge of cecidia of Singapore. Gardens’ Bulletin. 27: 17-65. BALICK, MJ., D.G. FURTH and G. COOPER-DRIVER. 1978. Biochemical and evolutionary aspects of arthropod predation on ferns. Oecologia. 35: 55—89. BEARDSLEY, J.W. 1971. New genera and species of Hawaiian Pseudococcidae (Homoptera). Proc. Hawaiian Entomol. Soc. 27: 41—58. BEER, R.E. 1955. Biological studies and taxonomic notes on the genus Strongy/ogaster Dahlbom (Hymenoptera : Tenthredinidae). Univ. 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Hawaiian Entomol. Soc. 5: 57—65. TAKEMOTO, T., S. OGAWA, N. NISHIMOTO, S. ARIHARA and K. BUE. 1967. Insect moulting activity of crude drugs and plants. J. Pharma. Soc. Japan. 87: 1414—1418. THOMPSON, R.T. 1977. A revision of the New Guinean weevil genus Apisroca/us Pascoe (Coleoptera: Curculionidae). Bull. British Mus. (Nat. Hist.) Entomol. 36: 193—280. WATT, J.M. and M.G. BREYER-BRANDWIJK. 1962. The Medicinal and Poisonous Plants of Southern and Eastern Africa. Livingstone, Edinburgh and London. WIECZOREK, H. 1973. Zur Kenntnis der Adlerfarninsekten. Ein Beitrag zum Problem der Biologischen Bekaémpfung von Pteridium aquilinum (L.) Kuhn in Mitteleuropa. Ze/t. angew. Entomol. 72: 337—358. YAPP, R.H. 1902. Two Malayan ‘‘myrmecophilous” ferns, Po/ypodium (Lecanopteris) carnosum (Blume) and Polypodium sinuosum Wall. Ann. Bot. 716: 185-231. 46 FERN GAZETTE: VOLUME 12 PART 1 (1979) REVIEWS SPORES OF INDIAN FERNS by Santha Devi, pp. 228. New Delhi: Today & Tomorrow’s Printers and Publishers. 1977. 280 x 225 mm. Price RS. 75.00 (about £4.50). Dr. Devi starts her book with a brief preview of the history and recent studies on spore morphology and then describes systematically the spores of over 720 fern species in 134 genera. In describing techniques she rightly points out the dangers of only looking at external features with the S.E.M. There follows (pp. 99—122) a comparative analysis of the spore morphology, family by family, following B.K. Nayar’s treatment. Both description and discussion is illustrated by 172 light micrographs of varying quality, which are printed on the kind of paper we have come to expect in Indian publications. This is not Santha Devi’s fault, however, and she must be congratulated for carrying out the considerable amount of work that is needed to produce such a book. She is making a major contribution to fern palynology, for, although it is not apparent from the title of this book, more than half of the spores investigated were obtained from outside India. In fact when one reads Appendix II, which gives the source of spores, a large proportion of them are from cultivated botanical garden material of unknown provenance and some, one might think, of dubious name. A.C. JERMY FERNS OF THE OTTAWA DISTRICT by William J. Cody, Canada Department of Agriculture Publ. 1974, pp. 112: revised 1978. 210 x 133 mm. Available from Printing & Publishing Services, Canada, Hull, Quebec KA1 OS9. Price $3.90 (about £1.60). This is a little fern flora of 48 km (30 miles) radius of the centre of Ottawa — some 2,800 sq. miles; 18 genera, 36 species and one hybrid (Dryopteris x bootii (Tuckerm.) Underw.) are described and dotmaps and photographs (of pressed specimens, but very informative) are given. After the earlier edition, published in 1956 it was thought that little would be added except a few dots on the distribution maps. Nevertheless five new species were found for the area: Asp/enium platyneuron, Pellaea atropurpurea, P. glabella, Woodsia obtusa and W. oregana. |n the text, ecological notes are good and references are given to further reports. A.C. JERMY FERNS OF FLORIDA — AN ILLUSTRATED MANUAL AND IDENTIFICATION GUIDE by Olga Lakela and Robert W. Long, pp. 178; 1976. Banyan Books, Miami, Flo. 33143. Price $10.00 (about £4.70). This is an extremely attractive little book with 117, mostly full page, illustrations (mainly photographs). Many of the species described and illustrated are familiar to British horticulturists and to those interested in ferns. From a botanical point of view, the number of genera (53) is wide and gives a good insight into ferns generally. | thoroughly recommend it whilst it is still available. A.C. JERMY SS FERN GAZ. 12(1) 1979 47 THE PHILIPPINE PLAT YCERIUM GRANDE E. HENNIPMAN, G.J. DE JONCHEERE Rijksherbarium, Leiden, Netherlands and M.G. PRICE University of Michigan, Ann Arbor, U.S.A. ABSTRACT For the first time Platycerium grande (Fée) Presl, a rare, giant staghorn fern from the Philippines, is fully described, based upon cultivated material. INTRODUCTION The two first-mentioned authors remarked in an earlier publication (De Joncheere & Hennipman, 1970) on the confusion existing with regard to the application of the name Platycerium grande (Fée) Presi. Although typified by a Cuming collection from the Philippines, it became commonly attached to a well-known and extensively cultivated Australian staghorn fern. As that element proved to be different from the Philippine type, the Australian taxon had to be given a new name, viz. Platycerium superbum Jonch. & Hennipm. Both these species were regarded as closely related to P. wandae of New Guinea, and also to the new!y described P. ho/ttumii Jonch. & Hennipm. of Thailand and Indo-China. A difficulty encountered when reaching these conclusions was the surprising scarcity of authentic P. grande collections from the Philippines, and the poor quality of the few specimens present in large herbaria. Only one complete fertile leaf of P. grande (the type; BM) was traced. Although the taxonomy of the giant staghorn ferns is primarily based upon differences in shape and size of the foliage leaves, other features of interest such as the rhizome scales, and the shape of the nest leaves are also important and were unknown for P. grande. They could therefore not be taken into consideration earlier. Although our classification was accepted by others, e.g. Hoshizaki (1972), others expressed doubts. We long tried unsuccessfully to obtain a complete living or dried specimen of true P. grande. Obviously this species is very local. Quite accidentally, young specimens of P. grande said to have been collected from the wild in Davao, were traced in 1975 at a local nursery in Cebu, southern Philippines. One of these plants was sent to Leiden and another to Kew. The specimen at Leiden was placed in the tropical greenhouse of the Leiden Botanical Garden, and reached maturity in the summer of 1977. The cultivated material thus available makes it possible now to complete the description of P. grande proper, and to discuss its systematic position within the group of the giant staghorn ferns. THE CULTIVATED MATERIAL OF PLATYCERIUM GRANDE (FEE) PRESL Platycerium grande (Fée) Pres!l, Epim. Bot. (1851) 154; (J. Smith, Hook. J. Bot. 3 (1841) 402, nom. nud.). — Neuroplatyceros grandis Fée, Hist. Acrost. (1845) 103, p.p. — Alcicornium grande Underwood, Bull. Torr. Bot. Club 32 (1905) 594, p.p. — Lectotype: Cuming 157, Philippines, Luzon, Mt. Cristobal (BM). — figs. 1—2. Rhizome short, but massive, the apex densely covered by scales which are basally attached, 4—7 times as long as wide, 13—18 by 2—4 mam, stiff-herbaceous, light brown with the central and basal part somewhat darker coloured, margin with a c. 0.5 mm wide fringe of uniseriate (always?) unbranched multicellular hairs, apex acute. Nest /eaves upright, forming huge basket, sessile, 48 FERN GAZETTE: VOLUME 12 PART 1 (1979) FIGURE 1. Platycerium grande plant in cultivation showing the upper side of the bipartite fertile frond. Ma ee ? FIGURE 2. The same specimen, showing the lower side of the fertile frond, with the two conspicuous soral patches. HENNIPMAN, JONCHEERE AND PRICE: PLATYCERIUM 49 obliquely reniform, 80—90 by 150—180 cm, greyish green, soft-coriaceous, recurved and withered when old; basal part appressed, auriculate, semi-circular (radius c. 45 cm), nearly completely encircling the growing point and with a conspicuous, thin, projecting frill up to 8 cm long around the petiole base of the foliage leaves; the lateral margins entire, wavy; upper part lobed, lobes spreading, 3—5 times dichotomously divided, 15—25 cm wide at the base, 40—50 cm long, sinuses narrow, ultimate segments rather coarse with blunt tips. veins anastomosing with prominent veins running in pairs to the lobes, otherwise dichotomously forked up to the ultimate segments. Fertile leaves produced in pairs, c. 120 by 150 cm, fleshy-coriaceous, greyish green, stalk to 4 cm long, woody, striated, in cross-section elliptic, 1.5 by 1.0 cm, gradually widening distally; lamina pendent, symmetrically divided into two equal parts, separated by a long narrow sinus; each part c. 85 by 130 cm and c. 20 cm wide at the cuneate base (20—30 by 40—50 cm) which largely covered by a soral patch measuring c. 15 by 20—30 cm, distally divided into strap-like segments by 3—5 oblique, inwardly directed, dichotomous divisions, the lowest sinuses dilated, ultimate segments up to 25 by 2.5 cm; veins anastomosing with some prominent veins running parallel to the margin. Indument: leaves on both sides thinly covered by a felt of whitish, + sessile stellate hairs 300—600 um in diam., with 8—11 slender arms. Sporangia stalked, pyriform, 300—500 um long, with 18—22 indurated cells; spores bean-shaped, somewhat dented, 55—80 by 35—45 um, brown, smooth; paraphyses stellate, stalked, 150—200 pm in diam., with 9—11 arms. ' Notes. 1. An earlier remark (De Joncheere & Hennipman, |.c.) that the size of this species is smaller than that of its relative P. superbum is incorrect; the size of the specimens in cultivation at Leiden and Kew + equals that of P. superbum and P. ho/ttumii. 2. The rhizome scales are similar to those of P. ho/ttumii being flat (not + bullate as in P. superbum), and lacking a so-called midrib (present in P. wandae). Also, the marginal fringe of hairs on the scales is similar to that of P. ho/ttumii, differing from that of both P. superbum and P. wandae (hairs few-celled). 3. In our cultivated specimens, mature nest leaves of P. grande differ from those of P. holttumii; being less deeply cut, of a greyer colour, of a softer texture, and with less conspicuous veins. Further, the main veins in the nest leaves of P. grande run up to the ultimate lobes, in P. holttumii they run up to the penultimate lobes only. DISCUSSION AND CONCLUSIONS P. grande is a distinct species easily recognized by the shape of the fertile leaves. Those formed on our cultivated plant are similar in shape to the type specimen at BM, although much larger. This confirms the treatment of P. grande as a separate species. By its fertile leaves P. grande is easily distinguished from its relatives P. ho/ttumi/ and P. wandae (fertile leaves with two strongly unequal halves, each with one soral patch), and P. superbum (fertile leaf with one soral patch only). The rhizome scales of P. grande turned out to be similar to those of P. ho/ttumii, and different from P. superbum and P. wandae. The sterile leaves of P. grande show conspicuous differences from those of P. holttumi/. A discussion on the phylogenetic relations within the genus will be included in a forthcoming paper on the genus Platycerium by Mr. M.C. Roos, a graduate student from Leiden University. ACKNOWLEDGEMENTS It is a pleasure to express our gratitude to Mr. A. Mulder, who skilfully handled this beautiful ornamental fern in horticulture. The photographs were made by Mr. B.N. Kieft. REFERENCES JONCHEERE, G.J. DE, and E. HENNIPMAN. 1970. Two new species of Platycerium and the identification of P. grande (Fée) Presl. Brit. Fern Gaz. 10: 113—118. HOSHIZAKI, BARBARA JOE. 1972. Morphology and phylogeny of Platycerium species. Biotropica 4: 93—117. 50 FERN GAZETTE: VOLUME 12 PART 1 (1979) REVIEW EVOLUTIONARY PATTERNS AND PROCESSES IN FERNS by J.D. Lovis in Advances in Botanical Research, 4: 229-440, 1977 (ed. H. Woolhouse). Academic Press 228 x 155mm. Price: £16.80. This substantial paper is a review of up-to-the minute literature presented with a depth of knowledge and critical appraisal that few, if any, pteridologists could surpass. It is divided into six chapters. After an Introduction, Prof. Lovis reminds the reader of a number of trends seen in the fossil record (pp. 232-255). There follows a short introduction to the development of cytological studies in ferns followed by the largest chapter (pp. 258-319): cytology in relation to classification and phylogeny. Lovis accepts a single class (Filicopsida) and includes at subclass level the Psilotales as suggested by Bierhorst (1971); Osmunda and the Eusporangiates are given similar sub- class status. Within the Filicidae are nine orders containing 23 families, the Marsileaceae with chromosome numbers 10—20 are placed next to Adiantaceae with a base number 29 or 30, a situation which seems sound. Three families, Hymenophyllop- sidaceae, Loxsomaceae and Plagiogyriaceae are regarded as incertae sedis. At the subfamily level those accepted are similar to Crabbe et a/ (1975) except the ‘aspidiaceous’ group are placed under Dryopteridaceae and the latter authors are rightly criticised for not segregating Onocleoideae. A full list of genera and chromosome numbers is given with levels of ploidy, and number of species both known and counted. The gaps are most interesting and will hopefully lead to further work. Polyploidy in ferns is discussed in relation to systematic and geographical distribution. The rest of the work (pp. 330—400) is given over to a detailed account of the biosystematics of ferns describing as the basis for such work, genome analysis, species interrelationships in temperate genera: Asplenium, Cystopteris, Dryopteris, Polypodium and Polystichum; and tropical complexes: Adiantum caudatum, Asplenium aethiopicum and Ceratopteris. The role of introgression, breeding systems and apomixis generally is considered in the process of evolution. There are over 500 references and author and subject indices. This is a work that should be read by all taxonomists, especially pteridologists. Every word is carefully weighed, precise and meaningful, be it criticism or praise, and all points of view are fairly given. The reader is left in no doubt, however, of Professor Lovis’ own assessment. REFERENCES BIERHORST, D.W. 1971. Morphology of Vascular Plants. New York. CRABBE, J.A., JERMY, A.C. & MICKEL, J.T. 1975. A new generic sequence for the pteridophyte herbarium. Fern Gaz., 17: 141-162. A.C. JERMY FERN GAZ. 12(1) 1979 51 A FURTHER CHROMOSOME COUNT FOR CHRISTENSENIA (MARATTIALES) T.G. WALKER Department of Plant Biology, University of Newcastle upon Tyne ABSTRACT A chromosome count of n=40 is recorded for a diploid plant of Christensenia from Sarawak. The only previous determination showed a Solomon Islands representative to be tetraploid. The base number of x=40, so characteristic of the Marattiales, is thus confirmed. INTRODUCTION Christensenia is a small member of the Marattiales and whilst it shares many of the highly distinctive features of this order, it stands well apart from the other members in its palmately-divided fronds with round synangia scattered on the lower surface and with reticulate venation. In some systems of classification (for review and discussion see Pichi Sermolli, 1959, 1977) these differences have been considered sufficient to justify the erection of a separate family, Christenseniaceae (= Kaulfussiaceae). CYTOLOGY Christensenia occurs in Assam, Malaysia, through much of Indonesia and the Philippines, with a noteworthy extension of its range to Melanesia reported by Braithwaite (1977). One of the plants from San Cristobal in the Solomon Islands was cytologically investigated by Braithwaite and found to have n=80, 2n=-160. From these chromosome counts he concluded that he was dealing with a tetraploid plant based on x=40. This assumed base number is in conformity with the x:40 found in other members of the Marattiales that have been cytologically examined, namely Ang/opteris (Manton and Sledge, 1954; Mehra and Singh, 1955; Ninan, 1956; Ghatak, 1962), Danaea (Walker, 1966) and some species of Marattia* (Walker, in Manton, 1959; Walker, 1966). Whilst a member of the Royal Geographical Society Expedition to Gunong Mulu in Sarawak in the summer of 1978, | collected several plants of Christensenia for cytological fixation. Of these only one proved suitable for the purpose but it gave numerous cells clearly showing 40 pairs of chromosomes at meiosis. One such cell is illustrated in fig. 1. This count of n=40 thus confirms Braithwaite’s deductions as regards both the basic chromosome number of the genus and of the tetraploid nature of his plant in contrast to the diploid status of my Sarawak specimen. Christensenia has been considered to consist of a small number of species but Copeland (1947) states that these all may better be regarded as forms of the one species C. aesculifolia (BI.) Maxon. The presence of at least two cytotypes has now been demonstrated but whether or not these are recognizable on morphological grounds requires further sampling. ECOLOGY AND CULTIVATION All my specimens were gathered from several localities in lowland alluvial forest in or near the Gunong Mulu National Park in Sarawak and were growing in fairly deep shade in permanently damp or wet alluvial silt at c. 65 m (c. 200 ft) above sea level. Holttum (1954) records that it has been found up to 3000 ft altitude in Malaya. Although Schneider (1893) declares that this species (under the name Kau/fussia aesculifolia) is easy to grow in cultivation this has not been my experience. Plants sent in the past to Newcastle have not proved easy to establish and those that survived for a time were preferentially grazed by slugs which attacked and destroyed the growing apices. 52 FERN GAZETTE: VOLUME 12 PART 1 (1979) FIGURE 1. Meiosis in diploid Christensenia aesculifolia (B\.) Maxon from Sarawak (T13319) showing 40 bivalents, x 1000. Permanent acetocarmine squash preparation. ACKNOWLEDGEMENTS My thanks for financial assistance are due to the Research Fund of the University of Newcastle upon Tyne. REFERENCES BRAITHWAITE, A.F. 1977. A chromosome count and range extension for Christensenia (Marattiaceae). Amer. Fern J. 67(2): 49—50. BROWNLIE, G. 1961. Additional chromosome numbers of New Zealand ferns. Trans. Roy. Soc. New Zeal. Bot. 1: 1—4. COPELAND, E.B. 1947. Genera Filicum. Chronica Botanica, Waltham. GHATAK, J. 1962. Observations on the cytology and taxonomy of some ferns from India. Nucleus 5: 95-114. HOLTTUM, R.E. 1954. Flora of Malaya, II. Ferns. Government Printing Office, Singapore. MANTON, }!. 1959. Cytological information on the ferns of West Tropical Africa. /n A.H.G. Alston, The flora of West Tropical Africa, 2nd ed.: 75—81. Crown Agents, London. MANTON, | & SLEDGE, W.A. 1954. Observations on the cytology and taxonomy of the pteridophyte flora of Ceylon. Phil. Trans. R. Soc. (B) 238: 127—185. MEHRA, P.N. & SINGH, H.P. 1955. Cytology of Cyatheaceae, Woodsiae and Marattiaceae. Curr. Sep24 2425. NINAN, C.A. 1956. Studies on cytology and evolution of the pteridophytes |. Observations on the Marattiaceae. J. Indian Bot. Soc. 35: 233-239. PICHI SERMOLLI, R.E.G. 1959. Pteridophyta. /n W.B. Turrill (Ed.), Vistas in botany: 421, 493. London. PICH!| SERMOLLI, R.E.G. 1977. Tentamen pteridophytorum genera in taxonomicum ordinem redigendi. Webbia 37(2): 313—512. SCHNEIDER, G. 1893. The book of choice ferns. Vol. 2. Gill, London. WALKER, T.G. 1966. A cytotaxonomic survey of the pteridophytes of Jamaica. Trans. R. Soc. Edinb. 66: 169—237. WALKER, T.G. 1973. Evidence from cytology in the classification of ferns. /n A.C. Jermy, J.A. Crabbe and B.A. Thomas (Eds.). The phylogeny and classification of the ferns. Bot. J. Linn. Soc. 67, Suppl. 1: 91—110. *Footnote. The existence of x=39 in other species of Marattia has been confirmed (Ninan, 1956; Brownlie, 1961). This number has doubtlessly been secondarily derived from x=40 (Walker, 1966, 1973). FERN GAZ. 12(1) 1979 53 PELTATE SCALES IN SACCOLOMA G.B. NAIR Institut fur Systematische Botanik, Universitat Zurich, 8008 Zurich, Switzerland ABSTRACT Peltate scales are described in Sacco/oma elegans KIf. INTRODUCTION In ferns, peltate scales are mainly confined to essentially epiphytic groups. According to Alston (1956) ‘‘peltate scales seem to be found only in Polypodiaceae, Grammitidaceae and Davalliaceae’’. They occur also in terrestrial representatives of essentially epiphytic groups, but to my knowledge their occurrence is entirely terrestrial groups has not been reported before. This account is to report the presence of peltate scales on the rhizome and petiolar base of the terrestrial neotropical fern Saccoloma elegans K\|f. The genus Sacco/oma was established by Kaulfuss in 1820. On account of its pouch-shaped indusia it was usually associated with, or even combined with, Daval/ia, where peltate scales are of regular occurrence. At present, Sacco/oma (with its segregate genus Orthiopteris) is usually placed near Dennstaedtia and its relatives (e.g. Copeland, 1947: 49; Holttum, 1955: 305— 306; Tardieu-Blot, 1958: 14; Mickel, 1973), although it seems somewhat distinctive because of its scaly rhizome, the other genera of the Dennstaedtia alliance having consistently uniseriate, pluricellular dermal appendages usually called hairs (Nair, 1973). The presence of peltate scales in Sacco/oma was observed in the course of monographic work now in progress. MATERIALS AND METHODS These observations are based on study of the following specimens: Saccoloma elegans KIf. ECUADOR: Provincia Esmeraldes, Playa Rica, Parroquia de concepcion, Ynes Mexia 8429 (US, UC, U, NY, G, B). BRAZIL: Minas Gerais, I!heu, Ynes Mexia 4694 (UC, US, NY, GH); Vicosa, Ynes Mexia 4639 (US, NY, B, U, G); Amazonia, Territorio do Roraima, G.T. Prance et a/. 13611 (NY, U, US). GUYANA: S. Pakaraima Mts. Kopinang Falls, Bassett Maguire et a/. 46064A (NY, U, US). Dermal appendages were carefully removed after flooding the dried specimen with wetting agent for 15-20 minutes, washed, and mounted in lactic acid for observation. OBSERVATIONS The growing regions of the rhizome and the young circinately coiled fronds are covered by numerous scales. Hairs, however were found to be completely absent from the rhizome (cf. Copeland 1947: 49) but present on the young frond. Only soft- textured, multicellular, non-glandular hairs (5-12 cells long) have been found in Saccoloma. |n Dennstaedtia, on the other hand, both multi-cellular acicular or multi- cellular glandular and non-glandular hairs have been observed in all the species so far studied (Nair, 1973). In Saccoloma, the scales at the apex of the rhizome are densely crowded. The majority are peltate in structure, and these remain intact far from growing region on the parts of the thizome that are buried in the soil. Among such peltate scales, variously modified other scales of smaller size occur on the growing parts of the rhizome (‘basifixed scales’). These often show some degree of lateral expansion giving a cordate appearance near to the scale base (fig. 1: d,f & 54 FERN GAZETTE: VOLUME 12 PART 1 (1979) SS NAIR: PELTATE SCALES IN SACCOLOMA 55 h). Both types of scales occur abundantly also on the basal parts of the stipe. Mature peltate scales are 5-12mm long and 3-7mm wide at the broadest region. They are linear-lanceolate to lanceolate in shape. Sometimes the scales are smaller in size and almost circular in outline (fig. 1g). On the older parts they are dark brown in colour and on the younger parts light brown. The proximal end of some peltate scales shows a v-shaped insertion (fig. 1 : a & c), except when well-developed (fig. 1 : e & i). The stalk is dark brown, firm, 0.1-0.2mm long. The persistant scale bases (stalks) later develop into often backwardly pointed, firm, spine-like structures, particularly on the basal half of the stipe. The distance from the base of the point of attachment of the scale may vary considerably. The cells of the scale are uniforly thick in the central portion of the scale and the marginal cells pale and thin. The scale margins often bear lateral appendages of varying length (fig. 1 j). The distal part of the scale is frequently branched, their tips ending in a uniseriate row of cells (fig. 1 k-m). The cells of the marginal appendages are indistinguishable from the rest of the cells. The terminal cells of the branches contain a granular substance of unknown chemical nature. ACKNOWLEDGEMENTS | am greatly indebted to Professor K.U. Kramer for his suggestions, criticism and encouragement. | am also grateful to Professor C.D.K. Cook for reading the manuscript and to the Directors of herbaria from whom material was borrowed. REFERENCES ALSTON, A.H.G. 1956. The subdivision of the Polypodiaceae. Taxon. 5 (2) : 23—25. COPELAND, E.B. 1947. Genera Filicum, Waltham Mass: Chronica Botanica. HOLTTUM, R.E. 1955. A revised flora of Malaya, II. ('1954'') Ferns of Malaya. Singapore: Government Printer. MICKEL, J.T. 1973. The classification and phylogenetic position of the Dennstaedtiaceae. In A.C. Jermy etal. The Phylogeny & Classification of the Ferns. Journ. Linn. Soc. Bot 67: (Suppl.) 135-144. NAIR, G.B. 1973. Studies in the morphology and anatomy of some Dennstaedtioid ferns (Ph.D. thesis Kalyani University, India). TARDIEU-BLOT, M.L. 1958. Polypodiacées, in: H. Humbert, Fi. Madag. Com. Paris. FIGURE 1. Scales of Saccoloma elegans: a & c, peltate scales with incised base; b, peltate scale with an undivided base; d & f, basifixed scales; e & i, the proximal portion of a peltate scale showing point of attachement; g & h, basifixed scales from a circinately coiled frond; j, portion of a peltate scale showing the marginal appendages; k-m, distal portions of peltate scale showing the terminal granular cell and branchings of the scales. 56 FERN GAZETTE: VOLUME 12 PART 1 (1979) SHORT NOTES A NEW NAME FOR A EUROPEAN DRYOPTERIS The name Dryopteris pseudomas (Wollaston) Holub et Pouzar is illegitimate, its basionym of 1855 being predated by Nephrodium affine Lowe. A different taxon, “Dryopteris affinis’” Newman, was never validly published at the specific rank, but only as a variety, so the following new combination is made and replaces the name Dryopteris pseudomas: - Dryopteris affinis (Lowe) Fraser-Jenkins, comb. nov. Basionym : Nephrodium affine Lowe, Tr. Camb. Phil. Soc. 6 : 528 (1838). The type from Madeira in Key (K!) represents the atlantic European diploid taxon, common also in western Britain. C.R. FRASER-JENKINS British Museum (Natural History) London. DRYOPTERIS HYBRIDS : A CORRECTION Due to the temporary loss of Fraser-Jenkins’ original manuscript the name Dryopteris x sarvelae was inadvertently misseelt as Dryopteris x sarvelii (Fern Gaz. 11(5) : 339 (1977), as has been pointed out to us by Mr. Sarvela. The following new country records for D. x ambroseae were also omitted : Romania, Austria, Czechoslovakia, Sweden, Norway, North Turkey, West and Central Caucasus (all from collections of Fraser-Jenkins deposited in BM). C.R. FRASER-JENKINS & A.C. JERMY British Museum (Natural History), London. EQUISETUM VARIEGATUM NEW TO TURKEY The presence of Eguisetum variegatum Schleich ex Weber & Mohr has recently been confirmed from Turkey from a herbarium specimen (Davis & Coode 38422, E) collected in 1962 in Kastamonu Province, North side of Ilgaz Dag, which had been formerly regarded as E. ramosissimum Desf. in Flora of Turkey, vol. 1 : 32 (1965). The occurrence of this plant in north central Turkey (N. Anatolia) adds significantly to the range of E. variegatum, the nearest other stations to which occur in scattered localities high in the main Caucasus mountains of the USSR. This new Turkish station represents the most southerly known Eurasian outpost from the extensive northern circumboreal range of this species. This typically high-altitude streamside horsetail occurs at about 1350 m, in an area of Ab/es forest on the northern slope of the highest mountain in this part of Turkey, where mist and rain is frequent. It is of interest that the area also includes a number of other outlying Euro-siberian or circumboreal elements, such as all the Turkish species of Pyro/a, Orthilia secunda and Moneses uniflora (all Ericaceae) (P.H. Davis, personal communication). C.NSPAGE Royal Botanic Garden, Edinburgh. 7... ——— EEE, ————eE eer” rerererorrrereae SHORT NOTES 57 EQUISETUM X TRACHYODON IN WESTERN SCOTLAND For more than 100 years, Equisetum x trachyodon A. Br. (= E. mackaii Newm. of older literature, the presumed hybrid between &. hyemale L. and E£. variegatum Schleich. ex Web. & Mohr), has been known in Scotland only as a single station” on the banks of the River Dee in Kincardineshire (VC 91), where it was first reported growing in the mid 19th-century (Brichan, 1842). It occured in patches along several miles or river bank, with both parents nearby. —. x trachyodon is known to have still grown in the Kincardineshire locality in 1935 (Matthews, 1940), and is believed to have persisted into the 1950’s (J.G. Roger, personal communication), although more recent searches (since 1971, by the author and A.C. Jermy) have failed to find it. The correct identity of the Deeside plant is confirmed by herbarium material housed at Edinburgh (E). It is therefore of interest that in the last few years, no less than three further stations for undoubted Equisetum x trachyodon have been confirmed in Scotland. None seems to result from recent spread or formation of the hybrid. All appear to be long-established colonies which have been hitherto unrecognised as hybrids. All three are remote from the previously-known station and all occur in the Hebrides: one on the island of South Harris (VC 110), one on the Isle of Rhum (VC 104) and one in the north of the Isle of Skye (VC 104). Records for all three have been included in the ‘Atlas of British Ferns’ (Jermy et al., 1978), and the purpose of this note is to add some field observations and diagnostic details. Subsequent to these finds, E x trachyodon has been confirmed from a similar habitat in England, and is discussed by Barker (1979) (see below). The first of the new Scottish finds was that in South Harris, at a site to which my attention had been drawn by Dr. W.A. Clarke of Newcastle University, who had found, some years earlier, a horsetail resembling £. variegatum, but about the identity of which some doubts had been expressed at the time. With instructions, | was able to refind the exact locality near Nisabost in the field in June 1972, and established that this horsetail was indeed E. x trachyodon. At this South Harris locality the hybrid forms a colony several square metres in extent. Most shoots of the plant are little larger than those of typical E. variegatum in dune-slack habitats, and most are similarly decumbent for much of their length, becoming ascending or erect only towards their tips. These plants could clearly easily be overlooked for a form of E. variegatum. The colony occupies the steep ESE slope of a gradually eroding dune of calcareous shell sand, and the habitat is kept continually moist by percolating fresh water. The site is a maritime one, no more than 2-4 m above normal high-tide level and scarcely 2 m horizontal distance from the sea. Although sheltered from exposure to the west, it is unprotected in order directions, and the hybrid seems likely to receive some Salt spray, at least intermittently. The seepage water originates from acidic peaty moorlands, but doubtless becomes calcium-enriched as it percolates through the shell- sand of the dunes before emerging at the site of the hybrid. The slight erosion of the slope by the slowly seeping water keeps the habitat both a moist and a more or less continually open one, colonised only by this horsetail which continually pioneers the site. *An old Loch Tummel, Perthshire, locality given in the ‘Atlas of British Ferns’ as an open circle (Jermy, et al., 1978) remains unconfirmed and doubtful. 58 FERN GAZETTE: VOLUME 12 PART 1 (1979) The Isle of Skye station on the river Hinnisdal was discovered in November 1974 by Mrs. C.W. Murray, who sent a specimen to Edinburgh for confirmation. The site is possibly that from which an old record for E. variegatum exists, and the shoots of the E. x trachyodon are only a little larger than those of robust riverside forms of E. variegatum and could easily have been confused with this species, or taken for a small plant of E. hyemale. As with the Harris station, this Skye site is an essentially open one, with the hybrid occurring on two portions of collapsed bank of a small river, about half a mile (c. 1 km) from the sea and 25 ft (c. 8 m) altitude. The hybrid grows at the edge of the river in a basic sandy silt derived from basalt and the site is permanently moist. The habitat is kept open by gradual erosion by river water, which intermittently floods it (C.W. Murray, personal communication). The Isle of Rhum station was first determined in 1975 from a herbarium specimen collected by Dr. C. Ferreira in November 1960, and housed in the herbarium of the Nature Conservancy Council in Edinburgh. The specimen originated in an area of low-lying slack in machair sand at Kilmory dunes in the north of the island, near the mouth of a small river, about 150 m distance from the strand-line of the sea and about 1 m above the spring-tide level. The plant is probably under some influence of saline groundwater at least intermittently, but is also flushed by fresh-water seeping from sandstone slopes above, keeping the habitat more or less permanently moist. The seepage water flows through dunes of shell-sand and thus, as in the Harris station, doubtless becomes considerably calcium-enriched before entering the slack, the shell- sand soil of which also contains a thin organic horizon. Although partially sheltered from the sea by surrounding slightiy higher ground, the habitat is nevertheless rather exposed and most shoots of F. x trachyodon are nearly prostrate on the sand (C. Ferreira, pers. comm.). A further specimen collected in July 1978 by Dr. C. Ferreira from near this former site, but in Ammophila fixed-dune machair, also proved to be E. x trachyodon. Here too the plant is reported to be locally frequent and spreads into adjacent moist-slack grassland, associated with Festuca rubra, Bellis perennis, Lotus corniculatus and Galium verum. The hybrid thus appears locally widespread and well- established in this habitat. It is significant that in each of these new Scottish stations, &. x trachyodon occurs in open sites where one of both parent species are absent, at least in the immediate vicinity. Near the Harris station, E. variegatum is known in a sand-dune slack about 4 km away, but &. hyema/e remains unknown in the whole of the Outer Hebrides. In Skye, both £. variegatum and E. hyemale are present on the island, but both are rare, and absent near to the hybrid site. In Rhum, both parents are recorded for the island, but only E. variegatum grows near the E. x trachyodon. Specimens from the Harris and Rhum stations have somewhat smaller shoots (mostly less than 20 cm high and less than 2 mm stem diameter) than those from Skye and Deeside (which reach 20-30 cm in height and up to 3 mm stem diameter), but are otherwise morphologically similar. The size differences probably result im part from the more exposed habitat of the Harris and Rhum colonies, as well as their probably involving a smaller dune ecotype of F. variegatum, rather than a larger river-side one, in the parentage of the hybrid. The following points of diagnosis may help in location of possible further sites for this hybrid. As seen at these stations, E x trachyodon is best distinguished from its parents by the following characters. The sheaths typically have a much broader black band (occupying most of the upper 1/2 to 2/3 of each sheath), and end in teeth which are long, dark, slender and flexuous and have only a very narrow white margin to their slightly dilated bases. The teeth normally persist either indefinitely on the shoot, or at SHORT NOTES ag least for a time after the shoot has fully expanded (depending on degree of exposure of the habitat), but as they become lost, break along preformed abscission zones at the top of the fused portion of the sheath to leave an irregularly crenate sheath margin. This latter character contrasts strongly with both parents: in F. variegatum the teeth are almost always persistent, in E. hyemale they are almost always shed during the actual expansion of the shoot (during which process they become torn off to collectively form small pagoda-like caps on the tips of the shoot). The shape of the teeth, the broadness of the black band and the absence of broad white scarious teeth margins thus distinguish most shoots of E. x trachyodon even from large specimens of E. variegatum, whilst the persistence of the teeth, as well as absence of the ash-grey coloured band around the upper sheath margin, more clearly biangulate internodal ridges and teeth which are clearly 2-ribbed, separate most shoots of E x trachyodon even from small specimens of &. hyema/e. In the hybrid, sheaths are longer in relation to their width than in either parent. Cones, where present, contain only highly mis- shaped abortive spores, thus confirming its hybrid status. Herbarium specimens from each of these three localities are deposited at R.B.G., Edinburgh (E). REFERENCES BARKER, M. 1979. Equisetum x trachyodon in Cheshire, new to the English flora. Fern Gaz. 12: 59—60. BRICHAN, J.B. 1842. Description of Equisetum hyemale, Mackaii and variegatum, as found on the Banks and in the bed of the River Dee; with additional observations. Phyto/ogist 7: 369— So Tr: JERMY, A.C., ARNOLD, H.R., FARRELL, L.A. & PERRING, F.H. 1978. Atlas of Ferns of the British Isles. London. MATTHEWS, J.R. 1980. Equisetum trachyodon as a Scottish plant. Trans. and Proc. Bot. Soc. Edinb. 33: 29—32. NEWMAN, E. 1844. A History of British Ferns and Allied Plants. London. C.N. PAGE Royal Botanic Garden, Edinburgh EQUISETUM X TRACHYODON IN CHESHIRE, NEW TO THE ENGLISH FLORA Some specimens of Equisetum, recently sent to C.N. Page, were determined as the hybrid Equisetum x trachyodon A. Br. (Equisetum hyemale L. x Equisetum variegatum Schleich. ex Web. & Mohr). These specimens came from the Cheshire Conservation Trust’s Red Rocks Nature Reserve and the adjoining Royal Liverpool Golf Course at Hoylake, Merseyside, (v.c. 58, Cheshire). Here, the E. x trachyodon grows on the leeward side of a slightly brackish marsh at the base of some ‘yellow’ sandhills, which are still reached by blown sand from the foreshore. The colony is extensive and abundant in rough grassland at the base of the sandhills, and stretches for 1.3 km south from the Red Rocks. The dominant vegetation of the habitat is a turf of Festuca rubra L. and Ammophila arenaria (L.) Link. It is the density of the Ammophiletum rubrae community which effectively limits the growth of &. x trachyodon at this site. Consequently, it is most vigorous where this community is sparse or where the ground has been recently burnt. In such bare sandy situations associated plants are: Agrostis stolonifera ., Myosotis ramosissima Rochel., Ga/ium verum L., Rubus caesius L., Rose pimpinellifolia L. and Taraxacum officinale agg. (predominately 7. brachyglossum (Dt.)Dt.). 60 FERN GAZETTE: VOLUME 12 PART 1 (1979) The morphology of &. x trachyodon at Red Rocks varies within the population. In the north of the site there is a form resembling E. variegatum (and this is where the only pure &. variegatum, in a small quantity, grows at Red Rocks), while in the south it resembles &. hyemale more closely. Dr. Page has commented that the Red Rocks E. x trachyodon is unusual in that it branches and that these branches, because of their small size, have a morphology close to that of E. variegatum (and in isolation could easily be confused with it), although the main stem exhibits the characters of E. x trachyodon. This colony is of particular interest because it is the first station found in England. Elsewhere in Britain, it has a mainly Atlantic distribution, in North and Western Ireland and Western Scotland (Jermy et al., 1978; Page, 1979 — see above). This new site extends the range of the species now to the coast of north-west England. The other notable feature of the colony is that, apparently, only one of its parents, E. variegatum, still grows with it. In both Floras of Cheshire (Newton, 1971; Warren, 1899) E. variegatum is recorded from Wallasey on the North Wirral coast and along the Dee estuary to Gayton (which includes the Red Rocks site). It is also recorded in the earlier Liverpool Floras from the North Wirral coast (Hall, 1838; Dickinson, 1851; and Liverpool Naturalists Field Club, 1872.) However, the other parent, E. hyema/e was first recorded at Red Rocks in 1968 by A. Newton (Newton, 1971). The voucher specimen for this record has not been obtainable and recent intensive searches have failed to find E. hyemale at Red Rocks. The only verified record for E. hyemale from the Wirral is from, ‘sandhills, Heswell Cheshire’, collected by Brown in 1871 (LIV), a site 7 km south of Red Rocks, along the River Dee shore. The 19th century Liverpool Floras (Dickinson, 1851; Liverpool Naturalists Field Club, 1872), also record E. hyemale from Parkgate and Thurstaston (sites which are also south of Red Rocks along the Dee shore) but neither of these records are supported by herbarium specimens. Recent searches of all these localities, have failed to find E. hyemale, but it is possible that it may yet be found in the area. ACKNOWLEDGEMENTS | am indebted to Mr. & Mrs. E.F. Greenwood, who first introduced me to the population of Equisetum at Red Rocks and for their subsequent help, and to Dr. H.A. McAllister and Dr. C.N. Page for their advice. REFERENCES DICKINSON, J. 1851. The Flora of Liverpool. HALL, T.B. 1838. A Flora of Liverpool. JERMY, A.C., ARNOLD, H.R., FARRELL, L.A. & PERRING, F.H. 1978. Atlas of Ferns of the British Isles. B.S.B.1. & B.P.S. London. LIVERPOOL NATURALISTS FIELD CLUB. 1872. The Flora of Liverpool. NEWTON, A. 1971. Flora of Cheshire. Cheshire Community Council Publications Trust Limited, Chester. PAGE, C.N. 1979. Equisetum x trachyodon in western Scotland, Fern Gaz. 12: 57—59. WARREN, J.L. 1899. The Flora of Cheshire. Ed. S. Moore. London. MARIAN BARKER 9 South Drive, Irby, Wirral, Merseyside SHORT NOTES 61 A SECOND BRITISH RECORD FOR EQUISETUM x FONT-QUERI, AND ITS ADDITION TO THE ENGLISH FLORA In June 1978, an unfamiliar Equisetum was noticed by one of us (R.H.R.) on the site of a disused railway near Evesham, Worcs., v.c. 37. At a first glance it looked like a somewhat slender &. te/mateia Ehrh., but closer inspection showed that cone-bearing shoots were present at this time of the year. These cone-bearing shoots had numerous whorls of long, green branches and seemed almost identical with the vegetative shoots. No unbranched, fertile stems, such as are characteristic of eo te/mateia, could be found despite a prolonged search. The plant appeared to answer closely to the description of E. x font-queri Rothm., first reported for the British Isles by Page (1973). A representative sample of freshly-gathered material was sent to Edinburgh, where its identification was confirmed (by C.N.P.) as the second British station for Equisetum x font-queri Rothm. (Equisetum palustre L. x E. telmateia Ehrh.), and the first for England (the previously known British station being on the Isle of Skye). The Worcestershire site on which the hybrid Equisetum grows is a disused railway embankment by the River Avon meadows near Evesham. A comparison between the Evesham plants of £&. x font-queri and those known from Skye, shows the existence of strong overall similarities, yet some interesting differences in detail. The overall shoot outline of the Evesham plants is very similar to that of the Skye plants and distinctly intermediate between those of E£. pa/ustre and E. te/mateia. Thus the outline is slender, and tapers in most specimens to a long terminal branchless portion in the upper 10 cm or so of the shoot (see illustrations in Page 1973) — this is probably the best feature to look-out for in spotting plants of this hybrid. The overall size of the Skye and Evesham plants is similar, both being much larger than E. palustre (reaching 60-70 cm in the Evesham plants and about 65 cm in the Skye . ones). In both Evesham and Skye material the whorled branches are abundant throughout the middle two-thirds of the shoot, and are quite strongly ascending (inheriting this feature from the &. pa/ustre parent). Both have about 8-12 ridges to the main stem. The Evesham plants however differ from the Skye ones in having even more conspicuously white internodes, and longer free teeth to the main stem sheaths, and in these characters the Evesham plants approach the condition found in &. telmateia more than do those from Skye. However, the Evesham plants also have less distinctly bi-angulate and more rounded branch internode ridges, and less strongly 2- ribbed teeth to both the main stem and branch sheaths than do the Skye plants, and in these characters the Evesham plants approach nearer to the condition of &. pa/ustre than do the Skye ones. Despite these small morphological differences, there can be little doubt that the Evesham plants, like the Skye ones, are the result of crossing between E. pa/ustre and E. telmateia. The differences, if not environmentally induced (which cultivation alongside the Skye plants at Edinburgh will determine), might either reflect differences in the relative genetic dominance of the particular characters in the original parents, or might result from the cross having occurred in the other direction, or a combination of both phenomena. In the Evesham locality, the single colony of the hybrid forms a dense stand for a distance of about 75 metres on both sides of the top of a railway embankment, which is capped by a layer of ballast composed of coarse limestone chippings. As the habitat is entirely man-made, the origin of the hybrid Equisetum is problematic. It seems rather unlikely that it could have arisen on this site, for neither of the putative parents occurs anywhere near it at present. The ten-mile section of the old L.M.S. railway from Evesham to Alcester was opened in 1866 (Evesham Journal, July 14, 62 FERN GAZETTE: VOLUME 12 PART 1 (1979) 1866). The remaining section from Alcester to Redditch and Birmingham was opened later. It is most likely that the material used to build the embankment came from the large cutting between the embankment and the station in Evesham (B.G. Cox, pers. comm. 1978). The cutting was made through a ridge of well-drained soil and hence this too seems a most unlikely source of origin for the Equisetum. A more likely explanation is that a piece of rhizome was introduced with the ballast brought in at a later date, as the material making up the embankment became consolidated. This implies that another colony of E. x font-queri existed (and perhaps still exists) elsewhere in Worcestershire, or in one of the adjoining counties. The Evesham to Redditch railway was closed in 1963 (Cox 1977: 109), and parts of the disused track to the north of the embankment have been sold. But the embankment itself, which at this point is about 4 metres above the level of the ground on either side, has so far remained undisturbed. The dense and vigorous stand of the hybrid in the Evesham locality, in a habitat which does not appear particularly favourable to the growth of either E. pa/ustre or E. telmateia, lends support to the observation already made on the very large colony on the Isle of Skye (Page 1973) that this hybrid is very vigorous-and, once established, is capable of considerable vegetative spread even into habitats in which the occurrence of the parents would be unlikely. Specimens of &. font-queri from Evesham have been deposited at BM, E, and P. REFERENCES COX, B.G. 1977. The Book of Evesham. Chesham. PAGE, C.N. 1973. Two hybrids in Equisetum new to the British flora. Watsonia 9: 229—237. R.H. ROBERTS 51 Belmont Road, Bangor C.N. PAGE Royal Botanic Garden, Edinburgh OBSERVATIONS ON LIGHT RESPONSE IN MARSILEA CRENATA Ferns, like other plants, are phototropic — their fronds tend to turn towards the direction of greatest illumination. In addition, different species have distinct preferences for particular conditions of light and shade. Marsilea crenata Presl is a small aquatic fern which grows in full sunlight, rooting in the mud of rice fields, ponds and irrigation ditches. Where illumination is restricted, at the edge of a rice field or by competing plants, the fronds lean in the direction of maximum light intensity. During the wet season the rhizome is long-creeping, producing fronds at intervals of 4-6 cm, but during the dry season it is short-creeping, producing small fronds close together. The lamina consists of two close pairs of pinnae arranged in a square at the apex of the stipe. Laminae of submerged plants either float on, or project above, the surface of the water. The following observations have been made in Malaysia. In the field, under conditions of full illumination, the stipes are either vertical or lean with the flow of the water. The pinnae of projecting mature laminae lie in a horizontal plane during the middle of the day. Early or late in the day the lamina tilts to face towards the sun. When the light intensity decreases towards sunset, or when a heavy storm approaches, the pinnae fold together and the closed lamina assumes a nodding position. At dawn, or when the storm has passed, the lamina re-opens. SHORT NOTES 63 The daily variation in water temperature in rice fields can be more than 20°C, but this appears to have little or no effect on the opening and closing of the lamina, except that at temperatures greater than 40°C the stipes tend to wilt and the laminae then close. Both open and closed laminae were observed throughout the range of temperatures from 23° — 39°C. In order to study the sensitivity of fronds of Marsi/ea crenata to light intensity and direction of illumination under more controlled conditions, plants were taken from a wet rice field near Rembau in Malaysia and grown in an aquarium tank. The rhizomes were planted at random, their elongating tips facing in all directions. Light intensities were measured with a photographic light meter and converted to foot- candles, using the formula quoted in Hoshizaki (1976): 20 (f) @ cS where f = aperture in f stop, T = shutter speed in seconds, and S = film speed in ASA units. The position of the tank was changed from time to time and the various responses were recorded. The following observations were made: 1. During the day the stipes of mature fronds assumed a position pointing in the direction of illumination. The open laminae were at right angles to this, tilted towards the east in the morning and gradually changing their angle through the day until, in the early evening, they tilted towards the west. 2. Towards sunset, when the intensity of illumination decreased to 294 foot- candles, the laminae started to close (fig. 1 a-b). As the light dimmed, first the upper pair of pinnae gradually moved together, their upper surfaces facing. Then the lower pair started to close around them. Movement originated at the point of attachment of each pinna to the stipe apex. Finally, just before it was fully closed, the lamina changed position. Fronds facing both the elongating tips of their rhizomes and the direction of illumination during the day nodded in that direction. Fronds facing away from the elongating tips of their rhizomes either nodded in its direction or took up a vertical position. The process of closing, which took place over a period of 1-1.25 hours, was complete before sunset. 3. At dawn, when the light intensity reached 294 foot-candles, the laminae started to open again. First the lower or outer pair of pinnae moved apart, then the upper or inner pair, followed by tilting of the lamina towards the sun. Opening was complete after 1-1.25 hours on a normal sunny day. 4. As storms approached during the hours of daylight and the light intensity dropped below 294 foot-candles, the lamina closed as at night. The length of time taken depended on the speed of reduction of the light intensity. As soon as the storm passed and illumination increased again, the laminae re-opened. When prolonged heavy rain caused the light intensity to remain low, the laminae stayed closed until the following day. The critical light intensity for this response thus appears to be of the order of 294 foot-candles. 5. Placing the aquarium tank in a dark cupboard overnight delayed the opening of the lamina until such time as it was replaced in sunlight. The period taken for the lamina to open was dependent on the intensity of the sunlight when the tank was replaced. Removal of the tank from darkness to very bright sunlight, about 4500 foot- candles, reduced the time taken to half an hour. 6. Removal of the tank into the dark cupboard during the day also caused the laminae to close. They re-opened when the tank was replaced in sunlight. This effect was repeated several times inoneday. | Foot candles = 64 FERN GAZETTE: VOLUME 12 PART 1 (1979) FIGURE 1. Lamina of Marsilea crenata (X 2 approx.): a, upper pinnae folded together; b, lower pinnae enclosing upper pair and the whole assuming a nodding position. (Photos C.J. Piggott) 7. The artificial illumination provided by ordinary electric lights was insufficient to cause the laminae to remain open at dusk, but by placing a powerful spot-light close to the tank it was possible to cause the laminae to re-open during the hours of darkness. They closed again when the spot-light was withdrawn. 8. During the observation period, new fronds were produced regularly. Very young fronds started to uncoil facing towards the tip of the rhizome. As elongation took place, the stipes twisted so that the fronds uncoiled facing the direction of illumination. 9. The water level in the tank was raised until several of the open laminae were floating on the surface. At dusk, or when placed in the dark, the smaller floating laminae remained open. Those with larger pinnae managed to overcome the effect of surface tension and close. Sometimes only one, two, or three, of the four pinnae were able to assume the folded position. Thus, where the laminae are not projecting but are floating on the surface of the water, only those fronds with large pinnae are able to overcome the surface tension and close their laminae. 10. Some small plants with short-creeping rhizomes were collected from the dry bank of an trrigation ditch and added to the tank. The small fronds of these dry season plants showed little response to changes in direction of illumination or to changes in light intensity. As the now submerged rhizomes elongated and produced larger fronds, the laminae became sensitive to such changes. REFERENCE HOSHIZAKI, B.J. 1976. Fern Growers Manual. New York. A.G. PIGGOTT 29, Jalan Dungun, Damansara Heights, Kuala Lumpur 23-05, Malaysia. THE FERN GAZETTE Original papers, articles, or notes of any length on any aspect of pteridology will be considered for publication. Contributions will be particularly welcomed on floristic, biogeographical and ecological aspects of ferns and their allies, worldwide. Members are especially encouraged to make use of short notes for reporting pteridophyte field observations and records. All may be illustrated with black and white photographs of good technical quality and line drawings. Contributions should be sent to: C.N. Page, Royal Botanic Garden, Edinburgh, U.K. The /ast date for receiving notes and articles to make the following early summer number is: 30 November each year Authors should follow the general style of this number. Close adherence to the following notes will help to speed the publication of contributions. NOTES FOR CONTRIBUTORS Manuscripts: Copy should be in English and submitted in double-spaced type with adequate margins, on one side of the paper only. Abstract: All papers, other than short notes should include a short abstract, to be set at the head of the main text, indicating the scope of the topic and main conclusions. Headings and sub-headings: These should follow the style of this number. (Primary sub-headings are centred capitals. Secondary sub-headings U and L case side roman. Tertiary sub-headings, if necessary, are U and L case side italic.) Numbering of sub-headings should be avoided. Latin names: Quote the authority at (usually) the first mention only, in the main text but, unless unavoidable for clarity, not in the title. All latin names should be underlined throughout the typed copy. I/lustrations: Any number and combination of line and half-tone illustrations (original drawings or diagrams in ink, or black and white photographs of good technical quality) can be included with a manuscript where these help to augment or amplify the text. Photographs should be of the required magnification or larger and need not be made up into full page plates. Each drawing or photograph should be marked on the back with details of author and figure number, and the top edge clearly marked “‘top”’. Figure numbering: Grouped illustrations should follow the numbering system, fig 1a, fig 1b, fig 2a, fig 2b, fig 3, etc. Figure numbers should be applied to illustrations in pencil only or on a transparent overlay. Final lettering will be added by the editor in a style and size consistent with the journal. Figure captions: Type on a separate sheet from the manuscript — include any necessary details of magnification as submitted (the editor will apply any correction for reduction). Reference lists: Please follow closely the style of this number to speed publication. Lists in other styles may have to be returned to authors for re-typing. Reprints: Twenty-five reprints are supplied free of charge to authors, who may order in advance further reprints which will be supplied at cost (plus postage) if requested at time of returning the first proofs. BOOKS FOR REVIEW Books for review in the Fern Gazette or B.P.S. Bulletin should be sent to A.C. Jermy, Botany Department, British Museum (Natural History), Cromwell Road, London SW7 5BD. THE FERN GAZETTE VOLUME 12 PART 1 1979 CONTENTS Page MAIN ARTICLES The Killarney Fern, Trichomanes speciosum, in Wales — R.H. Roberts | 1 Preliminary investigation of two south-west England populations of the Asp/enium 5 adiantum-nigrum aggregate and the addition of A. cuneifolium to the English flora — C.N. Page & Frances M. Bennell An ecological survey of the ferns of the Burren, Co. Clare, Eire — A. Wilimot 9 The association between pteridophytes and arthropods — Uri Gerson 29 The Philippine Platycerium grande — E. Hennipman, G.J. de Joncheere & M.G. Price 47 A further chromosome count for Christensenia (Marattiales) — 7.G. Walker 51 Peltate scales in Sacco/oma — G.B. Nair 5S SHORT NOTES A new name for a European Dryopteris — C.R. Fraser-Jenkins 56 Dryopteris hybrids : a correction -- C.R. Fraser-Jenkins & A.C. Jermy 56 Equisetum variegatum new to Turkey -- C.N. Page 56 Equisetum x trachyodon in western Scotland — C.N. Page 57 Equisetum x trachyodon in Cheshire, new to the English flora — Marian Barker 59 A second British record for Equisetum x font-queri and its addition to the English flora 61 — R.H. Roberts & C.N. Page Observations on light response in Marsilea crenata — A.G. Piggott 62 REVIEWS The eightieth birthday of R.E. Holttum 28 Spores of Indian ferns . 46 Ferns of the Ottawa district 46 Ferns of Florida — an illustrated manual and identification guide : 46 Evolutionary patterns and processes in ferns 50 (THE FERN GAZETTE Volume 11 Part 6 was published 15th January 1979) Published by THE BRITISH PTERIDOLOGICAL SOCIETY, c/o Department of Botany, British Museum (Natural History), London SW7 5BD. Printed in England by he COMPASS COLORPRINT **:* Telephone: 01-571 0991 MSO ISSN 0308—0838 | THE FERN GAZETTE VOLUME TWELVE PART TWO 1980 THE JOURNAL OF THE BRITISH PTERIDOLOGICAL SOCIETY President Vice Presidents _ General Secretary Treasurer Membership Secretar os | Meetings Secretary Editors: Fern Gazette ; 5 al Bulletin os ‘ tereent . , see 7] = British Fern Distribution Recorder i ee , oe. of Ast mn 3 Spore Exchange Organiser a ee : Committee ee JA abe. Ay K. , ee, ee Bo 3 AW. eres ay The Fern Gazette and the Bulletin are 2th The Society has a miele eNbeninie eae hela gar amateur and professional, in Britain and overseas. Ar for received from, and reflect the interests of, pane meee ae : cigist ee ; Membership is open to all rhtereakedit in petits aud: fern-allies, worl Y 1st January each year) are: Full Personal. Members £4; Personal Gazette £3; Student Members (full-time students) £3; Subscrit membership should be sent to the Membership: Secretary (ad details can be obtained. (Remittances made in currencies hi cover bank conversion charges.) Sie iM at Ae: Back ndnibars of the Gazette and Bulletin are available for purchase above), from whom further details can be obtained. 3 ; . FERN GAZ. 12(2) 1980 65 HISTORICAL NOTES ON WOODSIA ILVENSIS IN THE MOFFAT HILLS, SOUTHERN SCOTLAND J. MITCHELL 22 Muirpark Way, Drymen, by Glasgow. ABSTRACT The main part of the account, which is based on literature sources, traces the historical background to the discovery and subsequent virtual extirpation of the rare fern Woodsia ilvensis in the Moffat Hills, Southern Scotland, during the second half of the 19th century. In 1977, an investigation was carried out by the author on behalf of the Nature Conservancy Council into the past and present status of Woodsia i/vensis (L.)R.Br. in the Moffat Hills, Southern Scotland. It was found that the initial examination of literature sources gave not only the required list of published records, but also a fascinating insight into the background to the fern’s virtual extirpation from its former principal station in the British Isles. The discovery of a rare plant or animal species occuring abundantly at a hitherto unknown locality, only to be brought to the brink of extinction within a few years by over-collecting is an only too familiar story. Britain’s native ferns have been no exception to this form of wanton destruction, and the few scattered populations of W. ilvensis suffered worse than most. The beginning of what was almost the end for this rare fern in the north-eastern portion of Dumfriesshire (v-c. 72) can be precisely, dated to 1848, the year of completion of the Caledonian Railway linking Beatock (2 miles from Moffat) with Edinburgh, Glasgow and Carlisle. Moffat’s hotel proprietors vied with one another for the immediate influx of visitors, many of who came seeking relief from their aches and pains at the famous but up to then rather remote local mineral wells, meeting each incoming train with horse-drawn omnibuses to convey prospective guests to their respective establishments (Turnbull, 1871). It was one such visitor to the area, a botanist named Stevens, who chanced upon W. //vensis ‘in considerable abundance’ at White Coomb in the nearby Moffatdale Hills (Stevens, 1849). In the following year (1849) a second station for the Woodsia was found just north of Moffat at the Devil’s Beef Tub (Oliver, 1850). With several additional localities being discovered in the district during the next few years, including a ravine near Loch Skene (Newman, 1854), Hartfell (Balfour, 1858) and the head of the Carrifran Burn (Sadler, 1857), the Moffat Hills became recognised as the headquarters for W. i/vensis in the British Isles. Steven's disclosure that W/. i/vensis was apparently to be found in some quantity at Moffat could not have been more unfortunately timed, coinciding as it did with an impending wave of ‘Pteridomania’ or ‘Fern fever’ that was to grip Victorian Britain for at least two decades from the late 1840s. The middle-class’s obsession with obtaining and cultivating hardy native ferns proved insatiable, the unprecedented demand resulting in the emergence of a new type of fern dealer/nurseryman who scoured the countryside for much of his stock (Allen, 1969). In the Moffat area, the effect of the more selective trade in wild ferns was felt first at the Devil’s Beef Tub, for, within three to four years, professional collectors had completely eradicated all the W/oodsia plants from this readily accessable site (Keddie, 1854). Fern dealers’ activities in the district evidently continued, shown by an article that appeared in the Moffat Register and Annandale Observer on 6th October 1860 accusing York nurseryman James Backhouse (not mentioned by name, but see Backhouse, 1860) of unscrupulously 66 FERN GAZETTE: VOLUME 12 PART 2 (1980) uprooting a number of W. //vensis plants and disposing of them to his clients at half-a- guinea a time (Sadler, 1860). It is probable this indignant outburst drawing attention to the high monetary value being placed on the fern did not have quite the effect intended, for Hope (1929) states that the local hill shepherds (seemingly not slow in sensing a remunerative side-line), joined in the gathering of Woodsia plants for sale. Despite the inclusion in the fern nurserymen’s catalogues of W. //vensis imported from abroad, plants of British origin continued to sell at relatively high prices throughout the 1860s. Collecting WV. i/vensis from the wild may have been profitable but was not without its dangers, for several lives were apparently lost in its pursuit on the precipitious cliffs of Glyder Fawr in North Wales (Step, 1908). In the Moffat Hills, the treacherous broken rocks above the Carrifran falls almost claimed a future Curator of the Royal Botanic Garden, Edinburgh, as he unsuccessfully attempted to obtain a tuft of W. ilvensis just beyond his reach (Sadler, 1860). Nurserymen and gardeners were not alone in casting covetous eyes on this attractive miniature fern, for good specimens were much in demand for private and institutional herbaria. Such was the appeal of the rare WV. //vensis to collectors, it was to hold a permanent place in the Botanical Exchange Club’s ‘List of Desiderata’ from the 1850s right up to the turn of the century, contributors to the scheme being encouraged to submit a minimum of ten specimens for distribution to other club members. To judge from the few surviving sheets of W. //vensis housed in University and Museum herbaria (see Rickard, 1972), the peak period of collecting from the Moffat Hills for ‘scientific purposes’ occured in the 1850s. Visiting botanical parties were often extremely large, and it is recorded that one such well attended excursion led by Professor J.H. Balfour in July 1856 gathered W. //vensis ‘in considerable quantity’ (Balfour, 1858). Amongst the proliferation of books on British ferns that appeared in the 1850s and 1860s was a fairly recent innovation popularly known as a ‘Fern Album’. These albums consisted of annotated sets of pressed ferns from named localities in various parts of Britain and elsewhere. The ‘Moffat Fern Album’ (Carruthers, 1863), or under its correct bibliographical title The Ferns of Moffat, first made its appearance in 1863. Written by Dr. W. Carruthers, a native of Moffat employed at that time as Assistant Keeper in the Department of Botany at the British Museum, this unsigned descriptive work appears to have been issued with the specimen pages left blank for the visiting fern enthusiasts to insert their own self-collected examples. It is likely however, that enterprising local entrepreneurs would take on the task of filling-in the blank spaces with specimens, in order to re-sell the completed or part-completed album at a suitably increased price to the less energetic or invalid visitor. Of five copies of the Moffat Fern Album examined by the present writer, two contained pressed specimens of W. ilvensis on the appropriate page. Under intense collecting pressure from locals and visitors alike, Moffat’s once thriving population of W. //vensis — estimated at ‘hundreds of plants’ in 1856 (Scott- Elliot, 1896) — was eventually reduced to just one known solitary tuft by 1891 (Johnstone, 1893). Recriminations inevitably followed, and in The Flora of Dumfriesshire (Scott-Elliot, 1896) the author firmly places the blame on ‘the ravages of the Innerleithen Alpine Club’. On the face of it this accusation seems a little unjust, as the club was not formed until 1889, long after most of the damage is known to have been done. It is possible however that the members were responsible for administering the coup de graCe to a remnant colony of W. i/vensis at the head of the Midlaw Burn, near Loch Skene, when the Innerleithen Alpine Club visited the upper glen in strength in 1889 and again in 1894, returning with ‘vasculum crammed’ and ‘stuffed vascula, MITCHELL: WOODSIA ILVENSIS HISTORICAL NOTES 67 quite pleased if not happy with the spoils’ (Todd & Young, 1895). What appears to have been the last recorded incident in the whole sorry story involved G. Claridge Druce, a leading field-botanist of the period, when in August 1909 he was shown the supposed one remaining tuft of W. i/vensis in the Moffat Hills (Druce, 1910). Druce wished the sole surviving plant well for the future, but could still not resist helping himself to a few fronds before taking his leave. Looking back with the advantage of hindsight, the almost complete extirpation of W. i/vensis from its British headquarters in the Moffat Hills was inevitable. As Raven & Walters (1956) so aptly describe the fate of the now extinct colony of W. //vensis in upper Teesdale, North-East England — it ‘suffered the misfortune of too much publicity’. To those charged with attracting and sustaining the lucrative tourist trade to Moffat during the second half of the 19th century, the local ferns (in particular the rare W. ilvensis) would have seemed an obvious natural resource to be promoted in much the same way as the alleged healing properties of the local mineral waters had been for the previous hundred years. Successive editions of the Moffat guide book referred in glowing terms to the choice ferns and wild flowers that were to be readily found in the surrounding countryside. Even when the current enthusiasm for collecting and cultivating ferns eventually waned, Moffat’s publicists appeared reluctant to accept the change in Victorian taste. The Moffat Fern Album was still being advertised for sale in 1894 and as late as 1902 the 5th edition of the guide book was suggesting that W. //vensis (‘much valued by collectors’) might still be found at the Devil’s Beef Tub, although it was known to have been collected-out from this particular spot fifty years earlier. Yet by this time the fern craze had been over for a good number of years, exemplified by a leading nurseryman’s catalogue price for the once much sought after W. //vensis having slumped to a shilling. With the death of veteran Moffat botanist J.T. Johnstone in 1922, the last personal knowledge of the exact where-abouts of some of the former WW. //vensis sites in the district appears to have been lost. Nothing more was seen or heard of the fern in the Moffat Hills until 1954, when a small population of about 25 plants was found by D.A. Ratcliffe while undertaking a botanical survey of the upland area (Ratcliffe, 1959). Eighteen years later, a second small colony was turned-up by M.H. Rickard during a field meeting of the British Pteridological Society in 1972 (Dyce, 1972). Curiously, the re-discovery of W. i/vensis in the Moffat area again coincided with an awakening of public interest in the collecting and growing of ferns. In the last few years a number of popular books and articles on cultivating ferns have been published, some of them even advocating a revival of the once discarded and almost forgotten 19th century Wardian cases and bell-glasses as a solution to the modern fern-grower’s problem of centrally-heated flats and houses. It is of regretable necessity therefore, that the precise localities of the two recently found W. i/vensis colonies in the Moffat Hills must remain a closely guarded secret for at least the foreseeable future. ACKNOWLEDGEMENTS | would like to express my appreciation for the assistance given in tracing published literature on the occurrence of Woodsia i/vensis in Moffatdale to the library staffs at the Nature Conservancy Council Headquarters for Scotland, the Royal Botanic Garden, Edinburgh, and the Ewart Library, Dumfries. Also to John H. Harvey and Jeannette Ratcliffe for readily making available the results of their researches into early horticultural catalogues specialising in ferns. It is a particular pleasure to thank Dr. Derek Ratcliffe, not only for the original suggestion of drawing together this historical account, but for kindly reading through the initial draft. 68 FERN GAZETTE: VOLUME 12 PART 2 (1980) REFERENCES ALLEN, D.E. 1969. The Victorian Fern Craze. London. BACKHOUSE, J. 1860. Botany of the Breadalbane Mountains. The Phytologist 4(NS) : 65—67. BALFOUR, J.H. 1858. Notice of a Botanical Trip to Moffat in July 1856. Transactions of the Botanical Society of Edinburgh 5 : 166—169. CARRUTHERS, W. 1863. The Ferns of Moffat. Moffat. DRUCE, G.C. 1910. Plants of some Southern Scottish Counties. The Annals of Scottish Natural History 30 : 39—43; 95—102. DYCE, J.W. 1972. Report of a field meeting at Teesdale and Moffat, 21—30 July 1972. British Pteridological Society Newsletter No. 10, pp 11—15. HOPE, M.I. 1929. Report of a field meeting at Loch Skene, 25 July 1928. History of the Berwickshire Naturalists’ Club 26 : 306—308. JOHNSTONE, J.T. 1893. Notes on the Flora of Moffat District for 1891. Transactions of the Dumfries & Galloway Natural History & Antiquarian Society 8(NS) : 18—21. KEDDIE, W. 1854. Moffat: Its Walks and Wells. Glasgow, Edinburgh & London. NEWMAN, E. 1854. A History of British Ferns. London. OLIVER, D. 1850. Note on Serratula tinctoria, Alsine stricta, and a species of Woodsia. The Phytologist 3 : 775. RATCLIFFE, D.A. 1959. The Mountain Plants of the Moffat Hills. Transactions of the Botanical Society of Edinburgh 37 : 257 — 271. RAVEN, J. & WALTERS, M. 1956. Mountain Flowers (New Naturalist Series). London. RICKARD, M.H. 1972. The distribution of Woodsia ilvenis and W. alpina in Britain. British Fern Gazette 10 : 269 -- 280. SADLER, J. 1857. Narrative of a ramble among the Wild Flowers on the Moffat Hills in August 1857. Moffat. SADLER, J. 1860 A Botanist’s ramble up Moffatdale. The Moffat Register and Annandale Observer (Saturday 6th October 1860). SCOTT-ELLIOT. G.F. 1896. The Flora of Dumfriesshire. Dumfries. STEP, E. 1902. Wayside and Woodland Ferns. London & New York. STEVENS, W. 1849. Notes on the Flora of Dumfriesshire. The Phytologist 3 : 390—393. TODD, J. & YOUNG, T. 1895. Principal Excursions of the Innerleithen Alpine Club during the years 1889—1894. Galashiels. TURNBULL, W.R. 1871. History of Moffat. Edinburgh. FERN GAZ. 12(2) 1980 69 POLYPODIUM MACARONESICUM AND P. AUSTRALE : A MORPHOLOGICAL COMPARISON Rh. ROBERTS 51 Belmont Road, Bangor, Gwynedd ABSTRACT Some of the morphological characters of Po/lypodium macaronesicum Bobrov have been studied and a comparison with those of P. australe Fée has shown that several of them can be used to separate these taxa unequivocally. In particular the characters of frond serration, shape of the rhizome scales, the size and branching of the paraphyses and the structure of the sporangium have been found useful to differentiate between them. The recognition of P. macaronesicum as a species distinct from P. australe is considered to be justified. INTRODUCTION Polypodium macaronesicum Bobrov occurs on the Atlantic islands of the Azores, Madeira and the Canaries (Page, 1977; Ward, 1970). It looks superficially like a well- grown P. australe Fée with larger, proportionately broader fronds and more coarsely serrated pinnae. The Azorean plants have been referred to as P. vulgare L. var. serratum Willd. (Seubert, 1844), P. vulgare L. subsp. azoricum Vasc. (Vasconcellos, 1968) and even as a distinct species P. azoricum (Vasc.) R. Fernandes (Fernandes, 1968). However, Fernandes has based her conclusions on only one specimen of P. macaronesicum and observations on a wider range of material have shown that the characters used by Fernandes to distinguish P. azoricum are unreliable, as was also noticed by Ward (1970). In plants from both the Canary Islands and the Azores these characters are much more variable than this author assumed. The azorean plants are thus better regarded as conspecific with P. macaronesicum, although there are some indications which suggest that it may eventually be possible to separate them as a subspecies of this taxon. Nardi (1977) has also concluded that the separation of P. azoricum from P. macaronesicum at species level is not justified. Recently P. macaronesicum has been reported from the extreme southern tip of the Iberian peninsula (Garretas & Salvo-Tierra, 1979). This is the first recorded occurrence of this species from continental Europe. Because of their superficial resemblence to P. australe these plants have sometimes been recorded as this species, for example, in Flora Europaea (Valentine, 1964), or placed under it as a subspecies (Nardi, 1977). However, there are several characters by which P. macaronesicum can be separated quite unambiguously from P. australe and these are summarised below. FROND CHARACTERS In P. macaronesicum the fronds are normally very large with a broadly triangular to ovate-triangular blade (fig. 1a). Although there is a considerable overlap in frond shape between it and P. australe, the ratio of blade length to breadth (5) differs in the two species. In P. macaronesicum this ratio varies from 0.9 to 2.2, in P. australe from 1.2 to 2.4. The pinnae of P. macaronesicum are also very coarsely serrate with blunt teeth, or in some specimens crenate, while those of P. australe vary from prominently to shallowly serrate (occasionally with no appreciable serration) and with more acute teeth. An additional difference which became evident during the course of this study 70 FERN GAZETTE: VOLUME 12 PART 2 (1980) 8 a\® ale al Lis > | ILS ‘S d Sy ® EP NSE SS Sos” fan8 8 wee ¥ LSS TES “as p & Gat sg Sg a5 i) 5 4 OB oy A > nS es & 3 pa B Ne By Rey Ss ae Hi a3 g 5 2g Go as Sa" ®os"s a @ oS a SS) 2 Baa. ae. A ay SB “a 2 fe 2 DP N ~' i i pe fF FIGURE 1. Polypodium macaronesicum and P. australe compared: a, whole frond of P. macaronesicum; b, tip of pinnae of P. australe; c, tip of pinna of P. macaronesicum; d, paraphyses of P. macaronesicum; e, paraphysis of P. australe. is that the pinnae of P. macaronesicum are almost constantly simply serrate, while those of P. australe are doubly serrate. These differences can easily be observed with a X 10 lens (fig. 1b, c). RHIZOME SCALES There is a very noticeable difference between P. macaronesicum and P. australe in the shape of the rhizome scales. Those of the former are more broadly ovate-lanceolate (up to 13.5mm long and 4mm wide) than those of the latter (up to 16mm long and 3.2mm wide), and their margins are more coarsely denticulate. Bobrov (1964) described a dark central stripe in the rhizome scales of P. macaronesicum, but Fernandes (1968) could not find it in those of the azorean plants. Ward (1970), however, observed that the scales of some specimens from Flores had a ROBERTS: POLYPODIUM MACARONESICUM 7 faint stripe. This feature seems to be so variable as to have little, if any, taxonomic value. SPORANGIUM CHARACTERS The annulus of the sporangium in P. macaronesicum is very similar to that of P. australe, although there is a small difference in the ranges of annulus width. This varies from 60 to 96um in the former species; from 70 to 105um in the latter. On the other hand the mean lengths of the indurated cells of P. macaronesicum (measured when the annulus of a dehisced sporangium is flattened between two glass slides), were found to be nearly identical with those of P. australe (Roberts, 1966). The two species also differ in the ranges of the number of indurated cells in the annulus : (7-) 9-20 in P. macaronesicum, (2-) 4-18 in P. australe. The following figures give the mean number of indurated cells based on 50 sporangia from each plant: P. macaronesicum, 10-14; P. australe 5-11. Badré & Prelli (1978) have found similar values for this character in P. austra/e during a recent study of the P. vu/gare aggregate in the Armorican Massif, France. The basal cells of the sporangium, however, provide an absolute criterion for separating the two species. In P. macaronesicum the number of these cells is (O-) 1-2; in P. australe (2-) 3-4 (-5). This character is most easily observed in dehisced sporangia (fig. 2a, b) and examination of 5 or 6 sporangia is then sufficient to give a clear separation. ~7e"eTe".S POON Py FIGURE 2. Dehisced sporangia of (a) P. australe; (b) P. macaronesicum. 72 FERN GAZETTE: VOLUME 12 PART 2 (1980) PARAPHYSES Fernandes (1968) was uncertain whether true paraphyses occurred among the sporangia of P. macaronesicum. However, they have been found in all specimens of this species examined during the present study and in those seen by Garretas & Salvo- Tierra (1979). The paraphyses of P. macaronesicum differ from those of P. australe in being much shorter and either unbranched or with a few, very short branches (fig. 1d, e). They are also not so abundant as in most plants of P. australe, though it should be noted that the abundance of paraphyses in this species is very variable: occasionally plants are encountered with only 2 or 3 paraphyses in each sorus. The following figures show the difference between the two species in the length of the paraphyses: P. macaronesicum, (200-) 300-400 (-800) um; P. australe, (400-) 600-1,200 (-1,800) um. SPORES Shivas (1961) studied spore length in the P. vu/gare aggregate. She measured samples of 50 spores from each plant and found that the range of mean values for P. vulgare (56-68 um) shows a large overlap with the range for P. australe (62-74 um). Those of P. interjectum alone (74-88 um) were large enough to be of taxonomic use. Only a small number of spore samples of P. macaronesicum have been available during this study. Random samples of 50 have been measured and the mean values obtained from them were seen to vary from 57 to 63 um. These figures correspond most nearly to those of P. vu/gare given by Shivas, but they also overlap with her values for P. australe. It thus seems that measurements of spore length will have at best only a limited taxonomic usefulness. Further measurements of spores from a larger sample of plants from the Atlantic islands and the Iberian peninsula are desirable. HABITAT Page (1977) has described the habitat preferences of P. macaronesicum in the Canary Islands. He found that, with Asp/enium onopteris L. and Davallia canariensis (L.) Sm., it is one of the three most abundant ferns of the heath forests, growing extensively along the mossy forest floor, on exposed roots, over rocks‘and moss-covered boulders, upon the trunks of fallen trees, and as an epiphyte along large branches of mature tree heaths. It also grows on exposed rock faces and cliffs, situations which dry out in summer. Page observed that the fern can adapt to such habitats by shedding its fronds and surviving the summer drought as thick, scale-covered rhizomes. In light shade it tends to remain evergreen. In Flores Ward (1970) found this fern commonly growing on the tops of walls, on roadside banks and as an epiphyte on trees. Garretas & Salvo-Tierra (1979) have found that in the Iberian peninsula it shows similar habitat preferences, being found in acidic, shaded situations and epiphytic on trees. P. australe, on the other hand, is well known to be a calcicole and is mainly found in crevices of limestone or other base-rich rocks, on mortared walls, and occasionally as an epiphyte on trees in limestone woodland. The two species thus not only differ morphologically, but also in their habitat preferences. Moreover, Garretas & Salvo-Tierra (1979) have found several specimens showing a mixture of the characters of the two species and with aborted sporangia. They grow in the ecotone between the habitats of the species and these authors suggest that they may be hybrids. Their existence opens the way for experimental work which may throw further light on the relationship of P. macaronesicum and P. australe. ee ROBERTS: POLYPODIUM MACARONESICUM 73 KEY TO THE EUROPEAN SPECIES OF POLYPODIUM The main characters which can be used to separate the four European species are summarised in the following key. This is based mainly on the work of Shivas (1961), Benoit (1966), Roberts (1966; 1970) and the results reported here. 1. Rhizome scales 5 to 13.5mm long (or more); sori with paraphyses; blade of frond usually broadly triangular-ovate (ratio of length to breadth 0.9 - ads Mags Gin eit Se we iN tile Pe seco eiy ake Ra: + nO emer eee ee ea 2 Rhizome scales 3 to 10mm long; sori without paraphyses; blade of frond usually lanceolate (ratio,of length.to breadth 1.4 --4.3). 0... ve ele dee 3 a Sori with large, branched paraphyses; number of basal cells of the sporangium (2-) 3 - 4 (-5); rhizome scales narrlowly lanceolate, up to 15mm long. ee SR Mt ee es a, a Blicalg ha Mes es cara, ce tie P. australe Fée Sori with small, unbranched or shortly branched paraphyses; number of basal cells of the sporangium O - 1 (-2); rhizome scales broadly lanceolate, up to SST WL Oho gine ie ee i ie ee ae ee P. macaronesicum Bobrov 3. Number of indurated cells in the annulus (4-) 7 - 10 (-13); ripe annulus colourless or pale golden-brown; number of basal cells of the sporangium 2 - 3; rhizome scales broadly ovate-lanceolate, up to 10mm long 100 individuals 1% 2m very frequent > 100 individuals 5% 2a numbers of individuals various, but cover 5-12% 2b 12-25% 3a 25-36% 3b 37-50% 4a 50-62% 4b 62-75% Ba 75-87% 5b 87-100% Following the field description there are five steps in the characterisation and classification of the quadrats to enable neat presentation and rapid comprehension of the data (Shimwell 1971): aggregation of the field data into tables to represent the local variations in vegetation; checking in the field, the ecological reality of the units extracted from the tabulation by simple field observations or measurement of environmental features; are the units described in the table readily recognisable in the field? investigation of similar patterns in other localities, thereby obtaining an overall pattern of variation within a particular vegetation type; do the patterns recur in different geographical regions? erection, differentiation and characterisation of associations; what species serve to label the communities and define the main lines of variation within them? classification of the associations into a hierarchical system of alliances, orders and classes of vegetation by reference to previously published work using an internationally accepted code of nomenclature. (Moravec 1968). EN The general approach to description and classification is probably best exemplified by a specific example, as follows. . The coincidence of Selaginella denticulata and Anogramma leptophylla in rock and wall crevices in the more humid localities of several islands and coastal regions in the Mediterranean region from Crete to Sardinia (Zaffran 1970, De Joncheere 1963), and on the Atlantic islands of Sao Miguel in the Azores (Wilmanns & Rasbach 1973) and Madeira (Benl 1970) is quite a well known phenomenon. But despite this familiarity, the only detailed records available are those of Molinier (1937) from the Isle of Hyeres, France, of Braun-Blanquet (1966) from Guipuzcoa Province, northern Spain and of Segal (1969) from the Island of Elba, Italy and Ramatuelle in the Var district of southern France. This lack of data may partly be due to the persistence of the annual A. /eptophy//a only through the humid period of the year from November SHIMWELL: FERN DOMINATED WALL COMMUNITIES rar to May and its subsequent desiccation, along with other cryptogams. During a visit to the island of Corfu in late April, it became apparent that this mural community was probably at the best stage of its annual development and some 21, one sq. m. quadrats were thus collected as a contribution to an understanding of fern-dominated vegetation in the Mediterranean region and to clarify the plant sociological affinities of this community. !n France. the community is known to plant sociologists as the association Selaginello-Anogrammetum, but as yet, it has not been described from the eastern Mediterranean region. FIGURE 1. Distribution of Mediterranean and Lusitanian fern-dominated, wall and rock vegetation types: 1, Association Selaginello-Anogrammetum leptophyllae; 2, Alliance Cheilanthion hispanicae; 3, Alliance Asplenion fissi; 4, Alliance Asplenion glandulosi. * = Corfu. SELAGINELLA-ANOGRAMMA COMMUNITIES IN CORFU Representative stands of the vegetation were recorded mainly on south-west-, west- and north-west-facing calcareous walls built from rocks of the Cretaceous period in the west of the island, and on the mortar walls of Corfu Town. The best developed stands occurred on the terrace walls of the olive groves between Lakones and Palaeocastritsa, where the dense canopy of O/ea and, occasionally, Quercus coccifera reinforced the humid winter microclimate of the island. With an annual rainfall of between 1000 and 1200mm and between 90 and 100 rain days, the island and the other lonian Islands represent the wettest coastal areas of Greece and the central Mediterranean region. Similarly, with a mean annual temperature of 18°C and a maximum monthly mean of 78 FERN GAZETTE VOLUME 12 PART 2 (1980) 25.5°C in July, the island also experiences the coolest coastal climate (Kayser & Thompson 1964). Even so, the summer temperatures and microclimate of the olive groves are insufficient to prevent the desiccation of many components of the vegetation, including Anogramma and Selaginella, which behave as winter annuals in a community with its best development in March and April. The vegetation was recorded in the crevices and spreading over the faces of calcareous walls between 10 and 250 metres altitude, with the pH of associated lithosols ranging between 6.8 and HAS): The association is characterised by the presence and dominance of Selaginella denticulata) and Anogramma leptophylla along with Ceterach officinarum ssp. officinarum, Parietaria lusitanica and/or P. judaica (Table 3). Other species which are widespread in related mural communities include Umbilicus rupestris, Mycelis muralis, Parietaria officinalis, Veronica cymbalaria and the moss, Tortula muralis. Asplenium adiantum-nigrum is the only other commonly-associated pteridophyte. Companion species of high constancy which give the vegetation a distinct appearance are the Mediterranean bryophytes Scorpiurium circinatum and Fossombronia angulosa; ubiquitous species of dry calcareous rocks and walls, such as Geranium robertianum and Cardamine hirsuta, and shade species such as Poa nemoralis, Smilax aspera and Arisarum vulgare. These overall characteristics confirm that this vegetation type may be referred to the association Selaginello-Anogrammetum. Three distinct sub-associations that reflect different ecological conditions within the association are recognisable. 1. The typical sub-association has the basic features described for the association and is generally rich in species characteristic of wall vegetation. It occurs both under olive groves and in the open, and predominantly on north- and west- facing walls with a full range of exposure from SW (220°) through to NNE (209). The major variation within the sub-association is to be seen in a community dominated by Po/ypodium vulgare, found on shady N and NNE-facing walls near the village of Krini. In plant sociological terms this community may be referred to as the sub-association typicum. 74, A second community is found on mortared walls in deep shade, where there is always some surface seepage during periods of precipitation. The community is recognisable by the predominance of the rare Mediterranean leafy liverwort Southbya tophacea and the three calcicolous mosses A/oina aloides, Gymnostomum calcareum and TJortella nitida, which often deposit a thin tufaceous crust on the vegetation and tend to eliminate the more xeric wall species. This mesic, almost sub-hydric community is virtually a transition to tufa-forming, rock vegetation typically characterised by Adiantum capillus-veneris, and it is noteworthy that Adam et al. (1972) describe a community of Adiantum and Eucladium verticillatum from Montenegro in which Southbya tophacea is a prominent species. This sub-assocation is thus known as southbyetosum. TABLE 2: LOCALITIES OF QUADRATS 1-4 Roadside walls between Ayios Simeon and Lakones 5-7 Walls at Krini 8 Angelokastron 9-14 Walls and rocks beneath olive groves, Palaeocastritsa 15-18 Walls between Ayios Simeon and Dhoukadhes 19-21 Mortared walls of Kerkira (Corfu Town) SHIMWELL: FERN DOMINATED WALL COMMUNITIES 79 TABLE 3: SELAGINELLO-ANOGRAMMETUM LEPTOPHYLLAE MOLINIER 1937 Releve Nr. LO; 27 UL 5, 15 XL 2. 18 5 S16 6 7 8 20 19 21 30 lS 2 3 93 Exposition (Degrees) 330 20 360 310 230 220 300 270 350 330 20 15 350 250 310 270 250 310 140 160 200 Vegetation Cover (%) 60 70 50 50 70 50 90 70 90 60 90 80 80 70 80 50 70 40 50 80 60 Olea Cover (%) 100 - 100 100 100 50 100 - 100 - - - 60 109 100 100 - 10 - - - Species Nr. 15S Onset 25 15, 14 920) 5 Mug SF 7. us Lees 1G 19 9) 13) 18h 19 Association, Alliance & Class Character Species Selaginella denticulata 4a 2a 3b 3b 3b +a 2a 3a 4a +b 2b +p 3a 2b 2b 2b 2b +b +b 3b 3a Anogramma leptophylla 4 +b +b +b +. +b +b +p +a +p ,. +b 2m +b +b “ED 2D) 6 ° 2a Ceterach officinarum = ib Pebie Za -2ay 2b) Ib Tae Lb) 5 O 2m +b +b +a 2b 3b 2a 2a Parietaria lusitanica +a. la +p 2m +a 1b. Vay ey) cha) Ebi. ; +p 2m +b la la +a , P, judaica . Zay * Lay cs 5 5 2b +b 3a 2b 2a 2b : +p +p : 7} +a. 5 Umbilicus rupestris ° . e - +b 1b +b +b +p +b 2a +b +b ° . ° . ° . ° la Mycelis muralis ~ a 35E 4) og aRE O 5 2 . Fy +p . + S ° ° ° +b +b Parietaria officinalis 6 a - * aH) < +b +p 2b ,. CDies a O 5 5 - : ° la Tartula muralis ° . ° . 5 [ED CC Deets ie 2m. ° +b ° e ° . e ° +b . Veronica cymbalaria 5 4 rs ° ° e Epi cep 2a. +b) -ED) = A * e e . ° ° ° Asplenium adiantum-nigrum ° . ° . . . . . . Bare anh 402) . ° . . De mea on Beare! Cymbalaria muralis 5 5 5 ° . ° 2 . e 2a. ° . ° e 5 e e ° 5 ° Differential species of Variant Polypodium vulgare ° . ° ° . ° . . ° Differential species of sub-ass, southbyetosum Southbya tophacea . ° e . . . . . . . . . . Aloina aloides . e . ° « . . . . . . . . Gymnostomum calcareum . e ° ° ° . . . . . . . . Tortella nitida ° ° ° . . . . ° . . . . . Differential species of sub-ass, targionietosum Targionia hypophylla co s . . zs ° . e e . ° é 2 e “ 5 2a) Za) Za S2b) 2 Companion species Scorpiurium circinatum by e2ae FD is) br (2a). Sb 2b 3a 2a 5 2 2 Deets 2a 2b 2b Poa nemoralis Wey” lb +b lb +a +p . +p . +p +p . +p . +p . 4 2a. ye Geranium robertianum rb --p +b). ° Daa a) “ cn +a er he og ° . . fa Mg Fossombronia angulosa - peetD (Zar o> A 96 : 5 aR) ane) og +b 2a 2a Ube es © la. Cardamine hirsuta . EDIe . tp Ee pp ha. +p +p +p . . . . . . . +a Lunularia cruciata é é 5 2a. = +p +p +p . “Pe 4 +p) (2b v. +b. 4 = = Barbula cylindrica 2a. : 2a Zio Dit . . O. 3AD » co 5 ZIAMs zal 3 2a Sonchus asper - Ir 6 5 : 28F Og e A aR) s 5 . A + 5 +a. +a +b +a Arisarum vulgare ans SFE GHD GPE 5 ans! Js ° . . . . . . . . . e +a. Quercus coccifera . ° os “r= 3bi 2ay & A ° . . +r Cl ° e . . . . ° Sherardia arvensis 5 5 +r O«w rs 2 +p. A * ny A A 5 A 4 +b . +b . Anthemis chia = aul ° . . . 7e . . . . . . . 2a +b. Seta Euphorbia peplus ae ° +p +p . . . . . . ° . . . ° lb +b . ° ° Thalictrum minus +b . 2a +b . . . . +b ,. - + 5 A 4 4 5 5 a > o Smilax aspera . ° are) sia) feo) ang 7 a c a + 5 e5 G c . ° ° ° ° Sonchus oleraceus F -r 5 sree . . . . . . . . ° . ° . e . ° +a Brachythecium rutabulum . ° . . . . . . 2a, tral (2a —5 . ° ° . ° ° ow sees Camptothecium sericeum . ° . . ° 5 FRE aN gg . . . . . . : . Leiocolea turbinata tale * is - = 6 Fi = - “ = 3 = e tbl ts e e +a Geranium rotundifolium A . ° e J e Bars! 5 > a 5 5 ° ° ° Fa wef octets . Brachypodium sylvaticum - A ° ° ° ° ° ° . . e é Py 5 . e . nn) che) hale ee: Vicia ervilia 5 . . e ° . . a e ° . ° ° a 6 . eo ote oe RG te Ornithogalum montanum a c ‘ . a ° e ° ° SP apc ag A . ° Fs ° ° . 2a +b Clematis flamnula . ° +r 28) . se ° ° . . . ° . . . . . . . Barbula unguiculata 3 ‘ = PED) a A - - A 4 A a = - A se G . . +b Bryum capillare ean ee 5 + 5 ° ° . rc ° - = : A . . oa ote) ts Fissidens taxifolius ‘ 5 DareZan ~< = +p . . ° ° e e e » ° . . . . . F, adianthoides aeE ge 2A eae = - ane e ° e ° . ° . ° ° e . . . Ditrichum subulatum +p . ‘3 ib =. ° ° ° ° . . . . . ° . . . . . . Cyclamen hederifolium ~ ao amy 22 . ° ° ° ° ° e e ° . . . . . . . Also in Relevé: (1) Rubia peregrina 2a, Tortula muralis aestiva +p; (2) Allium roseum +r; (4) Holcus lanatus +p, Carthamnus lanatus +r;, Helicrysum siculum +p, Barbula tophacea 2a; (5) Dryopteris carthusiana +p, Geranium lucidum +a, Catapodium rigidum +r; (6) Myosotan aquaticum 2a, Arabis verna +r; (9) Stachys sylvatica +p, (10) Leontodon taraxacoides +r, Cladonia sp, +a; (11) Lapsana communis +r, Hypericum montbretii +r, Carex sp, +a, Anemone pavonina +p, Hypnum cupressiforme 2a; (15) Fossombronia foveolata +b; (16) Erigeron mucronatus +r, Poa annua +b, Senecio vulgaris +, Stellaria media +b; (17) Phagnalon rupestre +r, Antirrhinum sp, +r; (19) Saxifraga tridactylites +p. Nomenclature: Phanerogams - TUTIN,T,G. et al, 1964-76, Flora Europaea. Vols. 1-4, Cambridge University Press, Mosses - WARBURG,E.F, 1963, Census Catalogue of British Mosses, BBS Publ., Ipswich. Liverworts - PATON,J.A, 1965. Census Catalogue of British Hepatics, BBS Publ., Ipswich. 3: The most xeric of the three sub-associations is found mainly on SE and SW (140--310°)- facing walls in open situations that receive direct sunlight for at least part of the day. The thallose liverwort Targionia hypophylla acts as a differential species, Ceterach officinarum assumes greater importance in terms of cover, and weeds of cultivation such as Sonchus asper, Vicia ervilia and Anthemis chia are more common. This sub-association is known as targionietosum. 80 FERN GAZETTE: VOLUME 12 PART 2 (1980) THE PLANT GEOGRAPHICAL AND SOCIOLOGICAL AFFINITIES OF THE ASSOCIATION SELAGINELLO-ANOGRAMMETUM In his numerical analysis of the geographical distribution of European pteridophytes, Birks (1976) recognises a ‘Se/aginella denticulata element’, a group of ten species which includes Anogramma leptophylla, Asplenium onopteris and A. petrarchae. \t has a distribution which is centred on the central and western Mediterranean and Lusitanian regions. The predominantly coastal distribution pattern of S. denticulata as mapped by Jalas & Suominen (1972), ranging from Crete and the Greek Islands, along the Adriatic coasts of Albania and Yugoslavia, western Italy and southern France, and with concentrations in Corsica, Sardinia and Portugal provides a good indication of the main centres of distribution of the species group. The majority of the species favour a series of humid microclimates such as those provided by the narrow-sided valleys beneath Platanus, Castanea and Cupressus woodlands in Crete (Brownsey & Jermy 1973), Olea and Quercus coccifera in Corfu to moderately open situations in the Azores and Madeira where the annual precipitation is in the range 2000—3000mm (Wilmanns & Rasbach 1971; Benl 1971). But of the ten species, only Anogramma and Selaginella commonly occur together, especially in calcareous, coastal localities throughout the region (Molinier 1937, Braun-Blanquet 1966, Segal 1969, Terretaz 1964). In most of the reported localities, the Anogramma and Selaginella are accompanied by Parietaria /usitanica, Umbilicus rupestris, Scorpiurium circinatum and Targionia hypophylia. Because of the general lack of research on fern-dominated wai! communities, there is considerable confusion in the plant sociological classification of such vegetation types. All researchers are in agreement that the majority of rock and mural vegetation types should be classified in the Class Asplenietea rupestris (Braun-Blanquet 1948). But beyond this, opinions on the naming of orders and alliances vary widely (cf. Rivas Martinez 1960, Oberdorfer et a/. 1967, Segal 1969), and there is no universally accepted system for the classification of associations. The popular trend is to refer wall and chasmophyte vegetation in which species of Asp/enium and Parietaria are predominant to the Order Parietarietalia muralis (Rivas Martinez 1960). But Segal (1969), in his extensive overview of European wall vegetation prefers to erect the Order Tortulo-Cymbalarietalia and the Alliance Parietarion judaicae to include all Parietaria judaica wall vegetation of the Mediterranean region and the Selaginello- Anogrammetum. This is probably a more preferable classificatory position for the association than either of those given by Braun-Blanquet (1966) or Rivas Martinez (1960). The various Mediterranean and Lusitanian communities of rock and wall vegetation in which pteridophytes are predominant may be classified in the following system whose taxa unite communities that are primarily similar in floristic characteristics but which also have similar ecological and geographical distribution patterns. Ae Class Asplenietea rupestris Br.-BI. 1934. Ae Order Androsacetalia vandellii Br.-Bl. 1926 — chasmophytic vegetation of siliceous rocks in the sub-alpine and alpine regions of central Europe, the Pyrenees and Sierra Nevada. The order is named for the uncommon calcifuge, Androsace vandellii, and Braun-Blanquet 1948) considers Woodsia ilvensis and Asplenium septentrionale as good regional character species. In view of the wider distribution of the latter species, Oberdorfer et a/. (1967) suggest that Asplenietalia septentrionalis is a better name. 1.11 Alliance Asplenion septentrionalis Gams 1927 (Androsacion vandellii Br.-Bl. 1926) — an alliance containing two main associations, Asplenietum septentrionali-adianti-nigri Oberd. 1938 and Woodsio-Asplenietum septentrionalis R.Tx.1937, which are recorded in the mountain regions of central Europe and are probably to be found in the Lusitanian region. 1.12 Alliance Cheilanthion hispanicae Rivas God. 1955 — the most oceanic alliance in the order, ee SHIMWELL: FERN DOMINATED WALL COMMUNITIES 81 found on shallow soils over siliceous rocks in the Sierra Morena, Spain and in eastern Portugal. Cheilanthes hispanica is a good character species. 1.13 (Alliance Anogrammion leptophyllae Bellot & Casaseca 1959 — described by Rivas Martinez from the western mountains of Spain and of doubtful status; Anogramma communities of natural rock-crevice sites). 1.2 Order Asplenietalia glandulosi Br.-Bl. & Maier 1934. 1.21 Alliance Asplenion glandulosi Br.-B!. 1926. The most interesting fern-dominated association in this group of Mediterranean calcicolous vegetation types is probably Phagnalo- Asplenietum glandulosi Br.-Bl. (1931) 1952, named for the occurrence of Asplenium petrarchae (syn. A. glandulosum) and Phagnalon sordidum. The association is described by Sutter (1973) mainly from south-facing calcareous rocks in lower Languedoc and probably occurs throughout the range of A. petrarchae. Cheilanthes fragrans is the only other rare fern found in the association and communities in which the calcicole Asplenium fontanum dominates and probably also belong to this alliance. 1.22 (Alliance Asplenion fissi — vegetation of fissures and crevices in montane calcareous rocks of SE Europe in which Asplenium fissum and A. lepidum are common may form an eastern vicariant of Asplenion glandulosi). 3 Order Tortulo-Cymbalarietalia Segal 1969. .31 Alliance Parietarion judaicae. Association Selginello-Anogrammetum leptophyllae Molinier 1937. 1.32 Alliance Tortulo-Asplenion — includes two associations which are widespread in Europe — the ubiquitous Asplenietum trichomano-rutae-murariae R.Tx. 1937 and Asplenio- Cystopteridetum Oberd. (1936) 1947, a sub-montane and montane association characterised by the presence of Asp/enium viride and Cystopteris fragilis. 2: Class Adiantetea Br.-Bl. 1947. a4 Order Adiantetalia Br.-BI. 1931. 2.11 Alliance Adiantion Br.-BIl. 1931 — communities dominated by Adiantum capillus-veneris, occurring On moist, shaded, often tufaceous, limestone rocks, or in humid caves in the Mediterranean region and on the southern Atlantic coast of Europe. The most common association, Eucladio-Adiantetum is named for the fern and the small, tufa-forming bryophyte, Eucladium verticillatum. The above classification is by no means complete. It is simply a contribution to a better understanding of the sociology and ecology of fern-dominated vegetation of rocks and walls. There is a great need for further research by plant sociologists and pteridologists and this paper presents a standard method and classificatory framework as stimuli for such studies. REFERENCES ADAM, P., BIRKS, H.J.B. & WALTERS S.M. 1972. A contribution to the study of the flora and vegetation of the Budva area, Montenegro. Glas. Republ. Zav. Zast. Privode. 4: 41-72. BENL, G. 1971. Fern hunting in Madeira. Brit. Fern Gaz. 10: 165-174. BIRKS, H.J.B. 1976. The distribution of European pteridophytes: a numerical analysis. New Phytol. 77: 257-287. BRAUN-BLANQUET, J. 1948. Ubersicht de Pflanzengesellschaften Ratiens. |. Vegetatio. 7: 29-41. BRAUN-BLANQUET, J. 1966. Vegetationsskizzen aus dem Baskenland mit Ausblicken auf das weitere Ibero-Atlantikum. |. Teil. Vegetatio 73: 117-147. BROWNSEY, P.J. & JERMY A.C. 1973. A fern collecting expedition to Crete. Brit. Fern Gaz. 10: 331-348. DE JONCHEERE, G.J. 1963. Ferns of Sardinia. Brit. Fern. Gaz. 9: 114-116. JALAS, J. & SUOMINEN, J. 1972. Atlas Florae Europaeae |. Pteridophyta. Helsinki, 121 pp. KAYSER, B. & THOMPSON, K. 1964 Economic and Social Atlas of Greece. National Statistical Service of Greece, Athens. MOLINIER, R. 1937. Les Iles d‘'Hyéres; étude phytosociologique, Ann. Soc. Hist. Nat. Toulon. Nr. 21. MORAVEC, J. 1968. Zu den Problemen der pflanzensoziologischen Nomenklatur. In TUXEN, R. 1968 (ed.) Pflanzensoziologische Systematik Ber. Int. Symp. 1964, Stolzenau/Weser. Springer Verlag. OBERDORFER, E. et a/. 1967. Systematische Ubersicht der westdeutschen Phanerogamen — und Gefasskryptogamen — Gesellschaften. Schrift. Vegetationskunde 2: 7-62. 82 FERN GAZETTE: VOLUME 12 PART 2 (1980) RIVAS MARTINEZ, S. 1960. Roca, clima y communidades rupicolas; sinopsis de las allianzas hispanas de Asplenietea rupestris. Annales Real Acad. Farm. 1960: 153-168. SEGAL, S. 1969. Ecological notes on wall vegetation. Junk. The Hague. 325 pp. SHIMWELL, D.W. 1971. Description and Classification of Vegetation. Sidgwick & Jackson, London. z SUTTER, R. 1973. Uber Vorkommen und Vertbreitung von Asplenium petrarchae (Guerin) DC. und die Assoziationen Phagnalo-Asplenietum petrarchae. Bot. Jahrb. Syst. 93: 568-602. TERRATAZ, J-L. 1964. Anogramma leptophylla en Valais et en Ossola. Trav. Soc. Bot. Geneve. 7: 20-30. WILMANNS, O. & RASBACH. H. 1973. Observations on the Pteridophyta of Sao Miguel, Acores, Brit. Fern. Gaz. 10: 315-329. ZAFFRAN, J. 1970. Les Pteridophyte Cretoises. /srae/ J. Bot. 19: 236-244. REVIEW OVERLAYS OF ENVIRONMENTAL AND OTHER FACTORS FOR USE WITH BIOLOGICAL RECORDS CENTRE DISTRIBUTION MAPS. Published by the Institute of Terrestrial Ecology, 68 Hills Road, Cambridge CB2 1LA, England. 1978. 390 x 212 mm. Price £2.00. (Available, by post, from the Publications Officer at the above address.) When | had the pleasant task of reviewing the ‘Atlas of Ferns of the British Isles” (see Fern Gaz. vol. 11 p.399-400, 1978), | mentioned that there were no overlays which could be used to compare fern species ranges with environmental factors. Since then, | have been sent a set of transparent overlays which can be used in conjunction with the Atlas. It must be added that these overlays are not an exact fit on the Fern Atlas maps as they are not designed specifically for this purpose. But they are sufficiently near in size to be useful, for, with a bit of manipulation, appropriate areas can be brought into reasonably close superposition (overall the overlays are about 3 mm longer top to bottom, than the scale of the Atlas maps). The set comes flat in a card folder which contains some information about the features used on each overlay. There are 12 overlays. These include: altitude, geology (chalk and limestone), rivers, upland moorland, western oakwoods, February minimum temperature, January mean temperature, July mean temperature, annual rainfall, wet days, vice-county boundaries, county boundaries. Despite the slight discrepancy in map size, this seems a very useful set of overlays to use in conjunction with the Fern Atlas. C.N. PAGE FERN GAZ. 12(2) 1980 83 FERNS OF NAINITAL (WESTERN HIMALAYAS): AN UPDATED LIST S.C. VERMA and S.P. KHULLAR Department of Botany, Panjab Univerisity, Chandigarh, India. ABSTRACT Nainital, situated about the easternmost limit of the western Himalayas, is one of the most beautiful hill stations in the Himalayas (India). It abounds in fern species. This account endeavours to list all the species of ferns which occur in and around Nainital. The list, largely based on our own collections, includes 96 species distributed over 33 genera. INTRODUCTION During the monsoons, ferns are commonplace at every hill station in the Himalayas. Nainital is a beautiful hill-station of the Kumaon hills (India), deriving its name from the large ‘eye-shaped’ lake (Naini = eye, Tal = lake) which it surrounds at an altitude of 1850 m. It is located at 29°23 N, 79°30 E and forms the easternmost limit of the western Himalayas. In addition to the main lake, the area includes several lakes at lower altitudes, such as Sariya-Tal (1770 m), Khurpa-Tal (1635 m), Bhim-Tal (1371 m), Naukhuchia-Tal (nine-cornered, 1219 m) and Sat-Tal (a complex of seven lakes, 1371 m). It also includes several types of forest formations covering an altitudinal range from 900 m (below Bhujia Ghat) to 2611 m (Cheena peak or Naini peak). Duthie (1918) was the first to catalogue the ferns of Nainital, together with those of the Kumaon region and the adjacent frontiers of Garhwal and Tibet, covering a total area of 11,500 sq. miles. From this vast area, a total of 185 species in 30 fern genera, and 15 species of fern-allies in 6 genera were recorded. But only 24 species of ferns, based on collections made by Strachey and Winterbottom between the years 1846-1849, were referred to Naintal. Loyal and Verma (1960) made fresh collections and recorded a total of 61 species for Nainital. Subsequently more collections were made by S.C. Verma (in 1962 and 1975), S.P. Khullar (in 1965, 1967, 1972 and 1975), R.P. Nagpal (in 1970) and Vijay Bala (in 1974), all from the Department of Botany, Panjab University. We feel that these much more complete collections justify an updated list of the ferns of Nainital. The present updated list brings the total number of ferns for the Nainital area to 96 species in 33 genera. LOCATIONS VISITED With headquarters at Nainital, field trips to most surrounding different types of vegetation can be made. Ferns are particularly abundant in the oak forests and especially in the very humid ravines and streams within them. Notably rich areas are (i) the pony road to Kilbury, (ii) Naintital to Dhobi Khad (towards the Laria Kanta side) following the old Nainital-Almora pony road, (iii) the area between Bhujia Ghat and Jeolikote, and (iv) along the path passing through the forest from Land’s End to Dorothy’s Seat (Tiffin Top) (see fig. 1). 84 FERN GAZETTE. VOLUME 12 PART 2 (1980) I 79°30 ot © Kilbury ( 2,194 M) ris NAINA PEAK (2610 M) Y) —<> DS Qe VIEW (2270 M) DHOB! KHAD ® LARIA KANTA @ (2481M) N He 27-007 7 80 oe votes . BAKO QO o> . ee . e eh >» oo . oes b as she oe ey es ° To Eh . Sariyatal oe seamtarae ewe Lefstes AS f' s & Se ; rac) OTHY SEAT tee (2290 M) a a io Ory Do Age ar O 0 SOE = eM —— To Bhowali (Sat-Tal , Bhim -Tal , Naukuchia -Tal ) 1920M to 1996M 1996 M tc 2073M 2073M to 2149M Motorable Road Non-motorable Road © Jeolikote (1,219 M) or? @ Do-gaon Bhujia Ghat To Kathgodam FIGURE 1. Map of Nainital area showing relief, roads and main localities mentioned in the text. THE LIST OF FERNS Species are arranged in families following the system proposed by Mehra (1961). For canvenience, the genera and species within families are listed alphabetically. Eleven species, based solely on Duthie’s catalogue are included (marked ‘’D’’), and these need to be located afresh. While listing the ferns, the principal localities of only those species considered relatively restricted in distribution are stated, and such localities are numbered on the map (fig. 1). All other species are more or less common around Nainital. Preparation of a fully illustrated and keyed fern flora of Nainital is in progress. Botrychium lanuginosum Wall. ex Hook. & Grev. var. /anuginosum. Fairly common throughout Nainital, usually on exposed sloping surfaces in the forest and road sides, highly variable in size and pubescence. B. ternatum (Thunb.) Sw. Collected by Prof. P.N. Mehra in 1949 from Laria Kanta side, also reported by Duthie (1918). Ophioglossum petiolatum Hk. Restricted to grassy slopes, above Khurpatal, grows in patches protected and concealed under small shrubs (fig: 8), very variable. O. vulgatum L. (s.|.). Rare, found with difficulty in the grassy open land below Tiffin Top. A larger form, more near to O. reticulatum L., occurs in abundance about the Kalika temple (Ranikhet, near Nainital, fig. 9). Lygodium flexuosum (L.) Sw. Fairly common below Bhujia Ghat, about the edges of broad- leaved forests, climbing on shrubs, in relatively open places. Adiantum capillus-veneris L. The cosmopolitan Venus-hair fern occurs ‘everywhere’ near water, particularly along stony drainage courses, variable. A. edgeworthii Hk. Restricted, found behind Kainchi temple, about Bhowali and Bhim Tal — Sat Tal area. VERMA & KHULLAR: FERNS OF NAINITAL 85 A. incisum Forsk. Abundant at low altitudes. Occurs in at least two forms: an erect or semi- erect one with large dark-green fronds and a smaller prostrate form (usually more hairy) often completely matting sloping forest margins and the masonary work. Both forms are diploid (n=30), unlike the African A. incisum, which is tetraploid. A. lunulatum Burm. More often called A. philippense, but the name A. /unulatum is preferred (Verma, 1961). Common at low altitudes on shady and moist slopes. Variable in frond size, pinnule form and size, and at least the Bhujia Ghat population seems to be uniformly triploid apogamous (n=2n=90) (cf. Mehra and Verma, 1963). A. venustum Don. Abundant on very humid shaded slopes, usually in the forests, between 1800-2400 m. Onychium contiguum Hope. Abundant, covering large areas in both shaded and open localities between 1800-2400 m (fig. 3). The possibility of the occurrence of O. /ucidum and O. siliculosum, which are met with in the adjoining regions eastwards, needs to be explored. O. siliculosum has been collected from Thal, a place 140 miles from Nainital, in the interior of Kumaon hills. *Cheilanthes albomarginata Clarke (= Aleuritopteris). The most common and the most variable fern, along roadsides and forest margins in and around Nainital, descending to 1300 m. Fronds scaly all over (stipe, primary and secondary rachises and costae). C. anceps Blanf. Rather rare, collected near Jeolikote along the roadside. C. anceps Blanf. var brevifrons Khullar. Shares with C. anceps only the extension of scales on the stipe and the principal rachis and better treated as a distinct species. Common in humid and shady or protected niches. Very common near Bhowali, slopes around Khurpa Tal, Do-Gaon- Jeolikote, and Bhim Tal. C. anceps aff. Locally common in Bhim Tal area, possibly hybrids between 12 and 13. The entire anceps group needs careful study. C. dalhousiae Hk. Very frequent at higher altitudes, particularly en-route to Cheena peak, Dhobi Khud and Kilbury. a non-farinose species of the A/euritopteris group. C. farinosa sens. lat. An extremely common fern up to 1500 m altitude, uniformly diploid (n=30), and a taxonomically a confused species. Most forms agree with C. farinosa sensu Blanford, but true C. farinosa of Yemen is triploid and apogamous. This is a complex group needing further study. C. subrufa Desv. Very common in crevices of slate rocks and masonary work between Dogaon and Bhujia Ghat (850 m). Anogramma leptophylla (L.) Link. A small annual fern, restricted to humid niches and concealed among mosses and liverworts (1900-2400 m), but by no means common; collected several times on way to Cheena peak and Dhobi Khud. Coniogramme affinis Hieron. Infrequent, gathered only in the Kilbury forest area. C. caudata (Wall.) Ching. Not common, found along the ravine about Dhobi Khud in the forest. Gymnopteris vestita (Wall.) Underw. Rather infrequent, found in the crevices along the road to Kilbury (fig. 2). Pteris cretica L. Extremely common (1600-1900 m) in both open and shaded locations on forest margins and inside the forests (fig. 5); very variable in overall size, serration on the margin, and fertility. P. excelsa Gaud. Local, in deep ravines, mostly in Dhobi Khud area. P. quadriaurita s.1.. Common polymorphic fern of forest floor and margin, at least three forms distinguishable, probably none equivalent to P. quadriaurita Retz. sensu stricto (see Verma and Khullar, 1965). P. vittata L. Common at low altitudes. Two forms occur, both tetraploid (n=58); one with a very long terminal pinna, which inhibits open roadsides and forest margins, and another with a small terminal pinna but fronds often very large, which inhabits shaded and damp habitats. The latter is closer to the Linnean type. A third form, which is infrequent, collected by S.C. Verma once below Bhujia Ghat, differs in having narrower pinnae and broader indusial flaps and is diploid (n=29). Further study is needed. Pe ea a eC ae age a ng ees *We follow Manton et a/. (1966: Kew Bull. 18(3): 553-565) in recognising the genus Che//anthes as many of the characters mentioned by Ching are not shared by all the species in the Himalayas. Excepting C. dalhousiae, other species (farinose on the under surface) could be placed under Aleuritopteris. 86 FERN GAZETTE: VOLUME 12 PART 2 (1980) Didymoglossum insigne van den Bosch (Crepidomanes insigne (v.d. Bosch) Fu). Plentiful very locally on moist dripping rock beside the bridge over a ravine near Dhobi Khud, otherwise rare. Dennstaedtia scabra (Wall.) Moore. Recorded by Duthie (1918). Pteridium aquilinum (L.) Kuhn var. wightianum (Ag.) Tryon. Rather localised (but not weedy), fairly common on way to Kilbury (fig. 4), Laria Kanta, and below Kalika temple (Ranikhet). Araiostegia pseudocystopteris (Kze.) Copel. An abundant epiphyte/lithophyte all around Nainital. Nephrolepis radicans (Burm.) Kuhn. The species has been reported by Duthie as WV. volubilis Ais Safle Crytomium caryotideum Presl. Collected from one locality beside a ravine near Dhobi Khud, not common. Dryopteris chrysocoma (Christ) C. Chr. Exceedingly common at the altitude of Nainital and above. D. cochleata (Don) C. Chr. A common Dryopteris of lower altitudes, particularly on the Dogaon-Bhujia Ghat area, Bhim Tal-Sat Tal, Kainchi, and in the pine forest about Kalika temple (Ranikhet); fronds dimorphic. D. marginata (Wall.) Christ. Common forest floor fern near Kilbury, forest between Land's End and Dorothy’s Seat, and on Bhowali road (near ravine). D. odontoloma (Moore) C. Chr. Very common along roadsides and on forest floor about Nainital (fig. 11). D. sparsa (Don) O. Ktze. Rare, collected only once opposite Sanatorium, Bhowali. Hypodematium crenatum (Forsk.) Kuhn. Exceedingly common at lower altitudes, on exposed rocks, particularly near the Khairna bridge on Kosi river; quite common on Kaladungi road (Nainital), variable. Polystichum acanthophyllum (Franchet) Christ. Rare, found on Kilobury road in dark shaded rocks, also recorded by Beddome (1892). P. indicum Khullar et Gupta. This distinct fern of the Himalayas has been commonly referred to P. aculeatum (L.) Roth by almost all the workers on Indian ferns, probably due to Beddome’s (1892) remark that P. acu/leatum occurs commonly in the Himalayas. Both at Kew, and the British Museum such samples have been filed under P. aculeatum Auct. (at Kew, Bliss Nos. 39, 165, 188, 282). These forms differ considerably from the European species (P. acu/eatum (L.) Roth), which is tetraploid and has compact, coriaceous fronds with a short stipe, and the pinnae are reduced towards the base. The Himalayan samples are diploid and have more or less lax, soft- subcoriaceous fronds with long stipe, and their pinnae are slightly or never reduced towards the base. The stipes are fibrillose and scaly, and the rachis is devoid of broad scales. To avoid confusion, Khullar and Gupta (1977) gave this new name to the common Himalayan form of the so-called P. aculeatum. P. indicum shows variation in length of stipe, shape, size and margin of pinnules, shape of fronds, colour of ramentae (light-brown to almost blackish) and texture of fronds. Quite common towards Kilbury, Tiffin Top forest, and Dhobi Khud area (1800—2100 m). P. nigropaleaceum (Christ) Diels. Somewhat similar to P. indicum, but differs in the presence of scales and fibrillae al! over the principal rachis. The species has been placed under P. aculeatum var angulare by many earlier workers. Dr. F.M. Jarrett identified the material collected from Kashmir by S.P. Khullar (Nos. 72 at Kew) and by NBGL (No. 96685 also at Kew) as P. cf. setiferum. Similar specimens from Mussoorie and Murree have been identified as P. nigropaleaceum (=P. setiferum (Forsk.) Moore ex Woynar var. nigropaleaceum Christ) by R.R. Stewart. At the British Museum, Schelpe’s specimens Nos. 3609 and 3582 from Parbati valley and No. 3172 from Manali have been identified as P. nigropaleaceum by A.H.G. Alston. The species is variable as regards the colour and density of scales and fibrillae on the rachis. Fairly common towards Kilbury, Tiffin Top, and Dhobi Khud area. P. obliquum (Don) Moore. Infrequent, on dark, dripping shaded rocks; locally common towards Kilbury and Dhobi Khud; variable in frond size. P. squarrosum (Don) Fée. Exceedingly common on the forest floor, forming dark-green baskets. Polystichum sp. (unidentified). Collected only once from an open dry rock on the road to Kilbury. Tectaria coadunata (J. Sm.) C. Chr. Common at lower altitudes, very variable. Athyrium anisopterum Christ. Common on moist shaded rocks along the Kilbury road (fig. TAR A. falcatum Bedd. Rare, collected once near Kilbury; also recorded from Ranikhet. VERMA & KHULLAR: FERNS OF NAINITAL 87 A. foliolosum (Wall. ex Clarke) Bedd. Found only near a streamlet on the pony track to Kilbury, particularly about the grove shown in fig. 16. A. oxyphylium (Hook.) Diels. Recorded by Duthie (1918). Collected from the Thal-Didihat forest (150 miles from Nainital) in Kumaon hills. A. pectinatum (Wall.) Presi. A low-level fern, locally common on the slopes above Khurpa Tal, Jeolikote, Bhowaii raod, Kainchi, Bhim Tal—Sat Tal area; near Kalika temple (Ranikhet). A. proliferum Moore. Recorded only from Ranikhet, can be expected in Nainital. A. drepanopterum (Kze.) A.Br. Infrequent, along the Kilbury road. A. rupicola (Hope) C. Chr. Common at higher levels, particularly along the Kilbury road. A. schimperi Moug. ex Fée. Exceedingly common on the forest floor. Athyrium nigripes (Blume) Moore. Recorded by Duthie (1918). Lunathyrium japonicum sens. lat. Recorded by Duthie (1918). Diplazium lobulosum Wall. ex Pr. Recorded by Duthie (1918). D. polypodioides Blume. Common near ravines, on Bhowali road, Kainchi, Bhim Tal—Sat Tal forest, and Kalika (Ranikhet). D. spectabile (Wall. ex Mett.) Ching. Not common, collected from the ravine near Dhobi Khud. Goniopteris prolifera (Retz.) Pres| (Ampelopteris prolifera (Retz.) Copel.). Common along the streamlets in the Naukuchia—Bhim Tal area. Christella dentata (Forsk.) Brownsey & Jermy (Cyclosorus dentatus (Forsk.) Ching). Common along ravines, water channels in Khurpa Tal, Dogaon—Jeolikote and Sat Tal areas. Macrothelypteris bukoensis (Tagawa) Pic. Ser. (The/ypteris brunnea (Wall.) Ching). Locally common, near water channels and humid areas, particularly Kilbury, about the place in fig. 16. Glaphyropteridopsis erubescens (Wall. ex Hook.) Ching (Thelypteris erubescens (Wall.) Ching). Common along ravines at lower elevations, particularly along the Bhowali road. Pseudocyclosorus repens (Hope) Ching (The/lypteris repens (Hope) Ching). Common along ravines and ‘swampy’ areas of Dogaon, Bhowali—Jeolikote area. Cyclogramma auriculata (J. Sm.) Ching (Thelypteris subvillosa (Moore) Ching). Recorded by Duthie (1918). Pseudocyclosorus tylodes (Kunze) Ching (7. xylodes (Kunze) Ching). Recorded by Duthie (1918). Asplenium dalhousiaé Hook. The most common fern, extending from low altitudes (Kathgodam area) to Cheena peak; very variable in form and size. A. ensiforme Wall. ex Hook. & Grev. Not common, lithophyte in very humid localities, collected on way to Kilbury and from Dhobi Khud ravine. A. exiguum Bedd. Rare, on moist shady rocks in the Kilbury area. A. indicum Sledge. Belongs to A. /aciniatum complex, not infrequent, in quantity opposite Sanatorium Bhowali, towards Kilbury, and near Rattighat. A. unilaterale Lamk. Recorded by Duthie (1918). A. varians Hook. et Grev. Abundant all around Nainital, on moist shaded rocks, and along roadsides. Arthromeris wallichiana (Spr.) Ching. Infrequent, found in Dhobi Khud area. Drynaria mollis Bedd. Common epiphyte in the Cheena peak area, Kilbury, towards Land's End, Dorothy's Seat and Laria Kanta. Lepisorus excavatus (Bory) Ching (=Pleopeltis excavata (Willd.) Sledge). An abundant epiphyte/lithophyte all around Nainital (fig. 18). L. aff. excavatus. A larger form from Cheena peak area, needs further study. L. kashyapii (Mehra) Mehra (=Pleopeltis kashyapii (Mehra) Alston). Common epiphyte around Nainital, (fig. 14). L. kuchanensis (Wu) Ching. Common epiphyte, particularly in the Tiffin Top area. L. nudus (Hk.) Ching. Fairly common epiphyte, about Tiffin Top, en route to Snow View, Naintal lake-side, and Nainital-Ranikhet road. Lepisorus aff. nudus (Hook.) Ching. A very characteristic fern, and the most common epiphyte at lower altitudes, particularly Khurpa Tal, Saria Tal, and below Jeolikote (fig. 12). Also occurs about Dhobi Khud and Tiffin Top. It is characterised by simple leathery fronds (subcoriaceous), yellowish-green colour and scales over the undersurface. R.R. Stewart has placed similar material from Mussoorie, at Kew, under L. nudus (Hook.) Ching. The material differs from the description of L. nudus provided by Bir and Trikha (1969) in possessing the scaly undersurface. Further studies are necessary. L. scolopendrium (Don) Mehra and Bir (=Pleopeltis scolopendrium (D. Don) Alston & Bonner). Infrequent, found towards Cheena peak and Kilbury. 88 FERN GAZETTE: VOLUME 12 PART 2 (1980) Loxogramme involuta (Don) Presl. Rather rare, gathered on Nainital-Saria Tal road below Cheena peak road, and from Dhobi Khud. Microsorium membranaceum (Don) Ching. Fairly common on moist shady rocks, locally abundant below Jeolikote (fig. 6). Phymatodes oxyloba (Wall. ex Kunze) Presl. Infrequent lithophyte, found near Bhowali Sanatorium and beyond, Bhim Tal — Sat Tal area, on the way to Rattighat, and beyond Bhowali towards Kainchi. P. stracheyi Ching. Rare lithophyte, found in large numbers on the pony track to Kilbury (locally common, fig. 13). Polypodium amoenum Wall. ex Mett. A common epiphyte or lithophyte of Nainital. P. argutum Wall. ex Hook. Infrequent, found as epiphyte on way to Kilbury, Tiffin Top, and Dhobi Khud (fig. 19). P. lachnopus Wall. ex Hook. Common epiphyte/lithophyte of Nainital, characterised by black ‘hairs’ on rhizomes. P. microrhizoma Clarke ex Bak. Epiphyte/lithophye, very common all around Nainital. P. subamoenum Clarke. Epiphyte/lithophyte, common all around Naintal. Pyrrosia beddomeana (Gies) Ching. Common epiphyte near Bhim Tal. P. flocculosa (Don) Ching. Frequent to common epiphyte/lithophyte, particularly about Khurpa Tal and below Jeolikote. P. stictica (Kunze) Holtt. Common about Dogaon, Bhujia Ghat, and on way to Rattighat. REFERENCES BEDDOME, R.H. 1892. A Handbook to the Ferns of British India, Ceylon and Malaya Peninsula (with Supplement). Thacker Spink & Co., Calcutta (India). BIR, S.S. and TRIKHA, C.K. 1969. Taxonomic revision of the polypodiaceous genera of India- IV. Polypodium lineare complex and allied species. Bul/. Bot. Surv. India. 117: 260—276. DUTHIE, J.F. 1918. Catalogue of the Plants of Kumaon and the Adjascent Portions of Garhwal and Tibet. (Catalogue originally prepared by Strachey in 1852 on collections made by Strachey and Winterbottom between the years 1846—1849), Lovell Reeve & Co. Ltd., London. KHULLAR, S.P. and GUPTA, S.C. 1977. On the occurrence of Polystichum aculeatum in the Himalayas. Symp. Recent Res. Pl. Sci. (Punjabi University), Patiala, Abstr. (sect. 2): 11—12. LOVE, A., LOVE, D. & PICHI-SERMOLLI, R.E.G. 1977. Cytotaxonomical Atlas of the Pteridophyta. J. Cramer, Vaduz. LOYAL, D.S. and VERMA, S.C. 1960. Ferns of Nainital. J. Bombay Nat. Hist. Soc. 57: 479—490. MEHRA, P.N. 1961. Cytological evolution of ferns with particular reference to Himalayan ferns. (Presidential address). Proc. Indian Sci. Congr. (Roorkee) part 11: 130—153. MEHRA, P.N. and VERMA, S.C. 1963. Polymorphicity and cytogenetics of Adiantum lunulatum complex. Maheshwari Comm. Vol., J. Indian Bot. Soc. 42A: 110—121. VERMA, S.C. 1961. Taxonomic status of Adiantum lunulatum Burm. Nova Hedwigia 3: 463—468. VERMA, S.C. and KHULLAR, S.P. 1965. Cytogenetics of the western Himalayan Pteris cretica complex. Ann. Bot. (n.s.) 29: 673—681. FIGURE 2. Gymnopteris vestita, Kilbury road area (c. 1/7 natural size). VERMA & KHULLAR: FERNS OF NAINITAL * FIGURE 3. Onychium contiguum, towards Cheena peak (c. 1/20 natural size). eo ~ % 02 oo As ae > soli he > _ * , 4 Paar 4 be As TAS ects 2: x a ds a . “a ou ¥ ‘ eS oe) a ; —* Ye eky f te ae . 4 a FIGURE 5. Pteris cretica, Kilbury road area (c. 1/12 natural size). 89 90 FERN GAZETTE VOLUME 12 PART 2 (1980) PE SSS r FIGURE 6. Microsorium membranaceum, Saria-Tal road (c. 1/7 natural size). FIGURE 7. Dryopteris marginata, near Kilbury (c. 1/7 natural size). FIGURE 8. Ophioglossum petiolatum, above Khurpa-Tal (c. 1/7 natural size). FIGURE 9. Ophioglossum vulgatum (large form), near Kalika temple, Ranikhet (c. 1/10 natural size). FIGURE 10. Asplenium varians, Lakeside, Nainital (c. 1/7 natural size). FIGURE 11. Dryopteris odontoloma, Laria Kanta (c. 1/20 natural size). VERMA & KHULLAR: FERNS OF NAINITAL 91 ee Lae «eee ad eee +? Aeh) if HN \ FIGURE 12. Lepisorus amaurolepida, midway to Saria-Tal (c. 1/5 natural size). FIGURE 13. Phymatodes stracheyi, beside pony road to Kilbury (c. 1/10 natural size). FIGURE 14. Lepisorus kashypaii, near Kilbury road (c. 1/8 natural size). FIGURE 15. Polypodium microrhizoma with Athyrium anisopterum beneath, beside Kilbury road (c. 1/10 natural size). 92 FERN GAZETTE: VOLUME 12 PART 2 (1980) x “oe Rie FIGURE 16. A typical fern grove on the pony track to Kilbury, showing mainly The/ypteris brunnea (in front) (c. 1/50 natural size). FIGURE 17. Athyrium anisopterum, Kilbury road area (c. 1/8 natural size). FIGURE 18. Lepisorus excavatus, Tiffin Top forest (c. 1/8 natural size). FIGURE 19. A branch full of epiphytes with many fronds of Po/ypodium argutum (c. 1/20 natural size). FERN GAZ. 12(2) 1980 93 THE FERN FLORA OF OILPALM PLANTATIONS IN WEST MALAYSIA A.G. PIGGOTT 21 Jalan Dato Klana, Seremban, Malaysia ABSTRACT The environment of scientifically-controlled commercial plantations of Elae/s guineensis on the Malaysian peninsula is described and discussed as a habitat for ferns. INTRODUCTION Elaeis guineesis Jacquin, a palm of the sub-family Cocoideae, is a native of West Tropical Africa and has long been exploited there on a village scale for its oil. The tall, stout, upright trunk bears a crown of pinnate leaves which have spines along the edges of their leaf-stalks. Male and female inflorescences are produced on short stalks amongst the leaf bases of the crown, as subsequently are bunches of fruit. Edible oils are manufactured from both the pericarp and the kernel. Large plantations of this species are cultivated in wet tropical lowlands through- out the world, within about 16° of the equator. By 1968, there were over 120,000 ha of oilpalms in Malaysia alone. The natural vegetation in Malaysia is tropical rain forest. Annual rainfall in the lowlands is about 2,500 mm, with precipitation more or less throughout the year. Humidity is high. Palms and ferns contribute significantly to the natural flora, and wild palms which retain their leaf-bases often support epiphytes, most commonly ferns. Scientific plantation agriculture has produced a new habitat for these, extending their previously limited distribution. The author has long had associations with the palmoil industry, and this article is compiled from personal field observations. Pteridophyte nomenclature is based on Holttum (1966), with some revisions. Further useful information on palms in general is given in Corner (1966) and Whitmore (1977), and oilpalms by Hartley (1977). Fern specimens have been deposited at Kew (K), and duplicates of Malaysian specimens in Singapore (SING). YOUNG OILPALMS High quality oilpalm seed is germinated under laboratory conditions and, after careful selection, planted in nurseries. Seedlings develop until they are of sufficient size for planting in the field. By this stage, the palms have not produced trunks and there are no epiphytes. In the field, the young palms are at first surrounded by leguminous cover crops. Clean circles are maintained around each palm, either manually or by use of herbicides, and the cover crop itself is weeded. Then ferns begin to grow. Nephrolepis biserrata often establishes on the now-developing trunks, but such plants are continually removed during plantation maintenance operations. After about 30 months the first palm fruit crop is taken. The short trunk is now recognisable and the leaf-bases are beginning to accumulate debris. NV. biserrata is still cleaned away regularly, but some of the rhizomes have become so well-established by this stage that fronds soon reappear. Where wetter conditions prevail and Stenochlaena palustris occurs in the surrounding areas, it too may make its appearance early. Conditions 94 FERN GAZETTE: VOLUME 12 PART 2 (1980) within the plantation are still fairly open, but as the palm canopy closes and the cover crops die out, and terrestrial ferns gradually appear. Although the trunk ts regularly cleaned until it is 6 or more years old, larger numbers and more species of epiphytic ferns then establish on it. WEST MALAYSIA THAILAND ol Penang Kelantan £ : O & Trengganu O O STRAITS F ie} MALACCA Negeri Sembilan KEY Railway State boundary National boundary ——-—-— Plantation with epiphytes @ Plantation with no significant epiphytes oO Johore SINGAPORE FIGURE 1. Map of West Malaysia showing the distribution of oilpalm plantations and small- holdings where the fern flora was recorded. PIGGOTT: FERNS OF OILPALM PLANTATIONS 95 MATURE OILPALMS Rank upon rank of oilpalms with their neatly tailored trunks cover large areas of the Malaysian lowlands (fig. 2). The canopy meets overhead giving shade and cooler, but more humid, conditions. The ground is often covered with a moderate grassy under- growth, not permitted to grow out of control. Around the base of each trunk, the cleared circles are maintained. Epiphytic growth on the lower part of each trunk is kept to a minimum, but above this is often a profuse epiphytic flora, mostly ferns (fig. 4). Such plants cause no harm to the oilpalms themselves, although they may reduce the crop by concealing loose ripe fruit. Long-climbing plants are discouraged, since they may reach into the crowns and interfere with pollination. Figs and other woody plants are removed to prevent loss of vigour in the host palm. es Se SS RS FIGURE 2. Plantation of oilpalms, nine years old, in Johore, showing pruned leaf-bases. Most of the ferns begin growth in the pockets of debris and humus accumulated in the palm leaf-bases (fig. 3) and which retain sufficient moisture for their continued growth except during unusually dry periods. Once established, those species with creeping rhizomes or runners soon spread. The small-fronded epiphytes with long- creeping rhizomes appear around the tenth year, by which time spores from terrestrial species on the floor of the plantation have also begun to grow epiphytically on the trunks. After 18-20 years rotten leaf-bases begin to fall, revealing patches of the surface of the trunk. By 23-25 years large expanses of bare trunk surface can be seen (fig. 5) or the trunk may be smooth for the greater part of its length. Long-creeping rhizomes tend to bind the leaf-bases and prevent them falling, often leaving masses of vegetation suspended. The flora of the trunk changes as the leaf-bases fall. Many epiphytes drop to the ground — some continue to grow there, others do so where they had been thrown on heaps of decaying matter in the inter-rows. 96 FERN GAZETTE. VOLUME 12 PART 2 (1980) FIGURE 3. Trunk of mature oilpalm with an abundant growth of epiphytes, predominantly Davallia denticulata and Nephrolepis biserrata. Some of the oldest of the original oilpalm plantings, dating from 1917-20, in Johore and Selangor were visited in 1976. The trunks of these palms varied, presumably with the planting material used. Some were quite smooth up to the crown, whilst others had retained all their leaf-bases which were not pruned as short as present-day standards demand. The smooth trunks had very few epiphytes growing on them but had retained some suspended tangled masses. Growth on the trunks with the longer leaf-bases was not luxuriant. Over a period of three years, 271 plantations and small-holdings, varying from recent plantings, to ones of 59 years old, were visited in West Malaysia (fig. 1). Of these 139 supported a significant flora of well-grown epiphytic ferns, 54 had trunks still being regularly cleaned of epiphytes, and the remainder were too young to have developed a significant fern flora. Most plantations cultivated palms of a range of ages, each block consisting of palms of the same age, and ground conditions varied from place to place in the plantation. The epiphytic ferns of groups of ten or more adjacent oilpalms, situated well within the boundary of the plantation, were recorded at each site and notes made of the terrestrial fern flora there. The total of 55 ferns and two fern-allies recorded in mature oilpalm plantations and small-holdings are listed in table it DEVELOPMENT OF THE EPIPHYTIC FERNS WITH OILPALM AGE The trunk of a mature oilpalm elongates at the rate of 30-60 cm per annum and can attain a height of more than 24 m. The surface of the trunk is not immediately exposed because the dying leaves are unable to form a clean abscission layer and they break off some distance above the base of the stalk. Under natural conditions these long leaf-bases remain attached to the trunk, slowing rotting until, about 20 years later, they fall and expose the trunk surface. In commercial plantations, trunks are tidily trimmed until they resemble elongated pineapples. Debris from the crowns of 2 | PIGGOTT: FERNS OF OILPALM PLANTATIONS ul = yuNnd} Buimoib ‘nuebbuas | wyjedjio ue uo esosgaje; eayeAD AyjeoizAudida “) JYNOlsA ‘snpiu wniuadsy si ayAydida usa} ulew ayy "QdejiNs UNI} ay} BuisOdxea “uajjey aney saseq “J28) UOJOA YOIYM WO1y WIedIIO "G AYNOIS P oe * "yUNI} WuyedjIO Ue JO Saseq-jea| JOYS ay} ul payejyNuinose snwiny ay} ul Buimoib epesasig sidajosydanyy yO sjuejd asinjewuy “p AYNO!4 98 FERN GAZETTE: VOLUME 12 PART 2 (1980) the palms accumulates in the short leaf-bases, gradually rotting to form humus. The pinnate leaves allow rain to penetrate the canopy and their bases channel it towards the trunk. Many ferns and small climbing plants, some figs and occasionally orchids develop as epiphytes in this habitat. Rats, snakes and other small animals frequently make their homes there too. Carefully calculated quantities of fertiliser are regularly applied to the palms to ensure optimum growth and maximum yield. In the early years of the industry the TABLE 1: TOTAL NUMBERS OF FERNS AND FERN-ALLIES RECORDED GROWTH GROWTH HABIT SPECIES HABIT SPECIES aT Acrostichum aureum L. E itt N. biserrata (Sw.) Schott t A. speciosum Willd. E N. radicans (Burm. f.) Kuhn ) T Adiantum latifolium Lam. E Ophioglossum pendulum L. e T Amphineuron opulentum e Phymatodes longissima (B1.) (Kaulf.) Holtt. J.Sm. E Asplenium glaucophyllum e P. nigrescens (BI.) J.Sm. v.A.v.R. E P. scolopendria (Burm.) Ching a A. longissimum BI. e < Pityrogramma calomelanos (L.) 5 A. nidus L. Link e A. pellucidum Lam. e ah Pleocnemia irregularis (Pres|) A Blechnum indicum Burm. f. Holtt. Af B. orientale L. e T Pronephrium triphyllum (Sw.) e T Christella arida (Don) Holtt. Holtt. e lk C. dentata (Forsk.) Brownsey t Pteridium caudatum (L.) Maxon & Jermy. var. yarrabense Domin e al C. parasitica (L.) Lev. if Pteris biaurita L. e 1h C. subpubescens (B\.) Holtt. e t P. ensiformis Burm. e T Cyathea /atebrosa (Wall.) Copel. fr) t P. longipinnula Wall. i Cyclosorus gongylodes (Schkur) ii P. semipinnata L. Link e Li P. tripartita Sw. E Davallia denticulata (Burm.) e T P. vittata L. Mett. e Pyrrosia angustata (Sw.) Ching t Dicranopteris linearis (Burm.) e P. longifolia (Burm.) Morton Underwood var. subpectinata t Schizoloma ensifolia (Sw.) (Chr.) Holtt. J.Sm. e t Diplazium asperum Bl. e T Sphaerostephanos heterocarpus e aT D. esculentum (Retz.) Sw. (BI.) Holtt. E Drymoglossum piloselloides (L.) E ili S. polycarpus (B\.) Holtt. Pr. E aly Stenochlaena palustris (Burm.) e Drynaria sparsisora (Desv.) Bedd. Moore e aly Taenitis blechnoides (Willd.) E Goniophlebium verrucosum Sw. (Hook.) J.Sm. t Tectaria vasta (BI|.) Copel. 7 Ap Lygodium flexuosum (L.) Sw. E Vittaria elongata Sw. © Lh L. longifolium (Willd.) Sw. E V. ensiformis Sw. e T L. microphyllum (Cav.) R.Br. E aT Microlepia speluncae (L.) Moore FERN ALLIES E Microsorium punctatum (L.) e Psilotum flaccidum Wall. Copel. E P. nudum (L.) Beauv. E Nephrolepis acutifolia (Desv.) Chr. ‘E’ denotes the species as a common epiphyte on oilpalm trunks: abundant, locally abundant or of frequent occurrence. ‘e’ denotes an epiphyte of less frequent occurrence. ‘T’ denotes a terrestrial fern common on the floor of the plantation: abundant, locally abundant or frequent. ‘t’ denotes a terrestrial fern of less frequent occurrence. Epiphytes which had obviously recently fallen from older palms were discounted. PIGGOTT: FERNS OF OILPALM PLANTATIONS FIGURE 9. Vittaria elongate and V. ensiformis FIGURE 8. Fertile plant of Pityrogramma calomelanos growing as an epiphyte on an FIGURE 7. Goniophlebium verrucosum grow- concealing an oilpalm trunk. ing abundantly on an oilpalm in Johore. oilpalm. ole 100 FERN GAZETTE: VOLUME 12 PART 2 (1980) palms were cultivated until they were too tall to harvest, but today it is considered economic to replant when the palms are about 25 years old. The debris in the leaf-bases of very young palms consists almost entirely of soil-splash. As humus, mostly derived from the inflorescences, accumulates and as the habitat becomes shadier, the number of fern species growing there increases. At first they are the ferns commonly recognised to be epiphytic. Then some normally terrestrial species appear, growing in the lowest leaf-bases first. When the palms are about ten years old, the distribution of species is fairly uniform on all trunks. Nephrolepis biserrata, Goniophlebium verrucosum and Davallia denticulata are, however, more abundant than other species. Then the lower trunks often become mossy and the number of plants of Vittaria spp. increases. N. radicans, N. acutifolia, Asplenium longissimum, Microsorium punctatum and Phymatodes scolopendria become conspicuous and the distribution of species changes. Some trunks support a wide variety of ferns, whilst others support few. The normally terrestrial species begin to disappear. By the eighteenth year, many of the leafbases have become rotten. Slow at first, they begin to fall away. The process accelerates and the trunk surface becomes exposed. The long rhizomes of some epiphytes are so tightly bound round the trunks that they remain, others are displaced and fall to the ground. Stenoch/aena palustris, if present, and WV. biserrata usually continue to grow in and near the crown of the palm. G. verrucosum and D. denticulata frequently remain high on the trunks too. Pyrrosia spp., Phymatodes spp. and M. punctatum may continue to climb on the smooth surface. In some areas Drymoglossum piloselloides adheres patchily. Vittaria spp. continue to grow amongst mosses near ground-level. Small young plants of A. nidus are common on oilpalms. In well-maintained plantations they are removed before they reach any appreciable size, as large ‘nests’ would trap too much loose fruit and reduce the crop. Similarly, well-grown plants of Drynarta sparsisora are rarely seen. Ophioglossum pendulum could be more common than has been recorded here. The limp fronds hang beneath the stiffer fronds of other ferns and are not easily seen. They are particularly difficult to detect when growing amonst the ribbon-shaped fronds of Vttaria spp. Table 2 gives ages of palms when epiphytic fern species were recorded. A charcteristic of the genus Nephrolepis is its ability to reproduce vegetatively by means of runners and spreads rapidly. Both WV. biserrata and N. acutifolia were recorded on oilpalms long after the leaf-bases had fallen. The oldest palms Gn which N. radicans was recorded were still losing their leaf-bases. Further observations are needed to determine if this species remains on palms with smooth trunks. With the exception of three species, all the fern epiphytes on old palms with smooth trunks possess runners or long-creeping rhizomes which carry them into the crown of the tall palm or bind them to the trunk. The exceptions are A. nidus, which has a very tenacious growth habit, and the two V/ttaria spp., which root in the mosses growing on the lower part of the palm trunk. ORNAMENTAL OILPALMS AND ‘ESCAPES’ Specimens of E/aeis guineensis are grown in many botanical gardens, out-of-doors in the tropics and in hot-houses in temperature climates. They are also common as ornamentals, often under conditions which would horrify a professional planter. Others have appeared on roadsides, where fruit has fallen as the crop was being taken to the factory. PIGGOTT: FERNS OF OILPALM PLANTATIONS 101 Such palms may grow well, but they are not subject to the strict regime of the plantations. In gardens the dead and dying leaves are not removed unless they become too unsightly, and the ‘escapes’ receive no attention at all. Often the base of the trunk is overgrown with terrestrial plants. Stenoch/laena palustris, Dicranopteris linearis, Lygodium flexuosum and L. microphyllum scramble and climb on the trunks, some- times reach into the crowns. Debris and humus do accumulate, but in the deep shade of the long, broken leaf-bases. Few epiphytes find this an ideal habitat and only WN. biserrata, Goniophlebium verrucosum, Davallia denticulata’ and Microsorium punctatum have been recorded. TABLE 2: DISTRIBUTION OF EPIPHYTIC FERN SPECIES IN RELATION TO AGE OF OILPALM Age of palm in years Occurrence (%) SS eS in mature ; Earliest Commonly Latest plantations epeciss record recorded record visited Nephrolepis biserrata 5 5 59 99.3 Stenochlaena palustris 5 5 40+ 60 Asplenium nidus 5 8 59 58 A. longissimum 5 8 26 50 Goniophlebium verrucosum 5 6 59 KT Vittaria elongata 5 10 40+ 54 Davallia denticulata 6 7 59 80 Pityrogramma calomelanos 6 7 12 20 Sphaerostephanos polycarpus 6 7 15 12 Microlepia speluncae 6 8 23 23 Phymatodées spp. 6 10 59 43 Nephrolepis radicans 6 8 25 7 Pteris spp. 7 9 13 8 Asplenium glaucophyllum 7 12 20+ 9 Drymoglossum piloselloides 7 12 30+ 12 Drynaria sparsisora 7 12 40+ 8 Vittaria ensiformis 8 12 40+ 29 Microsorium punctatum 9 16 30+ 11 Nephrolepis acutifolia 10 12 40+ 11 Pyrrosia spp. 10 16 40+ 11 Ophioglossum pendulum 10 15 25 4 Asplenium pellucidum = 12 = 0.7 Other Thelypteridaceae 5 7 15 18 Other ‘terrestrial’ spp. 5 8 15 14 Psilotum flaccidum — 20 = 0.7 P. nudum 15 25 - 2 102 FERN GAZETTE: VOLUME 12 PART 2 (1980) ACKNOWLEDGEMENTS | am most grateful to Dr. R.E. Holttum for checking, and where necessary correcting, the identification of the specimens and for his interesting comments. | wish to thank the Agricultural Corporation of Burma, Kenram Philippines Inc., Kumpulan Guthrie Sdn. Bhd., Menzi Agricultural Corporation and many others who, knowingly or not, co-operated and contributed to this article. In particular | thank Mr. and Mrs. A.L. Africa Jr., Mr. and Mrs. O. Grino, U Myint Maung and, last but not least, Mr. C.J. Piggott for their interest and help. REFERENCES CORNER, E.J.H. 1966. The Natural History of Palms. Wiedenfeld and Nicolson, London. HARTLEY, C.W.S. 1977. The Oil Palm. Longman, London. HOLTTUM, R.E. 1966. Ferns of Malaya (Revised Flora of Malaya, Vol. Il) 2 ed. Govt. Print. Office, Singapore. WHITMORE, T.C. 1977. Palms of Malaya, 2 ed. Oxford University Press, Kuala Lumpur. REVIEW THE PTERIDOPHYTES OF SURINAME by K.U. Kramer, 1978. Naturweten- schappelijke Studiekring voor Suriname en de Nederlandse Antillen, No. 93. Utrecht. 24 x 16cm, 198pp. This work includes over 300 species, subspecies and varieties in 63 genera of which about one quarter are described as being rare. Although there are no specific descriptions as the subtitle indicates (An Enumeration with Keys of the Ferns and Fern Allies) each entry is a rich source of references to the literature. The genera are keyed out in one of two ways: by a general key which distinguishes nine of the most distinctive groups such as the lycopods, aquatics, filmies and the gleicheniaceous members, whilst the great majority are treated by a multi- access key which is an innovation in fern literature. Every genus has 7 or 8 alternate characters each represented by a letter of the alphabet, thus giving a letter formula. This has the advantage that it very clearly brings out the features in common or separating genera. However, the convention used is not sufficient to characterize every genus by a different formula and in such cases an additional note is given to distinguish the components. For example, Asp/enium and Blechnum both have the same letter formula ACGIOSWX but are finally separated from one another by the presence of the sorus On a free vein in the former case or on a commissure parallel to the costa in the latter. The keys to the species are of a conventional type. Recent important literature published on a genus is quoted and in a number of cases where full monographic work is not available geners such as Po/ypodium, Thelypteris, Hymenophyllum and Trichomanes are treated in the broad sense. For each taxon, in addition to relevant literature and synonomy, the geographical distribution is given, together with a sentence on its ecology in Suriname and an indication of its abundance. A valuable feature which should help to stimulate collectors is the inclusion of several taxa which have not yet been found in Suriname but whose presence may be reasonably expected from the geographical distribution. T.G. WALKER SS FERN GAZ. 12(2) 1980 103 ON THE REPORTED OCCURRENCE OF ASPLENIUM CUNEIFOLIUM AND A. ADIANTUM-NIGRUM IN THE BRITISH ISLES ANNE SLEEP Department of Plant Sciences, The University, Leeds, LS2 9JT ABSTRACT Preliminary experimental evidence is presented which shows that British serpentine populations of “A. cuneifolium” are allotetraploid. In Europe, the Asplenium adiantum-nigrum aggregate consists of a complex of three closely-related taxa, two diploid species, A. onopteris L. and A. cune/folium Viv. (with n = 36), and an associated allotetraploid, A. adiantum-nigrum L. (n = 72), which has been shown by Shivas (1969) to be derived from a cross between these two diploid taxa followed by chromosome doubling. A. adiantum-nigrum is a very variable species and it is found throughout most of Europe except the east and the extreme north (Jalas & Suominen, 1972). A. onopteris is a Mediterranean species with an outlier in south-west Ireland, while A. cune/fo/ium, which occurs in central and eastern Europe, has particular edaphic requirements, being found exclusively on serpentine and other ultrabasic rocks. Although it has been pointed out in the past that A. cune/fo/ium could possibly occur in suitable habitats in Britain, the first confident claim of its occurrence in this country was by Roberts & Stirling in 1974. These authors collected material of the Asplenium adiantum-nigrum aggregate from 11 Scottish serpentine localities; three plants from two of these sites (Glen Lochay, v.c.88 and Glendaruel, v.c.98) were examined cytologically and were said to have given diploid counts of n = 36. Plants from other serpentine localities in Scotland were reported to be morphologically identical with these two populations, and were therefore assumed to be diploid also. Roberts & Stirling pointed out the morphological similarity between all of their material and plants of A. cune/folium from Europe, and, despite the inadequacies of their cytological data, unhesitatingly referred all the material from the Scottish serpentine localities to Asp/enium cuneifolium Viv. Subsequently, at the request of these authors, chromosome counts were made in Leeds on material collected by Stirling from the Scottish localities listed in his joint paper. The diploid counts previously reported by them could not be corroborated; tetraploid counts of n = 72 were consistently obtained (Sleep et al., 1978). To date, no diploid counts of the Asp/enium adiantum-nigrum aggregate from Scottish serpentine localities have been recorded. Counts have now been made from 8 of Roberts & Stirling’s listed sites (including the two populations from which diploid counts were previously reported) and all have been clearly tetraploid. Concurrently, a plant of the A. adiantum-nigrum aggregate from Corsica, which on morphological grounds one would have no hesitation in referring to A. cuneifolium Viv., was reported to be tetraploid by Deschatres, Schneller & Reichstein (1978). This plant was so close to the central European populations of A. cune/folium Viv. in general appearance that it was suggested that the Corsican plant could be an autotetraploid derivative of A. cune/folium. In gross morphology, spore size and chromosome number the Scottish plants corresponded well with material of this Corsican tetraploid, and it was thought possible that the Scottish material could also be regarded as an autotetraploid form of A. cuneifolium. However, without further evidence, it could equally well be 104 FERN GAZETTE: VOLUME 12 PART 2 (1980) interpreted as a form of A. adiantum-nigrum which, on serpentine, could produce a type of morphology closely resembling A. cune/folium (Sleep et al., 1978). In order to determine which of these two possibilities is correct, a hybridization programme was set up at Leeds using material of putative autotetraploid A. cune/folium from Scotland and Corsica. Preliminary results from this experimental investigation are now available and are reported below. In order to determine if a plant is a//otetraploid (i.e. derived by hybridization between two distinct diploid species, followed by chromosome doubling) or autotetraploid (i.e. derived by chromosome doubling from a single diploid species), attempts are made to synthesize “‘wide hybrids’, i.e. a hybrid between the tetraploid under investigation and either an unrelated diploid (e.g. Phy/litis scolopendrium) or an unrelated allotetraploid species of known parentage. If the Scottish serpentine material is indeed an autotetraploid derivative of A. cuneifolium, it can be expected to show at meiosis chromosome pairing between its constituent genomes (autosyndesis), and this pairing can be expected to be constant in any hybrid between it and another unrelated species. If, however, the material under investigation is an allopolyploid which has arisen as a result of hybridization between two cytologically distinct diploid species, then it can be expected to show complete failure of chromosome pairing in any hybrid which involves a species with which it is unrelated. This is shown diagrammatically below: The genomes (chromosome sets) involved can be represented by letters, as follows: C C=A. cuneifolium CCC’ C’ = autotetraploid A. cuneifolium O O=A. onopteris CCO O =A. adiantum-nigrum Base number in Asplenium = 36. A. Comparative behaviour shown by crosses between (a) an autotetraploid and (b) an allotetraploid and an unrelated diploid species (PP). (a) (b) ELSES Beam RP SS MCI Oe Triploid | | Triploid hybrid CC’P Gor hybrid 36 pairs + 108 univalents 36 univalents B. Comparative behaviour shown by crosses between (c) an autotetraploid and (d) an allotetraploid and an unrelated allotetraploid species (XXYY). CCC C’ x XXYY Xx CCOO — Tetraploid | | Tetraploid hybrid CC’'XY KY GG hybrid 36 pairs + 144 univalents 72 univalents The underlining bars indicate the constitution of bivalents formed at meiosis. The chromosome pairing behaviour in wide hybrids can thus be used with a high degree of certainty in determining if a given tetraploid is auto- or allopolyploid. Attempts were therefore made to cross cultures of putative autotetraploid A. cuneifolium and A. adiantum-nigrum with unrelated species. Several different hybrid SLEEP: ASPLENIUM CUNEIFOLIUM AND A. ADIANTUM-NIGRUM 105 combinations have been synthesized, but the crosses which specifically concern us here involve A. kobayashii, a rare Japanese allotetraploid that is derived from a cross between Camptosorus sibiricus Rupr. and Asp/enium incisum Thunb. This species has already proved itself to be a valuable standard since, in investigations of this kind (Lovis, Brownsey, Sleep & Shivas, 1972), it both hybridises easily with other unrelated species and, because of its spectacular inter-generic origin, shows no homology between its constituent genomes. This preliminary report concerns hybrids which were synthesized between A. kobayashii and tetraploid plants identified as A. cune/folium from the upper Deveron valley, Bridgend, on the Banff/Aberdeenshire border (the 7th locality listed by Roberts & Stirling, 1974). This cross proved easy to produce, and 91 hybrid plants were obtained. The hybridization technique used is now standard at Leeds and was described in detail by Lovis (1968). In the cytological investigation, developing sporangia were fixed, stained, and the preparations made permanent according to Manton (1950). Meiosis has been examined in the nine of these hybrids which have so far become fertile. These plants incorporate two different stocks of A. kobayashii, and three different parental plants of “A. cune/folium”, although all from the single locality mentioned above. Material attributed to A. cune/folium from different places has been incorporated into other hybrids which have not yet become fertile. All of the hybrid plants studied have proved to be tetraploid (as was expected) and show almost complete failure of pairing at meiosis, most of the chromosomes (132-144) appearing as univalents. The number of bivalents observed ranges from O to a maximum of 6 per cell. There is thus no effective homology between any of the four genomes present in this hybrid (two from the “cune/folium” parent and two contributed by A. kobayashii), from which it follows that both parents must be of allopolyploid origin. This result, besides confirming allopolyploidy in A. kobayashii, shows unequivocally that the culture of “A. cune/folium” investigated must be an allotetraploid. The number of bivalents observed (0-6, out of a possible maximum of 36) is too few to invalidate this conclusion. The evidence presented here from the chromosome pairing behaviour in synthetic hybrids thus demonstrates clearly that one of the Scottish serpentine populations which has been claimed to be A. cune/folium by Roberts & Stirling (1974) is not, as had been suggested, autotetraploid, but is allotetraploid in origin. This observation has obvious relevance to the several recent references to the occurrence of A. cuneifolium in the British Isles (Roberts, 1979(a) and (b); Scannell, 1978). For example, A. cuneifolium has now been claimed to occur also on serpentine outcrops at the Lizard in Cornwall (Page & Bennell, 1979). Plants from these Cornish serpentine populations have not been examined cytologically, but the spore measure- ments given in this paper (range 37-51u) agree very well with those published by Roberts & Stirling (range 36-46) for their material from the Scottish serpentine localities which is now known to be tetraploid. It therefore seems highly probable that these Cornish plants from Kynance Cove will also prove to be tetraploid. Additional evidence, albeit indirect, is available which suggests very strongly that these populations are in fact allotetraploid. Lovis & Vida (1969), in their elegant experimental resynthesis of X Asp/lenophyllitis jacksonii, used material of the A. adiantum-nigrum aggregate from Kynance Cove in their hybridization programme. | have had the opportunity to study Professor Lovis’ collections of A. adiantum-nigrum from this locality; his specimens bear a striking resemblance to the plants from Kynance Cove described by Page & Bennell as being attributable to A. cune/fo/ium, as well as to Scottish serpentine material attributed to the same taxon, and, although it has not yet been possible to trace the Lovis voucher specimen, it seems very likely that he 106 FERN GAZETTE: VOLUME 12 PART 2 (1980) incorporated into his hybridization programme a specimen of the same material which Page & Bennell are now referring to A. cune/folium. \n order to check this, one of Professor Lovis’ specimens was sent to Dr. Page, who confirmed the similarity of the material. Lovis & Vida obtained two hybrids between A. adiantum-nigrum from Kynance Cove and Phyllitis scolopendrium, both of which show almost complete failure of chromosome pairing at meiosis. There is clearly no homology between either of the genomes contributed by the Asp/enium parent, which must therefore be an allotetraploid. What, then, is the nature of this allotetraploid? In view of its morphology, there seems little doubt that it is an extreme form of Asp/enium adiantum-nigrum L. Evidence for this view may be obtained by crossing the material suspected of being A. adiantum-nigrum with the typical form of that species. If the two are in fact the same, the hybrid between them can be expected to show complete pairing of the chromosomes at meiosis. Material of the plants identified as A. cune/folium from the serpentine at Bridgend (which had already been incorporated into the hybrids with A. kobayashii/) was crossed with A. adiantum-nigrum from Guernsey. Hybrids proved very easy to obtain, and no less than 104 were produced from the same number of inseminations. In this type of cross, it can be very difficult to determine if the plants obtained from a hybridization attempt are in fact hybrids, or just selfed progeny arising from self-fertilisation of the female parent. In this case, the parental stocks were of distinctive morphology, and hybrids between them could be recognised by their intermediate appearance as well as by hybrid vigour. Some specimens also showed a small amount of spore abortion, although others, despite their intermediate morphology, produced uniformly good spores. A full report of this investigation, with photographs, will appear elsewhere, but the preliminary results, since they are clearly pertinent to the problem under discussion, can be presented here. Two hybrids have so far been analysed: both yielded several cells which showed virtually complete chromosome pairing at meiosis, a result which is completely in agreement with the belief that the Scottish serpentine populations are neither A. cune/fo/ium nor an autotetraploid derivative of that species, but a form of A. adiantum-nigrum. The evidence presented above clearly shows that the serpentine populations from Scotland and Cornwall which have been identified as A. cune/folium are not that species, but are Asp/enium adiantum-nigrum L. These populations do appear to show a remarkable degree of morphological homogeneity, and it seems most likely that they represent a specialised serpentine form of A. adiantum-nigrum. How far they merit taxonomic recognition (perhaps at the varietal level) is a vexed question which is currently under review. A. adiantum-nigrum is an exceedingly adaptable species. It is of wide distribution, occurring also in North America and in the mountains of central and southern Africa, and it may be found in a wide range of habitats, both natural and man-made. As may be expected of a vigorous allotetraploid, it shows a wide spectrum of morphological variation, completely overlapping with the typical forms of both its parents. | have experienced great difficulty in separating plants of A. adiantum-nigrum and A. onopteris in the Mediterranean region because of their similar appearance. Without a detailed and quantitative morphological study to determine the precise range of variation in A. adiantum-nigrum and its progenitors, it seems premature to make a decision to separate the British serpentine populations of this species as a distinct taxon. It is hoped that further work in progress at Leeds will help to resolve this question. In the meantime, however, it is clear from the present study that the name of A. cuneifolium has been erroneously applied to serpentine populations of A. adiantum-nigrum in the British Isles. All records of A. cune/folium from Britain i a 2 ee ee SS SLEEP: ASPLENIUM CUNEIFOLIUM AND A. ADIANTUM-NIGRUM 107 should therefore be treated as suspect. | am indebted to Dr. Janet Souter for her help in making the cytological preparations used in this investigation. REFERENCES DESCHATRES, R., SCHNELLER, J.J. & REICHSTEIN, T. 1978. A tetraploid cytotype of Asplenium cuneifolium Viv. in Corsica. Fern Gaz. 17: 343-44. JALAS, J. & SUOMINEN, J. (Eds.). 1972. Atlas Florae Europaeae 1. Pteridophyta. Helsinki. LOVIS, J.D., 1968. Fern hybridists and fern hybridising Il. Fern hybridising at the University of Leeds. Fern Gaz. 10: 13-20. LOVIS, J.D., BROWNSEY, P.J., SLEEP, A. & SHIVAS, M.G., 1972. The origin of Asplenium balearicum. Fern Gaz. 10: 263-68. LOVIS, J.D. & VIDA, G. 1969. The resynthesis and cytogenetic investigation of X Asplenophyllitis microdon and x A. Jacksonii. Fern Gaz. 10: 53-67. MANTON, |. 1950. Problems of cytology and evolution in the Pteridophyta. Cambridge. PAGE, C.N. & BENNELL, F.M. 1979. Preliminary investigation of two south-west England populations of the Asp/enium adiantum-nigrum aggregate and the addition of A. cuneifolium to the English flora. Fern Gaz. 12: 5-8. ROBERTS, R.H. 1979a. Spore size in Asplenium adiantum-nigrum L. and A. onopteris L. Watsonia f25 230-250. ROBERTS, R.H. 1979b. The serpentine Black Spleenwort in Norway. Br. Pterid. Soc. Bull. 2: 22. ROBERTS, R.H. & STIRLING, A. McG. 1974. Asplenium cuneifolium Viv. in Scotland. Fern Gaz. 77: 7-14. SCANNELL, M.J.P. 1978. Asplenium cune/folium Viv. in West Galway. /rish Nat. Journ. 19: 245. SHIVAS, M.G. 1969. A cytotaxonomic study of the Asp/enium adiantum-nigrum complex. Fern Gaz. 10: 68-79. SLEEP, A., ROBERTS, R.H., SOUTER, JANET |. & STIRLING, A. McG. 1978. Further investiga- tions on Asplenium cuneifolium in the British Isles. Fern Gaz. 77: 34548. REVIEWS EXPERIMENTAL BIOLOGY OF FERNS, Edited by A.F. Dyer, 1979, 657. 157 x 234 mm, Academic Press, London. Price £37.50. As Adrian Dyer says in his introduction this is the first review of the experimental investigation of the structure, function, development and evolution of ferns for 40 years. The book has two main aims: to make more botanists aware of work already done; and to encourage further experimental work by drawing attention to problems in fern biology. There is no doubt that these aims are achieved. The book begins with a chapter on the Diversity of Ferns: An Ecological perspective (C.N. Page). There is nothing experimental about this — experimental aspects of ecology are dealt with later, by the same author; it is an interesting account of ferns in different habitats throughout the world. Physiologists and other experimentalists, | suspect, would have liked more details about the adaptive biology and morphology seen in the many genera mentioned in this text. P.R. Bell follows giving a stimulating comparison of Fern and Vascular Plant Life Cycles and T.G. Walker in reviewing the Cytogenetics of Ferns, gives a readable account from 160 references tempered with his considerable experience of tropical studies. E.J. Klekowski in Genetics and Reproduction brings the reader into more experimental studies with a bang and plenty of jargon. Ultrastructure has its place: in gametophyte cells (G.D. Cran); and spore wall morphogenesis (J.M. Pettitt) — an account which, like Walker’s above, is contributing 108 FERN GAZETTE VOLUME 12 PART 2 (1980) substantially to fern phylogeny. The next six chapters deal with aspects of the gametophyte which as one might expect lends itself to experimental studies. The Culture of Gametophytes (A.F. Dyer) is dealt with clearly and in depth. Cell division in Protonemata (A.F. Dyer and M.A.L. King), Gametophyte Differentiation and Development (D.L. Smith), Gametophyte Photomorphogenesis (G.P. Howland and M.E. Edwards) and Antheridiogens and Antheridial Development (U.Naf) are highly specialised topics and unavoidably overlap in marginal areas. Some fundamental features of Gametogenesis and Fertilisation are put over in a second paper by P.R. Bell where these processes are reviewed again ultrastructurally. R.A. White’s account of Sporophyte Development is traditional and clear. Apart from a chapter on the Biology and Control of the Bracken Fern (W.W. Fletcher and R.C. Kirkwood), that seems superfluous and out of context, C.N. Page has the closing words on Experimental Aspects of Fern Ecology. It is a very difficult chapter to write and one thing becomes clear — that much “‘experimental”’ (i.e. observational?) ecology was done over 50 years ago. If ever there was a field for experimental research it is in the biology of ferns in relation to their environment. This book was a mammoth task to edit and Adrian Dyer is to be congratulated on having the courage to take it on. From the dates of the bibliographies it took over two years to get through the press, possibly too long for some actively working authors. Sometimes | felt ‘experimental’ meant looking in more detail with modern equipment such as the EM. It would have been worth having perhaps two parts to the book — description and synthesis, the latter possibly discussing subjects like ‘apogamy’ from several angles. Many must feel a lack of descriptive background; for instance | can see no references to Atkinson and Stokey’s life-long studies on gametophytes. On the whole, however, authors have done their homework. | estimate in the region of 1750 references are quoted and even allowing 10% duplicity this is still a lot. Aimed, it is said, at senior undergraduates, postgraduates and established botanists, (all underpaid sectors of the community!) this book may seem expensive but it is really good value and some libraries may need two copies. A.C. JERMY SYSTEMATICS OF POLYSTICHUM IN WESTERN NORTH AMERICA, NORTH OF MEXICO by David Wagner. Pteridologia 1, 64 pp.; 1979. (American Fern Society Special Publication.) In the pteridological world, it has been customary to publish monographs and the longer detailed studies based on theses in either a general journal, or, as in the U.S.A. and Japan, in the publication of the institution where the work has been carried out. Now, for the first time, a new journal, Pteridologia, issued by the American Fern Society, is to be devoted to a broad range of topics specifically concerned with ferns and fern-allies. The ‘Systematics of Polystichum in Western North America North of Mexico” by David Wagner constitutes the first number of this new journal. This article is derived from the author’s Ph.D. dissertation, submitted to Washing State University in 1976; it sets out very clearly the results of a thorough and detailed morphological study, based on the examination of more than 5,000 REVIEWS 109 herbarium specimens, of the Po/ystichum species of western North America as delimit- ed above. Some interesting conclusions are reached, two new taxonomic combinations made, a hitherto-forgotten taxon rediscovered, and a species new to North America postulated. The present study is an extension of the preliminary work of Professor W.H. Wagner on reticulate evolution in the Holly Ferns of the western United States and adjacent Canada, published in 1973. Dr. Wagner begins by giving us his reasons for splitting the western American Sword Fern into two distinct species as P. munitum and P. imbricans. Despite some deficiencies, both in the quantitative measurements and in the presentation of the data, there still seems to be a strong case for his treatment of these two simply-pinnate polystichums; their separation as distinct species is further supported by the virtually complete failure of chromosome pairing shown by a wild hybrid between them, as well as by their occurrence in contrasting ecological habitats. Polystichum andersonii is a tetraploid species bearing a characteristic proliferous bud and it occurs only in western North America; according to W.H. Wagner, it could be interpreted either as an autopolyploid or as an allopolyploid having P. munitum as one parent. David Wagner postulates a bipinnate second parent for his species and claims to have found a herbarium specimen representing it; it should be noted, however, that allopolyplgidy still has to be proved for this species. Polystichum andersonii is a tetraploid species bearing a characteristic proliferous bud and it occurs only in western North America, and according to W.H. Wagner, could be interpreted either as an autopolyploid or as an allopolyploid having P. munitum as one parent. David Wagner postulates a bipinnate second parent for this species and claims to have found a herbarium specimen representing it, although allopolyploidy has still not been proved. Some confusion in the past about the nature and relationships of the large, divided polystichums of the P. braunii/P. andersonii group has been resolved by Dr. Wagner’s discovery of the existence of P. setigerum (= P. alaskense Maxon), a species which was described by Presl in 1825 but which has remained un-noticed by generations of pteridologists until now. Living material corresponding to this species was collected and cytological investigation showed 123 regular paired chromosomes at meiosis. P. setigerum is the first sexually-reproducing hexaploid to be reported in Polystichum (although an octoploid, P. fa/cine/lum, occurs in Madeira), and Dr. Wagner is to be congratulated on its discovery. It is considered possible that P. setigerum arose as a cross between the bipinnate P. braunii and P. munitum, followed by chromosome doubling; my own studies on the synthetic hybrid between these two species and on herbarium specimens of P. alaskense confirm this suggestion. It is, however, surprising to find that no wild hybrids within this group have ever been collected. | would expect the triploid hybrid between P. munitum and P. braunii to occur in nature, particularly in view of the ease with which it can be synthesized in the laboratory. Presumably routine spore examination to detect possible hybridity has been carried out on all the thirty specimens of P. setigerum cited as good species. It would also be interesting to know if spore characters can provide a useful indication of the level of ploidy, or confirm the suggested origin from P. braunii and P. munitum. The proposed ancestry of three western North American amphidiploid species is discussed in detail, and the conclusions regarding the phylogeny and relationships of the five diploid and five tetraploid species under consideration are then summarised; a brief survey of their ecology and phytogeography is also given. The work concludes with a usable key to Po/ystichum in western North America, followed by a detailed account of the taxonomy of the thirteen species within the area studied. Useful maps showing the American distribution of these species are included, as well as a list of 110 FERN GAZETTE: VOLUME 12 PART 2 (1980) representative specimens of each taxon. This is a very carefully executed and detailed taxonomic study, and many of the problems which caused confusion in the western American polystichums have now been unravelled. | would, however, like to see some experimental hybridization work and genome analysis carried out on this fascinating and closely-related group of species. It could confirm many of the findings presented here as well as clarify some of the points which at present must remain uncertain. ANNE SLEEP THE PTERIDOPHYTES OF KANSAS, NEBRASKA, SOUTH DAKOTA AND NORTH. DAKOTA by Aleta Jo Petrik-Ott, Beihefte zur Nova Hedwigia 61:5-332; 1979. J. Cramer, Vaduz, Price DM100,-- (£25.00). This book is a local Flora of four States in central USA and its aim is to collate information on the distribution of all pteridophytes (some 90 taxa) in those States from 24 major works and many smaller papers (listed in the bibliography, pp. 299-309). The author, herself, has contributed much in field-work in recent years and maps of the States, showing presence on a county basis is an effective method of presenting the data. Hopefully this will stimulate local botanists to fill in many new localities and in the reviewer's opinion this is the gist of, and the most useful part of the book. The distribution of each species is also given on a map of the N. American continent but as this is on a State for State basis, the maps are misleading often showing a continuous distribution from east to west for example, when there is really a disjunction caused by the Mississippi valley. Each species is well-drawn (by the author). The text gives verbose and not always clear descriptions of each species; genera are not described so many generic characters are repeated. This aspect has been covered of course in several recent books and much better. The text also includes detailed localities of all specimens seen which to my mind are totally unnecessary. This is a clearly printed, well bound book and although much of the cost will be absorbed by subscribers who will get the book in the Series, whether they want it or not, it is still expensive to the individual purchaser. THE PTERIDOPHYTA OF FERNANDO PO (Contributions to a Flora of the island) 1 by G. Benl in Acta Bot. Barcinonensia 31:1-31; 1978 (publ. 1979). The first part of this work covering Lycopodiaceae, Selaginellaceae, Psilotaceae, Schizaeaceae, Gleicheniaceae, Osmundaceae, Cyatheaceae contains often detailed ecological notes resulting from the author’s own field work. A.C. JERMY ee Ne ieee a See ee ee REVIEWS 111 FLORA OF THAILAND Vol. 3 Part 1 Pteridophytes by M. Tagawa and K. |watsuki, pp. 128; June 1979. Obtainable from Thai National Documentation Centre, Thailand Institute of Scientific and Technological Research, 196 Phahongothin Road, Bankok 9; price U$ $8.00. This volume covers all families up to and including Dennstaedtiaceae of the 31 families listed for Thailand by the authors in Mem. Fac. Sci. Kyoto Univ. Biol. 5: 67-88; 1972. A key to all families is given with keys to genera and species of the families dealt with in this volume. The work continues the high standard of this international project giving regional synonymy, clear and concise descriptions, detailed distribution in Thailand and briefly of the complete range. The ecological notes are often based on field experience of the authors in Thailand and elsewhere in S.E. Asia. CAPE PENINSULA FERNS by J.P. Roux, 66pp., 1979. Published and available from the National Botanic Gardens of South Africa, Kirstenbosch, Newlands, Cape Province S.A., price R.2.50 (about £1.50) [please remit in S.A. currency.] This is a well produced guide to the fern species of the Cape Peninsula, with line drawings and identification keys to all recognised species (56), by a horticulturist who has himself drawn the plates; most of which are very convincing. The text is clear, giving references to standard works, concise descriptions and a selected list of the few specimens examined. Hopefully this little book will encourage more local fern enthusiasts and as many of the species mentioned are sold as houseplants in Western Europe it should have a good sale here. FLORA DE LA PROVINCIA DE JUJUY, REPUBLICA ARGENTINA (ed. Angel L. Cabrera) Part I! Pteridopitas by Elias R. de la Sota, 275 pp., 1977. Coleccion Cientifica del Instituto Nacional de Tecnologia Agropecuaria (INTA) Vol. 13, Buenos Aires; price not quoted. This Flora covers the Andean Province of Jujuy in N.W. Argentina in country ranging from 500 to 6000 m altitude. 161 species of ferns and allies in 50 genera (and 23 families) are dealt with with good keys and full descriptions which are supplemented by excellent illustrations often to species not hitherto illustrated before. This book is worth obtaining for its illustrations alone, although the Spanish text is easy to get to grips with. Part | of this series gives the background and phytogeography of the area; the rest of the 3,500 vascular plants of the Province will be dealt with in eight future parts. A.C. JERMY 12 FERN GAZETTE: VOLUME 12 PART 2 (1980) SHORT NOTES ADDITIONAL FIELD CHARACTERS SEPARATING THE SUBSPECIES OF ASPLENIUM TRICHOMANES IN BRITAIN Habit differences observed by the authors between specimens of the Asp/enium trichomanes group in a number of Scottish stations provide a useful guide to the separation of the two subspecies in the field. Opportunity to discuss these observations with P.M. Benoit and J.W. Dyce and compare them with North Wales plants in the field (see Jermy in Dyce, 1979 : 17) showed similar differences to occur at Welsh sites known to contain the two subspecies. These differences are additional to those which can usually be seen in herbarium material, summarised by Lovis (in Jermy et al., 1978 : 58-60). When growing from steep or vertical rock faces (the most usual situation for both subspecies), specimens which morphologically can be referred to subsp. trichomanes, usually have a habit in which the fronds arch upwards and outwards, away from the rock face. In luxuriant specimens in deep shade they may also arch downwards, in a cascading fashion. By contrast, plants of subsp. quadrivalens, unless growing in very deep shade, usually adopt a habit in which the fronds are closely adpressed to the rock-face, over which they spread in all directions (but especially upwards) in a rather sinuous manner. In both subspecies, stipes and rachides on newly emerged fronds are normally coloured a glossy, deep blackish-brown. In subsp. trichomanes, this colour usually changes rapidly during the first season to a glossy coppery or bronze-red colouration throughout, whilst the blackish-brown colour persists much more permanently in rachides of subsp. quadrivalens. As fronds age, a third difference appears. In subsp. trichomanes, in most specimens the pinnae become particularly readily shed from their rachides (usually during their first autumn and winter), whilst remaining largely leafless, rachides are retained on the plants in sheltered places, apparently for several seasons. By contrast, in most specimens of subsp. quadrivalens, pinnae are retained on their rachides very much longer (usually at least throughout the first winter, except in very exposed situations), and are rarely completely shed before the rachides themselves eventually break away. Differences in environment can cause some modifications to these differences, plants of subsp. quadrivalens, for example, readily shedding their pinnae under conditions of severe drought. Plants of the rarer subsp. trichomanes thus give the overall appearance in the field, in addition to their greater delicacy of structure, of having a gracefully arching frond habit, with the current year’s fronds arising over persistent masses of thin, bronze-red stipes. At a distance, the habit of such plants can look very different from the spreading sinuous or sinusoidal habit of subsp. quadrivalens, and more closely resembles that of Asp/enium viride. Another field character which is worthy of observation is the plane of the pinna lamina. In subsp. trichomanes this pinna-blade is ridged or striate longitudinally often curling under along the side whilst turning up at the tip. Subsp. quadrivalens, on the other hand, has a flatter lamina without such ridges or raised tip (P. Benoit, pers. comm.). Although these differences do not form an infallible guide, they can be of help in separating plants of the two subspecies growing in most typical, broadly similar situations. They are also features which we recommend should be recorded as field annotations on herbarium specimens. SHORT NOTES Aves REFERENCES DYCE, J.W. 1979. [B.P.S.] Meetings in 1979: Criccieth, North Wales. B.P.S. Bulletin 2: 13-18. JERMY, A.C., ARNOLD H.R., FARRELL, L., & PERRING, F.H. 1978. Atlas of ferns of the British Isles. London. A.C. JERMY, British Museum (Natural History) London, and C.N. PAGE, Royal Botanic Gardens, Edinburgh. A NEW FIND OF EQUISETUM X TRACHYODON IN NORTH-WEST SCOTLAND In July 1978, Mr. J. Grant Roger and the author found a horsetail on the western shore of Loch Hope, W. Sutherland (VC108), which was assumed to be Equisetum variegatum Schleich. ex Web. & Mohr. Subsequently, a voucher specimen was sent to Dr. C.N. Page, who determined it as Equisetum x trachyodon A. Br. This record is therefore of some interest since it indicates a further extension in the known range of this hybrid. In Scotland, in addition to its original station in Kincardineshire (Matthews, 1940), it occurs on the Isle of Harris, Isle of Rhum and Isle of Skye (Page, 1979), and recently, Equisetum x trachyodon has also been found in England, on the Wirral coast (Barker, 1979). The present locality is about 150 km NW of the Skye station and it could well occur elsewhere in Northern Scotland. With regard to the local distribution of the putative parents of this hybrid, Equisetum variegatum is not known north of the Assynt district, but E. hyema/e L. occurs on the north side of Ben Hope (Ferreira, 1958), about 8 km SE of the present site. The site was revisited in July 1979, when the present observations were made. The general habitat is a SE-facing shore of Loch Hope (located at the level of full flood), which is irrigated by seepage water originating from calcareous rocks associated with the Moine Thrust Plane that crop out within Arnaboll Wood. The site is about 20 x 4 m in size and over 200 shoots of Equisetum x trachyodon (14 of which had produced cones) were seen, originating from about 60 separate rhizomes. The horsetail occurs here within two distinct plant communities (a) Carex panicea-Campylium stellatum flush and (b) Molinia — Myrica mire. Both communities are described by McVean and Ratcliffe (1962). (a) Carex panicea — Campylium stellatum flush This open flush vegatation has a cover of about 15% and grows amongst small stones up to 7 cm diameter. The sand/silt substrate is permanently moistened by the seepage water from the adjacent wood. The floristic composition of this flush was: *Equisetum x trachyodon (f) tT Ranunculus flammula (o) Molinia caerulea (f) Succisa pratensis (o) Carex demissa (f) Campylium stellatum (o) Nardus stricta (o) Agrostis tenuis (I) Leontodon autumnalis (o) Carex panicea (I) Pinguicula vulgaris (o) Eleocharis quinqueflora (I) Plantago maritima (o) Prunella vulgaris (r) (b) Molinia — Myrica mire Eroded tussocks of this dense mire vegetation occur within the area of Carex panicea — Campylium stellatum flush, on the shoreline. Here, the cover is 90-100% and a thin layer of peat overlies the sand/silt substrate. This community is flushed by the same water as in the previous one, but, in this case, by capillary action. The 114 FERN GAZETTE: VOLUME 12 PART 2 (1980) floristic composition of the mire was: *Erica tetralix (a)t Nardus stricta (o) Molinia caerulea (a) Pinguicula vulgaris (o) Calluna vulgaris (f) Selaginella selaginoides (I) Myrica gale (f) Carex pulicaris (I) Carex panicea (f) Eleocharis quinqueflora (I) Narthecium ossifragum (f) Juncus squarrosus (1) Potentilla erecta (f) Alnus glutinosa (seedlings) (r) Succisa pratensis (f) Plantago lanceolata (r) Equisetum x trachyodon (o) A detailed search of the shoreline for a kilometre both to the north and south of the site failed to locate any additonal material of this horsetail. *Nomenclature of the Flowering Plants & Pteridophytes according to Clapham, Tutin & Warburg (1962). t(a) = abundant, (f) = frequent, (o) = occasional, (I) = local and (r) = rare. ACKNOWLEDGEMENTS ! am much indebted to Dr. C.N. Page for the determination of Equisetum x trachyodon and for the stimulating discussions on the ecology and distribution of this hybrid in Britain. REFERENCES BARKER, M. 1979. Equisetum x trachyodon in Cheshire, new to the English flora. Fern. Gaz. 72: 59-60. CLAPHAM, A.R., TUTIN, T.G. & WARBURG, E.F. 1962. Flora of the British Isles. Cambridge. FERREIRA, R.E.C. 1957. Equisetum hyemale L. in West Sutherland. Trans. Bot. Soc. Edinb. 37: 220. MCVEAN, D.N. & RATCLIFFE, D.A. 1962. Plant Communities of the Scottish Highlands, HMSO, London. MATTHEWS, J.R. 1940. Equisetum trachyodon as a Scottish plant. Trans. & Proc. Bot. Soc. Edinb. 33: 29-32. PAGE, C.N. 1979. Equisetum x trachyodon in Western Scotland. Fern. Gaz. 12: 57-59. R.E.C. FERREIRA, St. Catherine’s, Windermere, Cumbria. PTERIS CRETICA AND SELAGINELLA KRAUSSIANA NATURALISED IN SCOTLAND Boturich Castle, Dunbartonshire, is situated at the south end of Loch Lomond, % mile (1.2 km) north of Balloch and approximately 400 metres from the shore of the Loch. Below the Castle is a large kitchen garden and orchard (now disused), derelict green- houses, coldframes and a potting shed. Recently, two alien pteridophytes, Pteris cretica L. and Selaginella kraussiana (Kze.) A. Br., were found to have become sufficiently well-established here that they can be considered to be naturalised. Both species have been frequently cultivated as glasshouse plants in Britain, and both have presumably established here from former cultivated plants grown in the castle green- houses, although these greenhouses have been disused for approximately 30 years. Selaginella kraussiana, a native to the Azores and tropical West Africa, is hitherto known to have established itself in the more oceanic areas of south and west Britain and Ireland where it can successfuly overwinter, but has only once been recorded to have done so in Scotland (recently in nearby Kintyre, V.C.101 — Kenneth 1979). Similarly, Pteris cretica has previously been recorded in scattered localities only in SHORT NOTES atk southern Britain (e.g. West Kent V.C.16, South Devon V.C.3 and Staffordshire V.C.36 — A.R. Busby, personal communication). Pteris cretica has become established on the interior mortared wall of the old potting shed at Boturich, in dense shade together with Phy//itis scolopendrium. The only sources of light are through the open doorway and through a skylight in the roof. Moisture supply within the shed is limited to that which can enter through the leaking roof. Selaginella kraussiana grows as a prostrate mat in dense shade below the staging of a lean-to greenhouse (back wall facing east). A tall overgrown beech hedge and naturally regenerating alder, sycamore, ash and hazel saplings shade the greenhouse itself. Glass missing from about half of the roof permits a constant supply of rain water to enter during wet weather (annual rainfall is circa 60 inches) and a water tank sunken into the floor in the corner below the staging helps retain humidity during dry conditions. The only other plants growing under the staging and associated with the Selaginella are Phyllitis scolopendrium, Dryopteris dilatata (one small plant) and the bryophytes Brachythecium velutinum, Amblystegium serpens, Ceratodon purpureus and Lophocolea bidentata. All these species have colonized pockets of soil in crumbling walls and in soil and detritus on the floor and amongst broken flower pots. Success of the Se/agine//a has been followed over three seasons and it appears to thrive, although restricted to the shaded area below the staging in the lean-to. It occurs in none of the other buildings. The Pteris seems to have a much more tenuous hold, and only three individual clumps, none looking particularly healthy, have been found. Both aliens suffered considerably in vigour following the particularly severe winter of 1978. One plant of the Pteris nevertheless survived, and the Se/aginel/la recovered health again by the following summer. Voucher herbarium specimens are to be deposited at R.B.G. Edinburgh (E). REFERENCE KENNETH, A. 1979. Plant Records. Watsonia 12: 347. ERIC BIGNALL, Nature Conservancy Council, Mid Danna, Tayvallich, Lochgilphead, Argyll. NEW RECORDS OF ASPLENIUM AND EQUISETUM HYBRIDS IN FRANCE Asplenium x ticinense D.E. Meyer (Meyer, 1961), the hybrid between A. adiantum- nigrum and A. onopteris, has only recently been shown to be a member of the French flora, having been identified (Roberts, 1974) from two herbarium fronds gathered at Vernet-les Bains (Pyrenees-Orientales). On September 1977 R. Prelli found A. x ticinense in northern Brittany near Trecelin, 5 km south of Cap Frehel (C6tes-du-Nord) with the parent species under the cover of Fraxinus excelsior and Quercus pendunculata. Two stocks have been discovered on a slope facing westward. Growing with them were Asp/enium scolopendrium, Polystichum setiferum, Hedera helix, Ruscus aculeatus, /ris foetidissima, Rubia peregrina, Brachypodium silvaticum. It seems likely that this hybrid might occur in other places where the altitudinal ranges of the two parent species overlap. In 1978 we identified two mature fronds of A. x ticinense found at La Blanche, between Saint-Etienne-de-Tinee and Isola (Alpes-Maritimes), at an altitude of c. 900 m, and between Col de Sorba and Ghisoni (Corsica). These specimens were respectively collected by A. Borel and G. Dutartre to 116 FERN GAZETTE VOLUME 12 PART 2 (1980) whom we are indebted for these new records. Specimens from the three sites are large, having a total length up to 60 cm, with the blade up to 27 cm long. The blade is tripinnate with the pinna segments long and acuminate. These segments are curved towards the apex of leaf, except in the specimen from Corsica. Mature sporangia contain only misshapen, defective spores. It is worth noting that records of A. onopteris in the western part of France are questionable (most being misidentification of A. adiantum-nigrum). Near Trecelin, spore size was also studied in the parent species. A random sample of 100 spores from each plant was measured, excluding the perispore. The spores of A. adiantum-nigrum are longer (mean values 32um to 38um) than those of A. onopteris (mean value 26um to 32um). Asplenium x sarniense Sleep (A. adiantum-nigrum x A. billotii), described from Guernsey (Sleep, 1971), has since been found by the authors (March 1978) in the southern coast of Britanny, at Kersicot, 2 km east of La Foret-Fouesnant (Finistre). The habitat of A. x sarniense at Kersicot is a wall partly covered with Hedera helix in the shade of a row of Castanea sativa trees. The parent species grow very near to the hybrid and occur widely in the neighbourhood. Associated species noted include Melandrium diurnum, Dactylis glomerata, Rubus sp., and Polypodium interjectum, all of which are regularly cut in the course of roadside trimming. As noted by Sleep, the hybrid and A. adiantum-nigrum have similar triangular, bi- to tripinnate blades, and triangular pinnae in which the proximal pinnules are large and subdivided. But the pinnules of the upper part of the blade, which are shortly but distinctly stalked, even towards the end of the pinnae, show the influence of A. billotii. These pinnules are oval in shape, rounded at the tips and bear short, mucronate teeth. Spores are abortive. It is interesting to point out that Litardiére (1924) described Asp/enium trojanii from Corsica. He suggested that the plant may be a hybrid between A. bi//otii and A. adiantum-nigrum. Although the locality has been visited, a more thorough investiga- tion of the site is needed. Even if the first parent is A. b///otii the possibility of another species of the A. adiantum-nigrum complex being one parent cannnot be overlooked. Equisetum x font-queri Rothm., the hybrid between &. palustre and E. telmateia, has been identified as a member of the French flora (Hauke, 1966; Page, 1973) from herbarium specimens gathered by Burnat at ‘‘entre le pont du Var et la mer; sables des alluvions du Var’’ (Alpes-Maritimes) on the 20th May 1872. Specimens from this locality can also be seen in herb. Burnat (G-BU !) at Paris (P !), at Lyon (LY !), at Kew (K) and at Stockholm (S-PA). Unfortunately the site has very probaly been destroyed, but R. Prelli found the hybrid in April 1977, at Cagnes-sur-Mer, west of the river Var (Alpes-Maritimes). The plant occupies about 150 sq m of a railway embankment near its junction with the river La Cagne. The Equisetum predominates amongst other norsetails (EF. arvense, E. ramosissimum, E. telmateia). Flowering plants noted included Hordeum murinum, Urtica dioica, Polygonum aviculare. The river side was in the past marshy, but today the stream has been confined by banks. E. pa/ustre has not been recorded in the site, but might have been there in the past. However it still grows 3-4 km away from this site. Another specimen of &. x font-queri from Pyrenees-Orientales is at Montpellier (MPU !). The labels read as follows: ‘‘abonde au rivage du Tech a Elne”’. All the &. x font-queri specimens agree completely with the descriptions given by Page (1973) and Roberts & Page (1979) for the two British stations, and are intermediate in size and morphology between the parent species. As noted by Page, a proportion of the spores are small, misshapen, lack good elaters and are obviously abortive. SHORT NOTES TAT REFERENCES HAUKE, R.L. 1966. A systematic study of Equisetum arvense. Nova Hedw. 13: 81-109. LITARDIERE, 1924. Ann. Soc. Linn. Lyon. n.s. 70: 131-133. MEYER, D.E. 1960..Sw Zytologie der Asplenien Mitteleuropus (XXIV-XXVIII). Ber. Deutsch. Bot. Ges. 73: 86-394. PAGE, C.N. 1973. Two hybrids in Equisetum new to the British Flora. Watsonia 9: 229-237. ROBERTS, R.H. 1974. Asplenium x ticinense D.E. Meyer from a French locality Fern. Gaz. 17: 55: ROBERTS, R.H. & PAGE, C.N. 1979. A second British record for Equisetum x font-queri, and its addition to the English flora. Fern Gaz. 12: 61-62. SLEEP, A. 1971. A new hybrid fern from Channel Islands. Brit. Fern Gaz. 10: 209-211. F.BADRE & R. PRELLI, Laboratoire de Phanerogamie, Museum National D’Histoire Naturelle, 16 Rue Buffon, F 75005 Paris. EQUISETUM X ROTHMALERI NEW TO JAPAN A herbarium specimen of Equisetum x rothmaleri C.N. Page (the very rare hybrid between &. arvense L. and E. pa/ustre L.) has been recently confirmed, collected from “environs de Sapporo” in the northern Japanese Island of Hokkaido. The specimen, at Paris (P), was collected by M. |’Abbe Faurie, on 26 May 1891 (Faurie 6850). Despite the abundance of the two parent species around the north temperate zone, this is only the second known station for this cross. The first (type) locality is on the Isle of Skye, western Scotland (Page 1973), and the Skye and Japanese plants match very closely. Separated by such a great distance, the two must clearly have originated independently. ReEPERENGE PAGE, C.N. 1973. Two hybrids in Equisetum new to the British Flora. Watsonia 9: 229-237. C.N. PAGE Royal Botanic Garden, Edinburgh MACROTHELYPTERIS, A FERN GENUS NEW TO MAINLAND AFRICA While collecting ferns in August 1979 on lan Garland’s farm “Twinstreams’ at Mtunzini, Natal, South Africa, Macrothelypteris torresiana (Gaud.) Ching was found to be locally common on the sides of ditches at the margin of swamp forest and sugar- cane fields not far above sea-level. Schelpe (1969) does not list this fern for southern Africa, nor does Holttum (1974) mention its presence in mainland Africa. (Both these authors have confirmed my identification.) Macrothelypteris torresiana is a wide-ranging species native to the Old World Tropics from the Mascarene Islands to Queensland and Hawaii, which has also become naturalised in the tropics of the New World. It is capable of growth in a wide variety of 118 FERN GAZETTE: VOLUME 12 PART 2 (1980) habitats, some unmodified, e.g. limestone screes in light shade, others of a secondary nature following man’s disturbance, e.g. roadside banks in forest and areas of forest clearance following logging. Although the nearest known localities are in Madagascar and Reunion it seems reasonable to assume that it is of natural occurrence in a secondary habitat at Mtunzini (which lies at approx. lat. 29° 57’ S and 329 45’ E, in the subtropical summer rainfall belt of South Africa) and that it probably occurs else- where along the coast of northern Natal and further north in Africa also. REFERENCES HOLTTUM, R.E. 1974. Thelypteridaceae of Africa and adjacent islands. Jour. S. Afr. Bot. 40: 123-168. SCHELPE, E.A.C.L.E. 1969. A revised checklist of the pteridophyta of southern Africa. Jour. S. Afr. Bot. 35: 127-140. B.S. PARRIS, Botany School, Downing Street, Cambridge. NEW COMBINATIONS IN SOUTH-EAST ASIAN FERNS Preparation of an account of Grammitidaceae for Flora Malesiana and of ecological papers based upon my recent fieldwork in Borneo, New Guinea and the Philippines necessitate the publication of the following new combinations. Crypsinus stenopteris (Baker) Parris comb. nov. Basionym: Polypodium stenopteris Baker, Jour. Bot. 17: 43 (1879). Ctenopteris barathrophylla (Baker) Parris comb. nov. Basionym: Polypodium barathrophyllum Baker, Jour. Bot. 29: 107 (1891). Ctenopteris rufidula (C. Chr.) Parris comb. nov. Basionym: Polypodium rufidulum C. Chr., Ind. Fil. Suppl. 3: 157 (1934) (nom. nov. for P. rufescens Brause (1912) non BI. (1829)). Diplazium angustisquamatum Holttum) Parris comb. nov. Basionym: Athyrium angustisquaamatum Holttum, Gardens’ Bull. . (Singapore) 11: 273 (1947). Diplazium latilobum (Copel.) Parris comb. nov. Zen 74250, (i960) nan, alae Basionym: Athyrium l/atilobum Copel., Univ. Calif. Publ. Bot. 18: 222 ( a Al Rasitde (778. Diplazium megistophyllum (Copel.) Perris-comb=-nev. Tay pnicedes 7: Basionym: Athrium megistophyllum Copel., Philip. Journ. Sci. 56: 475, t.7 (1935). Goniophlebium mehipitense (C. Chr.) Parris comb. nov. Basionym: Polypodium mehipitense C. Chr., Mitt. aus d. Inst. allgem. Bot. Hamburg 7: 159 (1928). Goniophlebium rajaense (C. Chr.) Parris comb. nov. Basionym: Polypodium rajaense C. Chr., Mitt. aus d. Inst. allgem. Bot. Hamburg 7: 159 (1928). Grammitis clemensiae (Copel.) Parris comb. nov. Basionym: Oreogrammitis clemensiae Copel., Philip. Jour. Sci. 12C: 64 (1917). Grammitis maxwellii (Baker) Parris comb. nov. Basionym: Polypodium maxwellii Baker. Kew Bull. 1893: 211 (1893). Grammitis papuensis (v.A.v.R.) Parris comb. nov. Basionym: Polypodium papuense v.A.v.R., Malayan Ferns Supp/. 521 (1917). Grammitis sparsipila (Copel.) Parris, comb. nov. Basionym: Polypodium sparsipilum Copel., Philip. Journ. Sci. 6C: 139 (1911). Grammitis sucklingiana (Baker) Parris. comb. nov. Basionym: Polypodium sucklingianum Baker, Annals Bot. 8: 128 (1894). Grammitis viridula (v.A.v.R.) Parris, comb. nov. Basionym: Polypodium viridulum v.A.v.R., Nova Guinea 14: 41 (1924). Grammitis vittariifolia (C. Chr.) Parris comb. nov. Basionym: Polypodium vittariifolium C. Chr., Ind. Fil. 574 (1906) (nom. nov. for P. minimum Baker (1879) non Aublet (1775) nec Brack. (1854) nec Moore (1857)). Humata subvestita (C. Chr.) Parris comb. nov. Basionym: Humata kinabaluensis var. subvestita C. Chr., Gardens’ Bull. (Singapore) 7: 232 (1934). ~ — SHORT NOTES 119 Phymatopteris glauca (J. Sm. ex Brack.) Parris comb. nov. Basionym: Drynaria glauca J. Sm. ex Brack., Expl. Exped. 16: 54 (1854). Phymatopteris taeniata var borneensis (Christ) Parris comb. nov. Basionym: Polypodium griffithianum var. borneense Christ. Ann. Jard. Bot. Buit. 20: 122 (1906). Pyrrosia dimorpha (Copel.) Parris comb. nov. Basionym: Cyclophorus dimorphus Copel., Journ. Arnold Arb. 10: 180 (1929). Pyrrosia stellata (Copel.) Parris comb. nov. Basionym: Cyclophorus stellatus Copel., Journ. Arnold Arb. 10: 179 (1929). Selliguea bellisquamata (C. Chr.) Parris comb. nov. Basionym: Polypodium bellisquamatum C. Chr., Brittonia 2: 313 (1937). Selliguea gibbsiae (v.A.v.R.) Parris comb. nov. Basionym: Pleopeltis gibbsiae v.A.v.R., Bull. Buit. 11 28: 37 (1918). Xiphopteris bryophylla (v.A.v.R.) Parris comb. nov. Basionym: Polypodium bryophylium v.Awv.R., Bull. Jard. Bot. Buit. 2. 16: 35 (1914). B.S. PARRIS Botany School, Downing Street, Cambridge THE FIRST SYMPOSIUM ON CHINESE PTERIDOLOGY The first Symposium on Chinese Pteridology which was organised by the Botanical Society of China was held from 19-23 October 1979, in West Tian-Mu Shan near Hangchow in Chekiang Province. There were thirty-seven delegates from botanical institutes, universities and natural history museums in fourteen provinces attending the meeting. Previous to the meeting an executive committee of five was elected: they are Chiu Pei-Shi, Shing Kung-Shar, Kung Shien-Shu, Wang Chu-Hao and Zhang Chao-Fang. The meeting was opened by executive chairman Shing Kung-Shar reading a speech by Professor Ching Ren-Chang, the Honorary chairman of the Botanical Society of China. After reviewing the history of research work carried out in Chinese pteridology in the past fifty years, Prof. Ching said: ‘‘Although we have made considerable headway in the past this is only the first step in our long march. We must make great effort to enlarge our team, undertake more thorough field exploration for ferns, and with the help of studies of anatomy, cytology and chemistry, gradually raise the level of Chinese pteridology until almost reaching the molecular level, thereby expounding the origin, evolution and systematic relationships of Chinese ferns, and also their economical uses, for the purpose of the ‘‘Four Modernizations” as proposed by our Central Government’’. These are the most inspiring and glorious tasks before us. | hope each of us will steadily march from victory to victory. Twenty-seven papers including taxonomy, ecology, spore morphology, cytology and chemistry of ferns were read and a warm discussion ensued in the following two days. From this it became clear that studies in Chinese ferns and the preparation for an all-embracing Chinese Fern Flora are in active progress. Taxonomists in several Provinces have been already collecting fern specimens and writing regional fern Floras, among which Fukien, Kwantung, Szechuan, Yunnan and Kiangsi are taking the lead in the work. From the papers read at the symposium one also can see that Chinese pteridologists are using new approaches, such as the SEM, in their work. Everybody present felt the symposium was successful. It has not only offered a chance to exchange scientific views and strengthen mutual connections, it also has laid a better foundation for closer co-operation between pteridologists. We also discussed ways and means of training young workers, popularising fern knowledge among the general public, and introducing wild ferns into cultivation for scientific 120 FERN GAZETTE. VOLUME 12 PART 2 (1980) study and as ornamentals in gardens. Delegates also exchanged opinions as to where and when the second symposium would be held. Two excursions were arranged after the meeting for the delegates to collect and observe ferns in nearby mountains, when an altitude of 1506 m was reached, and to visit the herbarium and fernery at the Botantical Garden, Hangchow. SHING KUNG-SHAR el ib Nae THE FERN GAZETTE Original papers, articles, or notes of any length on any aspect of pteridology will be considered for publication. Contributions will be particularly welcomed on floristic, biogeographical and ecological aspects of ferns and their allies, worldwide. Members are especially encouraged to make use of short notes for reporting pteridophyte field observations and records. All may be illustrated with black and white photographs of good technical quality and line drawings. Short papers can usually be incorporated more quickly than longer ones. Contributions should be sent to: C.N. Page, Royal Botanic Garden, Edinburgh, U.K. The /ast date for receiving notes and articles to make the following early summer number is: 30 November each year Authors should follow the general style of this number. Close adherence to the following notes will help to speed the publication of contributions. NOTES FOR CONTRIBUTORS Manuscripts: Copy should be in English and submitted in double-spaced type with adequate Margins, on one side of the paper only. Abstract: All papers, other than short notes, should include a short abstract, to be set at the head of the main text, indicating the scope of the topic and main conclusions. Headings and sub-headings: These should follow the style of this number. (Primary sub-headings.- are centred capitals. Secondary sub-headings U and L case side roman. Tertiary sub-headings, if necessary, are U and L case side italic.) Numbering of sub-headings should be avoided. 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Final lettering will be added by the editor in a style and size consistent with ‘the journal. Figure captions: Type on a separate sheet from the manuscript — include any necessary details of magnification as submitted (the editor will apply any correction for reduction). Reference lists: Please follow closely the style of this number to speed publication. Lists in other styles may have to be returned to authors for re-typing. Reprints: Twenty-five reprints are supplied free of charge to authors, who may order in advance | a © further reprints which will be supplied at cost (plus postage) if requested at time of returning the first proofs. BOOKS FOR REVIEW Books for review in the Fern Gazette or 8.P.S. Bulletin should be sent to A.C. Jermy, Botany Department, British Museum (Natural History), Cromwell Road, London SW7 5BD. THE FERN GAZETTE VOLUME 12 PART 2 1980 CONTENTS Page MAIN ARTICLES Historical notes on Woodsia i/vensis in the Moffat Hills, southern Scotland — J. Mitchell 65 Polypodium macaronesicum and P. australe; a morphological comparison — R.H. Roberts 69 Studies in the plant sociology of fern-dominated wall communities on the island of Corfu — D.W. Shimwell 75 Ferns of Nainital (Western Himalayas) : an updated list — S.C. Verma & S.P. Khullar 83 The fern flora of oilpalm plantations in West Malaysia — A.G. Piggott 93. On the reported occurrence of Asp/enium cuneifolium and A. adiantum-nigrum in the British Isles — Anne Sleep 103 SHORT NOTES Additional field characters separating the subspecies of Asp/enium trichomanes in Britain — A.C. Jermy and C.N. Page a 4 A new find of Equisetum x trachyodon in north-west Scotland — R.E.C. Ferreira 113 Pteris cretica and Selaginella kraussiana naturalised in Scotland — E.M. Bignal/ | 114 New records of Asp/enium and Equisetum hybrids in France — F. Badré and R. Prelli 115 Equisetum x rothmaleri new to Japan — C.N. Page 117 Macrothelypteris, a fern genus new to mainland Africa — B.S. Parris dle New combinations in south-east Asian ferns - B.S. Parris 118 The first symposium on Chinese Pteridology — K.S. Shing 119 REVIEWS 74, 82, 102, 107-111 (THE FERN GAZETTE Volume 12 Part 1 was published 31 August 1979) Published by THE BRITISH PTERIDOLOGICAL SOCIETY, c/o Department of Botany, British Museum (Natural History), London SW7 5BD. Printed in England by P.7 ¢ COMPASS COLORPRINNT =* Telephone: 01-571 0991 AS i | _ ISSN 0308—0838 VOLUME TWELVE PART THREE 4981 Sek . ; THE JOURNAL OF THE - BRITISH PTERIDOLOGICAL SOCIETY THE BRITISH PTERIDOLOGICAL SOCIETY Officers and Committee for 1981 President James W. Dyce, 46 Sedley Rise, Loughton, Essex, IG 10 tee : Vice Presidents R.E. Holttum, F. Jackson, R. Kaye, Irene Manton, S. Walker. General Secretary A.R. Busby, 42 Lewisham Road, Smethwick, Warley. 3 West Midlands, B66 2BS. Treasurer Dr. B.A. Thomas, Biological Sciences Department, University of London Goldsmiths’ College, New Cross, London, SE14 6NW. Membership Secretary Lt. Col. P.G. Coke, Robin Hill, Stinchcombe, Dae Gloucestershire, GL117 6BG. Meetings Secretary A.J. Worland, 102 Queens Close, Harston, Cambridgeoniem A> % Ae 4 1 3 . a ae of ie CB2 5QN. Publications Secretary Devon Masarati, 34 Archdale Road, London, SE22 SHS yi Editors: | . oe es % Fern Gazette Dr. C.N. Page, Royal Botanic Garden, Edinburgh, EH3 5LR. assisted by J.A. Crabbe, A.C. Jermy. — Bulletin M.H. Rickard, The Old Rectory, Leinthall Sparkes, nr. Ludlow, Shropshire. — assisted by J.W. Dyce. — ee British Fern Distribution Recorder A.R. Busby, Dept. of Biological Sciences, ee University of Aston, Gosta Green, Birmingham, B4 7ET. Spore Exchange Organiser R.F. Cartwright, 73 Perry Mill Road, Peontcieas . | Pershore. Worcestershire. Archivist N.A. Hall, 3 Sydney Road, Bramhall, Stockport, Cheshire, SK7 1H. Committee J.A. Crabbe, J.K. Cramp, Barbara S. Croxall, P. Edwardee Mary Gibby, J.W. Grimes, N.A. Hall, Gwladys Tonge, ay 2 L. Williams, J. Woodhams. 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Articles for the Society's: She publications are received from and reflect the interests of, Society members. ; Membership is open to all interested in ferns and fern-allies, worldwide. Subscription rates (dua Be RL on 1st January each year) are: Full Personal Members £7; Personal Members not receiving the ~ Fern Gazette £5; Student Members (full-time students) £5; Subscribing Institutions £8. Applications for membership should be sent to the Membership Secretary (address above), from whom further details can be obtained. (Remittances made in currencies other than Sterling are = i £1.00 extra, to cover bank conversion charges.) Back numbers of the Gazette and Bulletin are available for purchase from J.W. Dyce (address above), from whom further details can be obtained. ~, ¥ ae FERN GAZ. 12(3) 1981 121 DROUGHT TOLERANCE IN CHEILANTHES WITH SPECIAL REFERENCE TO THE GAMETOPHYTE HELEN QUIRK & T.C. CHAMBERS Botany School, University of Melbourne, Parkville, Victoria, Australia. ABSTRACT The gametophytes of a range of Australian species of Che/lanthes are shown, in comparison with other more mesic genera of ferns, to have the remarkable capacity to withstand severe water deficits and survive after drying and continued exposure to periods of low humidity. This ability to withstand severe desiccation is considered to be important in the establishment and survival of the genus in ecological niches not otherwise colonized by ferns. INTRODUCTION Only a few genera of ferns are noted for their drought tolerance and their ability to survive in habitats usually not colonized by pteridophytes. Some genera are able to Survive over extensive areas of open country otherwise hostile to most species of ferns. The almost cosmopolitan Pteridium is a good example of such a species as is Paesia scaberula in New Zealand. Such plants have become serious weeds of pasture country. To what extent the invasion by these species is associated with the biological adaptation of the gametophyte as against the adaptations of the rhizome of the sporophyte is not fully understood. Only a few of the genera of the lower vascular plants are able to extend into much more arid habitats than are these two examples; Pellaea and Cheilanthes are notable examples from the Leptosporangiate ferns; the resurrection plants Se/aginella lepidophylla and S. rupestris are well known examples from the Lycopsida. In Australia we have observed that the sporophyte of some species of Cheilanthes perennates during the summer months by means of a rhizome while the fronds completely die down e.g. C. tenu/folia and C. sieberi. Other species which extend even further into the arid parts of Australia maintain at least a few of their leaves in an extremely dehydrated but living state e.g. C. ve//lea and C. /asiophylla. There are a few species of Che//anthes which grow in the cooler coastal regions of Australia. But even in these regions they are never found in the moist, sheltered, constantly humid environments of most other ferns but rather they prefer rock crevices and shallow soil pockets in dry rocky hillsides e.g. C. distans. These species in these habitats sometimes perennate in the hot summer months by means of their rhizome, but under less severe conditions they may remain with some green fronds throughout the growing season. Hevly (1963) has described several adaptations of cheilanthoid fern genera of Arizona to their harsh desert environments. Many of these features are certainly also characteristic of the Australian species of Che//anthes in their sporophytic state. The sporophytic stages of Australian species exhibit microphylly and some have a dense indument of hairs and/or scales; the plants are small (generally below 30 cm in height) and either have narrow fronds (C. s/eberi, C. lasiophylla) or if with ovate fronds then these are finely dissected (C. tenu/folia) or they may have very narrow pinnules (C. caudata). Hevly suggests that the dissection is a reduction in leaf size which may reduce water loss by reduction of total leaf surface. The rhizomes of all species of Che//anthes in Australia are densely scaly and a cover of hairs and/or scales is found on the stipe and lamina of C. /asiophyl/la, C. distans, C. vellea, C. hirsuta, C. contigua and an as yet undescribed sub-species of C. s/eberi. That this thick indument is an inbuilt genetic feature of the plant is demonstrated in the observation that species of C. /Jasiophy//a and C. distans taken from the field and 122 _ FERN GAZETTE: VOLUME 12 PART 3 (1981) grown under glasshouse conditions protected from harsh light and severe water stress, show no lessening in the thickness of the scale cover on the new fronds which developed, even after more than a year in cultivation. Also, sporophytic plants of some of these species raised from gametophytes in cultivation showed the same thick scale covering as did their wild parents. It seems likely that these scales and hairs may reduce water loss through transpiration by creating still air zones near to the plant surface in which the air remains at a relatively high humidity and through which diffusion from the interior of the plant to the exterior would be slow. It is also possible that the scales and hairs reflect significant amounts of solar radiation especially in species growing exposed on desert rocks, e.g. C. ve/lea and C. lasiophylla. Certainly those species which genetically have the most dense indument (C. /asiophylla, C. vellea) are found in the more arid regions of Australia. However, the more common form of C. s/eber/ is unusual in having a glabrous stipe and lamina and yet it is often found growing in the same arid localities as C. /asiophylla. Leaves of the Mediterranean cheilanthoid fern Notholaena marantae R.Br. were reported by Iljin (1931) to tolerate desiccation to relative humidities of less than 50%. This tolerance was attributed to the low leaf surface-to-volume ratio which Iljin suggested reduced the change in volume of the cells on de- and rehydration. In addition he reported the presence of a non-drying substance inthe cells, a stable body or ‘fester Korper’’ which could be differentially stained from the cytoplasm. The fronds of the Australian C. sieberi, C. tenu/folia, and C. “‘austrotenuifolia’™* wilt and die under hot conditions and low humidity. On rehydration, after a sustained dry period, completely new fronds are produced from the rhizome. This seasonal drying off may be at least partly a response to high summer temperatures and longer day length rather than summer water deficit. Evidence supporting this view is obtained from glasshouse-grown specimens of this species which die off each summer despite regular watering and sprout new fronds from the rhizome in autumn. On the other hand, C. /asiophyl/la in cultivation tends to die off to a lesser extent when glasshouse grown and it also does not normally produce new shoots during the hot months even if watering is maintained. We have obtained the best growth responses of these plants in cultivation by ensuring that they are subject to some water stress during the hot summer months. _ It can be seen therefore, that there are a number of adaptations of the sporophyte of Australian species of Che//anthes which include both stress avoidance (by reduction of moisture loss) and high temperature tolerance as seen by the ability to regenerate new fronds after what appears superficially to be death of the old plant by drought. However, these adaptations for many habitats in which these species occur would be in vain if the spores and the gametophyte generation were not also able to avoid or endure periodic xeric conditions. Such adaptations of the gametophyte could range from simple drought avoidance by rapid growth and maturation leading to early establishment of the sporophyte to either, morphologically distinctive xeromorphic features reducing water loss from the gametophyte or to truly physiological xerophytic adaptations of a cytoplasm which can withstand periodic desiccation. Spores of Che//anthes appear to be tolerant of extreme and prolonged exposure to low humidity. Working with the North American species C. graci/lima, D.C. Eaton, Pickett (1923) reported that the spores remained viable after a period of 16 months over anhydrous calcium chloride even though the cytoplasm by this time appeared “dried and shrunken to the sides of the spore’. Although we have not made a *The name ‘‘austrotenuifolia’’ has no status. It is used to designate populations of a species morphologically very similar to C. tenu/folia and up to the present referred to that species but with most distinctive spores. It is widely distributed through the southern half of the Australian continent and we are presently investigating its correct name. QUIRK & CHAMBERS: DROUGHT TOLERANCE IN CHEILANTHES 123 systematic study of the dessication tolerance of the spores of the Australian species we have found it possible to germinate spores of herbarium specimens of C. tenu/folia, 15 months after collection and spores of herbarium specimens of C. shirleyana, C. pumilio and C. caudata germinated and produced typical gametophytes two years after collection from the field. It is clear, therefore, that spores can remain viable for long periods after release. The gametophytes of those Australian species of Chei/lanthes which we have grown lack any obvious morphological protective structures; they are one cell thick for most of their area (except the central cushion region); they have no protective hairs and lack a conspicuous cuticle. They also lack the wax secretory glands reported for Notholaena trichomanordes (L.) Desv. (see Bower 1923) and for NV. stand/eyi Maxon (see Tryon 1947). It seems likely that the survival of the germinated spore and the gametophyte may also be assisted by two features, — first, a fairly rapid germination rate following hydration (usually within 10 days from when spores are sown in the laboratory on moist peat), and secondly by attaining sexual maturity within two months and this may be sooner under field conditions. While such development would assist the plant in avoiding drought, comparable speeds of germination and attainment of maturity have been observed in genera normally associated with more mesic conditions e.g. Phlebodium, Lastreopsis and Doodia. Another factor important in some species of Che//anthes is the presence of apogamy, which enables the fern to circumvent the need for free water droplets at the time of sexual maturity by avoiding the need for fertilization. Knobloch (1966) reported the occurrence of obligate apogamy in 19 species of Cheilanthes including C. tenuifolia and C. sieberi. We have observed apogamy to occur in at least the following 6 Australian taxa, — C. s/eberi, C. sieberi sub-species “pseudovellea”, C. distans, C. lasiophylla and C. pumilio. It seems likely, therefore, that the delicate gametophyte itself might be physiologically able to endure severe water stress and so establish itself in the varied arid niches that are characteristically occupied by members of this genus. The only study we are aware of was by Pickett (1923) who studied the drought tolerance of the Central American C. gracil/lima by drying gametophytes for 5 months in the laboratory followed by 13 weeks over anhydrous calcium chloride. Despite the severe water loss by such treatment he reported that a large percentage of prothalli revived when rehydrated and those which did not survive outright sometimes generated new prothalli on their margins. In view of the apparent complete absence of xeromorphic features associated with the gametophyte we were concerned to answer the following questions in relation to possible xerophytic adaptation:— (1) Are the gametophytes of selected Australian species of Chei/anthes drought tolerant? (2) Ifso, is drought tolerance in any way related to the stage of development at which they are dried? (3) How tolerant are gametophytes to drying at particular levels of relative humidity? (4) Are some spescies more tolerant than others? (5) Does drought tolerance vary with the climate of the area from which the specimen was collected? (6) Can prothalli of more mesic ferns tolerate drought in a similar way? 124 FERN GAZETTE: VOLUME 12 PART 3 (1981) MATERIALS AND METHODS Four species of Chei/anthes from five localities were used. These plants were in cultivation in the glasshouse and from these cultivated plants the spores were collected. The following voucher specimens have been lodged at MEL. (a) C. sieberi, Springhurst, N.E. Vict. E. Canning 4014E (b) C. sieberi, Big Desert, N.W. Vict. H. Quirk 111 (c) C. “austrotenuifolia”, Springhurst, N.E. Vict. E. Canning 4014E (d) C. distans, Dargo, East Vict. H. Quirk 1 (e) C. distans, Queensland, D. Gaff, s.n. (1972) (f) C. lasiophylla, Mootwingee, National Park, Far West N.S.W.1.G. Stone, s.n. (28th July, 1976) H. Quirk 108 (g) C. lasiophylla, Big Desert, N.W. Victoria (MEL 515004) In addition spores of the following more mesic species were similarly sown, Doodia media, R.Br., Lastreopsis shepherdii (Kunze ex Mett.) M.D. Tindale (from Victoria) and Phlebodium aureum J.Sm. (a S. American species in cultivation). Fresh pinnules bearing mature sporangia were surface sterilized by briefly soaking in calcium hypochlorite solution, followed by washing in running water. Spores were shed between clean sheets of white paper and were sown onto moist peat in small crystallizing jars sealed with a thin film of polyethylene plastic (““Gladwrap’’) to reduce the risk of contamination and prevent the cultures from drying. Cultures were grown indoors near a north facing window protected from high summer sun but allowed brief periods of direct sunlight especially in the Autumn months in the late afternoon. The humidity treatments were conducted in closed vessels using a range of concentrations of sulphuric acid following Wilson (1921) and making the assumption that the cultures would be kept in the temperature range of 25°C (in fact the temperature in this room fluctuated between 17°C and 28°C.) %H,SO, 70.5 58.5 50.9 43.4 36.0 DN RH at 25°C 5% 20% 35% 50% 65% 80% The concentration of acid was only known within the range 95-98% which could lead to minor errors. The dilutions were made assuming the acid concentrations to be 96.5%. The above range of humidities were selected to cover and extend below those likely to be encountered by field grown material. In arid Central Australia, humidity may drop to 15% or lower in the drier months whereas in Melbourne the lowest humidities in the summer are about 60%. In the field, humidities could vary greatly through each 24 hour period. No attempt was made to simulate such changing conditions. The vessels used for the humidity changes consisted of small screwtop glass jars 8 cm high x 5cmdiameter containing 33 mls of acid solution. Aluminium foil trays were suspended on wire attached to the lid, each tray being about 3.5 cm in diameter and divided into 4 sections so that prothalli from 4 different samples could be treated in each jar simultaneously. The gametophytes which were to be subjected to the humidity treatments were first allowed to dry slowly by taking 6 groups of prothalli (15-40 individuals) from a culture to be tested. Care was taken to see that the peat substrate remained attached to the prothalli so that damage to the rhizoides was minimal. These groups of species were placed in separate dishes of moist peat and exposed in a glasshouse for 7-10 days which allowed them to dry out gradually and reasonably naturally (the glasshouse was neither heated nor cooled but it was well ventilated). Each of the 6 groups now wilted and dried were then placed in one of the 6 humidity changes at 6 different relative humidities, sealed and wrapped in plastic and allowed to stand in the same room where they had been growing originally and where temperatures varied between 17° and 28°C. After 10 days of such treatment the prothalli were removed and, still attached to their original substrate (peat moss) placed in petri dishes and rehydrated with a few drops of water and allowed to stand overnight. the prothalli were then replanted into moist peat under the same conditons as those in which they had originally been grown, and periodically checked to determine whether they had survived the desiccation treatment and had retained the capacity to continue to grow and develop normally. From each experimental batch a few plants were tested to check whether they were alive by placing them for 4-5 hours in a 0.1% stock solution of neutral red made up 1 : 30 with O.1M CaCl, with a trace of dilute NaOH. Cells which were still alive took up the orange coloured solution while those which were permanently damaged did not. QUIRK & CHAMBERS: DROUGHT TOLERANCE IN CHEILANTHES 125 RESULTS Prothalli of the 7 samples of Che//anthes studied were dried at 4 different stages in their development:— Stage 1 3-4 days after germination, when prothalli are only a small filament or plate. Stage 2 2-3 weeks later when prothalli are cordate in shape but not yet fully grown. Stage 3 prothalli are fully mature with either antheridia or archegonia (in the case of C. “austrotenuifolia’’) or just beginning to give rise to apogamously produced sporophytes. Stage 4 Young sporophytes have developed 1-3 leaves. At this stage they are still very small (ca. 5mm high) and unprotected. Our first attempts to study the drought resistance of the gametophyte failed because we dried the gametophytes at their various stages of development immediately from the fresh state over the selected acid in the humidity controlled jars. This drying proved too severe and too rapid and very few gametophytes at any stage from any species survived after such treatment. All the experimental results which follow are based on gametophytes which were dried slowly by the method described. (a) C. sieberi (Springhurst) Stage % survival at various relative humidities 5% RH 20% RH 35% RH 50% RH 65% RH 80% RH 1 * * * * 100 * 2 6) @) 10 0 0 10 3 100 100 100 80 100 50 4 100 100 100 80 100 50 ae indicates points at which, due to loss from factors other than water stress (e.g. funal invasion) data is not available. (b) C. sieberi (Big Desert) Stage % survival at various relative humidities 5% RH 20% RH 35% RH 50% RH 65% RH 80% RH 1 100 100 100 100 100 100 2 100 100 100 100 100 100 3 at * * * * * 4 100 100 100 100 100 100 On replanting of stage 2 and 4 prothalli, all developed healthy sporophy tes. (c) C. “austrotenuifolia’ (Springhurst) Stage % survival at various relative humidities 5% RH 20% RH 35% RH 50% RH 65% RH 80% RH 1 a = ‘4 = 100 i 2 0 50 100 100 50 80 3 100 100 100 100 100 80 Replanted stage 2 prothalli were still alive 2 months later and had produced one sporophyte. 126 FERN GAZETTE VOLUME 12 PART 3 (1981) (d) C. distans (Dargo) Stage % survival at various humidities 5% RH 20% RH 35% RH 50% RH 65% RH 80% RH 1 0 30 100 20 100 0 2 80 100 80 25 10 5 3 0 0 10 0 0 0 4 100 100 100 100 100 100 Of the stage 2 gametophytes replanted, those dried at 5%, 20% and 50% RH produced sporophytes. Of the stage 3 gametophytes replanted, those dried at 5% and 50% RH produced sporophytes. Of the Of the stage 3 gametophytes replanted, those dried at 5% and 50% RH produced sporophytes. Of the stage 4 gamtophytes replanted, nearly all continued to develop more leaves on the sporophyte. (e) C. distans (Qld.) Stage % survival at various relative humidities 5% RH 20% RH - 35% RH 50% RH 65% RH 80% RH 1 100 100 100 0 100 60 2 80 100 100 100 100 12 3 0 0 100 20 0 6) 4 100 100 100 50 12 100 Of the replanted stage 2 and 3 prothalli, nearly all continued to grow and produce new sporophy tes. (f) C. lasiophylla (Mootwingee) Stage % survival at various relative humidities 5% RH 20% RH 35% RH 50% RH 65% RH 80% RH 1 100 i 50 Es % ee 2 5 100 100 100 100 100 3 80 75 100 100 100 20 4 100 100 100 100 100 80 Of the stage 2 and 3 prothalli replanted, all were still healthy 2 months later, but had not produced sporophytes. In contrast, those of the original culture had all produced 3-5 leafed sporophytes by this time. (g) C. lasiophylla (Big Desert) Stage % survival at various relative humidities 5% RH 20% RH 35% RH 50% RH 65% RH 80% RH 1 100 40 0 0 _ = z FAS) 100 14 0 50 100 3 100 100 100 — 100 6) Replanted stage 2, 3 and 4 prothalli continued to grow and produce healthy sporophytes. -- ec A QUIRK & CHAMBERS: DROUGHT TOLERANCE IN CHEILANTHES 127 (h) Doodia media (Stage 3 only) No prothalli survived at any of the humidities at which they were dried. (i) Lastreopsis (Stage 3 only) No prothalli survived at any of the humidities at which they were dried. (j) Phlebodium aureum (Only a few at stage 3 survived and these produced antheridia but not archegonia). Stage % survival at various relative humidities 5% RH 20% RH 35% RH 50% RH 65% RH 80% RH 3 6) 0 7 0 0 20 It is possible that the prothalli which did not survive after a period of desiccation were accidentally dried in the glasshouse at a faster rate than the others. It is also possible that some other unknown damage was caused to them by the nature of the experiment. It is equally possible that there is some genetically controlled variation in drought resistance of the gametophytes although we are not able to detect any definite pattern that would support such a view at this stage. The data here answer some of the questions that were asked. First it suggests that the stage of maturity of the gametophyte has very little if any effect on drought tolerance, several species showing 100% survival rates at both very early and late stages of development. Certainly the results with the Springhurst population of C. sieberi Suggest that it may be better adapted to survive such drought once it has reached the early sporophytic stage. In contrast the Big desert populations of the same species Survived at all stages of development. The absence of data for stage 3 was due to an experimental accident, — the invasion of fung). It might have reasonably been expected that he less severe the lowering of humidity the less damaging it would be to cells, but the data does not really support such a view. No pattern suggesting greater survival at higher humidities was observed. Death appears, to have occurred irregularly at all humidities in fact, there does appear to be a slightly greater likelihood for prothalli which were dried and treated at relative humidity 80% not to survive. This of course, may be a reflection of continued bacterial and fungal action, which presumably could in some instances continue at this relatively high humidity. For example, stage 2 prothalli of C. distans (both the Dargo and Queensland populations) show much lower survival rates at high humidities. Similarly in stage 3 prothalli of C. “austrotenu/folia” deaths occurred only at relative humidity of 80%. It is again possible that the higher humidities here allowed continued bacterial action. We do not have sufficient data to determine whether some species of Cheilanthes are more drought tolerant than others. Certainly, of the two Big Desert species studied, C. sieber/ had a greater survival rate than did C. /asiophy/la. However, the irregularity of the death pattern of C. /asiophy//a (100% of stage 2 prothalli survived 20% relative humidity, while 75% and 14% survived drying to relative humidities of 5% and 35% respectively), suggested that death may be due to some unknown factor. It appears that gametophytes of the more arid zone species from the Big Desert and Mootwingee showed an overall greater survival rate than did those of C. “austrotenuifolia” and C. distans (Dargo) although some other specimens of C. “austrotenuifolia” and C. distans did exhibit some 100% survival of the very young stage 1 (prothalli) and of prothalli dried at 5% relative humidity. 128 FERN GAZETTE. VOLUME 12 PART 3 (1981) Comparison of the data for C. s/eberi cultures from the Big Desert plants, with those from specimens collected at Springhurst suggests that the prothalli from the Big Desert (the more arid areas with 210mm precipitation per annum) may be more drought tolerant than those from near Springhurst (annual precipitation 560mm). If this interpretation is correct, then the Big Desert plants are genetically better adapted to endure server periods of water stress than those of the more humid Springhurst district. However the Springhurst data is not easy to interpret in view of the 100% survival of stage 1 prothalli at 65% RH. So far as the prothalli fo the more mesic fern species are concerned it is conclusively shown that the species of Doodia and Lastreopsis investigated are completely without tolerance to the levels of desiccation in which Cheilanthes can survive. The few prothalli of the epiphytic Ph/ebodium aureum which survived drying at 35% and 80% RH suggests that the species has some degree of adaptation to temporary water deficits. DISCUSSION The almost zero survival of prothalli subjected to sudden drying (1-5 hours) by direct introduction into the controlled humidity chambers suggests that changes which need to take place in a living cell as the cytoplasm dehydrates require a reasonable time. Given slow drying conditions (5-7 days) the physiological changes to the cytoplasm which may take place allow the cells to become adapted to a sustained period of severe moisture stress. Somewhat comparable experiments on the moss Tortula muralis by Gwozdz and Bewley (1975) showed that gradual drying apparently protects the cell from the injury caused by rapid drying; further they reported the fine structural changes which suggested that the slowly dried moss plant retained fewer by more active polyribosomes than did the rapidly dried plant and that the protein synthesis activity of the polyribosomes increased more rapidly on rehydration in these slowly dried plants. When gametophytes were treated with neutral red dye to determine which cells remained alive, we found that those parts most crucial to the survival of the plant, — the rhizoids, meristem, reproductive organs and the very young sporophyte had a greater Capactiy to Survive rapid severe drying than did the remaining vegetative cells. The experiments reported in detail in this paper all involved slow drying of the gametophyte prior to treatment and the data undoubtedly demonstrates that the species of Che/lanthes studied have the ability to tolerate and survive drying to humidities well below those which would normally be encountered in the natural environment. Under these conditions of drying the gametophytes retain their chlorophyll pigment but otherwise appear shrivelled and severely damaged. At the lower humidities they become brittle to the touch. On rehydration those which Survived appeared healthy again and indistinguishable from those which had never been dried. Most of the prothalli which survived the drying went on developing to produce healthy sporophytes, growth being merely interrupted during the period of drought. It may be significant that the dehydrated prothalli although shrivelled and brittle remained green. It has been reported for other plant groups (Gaff & Hallam 1974) that those which retain their chlorophyll during desiccation (as does Che//anthes) suffer less change of structure than those plants which do not retain their green pigments. Comparative fine structural analysis of the cytoplasm of the gametophytes of Cheilanthes with drought intolerant species during both dehydration and rehydration could possible explain some of the events taking place during water stress. QUIRK & CHAMBERS: DROUGHT TOLERANCE IN CHEILANTHES 129 ACKNOWLEDGEMENTS We are grateful for the advice of Dr. D. Gaff of the Botany Department of Monash University who pointed out the requirement of slow drying in experimental studies of drought tolerance. One of us (HQ) acknowledges the support for this study of a Melbourne University Post-Graduate Award. REFERENCES BOWER, F.O., 1923. The Ferns Vol 1. Cambridge Univ. Press, Cambridge. GAFF, D.F and HALLAM N_.D., 1974. Resurrecting desiccated plants. R.Soc. N.Z. Bull. 12: 389- 393. GWOZDZ, E.A. and BEWLEY, J.D., 1975. Plant desiccation and protein synthesis. Plant Physiol. 55: 340-345. HEVLY, R.H., 1963. Adaptations of cheilanthoid ferns to desert environments. J. Arizona Acad. Sci. 2: 164-175. ILJIN, W.S., 1931. Austrocknungsresistenz des Farnes Notoch/aena marantae R.Br. Protoplasma 13: 322-330. KNOBLOCH, I.W., 1966. A preliminary review of spore number and apogamy within the genus Cheilanthes. Am. Fern J. 56: 163-167. PICKETT, F.L., 1923. An ecological study of Che//anthes gracillima. Bull. Torrey. Bot. Cl. 50: 329- 338. TRYON, A., 1947. Glandular prothalli of Notholaena standleyi. Am. Fern J. 37: 88-89. WILSON, R.E., 1921. Humidity control by means of sulfuric acid solutions, with critical compilation of vapour pressure data. J. /ndus. and Engin. Chem. 13: 326-333. 130 FERN GAZETTE: VOLUME 12 PART 3 (1981) REVIEWS SELECTED PTERIDOLOGICAL PAPERS 1952-1978 OF PROFESSOR S. KURATA. Edited by T. Nakaike. Supplement No. 7 to the Journal of the Nippon Fernist Club Volume 2 1979. 190 x 265 mm. No price stated. Sataru Kurata was born in 1922. He died in the latter part of 1978 having published over 65 papers on the taxonomy, ecology, distribution and nomenclature of Japanese ferns particularly on what has been generally called Aspideaceae although one substantial paper is on Lep/sorus in Japan. For much of this time Kurata was Professor in the Institute of Forest Botany, Faculty of Agriculture, University of Tokyo. The papers appeared in the Hokuriku J. Bot. (latterly the J. Geobot.): Notes on Japanese Ferns (1) — (50) (1952-1970). Other major papers are from Sc. Rep. Yokosuka City Mus. andthe Sci. Rep Téhoku Univ. Ser. !V (Biol). Other major papers are from Sci. Rep. Yokosuka City Mus. All are facimile reproductions excluding pagination but the collected work is additionally paginated to accommodate the index to Japanese and Latin names. There are substantial keys, in English, for Cyrtomium, Lepisorus and the Polystichum polyblepharum group. Two other keys, to Arachniodes and the Athyrium otophorum group are in Japanese. Perhaps Dr Nakarke, who saw this through the press will one day translate these so they may have greater use. A.C. JERMY FERNS OF FLORIDA - AN ILLUSTRATED MANUAL AND IDENTIFICATION GUIDE by Olga Lakela and Robert W. Long ix + 78pp. Banyan Books, Miami, Florida, 1976. ISBN 0-91 6224-08-2. Price $13. This book is intended as a field guide with identification keys and descriptions of the 135 species of native and naturalised ferns. 115 are illustrated, nearly all as photographs of herbarium specimens. The photographs are generally good although some additional detailed drawings would be very acceptable. There is a short introduction attempting to summarise the local ecology of ferns, pteridophyte classification, taxonomic characters of ferns, polyploidy in ferns, and even growing ferns from spores. There is also a glossary of nearly two hundred terms. Apart from people contemplating a visit to Florida, it is essentially a flora for those who are interested in sub-tropical and tropical ferns. Although a few species have wide distributions in North America, the vast majority are restricted to the south- eastern States or even Florida while also being common in the Caribbean and tropical Central and South America. These readers would appreciate having more notes on the distribution and ecology of the Florida species together with additional information on the local climate and geology. B.A. THOMAS FERN GAZ. 12(3) 1981 131 TROPISMS IN REGNELLIDIUM DIPHYLLUM LUIS D. GOMEZ Herbario Nacional, Museo Nacional de Costa Rica, C.A. ABSTRACT Diaphoto- and paraheliotropisms are reported from plants of Regnellidium diphyllum under cultivation. The possible significance of these physiological reactions on the ecology and distribution of this fern is briefly discussed. OBSERVATIONS The movement of plants or their parts attracted the attention of Darwin (1880) and many others after him. Nevertheless, with the exception of occasional mentions, the movements of ferns has remained a rather void aspect of pteridophyte physiology, the last detailed account of them is that of Du Buy & Nuernbergk (1938). The expansion and contraction of the leaflets of Marsi/ea has been recently observed by Piggott (1979) who also mentions the heliotropic movement of its fronds. Another genus in the Marsileaceae, Regnellidium, shows a _ remarkable diaphototropism or light-tracking ability. Plants cultivated in tanks at 20°C and placed near a light source (75 Watt at 3 feet) will turn all the fronds towards the light in 90-120 minutes, the curvature taking place near midlength of the petioles. The same plant, turned around 180° will initiate reorientation towards the light in 15-30 minutes until all leaf surfaces are perpendicular to the light source. If a plant is exposed to light sources of equal intensity, one at each side of the tank, some fronds will face in one way while the rest face the other, except for the young unexpanded ones which will remain more or less straight and oblique to the path of light. The shoot apex of Regnellidium is also light sensitive. Developing rhizomes are directed to the light source but their reaction to changes of position of the light is not as rapid as that of the leaves. The movement of the apical portion of the rhizome and of the leaves is interindependent: if a portiom of the rhizome is placed in such a manner that some of its fronds are directed to one light source, and the rest to another, the shoot apex is still able to track a third source of light, even if it is of lower intensity. In Marsilea, a decreased illumination causes a closing-up of its leaflets (cupping or paraheliotropism). In Regnel/lidium this reaction is almost insignificant. Submerged plants do not exhibit cupping during the night or in artificial darkness but aerial leaves show very reduced paraheliotropism. This type of movement is only observed in fully expanded leaflets, never in the immature ones. The maximum width of the leaves is attained approximately 14-20 hours after the maximum petiole elongation is achieved, until then the leaflets remain appressed upwards, total divergence of the leaflets takes from 180-270 minutes. ECOLOGICAL INTERPRETATION The rare Regnellidium diphyl/um Lindm. is restricted to few localities in southern Brazil with one relictual population in Argentina (Schultz, 1949; de la Sota & Mitchell, 1970). It usually grows in shallow ponds of slightly acid (pH 6.0) water in association with grasses, aquatic phanerogams and Sal/vinia. In crowded conditions, a not uncommon situation in marginal lotic environments, effective sun-tracking mechanisms clearly allow for a maximum illumination since planar leaves would be continuously exposed to the maximum levels of incident energy. This, coupled to the fast petiolar growth (average rate is O.73mm/hour of light in our plants) increases the 132 FERN GAZETTE VOLUME 12 PART 3 (1981) potentiality of the plant to compete with other natant and emergent water plants. The cupping of the leaves in Marsilea and Regnellidium suggest a mechanism for the reduction of energy absorbtion via a decrease inthe angle of incidence of the source of radiation. In their natural environment the Marsileales are subjected to intermittent wet and dry periods. The stress resulting from such intervals of drought is counteracted through paraheliotropism, by a reduction of the energy levels, thus becoming a very significant adaption for the survival of the plants. This is particularly the case in the species of Marsilea, often found in xeric and sub-xeric habitats and whose sporocarps are quite adapted to a thorough drying and exposure to heat before dehiscense. The weak paraheliotropic reaction observed in Regnel/lidium suggests this plant requires longer periods of pronounced moisture and could, in all probability, account for its sparse highiy localized distribution (Gaudet, 1963). Whether the phototropisms of Regnellidium and other hydropterides are dependent on a phytochrome-mediated K-ion transfer as in other ‘‘sensitive’’ plants or not, remains to be studied as well as the possible role of the latex of this plant, and offers a fascinating field for exploration in the much-needed branch of fern sporophyte eco-physiology. ACKNOWLEDGEMENTS This research was partially financed by CONICIT, San José, Costa Rica. REFERENCES DARWIN, C., 1880. The power of movements in plants. 1st Ed. John Murray, London. DU BUY, H.G., NUERNBERGK, E.L., 1938. Growth, tropisms and other movements. In Fr. Verdoorn (Ed.) Manual of Pteridology. M. Nijhoff, The Hague. pp. 303-346. GAUDET, J.J., 1963. Marsilea vestita: Conversion of the water form to the land form by darkness and by far-red light. Science 140: 975-976. PIGGOTT, A.G., 1979. Observations on light response in Marsilea crenata. Fern Gaz. 62-64. SCHULTZ, A.R., 1949. Contribucoes ao cohecimiento de Regnellidium diphyl/um Lindm. Lilloa 17: 139-144. SOTA, E. de la, MITCHELL, D.S., 1970. Sobre la presencia de Regnellidium diphyllum Lindm. (Marsileaceae-Pteridophyta) en Argentina. Darwiniana 16(1-2): 408-409. FERN GAZ. 12(3) 1981 133 AN ECOLOGICIAL SURVEY OF THE FERNS OF BERWICKSHIRE, SCOTLAND A. WILLMOT Derby Lonsdale College of Higher Education, Derby ABSTRACT A systematic list of the ferns and fern allies of Berwickshire (v.-c. 81), south-eastern Scotland, is given, based mainly on records made in 1979 by the author. This includes notes on the distribution, abundance and habitats of the taxa. The list includes 32 species and 3 hybrids as recently recorded. The records for Equisetum x litorale and Dryopteris x deweveri are first county records. The status of species for which only pre-1950 records exist is discussed and two species accepted as native (Osmunda regalis and Asp/lenium ceterach) though only as casuals. The distribution of the commoner 18 species is shown by maps. Species lists are given for moorland, walls and deciduous woodland. Three species are suggested as primary woodland indicators in the area (Equisetum sylvaticum, Gymnocarpium dryopteris and Polystichum setiferum). The phytogeographical affinity of the flora on a European basis is described by dividing it amongst the floristic elements of Birks (1976). The majority of species belong to his widespread and Mediterranean/Atlantic elements and three species to upland or northern elements. The geographical distribution of species within the area is described and three major patterns of distribution are recognised: widespread; coastal plateau and northern-central; and eastern or coastal. INTRODUCTION Berwickshire is the south-easternmost vice-county (v.-c. 81) in Scotland, situated on the eastern end of the Southern Uplands, which are smooth rounded hills mostly between 450 and 600m in height (Whittow, 1979). However, the county is atypical of the Southern Uplands in that there are extensive areas of lowlands Lowland Areas NORTH SEA 7 Pott re! Carboniferous Uplands x Hee St. Abbs VZA O\d Red Sandstone a tH Ss Saeae FA Coastal Plateau 7 ISSSSSSeRSeeeeeee’, 6 {| SESSS828°4) | anneal ee aee8 Pad Lammermuir Hills Po 0 no Neo o°F} °° Kilometres FIGURE 1. Map of Berwickshire (v.-c. 81) showing the five topographical regions and the 10 km grid squares of the National Grid of the Ordnance Survey. 134 FERN GAZETTE: VOLUME 12 PART 3 (1981) (Sissons, 1976). The county is bounded to the north by the Lammermutr Hills, to the east by the North Sea, to the south by the River Tweed and to the west by the Moorfoot Hills. It is one of the smaller vice-counties, extending at most only 40km north/south and 60km east/west. Botanically it has no particular claim to fame, having no concentrations of rarities or nationally rare habitats, though it has a wide range of habitats and species to match. Sissons (1976) divides the area into five topographical units (Fig. 1). In the north is the highest ground (400-500m), which is part of the Lammermuir Hills. These produce typical Southern Upland scenery of rounded hills and smooth slopes with few outcrops of rocks. The solid geology (Greig, 1971) is mostly Silurian, sedimentary rocks which weather to produce poor, acid soils. To the south is the Tweed Basin which consists of three morphological units related to the geology of the region. The lowest ground, which is mostly below 100m, is in the south east. This is known as the Merse and is underlain by Carboniferous sedimentary rocks, concealed by a thick layer of strongly drumlinised drift, which produces high quality farmland. The Merse is separated from the Lammermuir Hills by low uplands of Old Red Sandstone interrupted by a series of igneous intrusions that form conspicuous hills. In the extreme south of the county is a small area of Carboniferous larvas that also form low uplands. The fifth topographical unit is a plateau of Ordovician and Silurian rocks (200- 240m), which occurs along most of the short coastline of the county forming steep, grassy sea-cliffs with numerous outcrops. The last three topographical units ali have acidic soils that produce farmland intermediate in quality between the high quality farmlands of the Merse and the low quality ones of the Lammermuir Hills that mostly bear rough upland grazing. The climate of the area is described in Gregory (1976) who places most of the area inhis BD2 category along with Midland England and the eastern coast of northern England and Scotland. This region has a growing season of 7 to 8 months and less than 750mm of rain which falls mostly in the second half of the year. Inland the climate becomes colder in winter and merges into category BM2 on the hills, which have a higher average rainfall. Lacking rare species and unusual habitats the area has been botanically under- recorded. The only complete floras for the county being in the work of Johnston in the first half of the 19th century (Johnston, 1829 & 1853), though these both cover areas larger than the vice-county. New records for the county and species lists for the better known sites then occur fairly commonly in the History of the Berwickshire Naturalists’ Club up until about 1940. After this date botanical activity inthe county seems to have almost ceased, as judged by the number of records published in this and other journals. This cessation is so complete that during the Botanical Society of the British Isles’ field week to the county in July 1979 no one who actually lived in the county attended. This lack of activity is a shame as the county has many interesting sites that are in danger of being destroyed without even a proper record of what they contained. The lack of activity in the study of the modern flora of Berwickshire also extends to the study of the plant remains that must exist in the numerous Quaternary deposits in the area. Newey (1968) in his account of pollen analyses from south-east Scotland does not include a site from the area and the only account of Quaternary plant fossils in the area seems to be the note by Mitchell (1948) on Whitrig Bog. This records spores of Selaginella selaginoides* from late-glacial deposits. Despite the lack of records for the county itself Birks (1977) has reconstructed a preliminary picture of the natural vegetation of the area before man cleared it for agriculture. He places the county in his “Oak forest with birch’ region and considers these woods had U/mus glabra and Fraxinus excelsior on better soils and Cory/us avellana in more open areas. Surviving *Nomenclature follows Clapham et a/ (1962) for angiosperms and Jermy et a/ (1978) for ferns. ee WILLMOTT: THE FERNS OF BERWICKSHIRE 135 fragments of these woods on acid soils have a ground flora of Pteridium aquilinum and Deschampsia flexuosa when heavily grazed, and a richer flora including Dryopteris aemula, D. pseudomas and Blechnum spicant when ungrazed. These woods before they were cleared would have covered all but the highest parts of the Lammermuir Hills with the only other open areas being on cliffs, bogs and flood plains in the lowlands. FIGURE 2. View looking down into Pease Dean from the bridge (36/791.700), the classic locality for Polystichum setiferum in Berwickshire. The best sites for ferns in the area at present are steep, rocky hillsides which, because they are unsuitable for grazing, have retained remnants of the native oak/birch woods described above. These occur mostly in cloughs (narrow, steep-sided valleys) around Duns and Abbey St. Bathans and in deans near the sea, for example Dowlaw and Pease Dean (Fig. 2). Otherwise the distribution of ferns is determined by the layout of the topographical units, which provide markedly different opportunities for the occurrence of ferns. The lowland of the Merse with its intensively-cultivated, large fields, planted copses and shelterbelts provides habitats for only the commoner lowland ferns. Walls in this area, however, provide a habitat for ferns that would otherwise not occur. The uplands of the Lammermuir Hills provides habitats for the more upland and heathland ferns, particularly along the rocky banks of streams that flow through the large areas of undulating moorland. The rather lower Coastal Plateau contains areas of similar moorland alongside areas of more intensive agriculture and forestry. In this region the coastal cliffs with their rocky outcrops and grassy flushes provide suitable sites for some of the more interesting fern species. The low uplands of Old Red Sandstone and of Carboniferous larvas provide a mixture of the habitats that occur in the Merse and the Lammermuirs, and being less intensively-farmed provide more semi-natural areas in the form of marshy fields and boggy areas. This brief summary of the area provides rather too good a picture of the suitability of the area for ferns as changes in the form of farming are occurring that threaten to markedly alter the picture over large areas. These changes are part of the national 136 FERN GAZETTE: VOLUME 12 PART 3 (1981) trend from traditional methods of farming to modern more intensive methods that leave very little room for wildlife in crops and very few odd corners uncultivated. Inthe Merse these changes are virtually complete. In the low uplands and Coastal Plateau they are proceeding rapidly, such that orange coils of plastic drainage-pipe are a common site. In these areas, if the changes continue at the speed they are, in a few years there will be as few areas for wildlife as in the Merse. In the Coastal Plateau the changes appear to be a more abrupt and recent phenomenon with improved farmland directly replacing the moorland recorded by Johnston (1829 & 1853) leaving in many places only thin, roadside strips of moorland. The changeover is virtually complete in the southern section of the Plateau and proceeding fast in the northern section. Only the Lammermuir Hills, which because of their altitude do not provide the opportunities for agriculture that the other parts of the county do, are free from extensive improvement at present. METHODS The records (Table 1) made by the author given in the following sections were all made during July 1979 and refer to the vice county (v.-c. 81) as given in Dandy (1969). They were gathered on the basis that two to three examples of each major habitat type (Table 2) were visited in each 10 km grid square. At each habitat visited a subjective estimate of the abundance of each fern present was recorded on the Dafor scale. If a habitat spread over more than one one-kilometer square (referred to here as a monad), a separate record was made for each. If during the course of visiting one habitat other major habitat types were encountered, separate records were made for these. Habitats were selected either because there were old records for the sites or because they looked promising on maps or on the ground. No attempt was made to record habitats which contained no TABLE 1: FERN TAXA RECORDED FOR BERWICKSHIRE (Taxa recorded in present survey are marked ! and taxa, for which only doubtful or pre-1950 exist, are enclosed in brackets. Numbers in brackets refer to floristic elements in Table 3.) Lycopodium clavatum ! (1) Oreopteris limbosperma |! (3) Huperzia selago (1) Asplenium scolopendrium ! (3) Diphasiastrum alpinum (5) A. adiantum-nigrum | (3) (D. x issleri) (7) A. marinum ! (4) Selaginella selaginoides | (5) A. trichomanes agg. ! (2) (Equisetum hyemale) (1) A. trichomanes subsp. quadrivalens E. fluviatile | (1) A. ruta-muraria ! (2) E. arvense ! (1) (A. ceterach) (3) E. x litorale | Athyrium filix-femina ' (1) E. sylvaticum ! (1) Gymnocarpium dryopteris | (1) E. palustre | (1) Cystopteris fragilis | (1) E. telmateia | (3) (Polystichum lonchitis) (6) Botrychium lunaria (1) P. aculeatum ! (3) Ophioglossum vulgatum (2) P. setiferum | (4) (Osmunda regalis) (3) Dryopteris filix-mas ! (2) Cryptogramma crispa ! (6) D. pseudomas ! (3) Polypodium vulgare ' (2) (D. aemula) (8) P. interjectum ! (3) D. carthusiana ' (1) P. x mantoniae D. austriaca | (3) Pteridium aquilinum | (2) D. x deweveri ! Phegopteris connectilis | (1) Blechnum spicant | (3) WILLMOTT: THE FERNS OF BERWICKSHIRE 137 ferns. In all, records were obtained from 160 monads but due to the non-systematic nature of the survey there are a number of forms of bias in the results. The three major ones are: a bias towards monads near roads; a bias to monads below 250m; anda bias to monads in the eastern half of the county. Where the ecological or geographical distribution of taxa is discussed below, attempts have been made to correct for these forms of bias. TABLE 2: FREQUENCY OF THE COMMONER SPECIES IN THE MAJOR HABITATS (Figures in the body of the table refer to the number of monads in which a species occurred in the particular habitat. Heath includes moorland dominated by ericaceous shrubs and grasses while tall herb includes tall grasslands). MAJOR HABITAT TYPES No. of D. C. Tall Bare Monads with SPECIES Marsh Wood Wood Scrub Hedge Heath Herb Rock Walls Species Equisetum fluviatile 6 1 _ — — - ~ - _ 9 E. arvense 8 6 1 1 5 3 35 - 60 E. palustre 15 _ — — - — 9 = — 24 Polypodium vulgare agg. _ 2 _ a - _ - 18 23 43 Pteridium aquilinum 1 ZT 9 6 a es 21 3 1 63 Oreopteris limbosperma _ 6 1 1 = 11 = -- _ 18 Asplenium scolopendrium - 6 — _ _ _ 1 6 15 24 A. adiantum -nigrum = — — ~ ~ — — 1 7 8 A. trichomanes agg. — — _ — — _ — 4 25 23 A. ruta-muraria - — - — = _ - 1 28 29 Athyrium filix-femina 7 66 a3 7 1 ile: 9 5 11 100 Cystopteris fragilis — _ — — _ = — 1 8 9 Polystichum aculeatum _ 13 = _ 1 — 1 5 1 16 Dryopteris filix-mas 3 74 20 15 16 8 a7 13 45 132 D. pseudomas _ 46 7 q 3 8 a 4 1 76 D. carthusiana 1 6 1 _ — 1 _ _ ~ 7 D. austriaca 2 72. 19 8 3 7 10 9 10 107 Blechnum spicant _ 12 2 1 1 16 1 - _ a | No. of Monads with habitat 20 79 24 16 18 25 58 26 70 D. Wood = Deciduous C. Wood = Coniferous A survey was also made of the main works and data banks which contain fern records for the area; these included Jermy et a/(1978), the History of the Berwickshire Naturalists’ Club 7-40 (1834-1974); Johnston (1829-1853) and the Biological Records Centre, Monk’s Wood. As Jermy et a/ (1978) is based on 10 kmgrid squares and many of these on the borders of the area cross two or more vice-counties, a record from Jermy et a/(1978) was only accepted as from the vice-county if there was a record at Monk’s Wood for the grid squares in the vice-county. 138 FERN GAZETTE: VOLUME 12 PART 3 (1981) RESULTS Results are presented in four ways. The systematic list describes the status, ecology and distribution of all fern taxa ever recorded for the vice-county, while the remaining three ways refer only to records from this survey. Table 2 summarises the frequency of the commoner species in the more frequent habitats Figures 3, 5 and 7 give distribution maps for the commoner species on a tetrad basis; and the habitat lists detail the occurrence of ferns in selected habitats. SYSTEMATIC LIST OF TAXA Each taxon recorded during the survey is given with notes on (a) its geographical distribution, (b) its habitats and (c) its altitudinal range, as recorded in the survey. Where a taxon was recorded in only a few monads, an entry is given in the form (a-c) followed by notes on the individual occurrences. Reference in these notes to abundance in habitats refers only to abundance in examples of the habitat where the species occurred, how often the species occurred in the habitat type can be seen from Table 2. Notes are also given (d) for significant extensions to (a), (b) and (c) from other sources, particularly post-1950 10 km grid square records not reconfirmed here, and selected pre-1950 records that have not been refound recently. Taxa that have not been reconfirmed in this Survey are given only type (d) notes. Grid references prefixed with circa are my interpretation of localities originally given without a reference. Numbering of taxa follows Jermy et a/(1978). Abundance is recorded on the Dafor scale where D = dominant, A = abundant, F = frequent, O = occasional, R = rare and L = locally. 1.2. Lycopodium clavatum (Stag’s-horn clubmoss) (a-c) Recorded only once for Duns Castle reserve but at two localities. In both it occurred as a Sparse patch about one metre in diameter, at 150m. At 36/779.553 it bore cones and grew amongst short acid grass below beech trees (Fagus sy/vatica), while at 36/779.558 it was sterile and grew below heather bushes (Ca//una vulgaris) in heathland. (d) Recorded at BRC post-1950 for the 10 km grid squares 36/44, 45, 53, 54, 55, 64, 65, 74, 76 & 86. 3.1 Huperzia selago (Fir clubmoss) (d) Only post-1950 record is record at BRC for Earlston c. 36/56.38. There are, however, older records for Lamberton Moor c. 36/95.58, Dirrington Law c. 36/69.54 and Penmanshiel Moor c. 36/81.67 in Johnston (1853); and for Buncle Edge near the forts c. 36/80.60 and old roads on Coldingham Moor c. 36/86.67 (Lynn, 1895). The species therefore seems to be much less wide-spread than it was a hundred years ago. 4.1 Diphasiastrum alpinum (Alpine clubmoss) (d) Only post-1950 records are records at BRC for 36/64 & 65. There are, however, older records for Lamberton Moor c. 36/95.58 (Johnston, 1853); moor on ridge betwen Kyle Hill and Dogden Moss c. 36/70.50 (Anon, 1886); and Buncle Edge c. 36/80.60 and old roads on Coldingham Moor c. 36/86.67 (Lynn, 1895). The species therefore seems to be less wide-spread than it was one hundred years ago. 4.1xc D. x issleri : (d) There isa record for Lycopodium alpinum var. decipiens at Cattleshiel Moor near the Lesser Dirrington Law c. 36/68.53 (Anderson, 1915), however, no specimen from this locality has been traced and so the record must remain unconfirmed. 5.1 Selaginella selaginoides (Lesser clubmoss) (a-c) Recorded only once as rare but with cones in a grassy flush in heather (Ca//una vulgaris) moorland on Greenlaw Moor 36/721.485 at 210m. (d) Recorded at BRC for the 10 km grid squares 36/66 & 75 post-1950 and in Jermy et al (1978) for 36/64 pre-1950. There are also older records for Lamberton Moor c. 36/95.59 (Johnston, 1829), Penmanshiel Moor c. 36/81.67 (Johnston, 1853) and Ross Links c. 36/96.60 (Brown, 1939). Records therefore suggest species was previously more widely distributed, however, although some old sites have probably been lost due to land reclamation the species has probably been overlooked in other sites in recent years. WILLMOTT: THE FERNS OF BERWICKSHIRE 139 i, Equisetum hyemale (Rough horsetail) (d) There is a record in BRC for Westruther 36/56 dated 1956 but as this is unconfirmed it is not included in Jermy et a/(1978), but they do record the species as having been recorded pre-1950 for 36/95 & 96. The record for 36/95 is based on Johnston (1829) where it is recorded as plentiful on Lamberton Moor c. 36/95.58. However, this record is omitted by him in his later flora of the area (Johnston, 1853) which suggests that it too should be treated with suspicion. While the record for 36/96 appears to be based on a record for ‘36/96 half a mile south west of Burnmouth”. The authority for this record is given as Johnston (1853) but the record does not appear in this work. It is therefore very much an open question as to whether or not this species has ever been recorded for the country. 7.3 E. fluviatile Fig. 4a (Water horsetail) 7.4 7.4x3 = (a) (b) (c) (d) Widespread but local throughout the county. Locally abundant as an emergent at the side of ponds and rivers; frequent to occasional in marshes dominated by various species of monocotyledenous plants; recorded once as occasional in a sedge (Carex sp.) swamp at Lithtillum 36/803.409 and once as locally frequent in old peat workings overgrown by birch (Betu/a sp.) wood at Gordon Moss nature reserve 36/63.42. 0-400m though more common below 200m. Recorded post-1950 at BRC for 36/53, 54, 63, 65, 76, 77, 83, 86 and 95. E. arvense Fig. 4b (Field horsetail) (a) (b) (c) (d) (a-c) Widespread and common throughout the county though less common in the west of the county. The commonest habitat for the species was tall grass or herb communities, where it was occasional to locally abundant. Also occurred in marshes, hedges and deciduous woods, where it was less common and less abundant. Occurred finally as a casual in a range of habitats (Table 2) including arable and bare ground. 0-450m but less common above 100m and decreasing in abundance with-altitude being only recorded as rare above 300m. Recorded post- 1950 at BRC for 36/44, 54, 63, 73 & 94. Johnston (1829) records the species as ‘moist cultivated fields, too common’. The apparent reduction from this level to that in the present survey, where it was found on only a few occasions in fields, is probably a real decrease due to the intensive farming in much of the area. x litorale Recorded for only one monad. It occurred occasionally in a reed marsh (Phragmites communis) at 36/910.689 on the nature reserve at St Abb’s Head at 20m with both parents and £. pa/ustre. This record, which was confirmed by C.N. Page, is the first for the vice-county. 7.6 £. sylvaticum (Wood horsetail) oa 7.8 (a-c) (d) Recorded for only three monads, in each case in damp areas of deciduous woods on steep slopes either in a dean as at Raecleugh Head 36/740.527 (200m) or on the slope of a hill as below Edins Hall Brock 36/769.608 & 36/778.603 (170m). At each site the species occurred as small isolated patches a few metres in diameter. Recorded post-1950 at BRC for 36/54, 65, 74, 84 & 86 with older records for 36/53 (BRC), Newton Don c. 36/70.37 (Shaw, 1894) and Edington Hill Moor 36/90.56 (Henderson, 1875). E. palustre Fig. 4c (Marsh horsetail) (a) (b) (c) (d) Widespread but local throughout the county. Major habitat was marshes mostly dominated by rushes (Juncus spp.) where It was occasional to locally frequent and also occurred, but much less commonly, in tall grass and herbaceous communities where it tended to be more local picking out the wetter areas. 0-450m but commoner above 300m, though records were scarcer here. The increased frequency of the species at higher altitudes may be due to the prevalence of moorland with marshy areas here. Recorded post-1950 at BRC for 36/44, 56 & 94. E. te/mateia (Great horsetail) (a-c) Recorded for four monads, these were all within a kilometer of the sea and below 50m. Occurred in two distinct habitats, one was damp areas of deciduous woods in narrow, rocky valleys (Dunglass Dean 36/770.721 and Pease Dean 36/792.701) and the other damp areas in tall herbaceous communities on slopes overlooking the sea (Pease Bay 36/794.707, Catcairn Bushes 36/967.597 & Ross Point 140 8.1 9.1 10.1 FERN GAZETTE: VOLUME 12 PART 3 (1981) 36/962.606). At most sites the species occurred in small dense patches a few metres in diameter. (d) Recorded pre-1950 Jermy et a/ (1978) for 36/76 & 86. Botrychium lunaria (Moonwort) (d) Recorded post-1950 Jermy et a/(1978) for 36/64, 65 & 75 and pre- 1950 for 36/76, 95 & 96. There are also older records for Bemerside Hill c. 36/59.34 (Johnston, 1829), Lauder Common c. 36/50.46 (Kelly, 1874), and Penmanshiel Moor c. 36/81.68 (Johnston, 1853). Overall records indicate species has occurred widely but locally throughout county in grassland and moorland habitats. Ophioglossum vulgatum (Adder's-tongue) (d) Recorded post-1950 in Jermy et a/(1978) for 36/65 & 86 and pre-1950 at BRC for 36/77 & 85. There are also older records for ‘field above Wellrig Duns’ c. 36/76.53 (Falconer, 1932), ‘‘wood near Milne-Graden Mains” c. 36/87.44 (Donaldson, 1842), ‘‘near ruined church of St. Helens” c. 36/804.707 (Anon., 1932), Ross Links c. 36/96.60 (Brown, 1939) and Lauder Commonc. 36/50.46 (Kelly, 1874). Records therefore indicate that the species has occurred throughout the county but only locally. They further suggest the species is now more restricted in distribution than a hundred years ago but this may be because the plant has been overlooked. Osmunda regalis (Royal fern) (d) There are only three records for this species all well pre- 1950. The first record is for a plant at Coldingham Moor c. 36/86.67 (Shaw, 1872) and this is recorded by Kelly (1875) as having been taken away. Kelly (1875) also records the plant as rare ina moist boggy spot, at the side of a natural wood near Spottiswoode c. 36/60.49 and Hardy (1886) records three old plants-railed at Flass Old Wood near Wedderlie House c. 36/62.51. These two re€ords may refer to same site, in any case Shaw (1904) states the plants in the latter locations are thought to have been planted. The evidence for the species having occurred naturally in the county therefore is minimal. Cryptogramma crispa (Parsley fern) (a-c) Recorded only for the Black Hill of Earlston Fig. 3 (36/585.373). Here the species occurred in unshaded screes on the north side of the hill at 230m. The species was frequent in the habitat with fertile and sterile fronds present. (d) This site for the species has been known since at least 1853 (Johnston 1853) and is the only site where it has been seen since 1950. There are, however, pre-1950 records at BRC for 36/54, 36/64 (record dated 1930 for Houndslaw) and 36/76. This last record refers to that which is given in Johnston (1829) as south bank of the Whiteadder half a mile above Abbey St. Bathans (c. 36/74.61). FIGURE 3. Cryptogramma crispa growing in the screes on the northern side of the Black Hill of Earlston (36/585.373) in 1979. a ae ee Se $e WILLMOTT: THE FERNS OF BERWICKSHIRE 141 Be gy ge a: ooeeaeeec. ee ppp (f) FIGURE 4. The distribution of the following in Berwickshire as recorded in the present survey: a, Equisetum fluviatile; b, E. arvense; c, E. palustre; d, Polypodium vulgare agg.; e, Pteridium aquilinum and f, Oreopteris limbosperma. 142 FERN GAZETTE: VOLUME 12 PART 3 (1981) 16.1-3. Polypodium vulgare agg. Fig. 4d (Polypody) (a) (b) (c) (d) Widespread throughout the county except the Merse where it was more local. Rare to frequent on exposed and sheltered rock outcrops; occasional to locally abundant on drystone walls especially those with an earth capping or fill between the stones, but less common and less abundant (R) on mortared walls. A particular habitat was the top of beech (Fagus sy/vatica) banks, which are field boundaries made of linear mounds of earth and stones planted along their tops with beech trees, where the species was locally abundant among the beech roots. Minor habitats were unshaded screes where the species was occasional and as an epiphyte where the species was recorded once on a rowan (Sorbus aucuparia) tree. 0-400m but uncommon above 250m. Recorded post-1950 at BRC at 36/45 & 84. 16.1. P. vulgare (a) (b) (c) (d) Recorded throughout the area, this is the common segregate of the species complex in the area. Fifteen specimens were critically determined of these fourteen were this segregate. Rare to frequent on shaded and unshaded rock outcrops; frequent or locally abundant on drystone walls, particularly those capped with earth and beech banks but not recorded for mortared walls; and twice recorded for unshaded screes as. occasional to frequent. 0-400m. Recorded in Jermy et a/ (1978) post-1950 for 36/85 and in Long (1974) for ‘steep bank of Tweed below Bemersyde Hill 36/53”. 16.2 P. interjectum (a-c) (d) Recorded only once as frequent on damp, shaded outcrops for the deciduous wood in Pease Dean (36/79. 70) altitude 20m. Only other records are in Long (1974) for “36/53 steep bank of Tweed below Bemersyde Hill’ and a herbarium specimen dated 1960 at BM for humus on drystone wall at Howpark Burn (c. 36/81.65) collected by Bangerter & Hall. 16.2x1 P. x mantoniae (d) There is a pre-1950 record for 36/86 in Jermy et a/ (1978) and a herbarium specimen in BM dated 1960 for humus ondrystone wall at Howpark Burn (c. 36/81. 65) collected by Bangerter & Hall. 17.1. Pteridium aquilinum Fig. 4e (Braken) (a) (b) (c) (d) Widespread and common throughout the county except the Merse where the species was more local, probably due to the lack of habitats in this area of intensive farming. Most important habitat was moorland where it was dominant over large areas forming almost pure stands; less common and less abundant (LF-LA) but more widespread in tall herbaceous communities; and also widespread but less common and less abundant (O-F) in woods and scrub. Also recorded once or twice for the following habitats short grass, scree, rock outcrops and mortared walls. 0-400m but slightly rarer below 200m probably due to lack of habitats in lowland areas. Recorded at BRC post-1950 for 36/44, 55, 63 & 83. 19.1 Phegopteris connectilis (Beech fern) (a-c) (d) Recorded only once as occasional over a small area (c. 5m diameter) of rocky outcrops in ash/hazel (Fraxinus excelsior/Corylus avellana) wood on banks of the Whiteadder below Edin’ Hall Broch (36/772.604) at 170m (Fig. 6). Recorded at BRC post-1950 for 36/74, 75 & 77 and pre-1950 for 36/85. The species is therefore restricted to the centre of the county. The record for 36/85 probably refers to Hardy (1865) who recorded the species for Buncle Wood (c. 36/81.58), as this wood was being reclaimed for agriculture in 1979 it seems likely this site has been lost. 20.1 Oreopteris limbosperma Fig. 4f (Mountain fern) (a) (b) (c) Virtually confined to the Lammermuir Hills and the northern half of the Coastal Plateau where the species is widespread but local. Occurred most commonly and abundantly (O-F) in heather (Ca//una vulgaris) moorland, especially along the rocky banks of small valleys, and less commonly and less abundantly (R-O) in deciduous woodland often with a high abundance of Birch (Betula spp.). Also recorded once each for coniferous woodland and scrub. 100-400m but more common and abundant in higher half of this range probably due WILLMOTT: THE FERNS OF BERWICKSHIRE 143 (a) (d) (b) (e) (c) (f) FIGURE 5. The distribution of the following in Berwickshire as recorded in the present survey: a, Asplenium scolopendrium; b, A. adiantum-nigrum, ¢c, A. trichomanes agg.; d, A. ruta-muraria; e, Athyrium filix-femina and f, Cystopteris fragilis. 144 (d) FERN GAZETTE. VOLUME 12 PART 3 (1981) to the restriction of its preterred habitat to higher altitudes. Recorded post-1950 at BRC for 36/44 & 74 and pre-1950 for 36/54 & 77. 21.1 Asplenium scolopendrium Fig. 5a (Hart’s-tongue) (a) (b) (c) (d) Widespread but local to the east of a line from Cocksburnspath to Greenlaw, with only isolated occurrences to the south-west of this line. Recorded for two distinct habitats, mortared walls and woods. In the former the species was generally rare and occurred mostly in unshaded situations. In the latter the species was more abundant (R-F) and grew in damp, shaded situations. The woods in which it grew were all mixed deciduous woods on steep, rocky slopes and the plants grew either on the rock outcrops or on the woodland floor. Also recorded once each for earthed drystone wall and for tall grass habitats, though probably planted in the latter. 0-200m though more common below 100m. Recorded post- 1950 at BRC for 36/53 & 73 and pre- 1950 for 36/54. There is alsoa pre- 1950 record in Johnston (1829) for ‘‘cave between Fast-Castle and Redheugh by sea shore”’ (c. 36/86. 71). 21.2 A. adiantum-nigrum Fig. 5b (Black spleenwort) (a) (b) (c) (d) Widespread but local in eastern half of county, being commoner near the sea. Recorded as rare to frequent on unshaded, mortared, stone walls and once as occasional on unshaded, natural rock outcrops on sides of Dowlaw Dean (36/868.707). 0-100m. M. Braithwaite (Pers. comm., 1979) considers the species more common on natural rock outcrops than the above records suggest and this is borne out by an analysis of the published records for the species (Jerdon, 1866; Stuart, 1897 & Long, 1973). Recorded at BRC for 36/85 & 95 post-1950 and 36/54, 75 & 77 pre- 1950. There is also arecent record in Long (1973) for ‘Tweed bank opposite Old Melrose and below Gledswood, 36/53". 21.6 A. marinum (Sea spleenwort) (a-c) (d) Recorded only once as rare on unshaded rocks by sea at Cove Harbour (36/ 789.716) at 5m. Recorded post-1950 at BRC for 36/86, 87, 95 & 96 which are all coastal squares, the only record away from the coast is in Johnston (1853) ‘sandstone rocks by the Tweed above Ladykirk house — 7 miles from sea” (c. 36/89.46). 21.7 A. trichomanes agg. Fig. 5c (Maidenhair spleenwort) (a) (b) (c) (d) Widespread but local throughout the area. Most common on mortared walls where it was rare to frequent. It also occurred in the more natural habitats of natural rock outcrops in mixed deciduous woods at Stichill Linn (36/706.375) and Pease Dean (36/791.699 & 36/793.702) and unshaded natural outcrops in Dowlaw Dean (36/868.707). In these more natural habitats the species was only ever rare and below 60m. 0-200m but commoner below 100m. Recorded post-1950 at BRC for 36/55 & 95 and pre-1950 for 36/53. 21.7b A. trichomanes subsp. quadrivalens (d) Recorded post-1950 at BRC for 36/74, pre-1950 for 36/75, 77 & 85 all specimens checked by J.D. Lovis (Jermy et a/, 1978). The specimen for 36/74 came from an “old wall’’ which is a typical habitat of this subspecies. It would be interesting to check the records from the more natural habitats mentioned above to see if they represented the other subspecies. 21.9 A. ruta-muraria Fig. 5d (Wall-rue) (a) (b) (c) (d) Widespread but local throughout the area except the western half of the Lammermuir Hills. Occasional to frequent on mortared walls with one occurrence on a dry, unshaded rock outcrop in a quarry at Grantshouse (36/81.65) where it was occasional. 0-200m. Recorded at BRC post-1950 for 36/55 & 95. Older records suggest the species occurs more commonly on rock outcrops: Johnston (1853) “sandstone rocks by Tweed below Ladykirk House” (c. 36/88.45) and Stuart (1897) ‘‘large boulder at the Fastcastle side of Dowlaw Dean” (c. 36/86.71). 21.11 A. ceterach (Rustyback) (d) Recorded in Jermy et a/ (1978) for 36/86 as pre- 1950, this appears to be based ona record at BRC for ‘‘Renton near Cocksburnspath source C.O. Sonntag Flora of WILLMOTT: THE FERNS OF BERWICKSHIRE 145 (a) (d) (b) (e) (c) (f) FIGURE 6. The distribution of the following in Berwickshire as recorded in the present Survey: a, Polystichum aculeatum; b, Dryopteris filix-mas; c, D. pseudomas; d, D. carthusiana; e, D. austriaca and f, Blechnum spicant. 146 221 231 24.1 26.1 26.2 26.3 FERN GAZETTE: VOLUME 12 PART 3 (1981) Edinburgh’. The species is also recorded in Martin (1934) and Brown (1937) for Cocksburnspath (c. 36/77.71) and Evans (1917) for Renton House (c. 36/82.65) planted. It seems therefore that the species has been only seen in one locality in the area where it may be native. Athyrium filix-femina Fig. be (Lady-fern) (a) Widespread and common throughout the county. (b) The commonest habitat of the species was deciduous woods, where it was frequent to locally abundant. It was less common and less abundant (O) in coniferous woods, moorlands marshes and scrub. It also occurred sporadically, often as isolated plants, in tall herb, scree and hedge communities, as well as on mortared walls and shaded outcrops. (c) 0-450m though slightly more common above 100m. (d) Recorded post-1950 at BRC for 36/44 & 94. Gymnocarpium dryopteris (Oak fern) (a-c) Recorded five times. Four records were for mixed deciduous woodland on the side of a small rocky valley just above a steam — south bank of Dye above Longformacus (36/685.571), Eller Burn (36/768.612), bank of Whiteadder below Edin’s Hall Broch (36/772.604) and Drakemire Strips (36/794.608). The fifth record was for birch wood on a sphagnum bog at Coldingham Common (36/853.68 7). All localities were at about 200m and consisted of a single patch a few metres in diameter. (d) Other records suggest a sparse distribution throughout the county except for the Merse. Recorded at BRC post-1950 for 36/45, 53, 64, 74 & 75 and in Long (1973) for Chester Hill screes 36/54. There are also older records for 36/77 BRC pre- 1950, Edington Hill Moor (c. 36/90.56) in Henderson (1875) and Ale near Eyemouth (c. 36/91.63) in Hardy (1867). Cystopteris fragilis Fig. 5f (Brittle bladder-fern) (a) Widespread but local throughout the county though avoiding the south east and north west. (b) Occurred most commonly on mortared, stone walls that were dry and unshaded, though only a few plants in each locality. Recorded once for a natural, rock outcrop in mixed deciduous wood at Stichill Linn (36/ 706.375) where it was occasional and once as rare on the stoney bank of a ditch in a garden at Silverwells (36/879.664). (c) 50-200m. (d) Other records record plants in the south east of the county, thus there are post- 1950 records at BRC for 36/84 & 95 and older records for near Mains (c. 36/88.55) Johnston (1829) and for banks of the Leader Carolside (c. 36/56.39) Kelly (1874). Polystichum lonchitis (Holly fern) (d) There is a pre-1950 record in BRC for 36/76 which may refer to the other record for the county which is in Hardy (1885). Here the species is recorded for ‘‘the side of the Whiteadder at the base of Moonjee very dwarf but true’’, which sounds rather more like depauperate P. acu/eatum than P. /onchitis. Evans (1917) regards the species as an escapee in the Borders. The evidence is therefore minimal that the species ever occurred in the county. P. aculeatum Fig. 6a (Hard shield-fern) (a) Widely distributed but local in the east of the county. (b) Occurred most commonly in deciduous wood (O-LF), often in narrow rocky valleys and on natural, rock outcrops (F) in such woods. There were single occurrences in tall grass, a hedge and on a drystone wall. (c) 0-250m but commoner at lower altitudes particularly below 50m. (d) Other records fill in the species occurrence in the east of the county and extend its range to the south west. Recorded post-1950 at BRC for 36/54, 64 & 86 and as locally common on the steep bank of the Tweed below Bemersyde Hill 36/53 Long (1974). Anumber of old records refer to P. /obatumas distinct from P. acu/eatum and P. setiferum, thus Hardy (1879) records the three species for Pease Dean near the bridge (c. 36/79.70) and Stuart (1897) for Dowlaw Dean (c. 36/86.70). Polystichum lobatum is nowadays considered a form or variety of P. acu/eatum with narrower, less-divided fronds. P. setiferum (Soft shield-fern) (a-c) Frequent to locally abundant on the floor of the mixed deciduous woodlands in Tower Dean (36/78.69) and Pease Dean (36/79.69 & 36/79.70), and frequent ina hedge nearby (36/793.701). Also occurred twice on walls. Once at Mellerstain (36/651.384) where there is a small colony on a damp, mortared wall and once at WILLMOTT: THE FERNS OF BERWICKSHIRE 147 27.2 27.3 27.5 27.8 a js (d) Manderston (36/810.547), where there was a single plant on a drystone wall. This latter plant probably arose from cultivated plants nearby. Altitudinal range was O- 150m. Other records are all for sites within a few kilometers of the sea or coastal 10 km grid squares. Thus there is a post-1950 record at BRC for St. Abbs and area 36/96 anda pre-1950 record for 36/86. There are also older records for Dunglass Dean (c. 36/76.71) (Johnston, 1853) and for Dowlaw Dean (c. 36/86.70) (Stuart, 1897). Dryopteris filix-mas Fig. 6b (Male-fern) (a) (b) (c) (d) Widespread and common throughout the area. There were three major habitats for this species in the area: woods and scrub including hedges; tall grass; and walls and outcrops. It was commonest and most abundant (F) in deciduous woods; equally common but less abundant (F-O) in scrub and hedges; and both less common and less abundant (O) in conifer plantations. It was the only fern that was at all common or abundant (O) in stands of tall grass and herbaceous vegetation. It was about as common on walls and outcrops as in tall grass communities but less abundant (O-R). It was noticeably less common and less abundant (R) in heath communities and there were odd records for marsh, scree and short grass communities. 0-450m. It was equally common at all altitudes but became less abundant above 200m and only recorded.as rare above 300m. Recorded post-1950 at BRC for 36/44 & 56. D. pseudomas Fig. 6c (Scaly male-fern) (a) (b) (c) (d) Widespread and common in the Lammermuir Hills, Coastal plateau and the east of the Low Uplands of Old Red Sandstone; and widespread but more local in the rest of the county. The habitat in which the species was most common was deciduous woods, though it was only occasionally in abundance here. It was less common in heathlands but more abundant (F) in this habitat. It was also less common in coniferous plantations and scrub where it was rare in abundance. There were also odd occurrences in hedge, tall herb, scree, outcrop and mortared wall communities, where it was only recorded as rare in abundance. 0-450m but commoner and more abundant above 150m. Recorded post-1950 at BRC for 36/56, 64 & 94. D. aemula (Hay-scented buckler-fern) (d) Johnston 1853 comments under Lastraea foenisecii, which is a synonym for this species (Dyce, 1978), ‘| suspect it to be frequent in area’ but gives no localities. There is also a pre-1950 record at BRC for Dirrington Law 36/65 but the card states this record to be an error. D. carthusiana Fig. 6d (Narrow buckler-fern) (a) (b) (c) (d) Widespread but local thoughout the south and east of the county. Recorded most commonly for deciduous woods especially those dominated by birch (Betula spp.) for example Buncle Wood (36/813.585) and Silverwells (36/878.665). However, it was only more abundant than rare where there was a birch wood on a sphagnum bog, as at Coldingham Common (36/853.687) and Gordon Moss (36/63.42) where the species was occasional and frequent respectively. Also recorded as rare in a conifer plantation (Lauder Common 36/504.467), a rush marsh (Coldingham Common) and a gorsey heath (Buncle Wood). 50-300m. Recorded post-1950 at BRC for 36/45, 53 & 75 and in Henderson (1875) for Edington Hill Moor (c. 36/90.56). D. austriaca Fig. 6e (Broad buckler-fern) (a) (b) (c) Widespread and common throughout the area. The habitat in which the species most commonly occurred was deciduous woodland, where it was frequent in abundance. It also occurred commonly but less abundantly (F-O) in coniferous plantations and heaths. It occurred less commonly and less abundantly (O) in scrub and on drystone walls and outcrops; and there were odd occurrences in marsh, tall herb, short grass, scree, hedge and mortared wall communities. A particular habitat of the species was amongst the roots of beech (Fagus sylvatica) trees on banks and it was recorded twice as an epiphyte, once on oak (Quercus spp.) and once on alder (A/nus glutinosa). O0-450m but gradually more common with increasing altitude. 148 FERN GAZETTE. VOLUME 12 PART 3 (1981) 27.9x8 D. x deweveri (a-c) Recorded twice, once for Gordon Moss (36/63.42) as rare in a birch (Betula spp.) wood on a peat moss at 140m with both parents; and once for Duns Castle grounds (36/77.55) as rare in a mixed deciduous wood at 170m. At the latter site D. carthusiana was not recorded. Both records were determined by M. Gibby. (d) No previous records. 28.1 Blechnum spicant Fig. 6f (Hard fern) (a) Widespread but local in the Lammermuir Hills, the north section of the Coastal Plateau and the east of the Low Uplands of Old Red Sandstone. (b) Recorded as commonest and most abundant (F-O) in heaths, and less common and abundant (O-R) in deciduous woods especially those dominated by birch (Betula spp.) and/or on steep slopes. There were also odd records for conifer plantations, scrub, tall herb and hedge communities. (c) 100-450m but becoming commoner at higher altitudes. (d) Recorded at BRC post-1950 for 36/44, 54, 64, 77, 84, 87 & 96. HABITAT LISTS Species lists and notes are given for selected habitats. Abundance only refers to abundance in examples of the habitat in which the species in question occurred, and is recorded on the Dafor scale. How often the species occurred in the habitat is givenasa percentage frequency, rounded to the nearest ten percent. Moors This habitat includes all areas dominated by ericaceous undershrubs, and upland areas dominated by grasses used as open grazing. It was evident in the field that rocky areas of moorland along the sides of the small streams had a greater abundance of ferns than areas of general moorland. These two habitats were therefore recorded separately. The list refers to six examples of rocky moorland and twenty-two of general moorland. Species Abundance Frequency (%) General Rocky General Rocky Equisetum arvense O R 10 20 Polypodium vulgare agg. = R 30 Pteridium aquilinum LA F 70 50 Oreopteris limbosperma O-F F 30 70 Athyrium filix-femina O O 40 70 Dryopteris filix-mas R-O R 20 70 D pseudomas F F 10 80 D. carthusiana R - 10 0 D. austriaca O-F R-F 50 100 Blechnum spicant O-F O 60 50 The most important ferns of moorlands in the area were Pteridium aquilinum, Dryopteris austriaca and Blechnum spicant. The greater abundance of ferns in rocky areas was due to a greater abundance and frequency of the ferns present in the general areas. The only significant exceptions to this latter point were Po/ypodium vulgare agg. which did not occur in the general areas and Pteridium aquilinum which was less important in the rocky areas. The greater occurrence of ferns in the rocky areas was possibly due to crevices at the base of rocks providing habitats for prothalli. Although in the case of Po/ypodium vulgare agg. the provision of thin, dry, organic soils WILLMOTT: THE FERNS OF BERWICKSHIRE 149 over a Solid substrate, which is the preferred habitat of the sporophyte, was probably more important. The reason for the lesser occurrence of Pteridium aquilinum in the rocky areas is less obvious, though it may be that as these areas were near streams the water table was higher and the soils therefore too water-logged. Walls There are few outcrops of limestone or base-rich rocks in the county, the stone walls of the area are therefore made of siliceous rocks. The major source of variation in thes walls was the presence or absence or mortar/cement between the stones. The few brickwalls surveyed are included with the mortared walls in the list and dry stone- faced earthbanks with the drystone walls. The list refers to twenty examples of drystone walls and fifty-three examples of mortared walls. Species Abundance Frequency (%) Drystone Mortared Drystone Mortared Polypodium vulgare agg. O-F R-O 70 20 Pteridium aquilinum — R 0 40" Asplenium scolopendrium O R 10* 30 A. adiantum-nigrum _ R 0 TO A. trichomanes agg. O R-F 30 40 A. ruta-muraria O O-F 0 50 Athyrium filix-femina R R-O 1 20 Cystopteris fragilis ~ R 0 20 Polystichum aculeatum R _ 10" 0 P. setiferum R R 40* +0= Dryopteris filix-mas R-O R-O 60 60 D. pseudomas _ R 0 Tc D. austriaca 0 R 40 10 * Represents only one record The two most important ferns on walls were Dryopteris filix-mas and Asplenium trichomanes agg. Also important were Polypodium vulgare agg. and Dryopteris austriaca on drystone walls, and Asp/enium ruta-muraria on mortared walls. No other species were seen more than once on drystone walls but mortared walls had six other species which occurred more than once. The major reason for the greater species richness of the mortared walls was probably their relative base richness due to the mortar, though the presence of smaller crevices in mortared walls may favour colonization by ferns. One particularly noticeable feature of the ecology of wall ferns was the preference of Cystopteris fragilis and Asplenium scolopendrium for drier, less shaded habitats than these species prefer in other areas of Great Britain, for example the Burren (Willmot, 1979) and Derbyshire (Willmot, 1977). Although the records for A. scolopendrium were mostly for small, sterile isolated plants. This change in preference may be due to the relative coolness of the area, compared to the ares in Willmot (1977 & 1979), compensating for the drier nature of the habitats. It is also worth noticing the importance of walls for the existence of certain species in the county. Although no species were confined to walls, Asp/enium adiantum-nigrum, A. trichomanes agg., A. ruta-muraria and Cystopteris fragilis were much more frequent (Table 2) on walls than on their natural habitat of outcrops. 150 FERN GAZETTE: VOLUME 12 PART 3 (1981) Deciduous Woods It was noticeable during the survey that the fern flora of the deciduous woods varied greatly, with woods on steep slopes having a richer flora than woods and plantations of flatter ground. The list refers to thirty-four examples of wood on steep slopes and forty- five on flatter ground. Abundance Frequency (%) Flat Steep Flat Steep Equisetum fluviatile O _ 10* — E. arvense R-O O 10 10 E. sylvaticum — R-O 0 10 E. palustre O _ 10 0 E. telmateia = R O 10* Polypodium vulgare agg. — R-O 0 10 Pteridium aquilinum O-F O-F 30 50 Phegopteris connectilis — R 0 10* Oreopteris limbosperma R-O O 10 10 Asplenium scolopendrium R R-F 10* 20 Athyrium filix-femina R-F O-F 80 90 Gymnocarpium dryopteris R R-O 10* 10 Polystichum aculeatum = R-O 0 40 P. setiferum — = 0 10 Dryopteris filix-mas O-F O-F 100 100 D. pseudomas R-O R-F 40 90 D. carthusiana R _ 10 0 D. austriaca O-F O-F 80 100 D. x deweveri R _ 10* 6) Blechnum spicant O R- 10 20 * Represents only one record Ignoring Equisetum arvense and E. palustre, as casual species in woods and the four taxa recorded only once, there are fourteen species whose preference between the two types of wood can be judged. Three species (Athyrium filix-femina, Dryopteris filix-mas and D. austriaca) are very common in both sorts of wood. Amongst the remaining eleven rarer species nine showa preference for woods on steep slopes, one (D. carthusiana) a preference for woods on the flat, and one (Oreopteris limbosperma) no preference. To understand these preferences it is necessary to examine more closely the differences between the two groups of woods. Many of the woods on the flat were obviously plantations, as the tree species in them were introductions (e.g. Fagus sy/vatica) or because their layouts showed they were shelter-belits. The woods on steep slopes, on the other hand, often had very rocky floors. This suggests that they could never have been cleared for agriculture and therefore that they might be remnants of the ‘Oak forest with birch” which Birks (1977) considers the natural vegetation of the area. In fact many of these woods contained high proportions of Quercus sp. and Betula sp., along with U/mus glabra, Fraxinus excelsior and Corylus avellana which Birks (1977) considers were associates of the oak and birch. If the major difference between these two groups of woods is that those on the flat are plantations on cleared land and those on the slopes are woods on sites never cleared, there are two explanations for the preference of a species for the woods or slopes. The first explanation is that a species was once more widespread in the area and has been unable to recolonise the new woods. The second is that a WILLMOTT THE FERNS OF BERWICKSHIRE 151 species was always Commoner in the sites not cleared and has not spread to the newer woods because the conditions there are not suitable for it. Three species appear to fall into the second category (Po/ypodium vulgare agg., Asplenium scolopendrium and Polystichum aculeatum), as they have a preference for rocky outcrops which are only common in the woods on slopes. This leaves six species that might fall into the first category. Three of these species (Pteridium aquilinum, Dryopteris pseudomas and Blechnum spicant) show only a quantitative preference but three species (Equisetum sylvaticum, Gymnocarpium dryopteris and Polystichum setiferum) are virtually confined to woods on slopes. These latter three species should therefore be considered as possible primary woodland indicators, in the sense of Peterken (1974), in the area. The species with a preference for the woods on the flat (Dryopteris carthusiana) illustrates the fact that these woods were not a homogeneous group. The majority occurred on mineral soils but a few occurred on wet, peaty soils and this species occurred in the woods with this soil type, which is its preferred habit elsewhere. These soils do not develop on steep slopes in the area which explains the absence of the species from the woods on slopes. SUMMARY OF THE FLORA Thirty-two species and three hybrids have been recorded in the area since 1950 and six taxa before 1950 but not since (Table 1). The thirty-two species is an accurate reflection of the true fern flora of the area, as there are no species common in adjacent vice-counties which are absent from Berwickshire. However, hybrids are probably under-recorded due to the lack of study of them, and Dryopteris x tavelii and Polystichum x bicknellii could no doubt be added to the list if suitable habitats were examined. Division of the species into the floristic elements of Birks (1976) (Table 3) demonstrates the general biogeographic affinities of the flora. Eighteen species belong to elements widespread in Europe anda further eleven to elements that though more restricted in Europe (Mediterranean/ Atlantic) are still widespread in the British Isles. The eighteen species represent 80% of the former elements in the British Isles while the eleven only 50% of the latter elements. This lower percentage of the Meditteranean/Atlantic elements is to be expected as many of the species have a TABLE 3: THE PHY TOGEOGRAPHICAL AFFINITIES OF THE FERN FLORA OF BERWICKSHIRE (Based on the floristic elements of Birks (1976) Floristic Element Distribution in Europe Number of Species in Element in: (a) Berwickshire (b) (c) post-1950 (pre-1950) British Isles Europe 1. Cystopteris fragilis Widespread 12(1) 14 14 2. Pteridium aquilinum Widespread 6 8 8 3. Asplenium scolopendrium Mediterranean/Atlantic 9(2) 14 14 4. Adiantum capillus-veneris Mediterranean/Atlantic Z 6 6 5. Woodsia alpina Arctic-alpine 2 6 6 6. Polystichum lonchitis Widespread montane 1(1) 9 11 and sub-montane 7. Selaginella helvetica Alps, Pyrennes and (1) 3 11 lowlands of mainland 8. Hymenophy/llum wilsonii Western seaboard (1) 4 4 2 ee ee eee eee eee eee EE 152 FERN GAZETTE: VOLUME 12 PART 3 (1981) western distribution in the British Isles, and Berwickshire is on the eastern coast. The remaining three species (Diphasiastrum alpinum, Selaginella selaginoides and Cryptogramma crispa) are all rare in the county and belong to northern and/or upland elements. Thus overall the flora consists of species widespread in the British Isles with the absence of species showing a marked western distribution, and the presence of a few northern and/or upland species. The distribution of the six taxa for which only pre-1950 and often questionable records exist among the elements shed further light on the status of these species in the area. Three species (Equisetum hyemale, Osmunda regalis and Asplenium ceterach) belong to elements well-represented in the flora. There are no reasons to doubt the records for the latter two species, so these species at least were native in the area. However, the paucity of records for these species and their marked western distribution in the British Isles suggests they were only casual species. The remaining three taxa (Diphasiastrum x tssleri, Polystichum lonchitis and Dryopteris aemula) all belong to elements represented poorly or not at all in the extant flora. As the records for all three species are questionable, there is no reason to consider these species as ever having been native in the area. Examination of all records for each species reveals five patterns of geographical distribution for those species where sufficient records exist to construct a distribution. The ten species in the list below occur throughout the area: Equisetum fluviatile Asplenium trichomanes agg. E. arvense Athyrium filix-femina E. palustre Dryopteris filix-mas Polypodium vulgare agg. D. pseudomas Pteridium aquilinum D. austriaca Two species (Asplenium ruta-muraria and Dryopteris carthusiana) occur throughout the area except for the north-west, and three species (Lycopodium clavatum, Oreopteris limbosperma and Blechnum spicant) throughout the area except for the lower ground in the south and east. A further seven species, given in the list below, also avoid the lower ground in the south and east but are more restricted in their distribution on higher ground: Huperzia selago Botrychium /unaria Diphasiastrum alpinum Phegopteris connectilis Selaginella selaginoides Gymnocarpium dryopteris Equisetum sylvaticum These species occur in the Coastal Plateau and the northern-central part of the area, that is the eastern half of the Lammermuir Hills and of the low uplands of Old Red Sandstone. Phegopteris connectilis is rather anomalous here as it only occurs in the norther-central part. The final pattern of distribution is shown by the seven species below which are restricted to the eastern half of the area or to areas near the coast: Equisetum telmateia Cystopteris fragilis Asplenium scolopendrium Polystichum aculeatum A. adiantum-nigrum P. setiferum A. marinum These distribution patterns are demonstrated by the following figures: widespread Dryopteris filix-mas (Fig. 7b), widespread except north-west Asp/enium ruta-muraria (Fig. 5d), widespread except south and east Blechnum spicant (Fig. 7f), Coastal Plateau and northern-central is not shown well in the distribution maps here as they only refer to records from this survey, and eastern Po/ystichum aculeatum (Fig. 7a). The widespread species are all of broad ecological tolerance or species whose habitats are common throughout the area. The ‘“‘widespread except south and east”, and ‘‘Coastal Plateau and northern-central region” species are all species with well known preferences for upland environments. However, why the latter group are more WILLMOTT. THE FERNS OF BERWICKSHIRE 153 restricted in distribution is less obvious. The eastern species belong mostly to the Mediterranean/Atlantic elements of Birks (1976) and are probably restricted to the east of the area due to the milder climate there. Reasons for the apparent avoidance of the north-west by Asp/enium ruta-muraria and Dryopteris carthusiana are unclear. ACKNOWLEDGEMENTS My thanks are due to the field naturalists, both past and present, who have worked in the area and recorded ferns. Without their efforts this survey would not have been possible. In particular | should like to mention M. Braithwaite who also showed me many localities. My thanks are due to the following authorities for checking critical material in the genera indicated: Dryopteris, C.R. Fraser-Jenkins and M. Gibby; Equisetum, C.N. Page; Polypodium, J.A. Crabbe and R.H. Roberts; and Po/ystichum, A. Sleep. My thanks are also due to the Biological Records Centre, Monk’s Wood for access to their fern records for the area. Finally | wish to thank the following for technical help with the preparation of the manuscript: figures, A. Skinner; photographs, P. Holdcroft; and typing, Miss C. Pike. REFERENCES ANDERSON, A., 1915. List of the less common plants in the area of the club. Hist. Berwicks. Nats. Glab. 22° 227-272. ANON., 1894. Reports of meetings for 1892. Hist. Berwicks. Nats. Club 14: 9-86. ANON., 1932. Reports of meetings 1931. Ast. Berwicks. Nats. Club 27: 294-326. BIRKS, H.J.B., 1976. The distribution of European pteridophytes: A numerical analysis. Vew Phytol. 77: 257-287. BIRKS, H.J.B., 1977. The Flandrian forest history of Scotland: a preliminary synthesis. /n F.W. Shotton (Ed), British Quaternary Studies: Recent Advances: 119-135. Oxford. BROWN, J., 1937. Some plants found during 1937. Hist. Berwicks. Nats. Club 29: 294-296. BROWN, J., 1939. Some plants found during 1939. Hist. Berwicks. Nats. Club 30: 199-203. CLAPHAM, A.R., TUTIN, T.G., & WARBURG, E.F., 1962. Flora of the British Isles. Cambridge. DANDY, J.E., 1969. Watsonian vice-counties of Great Britain. London. DONALDSON, J.S., 1842. Selby address. Hist. Berwicks. Nats. Club 2: 261-269. DYCE, J.W., 1978. Fern nomenclature — old and new. Bulletin of the Brit. Pterido. Soc. 1: 261- 263. EVANS, A.H., 1917.Notes on plants found in the district worked by the Berwickshire Naturalists’ Club. Hist. Berwicks. Nats. Club 23: 217-235. FALCONER, A.A., 1932. Botanical notes. Hist. Berwicks. Nats. Club 27: 393-396. GREGORY, S., 1976. Regional Climates. /n T.J. Chandler & S. Gregory (Eds), 7he Climate of the British Isles: 330-342. London. GRIEG, D.C., 1971. British Regional Geology: The South of Scotland. Edinburgh. HARDY, J., 1865. Plant notes. Hist. Berwicks. Nats. Club 5: 232-236. HARDY, J., 1867. Anniversary address. Hist. Berwicks. Nats. Club 5: 371-406. HARDY, J., 1879. Reports of meetings. Hist. Berwicks. Nats. Club 9: 425-497. HARDY, J., (1885). Miscellanea. Hist. Berwicks. Nats. Club 10: 607-610. HARDY, J., 1886. Report of meetings for 1885. Hist. Berwicks. Nats. Club 11. 10-93. HENDERSON, G., 1875. On Chirnside parish: the estate of Edington. Hist. Berwicks. Nats. Club 7: 95-104. JERDON, A., 1866. Annual address. Hist. Berwicks. Nats. Club 5: 241-248. JERMY, A.C., ARNOLD, H.R., FARRELL, L. & PERRING, F.H., 1978. Atlas of Ferns of the British /sles. London. JOHNSTON, G., 1829. Flora of Berwick-upon-Tweed. Vol. 2. Edinburgh. JOHNSTON, G., 1853. The Natural History of the Eastern Borders. Vol. 1. London. KELLY, A., 1874. Habitats for some Berwickshire plants. Hist. Berwicks. Nats. Club 7: 125-126. KELLY, A., 1875. Localities for some border plants. Hist. Berwicks. Nats. Club 7: 327. LONG. A.G., 1973. Natural history observations during 1973. Hist. Berwicks. Nats. Club 39: 204- 209. LONG, A.G., 1974. Natural history observations during 1973-74. Hist. Berwicks. Nats. Club 40. 62-63. . LYNN, F., 1895. Buncle Edge forts. Hist. Berwicks. Nats. Club 15: 365-376. 154 FERN GAZETTE VOLUME 12 PART 3 (1981) MARTIN, I.H., 1934. Field-club Flora of the Lothians. Edinburgh. MITCHELL, G.F., 1948. Late-glacial deposits in Berwickshire. New Phytol 47: 262-264. NEWEY, W.W., 1968. Pollen analyses from south-east Scotland. 7rans. Proc. Bot. Soc. Edinburgh 40: 424-434. PETERKEN, G.F., 1974. A method for assessing woodland flora for conservation using indicator species. Biol. Cons. 6: 239-245. SHAW, W., 1872. Botanical notices. Hist. Berwicks. Nats. Club 6: 282-283 SHAW, W., 1894. Notes on some rarer plants. Hist. Berwicks. Nats. Club 14: 401-404. SHAW, W., 1904. Reports of meetings for 1901. Avst. Berwicks. Nats. Club 18: 17-122. SISSONS, J.B., 1976. The Geomorphology of the British Isles: Scotland. London. STUART, C., 1897. Lists of plants gathered in Dowlaw Dean. Hist. Berwicks. Nats. Club 16: 165- 166. WHITTOW, J.B., 1979. Geology and Scenery in Scotland. Harmondsworth. WILLMOT, A., 1977. A pteridophyte flora of the Derbyshire dales national nature reserve. Fern. Gaz. 11: 279-284. WILLMOT, A., 1979. An ecological survey of the ferns of the Burren, Co. Clare, Eire. Fern Gaz. 72: 9-28. FERN GAZ. 12(3) 1981 155 CHEILANTHES CORIACEA, AN AFRO-ARABIAN FERN SPECIES NEW TO IRAN C.R. FRASER-JENKINS c/o Botany Department, British Museum (Natural History), Cromwell Road, London, SW7, England. ABSTRACT Cheilanthes coriacea Decne. is reported for the first time from Iran, in the south- west of the country. It is the third known example of an Afro-Arabian fern species in Iran; the other species there being European elements. Some distribution patterns of western Asian species are pointed out in relation to Iran. INTRODUCTION A parcel of ferns was sent to Prof. K.U. Kramer at Zurich from the herbarium at Ariamehr Botanic Garden, Teheran, Iran (TARI) in connection with the account of the Pteridophyta for the Flora lranica project. Included among them was a small unidentified specimen of Che//anthes, labelled: ‘‘Iran, prov. Fars: Kazerun, Abgineh., S. slope on cliffs, hanging. In crevice of rocks. Leg. Foroughi and Assadi, no. 15098, 10 Oct. 1974." The two collectors are both at Ariamehr and have between them and also with Dr. P. Wendlebro of Goteborg, Sweden, collected a great number of useful and important specimens. Mr. Assadi specialises in ferns. Since the specimen came from Fars province, near the Persian Gulf in south-west Iran, it seemed possible that it might belong to the small Afro-Arabian element in the flora, rather than to the European one. The only other non-European fern species in Iran are Pteris dentata Forssk. subsp. flabe//ata (Forssk.) Runemark, known from the south Caspian coast in Mazanderan province, and Onychium divaricatum (Poir.) Alston (Syn.: O.me/anolepis (Decne.) Kunze, from south-west Iran; information about both these species is given by Wendlebo (1976), who also erroneously lists the specimen under consideration here as Cheilanthes marantae (L.) Domin, a slightly similar Iranian species which is part of the European element. A check through the African material of Che//lanthes at the British Museum (Natural History), London, immediately showed that this curious specimen belongs to Che//lanthes coriacea Decaisne, a species hitherto known only from north-eastern Africa and south-western Arabia (Somalia, Ethiopia, Sudan, Chad (Tibesti) and the Yemen). Thus it is a third and previously overlooked Afro-Arabian species in the Iranian fern flora. It’s presence in Iran suggests that other overlooked Afro-Arabian species might possibly be present there, especially in the rather little known regions of the southern Zagros mountains in Fars and adjacent provinces, though they would have to be species adapted to dry conditions. TAXONOMY Cheilanthes coriacea resembles C. marantae in being covered with closely adpressed, pale or pale-brown, lanceolate scales on the lower surface of the lamina, but may be readily distinguished by being a smaller plant in all its parts, with a long, thin, nearly glabrous stipe, usually as long as the lamina. The lamina is short, broadly ovate- lanceolate and widest towards the base; the pinnules are markedly narrower than in C. marantae and the basiscopic ones are lengthened towards the base of the lower pinnae. The fronds are normally up to c.10 cm tall. 156 FERN GAZETTE VOLUME 12 PART 3 (1981) PHYTOGEOGRAPHY Phytogeographical connections between the Iranian fern flora and that of north- eastern Africa and Arabia are few, involving only the three examples mentioned. But the rest of the Iranian fern flora is European in nature, with close connections to the Caucasus and Turkey. It is noticeable though that it is considerably impoverished compared to those areas, probably due to a combination of the present climate and the lack of a direct high mountain connection between them. There are 31 species of ferns and fern-allies in Iran that are common to Europe. Further afield, there are several European ferns that have spread into the West and Central Himalayas and are also common to Iran. All are species that are tolerant of a rather high degree of drought, though, apart from a few more highly xeromorphic exceptions (Chei/anthes persica (Bory) Mett. ex Kuhn, C. pteridioides (Reich.) C.Chr., Cystopteris fragilis (L.) Bernh. etc.), they are not now present in the desert gap of eastern Iran and western Afghanistan. A few species, such as Polystichum lonchitis (L.) Roth, occur in the Caucasus and West Himalayas (and Tien Shan), but apparently not in lran. There are no examples known of extant Himalayan fern species spreading westwards to Europe and Iran, but there are some indirect connections between the two regions, for example the probable relationship between the European and Iranian Dryopteris affinis (Lowe) Fras.-Jenk. complex and the Himalayan species D. paleacea (Swartz) Hand.-Mazz. (syn.: D. wallichiana (Spreng.) Hyl.) outlined by Fraser-Jenkins (1980), which suggests that there may have been connections during the Tertiary period. There are also some important connections between the Himalayan and African fern floras with some species or groups incommon, for exampie Athyrium schimperi Moug. ex Fée, which occurs in the Himalayas, Rajasthan and West and East Africa, Polystichum luctuosum (Kunze) J. Smith, from South Africa, Madagascar and the West Himalayas, with the very closely related P. tsus-simense (Hook.) J. Smith in south-east Tibet, China, Taiwan and Japan, or Adiantum reniforme agg. from Macaronesia, Madagascar, Réunion and Szechuan, West China (Lin, 1980). Several other species aggregates are common to Africa and the Himalayas. Unfortunately, in tentatively recording the existence in Iran of various species, Parsa (1950) and Parsa and Maleki (1978) did not take phytogeographical considerations into account and did not allow for the Iranian fern flora being considerably impoverished compared to Turkey and the Caucasus. Hence many of their records are spurious and misleading and they also make a number of erroneous records due to misidentification. An earlier work by Fedtchenko (1946) is more accurate, though it also over-records a few species, mainly those in complexes which were not fully elucidated at the time as no modern cytological investigations had then been carried out. An account by Wendlebo (1976) is in general the most reliable and accurate. ACKNOWLEDGEMENTS “ The author is grateful to Prof. K.U. Kramer of Zurich and Prof. T. Reichstein of Basel for encouraging the writing of this article. REFERENCES FEDTCHENKO, B.A., 1946. Ferns of Iran. Sborn. Nauch. Rabot. Bot. Inst. Komarov. Akad. Nauk S.S.S.R. 1945: 63-74. FRASER-JENKINS, C.R., 1980. Dryopteris affinis: a new treatment for a complex species in the European Pteridophyte flora. Wi//denowia 10: 107-115. LIN, You-Xin, 1980. New taxa of Adiantum L. in China. Acta Phytotax. Sinica 18: 101-105. PARSA, A., 1950. Flore de I'/ran. Teheran. PARSA, A., & MALEKI, Z., 1978. Flora of Iran (2 edn). Teheran. WENDLEBO, P., 1976. An annotated check-list of the ferns of Iran. J. Bot. 7: 11-17. FERN GAZ 12(3) 1981 157 A CHROMOSOME COUNT FOR ASPLENIUM ANCEPS FROM THE CANARY ISLANDS H. RASBACH & K. RASBACH Kurklinik Glotterbad, D-7804 Glottertal, West Germany and J.J. SCHNELEER Institut fur Systematische Botanik, Zollikerstr. 107, CH-8008 Zurich, Switzerland ABSTRACT A chromosome count for Asplenium anceps Lowe ex Hooker & Grev., is given for the first time from the Canary Islands. ASPLENIUM ANCEPS Asplenium anceps Lowe ex Hooker & Grev., a member of the Asp/enium trichomanes group, is a species endemic to the Macaronesian islands (Azores, Madeira, Canary Islands). It was in the last century that the species was recognised (Lowe 1831, Bolle 1866). Recently Benl & Sventenius (1970), Page (1971, 1977) and Lovis et al. (1977) have published various localities for this fern. Apart from Madeira, where the plant shows a wider distribution, only few localities are known, and from the Azores only two specimens of A. anceps have so far been reported. In the field A. anceps can easily be recognised by its rachis, which shows, in addition to the two adaxial wings present on A. trichomanes, a third abaxial wing (for further details see Lovis et al. 1977). All previously cytologically examined plants originated from Madeira and the Azores. They were all diploid with n = 36 pairs of chromosomes at meiosis (Lovis in Meyer 1969, Lovis et al. 1977, Lovis unpubl.). lt was thus of interest to see if the plants from the Canary Islands showed the same ploidy level. In the course of an excursion to the western part of the Canary Islands, H. and K. Rasbach visited the great Barrancos in the northern part of La Palma to west of Barlovento, an area particularly exposed to the northern trade wind (Page 1971). On May 21st and 22nd 1979 they found approximately 60 specimens of A. anceps in the Barranco Franceses at 1050-1110 m altitude (fig. 1). On 23rd May a further 12 plants were discovered in Barranco Gallegos at about 850 m. A. anceps grows on small ledges and steep slopes between high rock faces. Even here it is usually exposed to the north and growing in the shade of trees and bushes. The fern occurs in the moist, misty zone which is characterised by drizzle during summer. The plant grows in loose humus soil and it can be regarded as an element of the “‘Laurisilva’’ which is shown also by the species it is associated with. One plant taken from the Barranco Franceses (Ras-65) is now being cultivated in Basle by Prof. Dr. T. Reichstein (TR-5072). One frond of this plant was fixed in the field (absolute ethanol : acetic acid 3:1), the solution was changed after 24 hours and after 4 days the fixed material was brought into 70% ethanol and kept in a refrigerator. The chromosome count was made in Zurich by J.J. Schneller. Meiosis is regular and at metaphase |, 36 chromosome pairs can be distinguished (fig. 1, b and c). The chromosome numbers thus correspond to those already found in plants from the Azores and Madeira. All these plants are thus diploid. 158 FERN GAZETTE: VOLUME 12 PART 3 (1981) FIGURE 1. a, Asplenium anceps Lowe ex Hook. & Grev. from Canary Islands, La Palma, Barranco Franceses (Photo: H. & K. Rasbach); b, meiosis in A. anceps from Barranco Franceses, La Palma (Ras-65), show 36 bivalents (not all in the same plane) (Photo: J.J. Schneller); c, explanatory diagram of same cell; d, silhouette of whole frond. RASBACH, RASBACH & SCHNELLER: ASPLENIUM ANCEPS 159 ACKNOWLEDGEMENT We thank Prof. Dr. T. Reichstein very much for his stimulation and interest. REFERENCES BENL, G. & SVENTENIUS, E.R., 1970. Beitrage zur Kenntnis der Pteridophyten-Vegetation und -Flora in der Kanarischen Westprovinz. Nova Hedwigia 20: 413-462. BOLLE, C., 1866. Die Standorte der Farrn auf den canarischen Inseln, pflanzentopographisch geschildert. Ze/tschr. Ges. Erdk. 1: 209-238, 273-287. LOVIS, J.D.. HELGA RASBACH, RASBACH, K. & REICHSTEIN, T., 1977. Asplenium azoricum and other ferns of the A. trichomanes group from the Azores. Am. Fern Journ. 67: 81-93. LOWE, R.T., 1831. Primitiae faunae et florae Maderae et Portus Sancti. 7rans. Cambridge Phil. Soc.4: 1-70. MEYER, D.E., 1969. Asplenium newmani Bolle von den Kanarischen Inseln ist ein X Asplenoceterach. Willdenowia 5: 221-229. PAGE, C.N., 1971. Three pteridophytes new to the Canary Islands. Brit. Fern Gaz. 10: 205-208. PAGE, C.N., 1977. Anecological survey of the ferns of the Canary Islands. Fern Gaz. 17:297-312. 160 FERN GAZETTE: VOLUME 12 PART 3 (1981) REVIEW FERNS AND FERN ALLIES OF GUATEMALA : PART 2 POLYPODIACEAE by Robert G. Stolze. Fieldiana, Botany New Series no. 6,522 pp. 1981. Published by the Field Museum of Natural History, Chicago. 235 x 155 mm. This second part of the Ferns and Fern Allies of Guatemala lives up to the standard and expectations set in Part 1. Stolze deals with 69 genera of Polypodiaceae sensu /ato; they are arranged alphabetically. Two genera, Po/ypodium and Thelypteris, are divided into subgenera, the first into six subgenera (Campyloneurum, Microgramma, Niphidium, Phlebodium, Pleopeltis, and Polypodium). Thelypteris consists. of subgenera Amauropelta, Cyclosorus, Goniopteris, Macrothelypteris, Meniscium, Stegnogramma, and Steiropteris. Each entry consists of an ecological account, anda summary of its Guatemalan distribution followed by a general statement on its world distribution. The species is then described concisely without being pedantic, in a way which | find excellent for grasping the concept of a species that one has never seen. The entry is completed with notes on the history, taxonomy and nomenclature of the species in question. Frequently the author compares the species with close relatives, pointing out how it can be distinguished. The typography is clear and the book is again illustrated to the extent of 80 plates by five first-rate artists of which Richard Roesener stands out for his most unusual techniques. Bob Stolze saw fit to draw in two experts to help with this work. John Mickel (New York Botanical Garden) has contributed the account of Elaphoglossum (38 species) and Alan Smith (University of California at Berkeley), whose knowledge on New World Thelypteridaceae is extensive, has written up that family (59 species). There are good indented keys to all genera and to all species. This is a model fern flora and must be useful to anybody working on tropical American ferns. A.C. JERMY FERN GAZ 12(3) 1981 161 A CHROMOSOME COUNT FOR MACROGLOSSUM (MARATTIALES) T.G. WALKER Department of Plant Biology, University of Newcastle upon Tyne, England ABSTRACT A diploid chromosome count of 2n = 80 has been obtained for a plant of Macroglossum alidae from Sarawak. This is the first record for the genus and is based on x = 40, characteristic of the Marattiales. A diploid renresentative of Angiopteris evecta was also found with 2n = 80. INTRODUCTION Macroglossum is a small genus of Marattiales, consisting of a single, or at most two, species. It has a very limited geographical distribution, being recorded in the wild from Borneo and Sumatra under the name M™. alidae Copel. The problematical second species, M. smithi (Rac.) Campbell was erected by Raciborski (as Angiopteris smithii) on a smaller statured plant growing at the Botanic Garden at Buitenzorg (now Bogor) in Java. The status of this latter species in the wild appears to be somewhat uncertain. Whilst a member of the Royal Geographical Society’s Expedition to Gunung Mulu, Sarawak, in 19781 had the opportunity to examine a mature plant of MM. alidae growing near the Expedition’s base camp at Long Pala. Like Ang/opteris, which it most resembles, it had a massive rootstock somewhat more than 14m high and almost as much in width. The fronds were up to 3'4m long, inclusive of the thick stipe, and differed from those of Angiopteris in being simply pinnate. The pinnae were very broad — at least twice or up to three times the width of the pinnulesof Ang/opteris or Marattia and numbered up to 13 pairs per frond. A few of the fronds were fertile but all these, during July and August, bore only old, dehisced sporangia. The bases of the stipes had the thick fleshy stipules characteristic of the Order anda number of young plants were growing among the roots on the surface of the rhizome. Although their origin was difficult to determine with absolute certainty, it is believed that they were produced vegetatively. Despite a careful search being made no sporelings were found on the ground in the vicinity. ECOLOGY AND CULTIVATION This specimen of Macroglossum was growing in the permanently wet silt overlying limestone at an altitude of c. 60m above sea level in dense alluvial forest which casts a deep shade. Other herbaceous plants and shrubs were rather sparse in the vicinity of the fern. One of the offshoots was sent alive to Newcastle for further study. Here it thrives in one of the glasshouses under conditions favoured by Ang/opteris, putting on good growth in a potting mixture of coarse peat, loam and sand. Care has to be taken to avoid sudden drops in temperature and exposure to cold draughts which are anathema to many members of the Marattiales. CYTOLOGY It proved impossible to obtain a fixing in the field due to the lack of suitably young sporangia and the results reported here were obtained from the offshoot sent to Newcastle. This plant produced a large number of thick fleshy roots which were duly fixed in acetic alcohol. The chromosomes stained rather lightly with aceto-carmine (as did those of Angiopteris) and a typical cell is shown in Fig. 1a, together with its explanatory 162 FERN GAZETTE: VOLUME 12 PART 3 (1981) diagram (Fig. 2a). Several cellsshowed 2n = 80 unequivocally, indicating a diploid state on a basic chromosome number of x = 40. This base number is typical of other members of Marattiales viz. Ang/opteris (Manton and Sledge, 1954; Mehra and Singh, 1955; Ninan, 1956; Ghatak, 1962; Walker, press. comm. and unpub.), Danaea (Walker, 1966, 1973a), Christensenia (Braithwaite, 1977; Walker, 1979) and Marattia (Walker, in Manton, 1959; Walker, 1966 and unpub.). Other species of Marattia (Ninan, 1956; Brownlie, 1961) are based on x = 39, a number which Walker (1966, 1973b) considers to be secondarily derived from that of x = 40 found everywhere else in Marattiales from geographically widely separated areas. A sporeling of Angiopteris evecta (Forst.) Hoffm. was collected ina gully in mixed dipterocarp forest at c. 750m above sea level. In cultivation in Newcastle this yielded dividing root tip cells showing 2n = 80. The plant was thus diploid, in contrast to other specimens reported from Sri Lanka (Manton and Sledge, 1954) and India (Ninan, 1956; Ghatak, 1962) which were tetraploid. Like the chromosomes of Macroglossum those of Angiopteris did not stain deeply. A cell is shown in Fig. 1b together with an explanatory diagram (Fig. 2b). It is intended to carry out detailed karyotype analyses of members of this Order at a later date but suffice it to say that there are fairly obvious similarities in the distribution classes of chromosome lengths in the karyotypes of Macroglossum and Angiopteris. As the closeness of affinity between Macroglossum and Angiopteris is the subject of a current investigation at the British Museum (Natural History) where comparative anatomical and morphological studies are being made it is not proposed to pursue this matter further here, except to point out that Campbell (1914a, 1914b) noted that there are marked differences in certain features despite many overall similarities between the two genera. Figure 1. Root tip mitosis in (a) Macroglossum alidae, 713519; (b) Angiopteris evecta, T13401. Both x 1000. Permanent acetocarmine squash preparations. = 4 | anwar a vy 7> ota 47 SS os Fe = \Y ~ ant oS Z bs OV le5 Le Hag a4 Figure 2a & b. Explanatory diagrams to Figures 1a and 1b, respectively, x 1000. WALKER MACROGLOSSUM 163 ACKNOWLEDGEMENTS My thanks for financial assistance are due to the Research Fund of the University of Newcastle upon Tyne and to the Science Research Council. REFERENCES BRAITHWAITE, A.F., 1977. A chromosome count and range extension for Christensenia (Marattiaceae). Amer. Fern J. 67: 49-50. BROWNLIE, G., 1961. Additional chromosome numbers of New Zealand ferns. 7rans. Roy. Soc. New Zeal. Bot. 71: 1-4. CAMPBELL, D.H., 1914a. The genus Macroglossum Copeland. Philipp. J. Sci. C. Bot. 9:219-223. CAMPBELL, D.H., 1914b. The structure and affinities of Macroglossum alidae Copeland. Ann. Bot. 28: 651-669. GHATAK, J., 1962. Observations on the cytology and taxonomy of some ferns from India. Vuc/eus 5. 95-114. MANTON, |., 1959. Cytological information on the ferns of West Tropical Africa /n A.H.G. Alston, The flora of West Tropical Africa, 2nd ed.: 75-81. Crown Agents, London. MANTON, |.. & SLEDGE, W.A., 1954. Observations on the cytology and taxonomy of the pteridophyte flora of Ceylon. Phil. Trans. R. Soc. (B). 238: 127-185. MEHRA, P.N. & SINGH, H.P., 1955. Cytology of Cyatheaceae, Woodsiae and Marattiaceae. Curr. Sci. 24: 425. NINAN. C.A., 1956. Studies on cytology and evolution of the pteridophytes 1. Observations on the Marattiaceae. J. /ndian Bot. Soc. 35: 233-239. WALKER, T.G., 1966. A cytotaxonomic survey of the pteridophytes of Jamaica. 7rans. R. Soc. Edin. 66: 169-237. WALKER, T.G., 1973a. Additional cytotaxonomic notes on the pteridophytes of Jamaica. Jrans. R. Soc. Edin 69: 109-135. WALKER, T.G., 1973b. Evidence from cytology in the classification of ferns. /n A.C. Jermy, J.A. Crabbe and B.A. Thomas (Eds.). The phylogeny and classification of the ferns. Bot. J. Linn. Soc. 67, Suppl. 1: 91-110. WALKER, T.G., 1979. A further chromosome count for VAristensenia (Marattiales). Fern Gaz. 12: B1-52. 164 FERN GAZETTE: VOLUME 12 PART 3 (1981) REVIEW PALAEOBOTANY - AN INTRODUCTION TO FOSSIL PLANT BIOLOGY by Thomas N. Taylor xiii + 589 pp. McGraw-Hill Book Company, 1981. ISBN 0-07-062954-4. Price about £23 or US $29. The study of fossil plants is far too often thought of as an insular and over specialised field of botany by students of living plants. It can be a field that such people fear to enter because of the widely dispersed and seemingly over-detailed literature. Here, after a twenty year gap, is an up-to-date text book that should serve as an excellent introductory text for the serious reader. The book has a traditional layout in the sense that it has an introduction, dealing in general with fossils and their preservation, followed by chapters mainly devoted to major plant groups. About a quarter of the book deals with the three major groups of pteridophytes. The text is clear, the illustrations carefully chosen and each chapter has its own list of references. The text is however largely devoted to morphological studies with interspersed ideas of evolution. There is virtually no taxonomic information of value to those readers involved with this aspect of living pteridophytes and there is also no serious attempt to give ideas of palaeoecology. The book is clearly needed by all who are seriously interested in fossil plants. It is certainly not for the casual reader. B.A. THOMAS FERN GAZ. 12(3) 1981 165 AN ANALYSIS OF THE GRAMMITIS POEPPIGIANA-G.MAGELLANICA COMPLEX IN THE SOUTH ATLANTIC AND SOUTH INDIAN OCEANS B.S. PARRIS Botany School, Downing Street, Cambridge CB2 3EA, England ABSTRACT Grammitis poeppigiana (Mett.)Pic.Ser. and G. magel/lanica Desvaux subsp. magellanica are easily distinguished in part of their range (New Zealand Subantarctic islands and southern South America) but on Tristan da Cunha and Gough Island in the South Atlantic Ocean they appear more similar. However, they can be separated here and in all parts of their overlapping range by a combination of frond length and spore diameter. INTRODUCTION Grammitis poeppigiana (Mett.)Pic.Ser. (syn. G. kerguelensis Tardieu-Blot and G. armstrongii Tindale) is now regarded as a widely ranging circumantarctic species (Parris, 1975; Parris & Given, 1976) while G. mage//anica Desvaux subsp. magellanica (hereafter referred to as G. magellanica) has a more restricted distribution from the subantarctic islands of New Zealand through southern South America to Tristan da Cunha and Gough Island (Parris & Given, 1976). Fig. 1 shows more precisely the distribution of the two species. PACIFIC OCEAN INDIAN OCEAN FIGURE 1. Distribution of Grammitis poeppigiana (solid line) and G. magellanica subsp. magellanica (dotted line): 1. Macquarie |., 2. Auckland Is., 3. Campbell |., 4. Antipodes I|., 5. Falkland Is., 6. South Georgia, 7. Tristan da Cunha, Nightingale and Inaccessible Is., 8. Gough |., 9. Marion & Prince Edwards Is., 10. Crozet Is., 11. Kerguelen. I. 166 FERN GAZETTE: VOLUME 12 PART 3 (1981) There is no problem in distinguishing the two species in South America and New Zealand, where both occur, but on the islands of the South Atlantic and South Indian oceans discrimination on gross morphological appearance may be difficult, and judging from literature records and herbarium specimens, there has been much uncertainty as to which species occurs where throughout this region. Accordingly a variety of measurements were made on material of both species from New Zealand and South America so that those characters giving the best specific discrimination in areas where both species occur might be applied to the resolution of the status of material from the remainder of the species’ ranges. METHODS Wherever possible, ten fertile fronds from each collection were measured to provide the data on frond length and frond width; as the stipes of both species are indistinct and winged almost to base, frond length includes both stipe and lamina. For microscopic measurements of sporangia length and spore diameter, ten measurements of each character were made from each collection. Only mature sporangia, which had shed spores but which were closed, and spores which had not germinated were measured. Spores and sporangia were measured mounted in a glycerine solution. RESULTS It became apparant that the two characters of most reliable use in distinguishing G. poeppigiana and G. magellanica were frond length and spore diameter. Fig. 2 shows that with the exception of one aberrant New Zealand collection of G. magellanica a combination of these two characters gives absolute discrimination of the two species and this is also emphasised by the data in Table 1 which gives the range, mean and standard deviation of these characters for the South American and New Zealand populations of the two species. (CM) LENGTH FROND SPORE DIAMETER (/UM) FIGURE 2. Mean of frond length and spore diameter of G. poeppigiana and G. magellanica from New Zealand and South America: G. armstrongii New Zealand, @G. armstrongii South America, G. magellanica New Zealand, ©G. magellanica South America. PARRIS: GRAMMITIS 167 TABLE 1: RANGE, MEAN AND STANDARD DEVIATION OF FROND LENGTH AND SPORE DIAMETER IN G. POEPP/IGIANA AND G. MAGELLANICA: NEW ZEALAND, SOUTH AMERICA AND NEW ZEALAND/SOUTH AMERICA COMBINED DATA Frond length (cm) Spore diameter (uum) Range mean st.d. Range mean st.d. G. poeppigiana New Zealand 0.4-3.0 12 5 Our 31-65 46.4 eae ee) South America 0.6-2.9 1.6 OA ue) 35-58 46.5 Le ap combined 0.4-3.0 13 EP OED 31-65 46.5 t 66 G. magellanica New Zealand DIAZ 6.8 an D8 25-50 33.9 esa a | South America 83-4951 7.3 2 DA 25-50 35.0 = aA combined 2.9-429.4 7.1 Z 29 25-50 34.6 fe Aa Fig. 3 Superimposes on the overall values of the two characters of each species the appropriate mean measurements of all collections studied for the remainder of the species’ ranges. (CM) FROND LENGTH SPORE DIAMETER (UM) FIGURE 3. Mean of frond length and spore diameter of G. poeppigiana and G. magellanica from Atlantic and Indian Ocean superimposed on mean (+1 standard deviation) frond length and spore diameter of combined New Zealand and South American data for G poeppigiana and G. magellanica: © Falkland|s., v SouthGeorgia, @ TristandaCunha wv Goughls., # South Atrica, © Marion |s., OCrozetils ™® Kerguelen!., ® Australia, © Nightingale Is., ¥ Inaccessible |. 168 FERN GAZETTE: VOLUME 12 PART 3 (1981) This shows clearly that Falkland Islands, South Georgia, Marion Island and Kerguelen Island specimens are virtually identical with South American and New Zealand G. poeppigiana. Material from Inaccessible Island, Crozet Islands, Australia and South Africa, although tending to have slightly larger fronds, can only be regarded as G. poeppigiana. Tristan da Cunha specimens demonstrate convincingly that both species are present on this island and, apart from an anomalous intermediate specimen which cannot be assigned reliably the same would seem to be true of the relatively nearby Gough Island. Thus, particularly when viewed in the context of the variability of the two species in their chief ‘mainland’ areas of South America and New Zealand, identification of South Indian and South Atlantic Ocean specimens can be made with considerable confidence. The important distinctions between the species are summarised below: G. poeppigiana - rhizome usually long-creeping, sometimes short-creeping or + ascending, fronds usually less than 4cm long, sori less than 5 (-7) pairs, spores usually larger than 40 ym diam. G. magellanica - rhizome erect, rarely ascending, fronds usually more than 4cm long, sori normally more than 5 pairs, spores typically smaller than 40 yim diam. DISCUSSION The distribution of these two species fits very well the scheme for zonation of terrestrial vegetation in southern latitudes proposed by Wace (1960, 1965). His cool temperate zone extends south to the limits of tree or woody shrub growth and in this area (including southern South America, Tierra del Fuego, Tristan da Cunha group, New Zealand Subantarctic Islands) both the epiphytic G. mage//anica and the rupestral G. poeppigiana are found. Only the latter has so far been recorded from the Falkland Islands which are also placed in this zone. Wace’s subantarctic zone is defined by the absence of tree or woody shrub growth and the presence of closed herbfield vegetation. Only G. poeppigiana occurs on the islands included in this zone (South Georgia, Marion and Prince Edward Islands, Crozet Islands, Kerguelen Island, Macquarie Island). ACKNOWLEDGEMENTS | wish to thank the Keepers of the following herbaria for the loan of material and/or permission to examine collections: Arnold Arboretum herbaria, Harvard; Auckland Institute and Museum; Botany School, Cambridge; British Antarctic Survey, Cambridge; British Museum (Nat. Hist.), London; Botany Division, D.S.LR., Christchurch; Royal Botanic Garden, Edinburgh; Royal Botanic Gardens, Kew; Botanical Museum, Oslo; Museum d'Histoire Naturelle, Paris; Bolus Herbarium, University of Cape Town; National Museum, Wellington. REFERENCES PARRIS, B.S., 1975. A revision of the genus Grammitis Sw. (Filicales: Grammitidaceae) in Australia. Bot. Jour. Linn. Soc. 70: 21-43. PARRIS, B.S., & GIVEN, D.R., 1976. A taxonomic revision of the genus Grammitis Sw. (Grammitidaceae: Filicales) in New Zealand. N.Z. Jour. Bot. 14: 85-111. TARDIEU-BLOT, M.L., 1962. Sur les Grammitis des |les australes. Adansonia 2: 111-116. WACE, N.M., 1960. The botany of the southern oceanic islands. Proc. Roy. Soc. London B 152: 475-490. WACE, N.M.,1965. Vascular plants, pp. 201-266 in van Miegheim, J. and van Oye, P. (Eds.) Biogeography and Ecology in Antarctica. W. Junk, The Hague. FERN GAZ. 12(3) 1981 169 TWO NEW SPECIES OF SELAGINELLA FROM THE PHILLIPINES BENITO C. TAN Department of Botany, University of the Philippines at Los Banos College, Laguna Province, Philippines 3720 and A. CLIVE JERMY British Museum (Natural History), Cromwell Road, London SW7 5BD, England ABSTRACT Two new species of Se/agine/la, discovered by the senior author when undertaking a review of cone and spore morphology of the genus in the Philippines, are described and named as follows: S. atimonanenis Tan & Jermy from Luzon Island, and S. pricei Tan & Jermy from Samar Island. INTRODUCTION In the course of studying the sporangial distribution pattern and spore morphology of Philippine Se/agine//a species for a Masters thesis (Tan 1974) in botany, the senior author (B.C.T.) came across two new species of the genus, materials of which were collected from limestone areas, a habitat usually sparse in Se/aginella. The Philippine species of Se/agine//a were first summarized by Alston in 1 935. In that paper, a total of 46 species were listed with localities and a key tothe species was also included. Later, Alston (1937) made several corrections to nomenclature when he reduced S. ascendens v. Anderw.v.Ros. to synonymy of S. /ntermedia (BI.) Spring, S. peltata C. Presl to S. invol/vens (Sw.) Spring and S. springiana v. Anderw.v.Ros. to S. longiaristata Hieron. Subsequently, S. heterostachys Baker and S. uncinata (Desv.) Spring were reported by Tan (1975) as new to the country. The latter species, which is of Chinese origin, has been observed to become naturalized in Laguna Province. In addition, S. plana (Desv.) Hieron. from the West Indies, and S. wi//denovii Baker, an eastern Asian-Malasian species, have also been reported as ornamentals in many local gardens and greenhouses (Price 1974). KEY AND DESCRIPTIONS The two presently new species of Se/agine//a from the Philippines described below are related to the S. /ntertexta group which may be distinguished by the following key: 1. Lateral leaves broadly ovate to orbicular, less eI S AS AONE AS ORG 2.6 2 anno esis eecna ate ib lo pe iar oh atm Slog wm Gas oS Shes wee Z 2. Lateral leaves = contiguous and opposite; Blan Sh arsely Ane ADEM Y VANCMeG so flees 868. oy on Se. Sat ae ies 2 oe S. pricei 2. Lateral leaves spaced, not opposite; plant often much branched and forming intertwining mats ................200005: 3 3. Lateral leaves symmetrical, margins EACH RANA SE OE ase Fa 5 ene cas 5 Mand Ps oo Moca A os mvs Cw iene S. nummularia 3. Lateral leaves asymmetrical, margins PEAR eA eee od 5 Pa Ak 2 ale icn wc PERE emg Pasion © wes 'e S. atimonanensis 1. Lateral leaves broadly elliptic to narrowly ovate, Rt RE tris RIC ret FOGG ity vine ait nosis! eee = ate BinewiaihunSe es it's Ries wdiniele, © oo 8 edge Race a 4. Marginal cilia of all leaves long; median DEE TEE AT SE FO ne eg ER oo ne aaa a eae ee S. apoensis 4. Marginal cilia absent or, if present, inconspicuous and short; AE Eye TUNA TID), gexec ania. tes ag ve.iase = pnhdsn. 8 ainin es aim wipe tia aw we mip, S. intertexta 170 FERN GAZETTE: VOLUME 12 PART 3 (1981) Selaginella antimonanensis Tan & Jermy sp. nov. (Figs. 1-6) Caulis primus repens, inaequaliter ramosus, tegetes laxas formans, interdum radicans, rhizophoro filiformi instructus. Fo/ia omnino dimorpha; folia /ateralia distantia, 1-1.5 x 0.8- 1.3mm, late ovata vel plus minusve orbicularia, saepe asymmetrica, marginibus posterioribus ellipticis longe ciliatis, anterioribus autem hemisphaericis breviter ciliatis, apice acuto vel breviter cuspidato mucronatove, basi rotunda; fo/a axi//aria ovata vel elliptica, lateralibus dimidio minora, breviter cuspidata, marginibus ubique ciliatis; fo/ia intermedia circa 0.25-0.75mm longa, 0.2-0.5mm lata, ovata, ea in caulibus primis asymmetrica, ea in ramis plus minusve symmetrica, apice 0.25mm longo, acuminato vel aristato, basi cordata, marginibus ubique ciliatis. Strobili complanati, plerumque longi minus quam 1.0cm. Sporophy/la dimorpha, ciliata, ea dorsalia ovata vel lanceolata, dimidio minimum minora quam folia lateralia, apice acuto-acuminato, ea ventralia lanceolata, apice acuminato. Megasporae in siccitate albidae vel flavidae, c. 200 jim diametro, cerebriformes, sporodermate minute papillato; microsporae aurantiacae, 30-34 jim diametro, verrucatae, verrucis tenuiter spinulosis. Typus: on limestone cliff along trail to summit, 300-400 m alt., Atimonan National Park, Quezon Province, Luzon, B.C. Tan 77-2005 (holotype, CAHP; isotypes, BM, KYO, L, US). Paratypes: in limestone crevices, near summit, 428 m alt., same locality, B.C. Tan 77-2006; on moist limestone wall, near Minador Cave, Atimonan National Park, c. 250 m, B.C. Tan 77-500; 77-501. A creeping, irregularly branched plant forming a loose mat, stems (including the lateral leaves) 2-3mm wide, rooting at intervals, rhizophores filiform. Leaves dimorphic throughout; lateral leaves 1.0-1.5 x 0.8-1.3mm, distant below, becoming more crowded towards the apex and strobili, superficially broadly ovate to almost orbicular, but asymmetric with the posterior margin elliptic and the anterior t hemispherical, apex acute to shortly cuspidate or mucronate, margins ciliate throughout, cilia O0.2-0.4mm and longer on the anterior margin, marginal cells papillate and conspicuous on the underside; axillary leaves ovate to elliptic approximately equal in size to the lateral leaves of the branches they subtend, apex shortly cuspidate, margin ciliate; median leaves 0.25-0.75 x 0.2-0.5mm, ovate, with long acuminate to aristate apices, those on the primary stem + asymmetrical, those on the branches symmetrical, bases cordate to auriculate, margins ciliate. Strobili complanate with 1-3 megasporangia near the base or occasionally almost entirely megasporangiate with 1-3 microsporangia near the apex; sporophylls of two sizes, those larger ones, dorsal and sterile, margins ciliate, those ventral and median, narrower, usually all containing sporangia. Megasporesc. 200 um, sporoderm minutely papillate folded into an irregular brain-like appearance, white or yellowish when dry; microspores 30-34 ym in diameter, orange, verrucate, verrucae finely spinulose. Among the local species, the new taxon is closely related to S. intertexta Spring and S. nummularia Warb. It differs from the former in being shortly creeping and having strongly ciliated vegetative leaves and sporophylls, and also in the almost suborbicular lateral leaf shape. From the latter, which it resembles closely in habit and leaf shape, it can be easily recognised by the presence of cilia on the lateral leaves which are asymmetrical. The broadly ovate to near orbicular leaf outline separates it also from the other ciliate species in the group, S. apoensi/s Hieron. Selaginella pricei Tan & Jermy sp. nov. (Figs. 7-10) Caulis primus repens, longus, tenuis, brevibus ramis distantibus, interdum radicans, rhizophoro filiformi instructus. Folia /ateralia 1.2-1.6mm x 0.8-1.2mm in caule ramisque conferta, late elliptica, basi truncato-cordata, ad apicem breviter acutum vel acuminatum contracta, margine serrato, ciliis 0.1-O.3mm longis basi parce sparsis, secondum autem margines superiores frequentioribus et conspicuis; fo/ia axi//aria ovata vel elliptico-lanceolata, quam lateralia plus minusve tertia parte minora, non auriculata, apice acuto vel acuminato, in marginibus ambobus et basem versus ciliata; fo/ia intermedia eis lateralibus plus minusve triplo minora, ovato-lanceolata, aristata, basi leniter cordata, asymmetrica, exterioribus lobis evidentioribus, basi ciliata et manifeste secundum margines interiores integros. Strobili complanati, 1-2cm longi, plus minusve 1.5mm lati. Sporophy//a dimorpha, ciliata, ea dorsalia late lanceolata plus minusve 1.5mm longa, gradatim acuminata, non carinata, ea ventralia minora, lanceolata, acuminata, carinata. Megasporae non visae; microsporae aurantiacae 28-35 um diametro. TAN & JERMY: TWO NEW SPECIES OF SELAGINELLA 171 FIGURES 1-6. Selaginella atimonanensis Tan & Jermy : 1. portion of main branch from above, x 50. 2. median leaf, from above, x 100. 3. branchlet from below, x 20. 4.detail microspore wall, x 5500. 5. megaspore distal face, x 300. 6. detail of megaspore wall, x 2200. a = axillary, | = lateral, m = median leaves. 172 FERN GAZETTE: VOLUME 12 PART 3 (1981) FIGURES 7-10. Se/aginella price; Tan & Jermy : 7. main branch from above, x 40. 8. branchlet from below, x 20. 9. cilia on axillary leaf, x 300. 10. upper surface of median leaf. a = axillary, | = lateral, m = median leaves. TAN & JERMY: TWO NEW SPECIES OF SELAGINELLA 173 Typus: on limestone cliff, 200 m alt., in forested trail to Concord municipality from Bagacay, Hinabangan, Western Samar, 28 June 1975, B.C. Tan 75-414 (CAHP; isotypes BM, KYO, L, US). Paratype: creeping on limestone cliff inside primary forest, municipalities of Gandara and Matuguinao, Western Samar. Price & Hernaez 119 (CAHP). A species with a long slender creeping main stem with distantly spaced short branches, rooting at intervals with filiform rhizophores. Lateral leaves c. 2 x 1mm, base truncate-cordate, broadly elliptic, those on the main branches becoming ovate-elliptic, abruptly acuminate at apex, margins sparsely serrate, ciliate around the base especially on the anterior margins; axillary leaves ovate to elliptio-lanceolate, c. 74 the size of the lateral leaves, not auriculate, apices acute to acuminate serrate above, ciliate below; median leaves 0.8-1.2 x 0.3mm lanceolate-ovate, aristate, weakly cordate at the asymmetric base, the outer lobes being larger, cilia sparse along inner margin. Strobilus flattened, 1-2cm long, c. 1.5mm broad; sporophylls dimorphic, those of the upper plane broadly lanceolate, + 1.5mm long, acuminate, not keeled, ciliate throughout, those of the lower plane smaller in dimension, similar but keeled. Megaspores not seen; microspores 28-35 um, orange. ‘The species is named after Michael G. Price of UP at College, whose studies on Philippine ferns warrant special recognition. Superficially, the taxon looks like a large form of S. apoensis Hieron. or a small form of S. boninensis Baker. The leaf sizes and the broader lateral leaves, however, distinguish it from S. apoensis. The fewer and shorter cilia (0.1-O.3mm) along the leaf margins and bases, plus the laxer and more forwardly directed sporophylls of the upper plane are additional distinctive characters for the newtaxon. From S. /ntertexta Spring it can be segregated by the aristate median leaves, and the long, creeping stem habit with only a few short branches. ACKNOWLEDGEMENTS We should like to thank Miss Alison Paul for taking the SEM micrographs and Miss Kathryn Kavanagh for help with the Latin descriptions. REFERENCES ALSTON, A.H.G., 1935. The Philippine species of Se/aginel//a. Philip. J. Sci. 58: 359-383. ALSTON, A.H.G., 1937. The Selaginellae of the Malay Islands Il. Sumatra. Bull. Jard. Bot. Buitenzorg, ser. 3, 74: 175-186. PRICE, M.G., 1974 Pteridophytes of Mount Makiling and vicinity. M.Sc. Thesis Univ. Philippines, Los Banos (unpublished). TAN, B.C., 1974. Sporangial distribution and sporal morphology of Philippine Selaginella. M.Sc. Thesis Univ. Philippines, Los Bafos (unpublished). TAN, B.C., 1975. Two Selagine/la species new to the Philippines. Ka/ikasan, 4: 42-46. 174 ~ FERN GAZETTE: VOLUME 12 PART 3 (1981) REVIEW THE EVOLUTION OF PLANTS AND FLOWERS by Barry Thomas. 1981, 220 x 28Omm, 116 pp. Peter Lowe Ltd., London. Price: £5.95. This is a Semi-popular book about the evolution of the plant kingdom. It presents not so much the theoretical side of evolution but a synopsis of the main stages of plant life on earth as known from fossil evidence. It is profusely illustrated throughout with coloured diagrams, restorations of the appearance of the fossils as whole plants, and photographs of living members of each group. As one might expect, pteridophytes, together with conifers, are well represented, the two groups having dominated the land flora of this planet for solonga period of evolutionary time. The book gives a synopsis of the main pteridophytes present from the late Devonian period through to the Mesozoic in particular, setting them against a background of the kind of vegetation and landscape in which they grew. The text is written in a readable and simple style which should be easy to understand by anyone without previous knowledge of the subject. An important feature of the book is certainly its illustrations, which are well-chosen, and in most cases, very well done. They greatly help in bringing the subject to life. Particularly informative are the diagrams of life-cycles, as well as the small charts showing the interrelations of the groups discussed and the period of time for which they survived. So often, fossil plants leave the impression of dry and dusty things. | found this book a valuable stimulant in setting them in the dynamic perspective of green and living things. It is a good book for the beginner, and will probably leave him wanting to delve further. C.N. PAGE FERN GAZ. 12(3) 1981 ia i) A FORKED VEIN AND FOLIAR FIBRES IN SELAGINELLA R.N. MUKHERJEE and U. SEN Department of Botany, Kalyani University, West Bengal, India. ABSTRACT The presence and significance of a forked vein and foliar fibres is described in Selaginella. INTRODUCTION The leaves of Se/agine/la are small, scaly and provided with a single unbranched nerve. As the leaf trace is small and unbranched, its departure has no influence inthe stelar cylinder, and there is no formation of a leaf gap. A survey through literature, however, reveals the existence of more than one nerve in some species of the genus. Alston (1945) noted a tri-nerved condition in Se/aginella griffithii Spring. Panigrahi and Dixit (1967) described bi-nerved leaves in S. invo/vens (Sw.) Spring. Grambast and Rosello (1965) found forked veins in S. wi//denowi (Desv.) Baker. The presence of more than one nerve was also reported by Webster (1970) in aberrant leaves on excised stem of S. martensii Spring grown in culture. These leaves differed from normal ones in having lobed tips, more stomata and sometimes more than one ligule associated with each leaf. OBSERVATIONS When making a systematic survey of the anatomical features of the genus, we could not confirm the occurrence of more than one vein in the normal leaves of S. willdenowi, S. griffithii and S. involvens. |n the latter two species we found, however, a sclerenchymatous band composed of fibres with warty outgrowths on the outer periclinal surface extending along either side of the midrib on the abaxial epidermis of the lateral leaves (figs. 1a and 1f). Tracheids and sieve cells are never associated with these sclerotic bands. These two sclerotic bands together with the midrib give a deceptive tri-nerved appearance in dried leaves of S. griffithi.In S. invo/vens the midrib iS inconspicuous and as such the two median sclerotic bands appear as two nerves of the leaves. The occurrence of similar false veins was reported in S. cau/escens Spring from Java by Baker and Posthumus (1939). Median sclerotic bands also occur in the abaxial epidermis of the lateral leaves of S. intermedia (Bl.) Spring, S. chrysocaulos (Hook. and Grev.) Spring and S. tenera (Hook. and Grev.) Spring (figs. 1b-d). The foliar fibres of S. intermedia are so thick-walled that their lumina are almost obliterated. In S. tenera small patches of fibres form almost a continuous strip extending more than half the entire length of the lamina on the acroscopic side, but in the basiscopic half only a few discrete patches occur (fig. 2). Sclerotic fibres occur regularly at the margins of the lamina of the median leaves of S. he/feri Warb., S. intermedia, S. ciliaris (Retz.) Spring, S. chrysorrhizos Spring, S. denticulata (Desv.) Alston and S. vaginata Spring. Similar occurrences of fibres at the margins of the leaves was also reported by Mital (1965) in many species of Se/aginel/la from India. S. adunca A.Br. ex Hieron is the only species we have come across in which the vein is really forked (fig. 1e). The leaf trace after entering the vegetative and reproductive leaves undergoes two successive dichotomies within a short distance. 176 FERN GAZETTE: VOLUME 12 PART 3 (1981) Lateral leaves of Selaginella. a, S. griffithii x 50; b, S. tenera, x 33; c, S. interdemia, x 33; d, S. chrysocaulos, x 33; e, S. adunca, x 33;, f, foliar fibres in the abaxial epidermis of S. chrysocaulos, x 450 (ec = epidermal cell, ff = foliar fibre, sb = sclerotic band, sp = sclerotic patch, v = vein). MUKHERJEE & SEN. SELAGINELLA 177 Yet the trace creates no impact in the vascular cylinder of the stem and no incipient gap can be recognised just above the departing trace. The occurrence of a branched vein in a relatively primitive species like S. adunca appears significant in connection with Zimmermann’s (1959) hypothesis of origin of microphyllous leaf through reduction REFERENCES ALSTON, A.H.G., 1945. An enumeration of Indian species of Se/aginella. Proc. Nat. Inst. Sci. India, B(11): 211-235. BACKER, C.A., and POSTHUMUS, O., 1939. Varenflora voor Java. Java. GRAMBAST, L., and ROSELLO, S., 1965. Les notions de microphylle et de mégaphylle et leur signification phylogénétique. Compt.Rend Acad. Sci., Paris, 261: 5183-5186. MITAL, P.L., 1969. Epidermal studies in the genus Se/agine//a Beauv. Bull. Bot.Surv.India, 11: 150-160. PANIGRAHI, G., and DIXIT, R.D., 1967. Studies in the systematics of Indian Se/aginella — II. J. Indian Bot.Soc., 36: 102-108. WEBSTER, T.R., 1970. Aberrant leaves on angle-shoots of Se/aginella martensii Spring. Amer. Fern J., 60. 1-6. ZIMMERMAN, W., 1959. Die Phylogenie der Pflanzen. \\. Aufl. Stuttgart. 178 FERN GAZETTE: VOLUME 12 PART 3 (1981) SHORT NOTES DRYOPTERIS X SARVELAE IN SCOTLAND; A NEW HYBRID FOR THE BRITISH ISLES Dryopteris x sarvelae Fraser-Jenkins and Jermy is the hybrid D. carthusiana x expansa. Until now, the only record for this hybrid was from Espoo, Finland, where it was found by J. Sarvela (Widén, Sarvela and Ahti, 1967). In 1978 the hybrid was discovered in Argyll, Scotland (V.C. 101) by H. V. Corley, and its identification was confirmed on cytological and morphological evidence by M. Gibby in 1980. The hybrid is easily confused with D. x deweveri (Jansen) Jansen and Wachter and D. x ambroseae Fraser-Jenkins and Jermy, but it can be recognised by examination of the chromosomes during meiosis. D x sarvel/ae is triploid (2n = 123); during the first metaphase of meiosis most of the chromosomes fail to pair (Gibby and Walker, 1977). This distinguishes it from D. x ambroseae, which is also triploid, as equal numbers of bivalents and univalents are seen in meiosis in this hybrid. D. x dewever is tetraploid (2n = 164). D. x sarvelae could form wherever the two parents grow together, and it may well occur at other sites in Britain but has previously been overlooked. REFERENCES GIBBY, M., & WALKER, S., 1977. Further cytogenetic studies and a reappraisal of the diploid ancestry in the Dryopteris carthusiana complex. Fern Gazette 17: 315-324. WIDEN, C.-J., SARVELA, J., & AHTI, T., 1967. The Dryopteris spinulosa complex in Finland. Acta Botanica Fennica 77: 1-24. H.V. CORLEY, Pucketty Farm, Faringdon, Oxfordshire and M. GIBBY. British Museum (Natural History), London. A NEW NAME FOR A HYBRID HORSETAIL IN SCOTLAND The wild hybrid between the widespread horsetails £. fluviati/e L. and E. pa/ustre L. was first tentatively reported by the author from the island of Harris, Outer Hebrides, where it had been found in July 1962 (Page 1963). Further observations and finds have supported the original diagnosis of the parentage. J.G. Duckett (see Duckett & Page, 1975: 100-101) has subsequently also ably shown that this cross can, indeed, be achieved in synthetic culture, using &. fluviatile as the female parent, and independentiy has also found plants attributable to this parentage wild in Scotland (Duckett 1979). This hybrid may well yet prove more widespread than hitherto recorded. For convenience of reference, a binominal name for it is thus proposed. Equisetum x dyce: C.N. Page hybr. nov. (E. fluviatile L. x palustre L.). Ic.: C.N. Page, Brit. Fern Gaz. 9 (4): 118 (1963). Hybrida inter £&. fluviatile et E. palustre ramificationis habitu, vaginis, numero et profunditate vallecularum carinarumque, et ratione cavi centralis diametro cauli intermedia. Caules 15-45cm alti, virides, annul; internodii endodermididibus individuis provisi. Internodii rhizomatis ut in E. fluviatile excavati; tubera rhizomatis ut in £. pa/ustre adsunt. Strobili parvi, tenues, nigrescentes, in caulibus vegetativis terminales. Sporae abortivae. Typus. C.N. Page No. 873: Loch-a-Mhorghain, Harris, Outer Hebrides, July 1962 (E). SHORT NOTES 172 Hybrid intermediate between &. fluviatife and £. palustre in branching habit, sheaths, number and depth of ridges and furrows and in the ratio of the central hollow to the diameter of the stem. Stems 15-45cm high, green, annual; stem internodes with individual endodermises. Rhizome internodes hollow as in &. fluwatile; rhizome tubers present as in £. pa/ustre. Cones small, slender, brackish, terminal on vegetative shoots. Spores abortive. Plants differ clearly from both parents in their strong intermediacy of structure, both in overall appearance and in proportions of the internai cavities. They resemble E. fluviatile in the hollow rhizomes and individual stem endodermises (both features unique in subgenus Fquisetum to E. fluviatile and its hybrids), and resemble E. palustre in the presence of tubers and dark, slender cones. The much weaker habit of growth, fewer, more ascending branches, and black cones which are small, slender, not barrel-shaped, and much more numerous, help distinguish it from E. x /itorale (E. arvense x fluviatile). Plants are currently known from a small number of scattered locations in Scotland. The plant is named in honour of J.W. Dyce, who has done much, over many years to stimulate pteridophyte recording work in Britain. ACKNOWLEDGEMENTS | am grateful to Dr. Robert Mill for the latinisation of the description. REFERENCES DUCKETT J.C., 1979. An experimental study of the reproductive biology and hybridisation in the European and North American species of Equisetum. Bot. J. Linn. Soc. 79: 205-229. DUCKETT, J.G., & PAGE, C.N., 1975. Equisetum pp 99-103, in Stace, C.A. (Ed.) Hybridisation in the British Flora. Academic Press, London. PAGE, C.N., 1963. A hybrid horsetail from the Hebrides. Brit. Fern Gaz. 9: 117-119. C.N. PAGE, Royal Botanic Garden, Edinburgh. EQUISETUM X TRACHYODON IN SKYE, WESTERN SCOTLAND In November 1974, an odd-looking Equisetum was found in Skye, Inverness-shire, Scotland. It was growing on a lump of bank that had slipped into the Hinnisdal river, on the section of the river half-a-mile west of the bridge carrying the road between Portree and Uig, in the north end of Skye (in square 18/35). It was determined as E. x trachyodon (E. hyemale x variegatum) by Dr. C.N. Page, at RBG Edinburgh. At the time, the nearest known £. hyemale was in Geary (18/26), across Loch Snizort, or in Glen Brittle (18/41), and the only £. variegatum record was that shown in the ‘Atlas’ (18/36), for which Monkswood had no details of locality. Succeeding winters have added some interesting facts, as we kept an eye open for Equisetum specimens which are evergreen. £. hyemale was in Trotternish, at two sites on opposite sides of the Storr ridge (two places on a river bank in 18/45 and one where a seepage area dripped over a sea-cliff in 18/55). Drs. C.N. Page and R.E.C. Ferreira visited Skye in July 1979, found several pieces of EF. x trachyodon upstream from the original site, and suggested that there could be yet more of it further up the glen. The river banks just east of the road bridge had been investigated soon after the 1974 discovery, but in November 1979, a search starting below Peinha (11/, miles east of road bridge) turned up dozens of little plants, where the banks were either sandy or gravelly, along a 1'/% mile stretch eastwards. On a later 180 FERN GAZETTE: VOLUME 12 PART 3 (1981) visit, above the join of the streams below Glenuachdarach, nothing was found. Meanwhile, inthe summer of 1978, Mr. and Mrs. J. Bowman had found the first localised record of £. variegatum on Skye — from a corrie below the SE end of the Cuillin ridge (18/41) — so both parents of the hybrid are on the island. The winter of 1979-80 was spent walking sections of various rivers, in Trotternish and elsewhere, in search of further Equisetum x trachyodon, or E. hyemale, but without success. The question now is: why is the £. x trachyodon confined to the Hinnisdal valley, and how did it get there in the first place? And the old record for E. variegatum in 18/36 (the neighbouring square) may well be our plants, since a similar record (same recorders) for EF. variegatum at Nisibost in Harris (V.C. 110) has been re-determined by Dr. Page as E£. x trachyodon. E. x trachyodon has also been added recently to the list for Rum. C.W. MURRAY, Prabost, Isle of Skye. MORE NEW COMBINATIONS IN SOUTHEAST ASIAN FERNS Preparation of the Flora Malesiana account of Grammitidaceae and other papers on Southeast Asian ferns necessitates publication of the following new combinations: Ctenopteris brooksiae (v.A.v.R.) Parris, comb.nov. Basionym: Polypodium brooksiae v.A.v.R. in Bull. Jard.bot.Buitenzorg Ser.2, 23: 19, t.2, f.2 (1916). Grammitis intromissa (Christ) Parris, comb.nov. Basionym: Polypodium intromissum Christ in Verhand/ungen der Naturforschenden Gesellschaft in Basel 11: 440 (1896). Grammitis plana (v.A.v.R.) Parris, comb.nov. Basionym: Polypodium planum v.A.v.R. in Bull. Jard.bot.Buitenzorg Ser. 2, 16: 32 (1914). Grammitis subdichotoma (Racib.ex v.A.v.R.) Parris, comb.nov. Basionym: Polypodium subdichotomum Racib.ex v.A.v.R. in Bull. Dept.Agric.Ind.Néerl. 18: 20 (1908). Grammitis torricelliana (Brause) Parris, comb.nov. Basionym: Polypodium torricellianum Brause in Bot.Jahrb. 49: 45, f.3A (1912). Pteridium caudatum subsp. yarrabense (Domin) Parris, comb.nov. Basionym: Pteridium aquilinum var. yarrabense Domin in Bib/. Bot. 85: 161, f.32 (1916). B.S. PARRIS, Botany School, Downing Street, Cambridge CB2 3EA, England. THE OCCURRENCE OF A LAMINAL FLAP IN SELAGINELLA The occurrence of a laminal flap, developed as a vertical projection on the adaxial surface of sporophylls or bracts, has been found in Se/aginella bisulcata Spring, S. reticulata (Hook. and Grev.) Spring, S. tenera (Hook. and Grev.) Spring and S. subdiaphana (Wall.) Spring. In each of these species, the laminal flap is isobilateral and is situated next to the ligule. While the ligule is a distinct structure and is embedded within the leaf tissue only at the basal region, the flap is a continuation of the lamina and extends from the base towards the distal region of the bract or the sporophyll along the midrib. The epidermal cells of the flap are elongate and resemble those of the adaxial surface of the SHORT NOTES 181 lamina proper. Stomata occur on both the surfaces of the flap, and are structurally similar to those on the surfaces of the sporophyll and the bract. Round cells with walls of tracheidal thickening often occur at the base of the ligule, but no tracheid enters the flap. The margin of the laminar flap in S. subdiaphana is markedly distinct being long ciliate, but in the other three species it resembles the margin of the sporophyll or the bract. FIGURE 1: Se/aginella reticulata, a bract with laminal flap, x 35; b, S. subdiaphana, a large sporophyll with laminal flap (megasporangium removed), x 70. In S. reticulata the laminate flap only develops on the bracts associated with the sporophylls (fig. 1.) In S. tenera and S. subdiaphana only the larger sporophylls bear the flap (fig. 1b.). ; In S. subdiaphana as well as in S. reticulata, the foliar vein extends beyond the termination of the flap, while in St. tenera and S. bisulcata the flap extends beyond the vein ending. The flap occurs on the smaller sporophylls and larger bracts. of S. bisulcata. No laminal flap has yet been found on the vegetative leaves of any species of Selaginella. The laminal flap probably increases the photosynthetic surface of the sporophyll and the bract since chloroplasts occur within their cells. But it seems significant that it is always associated with the sporophylls or the bracts protecting the sporophylls. An alternative interpretation of its function might be that the flap with its cilia might be a contrivance for capturing spores micro- and megaspore can germinate and develop in close proximity; but clearly further observations on this are needed. It is possible that this flap is homologous with the ensheathing velum and mass of tentacles surrounding the macrosporangium of the carboniferous genus Miadesmia, since the velum had its origin from a region between the ligule and the axil of the sporophyll. As the three living species in which the flap has been found are members of the subgenus Heterostachys it would be interesting to see whether the occurrence of similar laminal flaps are a more extensive feature of this subgenus. R. MUKHOPADHYAY & U. SEN, Department of Botany, Kalyani University, Kalyani, West Bengal, 741235, India. 182 FERN GAZETTE: VOLUME 12 PART 3 (1981) A NEW SPECIES OF PLAGIOGYRIA FROM MALAYA During the examination of herbarium sheets of the genus Plagiogyria (Kunze) Mett. present in the Central National Herbarium, Howrah (CAL), the authors have come across a new species of the genus, which is described below. The specimens had been identified earlier as P. euph/eb/a but the new species is distinct from it in the following characters: 1. Hook-like aerophores are present throughout the stipe and on the rachis at the base of insertion of the lateral pinnae 2. The lamina ts leathery in texture oS: The pinna margins are strongly reflexed, the apex acute and undulate, the lower surface pale-brown, and the upper surface dark brown in colour. The present species can be readily separated from P. tuberculata Copel. by the presence of terminal leaflet simple like other lateral pinnae and not pinnatifid as in P. tuberculata Copel. Plagiogyria malayensis R.D. Dixit & Anjali Das sp. nov. Fig. 1. Aerophora unciformia prominentia praesentia in stipitibus et ad bases pinnarum, apicalibus paucis exceptis. Pinnae terminalis lateralibus similes, pinnae in 15-20 paribus, erecto-patentes, oblongolanceolatae, 12-14 x 1.8-2cm, basi cuneatae, apica acutae, undulatae, ad marginem integrae, tenues, reflexae, coriaceae, Supra atro-brunneae, et infra pallido-brunneae. HERBARIUM MUS. PERAK Rosi? pe AAR ARE Sohn es Bah rect i POP Game ey FOE abt, * Bates Ci llees. te WHAY fe. FIGURE 1. Holotype of Plagiogyria malayensis R.D. Dixit & Anjali Das sp.nov. — SHORT NOTES 183 Stipes of sterile fronds 20-30cm, dark reddish-brown, and rachis pale-brown in colour. Aerophores prominent, hook like, present throughout stipe and on rachis at the point of insertion of the lateral pinnae. Lamina 40-55 x 15-20cm pinnate, terminal leaflet simple like other lateral pinnae; lateral pinnae 15-20 pairs, alternate, 1.5-2.5cm apart, erecto-patent, shortly stalked except a few adnate pinnae in the apical region, leathery in texture, upper surface dark brown and lower surface pale brown, middle pinnae 12-14 x 1.8-2cm, oblong-lanceolate, cuneate at base, apex acute, undulate, margins thinned, entire, and strongly reflexed: midrib raised on the lower surface, obscure on the upper surface; veins free, immersed, usually simple, sometimes furcate. Fertile pinnae present in paratype specimen, 10-14cm x 2-3mm, coiled, ascending. Holotype: MALAYA — Gunong Batu Paleb, 6700ft., Wray Jr. 3177 (CAL). Paratype: MALAYA — without exact location Scortechini 397 (CAL). ACKNOWLEDGEMENTS Grateful thanks are due to Dr. S.K. Jain, Director, Botanical Survey or India for the encouragement. Our thanks are also due to Dr. N.C. Majumdar, Central National Herbarium, Howrah for the Latin translation. R.D. DIXIT & ANJALI DAS Botanical Survey of India, Howrah-711103, India. Present address: Botanical Survey of India, Allahabad. NOMENCLATURAL NOTES ON DRYOPTERIS — 5 C.R. FRASER-JENKINS c/o Botany Department, British Museum (Natural History) Cromwell Road, London, SW7 Fraser-Jenkins (1977) described D. x cebennae, from the Cevennes inthe Department of Ardeche, France, which was assumed on morphological grounds to be a hybrid of D. affinis agg. x D. villarii agg. and reported to be a tetraploid apomict. However it has now become clear that a mixed concept was involved and that there are two unusual species present in the area of the type locality. A fine specimen recently found in the herbarium at Toulouse (TL) labelled “Po/ystichum abbreviatum. Par les falaises basaltiques (orgues des géants), Pont de Labeaume (Ardéche), vallée de | Ardéche vers 400 m. J. Revol 998, 4/7/1907" isclearly a specimen of D. tyrrhena Fraser-Jenkins & Reichstein, with characteristic sparse, wide and pale stipe-scales, highly glandular axes and lamina, somewhat deeply lobed pinnules and long-acuminate, almost Slightly aristate pinnule-teeth. As this was an earlier collection by Revol, the collector of the type specimen of D. x cebennae, it was necessary to re-examine the type in Manchester (MANCH). On reinvestigation the Manchester specimen was found to be also D. tyrrhena though not quite such a well-developed specimen as the one in Toulouse. The earlier report (Fraser-Jenkins 1977) that the type had a mixture of good and abortive spores was evidently due to spore immaturity and should have referred only to the author’s recent collections. D. x cebennae must therefore become a synonym of D. tyrrhena and the apomictic tetraploid taxon previously assumed to have been a hybrid is therefore without a name and is described below as D. ardechensis. Its description is emended to exclude D. tyrrhena, from which it differs in several important features (especially the scales, glandularity and pinnule teeth). It now seems highly likely that the parentage of D. ardechensis is not as originally thought, but D. affinis subsp. affinis (which is present in the area) crossed with D. tyrrhena, afar more acceptable explanation because the D. vi//arii agg. does not occur anywhere near the area and, as pointed out by Prof. C. Bange of Lyon University, (pers. comm. 1979) the 184 FERN GAZETTE: VOLUME 12 PART 3 (1981) Cevennes are a range of siliceous granitic mountains where the D. vi/larii agg. is unlikely ever to have occurred, while D. tyrrhena would be more likely. The plant is now treated as a species and nota hybrid because rather than arising de novo, it behaves as a species in maintaining itself in several populations in the area, producing a high proportion of good spores. Prof. T. Reichstein has also found that it reproduces itself freely from spores (personal communication, 1979). It is thus the only known example of an apomictic allotetraploid species in the European fern flora. This treatment is in contrast to that of some recent American authors (Wagner 1969 & 1971, and Knobloch 1977) who have introduced a multiplication sign in front of the specific name treating as hybrids all allopolyploid species once their genomic origin has been ascertained, which is the case in almost all the known European and North American polyploid Dryopteris species, a treatment with which the author does not agree. Dryopteris ardechensis Fraser-Jenkins, sp. nov. Diagnosis: Morphologia intermedia inter D. tyrrhena et D. affinis subsp. affinis Frondes usque ad 50cm longae; petiolus media longitudine laminae, paleis angustis uniformibus pallide fulvis costam versus angustissimis piliformis bene obsitus; lamina triangulari-lanceolata, ad axes pauce glandulis interdum obsita, simpliciter pinnata sed par inferius pinnularum cujusque pinnae iterum pinnatum; pinnulae basi lata sessiles, pars infimo cujusque pinnae excepto quod bene petiolulatum; pinnula infima basiscopica paris infimi pinnarum longissima et ad marginem aliquanto lobata; latera pinnularum aliarum parallela parce tantum dentata nisi apicem versus, apices rotundati et dentibus aliquanto longis et acutis sed a basis latioris quam in D. tyrrhena nonne interdum mucronatis. Indusia cinerea, pallida fulva et fusca eanta, margine inflexo, glandulas ferrentas. Sporae partim abortivae, partim bene evolutae. Cytotypus tetraploideus, apogamus. Type. Ex hort. T. Reichstein (sub TR 3443); coll. 26. Aug. 1979 C.R. Fraser-Jenkins No. 9784. - Origin from spores coll. CRF-J (no. 3602) above Pont de Labeaume, near Aubenas, Ardeéche, France 17 Jan. 1972 (Holo: BM, Iso: CANU). Plant a tetraploid apomict (G. Vida pers. comm. 1975, and J.J. Schneller 1980, pers. comm.). Differs from the original description of D. x cebennae in being well clothed with stipe and rhachis scales, lamina occasionally bearing a few glands on the axes, teeth at the pinnule apices with wide bases somewhat long and acute and not tending to become aristate. Indusia grey, becoming pale-brown and then mid-brown, bearing glands. Its morphology is thus rather more intermediate between D. affinis subsp. affinis and D. tyrrhena than D. affinis and D. submontana, as far as can be extrapolated. Three new records have now been verified as follows: ip Ardéeche, Thins. C. Bange, 1978, nos. 78.313, 78.317, 78.319, 78.322 and TG a2 5 (LY 1): 2 Ardéche, Gravieres, bois a 1 km a val, 235 m ™. Breistroffer, 30/10/1970. (GRM!). o: Ardéche, Malarce aux Eynes, ravin ombragé. M. Breistroffer, 4/7/1970 (GRM!). REFERENCES FRASER-JENKINS, C.R., 1977. Three species in the Dryopteris villarii aggregate. Candollea 32 (2): 305-319. KNOBLOCH, I.W., 1977. Pteridophyte hybrids. Pub/. Mus. Michigan State Univ., Biol. ser., 5 (4): 1- 352 (and addend. & corrigend.) WAGNER, W.H., 1969. The role and taxonomic treatment of hybrids. Bio Science 19 (9): 785-789. WAGNER, W.H., 1971. Evolution of Dryopteris in relation to the Appalachians, in P.C. Holt (Ed.), The Distributional History of the Biota of the Southern Appalachians II. Flora. Research Div. Monograph 2, Virginia Polytech. Inst. & State Univ. THE FERN GAZETTE ~ Original papers, articles, or notes of any length on any aspect of pteridology will be 2 considered for publication. Contributions will be particularly welcomed on floristic, biogeographical and ecological aspects of ferns and their allies, worldwide. Members bg e Bar. are especially encouraged to make use of short notes for reporting pteridophyte field -_ observations and records. All may be illustrated with black and white photographs of a : y - good technical quality and line drawings. Short papers can usually be incorporated % ' more quickly than longer ones. i . 7 bath a s * ye 4 Contributions should be sent to: ; Re. Py a C.N. Page: Royal Botanic Garden, Edinburgh, U.K. ee 30th November each year Be) Ae ne ehinislcl follow the general style of this number. Close adherence to the Evy. ss fee notes will help to speed the publication of contributions. aa 5. as NOTES FOR CONTRIBUTORS a 24 ter "a : is: Copy should be in English and submitted in double-spaced type with adequate ma ins, ‘on one side of the paper oniy. _ Abstract: All papers, other than short notes, should include a short abstract, to be set at the head aN of he Baad text, indicating the scope of the topic and the main conclusions. Heaa ne lings and sub-headings. These should follow the style of this number. (Primary sub-headings are recent capitals. Secondary sub-headings U and L case side roman. Tertiary sub-headings, if Wises re ee ry, are U and L case side italic.) eee of sub- jstagrka should be avoided. “ag be Reprints: Twenty-five reprints are supplied free of charge to authors, who may order in advance _ further reprints which will be supplied at cost (plus postage) if requested at time of returning the fi irst proofs. sy , - "1 BOOKS FOR REVIEW oy Books for review in the Fern Gazette or B.P.S. Bulletin should be sent to A.C. Jermy, ve Botany Department, British Museum (Natural History), Cromwell Road, London, _- SW7 5BD. FERN GAZI VOLUME 12 PART 3 CONTENTS MAIN ARTICLES Drought tolerance in Cheilanthes with special reference to the « > ge -—H. Quick & T.C. Chambers | ‘ vis) a aeabe tN Tropisms in Regnellidium diphyllum - i D. Gontan pa haat e ae An ecological survey of the Ferns of Berwickshire, Scotland - Cheilanthes coriacea, An Afro-Arabian species new to Iran - A chromosome count for Asplenium anceps from me Canary Islands - H. Rasbach, K. Rasbach & J.J. Schneller — A chromosome count for Macrogolossum (Marattiales) - re cs: Walke ee A An analysis of The Grammitis poeppigiana-G. magellanica complex it in h ic South Atlantic and South Indian oceans - B.S. Parris basea Two new species of Selaginella from the Phillipines - B. og Tan & A c A forked vein and foliar fibres in Sergenes R.N. Mukherjee & U. Ser SHORT NOTES Dryopteris x sarvelae in Scotland; a new hybrid for the British Isles e - H.V. Corley & M. Gibby MS A new name for a hybrid horsetail in Scotland - re, N. “Page Equisetum x trachyodon in Skye, Western Scotland - C.W. Murray — More new combinations in Southeast Asian Ferns - B.S. Parris The occurrence of a laminal flap i in Selaginella - R. Mukhopadhya } & U A new species of Plagiogyria from Malaya - A.D. Dixit &A. Das : ey | Nomenclature notes on Dryopteris - CR. Fraser-Jenkins ae Sere REVIEWS he Ne ca iee near eet (THE FERN GAZETTE Volume 12 Part 2 was s published in ahi PLL ge A ‘ # - : ike 4 ) meee - ?. ey METLOC PRINTERS LTD - Old Station Road - Loughton Essex vs ii ae a . ¥ VOLUME TWELVE PART FOUR. as ‘. = OOS rac ISSN 0308—0838 f/ : fA AL 4™0. 4009 IAN T © Woo = © = ir mit R ~~ (BRAN EY ‘ s \ THE JOURNAL OF THE Tea Ae BS ISH PTERIDOLOGICAL SOCIETY a a y othe ey ’ ws. A Parr ain? ms at en THE BRITISH PTERIDOLOGICAL SOCIETY | Officers and Committee for 1982 NN hil ee pala Ne est Pas Ton _ President Vice Presidents General Secretary Treasurer Membership Secretary K. Kava nagh, Botany Pepsieclan British M i ¥ Meetings Secretary useur 4 (Natural eipabieg rey. Road, London, sv ene eI si | ne Pea: Publications Secretary Editors: Fern Gazette Archivist Committee M. Barker, J.A. Crabbe, J.K. Grane Barbara S. Croxall, Pp. awa Mary Gibby, N.A. Hall, ape aek segs R. Rush, J Woo T a P eee — ee ‘ace 5 The Fern Gazette and the Bulletin are the journals of the British Pteridological Society, Gees eee the Bulletin topics of more general appeal. (See inside back cover for notes to contributors). . i a rn 4 eee BN hy ; ee Fe ae gir, 4 The BRITISH PTERIDOLOGICAL SOCIETY was founded in 1891 and today cannrade asarenuel or ; fern enthusiasts. It provides a wide range of information about ferns through the medium of its publications and available literature. It also organises formal talks, informal discussions, field meetings, garden visits, searches for rare fern books, plant exchanges and a spore exchange — scheme. The Society has a wide membership which includes gardeners, nurserymen and — botanists, both amateur and professional, in Britain and overseas. Articles for the Soc < 's ae publications are received from and reflect the interests of, Society members. “- she Membership is open to all interested in ferns and fern-allies, worldwide. Subscription Be a, on 1st January each year) are: Full Personal Members £7; Personal Members not receiving the Fern Gazette £5; Student Members (full-time students) £5; Subscribing Institutions £8. — Applications for membership should be sent to the Membership Secretary (address above), from whom further details can be obtained. (Remittances made in currencies other than Sterling are £1.00 extra, to cover bank conversion charges.) Back Butter of the Gazette and Bulletin are available for purchase from m W. Dyce address above), from whom further details can be obtained. FERN GAZ. 12(4) 1982 185 THE CONTINUING NEED FOR MORE MONOGRAPHIC STUDIES OF FERNS R. E. HOLTTUM c/o Royal Botanic Gardens, Kew, Richmond, Surrey, England In 1972 | was asked to give an opening address (published 1973) at a meeting of pteridologists in London. Though much new investigation and thought has developed since then, my attitude to the general situation is still much the same. The need still is for more monographic work on tropical species which represent the great majority of all ferns. Until we have a better knowledge of them we cannot formulate adequate generic concepts which may lead to better suprageneric ones, nor understand the findings of morphologists who study a few species. Experimental workers of all Kinds, including cytogeneticists, need a good taxonomy to indicate likely subjects for fruitful work in their own disciplines and to enable them to assess its significance in relation to the whole. Such workers hitherto have largely dealt with temperate-zone species and need to be more aware of wider possibilities. Questions of nomenclature are also basic; aname must have a definite meaning, the lack of which in earlier literature has led to confusions. Monographic work, including re-examination of all types, is needed to clarify this situation. | would like briefly to summarize the history of my own involvement with ferns. As a student | was introduced to the morphological studies of F.0. Bower and to fossil ferns and their presumed living relatives, but hardly at all to fern taxonomy in general. When | went to Singapore in 1922 | was surrounded by ferns of genera and species many of which were new to me. | soon realized that fern taxonomy was in a confused state, and | gradually came to realize that some of Bower's generalizations (Summarized in his volume of 1928), based on few species, too great an emphasis on vascular anatomy and on the taxonomy of the 19th century, were untenable. Classifications of the 19th century were in most cases unnatural because they were based on rather crudely defined structures which modern understanding shows to have evolved on several different lines. For example, anastomosing veins of similar pattern were used to associate genera which are now seen not to be closely related, such as Acrostichum and Lomogramma. On the other hand, species with free and anastomosing veins, though in most other characters similar, were placed in different genera; Diels (1899) copied many such errors. My first major investigation was on the ferns then still named Stenoch/aena in Malaya (1932). After examining their external morphology, anatomy, scales and spores, | realized that they constituted a mixture of quite unrelated species; also that this had been recognized by Mettenius (1869) and John Smith (1875, p.140) but ignored by Diels and by Bower (1928, pp 175—177). Thus | came to understand that most 19th century taxonomists ignored important characters which need to be considered; also (like John Smith) that one can learn much from living ferns that is not clearly evident from dried specimens. | had ready access to the species of Malaya (about 500) and from them | learned a great deal. In this | was greatly helped by Carl Christensen with whom | corresponded regularly from 1925 to 1940; | also met him personally and had long talks with him in 1930 (after the Cambridge Congress), 1934 and 1938. He was the founder of modern fern taxonomy, but when | was a student | never heard his name. * This paper is the substance of an address given by Dr Holttum on his election to the Presidency of the International Association of Pteridologists, 24 August 1981. 186 FERN GAZETTE: VOLUME 12 PART 4 (1982) In the course of writing a fern flora of Malaya, which by good fortune | was able to do during the Japanese occupation of Singapore, | attempted to carry my thoughts beyond Christensen’s summary of 1938; | tried to express my ideas in two papers dated 1947 and 1949 and in my book on Malayan ferns (1955). It was at this point that | came into contact with Prof. Manton’s cytological studies (1950) and! began to realize the importance of this in fern taxonomy, how it confirmed some of my ideas (e.g. on Thelypteridaceae) and pointed to the need for more thought on others. | was fortunate in being able to help her to extend her studies to Malayan species, partly through fixations made in the field in Malaya and partly through plants sent to Kew for Cultivation. . In 1954 | retired from my appointment in Singapore and came to work at Kew, having as my objective the preparation of a taxonomic account of the ferns in the whole Malayan region (Malesia) for Flora Malesiana. | began with Gleicheniaceae (1959) which are abundantly represented in Malaya. Here | found that Diels had followed Pres! and Hooker in making a primary division, of Eu-G/eichenia from the rest, on the terminal or non-terminal position of sori on the veins, though Mettenius (1863) had shown that the sorus is not terminal in the former; Diels ignored the excellent subdivision made by Mettenius and so obscured the basic similarity between Eu- Gleichenia and Diplopterygium which Bower never understood. It seems to me abundantly evident that Dip/opterygium represents the primitive element in the family as it now exists. | proceeded to Cyatheaceae (1963) concerning which work for Flora Malesiana greatly widened my horizons. Instead of the sixteen species of Cyathea in Malaya, | had to deal with 190 species, for a better understanding of which | also examined the species of mainland Asia (1965) and the Pacific (1964). This acquainted me with a considerable proportion of the world total and enabled me to recognize infrageneric groups which were not so distinct in the few species of Malaya; | came to see the latter in a better perspective. | was fortunate to have the cooperation of Dr. Uday Sen while! was studying Cyathea. It was his anatomical study, published in a joint paper (1961) which led me to see the similarities between Cyathea and Dicksonia which, following Bower, | had previously seen as wide apart. | also saw how very different Dicksonia is from Culcita, though their sori are superficially similar. | proceeded to look at Thelypteridaceae, a very distinct family of nearly one thousand species which was not recognized as a natural group before the work of Christensen, extended by Ching (1936, 1938) to the species of Asia. When preparing my book on the ferns of Malaya | discovered that existing subdivisions of the family were unsatisfactory (Copeland's Lastrea and Cyclosorus of 1947 are both confused mixtures) but | could see that study of a much wider range of species was necessary before | could understand the situation better. Such a study occupied twelve years, and has been extended to all species of the Old World, including Africa, where | found that two groups of species are mainly American in distribution. The family is much more diversified in the Old World than in the New, and in the Old World more diversified in Malesia than in mainland Asia, to which the studies of Ching and Iwatsuki (1964, 1965) had been almost confined. | was led to establish new genera and to re-define old ones (1971), incidentally showing that five European species of the family belong to five distinct genera when viewed on a world scale. Having done this, | tried to consider possible groupings of genera, and have set them out ina conspectus in the next part of Flora Malesiana, which | hope will be published this year. But it is evident that more observation and thought are still needed. The object of preparing such a conspectus is to show where new thought is called for. In the course of all this work it became evident to me that in any group of ferns R.E. HOLTTUM: CONTINUING NEED FOR MORE MONOGRAPHIC STUDIES OF FERNS ~ 187 there are distinctive characters whereby species and species-groups can be recognized. As noted above, such characters were mostly not described by taxonomists in the 19th century, which is the main reason why they did not arrive at an acceptable taxonomy for ferns in general. In different groups, different characters may be important; one cannot tell in advance. Such characters may appear trivial (as they did to William Hooker), but where two or three, based on different parts of a plant, are found constantly associated, one invariably finds that other, less clearly definable, characters are also associated. Christensen made this observation in his paper on Stigmatopteris in 1909 (p.292), but Schkuhr had already shown, a century earlier, the difference between the hairs on Thelypteroid and Tectarioid ferns. Effective taxonomy is not a routine matter of recording a limited number of characters, but a scientific discipline requiring imaginative insight just as much as in any other field of work. This is what many botanists who are not concerned with tropical plants do not realize. New morphological characters, if recorded by the use of highly complex instruments, are considered to be of scientific importance; but the significance of such observations (e.g. SEM photographs of spores) is only evident if the object under investigation can be seen against a good taxonomic framework, and observations made by relatively simple optical equipment are still important in establishing that framework. Modern methods in cytogenetics, still little applied to tropical ferns, can only be fully exploited where taxonomy shows how to distinguish natural groups of species. So the moral of all this may be summarized as follows. A taxonomist who wishes to attain a better understanding of genera which appear not to be well defined must examine all species (some of them may have been previously placed in other genera). A morphologist cannot do this, but his observations may indicate to taxonomists new characters which may be used in their comparative studies. Morphologists cannot understand the significance of their work without the cooperation of taxonomists, and conversely, taxonomists must take into consideration the evidence of morphologists and use it to find new characters by which to improve their systems. The two processes interact. My impression is that the greater need at present is for morphologists to try to have a better understanding of taxonomy. As | remarked in 1973 (p.3) the Code of Nomenclature is responsible for much misunderstanding. The basic rule is that no new species can be validly named and described unless it is assigned to a genus. This implies that the correct genus is known. But the earlier taxonomists could not know how future additions to knowledge might modify generic concepts; this is especially true of ferns. The Code thus makes the impossible a necessary condition for action. Taxonomists have to make the best judgements they can, and if subsequent change of genus for a species is indicated, the Code makes rules as to how the change is to be effected. Morphologists are too apt to think that a generic name has a definite meaning and that they can take any species as representing the characters of the genus the name of which they find attached to it. In fern taxonomy name changes have been many, often uncritically made, and we have not reached finality. There are many tropical groups which need new monographic treatment. The process should be: (1) a careful study in detail of living plants of a varied selection of species, to discover what are distinctive characters; such a study of living plants helps to understand dried specimens; (2) re-description of all type specimens for record of characters not noted by the original authors or later workers; (3) examination of all collections of specimens in the major herbaria, and as much new field work as may be possible; (4) assessment of possible new generic concepts; (5) nomenclatural problems. Note that nomenclatural problems come last in this series. There must be 188 FERN GAZETTE: VOLUME 12 PART 4 (1982) rules to stabilize binomials; | am doubtful of the need for any rules beyond that. NOTES ON SOME TROPICAL GROUPS IN NEED OF MONOGRAPHIC STUDY Polypodiaceae sensu stricto. Nearly all species were formerly lumped in one genus Polypodium. Genera were split off mainly on the basis of shape and position of sori, venation and frond-form. As regards venation, it is evident that in many groups of ferns species with free and anastomosing veins can be allied, and | believe that this has not been sufficiently considered in the Po/ypodium alliance. Characters of scales are recognized to be important when distinguishing between allied species; generic characteristics of scales are not easy to define but | suggest that some may be found. It is evident that in India Copeland's definition of Po/ypodium applies mainly to species related to his Goniophlebium but not nearly related to Polypodium vulgare. | have suggested (1947, p. 128) that primitive Polypodiaceae had fronds with much- anastomosing veins; on such a view the many diverse tropical American species now in Polypodium are advanced and specialized. In 1973 (p. 7) | suggested that Polypodiaceae are related to Davallia, though not very closely. | also think that both are in some way related to Asp/enium. A curious character in common to the three groups its the lateral union of the adaxial vascular strands near the top of the stipe; | have never seen this in any other ferns and suggest that it may be significant. Dennstaedtia, Microlepia and Hypolepis. | believe that these three genera are still not clearly distinguished, and that Dennstaedtia, as at present constituted, represents a mixture of distinct genera. R.M. Tryon (1960) has recognized eleven species of Dennstadetia in the Neotropics, dividing them into two groups, but he also added a list of other names the types of which he had not seen. There are a considerably larger number of species in the Palaeotropics, mostly in the Philippines, New Guinea and the Pacific; most of them have not been adequately described or distinguished, and their large fronds are not well represented by fragments on herbarium sheets. Chromosome counts indicate diversity as between Palaeotropic and Neotropic species, also within the former. R.M. & A.F. Tryon (1980) have stated that Dennstaedtia flaccida (Forst.) Bernh., type species, has spores like those of Microlepia, but one cannot define genera on spores alone. This group of genera seems to me to have an important place in the classification of ferns as a whole. It includes Pteridium, | am still not convinced that Pteris is quite unrelated to Pteridium. Asplenium. Copeland (1947, p. 164) stated that there were about 700 species of this genus. He included Phy//itis and Diplora as synonyms but expressed the opinion that the types of these two genera, though agreeing in their peculiar arrangement of opposite pairs of sori, were not closely related, and | believe he was right. He further stated that this arrangement of sori had arisen at least six times independently. Hooker had placed all ferns with this arrangement ina ‘‘suborder’’ Scolopendrieae, and Bower (1928, p. 188) placed them with the Blechnoid ferns, denying their near relationship to Asplenium (p. 197). Dr John Mickel has recently proposed a new generic name Sinephropteris for one of them. No one has yet produced a good subdivision of Asp/enium. Until this is done the position of Phy//itis and others with similar soral arrangement will not be clear. In my book on Malayan ferns (1955, pp 414-415) | made suggestions for a subdivision, and | believe that the form of the rachis in relation to the attachment of pinnae provides useful evidence. But to make a good subdivision will require a study of all species. European species have been studied in great detail but without reference to their possible relationships among tropical species. In Europe there is evidence of polyploidy and hybridization; in the tropics there is high polyploidy in some groups of species. These considerations make the discrimination of individual species difficult, R.E. HOLTTUM: CONTINUING NEED FOR MORE MONOGRAPHIC STUDIES OF FERNS ~ 189 but for purposes of subdivision of the genus such detailed problems need not be involved. A special problem is presented by the allied genus P/eurosorus which has a very peculiar distribution. In Spain there is a species Asp/enium petrarchae (Guérin) DC which is similar in its general aspect to Pleurosorus hispanicus (Cosson) Morton and the two species will hybridize. lt seems to me probable that they have been derived from a common ancestor, now no longer existent. The other species of P/eurosorus are in Australia, New Zealand and South America. | suggesi that these species may have separately evolved from species of Asp/enium in southern regions. The Tectaria group of genera. The distinctions between Jectaria and Ctenitis appear not to be very clearly defined, and Ctenitis, as at present interpreted and after removal of Lastreopsis, appears to be an unresolved complex. The position of Heterogonium in relation to Jectaria also needs re-consideration, and the fact that natural hybrids between Heterogonium and Stenosemia have been found. | think that in this group of genera characters of scales may be important as indicators of relationships. Athyrium, Diplazium and related genera. This is a large and complex group in the Palaeotropics. In Malesia the many species of Dip/azium have not yet been clearly distinguished and some are undescribed. Ching’s work (1964) on the genera and species of mainland Asia needs to be studied in relation to those of Malesia. REFERENCES BOWER, F.O., 1928. The Ferns, Vol. 3. Cambridge University Press. CHING, R.C., 1936, 1938. A revision of the Chinese and Sikkim-Himalayan Dryopteris with reference to some species of neighbouring regions. Bul/. Fan Mem. Inst. Biol. Bot. 6: 237-352, 8: 157-268. CHING, R.C., 1964. On some confused genera of the family Athyriaceae. Acta Phytotax. Sinica 9: 41-84. CHRISTENSEN, C., 1909. On Stigmatopteris, a new genus of ferns, with a review of its species. Bot. Tiddskr. 29: 291-304. CHRISTENSEN, C., 1938. Filicinae. Chapter 20 in Fr. Verdoorn (Ed.) Manual of Pteridology, The Hague, Nijhoff. COPELAND, E.B., 1947. Genera Filicum. Waltham, Mass. Chronica Botanica. DIELS, L., 1899. Cyatheaceae (pp 113-139), Polypodiaceae (pp 139-339) and Gleicheniaceae (pp 350-355) in Engler & Prantl, Die Naturlichen Pflanzen-familien |, Abt. 4. Leipzig, W. Engelmann. HOLTTUM, R.E., 1932. On Stenochlaena, Lomariopsis and Teratophyllum in the Malayan region. Gard. Bull. Str. Settl. 7: 245-312. HOLTTUM, R.E., 1947. A revised classification of Leptosporangiate Ferns. J. Linn. Soc. Bot. 53: 123-158. HOLTTUM, R.E., 1949. The Classification of Ferns. Biol. Reviews 24: 267-296. HOLTTUM, R.E., 1955. A Revised Flora of Malaya, 2. Ferns of Malaya. Singapore, Government Printer. HOLTTUM, R.E., 1959. Gleicheniaceae, in Flora Malesiana, Ser. || Pteridophyta, 1: 1-36. HOLTTUM, R.E., 1963. Cyatheaceae, in Flora Malesiana Ser. || Pteridophyta, |: 65-176. HOLTTUM, R.E., 1964. The tree-ferns of the genus Cyathea in Australasia and the Pacific. Blumea 12: 241-274. HOLTTUM, R.E., 1965. Tree-ferns of the genus Cyathea in Asia (exclusing Malesia). Kew Bull. 19: 463-487. HOLTTUM, R.-E., 1971. Studies in the family Thelypteridaceae, Ill. Anew system of genera in the Old World tropics. B/lumea 19: 17-52. HOLTTUM, R.E., 1973. Posing the Problems, (A.C. Jermy et al (Eds.) in The Phylogeny and Classification of the Ferns, Suppl. no. 1 to the Bot. Journ. Linn. Soc. Vol. 67. HOLTTUM, R.E., 1978. A commentary on some Thelypteroid ferns of Eastern Asia and North America. Acta Phytotax. Geobot. 29: 16-17. HOLTTUM, R-E., and SEN, U., 1961. Morphology and Classification of the Tree-ferns. Phytomorphology 11: 406-420. IWATSUKI, K., 1964. Taxonomy of the Thelypteroid ferns, with special reference to the species of Japan and adjacent islands, Ill, Classification, Mem. Coll. Sci. Univ. Kyoto B, 31: 11-40. 190 FERN GAZETTE: VOLUME 12 PART 4 (1982) IWATSUKI, K., 1965. Taxonomy of the Thelypteroid ferns, with special reference to the species of Japan and adjacent islands, IV. Enumeration of the species of Japan and adjacent islands. Mem. Coll. Sci. Univ. Kyoto B, 31: 125-197. MANTON, I., 1950. Problems of Cytology and Evolution in the Pteridophyta. Cambridge University Press. METTENIUS, G., 1863. Gleicheniacea. Ann. Mus. Bot. Lugd.-Bat. 1: 46-51. METTENIUS, G., 1869. Lomariopsis (p. 294), Teratophyllum (p. 296) in M. Kuhn, Filices. Ann. Mus. Bot. Ludg.-Bat. 4: 276-300. MICKEL, J., 1976. Sinephropteris, a new genus of Scolopendrioid ferns. Brittonia 28: 326-328. TRYON, R.M., 1960. A review of the genus Dennstaedtia in America. Contr. Gray Herb. 187: 23- 52. TRYON, R.M. and TRYON, A.F., 1980. Proposal to conserve the name Dennstaedtia T. Moore (1859) against Dennstaedtia Bernh. (1801), Sitolobium Desv. (1827), Patania Presl (1836) and Adectum Link (1841). Jaxon 29: 512. REVIEW FERNS, FERN ALLIES AND CONIFERS OF AUSTRALIA by H.T. Clifford and J. Constantine. University of Queensland Press, distributed by Prentice-hall International, London. September 1981. 246 x 186mm, 36 tables + 24 plates. Price £12.95. This slim but valuable book provides a synopsis of the Australian non-flowering vascular plants. Botanical descriptions are given of all families and genera, with short and straightforward keys to families, genera and species. Information on the total range of taxa is provided, together with their distribution within Australia. Many of the more technical structural details of plants mentioned inthe text are illustrated by good line drawings. The choice of concentrating on generic descriptions rather than specific ones at first seems unusual. But with 387 species of Pteridophyta keyed (probably a conservative estimate) belonging to no less than 123 genera, and witha similar ratio amongst the conifers, a working knowledge of genera present in the native flora is paramount to the Australian senior schools level and university biology student, at whom the book is primarily aimed. It should, however, give a very sound generic grounding to many a fern enthusiast far outside Australia, for all but a handful of the pteridophyte genera described range far through the tropics and very many are to be found in cultivation in Europe. To anyone also interested in the wider issues of the plant geography of this fascinating continent, this book provides a valuable and highly recommended foundation. C.N. PAGE FERN GAZ. 12(4) 1982 191 A PRELIMINARY SURVEY OF THE PTERIDOPHYTA OF TRANSKEI, SOUTHERN AFRICA J.P. ROUX Drakensburg Botanic Garden, P.O. Box 157, Harrismith 9880, South Africa ABSTRACT The results of a preliminary survey of the pteridophytes on the Transkei coast and midlands are described. Sixty-five species of pteridophytes are recorded. INTRODUCTION Transkei is situated on the east coast of Southern Africa between the 29°30’ and 32°S latitude and 27°30’ and 30°E longitude (Fig. 1). The 250 km long coast forms the eastern border, and the southern foothills of the Drakensberg the northern and western borders. South Africa U7, © J MKAMBATI BIG UMNGAZI RIVER fe / e J J ae: ~/PORT ST.JOHNS A BASHEE RIVER pape { _7% A PJDWESA FOREST : a / a ¢ SS KE! RIVER thea Ss 0 FIGURE 1. Map showing the location of Transkei, Southern Africa. 192 FERN GAZETTE: VOLUME 12 PART 4 (1982) The landscape is much dissected, rising steeply from the coast to the upper plateau, and there are many short rivers. The climate along the coast is sub-tropical but a mediterranean climate is experienced in the midlands. The coastal belt (0-600 m) receives an average annual rainfall of 1,027- 1,592mm per annum, with the main rainy season from October to March, but some rain occurs throughout the year (184mm October to March, 43mm April to September). June is the driest month. Temperatures of the coastal regions are more constant than inland. In the south, the maximum average temperature in February (which is the warmest month) may reach 25.6°C at Bashee in the south and 26.5°C at Port Edward in the north. July is the coldest month and temperatures may drop as low as 11.8°C in the south and 11.7°C in the north. The midlands vary slightly from the coastal belt for here the warmest month is during January when temperatures of 27°C can be measured. Temperatures drop as low as 0.7°C in July in this region. VEGETATION Typical Coastal Belt Forest This vegetation forms the major part of the coastal forests and is found in the form of riverine forests in deep dark gorges and valleys. It is usually a dense tangle near water, but becomes more open on the higher slopes. This broad-leaved evergreen forests in some instances cover vast areas. The palm Phoenix reclenata on stream or riverbanks is characteristic of this forest type. Trees of Mi/letia grandis, Protorus longifolia, Vepris undulata form the main canopy, Grewl/a occidentalis, Uvaria caffra and Rhoicissus tomentosa the main undergrowth and climbers and Oplismenus_hirtellus, Plectranthus ecklonii and Cyperus albostriatus the common forest floor vegetation. Transitional Coastal Forest This broad-leaved evergreen forest is more common inland in drier areas, and Is confined to steep riverbanks and valleys. The forest is not as high as the typical coastal belt forests. The main trees are Schotia spp., Cassine spp., Euphorbia grandiceps and Strelitzia nicolai which is characteristic of this forest type. Dune Forest ; The dune forest occurs intermittantly in a narrow strip on the dunes running down the coast, especially around estuaries. This evergreen forest is usually stunted on the seaward side but much higher (12m) on the landward side, where protected from prevailing winds. The main canopy is formed by Mimusops caffra, Euclea natalensis, Canthium obovatum and Apodytes dimidiata. The understory is formed by Scutia myrtina, Draceana hookerana and Eugenia capensis. Main forest floor and margin species are Dactyloctenium australe, Oplismenus hirtellus, Scadoxus membranaceus. Characteristic of these forests north of Port St Johns is the abundance of Strel/itzia nicolai. Pondoland Coastal Plateau Sourveld This vegetation type occupies the plateau between 300-450m above sea level. The escarpment and gorges are tropical at the coast and sub-tropical on the higher elevations and these are forest-clad. The plateau itself is of dense perennial grass veld. This is a broad-leaved evergreen forest and is higher than the coastal tropical forests with trees of Strelitzia nicolai, Protorus longifolia and Schefflera umbellifera forming the main canopy, with other scattered trees and shrubs of Uvaria caffra, J.P. ROUX: PTERIDOPHYTA OF TRANSKEI 193 Smilax kraussiana, Dalbergia obovata and Flagelaria guinensis. Main undergrowth and margin species include Lobelia patula, Cyperus albostriatus and Setaria chevalieri. The dense plateau consists of grasses such as 7Themeda triandra, Tristachya hispida and Trachypogon spicatus. Zululand Thornveld This relatively dry type of evergreen forest occupies the escarpment of lower altitudes and the higher ridges. Although not a well developed forest it includes species such as Turaea floribunda, Rapanea melanophloeos and Apodytes dimidiata. THE FOREST PTERIDOPHYTES Epiphytes Two minor groups, the true epiphytes and the low level epiphytes, can be recognised. The true epiphytes probably require a high rainfall and humidity. Many occur in deep shaded forests. They include Pyrrosia africana, Microsorium punctatum and Pleopeltis macrocarpa which all have simple fronds and creeping rhizomes. The latter is fairly rare in the coastal forests but more common inland, where it grows with Polypodium polypodioides subsp. eck/lonii, a fern with a thin slender creeping rhizome which is totally absent in the coastal forests. Low level epiphytes include Asplenium sandersonii, a_proliferous species, A. rutifolium, A. prionitis and Microgramma lycopodioides. The latter is also often a lithophyte in coastal forests. FIGURE 3. Microgramma lycopodioides is a common lithophyte or low level epiphyte in broad-leaved evergreen forests. Lithophyes The majority of lithophytes are confined to wet coastal forest with a high humidity, and to very deep shade. Two of these species, Asp/enium rutifolium and Microsorium punctatum, may also occur as epiphytes. The latter, when sheltered, can have fronds 1.6m long. The true lithophytes are Lycopodium gnidioides, L. verticilatum with long pendulous dichotomously branched stems, Se/aginella mittenii, Trichomanes pyxidiferum var. melanotrichum, Microgramma lycopodioides, Oleandra distenta, a deciduous species, Asplenium gemmiferum, Rumohra adiantiformis and Vittaria isoetifolia. The latter becomes stunted with fronds as small as 6cm long, when growing under dry conditions. The Dry Forest Species These occur in areas of well drained soils within the forests. Although relatively dry, there is usually some moisture available through the year. Species growing along banks of annual streams, which are dry for most of the year, are also included here. Principal species are Anemia dregeana, Cheilanthes concolor var. kirkii, C. viridis var. macrophylla, Asplenium splendens, Ceterach cordatum, usually found as a 194 FERN GAZETTE: VOLUME 12 PART 4 (1982) xerophyte, Dryopteris inaequalis and Polystichum lucidum. All of these are common on the forest floor and usually have short erect rhizomes with the exception of the latter two species. Wet Forest Floor Included here are plants from all areas where the forest floor is more or less permanently wet from seepage, including most stream bank ferns. Light intensity is often much lower than in the areas of drier forest floor. Common species include the delicate Se/aginella kraussiana, Marattia fraxineavar. salicifolia, Pteris buchananii, P. catoptera var. horridula, Cheilanthes bergiana, Asplenium lunulatum, Thelypteris bergiana, Blechnum attenuatum var. giganteum and B. punctulatum var. krebsii. The Dune Forest These are areas of forest growing on dry sandy soils, often amongst dense vegetation on the landward side of dunes. Only one species, Microsorium scolopendria with a long slender rhizome occurs in this forest type. It often becomes epiphytic, and is usually absent from areas facing the sea. NON-FOREST PTERIDOPHYTES These include all pteridophytes growing in exposed conditions including those of forest margins which are absent from within the forest itself. Aquatic Species These include all habitats which are at least seasonally flooded. Only one aquatic, Marsilea macrocarpa, was recorded in two shallow pans which were almost dry by April. This plant species continues to grow, although not as vigourously, through the winter months. In exceptionally dry years the plants will die completely. As soon as FIGURE 2. Davallia caryophylloides as a non-forest lithophyte on the escarpment of the lower plateau. J.P. ROUX: PTERIDOPHYTA OF TRANSKEI 195 water fills these pans again the sporocarps will open letting the spores free which develop rapidly into vigorous growing plants. Lithophytic Species These include all pteridophytes of exposed rocky situations, not shaded by trees. Such lithophytic species are more common on the escarpment of the lower plateau areas, where the rainfall is high (about 612-592mm annuaily). These species succeed particularly well in these habitats: Se/aginel/la dregei, Pityrogramma calomelanos var. aureoflava and Davalia caryophylloides. Four others occur only where there is some slight protection from exposure: Se/aginella caffrorum, Cheilanthes inaequalis var. buchanani, Elaphoglossum macropodium and Blechnum punctulatum. One remaining species, Lycopodium gnidioides is also a forest dweller, which occurs only on a shaded part when growing under exposed conditions. Hygrophytic Species Although most of these species are common on wet bank situations, some occur along forest margins. Species include: Lycopodium clavatum var. inflexum, L. cernuum, L. carolinianum, Equisetum ramosissimum, Osmunda regalis, Todea barbara, Mohria caffrorum, Gleichenia volypodiodes, Alsophila dregei, Hypolepis sparsisora, Pteridium aquilinum, Adiantum capillis-veneris, A. radianum, Pteris vittata, Nephrolepis exaltata, Thelypteris confluens, T. totta, Blechnum sylvaticum and B. tabulare. Of these species, Mohria caffrorum will shrivel up or the fronds may die down entirely during a dry spell. Xerophytic Species Only four plants come into this category. These are Selaginella dregei, Schizaea pectinata, Cheilanthes viridis var. involuta and Pellaea calomelanos. All are common on the upper plateau in dry, exposed conditions. The latter species have the habit of dropping segments during long dry spells leaving the plant with a very twiggy appearance. Introduced Species In this study, three introduced species have been identified: Pityrogramma calomelanos var. aureoflava, Adiantum radianum and Nephrolepis exaltata. Any could rapidly develop into a threat to the local flora. Pityrogramma calomelanos var. aureoflava is spreading at a tremendous rate. Collections have been made on the Umtanvuna riverbed far from civilization. This species can develop into a pest, adapting freely to many weedy situations. Adiantum raddianum has been recorded at only two localities (both at Port St Johns) and in both cases it was growing on wet roadbanks. The third species, Nephrolepis exaltata, had invaded a patch of native wet forest. Habitats that seem potentially threatened by expansion of these introduced species include those of native Lycopodium cernuum, Thelypteris totta, Adiantum capillus-veneris and Microsorium punctatum (both as a lithophyte and epiphyte). N. exaltata is a garden escape and flourishes in the warm tropical climate of the coast. This species has spread at a tremendous rate threatening to establish itself in the deeper forest. CONCLUSIONS The Transkei has a rich pteridophytic flora. A range of specialized habitats is necessary to ensure the existance of these plants. The forest dwelling species are under ever increasing pressure through the continual deforestation by the local people. These species need certain micro- 196 FERN GAZETTE: VOLUME 12 PART 4 (1982) climates to survive and which only the forests can provide. Once these have been destroyed the plants will disappear entirely from these areas. Forest dwelling species cannot adapt to other conditions as can the xerophytes. Few coastal forest reserves which are relatively rich in fern flora already exist and is at present in a good condition. A definite need for forest reserves inthe midlands exists. Deforestation in these areas is at present very high. No endemics are however restricted to this region. Elsewhere Davalia caryophylloides populations can be tremendously reduced by frequent veld fires. These potential changes in the native species populations, and the possible expansion of introduced ones, underline the importance of continued close study of pteridophytes in Southern Africa. Much work is still needed for a complete pteridophyte survey of Southern Africa, and more intensive collecting in the southern foothills of the Drakensberg seems particularly necessary. REFERENCES ACOCKS, J.P.H., 1975. Veld Types of South Africa. Bot. Mem. S. Afr. 40. HOLTTUM, R.E., 1974. Thelypteridaceae of Africa and adjacent Islands. J./.S. Afr. Bot. 40: 123- 168. PALGRAVE, K.C., 1977. Trees of Southern Africa. Struik, Cape Town. REVIEW FLORA OF CHIAPAS: PART 1 INTRODUCTION by Dennis E. Breedlove; PART 2 PTERIDOPHYTES by Alan R. Smith, 35 + 370 pp., 82 figs. 174 x 254mm. California Academy of Sciences, SanFrancisco. 1981. Prices: US $2.50; $30.00. Alan Smith states in his preface that pteridophytes account for 7-8 per cent of the vascular plant flora of this state in southern Mexico. He estimates that thanks to the extensive collecting carried out over the past 10 years or so possibly 95 per cent of the ferns there have been recognised. Altogether, there are 99 genera of ferns and five of fern allies known from the State, giving a total of 609 species (563 ferns and 46 allies). This flora no doubt has its constraints imposed by the editorial committee but is comprehensive and traditional giving details of type (including synonyms) bibliographical location of illustrations, a full description (detailed and precise as we have come to expect from Alan Smith's pen) habitat, distribution and list of species seen. What is untraditional and greatly welcomed is, at the end of each genus, species to be looked for in Chiapas! and where potential new but undescribed taxa are at hand they are indicated. The end of the book contains line illustrations of 106 species (of 90 genera). The genera of ferns are arranged alphabetically and the arrangement of species within is similar with the exception of Grammitis, Thelypteris, Polypodium and Trichomanes which are first divided into subgenera. Thus no descriptions of, or keys to, families are given and indeed mention of them at all it omitted. This is a bold, but in the reviewer's opinion, wise attitude. At 5 cents a species and the keys and generic descriptions thrown in for good measure what could be better. FERN GAZ.: 12(4) 1982 197 DICTYMIA BROWNII (POLYPODIACEAE S.S.), AN ANCIENT AUSTRALIAN FERN T.G. WALKER Department of Plant Biology, University of Newcastle upon Tyne C.N. PAGE Royal Botanic Garden, Edinburgh ABSTRACT An account of the ecology, morphology and anatomy of Dictymia brownii from Australia is given. The probable long-isolation of the genus within the Australasian region make its morphology and cytology of particular interest for detailed study. Chromosome counts of n = 35, 2n = 70 are recorded, together with a karyotype analysis. INTRODUCTION Material of a relatively poorly known member of Polypodiaceae,. Dictymia brownii (Wikstr.) Copel., was collected in Australia in 1969 by CNP and sent to TGW for cultivation in the fern collection at the Department of Plant Biology in the University of Newcastle upon Tyne. Although epiphytic in the wild these plants thrive in an ordinary fern mixture in clay pots with good drainage and are in excellent condition after 12 years of this treatment. The cytology of the plants had been worked out for some years and it was felt that a more complete account of this interesting species was desirable. ae om Ps Hx ie ~~ es Sf - id FM On ee FIGURE 1. Dictymia brownii in the wild. A moderate-sized clump growing epiphytically on the lower part of a slender trunked rainforest tree, growing beside a stream ina light-gap at the top of a waterfall, near Kandallila Falls, south-east Queensland. (Photo: C.N. Page). 198 FERN GAZETTE: VOLUME 12 PART 4 (1982) DISTRIBUTION AND ECOLOGY Dictymia is a small genus of approximately four closely-allied species, present in Australia, New Caledonia, Fiji, and perhaps New Guinea (Domin, 1913; Copeland, 1929: Tindale, 1972; Brownlie, 1969; Jones & Clemisha, 1976). Throughout this geographic area, all the species are slow-growing epiphytes or lithophytes in humid rainforest vegetation. Beyond these few facts, very little seems to have been reported in the literature on the ecology of any one of them. FIGURE 2. Typical Queensland rainforest, in which Dictymia browni/ occurs epiphytically on trees high in the forest canopy, near Lake Eacham, Atherton Tableland, north-east Queenland. (Photo: C.N. Page). Dictymia brownii of Australia is the most widely ranging of the species with the Fiji plant and one of the two New Caledonian species closely allied to it. Its ecology is thus probably fairly characteristic of that of the group. It occurs from at least the Atherton tableland area of North Queensland (lat. 17°S.) through the discontinuous areas of rainforest that follow the Great Dividing Range southward to southern New South Wales (to about lat. 34°S.). Throughout this range it is a fairly local and seldom gregarious species, growing epiphytically or on rocks. In dense rainforest it is usually confined to epiphytic sites on the uppermost parts of branches of the tallest canopy trees. In patches of more open rainforest, it extends down tothe lower branches of the crowns of broadleaved trees, or occurs on the upper parts of the trunks of palms (mainly on robust, tall L/vistona australis insouth-east Queensland), which often form groves around the streams in the rainforest. Dictymia occurs on rocks mainly where these are tall and outcropping in the rainforest, such as on rocky cliffs, or where rocky bluffs occur in the upper rainforest margins on mountains, and there is frequent cloud and high humidity. Nevertheless, the plant seems able to withstand a moderate amount of desiccation, at least for periods during the middle of the day, and its coriaceous frond texture is doubtless of advantage in enabling it to do this. With excessive drought the long edges of the fronds roll down and inwards, protecting the lower, stomatiferous surface of the blade. As is typical of most of the epiphytic ferns of the high rainforest WALKER & PAGE: DICTYMIA BROWNII 189 canopy (Page, 1979), Dictymia seems to be considerably light-demanding, and it is probably this factor which prevents it normally occurring at lower levels in the forest canopy, whilst it is probably its humidity requirement that confines it to moist forests. In appropriate epiphytic habitats, plants eventually build up rounded, compact, intertwined rhizome masses forming collars around their supporting branches, and these give rise to many dozens of slender spreading-ascending or arching-descending fronds. In established clumps, the great majority of fronds bear abundant, large, oval, sori, although the total numbers of spores released do not seem very high. Nevertheless, established plants are probably long-lived, and the fronds are retained for lengthy periods, until they are finally cleanly shed from abscission zones a little above the rhizome surface. The developing rhizome and root masses eventually trapa certain amount of detritus, and are frequently colonised by mosses and sometimes by other epiphytic ferns and orchids, although usually the Dictymia remains the dominant member of such clumps. These masses probably provide something of a small reservoir of moisture for the plant during drier spells. PHYTOGEOGRAPHIC AFFINITIES AND PROBABLE HISTORY Endemic as a genus to the Australian-Polynesian region, Copeland (1947) suggests the taxonomic affinities of Dictymia lie generally with Po/ypodium, with little further comment about its origin. We feel, however, that its various morphological differences from Polypodium suggest that the two must have been separated for some considerable time. In a phytogeographic and evolutionary analysis of the Australian pteridophyte flora (Page and Clifford, 1981), Dictymia appears to belong to that part termed the “long-resident’’ element, along with such other characteristically Australian or part- Australian fern genera as Blechnum, Doodia, Culcita, Dicksonia, Leptopteris, Todea, Platyzoma, Neurosoria, Paraceterach, and Pleurosorus. \ndeed, Dictymia is the only genus of the Polypodiaceae sensu stricto represented in this probably ancient Australian pteridophyte assemblage. The element contains an array of habitat-types, from streambank denizens of moist rainforest, to plants of drier forest margins and exposed rocky clefts, most of which are now confined largely to eastern Australia and adjacent larger islands. This element seems to have long been present within the island continent of Australia during its lengthy period of separation from the rest of the Old World, since the early Cretaceous break-up of eastern Gondwanaland (Powell, Johnson & Veevers, 1981). As a whole, where links exist at all, the element has more distant and older links with New Zealand, South Africa and South America. This pteridophyte element is dominated by warm temperate members, some with species now extending well into tropical zones, perhaps adapting to warmer conditions with gradual northward migration of the continent during its plate tectonic and subsequent climatic history. It seems very likely too that the New Caldeonian-Fijian members have also been long- resident in their areas, like their equally old but much more migrationally sedentary conifer vegetation (Page & Clifford, 1981; Whitmore & Page, 1981) and more ancient angiosperm families (Johnson & Briggs, 1981; Beadle, 1981), as fragments of former more widespread Gondwanaland ferns, now present as few and isolated fragments on drifting tectonic rafts. MORPHOLOGY AND ANATOMY The account which follows is based on the plants growing in cultivation at Newcastle upon Tyne. We are grateful to Professor E. Hennipman, Utrecht, who kindly sent some pickled prothalli and baby sporophytes to supplement our material. 200 FERN GAZETTE: VOLUME 12 PART 4 (1982) FIGURE 3. Dictymia brownii, a, whole fronds; b, close-up of underside showing large, orange- brown, oval sori. WALKER & PAGE: DICTYMIA BROWNII 201 1) Rhizome The creeping, green, frequently branched rhizome bears numerous shortly- spaced fronds and is covered by abundant scales which are attached sub-basally. These measure c.5mm x 1.5mm and are light brown, with an entire margin which is only interrupted at the base by a few, for the most part, 2-celled, short hairs. The cortex is parenchymatous only, the cells of which are conspicuously pitted, although the pits tend to be scattered over the cell wai! rather than grouped into discrete pit-fields as is the case in the lamina (see below). The endodermis is very prominent with well-defined Casparian strips present on the radial walls only. No phlobaphene is present. The most noteworthy feature of the 6 or so meristeles which are arranged ina single ring is the exceedingly prominent pericycle. This is 2 or 3cells wide, each of the cells being several times larger than the phloem cells abutting them towards the centre and two or three times radially wider than those of the endodermis towards the outside. Fig. 4a diagramatically depicts the rhizome in TS. FIGURE 4. A. TS rhizome (part of root in lower portion); B. TS base of stipe; C.TS mid-stipe; D. & E TS upper stipe, showing fusion of 2 strands; F. Cyclocytic stomium. A-Ex 15 mag., Fxc.290 mag. 202 FERN GAZETTE: VOLUME 12 PART 4 (1982) 2) Frond The salient features of the general appearance and habit of the fronds have already been described and Fig. 3 shows the conspicuous sori which are deeply impressed in a single row on either side of the midrib. In cultivation the fronds measure some 30cm x 1-1.5cm. In a very small number of cases the frond may fork once, as has been noted in many genera having simple fronds. a) The Stipe The stipe is very short, only 1-2cm long, and seated on a phyllopodium having a dehiscence layer of smaller cells. At the lower end, the stipe is approximately oval in cross-section, having an eccentrically placed vascular strand with two larger ones flanking it (Fig. 4b). Nearest the rhizome all the ground tissue is parenchymatous but a well-defined complete outer band of sclerenchyma very rapidly forms and this persists throughout the remainder of the stipe. Almost immediately above the dehiscence layer the stipe becomes more semi-circular in section (Fig. 4c). The mid strand approaches and fuses with one of the lateral strands (Figs. 4d & e) and the resultant two strands both produce small offshoots towards the base of the lamina which form the main veins. FIGURE 5. A. Venation, position of sori indicated by broken lines; B, C & D. TS frond at base, mid and upper points respectively. Limits of sclerenchyma indicated by inner unbroken line at midrib and margin; E. TS lamina showing lack of defined spongy and palisade layers. Ax 1.3 mag., B-D x 15 mag., E x 50 mag. WALKER & PAGE: DICTYMIA BROWNII 203 b) |The Lamina The veins form costal arches on either side of the midrib and fork and anastomose to form several rows of areolae which normally lack included veinlets. Not all the veins fuse with one another and there is an overall appearance of irregularity (Fig. 5a). In section the main vascular strand and the veins show the very prominent pericycle noted in the rhizome. The lamina is thick and fleshy and its resistance te desiccation in the wild has already been referred to. If a frond is detached and left on a laboratory bench several days pass before there are signs of wilting and this may no doubt be correlated with the anatomical features. Transverse sections emphasise the thickness of the lamina, this being some 14-16 cells wide (Fig. 5b). The mesophyll cells are rounded with abundant intercellular spaces. They are not differentiated into spongy and palisade layers and are uniformly rich in chloroplasts. The remarkable feature of these cells is the presence of very conspicuous pit-fields in the walls (Fig. 6a). In some cases a single pit- field occupies virtually the whole of a wall as in the illustration, whilst in others two or three may be present. The epidermal cells have thick radial walls which are often heavily lignified and have prominent plasmodesmata, especially evident on the lower epidermis. Both upper and lower epidermal layers are covered with a very thick cuticle and the individual cells also show pit-fields. The surface of the lamina is devoid of appendages of any sort. FIGURE 6. A. cells of the lamina mesophyll showing pit-fields; B. & C. Venation of first-formed fronds. A x 500 mag., B. & C. x 5 mag. (Photo: T.G. Walker). 204 FERN GAZETTE: VOLUME 12 PART 4 (1982) A layer of sclerencyma is present on the upper and lower side of the midrib and also at the margins of the lamina, being interrupted by the presence of stomata onthe lower surface. The epidermal cells are particularly heavily lignified at these parts. The stomata are confined to the abaxial surface (Fig. 4f) and are mainly cyclocytic. They have a mean length of 40 um (range 41 to 52.5 pm). Cyclocytic stomata were reported in only a few genera of Polypodiaceae by both van Cotthem (1970) and Sen & Hennipman (1981), these being Lemmaphyllum, Paltonium, and Lecanopteris. \n addition, Sen & Hennipman recorded this type of stomata as being characteristic of Dicranoglossum (Eschatogramme) and Niphidium. By contrast the large majority of the genera of this family have polocytic or copolocytic stomata. The sori are deeply immersed and contain numerous, long-stalked sporangia which lack glands and have a mean number of 14 indurated annular cells (range 12- 18). Copeland (1947) states that paraphyses are lacking but this is an incorrect observation, probably due to examining dried specimens. In sections of fresh material small 2 or 3-celled paraphyses ehding in a large colourless cell are to be seen. The spores are yellow, monolete, with a mean length of 57 um (range 41-67 um). The markings are very inconspicuous (Fig. 7) and consist of a shallow dimpling of the surface. If a fresh fertile frond is put in a spore packet and allowed to dry the normal thick deposit of spores which would be produced by most other ferns is absent. Microscopic examination shows that the sporangia have indeed discharged and it would appear that a very much higher than usual percentage of shed spores are trapped among the sporangia without falling free. This simple experiment has been repeated several times with the same result. We cannot say whether or not there is any biological advantage for the species in this — the net effect may be effectively to spread over a longer period the dissemination of the spores but this is pure conjecture. FIGURE 7. SEM of spores x 1000 mag. (Photo: T.G. Walker). WALKER & PAGE: DICTYMIA BROWNII 205 3) ~~ Prothalli We did not follow the early stages of prothallial growth but inthe mature samples available the margins of the prothalli tended to be frilly and the apical notch was frequently very shallow. No glands were seen on the margins of the prothalli unlike the situation in many other members of the family. Antheridia were produced at an earlier stage and then archegonia were also found in great abundance with their necks pointing backwards. 10 9 8 FIGURE 8. A. Root tip cell of Dictymia brownii, C4535, showing 2n = 70, x 1000 mag. Permanent acetocarmine squash preparation. B. Karyogram of above cell with chromosome lengths given in units. Each unit length equals 0.33 um. (Photo: T.G. Walker). 1 x 4 206 FERN GAZETTE: VOLUME 12 PART 4 (1982) The young sporophytes initially produce a single small frond c. 1.5cm long, the number and size increasing with age. At first the main vein produces one or two branches (Fig. 6b) and later these branches increase in number and anastomose to form the costal areoles (Fig. 6c). With increasing size the venation becomes more complex. CYTOLOGY Three plants, Page C4400 & C4401, collected in the Eungella Mountains, north Queensland, and C4535 collected on Mount Cordaux, south-east Queensland, each gave good preparations of root tip cells showing 2n = 70 (Fig. 7a). Meiosis showed n = 35. These numbers, coupled with the direct observation of functional archegonia on the prothalli demonstrate that Dictymia browni/ is a sexual diploid species based on x= 35. This basic chromosome number is not common in Polypodiaceae, having been recorded in only 5 out of the 34 genera for which cytological information is available. These are Crypsinus, Pleopeltis (including Lep/sorus and Microgramma) Marginariopsis, Belvisia and Polypodium (including Marginaria). It is worth noting that apart from the monotypic Marginariopsis all these genera contain species with other base numbers in addition to the x = 35. A further point of interest is that it will be seen that none of these genera figure in the short list of those that have cyclocytic stomata. Fig. 8b is a karyogram of the cell illustrated in Fig. 8a. The chromosomes are grouped according to length, each group differing from its immediate neighbour on either side by one unit which represents 0.33 pm. The total range is from 2.67 ym to 6.33 um with a peak at 4.0 pm. The centromere positions have been calculated following the system of Levan, Fredya and Sandberg (1965). In this system when the centromere is in the exact centre of the chromosome this position is designated the median point M and when at the end, the terminal point T. The chromosome between these two points is divided into 4 equal regions, these being in order from the median point, the median region m, the submedian region sm, the subterminal region st and the terminal region t. Because it is exceedingly difficult to be sure that a centromere is actually at the terminal point T and not fractionally below it, it is probably wisest to combine the figures for the terminal region and terminal point. The analysis for Dictymia is shown in Table 1. It will be noted that the overwhelming majority of chromosomes (60 out of 68, excluding the satellited ones) have their centromeres located in the distal half of a chromosome arm and have only 2 at the median point. This general trend is found in some other members of Po/ypodiaceae although often not to quite such a marked extent (Walker, in preparation). TABLE 1 Centromere position in the karotype of Dictymia browni Centromere position No. of chromosomes Median point, M 2 Median region, m Zz Submedian region, sm 4 Subterminal region, st 10 Terminal region, t 7 R = 50 Terminal point, T 24 Satellited 2 WALKER & PAGE: DICTYMIA BROWNII 207 REFERENCES BEADLE, N.C.W., 1981. Origin of the Australian angiosperm flora, pp. 407-426 in Keast, A. (ed.) Ecological Biogeography of Australia, W. Junk. The Hague. BROWNLIE, G., 1969. Flore de la Nouvelle-Caledonie et Dependances No. 3. Pteridophytes. Mus.Nat.Hist.Naturelle, Paris. COPELAND, E.B., 1929. Ferns of Fiji. 8B.P. Bishop Museum Bull. 59: 1-105. COPELAND, E.B., 1947. Genera Filicum. Chronica Botanica, Waltham. DOMIN, K., 1913. Beitrage zur Flora und Pflanzengeographie Australiens: Die farnflora Queenslands. B/b/. Bot. 20 (85): 1-238. JOHNSON, L.A.S., & BRIGGS, B.G., 1981. Three old southern families — Myrtaceae, Proteaceae and Restionaceae. pp. 429-469 in Keast, A. (ed.) Ecological Biogeography of Australia. W. Junk. The Hague. JONES, D.L., & CLEMISHA, S.C., 1976. Australian Ferns and Fern Allies. A.H. & A.W. Reed, Sydney. KEMP, E.M., 1981. Tertiary palaeogeography and the evolution of the Australian climate. pp. 33- 49 in Keast, A. (ed.) Ecological Biogeography of Australia. W. Junk. The Hague. LEVAN, A., FREDYA, K., & SANDBERG, A.R., 1965. Nomenclature for centromeric position on chromosomes. Hereditas 52: 201-220. PAGE, C.N., 1979. The diversity of ferns. An ecological perspective pp. 9-56 in Dyer, A.F. (ed.) The Experimental Biology of Ferns. Academic Press, London. PAGE, C.N., & CLIFFORD, H.T., 1981. Ecological Biogeography of Australian Conifers and Ferns. pp. 471-498 in Keast, A. (ed.) Eco/ogical Biogeography of Australia. W. Junk. The Hague. POWELL, C.A., JOHNSON, B.D., & VEEVERS, J.J., 1981. The Early Cretaceous break-up of Eastern Gondwanaland, the separation of Australia and India, and their interaction with South-East Asia. pp. 17-29 in Keast, A. (ed.) Ecological Biogeography of Australia. W. Junk, The Hague. SEN, U., & HENNIPMAN, E., 1981. Structure and ontogeny of stomata in Polypodiaceae. B/umea 272.175-201. TINDALE, M.D., 1972. Pteridophyta. pp. 38-84 in Beadle, N.C.W., Evans, O.D., & Carorin, R.C. Handbook of the Vascular Plants of the Sydney District and Blue Mountains, Sydney. VAN COTTHEM, W., 1970. Comparative morphological study of the stomata in the Filicopsida. Bull. Jard.Bot.Nat.Belg. 40: 81-239. WHITMORE, T.C., & PAGE, C.N., 1981. Evolutionary implications of the distribution and ecology of the tropical conifer Agathis. New Phytol. 84: 407-416. 208 FERN GAZETTE: VOLUME 12 PART 4 (1982) REVIEW FLORA MALESIANA SER. 2 PTERIDOPHYTA VOL 1 PART 5: THELYPTERIDACEAE by R.E. Holttum. pp. (17), 331-599. 1982. 240 x 164mm. The Hague, Martinus Nijhoff. ISBN 90 247 2652 2. Price DFI. 145/US $63/f£ This is the result of ten years’ research on the Old World species of Thelypteridaceae by the doyen of tropical fern taxonomy, Dr R.E. Holttum, whose experience in Malesian botany now extends over 50 years. In a dedicatory preface to Carl F.A. Christensen, the Holttum says, ‘There can be no doubt that existing ferns have originated through a process of evolution. They have therefore an inbuilt classification, and our object is to find it; the nearer we get to it the nearer we are to the practical aspect of taxonomy.” This indeed is Holttum’s philosophy as his publications, totalling over 100, on orchids, bamboos and gingers, and ferns in particular, show. This Dedication is an excellent account of the history of fern classification from which the reader can put not only Christensen’s but also of course, Holttum’s work into perspective. Thelypteridaceae is a large family and Holttum’s many papers on it over the past ten years are well known. Twenty-two genera are defined and in the taxonomic discussion, Holttum groups these into seven units of which 7rigonospora Holtt. (3 spp.) stands out in the family in having trilate spores and Cyclogramma Tagawa (1 sp.) is distinct in having hooked hairs on lower surface of all axes and on the sporangia (although similar hairs are found in Sect. Grypothrix of Pronephrium Holttum thinks they are in no way related. The name Cyc/osorus is confined to a complex of possibly three pantropic species of which C. interruptus (Willd.) H. Ito is the only Malesian species. 7he/ypteris similarly is restricted to the 7. pa/ustris Schott complex of which the Malesian representative is 7. confluens (Thunb.) Morton. The question has been raised as to whether one blanket genus (/he/ypteris auctores plures) can usefully and logically be divided into 22 (on a world basis) genera. Certainly it is useful, although one must recourse to checking microscopic characters (glands or hairs on sporangia; spore ornamentation; septate or unicellular hairs) to identify genera. Holttum himself admits Nannothelypteris (5 Philippine spp.) may be included in Pronephrium. The largest genus by far is Sphaerostephanos with 152 species in Malesia (with another 33 elsewhere); Pneumatopteris (54) and Pronephrium (57) are other big genera. Although the author has published extensively on this family (mostly in B/umea or Kew Bulletin) | counted 100 new taxa described in this work. | think it is incredible that Eric Holttum can manually (mentally) manipulate the characters, so carefully observed, and come up with the taxonomy he has. And the value of each character is weighed with his long experience of field work in the Malesian tropics. There are a lot of species known only from the type or a few gatherings. Obviously, as more material becomes available a reassessment will have to be made and Holttum would be the first to admit this. This fifth part ends volume 1 and has Corrigenda and Emendata (in which 3 species of Cyathea, 2 /soetes and one Gleichenia are mentioned as having been described since those genera were published in Flora Malesiana). To this one may add a further species of Gleichenia from Gunung Mulu, Sarawak (B.S. Parris, Bot. J. Linn. Soc., 84: 00; 1982). Thanks to Holttum, and Hennipman and Kramer who have contributed substantially to other genera, F.M. Series Il is truly underway. A.C. JERMY FERN GAZ. 12(4) 1982 209 THE ECOLOGY AND CYTOLOGY OF PHANEROSORUS (MATONIACEAE) T. G. WALKER Department of Plant Biology, University of Newcastle upon Tyne A.C. JERMY Department of Botany, British Museum (Natural History), London ABSTRACT An illustrated account of the specialized growth and habitat of Phanerosorus sarmentosus (Baker) Copel. is given. Achromosome count of n=25 is recorded and brief comparisons are made between Matoniaceae and Gleicheniaceae. INTRODUCTION Baker (1887) described some plants collected by Charles Hose from the cliffs of the Niah caves in Sarawak as Matonia sarmentosus Baker; he later (1891) amplified this brief description with line drawings showing the branching habit of the frond and soral details. In 1908 Copeland raised this species to generic rank as Phanerosorus sarmentosus (Baker) Copel. on the grounds of the sympodial branching of the long pendulous fronds (see Figs. 2 & 3), the anastomising veins around the sori and on indusial and sporangial characters. Its generic distinction is not disputed, although there is no doubt of its closeness to Matonia, but a further species P. major was described by Diels (1932) from limestone cliffs on the coast of Waigeu Island, NW New Guinea, solely on its more robust character. The present authors, as have others (e.g. . Alston in herb. BM) believe this to be conspecific with the Bornean plant. Although more familiar from herbarium material, Phanerosorus is little known as a living plant because of its limited distribution and its failure to become established in cultivation, no doubt due to its very demanding ecological requirements. Consquently, a brief illustrated account of the plant in the wild was felt to be of interest. DISTRIBUTION AND ECOLOGY It is of considerable interest that P. sarmentosus is found on the NW tip of New Guinea (Waigeu Is.) and then again in N Borneo. The other member of Matoniaceae (Matonia pectinata R. Br.) is confined to peninsula Malaysia, Sumatra and Borneo. Although Phanerosorus has no close fossil relatives, those of MVatonia show possible origins in the Triassic or early Jurassic Periods and across the present Northern Hemisphere, i.e. Laurasia (Seward 1922). Both genera are plants of open places — ridge-tops or cliffs — and have a growth habit and leaf structure capable of withstanding temporary drought conditions and, although today associated with the ever-wet tropics, may not always have needed such constant moisture. Field observations were made on spore dehiscence in Gunung Mulu N.-P. in late August towards the end of the drier regime (no true dry season was experienced in this ever-wet climatic belt). Many of the mature sori had shed entire indusia leaving naked sporangia free to disperse spores into the air. Atno time apparently does the indusium shrivel as in dryopteroid ferns. In many cases the entire sporangium is dropped and in material collected for the herbarium almost all sori dropped on drying leaving receptacular ‘stumps’ on the leaf surfaces as a sole reminder that the leaf had been fertile. It seems that dispersal of these heavier soral units (10—14 sporangia and indusium), whilst having advantages at the breeding level once gametophytes have been established, may prevent a wider dispersal of Phanerosorus. Similar behaviour of dropping whole mature sori in the field is seen in Matonia pectinata. 210 FERN GAZETTE: VOLUME 12 PART 4 (1982) FIGURE 1. The limestone cliffs on the Melinau Gorge, G. Benarat, G. Mulu National Park, Sarawak, seen from the west. FIGURES 2 & 3. Close up of Phanerosorus sarmentosus on G. Benarat, G. Mulu National Park, Sarawak. WALKER & JERMY: PHANEROSORUS (MATONIACEAE) 211 Within Borneo the species has been collected in Bau and at Bidi in the 1st Division, and at Niah National Park, near Miri, and in the Gunung Mulu National Park, in the 4th Division. In all of these localities as in Waigeu the plant grows on exposed ‘limestone cliffs from 50m to 800m above sea-level. Figure 1 shows typical limestone cliffs, 400 metres high on G. Benarat in the Melinau Gorge, G. Mulu N.P. where the species was Studied by the authors in 1978. The Phanerosorus was seenon the lower areas of cliff by the cave Lubang Rendah Harimau above the lowland forest which was growing on the limestone scree slopes. The cliffs of G. Benarat contain a number of endemic species characteristic of limestone in Borneo, such as Paraboea banyengiana B.L. Burtt and P. candidissima B.L. Burtt, both Gesneriaceae, as is a related genus Boea of which five species have been reported on the open limestone cliffs. Other species in this habitat include Ficus tinctoria Forst. f. subsp. g/bbosa (BI.) Corner and Fagraea auriculata Jack subsp. borneensis (Scheff.) Leen. more usually as epiphytes (Anderson & Chai, 1982) and Neonauclea peduncularis Walp. ex G. Don. Ferns such as Adiantum malesianum Ghatak, Nephrolepis hirsutula (Forst.) C. Presl, Pteris multifida Poir., and Taenitis cordata (Gaud.) Holtt. are found here. In more shady spots Hypodematium crenatum (Forsk.) Kuhn, 7Tectaria brooksii Copel. and 7. devexa (Kze.) Copel. were frequent. Another Gesneriad, Monophyllaea, a distinctive plant of limestone in G. Mulu N.P., occurred in similar situations. Seven species were recorded for the limestone here: /V. Johannis-winkleri Kraenl. and M. beccarii C.B. Clarke were those associated with Phanerosorus. Phanerosorus is widespread throughout the Melinau limestone of the Benarat- Api massif and seen again in quantity on the exposed cliffs above Gua Payau in the south of the range. In almost alt the localities it was difficult to reach without artificial aid. It often forms hanging thickets some five metres across in both directions. MORPHOLOGY The affinities of Phanerosorus with Matonia are clearly shown in the concentric vascular steles, the dichotomous or pseudodichotomous branching of the fronds, and the soral features. The sessile, or almost sessile, sporangia are grouped into circular sori which are protected by very characteristic thick, persistent, umbrella-shaped indusia to which the sporangia are frequently adherent. Compton (1909) described the branching of the fronds as being of two types; either both branches of a fork may develop more or less equally, giving rise to elongate axes which repeatedly fork, or one of the branches may develop more fully whilst the other branch after producing two pinnae ends in an aborted bud — this latter being the more usual method. These systems of branching and the production of resting and aborted buds shows great similarities to those seen in Gleicheniaceae. The similarities in gross morphology of the fronds of Matoniaceae and Gleicheniacea also extend to stomatal details. The stomata of both Matonia and Phanerosorus are anomocytic as are those of some members of Gleicheniaceae — the others being diacytic (van Cotthem, 1970). The gametophytes of the two families also show great similarities (Stokey and Atkinson, 1952; Atkinson and Stokey, 1964; Atkinson, 1973) although it must be confessed our knowledge depends on the study of only a very few species. The prothalli of Matonia closely resemble those of G/eichenia in being large with a very thick cushion up to 11 cells or so deep and in being long-lived with a ruffled margin. In both genera the antheridia are of the large complex type with many cells in the walls and the necks of the archegonia are much longer and straighter than in more advanced types. It was hoped to be able to supplement this information with a description of the prothalli of Phanerosorus and to this end spores were gathered in the field and sown 212 FERN GAZETTE: VOLUME 12 PART 4 (1982) immediately on return to Newcastle a few weeks later. None germinated and this may have been due either to very limited viability or to highly specialized requirements. Similarly attempts to transport young plants and establish them on a variety of substrates in cultivation were unsuccessful. Stokey and Atkinson (1952) commented that not only were the spores of Matonia short-lived but that germination was poor even when the spores were still comparatively fresh. CYTOLOGY AND GEOLOGICAL HISTORY The first cytological record for a member of Matoniaceae was that of Stokey and Atkinson (1952) who observed in sections of a prothallus of Matonia pectinata that a few cells in polar view showed c.26 chromosomes. They also made the observation that many of the chromosomes appeared to have terminal or near terminal attachments (centromeres). This number was precisely determined by Manton (1954a, 1954b) who demonstrated the presence of 26 bivalents at meiosis in a spore mother cell of this species. FIGURE 4. Meiosis in Phanerosorus sarmentosus (113643) showing 25 bivalents, x 1000. Permanent acetocarmine squash preparation. Vouchers: Sarawak, 4th Division, Gunung Mulu National Park, G. Benarat, Lubang Rendah Harimau / 700m alt. 23 Aug. 1978, Jermy 14466(BM, K, L, SAR, Herb. Walker). Fixations were made in the field in Sarawak on young plants of Phanerosorus collected near the entrance of one of the many caves on the south face of Gunung Benarat. Two plants gave excellent preparations of a number of cells, one of which Is illustrated in Fig. 4 and showing n=25 without any doubt. Thus, the two genera of Matoniaceae have basic chromosome numbers which differ from one another by one, namely x=25 and x=26. It cannot, of course, be established with certainty for how long this aneuploidy has existed nor which is the original and which the derived number but there are a number of suggestive facts. According to Copeland (1947) the morphological differences seen in Matonia and Phanerosorus have been found in Matoniaceous fossil genera in the Mesozoic, suggesting a probable long history as distinct genera. Of the two genera there can be no question but that Phanerosorus is WALKER & JERMY: PHANEROSORUS (MATONIACEAE) 213 the more specialized as regards habit, habitat, etc. Both Matoniaceae and Gleicheniaceae are recorded back into the Mesozoic (Boreau, 1970; Seward, 1922 and Holttum, 1954, considers that as the two families have existed together for this very long period one must go back at least to the early Mesozoic for a common ancestor. Cytologically Gleicheniaceae shows a wide range of basic chromosome numbers, each number being characteristic of a particular genus. Table 1 lists the basic chromosome numbers found in the two families and it will be seen that the numbers reported for Matoniaceae fall in the bottom end of the range found in Gleicheniaceae. TABLE 1: BASIC CHROMOSOME NUMBERS IN MATONIACEAE AND GLEICHENIACEAE Matoniaceae Gleicheniaceae Phanerosorus x=25 Gleichenia x=20, 22 Matonia x=26 Diplopterygium x=28? (n=56) Sticherus x=34 Dicranopteris x=39 Acropterygium x=43 ACKNOWLEDGEMENTS Part of the work of TGW was carried out whilst held on SERC research grant and further assistance for travel was given by the Research Fund of the University of Newcastle upon Tyne. Our thanks must also be recorded to the Sarawak Department of Forests for allowing the Royal Geographical Society to carry out a major survey of the Gunung Mulu National Park and to those in the support teams who made the field work possible. REFERENCES ANDERSON, J.A.R. & CHAI, P.K.K., 1982. The vegetation, in A.C. Jermy & K.P. Kavanagh (Eds.) Gunung Mulu National Park, Sarawak: a survey of its biota and environment Sarawak Mus. J. 30 (51) Special Issue no. 2 (in press). ATKINSON, L.R., 1973. The gametophyte and family relationships. /n A.C. Jermy, J.A. Crabbe and B.A. Thomas (Eds.) The phylogeny and classification of the ferns. Bot. J. Linn. Soc. 67, Suppl. 1: 73-90. ATKINSON, L.R. & STOKEY, A.G., 1964. Comparative morphology of the gametophyte of homosporous ferns. Phytomorphology 14: 51-70. BAKER, J.G., 1887. On a further collection of ferns from West Borneo, made by the Bishop of Singapore and Sarawak. J. Linn. Soc. (Bot.) 24: 256-261. BAKER, J.G., 1891. Asummary of the new ferns discovered or described since 1874. Annis. Bot. 5-304 -332: BOREAU, E., 1970. 7raité de Paleobotanique \V (1) Filicophyta. Masson et Cie, Paris. COMPTON, R.H., 1909. The anatomy of Matonia sarmentosa, Baker. New Phytol., 8: 299-310. COPELAND, E.B., 1908. New genera and species of Bornean ferns. Philipp. J. Sci. 3C: 343-349. COPELAND, E.B., 1947. Genera Filicum. Chronica Botanica, Waltham. HOLTTUM, R.E., 1954. Flora of Malaya, 2, Ferns. Government Printing Office, Singapore. MANTON, |., 1954a. Cytology of meiosis in Matonia. Nature, 173: 453. MANTON, |., 1954b. Cytological notes on one hundred species of Malayan ferns. /nR.E. Holttum, Flora of Malaya, 2, Ferns: 623-627. Government Printing Office, Singapore. SEWARD, A.C., 1922. A study in contrasts: The present and past distributions of certain ferns. J. Linn. Soc. Bot. 46: 219-240. STOKEY, A.G. & ATKINSON, L.R., 1952. The gametophyte and young sporophyte of Matonia pectinata R.Br. Phytomorphology 2(2,3): 138-150. VAN COTTHEM, W., 1970. Comparative morphological study of the stomata in the Filicopsida Bull. Jard. Bot. Nat. Belg. 40: 81-239. 214 FERN GAZETTE: VOLUME 12 PART 4 (1982) SHORT NOTE ASPLENIUM PULCHERRIMUM IN BORNEO Asplenium pulcherrimum (Baker) Ching (syn. Dava/lia pulcherrima Baker, A. billeti Christ and A. ca/cicola Tagawa) is a small fern of limestone rocks which is known from China, Taiwan and Vietnam (DeVol & Kuo, 1975). This species has recently been discovered much further south, on Gunong Api in Gunong Mulu National Park, Sarawak, during the 1977-78 Royal Geographical Society Mulu Expedition. (Gunong Api is, atc. 1750 metres, the highest limestone mountain between northern Thailand and New Guinea (Hanbury-Tenison & Jermy, 1979). It was collected by A.C. Jermy (as No. 14115, in BM and SAR) from ‘soil detritus in crevices in shade and full sun on limestone pinnacles in montane limestone forest:at 1200 m’. Figure 1 shows three plants from this gathering. os A pulcherrium has a narrowly to broadly triangular to ovate 3- to 4- pinnatifid lamina dissected into linear segments, each with a single sorus, and dull black stipes usually as long as or longer than thetamina. It is most distinct from the other species of Asplenium in Borneo, but is morphologically similar to A. coenobiale Hance (syn. A. fuscipes Baker) of China and belongs to the mainland Southeast Asian element of the Bornean fern flora. FIGURE 1. Asp/enium pulcherrimum from Gunong Api, Sarawak, Borneo. (Jermy 14775, BM). REFERENCES DEVEL, C.E., and KUO, CHEN-MENG, 1975. Aspleniaceae, in Flora of Taiwan Volume 7, Eds. Hui- lin Li, Tang-shu Liu, Tseng-chieng Huang, Tetsuo Koyama and Charles DeVol. Taipei. HANBURY-TENISON, R., and JERMY, A.C., 1979. The RGS Expedition to Gunong Mulu, Sarawak, 1977-78. Geographical Jour. 145: 175-191. B.S. PARRIS, Botany School, Cambridge. Present address: Royal Botanic Gardens, Kew. FERN GAZ. 12(4) 1982 215 OBSERVATIONS ON THE VENATION PATTERNS IN OPHIOGLOSSUM, BOTRYCHIUM AND HELMINTHOSTACHYS S. BHAMBIE and PARKASH MADAN Department of Botany, Kurukshetra University, Kurukshetra - 132119, India ABSTRACT The Ophioglossaceae have three living genera showing two diverse types of venation patterns: (i) open dichotomous (Botrychium and Helminthostachys) and (ii) reticulate (Ophioglossum). The family lacks any fossil history and none of the living taxa are so far reported to have intermediate stages between the two venation types. Vein unions have been reported to occur in the open dichotomous venation of Kingdonia, Circaester, Utricularia, Ginkgo, Adiantum, Pteris etc, and are said to be of phylogenetic importance. A detailed study of the venation pattern in the Ophioglossaceae reveals that anastomoses are completely lacking in some species of Botrychium, but are of frequent occurrence in He/minthostachys zeylanica. |Inall, five different types of anastomoses leading to vein-islet formation have been observed. Other features of interest are multiple anastomoses, vein approximations, unconnected veinlets and blind vein endings of extrusive and intrusive type. It is »concluded that ,Botrychium, Helminthostachys and Ophioglossum appear to form an ascending series in so far as the complexities of venation pattern are concerned. INTRODUCTION ’ Foster (1952, 1969), Carlquist (1961) and Hickey (1973) have emphasized the significance of venation patterns in systematics and phylogeny. Open dichotomous venation is considered primitive and reticulate derived from it. Inthe last three decades various types of vein unions leading to areole formation have been described in open dichotomous venation of Kingdonia (Foster 1959; Foster and Arnott, 1960), Circaeaster (Foster, 1966, 1968), Utricularia (Taylor, 1964; Subramanyam, 1969), Ginkgo (Arnott, 1959), Adiantum and Pteris (Nair and Das 1974a, b; 1977). Venationin the family Ophioglossaceae is interesting, showing two extremes, viz., open dichotomous in Botrychium and Helminthostachys, and reticulate in Ophioglossum, and there has been only limited previous work on the venation patterns in this fern family (Mittal, 1968; Panigrahi and Dixit, 1969). MATERIALS AND METHODS A list of taxa investigated in this study is includedin Table 1. Plants studied were fixed in FAA, preserved in 70% ethanol, and cleared in 5% NaOH to display the venation. In Botrychium and Ophioglossum, ten mature leaves or pinnules from each source were so treated. In He/minthostachys, 110 pinnae from different leaves were examined against strong light and only those portions of pinnae were cleared, where some anastomosis, approximations etc., were recognised. The terminology of Hickey (1973), with some modifications, was followed for leaf architecture in general. In Ophioglossum, observations on shape, size and absolute number of vein-islets, vein endings etc., were made according to Gupta (1961). OBSERVATIONS The sterile segment of leaf or tropophyll is simple and small in Ophioglossum, quite big and ternately compound in Helminthostachys and pinnately decompound in Botrychium. In both species of Botrychium studied, venation is perfectly open-dichotomous. By contrast, He/minthostachys shows a number of vein unions, approximations, blind vein endings and detached veins. A typical reticulate venation is met within qQ ST ET ee eS es ee ee eee rep) — JOIJEM + oe oe ‘ : = ee v ” jEMjapuey>y AOJEWINY) a/eUsaYy2 “OC ‘OL As " " " " “u uNnpe4yeq “uUl] Wnze/NI/704 “OC 6 N = uede a i = rf a i a ‘1yoiew| 4q Ja00H Wn}ejO/lad ‘Q*8 = = Oo " " u" " us IYJBLUYIEd ‘uuly] a/neaipnu ‘OQ 7 > * i An a on b2 dW ‘4Ieye> ‘uur Wwngiueysn) ‘QO 9 N nr ao a fi ne IYJeWIYIIEd "PIIIM Winauiwelb “Q °G O SsNOWwOJp JOIJEMS Zz ce juesqy’y juasqy -0|N912810;AdWwed a4!}Ug a|AWIS ‘jeEMjapueU>d| winjej}so2 winssojbo!1ydO "YD LL snowoipo}dwesd sasOwOo}seue YIM SUOI}E4I9S "uulq ea/ue/Aaz $]9|UIBA aZeWAjN OF dA E SNOWOOYIIP UadO JOUIW YIM SNONUIS a}eusd | indyy7esoy sAyoe}SOylUIWw/ay 53 a}euUIdi} SOWWIJALWUOS S}JA|UIBA aJeWI}/N O} dF rf >, 2 a1e119S-paqo7 01 dn jeqiulepy ‘uulq, Wwnsoul6nue] “g vA snowojpo}dwesd “qunyuL S]9JUIBA a}zeLUI}Zj;NUEd 0} d~ }uUasaid SnOWOJOYIIP uadO asOja-pseqo7 a}euuidig 1yJCWYSe| winjeusa} winiyaAsz0g “p Bo i ae ee eee ee ee jJAydodo.} jpAydodo.} 39943N0S /eynuuid /ajnuuid jo uibsew ajnuuid widO4 /U01}3998)|09 y.eeys aypung Ul QI4ApliAy uol}eUua/ |e48uUas) _ [yAydodosL jAydodosy jO 39e]|d exe} JO sWeN “ON 'S i ee ES SS ae eee er ee ee ee ee VWHINZD JSYHL NI NYSLLVd NOILVNSA YOrVW GNV ADO 1OHdyOW 1T1AHdOdOUL -L 31aVvl 216 BHAMBIE & MADAN: VENATION PATTERNS 217 Ophioglossum. The venation is camptodromous in pinnae of Botrychium and Helminthostachys and campyloreticulodromous in Ophioglossum (Table 1). An examination of 110 pinnae from different leaves of He/minthostachys zeylanica revealed that one or more anastomoses were present in as many as 76 (69.90%) specimens. Some interesting cases of vein approximations, blind vein endings and detached veins are described below. Anastomoses: Based on the nature of contributing veins and their course after union, anastomoses were grouped into 5 major types and a few subtypes thereof. The frequency of different types is shown in Table 2. Type 1 includes all the anastomoses formed bya short confluence of two veins of the same order, which separate after union. It has two subtypes. In subtype 1a the anastomising veins are inner branches of first order dichotomy of the same costa (fig. 1d) and in 1b the anastomosing veins come from first order dichotomies of two adjacent costs (fig. 1e). Type 2 is again characterised by the union of veins of the same order but they become confluent to the margin. The type has three subtypes. In type 2a (fig. 1f) the nature of the contributing veins is the same as in Subtype 1a. Type 2b is formed by the union of inner branches of a second order dichotomy of the same costa (fig. 1g) whereas in 2c the anastomising veins come off as outer branches of first dichotomies of adjacent costa (fig. 1h). TABLE 2 FREQUENCY OF OCCURRENCE OF VARIOUS TYPES OF ANASTOMOSIS INHELMINOTHOSTACHYS ZEYLANICA Type 1 2 & 4 5 Sub- Multiple types a b a b c a b a b anastomosis %age of occur- rence FS. 4S 42 9 6 21 9 33 9 1 22.5 Anastomoses Type 3 and Type 4 are formed by the union of veins of different orders (figs. 1i-l). In Type 3 the veins become separated after union but in Type 4 they proceed as a single vein. Both these types show two further subtypes. Subtype 3a is characterised by the union and subsequent separation of a branch from first order dichotomy and the contiguous acruate vein from the same costa (fig. 11). In 3b the contributing veins come from two different costa (fig. 1)). Type 4a (fig. 1k) is formed by the union of an arcuate vein of first order dichotomy with a branch of the contiguous vein from the same costa, wheras in 4b, the uniting veins are of next higher order (fig. 11). Type 5 is discernible as the areole formed by the anastomising veins possesses a blind vieniet (fig. 1n). In addition to the types described above, multiple anastomoses formed by the contribution of veins from a single costa (fig. 1g) or two different costae (figs. 1g, p, r). have been commonly observed. 218 FERN GAZETTE: VOLUME 12 PART 4 (1982) FIGURE 1, a-b, cleared pinnule of Botrychium ternatum and B. lanuginosum showing dichotomous venation; c-r, portions of pinnae from different leaves of He/minthostachys zeylanica. C, approximation to anastomosis type 1a; d-e, anastomosis types 1a and 1b; f, anastomosis type 2a and an extrusive blind vein ending; g, anastomosis type 2b and multiple anastomoses; h, anastomosis type 2c; i-j, anastomosis types 3a and 3b; k-|, anastomosis types 4a and 4b; m, approximation to anastomosis type 4; n, anastomosis type 5; 0, an approximation anda detached vein; p, multiple anastomoses; q, an intrusive blind vein ending; r, multiple anastomoses and a blind vein ending. The broken line in a-c shows the bundle sheath. Codes: a, anastomosis; ap, approximation; bv, blind vein ending; co, costa, d/-7, 1st order dichotomy; d/-2, second order dichotomy; dv, detached vein; ebv, extrusive blind vein ending; /bv, intrusive blind vein ending; ma, multiple anastomoses, mv, mid vein; sa, secondary areole. BHAMBIE & MADAN: VENATION PATTERNS 219 Approximations. At times, two veins in open dichotomous system deflect from their path to come nearer but do not show true union. They form intermediate stages between open venation and various types of anastomoses. An approximation leading to the anastomosis Type 1a is illustrated in fig. 1c. Approximations shown in figs. 1m and o are formed by blind veins. They represent the intermediate stages before the anastomosis Type 4. An arcuate vein of second order dichotomy and a branch from the contiguous dichotomy forming an approximation are shown in fig. 1r. Blind Vein Endings and detached veins. \ntrusive (figs. 1q, r)and extrusive (fig. 1f) blind vein endings relating to various orders of dichotomies seem not uncommon, and have twice been seen to intrude into an areole (fig. 1n). A few detached veins, mostly present near the margins of pinnae, were also observed (fig. 1p). O FIGURE 2. Tropophyll outlines and venation in Ophioglossum: a-b, O. reticulatum, c-d, O. petiolatum; e-g, O. nudicaule,; h-i, O. thermale; i-k, O. /usitanicum; |-m, O. oramineum; n-o, O. costatum showing primary areoles only; p, O. costatum, two primary areoles magnified to show secondary areoles. 220 0°69 Ol 0€c'0 L8L°0 8S2'0 OCE'L 0'L6 ” ” v " ‘OL Oe9 Oc LSS‘0 80¢°0 808'0 Ove'l O'8Z " ” u ” 6 0:09 0°SZ 6cE£ 0 €€c2'0 06Z2°0 OLZ L OSL u " “ ” 8 a1eaund = ajyeuouonw Aji ybi\s -asnigo 0°8S 0°02 v6C'0 c6L'O v6L'0 092" O'EL 40 aynoVy 0} ayNoVy 31eAO gjnesipnu ‘OC V/s a}yeudJoNW asnigo -asniqo 9}e|O90ue| 0°68 0'°6S cCL'0 LO0S‘O 060'L 8L60 £18 0} ayNoVy 0} a1Noy 0} 919d1))5 a/BUl4ay} “OC 9 UNG aje(noiune -asnigo 31PAO apIM 0922 SG OOL 980'0 S90'0 L6L°0 6S1°S O99LL 40 ayepsiOy 0} a}NoVy 0} BuojqQ =winjejnzi1a4 ‘CO G asnigqg © 691 919 8cL 0 060°0 LEE0 OOL'E €9cS 0} BINoVy ayNnoVy aLAQ = =wnjejoljad ‘CO v 9}e|Oa0ue|qQ ajyeaund = = ayeulwunoe oT) L8v O'8L 9Ev0 6EE0 ELV L 8v78'0 ELV 81NoV ‘andy = OIA JQ ‘AIEAQ. = WNIJUeIISN/ ‘CO a 9}e9und ae nua}}e d1Ndi}|9 GLC OC 860'0 860°0 evel LcL0 v0 Aj, yB1|S JO ayNoVy “MOEN = Wntuswes6 “CE v4 a}yeuosJONW asnigg a1eA0gO 0909 0'C62 09Z'0 619°0 c89'l v6S 0 O'V8e aye9uny 0} apnoy 0} 3}2AD usnjej1so2 ‘OC =I | S}e|SIUIe@A §=6shulpusuIaA aod ajOo1V7 jo ‘ou jo ‘ou uu sajOaly ul ool }O azis zu ul ajynjosqy ainjosqy sbuipuseula/) sLejula/, /sajooiy ebe1isaAy ease yea] eseq-jea} xede-jeo7y edeys-jea] exe] ‘ON IINVIIGNN “O AO STIAHdOdO¥L LN3434ddI0 GNV WASSO7T9D0/HdO 4O SalOddS LNAY3dddIG NI AYNLOALIHOYV TIAHdOdOdL €418Vvl BHAMBIE & MADAN: VENATION PATTERNS 221 LEAF ARCHITECTURE AND VENATION IN RELATION TO SPECIES OF OPHIOGLOSSUM In the species of Ophioglossum studied, minor variation in the leaf (tropophyll) architecture occurred sufficiently consistently to suggest that this should be recorded as possibly of diagnostic use between different species. In all seven species of Ophioglossum included in this survey, the leaf (tropophyll) is simple with entire margins. But it shows variations in shape and size, as well as base and apex form, from species to species (see Table 3 and figs. 2a, c, e, f, h, J, |, n). Major venation in all the species is palmate, for more than one primary vein of moderate thickness emerges from the base (figs. 2g, m). These veins follow a sinuous course and lose their identity well before the margins (figs. 2g, m) — hence they are reticulodromous. However, in O. costatum there are 2-4 prominent primary mid veins, one of which passes to the apex. Terminal veins just reach the margins (fig. 20). Primary areoles are formed by veins of 1° and 2°. Areoles formed in all the Species are large, and are more or less elongated in the basal axial region and are smaller (and more tetragonal to hexagonal towards the margins — figs. 2g, i, k)except in O. gramineum where all the areoles are elongated (fig. 2m). The venation is double in O. costatumi.e., secondary areoles are regularly present (fig. 2p). Secondary areoles though formed, are also not frequent in all the other species (figs. 2b, d, g, i, k), except in O. gramineum where they are rarely observed (fig. 2m). Areoles are very large in O. reticulatum and O. petiolatum, \arge in O. nudicaule and smaller in the rest of the species (Hickey, 1973). The absolute number of areoles shows an approximate increase with leaf area in an individual species. The number of vein-islets per unit area of leafs however, seems fairly constant for each species (Table 3). DISCUSSION The importance of venation patterns on phylogenetic studies has been emphasised by Foster (1959, 1961, 1966, 1968) in the case of some Angiosperms. However, in Angiosperms there remains a controversy (Foster, 1968) whether the open, dichotomous venation is an archaic character (Lam, 1959; Subramanyam and Nair, 1972), or a result of reversion from a more complex vasculature (Foster, 1968). Gymnosperms have retained in general, a primitive open dichotomous venation (Zimmerman, 1959). Arnott (1959) has cited various types of anastomoses in dichotomous venation of Ginkgo and stressed their pioneer role in reticulation. Potonié (1912) has suggested that the predominantly reticulate venation seen in mesozoic plants has developed from the open dichotomous venation prevalent in palaeozoic floras by progressive vein unions. This view is supported by Bugnon (1925), Bower (1935) and Eames (1936), who considered open dichotomous venation to be characteristic of primitive living ferns and traced transitional stages between the dichotomous and reticulate venation in juvenile and successive adult leaves of some taxa. Foster (1961b) states that reticulate venation patterns probably originated independently from the open dichotomous venation of various lower groups of Pteropsida. Hara (1964) has traced the origin of reticulate venation in Onoclea by progressive vein-unions in open dichotomous venation. Bierhorst (1971) has also reported some anastomoses in dichotomous venation in the families Gleicheniaceae, Lygodiaceae, Cyatheaceae etc., and similar observations have been reported by Nair and Dass (1974a, b; 1977) in Adiantum and Pteris. These studies thus are in favour that in contrast to reticulate venation, open dichotomous venation is primitive. The observations made here show that in Botrychium, which is said to be primitive, particularly regarding its sporophyte, there is an open dichotomous venation. In He/minthostachys, where the sterile lamina as well as fertile spike shows a step towards simplification, the presence of various types of anastomoses in nearly 222 FERN GAZETTE: VOLUME 12 PART 4 (1982) 70% of the leaves (with up to 12 anastomoses and 3 approximations in a single pinna and some areoles with intrusive veinlets) can be taken as signs of the initiation of reticulation. Botrychium, Helminthostachys and Ophiglossum thus form a progressive series with respect to their venation patterns. Anastomoses have been differentiated as Greek alphabets (Melville, 1959), 5 types (Foster, 1966, 1968), 4 types (Arnott, 1959) and various types (Nair and Das, 1974a, b & 1977). In the present study major types 1, 2 and 3, 4canbe compared with types 1, 2 and 4, 5 of Foster. However, the fifth type where the areoles have an included blind vein ending, has not been reported so far in the open dichotomous venation of any plant. Subramanyam and Nair (1972) have even expressed doubt on the occurrence of such a type in plants possessing open dichotomous venations. Systematic importance of the venation observed It seems clear from this study that the venation pattern in the Ophioglossaceae differs substantially at generic level. In Ophig/ossum, it is also of taxonomic significance even at specific level as the vein islet number in a unit area remains fairly constant for a species. Wherever venation features show some overlap between species, other characters of leaf architecture provide further specific diagnosis. In O. thermale and O. /usitanicum, for example, where there is an overlap in number of areole per unit area, the former species possesses a greater number of free veinlets which are usually bifid, compared with the latter. Thus the present study supports the views of the systematic importance of venation patterns in the Ophioglossaceae, in which other good . characters are often rather few. ACKNOWLEDGEMENTS We are grateful to Professors V. Puri and S.K. Pillai for going through the manuscript and R.S. Mehrotra for facilities. Junior author is thankful to C.S.I.R. for a junior fellowship. ARNOTT, H.J., 1959. Anastomoses in the venation of Ginkgo biloba. Am. J. Bot. 46: 405-411. BIERHORST, D.W., 1971. Morphology of vascular plants. The Macmillan Company, New York. BOWER, F.O., 1935. Primitive land plants. MacMillan & Co., London. BUGON, P., 1925. la dichotomie Cotyledonaire, carracttre ancestral. Bull. Soc. Bot. France. 72: 1088-1094. CARLQUIST, S., 1961. Comparative Plant Anatomy. Hold, Rinchart and Winston, New York. EAMES, A.J., 1936. Morphology of Vascular Plants,;MacGraw-Hill Book Company Inc. New York and London. FOSTER, A.S., 1952. Foliar venation in Angiosperms from an Ontogenetic standpoint. Am. J. Bot. 39: 752-766. FOSTER, A.S., 1959. The morphological and taxonomic significance of dichotomous venation in Kingdonia uniflora Balfour, F. et W.W. Smith. Notes Roy. Bot. Gard. Edinb. 23: 1-12. FOSTER, A.S., 1961. The phylogenetic significance of dichotomous venation in Angiosperms. Recent Advances in Botany 2. 971-975. FOSTER, A.S., 1966. Morphology of anastomoses in dichotomous venation of Circaeaster. Am. J. Bot. 53: 588-599. FOSTER, A.S., 1968. Further morphological studies on anastomoses in dichotomous venation of Circaeaster. J. Arnold Arbor. 49. 52-57. FOSTER, A.S., and ARNOTT, H.J., 1960. Morphology and dichotomous vasculature of the leaf of Kingonia uniflora. Am. J. Bot. 47. 684-698. GUPTA, B., 1961. Correlation of tissues in leaves. 1. Absolute vein-islet number and absolute veinlet termination numbers. Ann. Bot. (London) 25: 65-70. HICKEY, L.J., 1973. Classification of the architecture of Dicotyledonous leaves. Amer. J. Bot. 60: ICSD HARA, N., 1964. Ontogeny of the reticulate venation in the pinna of Onoclea sensibilis. Bot. Mag. Tokyo. 77: 381-387. LEVIN, F.A., 1929. The taxonomic value of vein-islet areas based upon a study of genera Barsoma, Cassia, Erythroxylon and Digitalis. J. Pharm. Pharmacol. 2: 17-43. BHAMBIE & MADAN: VENATION PATTERNS 223 MELVILLE, R., 1959. Leaf venation patterns and origin of Angiosperms. Nature, London 224: 121-125. MITTAL, P.L., 1968. Ferns and Fern allies of Rajasthan-lll: Hitherto Unrecorded species of Ophioglossum from Rajesthan. Bull. Bot. Surv. India, 70: 171-176. NAIR, N.C., and DAS, A., 1974a. Studies on the venation pattern in ferns. |. Anastomoses in Adiantum incisum Forsk. (A. caudatum sensu Bedd. non. Linn. pro parte). Bull. Bot. Surv. Ind. 75: 108-117. NAIR, N.C., and DAS, A., 1974b. Studies on the venation pattern in ferns. Il. Further observations on Adiantum incisum Forsk. Acta. Bot. Ind. 6: 148-153. NAIR, N.C., and DAS, A., 1977. Studies on the venvation pattern in Ferns IV. Additional observations on Pteris vittata Linn. (P. /ongifolia sensu Bedd. non Linn.). J. Ind. Bot. Soc. 56: 267-274. PANIGRAHI, G., and DIXIT, R.D., 1969. Studies in Indian Pteridophytes IV. The family Ophioglossaceae in India. Proc. National Inst. Sci. India 35: 230-266. POTONIE, H., 1912.* Grundlinien der ptlanzen morphologie im Litche der Palacontologie. Jena. SUBRAMANYAM, K., 1969. Dichotomous venation in leaves of certain Indian Utriculariales, Proceed. XI. International Bot. Congress Seattle (Abstracts) 212. SUBRAMANYAM, K, and NAIR, N.C., 1972. Some aspects of dichotomous branching in certain _ Angiosperms. Biology of Land Plant. Sarita Prakashan, Meerut (India): 90-99. TAYLOR, P., 1964. The genus Utricularia L. (Lentibulariaceae) in Africa (South of Sahara) and Madagascar. Kew Bull. 78: 1-245. ZIMMERMAN, W., 1959. Die Phylogenie der Pflanzen. Stuttgart. * Not seen in original. REVIEW ILLUSTRATIONS OF PTERIDOPHYTES OF JAPAN, edited by Satoru Kurata and Toshiyuki Nakaike with the cooperation of the Nippon Fernist Club. 220 x 305mm. Vol. 1, 628 pp. (1979); Vol. 2, 646 pp. (1981). Tokyo University Press. ISBN O 86008 289 X. Price 50 yen each vol. These magnificent books illustrated by full page line drawings by various artists, but all to avery high standard, and a habitat photograph are something that all booklovers and fern enthusiasts will want to own. They are beautifully bound in simulated leather with a folded map of Japan in a holder on each back cover. Maps showing the dot- distribution in Japan are given for each taxon. Apart from the latin name of the species concerned all the text is in Japanese. The drawings, which say a great deal, vary in their coverage. In some species, epidermes and stomata; T.S. of stipe and rhachis, scales and fertile leaf are drawn in close up. Some things are not drawn, however, which might have been, e.g. the sporangia and paraphyses in Polypodium virginianum and P. vulgare but as identification guides the figures say all that needs to be said. It is unfortunate that a few new. nomenclatural combinations are made in this work, as they may get overlooked. | sincerely hope the publishers will bring out an English edition; |am sure such an important work warrants it. A.C. JERMY 224 FERN GAZETTE: VOLUME 12 PART 4 (1982) SHORT NOTE THE APPEARANCE AND DISAPPEARANCE OF A DRYOPTERIS CARTHUSIANA COLONY The following observations were made as part of the detailed recording (since 1965) of the flora of Perivale Wood Local Nature Reserve, Greenford, Middlesex. Prior to 1974 no D. carthusiana (Vill.) H.P. Fuchs had been recorded in this 11 hectare area of ancient oak woodland & permanent pasture. In early 1972 some cans of fuel oil spilled into the feeder stream which supplied a small woodland pond and an area of wet woodland covering approximately 35 x 20 m. The vegetation of the area considered predominantly of Salix cinerea L., ssp. o/e/folia Macreight, Prunus spinosa L., Populus tremula L., Rubus fruticosus agg., Carex pendula Huds., Ajuga reptans L., Scutellaria galericulata L., Glechoma hederacea L., Impatiens glandulifera Royle, Epilobium angustifolium L. and the moss Plagiomnium undulatum (Hedw.) Kop., which formed carpets to 12 cm high in places. A few Dryopteris dilatata (Hoffm.) A. Gray also occurred. The soil of the area was quite different from that of the rest of the wood, with no distinct A, horizon and much higher pH (7 surface readings in 1973 gave a range of 6.9 to 7.5; and in 1976 the range was 6.8 to 7.3). During 1974 three small but partially fertile sporophytes of D. carthusiana were found within the oiled area. During subsequent years more appeared within, and only within, the oil flooded area. The vegetation retained nearly all of its pre-1972 components, but all except Plagiomnium undulatum were somewhat redueed in vigour, and bare soil patches appeared. The extend of the flooded area and its associated thick brown oil was mapped and marked on the ground for future reference. In 1975 17 discrete individuals were located, and one of the 3 original plants had grown 25cm high x 5Ocm. 1976 saw an additional 10 plants, 1977, 18 more and 1978, 30 more. This total of 78 plants was not exceeded, and in fact numbers started to decline rapidly from 1979, until by 1981 just a single small individual survived for all the plants their demise followed a similar pattern, with only one or two fronds being produced after several years of apparent vigour (with 4 — 6 fronds), and these dying early, with the stock decaying visibly during the following summer. Dryopter/s dilatata increased in numbers to 23 over this period, and to this day are healthy. The Perivale site has probably been flooded occasionally each year for decades, the only obvious difference in 1972 being the oil accompaniment. Enrichment of the site by water from an increasingly leaky canal may also have taken place, as the feeder stream runs, via a culvert under the canal. As the periphery of the area has become almost impenetrable in places since 1977, due to dense bramble and elm sucker growth, other individuals may have persisted, though some painful searches have not revealed any! Similar oil contamination of a site 1 km away was also colonized by D. carthusiana from 1973 to 1979. No plants were recorded there in 1981, and the decline followed a similar pattern to that in Perivale Wood. Likewise there was only a single oil spill. Has this rise and fall of the species been observed before? If so was any form of pollution or enrichment intimated? Perhaps a temporary competitive advantage wasd had by the fern? | would be pleased to hear from anyone who has observed anything similar. REFERENCE ROBERTS, K.A. and EDWARDS, P.J., 1974. The Flora of Perivale Wood Nature Reserve. London Naturalist 53: 34-51. P.J. EDWARDS, Royal Botanic Gardens, Kew. FERN GAZ. 12(4) 1982 225 A NEW CONCEPT OF DRYNARIOID FERNS SUBHASH CHANDRA Pteridology Laboratory, National Botanic Research Institute, Lucknow-226001, India ABSTRACT Based on accumulated morpho-anatomical evidence, two new tribes of the subfamily Drynarioideae are proposed; Drynarieae with the genera Drynaria, Photinopteris and Aglaomorpheae with Ag/aomorpha, Drynariopsis, Holostachyum, Merinthosorus, Pseudodrynaria,Thayeria. INTRODUCTION The drynarioid ferns are an assemblage of large epiphytes of the Polypodiaceae family forming a taxonomically fairly discrete group. They are distributed from Africa and China to Australia and the Pacific islands. All the drynarioid ferns are distinguished by their creeping, fleshy, densely scaly rhizomes, specialisation of some of the entire fronds or bases of all fronds for collection and retention of detritus, texture and venation of the laminae, abscission of segments or pinnae from the rachises but not of stipes from the rhizomes (non-articulated fronds), and the possession of foliar nectiferous glands as translucent spots. SYSTEMATIC POSITION OF THE DRYNARIOID FERNS Several authors have given different treatments to the group as its systematic position is concerned. Christensen (1938) placed all the drynarioid ferns in the tribe Pleopeltideae of his subfamily Polypodioideae under the Polypodiaceae. Ching (1940) included them in the tribe Phymatodeae of the subfamily Pleopeltioideae. Dickason (1946), Holttum (1947, 1954), Copeland (1947, 1960), Alston (1956), Pichi-Sermolli (1958, 1959, 1977), Mehra (1961) and Bierhorst (1971) amalgamated the tribes and subfamilies of Christensen (1938) and Ching (1940), and included the drynarioid ferns in the single large family Polypodiaceae near Microsorium. Nayar (1970, 1974) established a new subfamily Microsorioideae of the Polypodiaceae and included all the drynarioid ferns along with microsorioids in that new taxon. Recently, Crabbe et a/. (1975) proposed a different classification in which drynarioid ferns were treated as a subfamily Drynarioideae of Polypodiaceae with out any further subdivision. Ching (1978) treated the drynarioid ferns as a very natural and distinct family Drynariaceae. In the present study, the classification of Crabbe et a/. (19275) has been followed. CONSTITUENT GENERA In 1929, Copeland accepted five genera (Ag/amorpha, Merinthosorus, Photinopteris, Thayeria, Drynaria) and subdivided the genus Ag/laomorpha into six subgenera (Drynariopsis, Aglaomorpha, Psygmium, Dryostachyum, Hemistachyum, Holostachyum). Later Ching (1940) recognized nine genera Ag/laomorpha, Drynaria, Drynariopsis, Dryostachyum, Merinthosorus, Hemistachyum, Holostachyum, Photinopteris, Pseudodrynaria). Copeland (1947) and Ching (1978) accepted eight genera (Drynariopsis, Pseudodrynaria, Aglaomorpha, Holostachyum, Thayeria, Merinthosorus, Photinopteris, Drynaria). Crabbe, et a/. (1975) accepted five genera (Drynaria, Photinopteris, Merinthosorus, Aglaomorpha, Thayeria). These genera are NBRI, Research Publication No. 101 (N.S.) 226 FERN GAZETTE: VOLUME 12 PART 4 (1982) considered to be of possible generic or subgeneric rank, but Dryostachyum, Holostachyum, Drynariopsis, Pseudodrynaria are tentatively reduced to Aglaomorpha. Recently, Pichi-Sermolli (1977) accepted nine genera, namely, DOrynariopsis, Pseudodrynaria Dryostachyum, Holostachyum, Aglaomorpha, Thayeria, Merinthosorus, Photinopteris, Drynaria. Thus, for the first time the drynarioid group was recognized as a conspicuous natural group of epiphytes (Crabbe, et a/ 1975: p.147). POSSIBLE GENERIC INTERRELATIONSHIPS The drynarioid ferns are quite different from most other polypodiaceous ferns and have often been considered as one of the most controversial with regard to their interrelationship and evolution. Varied views on this have been expressed by Goebel (1928), Copeland (1929, 1960), Christensen (1938), Ching (1940), Holttum (1947, 1954), Nayar and Kachroo (1953), Nayar (1954, 1955, 1959, 1965), Zamora and Vargas (1973 a, b) and Zamora (1975). In some cases, views expressed seem to have been based ona limited number of species or on selected morphological characters. An attempt has thus been made here to prepare a view of the interrelationships supported by a totality of evidence from as many morphological features of the plants as possible. The attributes included here are palea; rhizome; vegetative frond including epidermal cells, stomata, hypodermis, mesophyll, venation, fertile frond including sorus, sporangium and spores; and gametophyte. The main details of several of these separate features have been published previously (see Chandra, 1978, 1979a, b, c, Chandra and Zamora, 1979), but the table attempts to draw a broad comparison between them, as a basis for the taxonomic and phylogenetic views derived (see Table 1). On the basis of the evidence accumulated it is concluded that the contrasting combination of distinct characters and the degree of affinity (Table 1) among the drynarioid ferns supports the establishment of two natural, readily recognizable and definable groups, viz., Drynaria Group (Drynaria, Photinopteris) and Aglaomorpha Group (Aglaomorpha, Drynariopsis, Holostachyum, Merinthosorus, Pseudodrynaria, Thayeria). The affinity among some genera is clear, in others it is not so obvious consequently | prefer to limit myself to subdividing the subfamily Drynarioideae only into the tribes without any further subdivision and have adopted for them the names of Drynarieae and Aglaomorpheae. The purpose of the splitting of the drynarioid ferns was to give these two morphological groups a clearer place in the hierarchy of taxa. . Drynarieae. S. Chandra, trib. nov. Rhizoma repente; frondes in dua serie alternate, stipitata; folia nidi forma typice preaesens; hydathodus nullus; venatio reticulata, typice carens venulis inclusis liberis; sori typice punctiformes, totam frondem occupantes. Typus: Drynaria (Bory, Pro sectione Polypodii in Ann. Sci. nat. 5; 464; 1825), J. Smith in J. Bot., London 4: 60; 1841). Genera: Drynaria (Bory) J. Sm., Photinopteris J. Sm. Aglaomorpheae S. Chandra, trib. nov. Rhizoma repente; fronds in serie una, sessiles; folia nidi forma absentia; hydathodus typice praesens; venatio reticulata, areolae cum libra venula inclusa; sori typice non punctiformes, ad partem superiorem frondium restricti tantum. CHANDRA: DRYNARIOID FERNS Typus: Aglamorpha Schott, Gen. Fil.: 20; 1834. 227 Genera: Aglaomorpha Schott, Drynariopsis (Copel.) Ching, Holostachyum (Copel.), Ching, Merinthosorus Copel., Pseudodrynaria (C.Chr.) Ching, Thayeria Copel. These data have also helped to delineate possible lines of evolution and to comment on the relative primitiveness/ advancement of the member of this alliance. These views are summarized in Fig. 1, in which the ieft-hand half presents an interpretation of the views of Copeland (1929, 1947) and the right-hand half the views proposed here, based on the above evidence. TABLE 1 Distinguishing Features of the two groups of Drynarioid ferns Attributes Drynaria group Aglaomorpha group Paleae Structure Frond Arrangement Nest leaves Foliage leaves Hydathodes Hypodermis -Stomatal Size Stomatal Frequency Epidermal Cell Size Venation Pattern Nectary Type Leaf Shape Sori Form and Distribution Sporangial Size Spore size Peltate, lanceolate or shield- shaped. Marginal glandular hairs with cap-like waxy secretion Fronds in two alternating rows on the dorsal surface of the rhizome Present except in Photinopteris Stipitate Absent on the lamina surface except in Photinopteris Absent in fronds except in D. sparsisora and Photinopteris Varies from 29 y x 38 p to 42 yx 54 yp Varies from 39 to 79 per mm Large Finely reticulate, free-included veinlets found only in few areoles except in Photinopteris Epicostular or Hypocostular Fertile and sterile alike except in Photinopteris Usually punctiform throughout the frond except in Photinopteris Varies from 200 yp — 400 p x 210 p — 380 ps Varies from 26 — 42 p) x 40 — 70 yp Peltate or basally attached, lanceolate or sheild-shaped or ribbon-like. Marginal glandular hairs but without cap-like waxy secretion Fronds in one row on the dorsal surface of the rhizome Absent as a rule Sessile except in Aglaomorpha pilosa Present as a rule Always present Varies from 27 yp x 32 p to 34 yx 42 5 Varies from 50 to 140 per mm Smaller Broadly reticulate, free- included veinlets present in all the areoles Always hypocostular Fertile frond more contracted than sterile one except in Drynariopsis and Pseudodry- naria Coenosoral or acrostichoid, only at the upper portion of frond except in Dryna- riopsis and Pseudodrynaria Varies from 200 — 240 p: x 240 — 390 p Varies from 26 — 36 p x 42 — 56y FERN GAZETTE: VOLUME 12 PART 4 (1982) 228 4 + re) %, 2 = = 2 x as &Y =) wn a % a a > z < S) < iS) Bae lat fs x 4 Ir = Qa , la «aS YU Zz a ‘Oo\ le Vo ca Mira a & 5 ee Ee a {e) 9) ro} < o\|\|n ce) < AY a\ leh 7 ¥ z = Ww O\ Vy SS bs y § = \\ 0" ~, (@) Vv rm) fer a fe) 8 fe x x NG % (Vp) IE Ss Neg 4 A w a oS > A ) a. lu D5 ge L 7.% Q = o* i ao 9 Na ° < Ee pu as % Zz. ‘a TW 25 ~ Ss a Y oO AN co WE FIGURE 1. Schematic representations of the interrelationships of the drynarioid ferns; left, interpretation of the views of Copeland (1929, 1947); right, proposed revised scheme based on the evidence presented here. ACKNOWLEDGEMENTS It is pleasure to record here my gratitude to Dr. Prescillano M. Zamora, Professor, Department of Botany, University of the Philippines, Diliman, Quezon City, Philippines, for constant encouragement and helpful suggestions. Sincere thanks are due to Dr. Pablo K. Botor, Professor, Department of European Languages, University of the Philippines, for Latin rendering of the tribe descriptions, and to Dr. T.N. Khoshoo, Director, National Botanical Research Institute, Lucknow, India, for providing facilities. REFERENCES ALSTON, A.H.G., 1956. The subdivision of the Polypodiaceae. Taxon 5: 23-25. BIERHORST, D.W., 1971. The Morphology of Vascular Plants. Macmillan Co., New York. CHANDRA, S., 1978. Morphology and Phylogeny of drynarioid ferns. Ph.D. thesis, University of the Philippines, Diliman, Quezon City (unpublished). CHANDRA, S., 1979a. Gametophyte morphology of (Polypodiaceae). Amer. Fern Jour. 69: 111-118. CHANDRA, S., 1979b. Taxonomic use of foliar epidermis and hypodermis in drynarioid ferns. Kalikasan: Philipp. J. Biol. 8: 211-226. CHANDRA, S., 1979c. Morphological groups within the drynarioid ferns and their phylogenetic interpretation. Abstract: Third Indian Geophytological conference, Lucknow, Dec. 8-10. CHANDRA, S., and ZAMORA, P.M., 1979. Spore morphology in relation to taxonomy of drynarioid ferns. Phytomorphology 29: 252-255. CINGH, R.C., 1940. On the natural classification of the ‘‘Polypodiaceae’’. Sunyatsenia 5: 201- 269. CINGH, R.C., 1978. The Chinese Fern Families and Genera: systematic arrangement and historical origin. Acta Phytotax. Sin. 16: 1-37. CHRISTENSEN, C., 1938. Filicineae, in Verdoorn, F., Manual of Pteridology. Martinus Nijhoff, The the ferns genus Drynariopsis Hague. COPELAND, E.B., 1929. The Oriental Genera of Polypodiaceae. Univ. Calif. Publ. Bot. 16:45-128. COPELAND, E.B., 1947. Genera Filicum. Chronica Botanica, Waltham, Mass., USA. COPELAND, E.B., 1960. Fern Flora of the Philippines 3: 377-555. CHANDRA: DRYNARIOID FERNS 229 CRABBE, J.A., JERMY, A.C., and MICKEL, J.T., 1975. A new generic sequence for the pteridophyte herbarium. Fern Gaz. 11: 141-162. DICKASON, F.G., 1946. A phylogenetic study of the ferns of Burma. Ohio J. Sci. 46: 73-108. GOEBEL, K., 1928. Morphologische und Biologische Studien XIll. Weitere Untersuchungen uber die Gruppe der Drynariaceen. Ann. Jard. Bot. Buitenz. 39: 117-169. HOLTTUM, R.E., 1947. A revised classification of leptosporangiate ferns. J. Linn. Soc. Bot. 53: 123-158. HOLTTUM, R.E., 1954. Ferns of Malaya. Vol. ||. The Ferns. Govt. Print Press, Singapore. MEHRA, P.N., 1961. Cytological evolution in ferns with particular reference to Himalayan forms. Proc. 48th Indian Sci. Congr. Pt. Il Section Botany, 130-153. NAYAR, B.K., 1954. Studies in Polypodiaceae Il: Contributions to the morphology of Pseudodrynaria coronans (Wall.) C. Chr. Phytomorphology 4: 379-390. NAYAR, B.K., 1955. The gymnogrammoid and the pleopeltoid ferns: Asummary. J. Univ. Gauhati 6: 47-62. NAYAR, B.K., 1959. Studies in Polypodiaceae VI: Further observations on the morphology of Drynaria Bory. J. Univ. Gauhati 9: 95-103. NAYAR, B.K., 1965. The gametophyte and juvenile leaves of the drynarioid ferns.Bot. Gaz. 126: 46-52. NAYAR, B.K., 1970. A phylogenetic classification of homosporous ferns. Taxon 19: 229-239. NAYAR, B.K., 1974. Classification of homosporous ferns Pt. Il: pp. 111-201. In: B.K. Nayar and S. Kaur, Companion to R.H. Beddome’s Handbook of the Ferns of British India. Chronica Botanica, New Delhi. NAYAR, B.K., and KACHROO, P., 1953. Studies in Polypodiaceae |: Contributions to the morphology of Drynaria Bory: D. quercifolia (L.) J. Sm. and D. propinqua (Wall.) J. Sm. Phytomorphology 3: 411-423. PICHI-SERMOLLI, R.E.G., 1958. The higher taxa of the pteridophyta and their classification, in Hedberg, O., Systematics of Today. Uppsala Univ. Arsskrift. PICHI-SERMOLLI, R.E.G., 1959. Pteridophyta, in Turrill, W.B., Vistas in Botany. Pergamon Press, London. PICHI-SERMOLLI, R.E.G., 1977. Tentamen pteridophytorum genera in taxonomicum ordinem redigendi. Appendix | et Appendix II. Webbia 31: 481-500. WAGNER, W.H., Jr., 1973. Some future challenges of fern systematics and phylogeny. Bot. J. Linn. Soc. 67; suppl. 1: 245-256. ZAMORA, P.M., 1975. Additional notes on the nectary-costule associations in drynarioid ferns. U.S.T., J. Grad. Res. 4: 31-33. ZAMORA, P.M., and VARGAS, N.S., 1973a. Notable variations in leaf forms of Drynaria. Philipp. Agric. 57: 55-71. ZAMORA, P.M., and VARGAS, N.S., 1973b. Nectary-costule association in Philippine drynarioid ferns. Philipp. Agric. 57: 72-88. REVIEW LIST OF THE TYPE SPECIMENS IN THE HERBARIA OF JAPAN. FERN FAMILIES by K. lwatsuki60 pp., 1981. Kyoto University. 257 x 18Omm. Price not given. The families listed are Lycopodiaceae, Selaginellaceae, Ilsoetaceae, Equisetaceae, Ophioglossaceae, Maratiiaceae, Osmundaceae, Plagiogyriaceae, Gleicheniaceae, Schizaeaceae, Hymenophyllaceae, Cyatheaceae, Dicksoniaceae, Dennstaedtiaceae, Lindsaeaceae, Davalliaceae, Oleandraceae, Parkeriaceae, Vittariaceae, Pteridaceae, Aspleniaceae, Blechnaceae, Lomariopsidaceae. Full details of the type locality, collector, number and the herbaria where deposited are given. Kinds of type (syntype, isotype etc.) are given and in many cases lectotypes have been carefully chosen from among the syntypes. When the holotype appears to be lost a neotype has been selected. A.C. JERMY 230 FERN GAZETTE: VOLUME 12 PART 4 (1982) SHORT NOTE MORPHOLOGICAL AND CYTOTAXONOMIC OBSERVATIONS ON AZOLLA PINNATA Of the six extant species of Azo//a (Hall and Swanson, 1968), A. pinnata R. Brown is distributed over large geographic areas. In a recent taxonomic revision of the A. pinnata complex, Sweet and Hills (1971) have segregated two varieties, pinnata and imbricata, exclusively based upon the morphological features of the vegetative organs of the sporophyte. They also observed that plants from India and Ceylon were of intermediate status between these two varieties. Thus their findings raise the question as to whether all the morphological features by which the varieties were recognized, are taxonomically reliable. ECOLOGY Azolla pinnata is an annual, free-floating fern. In the area under investigation, the young sporophytes, developed from embryos, can be found by October and the vegetative phase lasts for nearly 2 months. The sporocarp is formed well before the recession of water in early summer, whence the sporocarp-bearing plants undergo burrowing in dry, cracking soil, or are buried through agricultural practices in paddy fields. The total life-span of the sporophyte is thus about 5-6 months. This species has a normal sexual reproduction and, each population breeds freely for as many generations in a given locality as is permitted by inundation during the successive rainy seasons. MORPHOLOGY The morphological characters by which the two varieties were recognized by Sweet and Hills (1971) throughout the geographical range of the species are given in Table I. As a basis for this study, we have obtained plants from 20 different populations occurring over an area of 200 sq. miles of the Punjab Plains, northern India. From each population, 100 plants were sampled and analysed. From this large sample, the following account of the morphology of the plants was prepared: Plants 1-2 cm wide, main axis pseudodichotomous, 2-5 leaves before the first lateral branch. Dorsal leaf lobe non-imbricate in the proximal part, slightly imbricate towards the distal part, lamina generally flat or only slightly curled, apex sub-rounded, hyaline of irregular width comprising 2-4 layers of radially-aligned, elongate cells (in sharp contrast to nearly equi- dimensional in var. imbricata), minutely serrate or undulate, pubescent in the basal half only, length-width ratio: 1.1-2.5 (Plate (1A-C). A comparison of this data with that of Sweet and Hills, shows that none of the populations examined, though widely separated from one another, fully resembles either of the varieties /mbricata or pinnata. Also, our present data provides strong grounds for believing that most, if not all, of the features associated with the vegetative organs of the sporophyte are indeed plastic. Under experimental as well as natural conditions, we have found that plants show particular variation in having. a dominant vs non-dominant main axis, dichotomous vs pinnate branch pattern, degree of leaf imbrication and flat vs curled lamina. Each of these features can be modified by environmental factors, and especially by the degree of spacing of individuals in any population in the field. Plants around the peripheries of such populations, for example, often become more dispersed by wind currents and action of water birds, whilst those in the centres of colonies may remain densely-packed together. We have verified this experimentally. We have found that individuals isolated from compact populations, when put into well-spaced conditions become laxer in 231 e . Po C FIGURE 1, A-C, Azol/la pinnata: A, cleared dorsal leaf lobe showing marginal features; B,C plants from compact (B) and from sparse population (C) after isolation - note the laxer branches and loss of imbrication in the latter case; D,E, Salvinia auriculata (=S. molesta), plants from compact population (D) and after isolation (E); F, Azo//la pinnata, somatic metaphase from a leaf tip cell (fuelgen reaction stained) showing 2n=44 chromosomes, x1600; G, Sa/vinia natans, somatic metaphase from a leaf tip cell (resorcin blue stained) showing 2n=18 chromosomes, x 1900. 232 SHORT NOTES TABLE I: DISTINCTIONS BETWEEN VAR. P/INNATA AND /MBRICATA Character Var. pinnata Var. imbricata 1. Main axis Dominant Non-dominant 2. Branch pattern Pinnate appearance Dichotomous-like type 3. No. of leaves before 4-16 (Average 8) 2-8 (Average 4-5) the first lateral branch 4. Dorsal leaf lobe: a) Imbricate vs Imbricate to highly imbricate Non-imbricate to slightly non-imbricate imbricate b) Length-width ratio 1.4-2.1 eas) c) Flat vs curled Curled Flat d) Apex Acute Rounded to sub-rounded e) No. of hyaline cell layers Up to 4 2-5 f) Width of hyaline margin § Irregular Regular g) Shape of hyaline cells Elongate Nearly equidimensional h) Margin Serrate Smooth growth and eventually appear pseudo-dochotomous or even pinnate. Similarly, the degree of curling of the lamina, is also markedly influenced, whilst the imbrication of the dorsal leaf lobe found in proximal portions of plants, becomes virtually absent and is onlyu slightly expressed in distal regions (fig. 1B-C). The same is true for another free-floating fern, Sa/vinia auriculata Abul. (S. molesta Mitchell), examined by us (fig. 1D-E). CYTOLOGY Loyal (1958) reported that the gametic chromosome number of Azo//la pinnata was n=22. The presence of 2n=44 chromosomes in leaf tip cells of our material (fig. 1F) agrees with this. A comparison of the chromosome size in Azo/la and Sa/vinia, eg, S. natans (fig. 1G), seems to show a phylogenetic reduction within the Salviniales. Also, the correlation of reduced chromosome size with certain highly specialized features of Azolla eg, the megaspore apparatus, is consistent with the view of Moore (1968) that ‘small chromosomes are frequently encountered in more derived plants than are large chromosomes’. The heterosporous genus Se/agine//a appears to present a remarkably similar correlation of reduced nuclear state and evolutionary specialization among the extant lycopods (Stebbins, 1966). REFERENCES : HALL, J.W. and SWANSON, N-P., 1968. Studies on fossil Azo//a: Azo/la montana, a cretaceous megaspore with many small floats. Amer. Jour. Bot. 55: 1055-1061. LOYAL, D.S., 1958. Cytology of two species of Salviniaceae. Curr. Sci. 27. 357-358. MOORE, D.M., 1968. The karyotype in taxonomy. In Modern methods in plant taxonomy. Edit. V.H. Heywood. Acad. Press, London and New York. STEBBINS, G.L., 1966. Chromosomal variation and evolution. Science 152: 1463-1469. SWEET, A. and HILLS, L.V., 1971. A study of Azo//a pinnata R. Brown. Amer. Fern Jour. 617: 1-13. D.S. LOYAL, A.K. GOLLEN and RAMAN RATRA, Botany Department, Punjab University, Chandigarh-160014, India. FERN GAZ. 12(4) 1982 233 FIELD OBSERVATIONS ON THE NECTARIES OF BRACKEN, PTERIDIUM AQUILINUM, IN BRITAIN C. N. PAGE Royal Botanic Garden, Edinburgh ABSTRACT Although bracken, Pteridium aquilinum L., is a widespread weed of economic importance in Britain, little attention has been paid to the occurrence of nectaries upon it and their possible biological significance. Field observations are given of the structure, distribution, activity and variation of bracken nectaries as seen in Britain and some views of their possible taxonomic and ecological significance suggested. INTRODUCTION The presence of nectaries on the fronds of bracken, Pteridium aquilinum L.,. was recorded as long ago as 1877 by Francis Darwin. In addition to bracken, a very few other, unrelated, ferns in other habitats and parts of the world have also been reported to have nectaries (see, for example, observations and comments made by Schremmer (1969), Holttum (1968) and Paterson (1982) ). These nectaries seem somewhat similar to the ‘extra-floral nectaries’ known more widely in flowering plants (eg Bentley, 1977). Further, Tryon (1941), Schremmer (1969) and Lawton (1976) have all also encountered the presence of arthropods, especially on young bracken fronds in association with these nectaries. However, despite these reports, remarkably little further attention seems to have been paid to their significance or their biology. This paper reports some preliminary observations concerning the distribution of nectaries on the frond, their variation with habitat, time of functioning and morphological structure. Their occurrence on different forms of bracken has been studied on herbarium material on a world basis. Further a comparison is made between bracken nectaries and those of other members of the Hypolepidaceae and some views about the possible biological significance of these nectaries are discussed. OBSERVATIONS Constancy and distribution of nectaries Examination of bracken fronds in Britain, through a wide range of localities and habitats from central Scotland to southern England, suggests that nectaries are to be found inall areas. Indeed, on all fronds, multiple nectaries of different sizes are present in the same characteristic position, but the degree to which these are developed varies somewhat with habitat (see below). The main nectaries occur as sub-opposite pairs at the points of junction of the top of the stipe and the base of the midribs of the first pair of pinnae, on the under surface of the bracken frond. Thereafter, further sets of nectaries occur in similar positions at the junctions of each subsequent pair of pinnae with the rachis. The nectary pairs typically become progressively smaller towards the upper end of the frond, as the pinnae diminish in size. A second set of nectaries occurs along the length of each pinna midrib, also on its underside. These occur at the junction of each pinnule midrib with the pinna midrib, thus mirroring in more reduced form, the positions in which they occur along the main frond axis. The total number of nectaries present on a single large bracken frond may thus be quite large, frequently more than fifty. External appearance and anatomical structure of nectaries All the nectaries seem to be of generally similar external appearance. The ones 234 FERN GAZETTE: VOLUME 12 PART 4 (1982) situated on the pinna-rachis junction are usually the largest and most obvious ones, often 2-3 mm in diameter forming rounded, tear-drop or shield-shaped shining bald patches. When fresh, they are flat, slightly raised or somewhat bulging, and have a glabrous and minutely puckered, clear pale-green coloured surface, with a slightly translucent quality. They usually seem at their most conspicuous on fresh, young, expanding croziers in spring, when the rest of the surrounding areas of stipe and rachis are covered in a brownish-white pubescence (see Figs. 1 and 2). They become less distinct (though are usually still detectable) as the hairs are shed from the Surrounding areas on the fully expanded frond. In spring and early summer, one or more minute, clear, colourless slightly sweet tasting droplets can often be found exuding from some of these areas. Anatomical investigations show the nectaries to be composed of small, isodiametric, parenchymatous cells, overlain by a particularly thin epidermis. These cells abut inwards on to parts of the stele, and merge into the vertically elongated cells of the cortex of the stipe and rachis. These small, parenchymatous cells, often with fairly dense protoplasmic contents, seem characteristic of secretory parenchyma as has been described in several flowering plants (eg Esau, 1965). Variation in nectary development with habitat Bracken colonies in Britain are typically most extensive in open areas of deep, well- drained, sandy soils, occurring through a wide range of altitude (eg Taylor 1980, Page 1982b). Less dense stands occur in a wide range of open, sandy woodlands and woodland margins, which before man’s interference were probably their main original habitats (Page 1982a). Field observations in Britain now show that although the position of nectaries is constant, their size and prominence varies with habitat. In some sites, poorly developed main nectaries can be less than 1 mm in diameter and nearly flush with the surface. In others, the main nectaries may exceed 5 mm and protrude considerably (by about 1 mm or more). Bracken in woodlands often has the least developed nectaries, whilst fronds in more open habitats frequently have the more prominent structures. The most conspicuous nectaries of all have been found in some sunny woodland margin sites, as well as in several open, sandy, habitats at low altitudes near the sea. In contrast, nectaries seem to show remarkably similar degrees of overall development in different fronds in any one site. Thus there seems to be a strong habitat correlation. It is hoped that a planned programme of transplant experiments may indicate how much of this variation is genetically determined, how much is environmentally induced, and what environmental correlations exist. Seasonal variation in nectary activity In the following observations, only those nectaries from which such sweet tasting droplets have been found to be exuding have been regarded as ‘active’. In all the bracken stands sampled for this purpose (principally in central and south-western Scotland and northern England), nectary activity occurs during the period of emergence of the fronds in late-spring to early summer. Such activity seems to be confined to this period, ceasing totally once the frond is fully expanded. Observations suggest that the nectaries mature and are active in succession, from the base of the frond upwards and outwards along each pinnule, with an overlap in the activity period between successive nectaries. Further, each main nectary pair appears to be active only during the period of expansion of its adjacent pair of pinnae, and to cease exudation and dry-up for good once these adjacent pinnae have become fully unrolled. Field estimates for the active life of each main nectary varied from about 6 to PAGE: PTERIDIUM AQUILINUM IN BRITAIN 235 FIGURE 1. Expa nding fronds of Bracken, Pteridium aquilinum, with active nectaries. Cumberland, early June (Photo C.N. Page). FIGURE 2. Expanding bracken fronds showing clearly the position of one of the paired nectaries adjacent to the lowermost pinnae. at this stage the nectary is about to become active. Cumberland, early June (Photo C.N. Page). 236 FERN GAZETTE: VOLUME 12 PART 4 (1982) 10 days, with the overall period of activity on one frond lasting probably 4 weeks or more. Daily variation in nectary activity There is considerable day-to-day variation. In general, the most regular nectary activity has usually been observed in the early morning, when bulging nectar droplets have sometimes been found to be present on almost every frond in a stand. These droplets have presumably accumulated slowly during the night so they are particularly apparent on cool, still, moist, dewy mornings. Their subsequent size diminishes with the increasing rate of evaporation. Daytime observations on nectary visitors Observations in Scotland and the north of England, made during the period of frond expansion, have shown that a considerable number of different insects are attracted to bracken nectaries. Passing insects generally seem to be most attracted to those nectaries which have obvious secretion of nectar. Very frequently, as fronds become more fully expanded, brown marks and scars can be found on the older nectary surfaces, presumably resulting from over-enthusiastic rasping or probing by insects. Ants have been observed to be very regular visitors to bracken nectaries in many sites. Additionally, small numbers of various flying insects (especially flies or beetles) have been seen flying directly to nectaries, and sometimes remaining there for many minutes. Observations on Pteridium from other parts of the world Herbarium material of Pteridium from most of its world range has been studied. However, as most herbarium material tends to be collected in mid to late summer after the fronds have fully expanded, the nectaries are much less distinct. This is accentuated by the necessary drying for preservation. Two other practical difficulties are encountered: Pteridium is usually extraordinarily poorly represented in most collections and when it is preserved, the pinna base and rachis are often absent. It is probably the combination of these factors which has made the presence of nectaries in dried Pteridium fronds so largely overlooked. Nevertheless, wherever sufficiently well-preserved herbarium material has been found, it has been almost invariably possible to show that nectaries were present. Specimens from northern and central Europe, Japan, China, Himalayas, Malaya, Australia, New Zealand, central and southern Africa, eastern and western North America, West Indies, Central and South America show nectaries. Furthermore, there is also evidence of frequent physical damage to nectaries in herbarium specimens. This was presumably the work of insects while the fronds were alive. These observations are supported for central Europe by Schremmer (1969) and for North America by Tryon (1941) and Victorin (1923). They also observed the fronds of local bracken stands to be visited in the spring by ants. It would be valuable to have further field evidence of nectary activity in expanding fronds of Pteridium from other parts of its range. Observations on juvenile Pteridium Fronds on most juvenile bracken plants look so totally unlike adult bracken that they could easily be mistaken for some other genus of fern. Their usually softly pubescent appearance, elongate frond shape, rounded segments and almost indefinite tip growth (see Page, 1982b), seem much more like specimens of tropical Hypo/epis than those of adult Pteridium. Nevertheless even at the most early stages of sporeling development, small nectaries are regularly present in the same axillary positions as on adult plants. PAGE: PTERIDIUM AQUILINUM IN BRITAIN 237 Observations on other near-allies of Pteridium Evidence of chromosome numbers as well as similarity of juvenile forms indicate the principal near-allies of Pteridium are most probably Hypo/epis and Paesia, together constituting the fern family Hypolepidaceae (Page, 1976). Examination of both living and herbarium specimens of Hypo/epis and Paesia has failed to show any structures which could be regarded as nectaries. Therefore it seems likely that nectaries are confined to Pteridium in the Hypolepidaceae. Their presence thus seems to provide a useful diagnostic character for this genus, even in juvenile material and raises the question of their possible significance in relation to this genus’ exceptional ecological success. Comparison with cyanogenic properties of bracken stands Research on bracken in Britain in recent years has shown that this plant can possess fronds which are cyanogenic and that stands of bracken differ in the percentage of cyanogenic and non-cyanogenic fronds present, even over quite small distances (eg Cooper-Driver & Swain, 1976; Cooper-Driver, 1978; Balick et al 1978). Field observations were made in an area of Kirkcudbright where both types of bracken were known to exist (Dyer, pers. comm.) Bracken fronds were tested for cyanogenic reaction (Jones, 1966) and for nectary activity. Nectaries were regularly found to be both present and active on cyanogenic as well as non-cyanogenic fronds. The presence of active nectaries thus does not seem to be merely an alternative defence mechanism of young croziers which are lacking in cyanogenic properties. It seems to be a phenomenon of bracken populations generally. POSSIBLE BIOLOGICAL SIGNIFICANCE OF NECTARIES IN BRACKEN It is known that bracken fronds contain large numbers of toxic substances (eg Cooper-- Driver, 1976). Lethal effects resulting from consumption of bracken fronds by such large herbivores as sheep, horses, pigs and cattle have been widely demonstrated by agricultural and veterinary scientists (eg |.A. Evans, 1976, 1982, W.C. Evans, 1976, Evans & Patel, 1972, Jarrett, 1982). There is evidence too, that the presence of some of these substances, eg tannins and cyanides, may actively deter grazing animals, including insects (eg Cooper-Driver & Swain, 1976, Cooper-Driver et al, 1977, Grierson, 1979). There are indications too, from the above literature, that the levels of some of these toxic principles are greatest when the fronds are fully expanded. It has been suggested that ants attracted to the fronds by nectaries play a directly defensive role in protecting the succulent young fronds against herbivores. Lawton (1976: 205-6) summarised this as his “set a thief to catch a thief” principle, although he recorded finding no differences in average numbers of bracken-specific herbivorous arthropods on fronds with ants present compared with those on fronds without them. The observations made here also indicate that the ant numbers on fronds can frequently be few and often sporadic even at the most active nectary stages. | want to suggest another strategy which might accrue from the presence of nectaries. By having fronds which produce such abundant nectar so early in the growing season, bracken could positively encourage ants to establish nests in its immediate vicinity. This could create frequent ground-patrols of ants throughout the whole area of bracken colonies, providing a strongly discouraging effect upon establishment of potential fern-herbivores such as slugs and snails or even against competing other plants. It seems possible that slugs and snails in particular might otherwise thrive beneath the dense, slowly-decaying litter layer of fronds that 238 FERN GAZETTE: VOLUME 12 PART 4 (1982) FIGURE 3. Some insect visitors seen at bracken nectaries; and and click beetle (above) and seven- spot ladybird and a fly (below). (Photos C.N. Page & Susan Paterson). PAGE: PTERIDIUM AQUILINUM IN BRITAIN 239 accumulates annually. Such a strategy might help explain the particular success of bracken ecologically in warm, deep, sandy soils, so conducive to certain terrestrial ants. It is hoped to initiate further research at Edinburgh on these biological aspects of this ecologically unusually successful fern. ACKNOWLEDGEMENTS | am grateful to Miss S. Arnot and Miss S. Paterson, for their observational help in the field, thereby considerably broadening the basis of this study. | am grateful too for valuable discussion of points arising by Dr A.F. Dyer, D.M. Henderson and Dr T.G. Walker, and to Miss D. Masarati for reading the manuscript. REFERENCES BALICK, M.J., FURTH, D.G. & COOPER-DRIVER, G., 1978. Biochemical and evolutionary aspects of anthropod predation on ferns. Oecologia 35: 55-89. BENTLEY, B.L., 1977. Extrafloral nectaries and protection by pugnacious bodyguards. Ann. Rev. Ecol. Syst. 8: 407-427. BRAID, K.W., 1959. Bracken — a review of the literature. Harley: Commonwealth Agricultural Bureaux. COOPER-DRIVER, G., 1976. Chemataxonomy and phytochemical ecology of bracken. Bot. J. Linn. Soc. 73: 35-46. COOPER-DRIVER, G., 1978. Insect fern associations. Ent. Exp. & Appl. 24: 310-316. COOPER-DRIVER, G. & SWAIN, A., 1976. Cyanogenic polymorphism in bracken in relation to herbivore predation. Nature 260. 604. COOPER-DRIVER, G.A., FINCH, S., SWAIN, T. & BERNAYS, E., 1977. Seasonal variation in secondary plant compounds in relation to the palatability of Pteridium aquilinum Biochem. Syst. Ecol. 5: 177-193. DARWIN, F., 1877. On the glandular bodies on Acacia sphaerocephala and Ceropia peltata serving as food for ants, with an appendix on the nectar-glands of the common bracken fern, Pteris aguilina. J. Linn. Soc. Bot. 75: 398-409. ESAU, K., 1965. Plant Anatomy. 2nd edition. Wiley: New York. EVANS, |.A., 1976. Relationship between Bracken and cancer. Bot. J. Linn. Soc. 73: 105-112. EVANS, |.A., 1982. The carcinogenic, mutagenic and teratogenic toxicity of bracken. Proc. Roy. Soc. Edin. 878: (in press). EVANS, W.C., 1976. Bracken thiaminase-mediated neurotoxic syndromes. Bot. J. Linn. Soc. 73: t3-131. EVANS, W.C., & PATEL, M.C., 1982. Acute bracken poisoning in homogastric and ruminant animals. Proc. Roy. Soc. Edin. 878: (in press). GERSON, U., 1979. The associations between pteridophytes and arthropods. Fern Gaz. 72: 29-- 45. GLIESSMAN, S.R., 1976. Allelopathy in a broad spectrum of environments as illustrated by bracken. Bot. J. Linn. Soc. 23: 95-104. HOLTTUM, R.E., 1968. Flora of Malaya. Vol. Il Ferns. Govt. Printer, Singapore. JARRETT, W.F.H., 1982. Bracken and cancer. Proc. Roy. Soc. Edin. (in press). JONES, D.A., 1966. On the polymorphism of cyanogenesis in Lotus corniculatus L. 1. Selection by animals. Canad. J. Gen. & Cytol. 8: 556-567. LAWTON, J.H., 1976. The structure of the arthropod community on bracken. Bot. J. Linn. Soc. 2s 187-216. PAGE, C.N., 1976. The Taxonomy and phytogeography of bracken — A Review Bot. J. Linn. Soc. 23.-1-34. PAGE, C.N., 1979a. The diversity of ferns. An ecological perspective. pp. 9-56 in Dyer, A.F. (Ed.). The Experimental Biology of Ferns. London: Academic Press. PAGE, C.N., 1979b. Experimental aspects of fern ecology. pp. 551-589 in Dyer, A.F. (Ed.). The Experimental Biology of Ferns. London: Academic Press. PAGE, C.N., 1979c. Macronesian Heathlands. pp. 117-123 in Specht, R.L. (Ed.). Heathlands and Related Shrublands of the World. The Hague: W. Junk. PAGE, C.N., 1982a. The history and spread of Bracken in Britain. Proc. Roy. Soc. Edin. 878: (in press). PAGE, C.N., 1982b. The Ferns of Britain and Ireland. Cambridge: Cambridge University Press. 240 FERN GAZETTE: VOLUME 12 PART 4 (1982) PATTERSON, S., 1982. Possible ant-involvement with several Tropical Rainforest Ferns. Fern Gaz. 12: 243-245. SCHREMMER, F., 1969. Extraruptiale Nectaries. Beobachtungen an Salix eleagnos Scop. und Pterdium aquilinum (L.) Kuhn. Ost. bot. Z. 777: 205-222. TAYLOR, J.A., 1980. Bracken: an increasing problem anda threat on health. Outl. Agric. 70: 298- 304. TRYON, R.M.J., 1941. A revision of the genus Pteridium. Rhodora 43: 1-31, 32-67. VICTORIN, F.M., 1923. Les Filicinées on Quebec. Contrib. Lab. Bot. Unv. Montreal No. 2. REVIEW AUSTRALIAN FERNS AND FERN ALLIES by D.L. Jones and S.C. Clemesha. 232 pp., 297 figures, 60 colour photographs. 185 x 265mm. A.H. & A.W. Reed, Sydney, Wellington & London. 1980. ISBN: 589 50 26 54. Price: £13.95. In a review inthe Fern Gazette in 1977 (Vol. 11: 341-2) the first edition of this book was hailed as the first comprehensive guide to the whole of the Australian continent. This second edition which has benefited by the considerable interest generated by the earlier book, is even more comprehensive; some 47 new species of ferns are added bringing the total to 358 species in 108 genera. It is estimated that at least 20 species remain to be described. The authors are good gardeners and have given us two good chapters on ‘Cultivation’ and ‘Propagation’. Dave Jones, with whom the reviewer has had the ~ pleasure of exploring the Queensland bush, is an acute ecologist who sums up the requirements of plants very quickly and this knowledge is given under each species throughout the book. Dipteris conjugata may, as Dave says, be “‘impossible to grow” but at least Kew has established two healthy, young plants recently. Fern allies, tree ferns, filmy ferns, ‘oddities’ (such as Azo/la, Ophioglossaceae, Marsileaceae and Schizeae) are treated in separate chapters. The rest of the book covers genera and species alphabetically. Nomenclature is up-to-date and the majority of species are illustrated by David Jones’ own line drawings and by Bruce Grey’s superb colour photographs. | found this book gave me immense pleasure to use in the field in Australia but it is also a book to browse and refer to here at home, especially when our Spore exchange contains so many of the species. The publishers too, can be congratulated, not only on an attractive book but for printing a second edition so soon. There are a few mistakes, e.g. the legends of colour figs. 33 and 34 are switched and occasionally the authors cross-refer to synonyms instead of the correct name used in the text, when discussing closely related species under ‘Confusing species’ — which adds to the confusion! But these are minor points; we have a lot to thank these authors for, especially the encouragement they give to others. A.C. JERMY FERN GAZETTE: VOLUME 12 PART 4 (1982) 241 SHORT NOTES ROOT CONNECTIONS IN A COLONY OF OPHIOGLOSSUM VULGATUM IN SOUTHERN ENGLAND Since 1967 | have been observing Ophioglossum vulgatum L. in two pieces of old permanent pasture at Perivale Wood Local Nature Reserve, Greenford, Middlesex (v.c.21). The two pastures consist predominantly of Cynosurus cristatus L., Alopecurus pratensis L., Festuca rubra L., Montia fontana L. ssp. amporitana Senn., Trifolium repens L., Ranunculus acris L., 8. repens L., Lotus corniculatus L., Bellis perennis L., and the moss Cal/liergon cuspidatum (Hedw.) Kindb. The pastures are 80 ft. above m.s.|. and are grazed by horses between May and October each year. They have apparently been grazed in this fashion for the last 60 years at least. The ground is often waterlogged between October and April. The colony extends over most of the western pasture, and a smaller area of the eastern one (see Fig. 1). The number of fronds produced seems to vary widely from year to year, and in the western pasture has varied between 230 and 550, with only 1 —2% of these being fertile in any year. A \ WOODLAND FIGURE 1. Showing the study area and extend of Ophioglossum vulgatum colonies. Hatched area. approximate extent of colony; /arge dots: groups of large fronded plants (protected from grazing). 242 SHORT NOTES E ° w N 1 © = FIGURE 2. An excavated long-distance root connection between two rhizomes of Ophioglossum vulgatum. During June 1978 hoof-cutting of the pasture by quarrelling horses provided a chance to gain some idea of what went on below ground as regards O. vu/gatum. Ina particularly densely cut (but not poached) part | examined some exposed rhizomes and roots of a patch of the fern occurring over an area of 2 mx 3.5 m. One of the 23 plants found was almost free of soil, and with careful teasing and turf-cutting with a knife | was able to clearly trace one of its intact yellowish-brown roots (Ogura, 1972) for 1- 15m to another rhizome with one leaf. The average diameter of this long-distance root was 2.1 mm (sample of 50 measurements using a fine micrometer, range 1.8 — 2.2mm). No branching was found along the course of the root, but one incipient bud was. With further work | was able to follow a major root, from this second rhizome for a further 0.65 m northwards until | lost contact with it (see Fig. 2). On route, only 3 fronds were seen within 1 m either-side of the ‘‘root-line’’. The course that the root followed was remarkably straight, and ran at a depth of 18 — 25 cm. | conjectured that there may just have been connections all the way to the seemingly isolated fronds up to 90 m north of the study area. Further excavations of major root connections between rhizomes gave lengths of 0.2 to 0.55 m, but unambiguous contact with the roots was lost in all cases. The smaller, adventitious roots emanating from the rhizomes averaged 1.62 mm diam., in a sample of 100 (range 1.1 — 1.7 mm). Just south of this study area of 2 m.x 3.5 mm is a hawthorn hedge with bramble scrub at the foot of a railway embankment. Up to 30 large fronds, mostly fertile, are found here each year. Conceivably these are connected with, and may just provide some sustenance to rhizomes in the pasture. The fronds of the latter regularly suffer full or partial decapitation each year. : As the interests of these fern colonies, and the state and composition of the turf are important considerations, | have not ventured to disturb the colony further. REFERENCE OGURA, Y., 1972. Comparative Anatomy of Vegetative Organs of the Pteridophytes. 2 Ed. Berlin. ROBERTS, K.A., and EDWARDS, P.J., 1974. The flora of Perivale Wood Nature Reserve. London Naturalist 53: 34-51. PETER EDWARDS, Royal Botanic Gardens, Kew FERN GAZETTE: VOLUME 12 PART 4 (1982) ons SHORT NOTES OBSERVATIONS ON ANT ASSOCIATIONS WITH RAINFOREST FERNS IN BORNEO The biology of myrmecophily, the association of ants with specialised ‘ant-plants’, has received considerable discussion in recent years (eg, Janzen, 1974; Huxley, 1980; Risch & Rickson, 1981). Although much of the attention has centred around flowering plant examples, these are paralleled by several ferns in which myrmecophilous associations with various species of ants are known. (eg, Yapp, 1902; Wagner, 1972; Gomez, 1974; Jermy & Walker, 1975). Observations made during a visit to the Mt Kinabalu National Park and the Sandakan areas of Sabah, Malaysia during 1981, showed a number of possible cases of ant involvement with several other tropical ferns not normally considered to be myrmecophilous species. Ants tend to be both abundant and widespread almost everywhere within these tropical forests. However, in all the following cases particular fern species and often particular ants seemed to be associated in ways which seemed more than just accidental or casual. All concern ferns growing terrestrially or epiphytically in a variety of different types of vegetation, from altitudes just above sea level to about 5500 ft (c. 1538 m). Although these apparent ant-fern associations have been observed over a wide range of altitudes, they seem most apparent in the low altitude mixed Dipterocarp forests of Sabah. The first case involves plants of Platycerium, in which ants seemed to be more or less permanently resident. The large, accumulating compost-masses of established epiphytic plants of this genus are previously known, in widely scattered places, to be the habitats of a range of animal life (including ants) and other epiphytic plants (Page 1979). Indeed it is possible, that the occurence of ants within wild Platycerium specimens may be regularly the case. In Borneo, Platycerium normally occurs high in the rainforest canopy, but in the Sandakan area of east Borneo, specimens of P. coronarium (Koenig) Desv. could occasionally be found low enough on trees to be accessible for observation by climbing. Field observation showed these wild plants to regularly be the habitats of numerous small ants, each about 2.5 mm in length. A number of wild-originating P. coronarium clumps which had been brought into semi- cultivation low on trees at the Forest Research Centre Garden, Sepilok, near the edge of the Sandahan forest, gave opportunity for closer observation to be made of the ant- activities within these plants. Most plants (6 specimens) had been attached to garden trees in much the same condition as they had been found in the wild. These plants appeared healthy, and in all of them their ant colonies seemed still very active. Ina seventh plant, attempts had previously been made to clear the clump of ants (for convenience in transportation) before being grown by brief immersion in water. Indeed, the treatment seemed to have worked, for no ants were found in this specimen, which also appeared in poor condition and, in contrast with the healthy specimens, had moss-covered basket leaves and crinkled spreading fronds, with evidence of herbivore attack. It is not possible to know if the demise in health of this particular plant was due to the lack of ants or tothe treatment, but observations on the healthy ant-colonised specimens showed that one of the differences was that in these, the ants mined numerous tiny holes from the interior of each plant through the base of the clasping leaves to the outside. The ants then used the holes as entrance and exit points as well as using the gaps around the edges of the leaves. Such holes thus presumably provided better drainage to the nest and compost mass within the plant — perhaps to the plant's advantage in these areas of high and frequent rainfall. One other activity observed which relates to the morphology of the plant, was that the ants were often seen pulling their mandibles, from base to tip, over the white, stellate hairs on 244 SHORT NOTES the outer surface of the clasping leaves (see Holttum 1968: 139 for an illustration of these hairs), sometimes removing some of the hairs and carrying them back via the holes to the nest. The ants present in each Platycerium clump appeared to be a single species, and have been identified as Phesdole sp. (Myrmicinae). The second case of apparent ant-involvement with a wild rainforest fern occurred in Drynaria. Two species of ant (Crematogater sp. (Myrmicinae) and lridiomyrmex sp. (Dolichoderinae) ) each about 3.5 mm in length, were sometimes found in more or less permanent association with plants of Drynaria sparsisora (Desv.) Moore, in low epiphytic sites. These two ant species also occurred widely through the Mt Kinabalu forests, as do many others, but only these species were found associating with this fern. Both seemed to form more or less passive and casual associations with D. sparsisora by colonising the small amounts of soil and debris collecting within its basket leaves. It is of interest to note in relation to these observations that Holttum (1968: 183) has previously recorded that Drynaria quercifolia and other species of this genus have nectaries on the lower surface near the base of the midrib of each lobe, seen as translucent spots in mature fronds. A third, non-residential, type of possible ant-association was seen on occasion with mature plants of Ang/opteris (Forst.) Hoffm. On this plant, when the spores wer mature, black ants about 6 mm in length were found to be present on the fronds. Observation showed their interests to be mainly centred on dehiscing sori, from which the ants were found to be carrying away spores. Also of interest were the activities of a large black ant, about 9 mm in length, which was found on several occasions in different places frequenting the fronds of Nephrolepis hirsutula (Forst.) Presl. These plants occurred mostly in roadside habitats © and the ants were seen frequently on their fronds. Observations showed the ants not to pay much attention to the sporangia, but instead to be attracted to the row of small, white-coloured spots around the periphery of the pinnules of most fronds. These structures appear to be those described by Holttum (1968: 374) as hydathodes, each with a white scale, situated at a vein tip. The ants were seen here mainly at very hot times in the day. The relationships between ants and ferns in rainforest thus seem diverse. These observations are, perhaps, not particularly exceptional, and observations elsewhere may well show other relationships between ants and ferns to exist. In each case observed here, particular species of ants seemed to associate with particular species of fern, and it is perhaps not unlikely that in the process, subtle biological advantages may be gained by each. In each case also, it seemed that the fern was providing something — a safe nesting place? food materials? nectaries? spores? hydathodes? — of immediate attraction to particular local ants. It seems not unlikely too that the fern is gaining some advantage out of these associations. Apart from the more or less physical change of having drainage holes put in the Platycerium, another advantage to all of the species may well be that by harbouring ant colonies or by encouraging ant visitations, additional protection from grazing animals might be gained. In the case of Angiopteris, or indeed, of the other species, it is not known whether the spores carried away ever germinated, and hence whether the ants might play any role in the local dispersal of the species. Such a_ possibility would need careful further experimentation. Indeed the complexity of biotic inter-relationships around these ferns may not even end with ants. Observations suggest that the fern Platycerium coronarium also provides a home for a jumping spider, Synemosyna formica (Salticidae), which imitates an ant in its morphology and was seen to take the mutualistic ant as prey. Further, each Platycerium fern usually also had a lizard, a species of skink, associated FERN GAZETTE: VOLUME 12 PART 4 (1982) 245 with it. These presumably also fed on the ants. Marking the skinks showed these animals to be highly territorial, each skink’s territory associated with a particular Platycerium clump. ACKNOWLEDGEMENTS During the period of these observations, | was a member of Aberdeen University Expedition to Kinabalu, and would like to record my thanks to the expeditions sponsoring bodies, especially to the Weir Fund of Edinburgh University and Fauna & Flora Preservation Society, as well as to the Director and Deputy Director of the Sabah National Parks for supplying practical help in the field. Thanks are due also to Dr D. Cosens, Edinburgh University Zoology Department for assistance in classification of the ants, and Dr C.N. Page, Royal Botanic Garden, Edinburgh, for suggesting the project and for discussion of the results. REFERENCES GOMEZ, P.L.D., 1974. The biology of the potato-fern, So/anopteris brunei.Brenesia 4: 37-59. HOLTTUM, R.E., 1968. Flora of Malaya, Voi. 2, Ferns. Singapore: Govt. Printer. JANZEN, D.H., 1974. Epiphytic myrmecophytes in Sarawak: mutualism through feeding of plants by ants. Biotropica 6: 237-259. JERMY, A.C., and WALKER, T.G., 1975. Lecanopteris spinosa — a new ant-fern from Indonesia. Fern Gaz. 17: 165-176. HUXLEY, A., 1980. Biol. Rev. PAGE, C.N., 1979. The diversity of ferns. An ecological perspective.pp 10-56 in Dyer, A.F. The Experimental Biology of Ferns. London: Academic Press. RISCH, S.J., and RICKSON, F.R., 1981. Mutualism in which ants must be present before plants produce food bodies. Nature 297: 149-150. WAGNER, W.H., 1972. Solanopters brunezi, a little-known fern epiphyte with dimorphic fronds. Am. Fern. J. 62: 33-43. YAPP, R.H., 1902. Two Malayan myrmecophilous ferns, Po/ypodium (Lecanopteris) carnosum Blume and Polypodium sinuosum Wall. Ann. Bot. 16: 185-231. SUSAN PATERSON, Department of Zoology, University of Edinburgh. REVIEW THE FLOWERING PLANTS AND FERNS OF ANGLESEY by R.H. Roberts. XV + 88 pp. 148 x 210mm. National Museum of Wales, Cardiff. CFl 3NP. 1982. Price £3.50 (£4.00 by post). [SBN O 72000241 a. R.H. Roberts, BSB! County recorder for Anglesey, and referee for Po/ypodium is no stranger to readers of the Fern Gazette. His studies in the Polypodium vulgare complex throughout Europe and Macronesia are well known and the Anglesey populations on the limestone of Plas Lligwy are indeed a challenge to any fern taxonomist. The fern records up-date the BSBI/BPS Fern Atlas (1978) showing G. Maw’s record of Dryopteris aemula of 1859 refound in 1979; a recent record (1976) at Mynydd LIiwydiarth failed to get in the At/as apparently. Both Gymnocarpium species and Phegopteris connectilis are missing — a challenge for the fern enthusiast but the stakes are high if RHR has looked already! The author and the publishers who have kept this little book attractive but cheap, are to be congratulated. A.C. JERMY 246 FERN GAZETTE: VOLUME 12 PART 4 (1982) SHORT NOTES ATHYRIUM GOERINGIANUM, A CONFUSED NAME The long confused name Athyrium goeringianum (Kze.) Moore, has been investigated, with a view to establishing its identity. The plant concerned was described as Aspidium goeringianum by Kunze (1848) and was stated to have been collected in Japan by Goering, the type collection, a single frond, being in Kunze’s herbarium. This was housed at Leipzig (LZ) before its complete destruction during the second world war; however a large number of duplicate types of Kunze’s survive at Berlin(B), witha few at Kew (K), Geneva (G) and elsewhere. At Berlin Dr D.E. Meyer was able to establish (pers. comm. 1980) that there were no original specimens there, but that specimens of Goering numbered differently from the type number 115 and determined as Asplenium goeringianum by Kuhn and others, but not by Kunze, were all Athyrium species, which we have been able to identify as the Japanese species, A. niponicum (Mett.) Hance, A. yokoscence (Fr. & Sav.) Christ and A. rupestre Kodama, a somewhat widely differing selection which suggests some confusion. Asimilar situation exists at Kew and Leyden. But as Kuhn may have been expected to be generally familiar with Kunze’s work and often made annotations on his specimens, it seemed likely that an Athyrium species was indeed the basis of Aspidium goeringianum and further information was sought in order to establish this. An illustration of Asp/lenium goeringianum (Kze.) Mett. by Mettenius (1866) was not sufficient to allow any precise identification and did not seem to fit any of the species mentioned with any degree of accuracy, though it is perhaps rather near to A. jseanum Ros., another Japanese species. However it is interesting that Mettenius was careful to distinguish between the true A. goeringianum (his fig. 11)and the A. goeringianum of other authors (his fig. 12). The former clearly has small, generally rounded indusia, becoming kidney-shaped and the latter has slightly more elongated indusia. The drawing of the type (but not fig. 12) would be more compatible with a Dryopteris than an Athyrium though it is impossible to be certain on the basis of this drawing. Although Mettenius had placed this species in Asp/enium, his concept of that genus also included Athyrium species among other genera. Identification of Aspidium goeringianum as an Athyrium species was first put forward by Moore (1860), who made the necessary combination and, we now assume, may have been influenced by specimens sent from Berlin or elsewhere, and in all probability may not have seen Kunze’s type because there was only the single frond at Leipzig. The evidence thus far in our investigation was therefore vague and indeed somewhat confusing. We next turned our attention to Japanese fern works, the most outstanding of which, the detailed and accurate ‘“‘Enumeratio Pteridophytarum Japonicarum’™ by Nakaike (1975), mentions the name Athyrium goeringianum and Athyrium iseanum as a name misapplied by various authors, but nowhere gives it in direct synonymy of any species or as a legitimate name. We were therefore most grateful to hear from Nakaike (pers. comm. 1980) why the name was not applied to any Japanese species. In common with other Japanese authors he was following Koidzumi (1929 and 1932), who had stated that Asp/dium goeringianum was not a Japanese species at all, but must have been recorded from Japan in error, nor was it an Athyrium, but a Dryopteris. In his 1929 paper Koidzumi made the combination, Dryopteris goeringiana (Kze.) Koidz., and by great good fortune illustrated a photographic silhouette of the type in 1932, some ten years before its destruction at Leipzig. We designate this photograph as the neotype, as it is the only extant record of the single original specimen that can be accepted as certainly representing Kunze’s concept. The photograph can be chosen as a type in accordance with the International Code of Botanical Nomenclature (Stafleu et al. 1978), article 9.1, and is accompanied by detailed drawings which depict the sori as SHORT NOTES 247 being circular and not somewhat elongated as would be expected in an Athyrium species such as A. /seanum or A. niponicum. Kunze himself does not mention the shape of the sori in his description, but stated that they are indusiate, a feature not drawn by Koidzumi but suggested by Mettenius’ drawing; the fact that Kunze chose the genus Asp/dium for this species implies that it had the rounded, kidney-shaped indusia characteristic of a Dryopteris species. On seeing Koidzumi’s photograph, it was immediately possible to recognize it, not as the slightly similar Athryrium iseanum, but as a well-known North Chinese, East Siberian and Japanese species at present known as Dryopteris /aeta (Kom.) C.Chr., described in 1901. This species has much coarser segments than an Athyrium, but has somewhat similar lowest pinnules, well developed on the basiscopic side of the pinnae, but reduced on both sides towards the bases of the lower pinnae. It was not known from Japan until 1949, well after the date of Koidzumi's publications and is rare there (see Tagawa 1949 and Hirabayashi 1974), which would explain his statement that it was an extra-Japanese species. As Kunze’s name is considerably earlier than Komarov’s it now becomes necessary to refer to this Dryopteris species as D. goeringiana (Kunze) Koidz., Bot. Mag. Tokyo 43: 386 (1929) and it is clear that many earlier authors were mistaken in applying the epithet goeringianum to various Athyrium species, as confirmed by Nakaike. This has considerable importance for British horticulturalists who have widely grown an Athyrium species labelled as A. goeringianum which contains some well- known varieties making fine and popular hardy garden-plants. Recently there has been some instability with regard to names since the Japanese rejection of the name has gradually become known, but the correct nomenclatural situation has not yet been clearly indicated. We therefore outline what we hope and expect will remain a stable situation. The Japanese and East mainland Asian Athyrium commonly grown in European gardens should be known as A. niponicum(Mett.) Hance. The other more finely dissect species, A. jseanum Ros., to which the name A. goeringianum has been commonly applied in Japan, is seldom grown in European gardens. A. niponicum occurs in two varieties; first the type variety, var. nijponicum, with a concolorous light green upper surface to the frond and a green or pink stipe and rhachis, and second the remarkable, var. pictum (Maxwell) Fraser-Jenkins, comb. nov. (basionym: Athyrium goeringianum var. pictum Maxwell, Gard. Chron. Agric. Gaz. 7867 (2): 25 (Jan. 1867); a clear description given in the editorial, by Maxwell who was the editor from 1865-1907, is sufficient to validate this name at the varietal rank, which the use of a trinomial in that periodical indicated. Synonym: Athyrium niponicum var. metallicum Makino, Bot. Mag. Tokyo 26: 178 (1912)). This has a rainbow effect of a pink and blue tinged central area tothe upper surface of the lamina and pink costae and it appears spontaneously in an approximately 50% ratio in natural populations of the species throughout part, but not all, of its range in Japan. The varietal rank is chosen here as it is of widespread occurrence in natural populations and has some geographical distinction as well. It is neither a monstrosity, nor an unusual form. Some variation occurs in it with some plants being transitional towards var. niponicum and some being paler in colour than others. Nakaike (1975) also points out the existence of a cristate monstrosity grown in gardens (and doubtless there may be others), which should now be known as A. niponicum var. niponicum forma cristato-flabellatum (Makino) Namegata & Kurata. It should be noted that A. niponicum has only one ‘p’, contrary to expectations. ACKNOWLEDGEMENTS We are most grateful to the late Dr D-E. Meyer of the Botanischer Garten und 248 FERN GAZETTE: VOLUME 12 PART 4 (1982) Botanisches Museum, Berlin-Dahlem, Dr T. Nakaike of the Botany Dept., University of Tokyo, A.O. Chater of the British Museum (Natural History), London and Mr R.D. Meikle of the Royal Botanic Gardens, Kew, for their invaluable help. REFERENCES HIRABAYASHI, H., 1974. Cytogeographic studies on Dryopteris of Japan. Tokyo. KOIDZUMI, G., 1929. Contributiones ad cognitionem Asiae Orientalis; Bot. Mag. Tokyo 43. 386. KOIDZUMI, G., 1932. Plantae Japonicae Rarissimae vel Dubiae; Acta Phytotaxonomica et Geobotanica 7: 232-233. KUNZE, G., 1848. Pteridographia Japonica etc.; Bot. Zeit: L 7848: 557. METTENIUS, G., 1859. Ueber einige Farngattungen etc., V/. Asplenium: 198, n. 293, t.6, f.11-12. NAKAIKE, T., 1975. Enumeratio Pteridophytarum Japonicarum. Filicales: 133-134 and 138-140. Tokyo. STAFLEU, F., et al. (Ed.), 1978. International Code of Botanical Nomenclature; Lenigrad. Utrecht. TAGAWA, M., 1949. Fern Miscellany (2); J. Jap. Bot. 23: 77. C.R. FRASER-JENKINS, 36 Fern Hill Road, Cowley, Oxfordshire. R. RUSH, 17 Toronto Road, Ilford, Essex. R.C. CHING, Institutum Botanicum, Academia Sinica, Peking, China. REVIEW SPORE FLORA OF TAIWAN by Tseng-Chieng Huang. 117 pp., 120 plates. 1981. 200 x 268mm. National Taiwan University. Obtainable from author at 24, Alley 3, Lane 30 Chou-shan Road, Tapei, 106 Taiwan. Price US $35. In this book the spores of 159 genera and 535 extant species are described and the locality and collector and number given for each voucher. The descriptive terms are typically those used by palynologists using light microscopy and | suspect contain descriptions of artifacts as seen under the light microscope. Their pictures are given on the plates varying from five to 17 per page. As photomicrographs go they are good but SEM micrographs could have been so much more useful. A list of the Miocene spores described from Taiwan is added. There is a very interesting illustrated glossary of terms used including a dichotromous key to sculptural patterns. A.C. JERMY > | x a8 « ' a xa to ee mat . Pe res ¢ +> - ¥: ad ! ata 3 gay i ey Ms >? “ ne ya hd E ‘7 THE FERN GAZETTE 7," . ‘ ao , . : » , ate vac, AMES aes | papers, articles, or notes of any length on any aspect of pteridology will be for publication. Contributions will be particularly welcomed on floristic, ical and ecological aspects of ferns and their allies, worldwide. Members encouraged to make use of short notes for reporting pteridophyte field and records. All may be illustrated with black and white photographs of cal quality and line drawings. Short papers can usually be incorporated y than longer ones. — pest | should be sent to: _ C.N. Page, Royal Botanic Garden, Edinburgh, U.K. ae wl te last date for receiving notes and articles to make the following early summer ast date for rece | g early er. Rh eet Sige ces = ee sores ee - 30th November each year follow the general style of this number. 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Each \otograph should be marked on the back with details of author and figure number, early marked “top”. _ , | versa FA Grouped illustrations should follow the numbering system, fig 1a, fig 1b, fig re numbers should be applied to illustrations in pencil only or on a transparent tering will be added by the editor ina style and size consistent with the journal. s: Type on a separate sheet from the manuscript — include any necessary details of s submitted (the editor will apply any correction or reduction.) vessel follow closely the style of this number to speed publication. Lists in other to authors for re-typing. ee Pe ea Ae i <0 aS as - 7 7 7 Coes ) ] } s x i AR a « % “ne NE Es eM aa nar aaah : | ev jiew in the Fern Gazette or B.P.S. Bulletin should be sent to A.C. Jermy, | epart ment, British Museum (Natural History), ‘Cromwell Road, London, ° ae ae ; 6 eee ey la = THE ~ FERN GAZETTE VOLUME 12 PART 4 : 1982 CONTENTS Page MAIN ARTICLES The continuing need for more monographic studies of ferns - R.E. Holttum 185 A preliminary survey to the pteridophytes of Transkei, Southern Africa - J.P. Roux ' 191 Dictymia brownii (Polypodiaceae s.s.), an ancient Australian fern - T.G.Walker & C.N. Page 197 The ecology and cytology of Phanerosorus (Matoniaceae) - 7.G. Walker & A.C. Jermy 209 Observations on the venation patterns in Ophioglossum, Botrychium and He/lminthostachys - S. Bhambie & Parhash Madan 215 A new concept of Drynarioid ferns - Subash Chandra 225 Field observations on the nectaries of Bracken, Pteridium aquilinum, in Britain - C.N. Page 233 SHORT NOTES Asplenium pulcherrimum in Borneo - 8.S. Parris | 214 The appearance and disappearance of a Dryopteris culthusiana colony - Peter Edwards 224 Morphological and cytotaxonomic observations on Azo//a pinnata - D.S. Loyal, A.K. Gollen & Raman Ratra 230 Root connections in a colony of Ophioglossum vulgatum in Southern England - Peter Edwards 241 Observations on ant associations with rainforest ferns in Borneo - Susan Paterson 243 Athyrium goeringianum, a confused name - C.R. Fraser-Jenkins 246 REVIEWS 190, 196, 208, 223, 229, 240, 245, 248 (THE FERN GAZETTE Volume 12 Part 3 was published 10th November 1981) Published by THE BRITISH PTERIDOLOGICAL SOCIETY, c/o Department of Botany, British Museum (Natural History), London SW7 5BD Metloc Printers Ltd., Old Station Road, Loughton, Essex. _ FERN GAZETTE VOLUME TWELVE PART FIVE 1983 OCT 17 1983 SS LBRARES YH THE JOURNAL OF THE > BRITISH PTERIDOLOGICAL SOCIETY THE BRITISH PTERIDOLOGICAL SOCIETY Officers and Committee for 1983 President A.C. Jermy, Botany Department, British Museum (Natural Hist Cromwell Road, London, SW7 5 Vice Presidents J.W. Dyce, R.E. Holttum, F. Jackson, R. ; Irene Manton, S. | General Secretary and A.R. Busby, 42 Lewisham Road, Smethwick, British Fern Distribution Recorder West Midlands, Treasurer Dr. B.A. Thomas, ‘Broleciea Sciences Department, Uni London, Goldsmiths’ College, Creek Road, London, SE. Membership Secretary Lt. Col. P.G. Coke, Robin Hill, Stinchcombe, | Gloucestershire, Gi Meetings Secretary Kathryn Kava neh Botany Department, British (Natural History), Cromwell Road, London, S Publications Secretary Devon Masarati, 27 Silsoe House, Park Ville ast, 3 | ) London, NW1 he Editors: . ory f Fern Gazette: Dr. C.N. Page, Royal Botanic Garden, Edinburgh, EH3 61. assisted by J.A. Crabbe, A.C. . Bulletin M.H. Rickard, The Old Rectory, Leinthall Starkes, nr. Ludlow, Sh assisted by J. Spore Exchange Organiser . as F. ail oe 13 Perry Mill Road, Pec Pershore, Worcest Archivist N.A. Hall, 3 Sydney Road, Bramhall, Stockport, ee Committee annually. The Gazette publishes matter chiefly of specialist interest on intarhationel pte the Bulletin topics of more general appeal. (See inside back cover for notes to corte fern enthusiasts. It provides a wide range of information about ferns through the meeilend publications and available literature. It also organises formal talks, informal discussions, fi scheme. The Society has a wide membership which includes gardeners, nurserymen a botanists, both amateur and professional, in Britain and overseas. Articles for the Society’ 5 publications are received from and reflect the interests of, Society members. Membership is open to all interested in ferns and fern-allies, worldwide. Subscription rates (due ott on 1st January each year) are: Full Personal Members £7; Personal Members not receiving the — aa 84 Fern Gazette £5; Student Members (full-time students) £5; Subscribing Institutions £8. ie Applications for membership should be sent to the Membership Secretary (address above), from : Bk « whom further details can be obtained. (Remittances made in currencies other than Sterling are — p <4 £1.00 extra, to cover bank conversion charges ) a e pers Back numbers of the Gazette and Bulletin are available for purchase from Dr. B.A. Thomas, — (address above) from whom further details can be obtained. ~ con Br eo = FERN GAZ. 12(5) 1983 249 AN ECOLOGICAL SURVEY OF THE FERNS OF THE KILLARNEY DISTRICT, CO. KERRY, IRELAND A. WILLMOT Derby Lonsdale College of Higher Education, Derby ABSTRACT A systematic list of the ferns and fern allies of the Killarney region of Co. Kerry, Ireland (parts of v.-c. H1 and 2) is given based mainly on records made in 1980 and 1981 by the author. This includes notes on the distribution, abundance and habits of the taxa. The list includes 41 species, hybrids and subspecies. Notes are also given on all other fern taxa recorded for the area. The records made by the author include the first Irish record for Dryopteris affinis (pseudomas) ssp. stillupensis; the first records for the area of D. oreades and Polypodium xmantoniae; and the first records since 1950 for Ophioglossum vulgatum and Oreopteris limbosperma. The status of doubtful species in the area is discussed: Equisetum xmoorei and Dryopteris expansa are rejected as errors for other species; while no conclusion is reached about the presence of Selaginella selaginoides and Gymnocarpium dryopteris in the area. The phytogeographical affinity of the flora on a European basis is described by dividing it amongst the floristic elements of Birks(1976). The flora consists mainly of widespread species, species with southern and oceanic affinities anda few species with montane or arctic/alpine affinities. INTRODUCTION The Killarney district in Co. Kerry is one of the more interesting areas for ferns in the British Isles being a mountainous area on the extreme south-western seaboard. The actual area studied is shown on Figure 1, the boundaries being defined by grid-lines of the Irish national grid. This area lies approximately at latitude 52°N and longitude 10°W and includes parts of two vice-counties, H1 South Kerry and H2 North Kerry. Although Co. Kerry is a maritime county only very short stretches of coast were included in the survey area, and these were estuaries. These stretches were to the north-west of Killorglin and to the south of Kenmare. 80 oo LY H2 N. KERRY | H1 S KERRY 70 FIGURE 1. The Killarney district surveyed. B: Barraduff, Ct: Carrauntoohil, Cw: Carrigawaddra, Kg: Killorglin, Kv: Kilgarvan, MM: Mangerton Mountain, PeM: Peakeen Mountain, PM: Purple Mountain, TM: Tomies Mountain. FERN GAZETTE: VOLUME 12 PART 5 (1983) 250 FIGURE 2. Killarney Woods. FIGURE 3. Typical upland vegetation. FERNS OF KILLARNEY DISTRICT 251 The topography of the area is dominated by a series of parallel ridges and valleys running east-west. These owe their origin to differential denudation of rocks thrown into anticlines and synclines by Armoracian folding (Aalen, 1978). Softer Carboniferous rocks were mostly removed to produce the valleys, leaving the older, more resistant Old Red Sandstone upstanding to form the ridges. These ridges include large areas of land over 300m (1,000 ft), a number of mountains over 600m (2,000 ft) and the Macgillycuddy’s Reeks. The Reeks are the highest range of mountains in lreland, with the highest peak in Ireland at 1,022m (3,414 ft) in Carrauntoohil. Glaciers in more recent times have accentuated this ridge and valley topography by deepening valleys and steepening cliffs. Overall this area is one of great contrasts from flat, well- wooded valley bottoms (Figure 2), at sea-level to bare, exposed mountain tops which are just shattered piles of rock. In between the majority of the area consists of rough hillsides with occasional boulders and outcrops of rock (Figure 3). Apartfrom this area of high ridges and low valleys, there is an area of generally lower, flatter topography in the north of the region. This area extends from north of Barraduff and Killarney to Killorglin. It drops gradually from around 210m (700 ft) in the east to near sea-level in the west. This area is covered in glacial drift and is a region of low undulating hills with large areas of poorly-drained land between. Throughout the region there are numerous lakes at all altitudes mostly due to the action of ice. Included in these are the famous lakes of Killarney. Due no doubt to the rich fern flora of the area, the area has been better studied than most areas of Ireland. There are numerous references in the literature at the end of the last century and the beginning of the present one to the ferns of the area (Bouskell, 1905; Hind, 1857 and Rasor, 1882) culminating in Scully's (1916) flora of Kerry. In the present century there has been somewhat of a lull in references to the ferns and flora of the area until recently when there has been an upsurge of interest (O'Mahony, 1980 and Curtis and Mhic Daeid, 1981). METHODS The records made in this survey were all made in the summers of 1980 and 1981 for the area defined in Figure 1, with mostof the recording for areas over 300m carried out in 1981. Records were collected on the basis that two or three examples of each major habitat Type (Table 1) were visited in each 10km grid square. In each habitat a subjective estimate of the abundance of every fern present was made on the Dafor scale. lf a habitat extended over more than one kilometre square, a separate record was made for each. Similarly if a habitat extended over a range of more than 75m, a separate record was made for each 75m altitudinal range. If during visiting one habitat, other major habitats were encountered, records were also made for these. Habitats were selected either because there were old records for the sites or because they looked promising on maps or on the ground. Where habitats contained no ferns, no records were made. In all, records were collected from some 200 altitudinaly- limited monads, but due tothe non-systematic nature of the survey there are a number of forms of bias in the results. The two major ones are: a bias towards monads near roads and buildings, and a bias to monads containing old records or ones that looked superficially interesting. Where the ecological or geographical distribution of taxa is discussed below, attempts have been made to correct for these forms of bias. A survey was also made of the major works and data banks which contain fern records for the area. These included Jermy et a/ (1978), the Biological Records Centre, Monk’s Wood, England and Scully (1916). The herbarium at the British Museum (Natural History) (BM) was also examined for specimens from the area. 252 FERN GAZETTE: VOLUME 12 PART 5 (1983) Table 1 : Fern taxa recorded for the Killarney Area Table includes all taxa recorded for the study area, those recorded in present survey are marked !, those for which only pre-1950 records exist are enclosed in round brackets ( ) and those for which only doubtful records exist are enclosed in square brackets [ ]. Numbers in brackets refer to floristic elements in Table 3. (Lycopodium clavatum) (5) (Lycopodiella inundata) (7) Huperzia selago! (5) (Diphasiastrum alpinum) (8) [Selaginella selaginoides] S. kraussiana! lsoetes lacustris! (7) !. echinospora (7) Equisetum hyemale (5) [E. x moore!] (E. x trachyodon) (E. variegatum) (7) E. fluviatile! (5) E. arvense! (5) E. x litorale! (E. sylvaticum) (5) E. palustre! (5) E. telmateia (3) (Botrychium lunaria) (5) Ophioglossum vulgatum! (6) Osmunda regalis! (3) Hymenophyllum tunbrigense! (4) H. wilsonii! (4) Trichomanes speciosum! (4) Polypodium vulgare! (6) P. interjectum! (3) P. x mantoniae P. australe! (2) Pteridium aquilinum! (6) Thelypteris palustris (6) Phegopteris connectilis! (5) Oreopteris limbosperma (3) Asplenium scolopendrium! (3) (A. x confluens) . adiantum-nigrum! (3) . onopteris! (1) . Marinum (2) . trichomanes agg! (6) ssp. quadrivalens A viride! (7) A ruta-muraria! (6) A. ceterach! (3) Athyrium filix-femina! (5) [Gymnocarpium dryopteris] Cystopteris fragilis! (5) Polystichum lonchitis (7) P. aculeatum! (3) P. x bicknellii P. setiferum! (2) Dryopteris oreades! (3) D. filix-mas! (6) D. affinis! (3) ssp. sti/lupensis! ssp. borreri! ssp. affinis! (D. remota) D. aemula! (4) D. carthusiana! (5) D. dilatata! (3) [D. expansa] Blechnum spicant! (3) Pilularia globulifera! (3) BBRBEB The bedrock of the ridge and valley area is mainly hard sandstone and conglomerates with thin occasional beds of shale of Devonian age (Wright, 1927). The only igneous rocks in this area are small areas of Felsite below Mangerton Mountain (Scully, 1916). There are also areas of glacial deposit of the same rock types, and small patches of Carboniferous Limestone in the Roughty Valley and around Lough Leane and Muckross Lake. The lowland area to the north is almost entirely covered in peat and drift derived from similar acid rocks to those that form the majority of the uplands to the south. FERNS OF KILLARNEY DISTRICT 253 The upland area was sufficiently high for ice sheets to have formed on it inat least two periods during the recent ice-age (Mitchell, 1976). These ice sheets formed the glaciers that deepened the valleys and deposited the drift deposits on the lowlands to the north. In the times of these ice sheets some mountain tops protruded through the ice to form nunataks and the sea-level was much lower. These ice sheets scraped the soil from much of the upland area leaving just bare rock or shattered piles of rock fragments. Elsewhere in the area, particularly onthe shallower slopes, blanket peats have formed since the ice retreated. Where mineral soils are present they are gley- podzols and brown podzolic soils (Ryan, 1963) which are poorly drained and acidic. In the lowland area to the north there are either peats, or acid brown earths (many with gleying), gleys and podzolised gleys (Ryan, 1963). These are again mostly poorly drained and acidic. The climate of the area is described in Scully (1916) and Aalen (1978), the main features of which are dampness and mildness. The rainfall is high at about 200mm (80 inches) per annum at 18m in the upland area rising to 250mm (100 inches) per annum and more on the mountains, but falling to only 100mm (40 inches) in the lowlands to the north. This rainfall is spread through the year with about 250 raindays. The mean air temperature is 6.0-7.0°C in January and 15.0-15.5°C in July. Frosts are rare at low altitudes but become commoner at high altitudes. The potential vegetation of the area at lower altitudes is mostly oak wood, which would have been similar to the remaining native forest vegetation in the Killarney district described by Kelly (1981). This would have covered all the lowlands except where impeded drainage allowed carr woods, dominated by alder to develop, or peat bogs. The natural altitude limit of forests in the area is uncertain but they would have given way on the hills, before clearance, to blanket peat and heather moors. These in their turn would have been replaced by open, rocky communities on the hill tops. This natural vegetation of the area has been greatly modified by man, particularly at lower altitudes. In particular the woods have been cleared to form mostly damp pastures leaving only patches of native woods around the Killarney lakes. Many of the peat bogs and blanket peats have been cut for fuel and the heather moors in many cases have been grazed to form further grasslands up to c. 450m (1,500 ft). However, the cutting of the forests has in some areas allowed blanket peats to spread (Aalen, 1978). The advent of man in the area has produced other habitats some of which are suitable for ferns, in particular, the shrub-topped hedgebanks and stone walls around fields and buildings. Man’‘s influence has not altered the numerous lakes in the area much as yet, except by clearing the vegetation back from the banks forming stoney shores. The vegetational history of the lowlands of the area since the last ice age has been examined in the peat bogs of the area (Jessen, 1949 and Watts, 1963). These studies have demonstrated that the vegetational history of the area has in general terms been the same as the rest of the British islands. That is open tundra type vegetation after the retreat of the ice, followed by the invasion of trees to form woodlands. These were at first dominated by birch, but this was replaced by pine which in its turn was replaced at the oak and birch of the woods we now consider the native vegetation of the area. RESULTS These are presented in two main ways, the systematic list described the status, ecology and distribution of all fern taxa recorded for the area, while Table 2 summarises the frequency of the commoner taxa in the commoner habitats. The possible origins of the flora and its biogeographical affinities are discussed in the 254 FERN GAZETTE: VOLUME 12 PART 5 (1983) summary section at the end. Nomenclature follows Jermy et a/(1978), but Dryopteris pseudomas has been changed to D. affinis and D. austriaca to D. dilatata; and Thelypteris palustris has been reinstated. SYSTEMATIC LIST OF TAXA Each taxon recorded during the survey is given with notes on (a) its geographical distribution, (b) its habitats, growth and ecology and (c) its altitudinal range, as recorded in the survey. Where a taxon was recorded in only a few monads, an entry is given in the form (a-c) followed by notes on the individual occurrences. Reference in this list to abundance in habitats refers only to abundance in examples of the habitat where the species occurred, how frequent the species was in the habitat type can be seen from Table 2. Notes are also given for significant extensions to (a), (b) and(c) from other sources, particularly post-1950 10km grid square records not reconfirmed here, and selected pre-1950 records that have not been refound recently, including those that have not been reconfirmed in this survey. Grid references prefixed with circa are my interpretations of localities originally given without a reference. Numbering of taxa follows Jermy et a/(1978). Abundance is recorded on the Dafor scale where D = dominant, A = abundant, F = frequent, O = occasional, R = rare and L = locally. The Biological Records Centre, Monks Wood, England, is referred to as BRC, Jermy et a/(1978)asAtlas, andScully(1916) as RS. Old and recent records refer to pre-1950 and post-1950 records respectively. Where unqualified, ‘grid squares” refer to 10 x 10km squares and ‘‘monads” referto 1 x 1km squares. Reference to all grid squares refers to all grid squares shown in Figure 1. 1.2 Lycopodium clavatum Only records are in RS for the Mangerton Mountain, Muckross Lake area and in At/as pre- 1950 for V/9.8 no doubt based on same records. 2.1 Lycopodiella inundata Only one recorded station (More, 1876 and RS) on margin of Lough Guitane covered at moth water (c. W/01.84). 3.1 Huperzia selago (a) Widespread but local in south and west of area due to its altitudinal requirements. Recorded for grid squares V/7.7, V/8.7, V/8.8, V/9.7 and V/9.8. (b) Most commonly on rock outcrops in upland vegetation where it was O-R, with a few records each for scree and heather moor where it was only R. It was seen to be more often producing bulbils than sporangia and was often present as broken off stem fragments which may have been acting as a means of asexual reproduction. (c) Not recorded below 240m, above, becoming more common with increasing altitude upto limit of survey at 810m. Recorded in Atlas post-1950 for V/7 8. RS records the altitudinal range of the species as sea level to 3,370 ft (1011m) which is almost the highest point in the area. 4.1 Diphasiastrum alpinum Only old records exist for Mangerton Mountain (Newman, 1844 and RS) V/9.8, and ridge south west of Curraghmore summit c. V/ 78.82 (RS). There are specimens in BM from Mangerton dated 1835 (ex herb. J.W. Curtis) and ex herb. H.T. Ryder (without date). 5.1 Selaginella selaginoides According to Praeger (1934) this is a most striking absentee from Kerry. There is a specimen in BM from Killarney VC H2 dated 1930 but no mention of this or any other record in Atlas. Considered doubtful as this record is only one for area 52 S. kraussiana (a-c) Recorded once on the banks of the Roughty River at V/963 723. alt. 15m. About 10 patches, each less than one square metre, in grass at base of hedge on north side of river, all plants sterile This site was well away from any houses or gardens, so plant was probably carried down to site by the river in flood. Species is recorded as new county record in O'Mahony (1980) at the same locality FERNS OF KILLARNEY DISTRICT 255 Sooo. ee Ge PEPE a Pads hak Isoetes lacustris "7 yp Equisetum fluviatile Equisetum xlitorale Equisetum arvense Athyrium filix-femina Osmunda regalis Dryopteris affinis Blechnum spicant 7 GZ G y L G aoenes Dryopteris dilatata Pteridium aquilinum Dryopteris aemula Dryopterlis filix-mas Polystichum setiferum ce Polystichum aculeatum Oreopteris limbosperma ‘od Huperzia selago Asplenium scolopendrium : Asplenium G % Z OLLLLL 2 LL ZZ A A v 4 G adiantum nigrum 3 ee ee a A Z| Asplenium trichomanes wm OY a ee, 7 60 Asplenium ruta-muraria j YF Asplenium ceterach of 100 Polypodium vulgare agg. %Z 0 77ZIZ2 rrrz7T# UK Z LO A Hymenophyllurn wilsonil me Ga Hymenophy 20 tunbrigense NUMBER OF MONADS4 | 99 | a2 | a | 49] 5 | w | 62 | 44 | 45 | 12 | 94 | 0 | oF | a7 | Table 2 256 ' FERN GAZETTE: VOLUME 12 PART 5 (1983) 6.1 lsoetes lacustris (a) Widespread but local thoughout the south and west of the area. Recorded for Cloon Lake V/70.78, Lough Caragh V/71.88, Hag’s Glen V/81.84, Lough Nambrackdarrig V/94.78 and Muckross Lake V/95.86. (b) All records mentioned in(a) are based on observation of megaspores. In all, 66 plants were examined for megaspores from 9 localities, of these 34 from 8 localities contained megaspores and were all referable to /. /acustris. Occurred amongst stones on the bottoms of lakes, sometimes as occasional plants, other times as extensive beds, some records based on plants or leaves washed up on shores of lakes. Plant occured as two growth forms, one with long, straight leaves in deep water (30-90cm) and the other with shorter, curled leaves in shallower water (15cm). Both forms had / /acustris type spores. (c) Occurred from sea level to 600m with equal frequency. The presence of clear lakes with stoney bottoms seems to determine its distribution rather than altitude. Recorded in RS up to 720m and in Atlas for V/7.9 and V/8.7. 6.2 1. echinospora Not recorded in present survey despite examination of numerous megaspores; see entry for /. lacustris above. Recorded (RS and At/as) as widespread but local, pre- 1950, throughout the area. There are also specimens in BM from area mostly pre-1950 but one from Lough Guitane (c. W/02.85) dated 1977 (A.C. Jermy No. 13591). The dearth of recent records suggests that some older records may in fact have been /. /acustris. 7.1 Equisetum hyemale Several old records in RS usually next to rivers. Only recent record is a specimen in BM from peninsular on west side of Caragh Lough (c. V/71.90) at edge of wood on foreshore (Miss B. Morgan s.n. 1952). See comments after E. variegatum. 7.1x2 E.x trachyodon (E. hyemale x E. variegatum) No recent records. RS gives two records both for riversides. There are several specimens of this hybrid from the area at BM (A.C. Jermy, pers. comm.) but see notes after FE. variegatum. 7.2 E. variegatum No recent records but several old records in RS and several old specimens in BM. These records are all either for Lough Caragh or Muckross Lake in the vicinity of Muckross House, or Lough Leane in the vicinity of Ross Island. They are therefore all for altitudes below 30m and by lakes or in ditches. Several authors (Atlas, Newman 1844, Praeger 1934 and RS) refer specimens of this species inthe area to EF. wi/soniiNewm. or E. variegatum var wilsoni Newm. The status of these is uncertain (Atlas) but at present they seem best treated as a lowland, upright and unbranched form of E. variegatum with smoother stems. The classic locality for this form seems to be ditches near lake, Muckross Demense, from where there is a specimen in BM, collected by W. Wilson. The last four Equisetum taxa plus E. variegatum var. wilsoni form a hybrid complex which needs further investigation to determine how many taxa of what status occur in the area (C.N. Page, pers. comm.). In this survey two localities for plants referable to this complex were found. One was amongst stones on shore of Sugarloaf Island, Lough Caragh (G.R. V/712.887) and the other stones on shore of Kilbeg Bay, Muckross Lake (V/958.860). These are being studied and will be reported on elsewhere. 7.3 E. fluviatile (a) Recorded for all grid squares except V/9.9. Widespread but occasional in area. (b) Emergent sides of lakes and slow moving rivers, variable in abundance here from rare to locally dominant. Recorded as occasional for a few areas of marsh. (c) Sea-level to 240m with scattered records up to 600m. Recorded in Atlas for V/9.9. 7.4 E. arvense (a) Widespread but local throughout the area. Recorded for all grid squares except V/7.7,V/8.8 and W/0.8. (b) Occasional in tall herb communities and hedge banks with individual records for a range of other communities including cultivated ground; probably restricted in area due to the prevalence of peaty soils which it avoids (At/as). (c) Sea-level to 60m with scattered occurrences up to 360m. This restriction to low altitudes is again probably due to its avoidance of organic soils. Recorded in Atlas for V/8.8 and W/0O.8. 7.4x3 E. x litorale (E. arvense x fluviatile) (a) Recorded for 8 monads around Muckross Lake, its inlet river, the southern shore of Lough Leane, and Ross Island plus an isolated record for Killorglin (V/78.96). (b) Occurs as O-R amongst rocks on lake shores, in damp, acid grassland (grass moor) and marshes, with individual records for tall herb and hedge communities. Occurred in same area as parents in intermediate habitats. The cluster of records round Muckross Lake area suggests hybrid only rarely forms de novo from parents but can be spread by fragmentation in water. (c) Sea-level to 60m. There are old records at BRC for Kilgarvan 1930 (c. W/01-73) and Reen Bridge (V/870.705) 1929. FERNS OF KILLARNEY DISTRICT 257 7.6 E. sylvaticum Recorded in RS as rare but scattered in several places around Killarney, the lakes, Mangerton Mountain plus the Roughty and Slaheny valleys. No recent records traced. vur E. palustre (a-c) Recorded three times for area: frequent in Phragmites marsh by road from Muckross to Dinnish cottage, alt. 27m (V/94.86); locally frequent on hedgebank at Aghadoe, alt. 90m (V/945.928); and locally frequent in marshy field near old railway station, now new school, at Kilgarvan, alt. 30m (W/009.737). Recorded in At/as post-1950 for W/0.9. 7.8 E. telmateia Numerous old records thinly scattered through area in RS, mostly at low altitudes associated with water and/or woods. Only recent records is in BRC for roadside bank one mile west of Killorglin 1967. (c. V/76.96). 8.1 Botrychium lunaria No recent records. Recorded in RS for a few sites scattered through area. 9.1 Ophioglossum vulgatum (a-c) Recorded once as rare in alder/ash/birch carr by side of track from Muckross House to Mines, alt. 25m (V/964.865). : Old records at BRC, and in RS and Bouskell (1905) record species as growing in woods and grassland at a few sites throughout the area. 10.1 Osmunda regalis (a) Recorded for all squares in area except W/0.9. Widespread and common throughout area but becoming less so in north-eastern corner of area. (b) The two habitats where the species was most commonly encountered were hedgebanks and stones by lakes. The species was more abundant (O-R) in the latter as compared to only rare in the former. The species occurred less commonly in deciduous woods, grass moor and on rock outcrops. It was generally only rare in these three habitats. Only small, sterile plants occurred on rock outcrops without seepages. It also occurred a fewtimes in each of a range of other habitats including peat cuttings, which it was seen to be colonising vigorously from spores. Overall appeared to prefer well-illuminated, damp habitats. (c) Sea-level to 360m with scattered records above up to limit of survey at 810m. Recorded in Atlas for W/0.9. 14.1 Hymenophylium tunbrigense (a) Recorded mostly for the area of the Killarney Lakes (V/9.8) with scattered out-lying records for: Mullaghanattin (V/73.77), Lough Caragh (V/71.90) and Looscaunagh Lough (V/88.79). (b) The habitat the species most commonly occurred in was rock outcrops both acidic and calcareous, that were shaded and covered in moss. It also occurred as an epiphyte on a variety of trees. In both situations it was generally O-R but where it occurred it grew as dense mats. Rasor (1882) and Turner and Watt (1939) also noted this species grew more commonly on rocks than as an epiphyte. Generally, but not always, more abundant than H. wil/sonii at lower altitudes. (c) Sea- level to 360m. Also recently recorded inAt/as for V/7.8, V/8.8, V/9.9,W/0.8 and W/0.9, and latterly, at BRC, for Carrigawaddra near Kilgarvan (c. W/04.74). 14.2 H. wilsonii (a) Thinly scattered throughout southern and central parts of area. Recorded for squares V/7.7, V/78, V/8.8, V/9.7, V/9.8 and at Crumagloun (W/02.72). (b) The two most favoured habitats for this species were rock outcrops and tree trunks. As in H. tunbrigense there were more sites for outcrops than tree trunks but the preference for outcrops was even stronger inthis species than inH. tunbrigense. This contradicts Turner and Watt (1939) who recorded H. wi/sonii as more of an epiphyte thanH. tunbrigense. The rock outcrops were damp, shaded, moss-covered and generally acidic. This apparent preference for acidic rocks was associated with the preference of this species for high altitudes, note calcareous rocks occurred at lower ones. The species was generally O-R on rocks and tree trunks, though where it occurred it was present as dense mats. Individiual records were also made for the species as rare on hedgebanks, screes, walls and rocks by water. It was commoner at higher altitudes than H. tunbrigense. (c) Occurred from sea- level to altitudinal limit of survey at 810m. However, it was only present in 10% of monads below 120m becoming gradually commoner at higher altitudes (70% of monads above 600m). Recorded in Atlas for V/9.9, W/0.8 and W/0.9. RS records the species as extending to 990m on the Reeks. 15.1 Trichomanes speciosum (a-c) Recorded only once during survey, shown to me by a local botanist who asked that | should not divulge the locality or give any details of it. The species was once widespread, but local in the Kiilarney area on shaded, damp, acid 258 FERN GAZETTE: VOLUME 12 PART 5 (1983) rocks often near waterfalls, (RS, and specimens in BM). However, it has been collected to extinction, or almost so, in many of its old localities. To discourage further depredations current localities are not published, so it is not possible to comment on its present status in the area. This species is now protected by the Irish Wildlife Act of 1976. 16.1-3 Polypodium vulgare agg. (a) Widespread and common throughout the area. Recorded for all squares. (b) Recorded most commonly for rocks, walls and as an epiphyte, with a number of records for hedgebanks and isolated records for scrub, moor, heath and woodland. More abundant (F-R) on calcareous outcrops and mortared or concrete walls than (O-R) on acidic outcrops and drystone or earthed walls. Generally recorded as occasional when an epiphyte, O-R on hedgebanks, but only as rare inthe remaining habitats. (c) Common from sea-level to 480m becoming less so above the limit of survey at 810m. See also entries for seggregates below, note that only some records for the aggregate were critically determined to seggregate species and altitudinal limits of seggregates not analysed as records too few. Overall P. vu/gare agg. was recorded for about 100 monads, P. vulgare, 17, P. interjectum 12, P. x mantoniae 1, and P. australe 6. 16.2 P. vulgare (a) Records widespread throughout area. Recorded for all squares except V/8. 7. (b) Recorded only for: acid outcrops 7 records (F-R); drystone and earthed walls 3.records (R); hedgebanks 3 records (O-R) and as an epiphyte 5 records (O). This avoidance of more base-rich substrates is in accord with the ecology of the species elsewhere. 16.2 P. interjectum (a) Records widespread throughout area, but less common than P. vu/gare. Recorded for squares V/7 8, V/8.7, V/8.8, V/8.9, V/9.7, V/9.8 and W/0.7. (b) Recorded only for: rock outcrops and scree 5 records (F-R); drystone walls and hedgebanks 2 records (O); mortared wall 1 record (O); and as an epiphyte 4 records (LA-O). Fhere was no tendency noted here for-the species to erequire either more lime or higher humidity than P. vu/gare as suggested in the At/as. 16.2 x 1 P. x mantoniae (a-c) Recorded once as rare on a hedgebank at Gearha (V/77.73) altitude 60m. No previous records traced. 16.3 P. australe (a) Restricted to areas of limestone outcrop around Muckross and the Kenmare Valley and uncommon even there. (b) Recorded only for: limestone outcrops 3 records(LF-R); mortared walls 3 records (LF-O); and once occasional as an epiphyte on oak. Selected records: limestone outcrop Innisfallen Island (V/93.89); mortared wall Dunkerron Castle (V/88.70) and epiphyte Muckross woods (V/95.86). 17.1 Pteridium aquilinum (a) Common throughout area. Recorded for all squares. (b) The four habitats the species most commonly recorded for were: heather moor (LD-O); grass moor (LD-R); deciduous wood (LA-O) and hedgebanks (LA-F). The species formed dense stands over large areas of moorland but fronds were smaller with spaces in between in deciduous woodlands. Species sometimes occurred ina wide range of other habitats. (c) Recorded in about 80% of monads below 360m but in less than 10% above and with no records made above 600m. 18.1 Thelypteris palustris (a-c) Only recorded for two adjacent monads at Muckross (V/95.86 and V/96.86) as locally abundant to rare in damp, tall herb and grass communities, altitude 25-30m, by side of road from house to mines. Other records record two more localities in vicinity of lakes, near western end of Ross Island (c. V/94:88) and near Cromaglen Bridge (c. V/92.82) (RS). At/as gives a post-1950 record for V/9.9 but no record for this could be traced at BRC. 19.1 Phegopteris connectilis ; (a-c) Recorded three times as rare on rock outcrops at higher altitudes near flowing water; Cloghera, alt. 120m (V/72.78); Torc Mountain, alt. 330m (V/95.84) and Horses’ Glen, alt. 300m (V/99.82). Other records give a similar picture (RS, At/as, and Curtis and Mhic Daeid, 1981), but record further localities. Records for marshes at Muckross (Newman, 1844 and RS) are probably erros for Thelypteris palustris. 20.1 Oreopteris limbosperma (a) Widespread but local in southern and central parts of region. Recorded for squares V/7.7, V/8.8, V/9.7, V/9.8, W/0.7 and W/0.8. (b) Recorded for heather moor (F-O); hedgebanks (O-R) and rocks by rivers (R). (c) 30-480m. FERNS OF KILLARNEY DISTRICT 259 21.1 Asplenium scolopendrium (a) Widespread throughout area (recorded for all squares), common in lowland area, especially around the Killarney Lakes, but much less so in upland areas. (b) Occurred most commonly, in order of decreasing frequency, on walls, hedgebanks, in deciduous woods and on rock outcrops. Much more frequent and abundant on limestone outcrops and mortared walls (F-R) than acidic outcrops and drystone walls (O-R). Plants smaller in drier habitats and in mountainous areas. Less abundant on hedgebanks (O-R) than in deciduous woods (F-R). (c) Becoming less common from sea-level to tops of mountains such that is occurred in more than 50% of monads below 60m but less than 10% above 240m. 21.1x7b A. x confluens Found once near Killarney in 1875 by Frazer (At/as) but not recorded since. (But see Rush, this - Volume, p. 301). 21.2 A. adiantum-nigrum (a) Widespread but local thoroughout area. Recorded for all grid squares. (b) Recorded for rock outcrops, walls and hedgebanks. Commoner and more abundant on acidic outcrops (F-R) than calcareous ones (R). Equally common and abundant (O-R) on mortared and drystone walls, and on hedgebanks. Fronds larger in more sheltered habitats and stipes longer when plants growing in deeper crevices. (c) Occurred from sea-level to limit of survey at 810m but becoming less common at higher altitudes. 21.3 A. onopteris (a-c) Recorded twice; once as rare on a stony hedgebank alt. 90m (Figure 4); and as rare on an exposed, acidic outcrop by side of road 30m. These specimens were determined on mean spore length (Roberts, 1979), which for ten spores from each specimen was 33um and 32um respectively. The mean lengths for 4 specimens of A. adiantum-nigrum from same area measured at the same time were 42, 37, 37 and 38um. Other records record that the species is more widespread in area, RS gives Loo Bridge, Glenflesk (c. W/08.81) and near the Sugarloaf, Caragh Lake (c. V/71.88) while At/as gives a pre- 1950 record for V/9.9 and a post-1950 record for V/7.9. 21.6 A. marinum Recorded for area and figured by Newman (1844) as numerous small plants on rocks by Upper Lake (specimen in BM); and by other authors since. Still present at site in 1980 (pers. comm. C. Mhic Daeid, 1980). A very unusual record for species as it normally grows on sea cliffs. Jermy et a/(1978) consider it requires salt-spray but | consider that it grows only near the sea because itis intolerant of frost. The mild climate of the area could then allow it to grow further inland than uSual. 2A7 A. trichomanes agg. (a) Widespread and common throughout area. Recorded for all grid squares. (b) Occurred most commonly on mortared walls and acidic outcrops with a few records for calcareous outcrops and hedgebanks (O-R). The apparently anomolous preference of this species for both base-rich and base-poor habitats is probably explained by the presence of two subspecies in the area (Atlas) though only one has so far been confirmed, see below. (c) Recorded for about 50% of monads below 457m but only for 10% from 480-810m. 21.7b_ A. trichomanes, subsp. quadrivalens Recorded (Atlas) for V/7.8, V/8.7, V/8.8, V/9.7, V/9.8 and V/9.9. 21.8 A. viride (a-c) Recorded once as occasional on damp rock outcrops at base of Devil's Ladder, Carrauntoohil (V/80.83) alt. 600m with Cytopteris fragilis. Old and new records indicate this is a very rare species of rock outcrops at high altitude. Curtis and Mhic Daeid (1981) record species for Mullaghanattin (c. V/73.77) and Mhic Daeid (pers. comm. 1989) records species in Horse’s Glen (c. V/99.81). RS records Torc and Mangerton Mountains while At/as vies a pre-1950 record for V/9.7. 21.9 A. ruta-muraria (a) Widespread throughout area but more common in the lowlands of the north and east than the higher land of the south and west. Recorded for all grid squares. (b) Practically confined to mortared walls where species was frequent to rare, with just two records each for acidic outcrops, drystone walls and calcareous outcrops. (c) Virtually confined to areas below 240m which is no doubt due in part at least to an absence of walls above this altitude. 21.11 A. ceterach (a) Records are widely but thinly scattered through the more lowland parts of the area. Recorded for grid squares V/7.7, V/7.9, V/8.9, V/9.7, V/9.9, W/0.8 and W/0:9. (b) Recorded 12 times as LF-R on mortared walls and once as abundant on a drystone wall. (c) Sea-level to 240m. 260 FERN GAZETTE: VOLUME 12 PART 5 (1983) 22.1 Athyrium filix-femina (a) Widespread and common throughout area. Recorded for all grid squares. (b) Equally abundant (F-O) in a range of habitats: hedgebanks, rocks by water; scrub; deciduous woodland and acid rock outcrops. Slightly less abundant (O-R) in heather moor and grass moor. Also present, but only spasmodically, in a range of other habitats including walls and conifer plantations. Species showed a marked preference for areas of habitats that were damp. It was much more abundant on rocks and walls here than in the drier eastern areas of the British Isles, eg. Willmot (1981). No specimens with red rachises were noted. (c) Decreased slightly in frequency with altitude but was still present in 50% of monads between 600-810m, the highest altitudinal range examined. 23.1 Gymnocarpium dryopteris Only record for area is Newman (1844) who gives Muckross, Kerry as a locality. This record considered an error as species is rare in Ireland and it has not been refound since. 24.1 Cystopteris fragilis (a-c) Recorded once as occasional on damp rock outcrops at base of Devil’s Ladder, Carrauntoohil (V/80.83) alt. (00m with Asplenium viride. .Recorded elsewhere as very rare at high altitudes on rocks: Mullaghanattin c. V/73.77 (Curtis and Mhic Daeid, 1981); Horse’s Glen c. V/99.80, Mangerton Mountain c. V/96.80 and one record for low altitude on rocks near Middle Lake, Muckross (all RS). 26.1 Polystichum lonchitis There are a few scattered old records for rocks at high altitudes in RS, At/as and at BRC. These are for: Horse’s Glen, Mangerton; above Lough Googh (c. V/84.83); Mullaghanattin(c. V/74.77) and cliffs south of Lough Erhagh (c. V/99.80). Recently Curtis and Mhic Daeid (1981) have reconfirmed the species in the Mullaghanattin area, and Mhic Daeid has communicated that the species is still in the Horse’s Glen (pers. comm. 1980). 26.2 P. aculeatum (a-c) Recorded for five monads in the region of Mangerton and Torc Mountains. Always only in small amount (R) amongst acid rock outcrops either unshaded as at Cloghfune (V/94.83) 375m, or in heather moor as at Horse’s Glen (V/99.82) 300m, or in deciduous woods as at Gortracussane (V/94.84) 120m and (V/95.84) 30m and at Torc Old Bridge (V/96.83) 180m. Recorded in RS for the Roughty Valley (V/9.7) and the Ballaghbeama Pass (c. V/76.77). 26.2x3 P. x bicknellii Recorded once for area post-1950 in Atlas for square V/9.7. 26.3 P. setiferum (a) Rare but widespread in lowland areas especially on limestone. Recorded for grid squares V/7.9, V/8.7, V/8.8, V/8.9, V/9.7, V/9.8, V/9.9, W/0.7 and W/0.9. (b) Occurred most commonly and abundantly on hedgebanks (LA-R) and in deciduous woodlands (F-O), with a few occurrences each on mortared walls and rock outcrops both acidic and calcareous. (c) Sea-level to 60m with one or two occurrences up to 240m but none above. 27.1 Dryopteris oreades (a-c) Recorded once (this survey) as a single patch at Dromderalough alt. 600m amongst rocks on heather moor, conf. C.R. Fraser-Jenkins, specimen in BM (Figure 5). No previous records traced for area, this is only fourth record in Ireland (At/as). 27.2 D. filix-mas (a) Widespread in lowland areas, recorded for all grid squares except V/7.7. (b) Most often encountered and most abundant in deciduous woods (O-R) and on hedgebanks (F-R), with isolated records at low altitude (R) in a range of habitats including walls, outcrops and moors. Generally less often met with than D. affinis, sporelings and young plants rarely seen. (c) Sea- level to 240m never recorded above. 27.3 D. affinis (a) Widespread and common, recorded for all grid squares. (b) Most commonly met with and most abundant (F-R) in deciduous woods, on hedgebanks and on acidic rock outcrops; slightly less common and less abundant (O-R) in grass moors and amongst rocks by water; with infrequent occurrences ina further range of habitats including coniferous plantations, mortared walls and drystone walls. (c) Occurred from sea-level to limit of survey at 810m but becoming less common at higher altitudes. Recently Fraser-Jenkins (1980) has separated the species into several subspecies; notes on specimens collected during survey and determined by him are given below. 27.3a subsp. affinis (a-c) Recorded for 10 monads scattered throughout the area; in deciduous woods, in grass heaths on acidic rock outcrops, on hedgebanks and on an earthed, drystone wall; altitudinal range 30- 375m. Selected records: occasional earthed, drystone wall Cloghera 120m (V/72.78); occasional hedgebank Gortnagan 150m (V/72.88); frequent acidic rock outcrops Boughil 240m (V/85.77); rare hedgebank Cummeenduvasig 150m (V/97.77); frequent oak wood near Torc Old Bridge 180m (V/96.83); and frequent grass heath near Loo Bridge 75m (W/07.80). FERNS OF KILLARNEY DISTRICT 261 27.3b ~~ subsp. stillupensis (a-c) Recorded once as rare amongst rock outcrops in Hag’s Glen (V/81.83) alt. 600m; first confirmed record for Ireland. 27.3c subsp. borreri (a-c) Recorded once as rare amongst rock outcrops on Mullaghanattin (V/ 73.77) alt. 600m. 27.4 D. remota Recorded once last century for area from Glen Flesk (specimen in BM, collected G. Wager) which is in W/0.8 not W/09 as indicated in Ati/as. 27.5 D. aemula (a) Widespread but local thoughout southern and western parts of area. Recorded for grid squares V/7.7,V/7.8, V/7.9, V/8.7,V/8.8, V/8.9,V/9.7,V/9.8 and W/0.7.(b) Most commonly found on rock outcrops, especially shaded and/or damp ones, where it was O-R in abundance, less common but equally as abundant on hedgebanks and in deciduous woods, with a few records each for moors, walls, scrub and conifer plantations. (c) Equally common from sea-level to limit of survey at 810m Recorded in Atlas for V/9.9 and W/08 post-1950. 27.8 D. carthusiana (a-c) Recorded as occasional in alder woods at eastern end of Ross Island alt. 24m (V/95.88). Records in RS, at BRC and in At/as provide about another five localities for species scattered through area, eg. records at BRC include an old record for Glen Flesk (c. W/06.85) anda recent one for Reen Point, Lough Leane (c. V/94.89). 27.9 D. dilatata (a) Widespread and common throughout area. Recorded for all grid squares. (b) Most common and most abundant (F-O) on hedgebanks, on acidic rock outcrops, and in deciduous woods; slightly less common and abundant (O-R) in conifer plantations, amongst rocks by water, in heather moors and grass moors; with scattered odd records for drystone walls, scrub and screes. (c) Equally common from sea-level to limit of survey at 810m. 27.10 OD. expansa Vida (1966) recorded this species from Kerry (V.C. H2) but Crabbe et a/ (1970) considered this record as extremely doubtful, and no other records are known. However, D. remota has been recorded which is an apogamous species probably originating from the hybrid between D. expansa and D. affinis (Atlas). D. expansa may then have occurred in area and may yet be found. 28.1 Blechnum spicant (a) Widespread and common throughout the area. Recorded for all grid squares (b)Occurred most commonly and abundantly (F-O) on acidic rock outcrops, on hedgebanks, in heather moors, in grass moors and in deciduous woods; occurred as abundantly but less commonly in conifer plantations and scrub; with a few records each for drystone walls and scree. (c) Recorded from sea-level to limit of survey at 810m, showed a slight increase in frequency with altitude. RS recorded species to 960m 29.1 Pilularia globulifera (a-c) Recorded once as locally frequent on bare mud at side of lake alt. 24m. All other records for area are around the Killarney Lakes. SUMMARY OF THE FLORA In this survey 38 species and hybrids of native British pteridophytes, nee one introduced speices (Se/aginella kraussiana), were found growing in the study area. Reliable references to a further 14 species and hybrids, either post-1950 or pre-1950, were encountered during the searches of data banks and herbaria. Thus overall some 52 species and hybrids have been found in the area which is about two thirds of the native species and hybrids that occur in Ireland (Jermy et a/, 1978). The area is thus rich in terns. This is further emphasised when the area is compared with the rest of Europe. Jalas and Suominen (1972) record the number of species and subspecies of ferns in 50km grid squares for the whole of Europe. They divide all their squares intoa number of classes of species richness, with the richest class having 43 or more. The study area falls into one of their squares which belongs to this richest class. Further fern taxa may be found in the area, particularly hybrids such as Dryopteris x tavel/ which are difficult to pick out in the field. However, it is considered that this total of 52 species and hybrids is an accurate representation of the true flora of the area. This is 262 FERN GAZETTE: VOLUME 12 PART 5 (1983) FIGURE 4. Asp/enium onopteris FIGURE 5. Dryopteris oreades FERNS OF KILLARNEY DISTRICT 263 thought to be so since there are no species which are common in adjacent or similar areas of Ireland that are not recorded for the study except for Se/aginella selaginoides. The reason for this species richness of the area is due to the presence of mountains near the western seaboard of Europe. This allows the coming together of species with montane, or arctic/alpine affinities with species of more southern or oceanic affinities (Table 3). It is notable that all the areas inthe British Isles that have the highest species numbers in Jalas and Souminen (1972) are also mountainous areas. However, examination of Table 3 shows that species of southern or oceanic affinities (Floristic elements 1-4) make a greater contribution to the species richness of the area than species of montane or arctic/alpine affinities (Floristic elements 7-8). Overall therefore the flora shows greater affinities with more southern and oceanic flora than more montane or arctic/alpine ones. This is further bourne out by the abundance of the species from the various floristic elements in the area. Five of the seven montane or arctic/alpine species have only one or two localities in the area whereas many of the southern or oceanic species have numerous localities. Although the montane, arctic/alpine influence on the flora is less important than the southern, oceanic influence, it is still significant. This is shown by comparing the Killarney fern flora with that of the Burren(Willmot, 1979). This is an area on the western seaboard of Ireland just to the north of the study area. It similarly shows a coming together of southern or oceanic species with montane or arctic/ alpine species. However, it does not have mountains and has only two species from floristic elements 7 and 8, whereas Killarney has 7 species. : Table 3 The phytogeographical affinities of the fern flora of the Killarney region Number of species " oe in: b Floristic elements Distribution (a) Killarney British (c) from Birks (1976) in Europe post-1950 (pre-1950) Islands Europe 1. Selaginella denticulata Mediterranean 1 4 10 2. Adiantum capillus-vener is Mediterranean/ 3 6 6 Atlantic 3. Phyllitis scolopendrium North Mediterranean/ 13 14 14 (= Asplenium scolopendrium) Atlantic 4. Hymenophyllum wilsonii Western seaboard 4 4 4 5. Cystopteris fragilis Widespread 9 (3) 14 14 6. Pteridium aquilinum Widespread 7 8 8 7. Polystichum lonchitis Widespread Montane/ 4 (2) 9 11 Sub-montane 8. Woodsia alpina Arctic/Alpine O (1) 6 6 Table includes all species reliably recorded for the study area which are included in Birks /1976). Examination of the relationship between the European distribution of the taxa noted as only doubtfully recorded for the area (Table 1) and those actually recorded might be thought to shed some light on the status of the doubtfultaxa. However, it does not. Two of the four taxa Equisetum x moorei and Dryopteris expansa were no doubt recorded by mistake for other taxa. The two other species have different distributions in Europe; Se/aginella selaginoides is an arctic/alpine species while Gymnocarpium dryopteris is widespread (Birks, 1976). This suggests that the latter is more likely to actually occur in the area than the former. However, the former is widespread in north Ireland while the latter has been recorded for only a few isolated localities in Ireland. 264 FERN GAZETTE: VOLUME 12 PART 5 (1983) The breakdown of the flora into its floristic elements (Table 3) allows a further analysis of it to be made. One can identify species that may turn up in the area by noting which species, of elements that occur in the area, are not present. If this is done for elements 1-6, there are 10 species that occur in the British Isles but not Killarney. These are: /soetes histrix (*) Asplenium billotii Ophioglossum lusitanicum (*) A. septentrionale O. azoricum (*) Gymnocarpium dryopteris Anogramma leptophylla (*) G. robertianum Adiantum capillus-veneris Dryopteris villarii It is not possible to suggest which of these species are more likely to turn up than others. All of them, except Adiantum capillus-veneris, are very rare in or absent from lreland. Four species, marked (*) above, are small plants, which are easily overlooked or mistaken for other species. Careful examination of the right habitats at the right times of the year might turn up species from this four. The origins of the Killarney flora is a problem that has interested botanists for a long time (Aalen, 1978; Jessen, 1948 and Mitchell, 1976). Discussion has centred around which plants if any survived the last ice age in the Killarney area and the routes taken by plants as they re-invaded the area after the ice. The geological evidence is that almost the whole of the upland part of the study area was covered in ice during the last ice age. This leaves two areas where plants would have survived within the area, the unglaciated area of the uplands (nunataks) and the lowlands in the north of the area. The climate in these would have been too severe for any species other than the montane or arctic/alpine ones to survive. Moreover there is no evidence in terms of fossils from this period that any of these species did survive in the region. A third area where species could have survived in the vicinity of the Killarney region is just off the present coast line of Kerry. Here there would have been areas of dry land due to the draw down of sea-level during the ice age. At present steep slopes extend off shore, so the draw down would have exposed sheltered, south-facing slopes. Mitchell (19 76) considers that these could have had warm enough microclimates for species other than montane or arctic/alpine ones to occur. However, Jessen (1949) considers that the pattern of the spread of species in Ireland, after the last ice age, in terms of timing and relative order or arrival of species offers no evidence of refugia for thermophilous species in Ireland. Overall, therefore, there is no conclusive evidence either way for the survival of species in the Killarney area or nearby during the last ice age. The question of routes taken by plants as they re-invaded the area after the ice age is as problematical as that of survival through the ice age. Two basic routes have been considered: overland through England and Wales or along an Atlantic seaboard exposed by the draw down of the oceans. However, the depth of water between southern Ireland and South Wales means that any route along an Atlantic seaboard would have been cut by the sea early in the post-glacial. It is unknown whether or not the climate of this Atlantic seaboard route would have been mild enough, before it was cut, to allow the southern elements in the flora to reach the area. However, the high representation of species liking warm and/or humid conditions in the area argues for an Atlantic seaboard route, at least for these species. The evidence of macrofossils and spores in peat bogs gives concrete evidence for the existence of certain species in the area later onin the post-glacial. Jessen (1949) records evidence for five species. Records of spores of Po/ypodium vulgare extend back into zone IV. Spores of Osmunda regalis and /soetes lacustris are recorded back to zone VI, while there is evidence of Dryopteris filix-mas and of Thelypteris palustris at Togherbane in zone VII. This last record is most interesting as this species only occurs at one locality now. FERNS OF KILLARNEY DISTRICT 265 ACKNOWLEDGEMENTS | should like to thank E.C. Mhic Daeid, Maura Scannell and D.A. Webb for advice on localities to visit, and Mary Gibby, C.R. Fraser-Jenkins, A.C. Jermy, C.N. Page and Anne Sleep for determining critical matter. | am grateful to the British Ecological Society who provided financial assistance for the 1981 field work. Finally | should like to thank for technical help in preparing the manuscript: A. Skinner, figures; P. Holdcroft, photographs and Mrs S.A. Bullock and Mrs P. Comber, typing. REFERENCES AALEN, F.H.A., 1978. Man and the landscape in Ireland. London. BIRKS, H.J.B., 1976. The distribution of European pteridophytes: A numerical analysis. New Phytol. 77: 257-287. BOUSKELL, F., 1905. Stray notes from South Kerry in 1903. 7rans. Leics. Lit. and Phil. Soc. 9: 39- 47. CRABBE, J.A., JERMY, A.C. & WALKER, S., 1970. The distribution of Dryopteris assimilis S. Walker in Britain. Watsonia 8: 3-15. CURTIS, T. & MHIC DAEID, C., 1981. Mullaghanattin, Glencar, Co. Kerry. 21st-23rd July. Watsonia 13: 261-262. FRASER-JENKINS, C.R., 1980. Dryopteris affinis: a new treatment for a complex species in the European pteridophyte flora. Willdenow/a 10- 107-115. HIND, W.M., 1857. Three days in Killarney. Phytologist (N.S.) 2: 25-28. JALAS, J. & SUOMINEN, J., 1972. Atlas Florae Europaeae Vol. 1 Pteridophyta. Helsinki. JERMY, A.C., ARNOLD, H.R., FARRELL, L. & PERRING, F.H., 1978. Atlas of ferns of the British Isles. London. JESSEN, K., 1949. Studies in late-Quaternary deposits and flora history of Ireland. Procs. Roy. Irish Acad. Ser. B. 52: 85-290. KELLY, D.L., 1981. The native forest vegetation of Killarney, south-west Ireland: An ecological account. J. Ecol. 69: 437-472. MITCHELL, F., 1976. The Irish landscape. London. MORE, A.G.. 1876. Lycopodium inundatum in Kerry. Journal of Botany 14: 373. NEWMAN, E., 1844. A history of British ferns & allied plants. London. O’MAHONY, A., 1980. Some recent plant finds in the Kenmare-Kilgarvan area of South Kerry (H1). Bull. /r.Biogeog.Soc. 4: 41-45. PRAEGER, R.L., 1934. The Botanist in Ireland. Dublin. RASOR, J., 1882. Notes on the Ferns of Killarney. Science Gossip 18: 162-3. ROBERTS, R.H., 1979. Spore size in Asp/enium adiantum-nigrum L. and A. onopteris L. Watsonia 12: 233-238. RYAN, P., 1963. The soils of Ireland. /rish Forestry 20: 46-60. SCULLY, R.W., 1916. Flora of County Kerry. Dublin. SIMON, T. & VIDA, G., 1966. Nene Angaben zur Verbreitung der Dryopteris assimilis S. Walker in Europa. Annis. Univ. Scient. bpest. Rolando Eotvos, Sect. biol., 8: 275-284. TURNER, J.S. & WATT, A.S., 1939. The oakwoods (Quercetum sessiliflorae) of Killarney, Ireland. J Ecol. 27: 202-233. . WATTS, W.A., 1963. Late-glacial pollen-zones in western Ireland. /r. Geog. 4: 367-376. WILLMOT, A., 1979. An ecological survey of the ferns of the Burren, Co. Clare, Eire. Fern Gaz. 12: 9-28. WILLMOT, A., 1981. An ecological survey of the ferns of Berwickshire, Scotland. Fern Gaz. 72: 133-154. WRIGHT, W.B., 1927. The geology of Killarney and Kenmore. Dublin. 266 REVIEW REVIEW A MONOGRAPH OF THE FERN GENUS PLATYCERIUM (POLYPODIACEAE) by E. Hennipman and M.C. Roos. Verhandelingen der Koninklijke Nederlandse Akademie van Wetenschappen, Afd. Natuurkunde, 2nd series, part 80. North-Holland Publishing Company, 1982. 126 pp., 4 colour and 8 monochrome plates, 31 figs, 170 x 243mm. ISBN 0-4448 5569-6. Price: Dfl. 75.-(c. £14.00). This is a beautifully produced and fully illustrated treatment of Stag’s-horn ferns. The first part covers the taxonomic history and systematic position of the genus, with details of the morphology, anatomy and habitats of the species. The phylogenetic relationships of the species are investigated and their geographical distributions are discussed in relation to the proposed phylogeny. The second part gives a helpful key, followed by full descriptions of all the species, which are presented in alphabetical order. Valuable notes cover points such as habitat, similarity to other species, native names and the fascinating uses of parts of these spectacular ferns: how! wish the authors had commented on the taste of the tea that can be made from Platycerium holttumii! Certainly those horticulturalists who talk to their plants will now be able to address their ‘Pepelegbuku’ or Tama halota’ in a more ethnic way. Two short sections also give useful advice on the collection of these ferns from the wild. The phylogenetic and biogeographic sections make use of the method of classification termed cladistics, which may appear daunting to anyone not already acquainted with the method. However, the clarity and consistency brought by this use of cladistics to the recording of characters (in this case features of morphology, anatomy and frond habit) are readily apparent. For example, the reason for assigning the so-called plesiomorphic (generalised) vs. apomorphic (derived) states for any character is given. Observations so recorded, in the form of a data matrix and diagram, can therefore be used readily by others when investigating further aspects, or even alternative methods, of classification. Additional data, for example on chromosomes, may be added easily to those provided by Hennipman and Roos. Indeed, information from biochemical analyses and cytology could be used to test their proposed phylogeny. In the developing discipline of cladistics, there is inevitably some difference of opinion as to how the most parsimonious cladogram should be constructed and interpreted. Hennipman and Roos have stated clearly their method and reasoning, and | would criticize only two points. Firstly, a rather important proof error seems to have escaped detection. The last figure on p. 36 should surely have a solid, rather than open, circle to symbolise ‘a’. Secondly, since two alternative transformation series are proposed for the base frond composite character ‘12,13’, it would seem preferable to have two cladograms (each including one of these alternatives) with any. difference between them discussed. This monograph will be of great interest both to herbarium taxonomists and horticulturalists. J. M. CAMUS FERN GAZ. 12(5) 1983 267 THREE NEW DAYOPTER/S HYBRIDS FROM SPAIN AND THE CANARY ISLANDS MARY GIBBY Department of Botany, British Museum (Natural History), Cromwell Road, London SW7 5BD. CARL-JOHAN WIDEN Institute of Pharmaceutical Chemistry, University of Kuopio, 70101 Kuopio, Finland. ABSTRACT Three new Dryopteris hybrids: D. x gomerica = aemula x guanchica and D. x cedroensis = guanchica x oligodonta from Gomera, Canary Islands and D. x fraser- jenkinsii = affinis with uncertain parentage, from Oviedo, N. Spain, are described with data on their chemistry, cytology and morphology. INTRODUCTION During collecting trips to the Canary Islands (1974) and Spain(1976), Fraser-Jenkins discovered three new Dryopteris hybrids. These have been investigated morphologically, cytologically and chemically in an attempt to determine their parentage. Dryopteris x gomerica Gibby & Widén, hybrid.nova. (D. aemula (Aiton) O.L Kuntze x D. guanchica Gibby & Jermy) Holotypus: Canary Islands, La Gomera, El Cedro, Mia Quemada, 1100m alt. With D. aemula, D. guanchica, D. oligodonta Pichi-Serm. 31 March 1974, C.A. Fraser-Jenkins 4212 (BM). Planta inter parentes putatos in morphologia intermedia. Lamina triangularis, glandulosissima. Squamae stipitis basi longae, contractae, brunneae. Sporae abortivae. The plant is intermediate in morphology between its putative parents. The lamina is triangular and very glandular. The scales at the stipe base are long, tapering and brown in colour. Spores are abortive. The hybrid is triploid with 2n = 123, and at first metaphase of meiosis shows approximately 41 bivalents and 41 univalents; D. guanchica is an allotetraploid species, and pairing in the hybrid is allosyndetic. Diploid D. aemu/a is one of the parents of D. guanchica (Gibby et a/, 1978). The phloroglucinol composition of the hybrid is similar to that of D. guanchica, and the characteristic compounds from D. aemula appear to be suppressed (Gibby et a/, 1978). During a cytogenetic investigation of D. guanchica in 1973, hybrids of the constitution D. aemula x guanchica were synthesized, prior to the discovery of this hybrid in the wild. The wild and synthesized hybrids are very similar in morphology, cytology and phloroglucinol composition. The site where they hybrid was discovered in La Gomera is the only locality in the Canary Islands where the two parents are known to growtogether. Both D. aemu/a and D. guanchica are recorded from north-west Spain, but as yet they have seldom been found growing together (Fraser-Jenkins, pers. comm.) Dryopteris x cedroensis Gibby & Widén, hybrid.nova. (D. guanchica Gibby & Jermy x D. oligodonta Pichi-Serm). Holotypus. Canary Islands, La Gomera, El Cedro, Mia Quemada, 1100m alt With D. aemula, D. guanchicaand D. oligodonta. 31 March 1974, C.R. Fraser- Jenkins 4211 (BM). 268 FERN GAZETTE: VOLUME 12 PART 5 (1983) Planta inter parentes putatos in morphologia intermedia. Frondes grandes, plus quam 1m longae, earum ad modum D. oligondontae \axae. Squamae in stipite usque ad 2cm longissimae. Pinnulae eis D. oligodontae plus divisae, segmentis ultimis D. guanchicae similibus dentatis. Sporae abortivae. The plant is intermediate in morphology between the putative parents. The fronds are large, over a metre in length, and lax like those of D. oligodonta, and the scales on the Stipe are long, up to 2cm. The pinnules are more divided than those of D. ol/igodonta, and the ultimate segments are toothed like those of D. guanchica. Spores are abortive. The cytology and phloroglucinol composition of this hybrid have been investigated (Gibby et a/, 1978). The plant is pentaploid with 2n = 205. This finding was unexpected; D. guanchica is tetraploid and D. oligodonta is diploid — a hybrid between these species would be expected to be triploid. The result obtained can be explained if the hybrid has resulted from the fusion of an unreduced gamete from D. guanchica (164 chromosomes) with a normal gamete (41 chromosomes) from D. oligodonta. The meiotic analyses obtained are compatible with such an hypothesis; these show up to 80 bivalents with 43 univalents. The phloroglucinol composition of the plant is similar to that of D. guanchica. Dryopteris x fraser-jenkinsii Gibby & Widén, hybrid. nova. Holotypus: Spain, Oviedo; beside road from Aviles to Ribadeo, by bend in road, 1km S. of Canero, E. of Luarca. 50m alt. Mixed Pinus, Quercus and Castanea forest, with D. affinis (Lowe) Fraser-Jenkins, D. dilatata (Hoffm.) A. Gray and D. guanchica. C.R. Fraser-Jenkins 4899, ex. hort. Chelsea Physic Garden, no. CPG 2189. Hybrida in morphologia D. affini similis. Pinnulae autem plus dissectae ad costam fissae, segmentis rotundatis dentatisque. Sporae pleraeque abortivae, vix 5% ut videtur bene evolutae et haec grandes, rugosae eis D. affinis similes sed nonnumquam subspineae vel spinulosae. The hybrid is similar in morphology to D. affinis, but the pinnules are more dissected, being cut at least halfway to the midribs, and the pinnule segments are rounded and toothed. Spores mainly abortive, less than 5% appearing normal, being large, rugose and like those of D. affinis, but often bear small spines or projections (Figure 1). The hybrid is tetraploid with 2n= 164. This has been confirmed by examination of root tip mitosis. Investigation of meiosis in spore mother cells reveals that the hybrid is partially apomictic, like D. affinis, and produces sporangia with eight spore mother cells that show 164 bivalents at metaphase |. More frequent are 16-celled sporangia, where each spore mother cell shows 164 univalents at metaphase | (Figure 2); anaphase separation in these cells is unbalanced and leads to the formation of abortive spores. The lowpercentage of good spores reflects the low frequency of eight- celled sporangia. Sporangial contents have been sown and germination of prothalli obtained, presumably from the unreduced spores (with 164 chromosomes) that result from eight-celled sporangia. The prothalli give rise to sporophytes apogamously. The phloroglucinol composition of D. fraser-jenkinsi/ is givenin Table 1, together with those of D. affinis, D. dilatata and D. guanchica. The hybrid is very different from D. affinis, and shows closest similarity with D. guanchica. The parentage of this hybrid cannot firmly be established. It is clearthat D. affinis must be one parent, as the hybrid is very similar to this in morphology, and it shows apomictic behaviour. This is typical of hybrids involving the apomictic D. affinis eg. D. x tavellii Rothm. (Manton, 1950). The second parent is in doubt, but probably belongs to the D. dilatata complex. Both morphological and chemical comparisons support this hypothesis. D. dilatata and D. guanchica, with which the hybrid was growing, are possibilities. Both are allotetraploid species, and hybridization of either with a diploid form of D. affinis would result in a tetraploid hybrid. During normal meiosis no pairing of chromosomes would be expected in such a hybrid; D. x fraser-jenkinsii shows this 269 NEW DRYOPTERIS HYBRIDS FROM SPAIN (%LZ'L) saseq adiis GQ"| Wo} pauleyqo }9e4}x9 Jauia BWE'QZ WOd} palpnis sem UOIZISOdWOD jOUION|BosojYyd au] (v "GZ6L “J2 18 UBPIN (Z "$}|NSOJ Ja! |429 0} S}UBWIPUaWIY (€ *(s@wopnasd ‘Gq Qqns) 9/61 “|e 18 UPI (L 4 L8LZ/p96L YL - (+) - g(t) + (+) — +4 +++ (+) ++ (+) — 991 eolyouent "Gg edoin3g os (+) <= e(t) + (+) +++ ++ +++ eo = (+) + v9 7 RERE LIP ‘a ++ + ett aici ay fe we @ = oe ae (+) €ZL UUs? O (DM) + + ett gt t+ = = = ROGL , S/Ulse ag €62 YL + + ett ett = = = = = = J) -ée ruse a 6812 DdO = oe (+) + ++ =e - + + + ++ + — »9l JIsulyUal-sJases, X “GQ 7 a) = : 3 : = = i, ~ ~ S Ry e ze > > > ig g S He 2) 5, a s ® © Af oh o > e 0. q ® 5 PB} a 3 3 £ o a a. a a o Q. =| = ry ie) Ss v, v. ame 3 3 G se se 2 ° 4 x) < a a a & a = = = fe) a om 3 so) a a =| = = a =r = oe a Oo 5 rot a e 5 bes 5 5 S| 3 a °. °. v Ww N = 2 9 o. o. oO : 3 : = a bo = -_- o ba © | my 2 S & *punoj }OU — ‘s}UNOWEe a9eJ} Ul JUaSsid (+) ‘%OL — G s}unowWe |jewWs Ul JUaSaId + '%OZ — OL S}UNOWe a}eJ9pOW UI! JUaSaId +4 /%OZ s}UNOWe abe] Ul JUASAId +44 :(UIpidse apnsd) a4n}xilW ;OUuLIDN|bHosojyd ay} 40 sabe}Usdsad se pa}e;nojed syUNOWe aAI}e}1}2UeNbD-1 Was "saideds jejUuased aAijeind pue //susyual-sases, Xx SI4A1GOAIG Ul SAAIWEALJap [OUIDN|Hos0jUd “L JIGWL 270 FERN GAZETTE: VOLUME 12 PART 5 (1983) FIGURE 1. Spore of Dryopteris x fraser-jenkinsi1, x 670. FIGURE 2. Meiosis in a 16-celled sporangium of Dryopteris x fraser-senkinsii (acetocarmine squash preparation showing metaphase 1), x 1000. result. Hybridization between a triploid form of D. affinis with a diploid species like D. aemula would also result ina tetraploid hybrid. However, on morphological grounds it is unlikely that D. aemu/a is a parent of the hybrid, nor was this species growing in the vicinity. ACKNOWLEDGEMENTS We are grateful to Kathryn Kavanagh for the Latin diagnoses. REFERENCES GIBBY, M., WIDEN, C.-J. & WIDEN, H.K., 1978. Cytogenetic and phytochemical investigations in hybrids of Macaronesian Dryopteris, (Pteridophyta: Aspidiaceae). Pl. Syst. Evol. 130: 235-252: WIDEN, C.-J., LOUNASMA, M., JERMY, A.C., EUW, J.v. & REICHSTEIN, T., 1976. Die Phloroglucide von zwei Farnhybriden aus England und Schottland, von authentischem “Aspidium remotum”’. A. Braun und von Dryopteris aemula (Aiton) O. SHALES aus Irland. Helv. Chim. Acta. 59: 1725-1744. WIDEN, C.-J., LOUNASMAA, M., VIDA, G. & REICHSTEIN, T., 1975. Die Phidragiveitle von drei Dryopteris-Arten sowie zwei Arten von Madeira und den Kanarischen Inseln zum Vergleich. Helv. Chim. Acta. 58: 880-904. FERN GAZ. 12(5) 1983 271 ASPLENIUM BOURGAEI A NEW ADDITION TO THE FLORA OF EUROPE W. GREUTER, R. PLEGER, TH. RAUS, B. ZIMMER & J.J. GREUTER Botanischer Garten und Botanisches Museum, Berlin-Dahlem ABSTRACT Asplenium bourgaei, hitherto known only from a few localities on the East Aegean islands, along the southern coast of Anatolia, and in a single site in Lebanon, has been discovered on the island of Karpathos. It is here illustrated, and notes on its ecology are given, notably on its remarkable poikilohydric habit. INTRODUCTION Asplenium bourgae! Milde (1866) is a seldom collected fern whose known distribution is confined to the area bordering the eastern Mediterranean basin, and to a few off- shore islands. Meyer (1962) has reviewed the literature dealing with this species, and has provided a distribution map of the few localities then known. Since, A. bourgaei has been found in various other localities in S. Anatoloia (Davis, 1965; Huber-Morath, 1966; Demiriz et a/, 1969; Sorger, 1971), and on two islets of the Kastellorizo group belonging to Greece (Greuter, 1979). Afurther locality on the island of Rhodes has, in all probability, been reported by Fink! (1962) under the certainly erroneous designation “Asplenium fontanum (L.) Bernh.”’ On the other hand, Mouterde (1966) has rectified an earlier erroneous record from Lebanon (Mt. Sannin), so that the long known locality in the Litani gorge remains the only one for that country. Rather suprisingly, no report from Cyprus has yet been published. The total known distribution of A. bourgaei, including our new finding (see below), is mapped in Figure 1. ea oe -*¥ > { Ss &G Se oN -Rhodo® Pe ae 0 ; Ga @ kar athos °@ P SaY OR WwA Cyprus LEBANON ° 50 100 150 200 250 300 on — FIGURE 1. The total known distribution of Asp/enium bourgaei. Milde (1867) speaks of Asplenium bourgaei as a ‘‘species pulcherrima”’, meaning an excellent species. It is indeed a taxonomically isolated, presumably old relic taxon, and has a diploid chromosome complement according to Reichstein (1981) who cites an unpublished chromosome count by Lovis based on material collected by Reichstein in 1964 near Cakirlar (misspelt ‘“‘Tcharyklar’’ on Bourgeau’s label of 1860 272 FERN GAZETTE: VOLUME 12 PART 5 (1983) and ‘‘Tscharydar”’ in Davis, 1965). Meyer believes that it is related to another very local old diploid, Asplenium sahandiezii (Litard.) Rouy from the Verdun gorge in southern France. It differs considerably, however, from the latter, not only in its more dissected foliar lobes with a less coarse dentation of a different shape, but also in the coloration of the rachis and stipe (see below). OBSERVATIONS Asplenium bourgae/ has been found by us in a single feecaltiys on the South Aegean island of Karpathos (Dodekanisos, Greece). This appears to be the first finding in Europe as defined, for floristic purposes, in ‘Flora Europaea” (Tutin et a/ 1964; see also Greuter et a/, 1981). The exact data are as follows: N. slopes of Mt. Asprovouno, between Mt. Kalilimni and the Apella Bay, 250m above sea level, 18 v 1982, W. & J.J. Greuter, Pleger & Raus 19162; ibidem, 9 viii 1982, W. & JJ. Greuter & Zimmer 19482. | Several individuals were growing in shaded clefts underneath big boulders of calcareous rock, inpermanent shade but without any special water or moisture Supply. The best developed specimens were considerably larger than any that had so far been found in this species (Figures 2 and 3). The frond length attained 28cm, whereas the normal measurements as given by Davis (1965) are 4-7cm. Such luxuriant specimens show particularly well the distinctive features of the species, especially the characteristic dissection of the frond. It is notable that the stipe and lower half of the 5 cm FIGURE 2. Luxuriant specimen of Asplenium bourgae/ from Karpathos (Greuter et al, 19162). ASPLENIUM BOURGAEI IN EUROPE 273 5 cm Po FIGURE 3. Single frond of the specimen in Figure 2. rachis is dark-coloured on at least one side, but may be green distally on the opposite side (sometimes on the ventral sometimes on the dorsal side). This feature, which seems to be constant within the species, offers an easy means to distinguish even its most dwarfed specimens fromthe somewhat similar Asp/enium jahandiezii where the whole rachis and stipe are green except at the very base. Asplenium bourgaei is, throughout its range, a typical representative of the coastal Mediterranean belt. With a single exception, to be discussed below, all its known localities lie between sea level and an altitude of 900m. Surprisingly Davis (1965) gives the altitudinal range as “‘1000-2000m”, although no single record from such altitudes yet exists. There is one completely aberrant indication of an occurrence at 2500m on Geyik Dag, which in our opinion is in need of confirmation(it might be due either to mis-identification, or to a confusion of feet with metres). FIGURE 4. a, dry, but living fronds of Asp/enium bourgaei collected in Karpathos in August 1982; b, the same fronds having recovered their normal shape after being kept for 2 days in moisture in a plastic bag. 274 FERN GAZETTE: VOLUME 12 PART 5 (1983) The second gathering made in August enabled us to observe a quite remarkable feature. The fronds were at that season completely dry and brittle, with their segments Spirally inrolled so as to conceal the sori and with the veins protruding as prominent, thread-like ridges (Figure 4a). They were nevertheless green, and completely recovered their normal form and function when kept under high moisture, in a plastic bag, for 2-3 days (Figure 4b). A similar observation was made by Reichstein(ms. note in the Berlin Herbarium) in November 1964 in the Cakirlar locality: after heavy rain two days earlier some plants, growing in rain shelter under rocks, were completely dry, while others growing nearby, where the soil had been soaked, had unrolled and turgescent fronds. Such a poikilohydric habit is known for several ferns colonizing dry open habitats such as Chei/lanthes and Ceterach species. It is usually associated with a dense scaly indumentum. Christ (1910: 83) mentioned this faculty of fern fronds to completely dry out without losing viability, while discussing epiphytic ferns, but referred to it only incidentally when commenting on xerophytes proper. In our opinion, the scaly cover is a protecting device against irradiation, not desiccation, whereas the poikilohydric habit is a genuine xerophytic feature — to be found, less pronouncedly, in other Asplenium species such as A. trichomanes. |It may be mentioned that fronds collected in August and soaked before pressing shed their spores in the herbarium, which apparently demonstrates that the maturation of the spores is completed, at least in part, in the desiccated stage. REFERENCES CHRIST, H., 1910. Die Geographie der Farne. Jena. DAVIS, P.H., 1965. Flora of Turkey and the East Aegean Islands. Vol. 1. Edinburgh. DEMIRIZ, H., TUTEL, B. & AYDIN, A., 1969. Turkiye flora ve vejetasyonu uzerinde arastirmalar: IV. Turkiye Pteridophyt'lerine ait yeni materyaller: Filicales. (Studia ad floram et vegetationem Turciae pertinentia: |V. New materials to the Pteridophytes of Turkey: Filicales.) /stanbul Univ. Fen Fak. Mecm. Seri B, 34: 137-181. FINKL, A., 1962. Beitrage zur Kenntnis der Flora der Insel Rhodos. Acta Albertina Ratisb. 24: 101- 120. GREUTER, W., 1979. The flora and phytogeography of Kastellorizo (Dhodhekanisos, Greece). 1. An annotated catalogue of the vascular plant taxa. Wi/ldenowia 8: 531-611. GREUTER, W., BURDET, H.M. & LONG, G., 1981. Med-Checkliist. |. Peteridophyta. Geneve & Berlin. HUBER-MORATH, A., 1966. Beitrage zue Kenntnis der anatolischen Flora Ill. Bauhinia 3: 7-45 MEYER, D.E., 1962. Uber neue und seltene Asplenien Europas. Ber. Deutsch. Bot. Ges. 75: 24 34. MILDE, J., 1866. Filices criticae. Dritter Artikel. Bot. Zeitung 24: 384-385. MILDE, J., 1867. Filices Europae et Atlantidis, Asiae Minoris et Sibiriae. Lipsiae. MOUTERDE, P., 1966. Nouvelle flore du Liban et de la Syrie. Vol. 1. Beyrouth. REICHSTEIN, T., 1981. Hybrids in European Aspleniaceae (Pteridophyta). Significance. recognition, genome analysis, and fertility; checklist of species and hybrids. Description of some new hybrids and cytology of several already known hybrids. Bot. Helv. 97: 89- 139. SORGER, F., 1971. Beitrage zur Flora der Turkei!. Mutt. Bot. Arbeitsgem. Oberosterr. Landesmus. Linz 3 (2): 1-98. TUTIN, T.G., HEYWOOD, V.H., BURGES, N.A., VALENTINE, D.H., WALTERS, S.,. & WEBB, D.A. 1964. Flora Europeae. Vol. 1. Cambridge. NOTE ADDED IN PRESS Since the above account was written, H. Runemark, Lund, has kindly communicated three additional localities for Asp/enium bourgaei from unpublished records. These are: SW Turkey, province of Mugla: Fethiye area, N. of Gocik, 100-300m, 1974, Runemark & Bentzer 29503; Fethiye area, 2km SE of Yelcegiz, 50-100m, 1974, Runemark & Bentzer 29452; Marmaris, NW of the town, 20-50m, 1974, Runemark & Bentzer 29414. All lie within the general species area already indicated on the map (Figure 1). FERN GAZ. 12(5) 1983 275 CONTRIBUTIONS TO THE MORPHOLOGY OF THE TECTARIOID FERN STENOSEMIA SUBHASH CHANDRA National Botanical Research Institute, Lucknow, India. ABSTRACT The monotypic genus Stenosemia Presl exhibits all the characters of Tectarioideae and has several characters in common with those of 7ectaria Cav. (especially the free-veined species) and tectarioid derivatives. Based on morpho-anatomical comparisons, it is suggested that Stenosemia is probably evolved in similar lines of reduction and simplification of characters as those of Camptodium Fée, Hemigramma Christ, and Psomiocarpa Presl. It has also been observed that Stenosemia is more nearly allied to 7ectaria than to Heterogonium Presl and possibly derived from a group of free-veined species of 7Jectaria and not from Ctenitis C.Chr. as suggested by Copeland (1947). Thus, it seems apparent that Stenosemia is a tectarioid fern. INTRODUCTION Stenosemia Presi is a genus of tectarioid fern ranging from Solomon Islands across Malaya and the Philippines(Copeland, 1960). It consists of three species (Ching, 1940; Copeland, 1947, 1960). Holttum (1949) transferred S. pinnata Copel. toHeterogonium: H. pinnatum (Copel.) Holtt. According to Holttum (1975) and Hennipman (1977) S. dimorpha Copel. (Copeland, 1955) is a Bo/bitis: B. quoyana (Gaud.) Ching. It appears thus that Stenosemia is probably a monotypic genus (Price, 1972), the type and sole species being S. aurita (Sw.) Presl. It is a rare and small limestone fern growing in shady places at an elevation of 250 metres. Various authors have given different taxonomic treatment to the genus. Christensen (1938) regards, Stenosemia a tectarioid derivative and placed it in his tribe Dryopterideae of the subfamily Dryopteridoideae. Ching (1940) lists this genus under the tribe Aspidieae of the family Aspidiaceae. Copeland (1947) amalgamated Ching’s tribes and placed Stenosemia along with many other genera in asingle large family Aspidiaceae. Holttum (1968), Nayar (1974), Crabbe et a/(1975)and Lovis(1977) placed the genus in a subfamily Tectarioideae. Copeland (1947), pointed out its relationship to Heterogonium and suggested its derivation directly from Ctenitis. However, Holttum (1968) considers Stenosemia to have been derived from a true 7ectaria. Hitherto, the morphology of the genus has not been known, except for a few taxonomic details (Copeland, 1905, 1929, 1947, 1960). Attributes of the sporophyte are here studied and assessed relative to the taxonomic status of the genus. MATERIAL AND METHODS The present study of sporophyte morphology is based on material collected by the author in April, 1978 from the Philippines (Luzon: Antimonan, Quezon National Forest Park; S. Chandra 126, PUH), fixed in F.A.A. and stored in 70% alcohol. Anatomical observations recorded here are based on microtome sections stained with safranin and Fast green. Stelar organisation of the rhizome has been studied mainly from serial transections and reconstruction based on camera lucida tracings of the outline of the vascular cylinder. Spore morphology is based on acetolysed samples mounted in glycerine jelly. OBSERVATIONS Rhizome The rhizome is erect, short (c. 5mm thick), infrequently branched, and covered NBRI, Research Publication No. 171 (N.S.) 276 FERN GAZETTE: VOLUME 12 PART 5 (1983) with closely placed dark brown paleae. The roots are thick, wiry, rust-coloured and profuse between leaf bases. The paleae are basally attached (Figure 1b), clathrate and with thin-walled hyaline marginal cells. The thickening of the walls gradually progresses from the apex downwards (Figure 1d). They are narrowly lanceolate with broad base and each one gradually tapers to a uniseriate club-shped glandular apex. Rarely the paleae are non-glandular (Figure 1c). The paleal margins (Figure 1e, f) are smooth or nearly so and bear long, unicellular as well as multicellular (2-4 cells) hairs FIGURE 1. Morphology of Stenosemia aurita (Sw.) Presi: a, portion of stelar cylinder of the rhizome; b, mature palea; c, apex of matue palea; d, young palea; e, margin of mature palea; f, margin of young palea; g, mature sporangium; h, foliar hairs; i, venation pattern, j, k, upper and lower foliar epidermis (‘'L’’ = leaf trace; ““LG’’ = leaf gap; ‘’R’’ = root trace; ‘’S”’ = third row of stalk cells; “VS” = vestigial strands). MORPHOLOGY OF STENOSEMIA 277 which are sparsely distributed. The terminal cells of the multicellular hairs are usually club-shaped and non-glandular (Figure 1f); rarely the terminal cell is glandular (Figure 1e). In young paleae (Figure 1d), such hairs are more profuse towards the posterior half and points towards the base of the palea. Surface hairs like those on the paleae of Ctenitis are absent. The development of the paleae is as in other tectarioid ferns (Kaur, 1973, 1978; Chandra, 1976). Structurally, the rhizome is soft and parenchymatous and contains starch deposits. The epidermal as well as ground tissue cells are thin-walled. Sclerenchyma strands as found in Ctenitis (Kaur, 1973) andin most species of 7ectaria (Chandra, and Kaur, 1976), are absent in Stenosemia as in free-veined 7ectaria (Chandra and Kaur, 1976), Camptodium (Chandra, 1976), Hemigramma (Chandra and Salgado, 1978), and Psomiocarpa (Zamora and Chandra, 1977). Vascular cylinder of the rhizome is a radiosymmetric dictyostele (Figure 1a) similar in basic plan to that of Ctenitis (Kaur, 1973), Tectaria (Chandra and Kaur, 1976), Camptodium (Chandra, 1976), Psomiocarpa (Zamora and Chandra, 1977), and Hemigramma (Chandra and Salgado, 1978). The leaf gaps(Figure 1a ‘‘LG’’) are closely placed and prominently overlapping so that the stele has 4-5 nearly cylindrical to narrowly ribbon-shaped meristeles in transectional view; they are ellipsoidal, nearly spindle-shaped openings in the vascular cylinder and taper upward to a broadly anterior end, while the posterior region slightly tapers downwards and becomes truncate because of the large median basal root trace. The leaf traces (Figure 1a ‘'L’’) are highly dissected by prominent longitudinal slits usually into 3 or 4 narrow, cylindrical leaf trace strands. The main leaf trace strands are given off one to either margin of the leaf gap near its middle plane, these further divide into four leaf trace strands; of which the abaxial ones unite to form a reticulate leaf trace consisting of 3 leaf trace strands(Figure 1a ‘’L’’). Thus the number of vascular strands in each leaf is much reduced in Stenosemia as in Psomiocarpa (Zamora and Chandra, 1977). Characteristic association of the root trace (Figure 1a “R‘’) on the posterior base of the leaf gap is similar to that reported for other tectarioid ferns (Kaur, 1973; Chandra and Kaur, 1976; Zamora and Chandra, 1977). Incontrast to Ctenitis, Tectaria and Camptodium, none of the root traces are associated with the leaf trace bundles as in Psomiocarpa. Roots other than those associated with leaves are absent as in most tectarioid ferns. A peculiar feature of the vascular cylinder of Stenosemia is the possession of occasional blind ending stump-like, short vascular strands (Figure 1a ‘‘VS"’) between the leaf trace strands; these strands resemble leaf trace strands at origin, follow a similar course a short distance parallel to them andend blindly. These abruptly ending vascular strands are apparently the remnants of reduced leaf trace strands as also observed in Jectaria devexa (Chandra and Kaur, 1976) and Camptodium (Chandra, 1976). The xylem tissue of the vascular cylinder is massive, 6- to 8-celled thick of rather large tracheids interspersed with few thin-walled parenchyma cells. A thin sheath (1 - 2 layers) of small parenchyma cells envelops the xylem tissue except at free ends. Protoxylem elements are usually present at the extreme ends. The phloem is not continuous and consists of usually one or two layers of narrow, thin-walled small cells. Pericycle is very prominent, consisting of usually one layer of large cells and is continuous around the vascular tissue. The endodermis is uniseriate and consists of radially compressed, thin-walled elongated cells. In contrast to Camptodium, no sclerenchyma sheath surrounds the meristele as in Psomiocarpa (Zamora and Chandra, 1977). 278 FERN GAZETTE: VOLUME 12 PART 5 (1983) Leaf Leaves are arranged spirally around the rhizome, stipitate, non-articulated to the rhizome and dimorphic (Figure 2). The stipes of the sterile fronds are dark-polished and shorter than the stipes of the fertile ones. In contrast to Ctenitis, the stipes of the sterile fronds bear sparse, mostly deciduous dark, linear paleae at the base only as in Jectaria, Camptodium, and Psomiocarpa. The stipes are cylindrical and beset with characteristic ctenitoid hairs as found on the lamina surface. Structurally, the stipe (at the base) is similar to the rhizome except for the presence of a distinct, sheath (4-6 layers) of thick-walled narrow cells beneath the epidermis. The cortical sclerenchymatous sheath is interrupted laterally (at the region of the aerating tissue) by parenchymatous tissue in the basal portion of the stipe. The vascular supply to the stipe consists of usually three cylindrical strands (Figure 3). The adaxial strands are much larger than the abaxial one. The lamina is thin, membranaceous drying dark or somewhat opaque, deltoid ovate, broadly trifoliate in plan (Figure 2); its central part deeply pinnatifid with entire, or obtusely serrate, lanceolate lobes; lateral pinnae very unequal sided, with lowest basiscopic segment pinnatifid, strongly developed as in Ctenitis, Tectaria (Holttum, 1968), Camptodium (Chandra, 1976) and Psomiocarpa (Zamora and Chandra, 1977). The lowest segments rarely becoming free pinnae; veins usually forming very narrow costal areolae only (Figure 11), the remaining veins free, without included veinlets. Similar ctenitoid hairs as found on the stipe occur profusely on both the surfaces of the lamina (especially on the veins) and the rachis (Figure 1h). In the leaf segments, both the epidermises are chlorophyllous and with thick outer walls. In surface view, the upper epidermis is composed of broad, regular hexagonal cells (Figure 1k). In most cases the walls between the two hexagonal cells are dissolved and they look like large, elongated cells with regular straight walls, i.e., the epidermal cell walls are not sinuous. In contrast, the lower epidermis is composed of large, narrow, elongated cells with irregular sinuous walls (Figure 1j). Stomata (measuring on an average 41 x 30 microns) are restricted to the lower epidermis and oriented along the long axes of the leaf lamina (average frequency 51/sq.mm). The guard cells are elongated, oblong and chlorophyllous. The subsidiary cells of the stomata differ in shape from the other epidermal cells. They are broader with similar wavy outlines and usually surround nearly 34, sometime, more of the circumference of the guard cells at the posterior end, whilst the anterior end is dovetail into the corners of usually 1-2 cells. A stoma encircled by a second subsidiary cell is not uncommon; the inner cell is narrow, small, and with smooth outline, the outer one is similar to those described above (Figure 1j). The foregoing stomatal type is classified as polocytic (Van Cotthem, 1970). Fertile fronds are distinct from sterile ones, long stalked and on the same plan (trifoliate) as the sterile. They are contrasted almost to wingless axes (Figure 2). The lamina area is reduced to linear segments bearing sori on both the surfaces. Similar hairs as found on the sterile frond lamina are present on the stipe but are smaller in size and are very Sparse. Sporangia The sporangia (Figure 1g) are of the common leptosporangiate type with a large, pear-shaped capsule borne on short stalk usually 2-3 cells long, 2-celled thick. There is a shortthird row of stalk cells (Figure 1g “S’’) at the base of the capsule which is formed secondarily during sporangial development as a downward protrusion of the basal cell wall and the thin-walled stomial cell. However, the stalk is only one-celled thick at its extreme base. The sporangial capsule (measuring on an average 245 x 235 microns) is 279 MORPHOLOGY OF STENOSEMIA ‘aJOdS jO MAIA |e41a}e| ‘p ‘aseq 94} }e adijs 9Yy} JO UOIDASUBI} :E ‘SpUu_Os} JO LIGeY :Z :|SA1q (“MS) eyune eiwasouals ‘p-7 SAYNOIS IC 280 FERN GAZETTE: VOLUME 12 PART 5 (1983) thin-walled and composed of one layer of 8-10 large, flattened cells with nearly smooth contours. The annular ring is well formed with a region of usually 12-14 highly indurated thick-walled cells protruding prominentfrom the surface of the capsule, and iS not continuous with the stomium. As in all other tectarioid ferns, the stomium is well-developed and possesses a pair of unthickened lip cells with thin-walled accessory cells above and below it. Sporangial stalk hairs as found in other tectarioid ferns (Kaur, 1973; Chandra, 1976; Zamora and Chandra, 1977; Kaur, 1978) are absent. Paraphyses are absent as in Camptodium (Chandra, 1976). Abortive sporangia are occasionally found. Spores The spores of Stenosemia, like those of all ferns of aspidioid affinity are of the typical monolete bilateral type, planoconvex in lateral view (Figure 4), oblong in polar view, light brown, and measuring on an average 25 x 35 microns (polar diameter X longest equatorial diameter exclusive of perine) with a distinct perine. Perine protruding c. 7 microns from the exine surface nearly hyaline, sparsely spinulose, and characteristically wrinkled into thin, elongated, sinuous, lobate folds which appear papillate in optical section. The exine is 3 microns thick, densely and minutely spinulose, light brown and usually clearly discernible into sexine and nexine, often nearly of equal thickness. The laesura is 15 microns long, tenuimarginate. On acetolysis there is no alteration in the size and shape of the spores. The general morphology of the spore is that of the 7ectaria group. DISCUSSION The morphology of the plant described in this paper shows all the characters of the Tectarioideae. As brought out in this study, Stenosemia closely resembles free-veined species of 7ectaria (Chandra and Kaur, 1976) and some of the tectarioid derivatives like Camptodium (Chandra, 1976), Psomiocarpa (Zamora and Chandra, 1977), and Hemigramma (Chandra and Salgado, 1978). These genera share several characters in common including the following: rhizome with spirally arranged fronds and dictyostelic vascular cylinder, lack of sclerenchyma strands in the rhizome, greatly reduced leaf trace strands (2-3 in Psomiocarpa, 3-4 in Stenosemia, 4-5 in Camptodium, 4-6 in Hemigramma, 5-6 in free-veined 7ectaria, 8-12 in most Jectaria species), root trace at the base of the leaf gap, basally attached clathrate paleae, glandular ctenitoid hairs on the paleal margin, lack of surface hairs on the paleae, articulated foliar hairs, small stature (like Psomiocarpa and Camptodium). | Though similar to 7ectaria in some respects (especially in form of frond), Stenosemia differs constantly from most species of 7ectaria in having much smaller stature; fronds trifoliate in plan (both sterile and fertile); fertile fronds contracted to linear segments with acrostichoid sori; greatly reduced leaf trace strands; lack of sclerenchyma strands; root traces not associated with leaf trace strands; unicellular papillate hairs on paleal margin; lack of sporangial hairs and paraphyses; free venation with costal areolae only; foliar buds in the axil of basal pinnae. Copeland (1947) considers Stenosemia to have been derived from Ctenitis but the present studies have shown that Stenosemia differs markedly from Ctenitis in having few number (3-4) of leaf trace strands; lack of root associated leaf trace strands; lack of sclerenchyma strands in the rhizome and stipes; occasional vestigial leaf trace MORPHOLOGY OF STENOSEMIA 281 lack of sclerenchyma strands in the rhizome and stipes; occasional vestigial leaf trace strands; lack of surface hairs on paleae; veins free with costal areolae only; stipes paleate at base only, fronds trifoliate in plan; fertile lamina contracted to linear segments with acrostichoid sori; lack of sporangial hairs; presence of abortive sporangia; much smaller stature. It has also been observed that Stenosemia though similar to the acrostichoid Heterogonium (H. pinnatum) in general appearance (Copeland, 1929, 1947; Holttum, 1968) differs from H. pinnatum in having much smaller stature; fronds always trifoliate in plan; the lateral pinnae having the basal basiscopic segments much the longest; upper surface usually more or less hairy all over; fronds much more translucent, drying browning colour; the fertile frond contracted to linear segments; buds in the axil of basal pinnae. The sum of the differences between Heterogonium and Stenosemia and the close similarity of Stenosemia with free-veined species of Tectaria and tectarioid derivatives, seem to warrant the separation of the two genera as also suggested by Holttum (1968, 1975). In view of the foregoing, it appears that the similarity between Stenosemia and Heterogonium is probably due to parallel line of evolution; the former is possibly derived from free-veined species fo Jectaria and the latter from Ctenitis. There exists a tendency towards reduction and simplification of tne venation pattern in Tectarioideae as shown by 7. devexaand TJ. fuscipes (Kaur, 1974, 1978). This group of species is considered by Chandra and Kaur (1976) as evolutionary advanced; phylogenetically linked with the derivatives of 7ectaria. It should be noticed that there is also a tendency towards reduction and simplification in several characters of Stenosemia, viz., much smailer stature, few leaf trace strands, fertile lamina reduced to linear segments; lack of sclerenchyma strands in the rhizome; lack of sporangial hairs, simplification of venation; suppressed leaf trace strands. On the other hand, Stenosemia shows similarities with a group of free-veined species of 7ectaria and tectarioid derivatives like Camptocium, Psomiocarpa, and Hemigramma_ as mentioned earlier. From the above conclusions it seems more probable that Stenosemia is also evolved along similar lines of reduction and simplification of characters as those of Camptodium, Hemigramma and Psomiocarpa, from free-veined species of 7ectaria. These results thus, support the earlier view of Christensen (1938) who considers Stenosemia a tectarioid derivative. The sum of these characters, together with close relationship of Stenosemia with the derivatives of 7ectaria point to a comparatively advanced position of the genus, and suggest, on balance, that Stenosemia is probably allied to the free-veined species of Jectaria and possibly derived from it and not from Ctenitis as suggested by Copeland (1947). ACKNOWLEDGEMENTS | am thankful to Dr. T.N. Khoshoo, Director, National Botanical Research Institute, Lucknow, India, for providing facilities and giving encouragement during the course of this study. Thanks are also due to Mr. Lalita Tiwari for the preparation of microtome serial sections. 282 FERN GAZETTE: VOLUME 12 PART 5 (1983) REFERENCES CHANDRA, S., 1976. Morphology of the adult sporophyte of Camptodium Fee (Aspidiaceae). Brenesia 9: 15-23. CHANDRA, S. & KAUR, S., 1976. Contributions to the morphology of Jectaria: Vascular organisation of the rhizome. Phytomorphology 26: 143-150. CHANDRA, S. & SALGADO, A., 1979. Sporophyte morphology in two species of Hemigramma. Kalikasan: Philipp. J. Biol. 8: 69-78. CHING, R.C., 1940. On natural classification of the family ‘Polypodiaceae’. Sunyatsenia 5: 201- 269. CHRISTENSEN, C., 1938. Filicineae, in Verdoorn, F. Manual of Pteridology, Martinus Nijhoff. The Hague. COPELAND, E.B., 1905. The Polypodiaceae of the Philippine Islands. \. Bureau Public Print., Manila No. 28: 1-139. COPELAND, E.B., 1947. Genera Filicum. Chronica Botanica, Waltham Mass., U.S.A. COPELAND, E.B., 1955. New Philippine Ferns XI: Philipp. J. Sci. 84: 161-165. COPELAND, E.B., 1960. Fern Flora of the Philippines. Vol. \\. Natl. Inst. Sci. Tech., Manila. CRABBE, J.A. et a/., 1975. Anew generic sequence for the Pteridophyta. Fern Gaz. 77:141-162. HENNIPMAN, E., 1977. A monograph of the fern genus Bo/bitis (Lomariopsidaceae). Leiden Bot. Ser. 2: 1-331. HOLTTUM, R.E., 1949. The fern-genus Heterogonium Presl. Sarawak Mus. J. 5: 156-166. HOLTTUM, R.E., 1968. A revised flora of Malaya. Vol. \l Ferns. Govt. Print. Off., Singapore. HOLTTUM, R.E., 1975. The genus Heterogonium Presl. Kalikasan: Philipp. J. Biol. 4: 205-231. KAUR, S., 1973. Morphology of two Indian species of Ctenitis. Bangladesh J. Bot. 2: 37-42. KAUR, S., 1974. The family Lomariopsidaceae (Filicopsida) and its probable ancestors. Bot. Jour. Linn. Soc. 68: 153-162. KAUR, S., 1978. Contributions to the morphology of 7ectaria. Venation pattern of sterile and fertile leaves. Phytomorphology 28: 14-19. KAUR, S., in press. Contributions to the morphology of 7ectaria: Morphology of the sporophyte. Phytomorphology. REVIEW UNA GUIA TAXONOMICA PARA HELECHOS DE EL SALVADOR by Ralph Seiler, 58 pp., 238 x 180mm, San Salvador, Ministerio de Educacion, 1980. Price not known but obtainable from the Director of Publications, of the above Ministry. This is a checklist of the ferns of El Salvador (279 species) with keys to families and genera. It is written by a keen pteridologist, who is professionally a geologist, who found himself seconded to this Central American country. It is an admirable effort and should encourage more local work in a tropical country where, like all tropical countries, every minute counts towards assessing what plants are to be found there before forests are destroyed. A.C. JERMY FERN GAZ. 12(5) 1983 283 THE CHANGING ROLE OF CHEMISTRY IN FERN CLASSIFICATION GILLIAN A. COOPER-DRIVER Dept. Biol. Sci., Boston University, Boston, MA 02215, USA CHRISTOPHER HAUFLER Dept. Botany, University of Kansas, Lawrence, Kansas KS 66045. ABSTRACT This article reviews the use of plant chemicals, especially secondary compounds, in systematic and evolutionary studies in the Pteridophyta. INTRODUCTION The use of chemical characters in systematics has had a very long history (Gibbs, 1963). In ancient times, many medicinal plants were classified according to their “virtues” indicating a close chemical relationship. Camerarius in 1699 noted that “’-- et que celles de la méme class ont aussi quelques rapports dans leur vertus--” (DeCandolle, 1816). But it was not until 20 years ago with the almost simultaneous publication of Alston and Turner's text on ‘Biochemical Systematics” (1963), the symposium on “Chemical Plant Taxonomy” in the same year (Swain, 1963) and the first volume of ““Chemotaxonomie der Pflanzen” (Hegnauer, 1962), that the discipline became recognized in its own right. Since then, the field has developed rapidly due to the exploitation of the newer methods of separating and identifying compounds onan ever increasingly smaller scale. The rationale of biochemical systematics is the same as in other taxonomic procedures. Plants, or other organisms, are regarded as being more closely related if they contain the same or biosynthetically closely related compounds. The substances examined include most macromolecules, especially proteins and nucleic acids, and a host of low molecular weight compounds (M W 1000 daltons), the so-called secondary products.such as alkaloids and flavonoids (Smith, 1976; Ayala, 1977: Gutfreund, 1981). (Table 1). TABLE 1: Chemical characters used in the classification of plants SECONDARY METABOLITES Flavonoids, alkaloids, terpenoids ISOZYME VARIATION Gel electrophoresis of plant enzymes PROTEINS Amino-acid sequencing of plant proteins SEROLOGICAL REACTIVITY Immunological cross reactions DNA Interspecific hybridization studies Because of their diversity, chemical stability and ease of separation and identification, secondary compounds have been the most useful in systematic and evolutionary studies, especially since their mode of biosynthesis is known with reasonable certainty and their important ecological roles are becoming increasingly apparent. Their use in evolutionary studies has recently been discussed (Swain and Cooper-Driver, 1981) and it is obvious that many new biosynthetic pathways have evolved with time leading to a whole range of new compounds such as the betalains, hydrolysable tannins and aromatic alkaloids, all of which are confined tothe flowering plants. Usually, the more ancient plants show much less chemical diversity than those which have evolved more recently (Table 2), but, as discussed below, there are exceptions. 284 FERN GAZETTE: VOLUME 12 PART 5 (1983) TABLE 2: Classes of secondary metabolites present in vascular plants Pteridophytes Angiosperms and Gymnosperms Cell wall components Monoterpenes Lignin Acetylenes Cutin Aromatic alkaloids Suberin Glucosinolates Diterpenoids Betalains Triterpenoids Hydrolysable tannins Lysine alkaloids Flavonoids Condensed tannins Of all the classes of secondary compounds which have been used in systematic studies, the flavonoids and related phenolic compounds have proved to be the most useful. This is because, in general, most plant leaves contain 10-20 of these compounds which can be readily separated by simple two-dimensional PC or TLC and their structures determined on a yg scale by UV spectrophotometry. Furthermore, they often show simple Mendelian genetical traits, and they were the first group of compounds to be used in biochemical genetics. The flavonoids show an increase in complexity with evolutionary time (Figure 1), the simpler flavones (often as C-glycosides or other C-linked compounds) occurring in more primitive plants while the more advanced ferns and gymnosperms have flavonols and the ubiquitous procyanidin tannins of immense ecological importance. In the more advanced ferns and seed bearing plants, flavones again prevail (Figure 2). There is also a greater number of species containing xanthones, (formed via a pathway involving 2 rather than 3 acetate additions to the original C, precursor), chalkones and isoflavones. It was shown nearly 25 years ago that many enzymes occur in multiple forms in the tissues of most organisms (Ayala, 1977), and these could be readily differentiated using gel electrophoresis. The technique was rapidly adopted by population biologists to measure genetic variation and even though there is still a great deal of controversy about the significance of this polymorphism, it is an extremely useful and relatively simple technique for detecting similarities and differences between taxa and for proposing origins of alloploid taxa. Electrophoresis is perhaps most useful, however, in detecting and analyzing variability within and between populations. Through such analysis it is possible to determine the level of interaction between individuals in a population and, by correlating electrophoretic data with laboratory study of breeding systems, to propose mechanisms by which this interaction takes place. The individual forms of the enzymes (or storage proteins) are separable on the grounds of shape, molecular weight and net charge: this is determined by movement in an inert gel at a given pH, and ionic strength under the influence of a standard electric field. The technique cannot distinguish all different isozymes since many vary because of changes in amino acid sequence/charge which have no effect on mobility. Variations in both DNA and RNA have also been used to examine evolutionary and systematic relationships (Ayala, 1977; Gutfreund, 1981). In prokaryotes, recent attention has been paid to gross sequence differences, but the homology between different nucleic acids (especially DNA) has also been determined by annealing procedures (Ayala, 1977) in which relatedness of two taxa is measured by the degree to which their single stranded DNA, separated by heating in buffer to ca. 90°C, reassociates on cooling. CHEMISTRY AND FERN CLASSIFICATION 285 A BB —__ FC ———_ D flavone isoflavone flavonol proanthocyanidins FLAVANONE —————> FLAVANONOL ———-> FLAVAN 3,4—DIOL<——> FLAVAN 3-OL FIGURE 1. Biosynthetic pathways of the flavonoids. p-coumaric acid ————— XANTHONES 3C5 CHALKONE FLAVANONE flavanonol flavonol ISOFLAVONES FLAVONES proanthocyanidins FIGURE 2. Reduction in the reaction sequence in flavonoids. 286 FERN GAZETTE: VOLUME 12 PART 5 (1983) CHEMISTRY OF FERN AND FERN ALLIES Most of the methods outlined in the introduction have been applied to ferns and fern allies (see Swain and Cooper-Driver, 1973; Giannasi, 1974) and in the last three or four years many new and exciting questions on fern taxonomy have been posed (Tryon and Tryon, 1982). The new chemical information has given us novel insights into this interesting group of ancestral plants. Flavonoids The flavonoids and related phenolic compounds have received most attention in fern systematics (Cooper-Driver, 1980). A three step scheme of biochemical evolution (Table 3) is as true for the pteridophytes (Cooper-Driver, 1980) as it is for the angiosperms (Gornall and Bohm, 1978). The more primitive taxa (Psilophyta, Lycopodophyta), contain only flavones, while flavonols and related 3-hydroxy compounds are found in Equisetum, eusporangiate and leptosporangiate ferns. In the more advanced fern taxa, the production of these components is suppressed and flavanones, chalkones, flavones and xanthones are more common (Cooper-Driver, 1980; Wallace et a/ 1982). (Table 4). TABLE 3: Biochemical evolution of the flavonoids Primitive Development of biosynthetic processes leading to the accumulation of flavones (including C-glycoflavones) Advanced Substitutional and skeletal diversification of compounds with 3-hydroxyl groups and other structural features (complex glycosylation) Highly advanced Greater utilization of primitive pathways and suppression of existing pathways TABLE 4: The distributions of flavonoid groups in the Pteridophyta Primitive Advanced Highly Advanced, Flavones Flavonols Flavonones, Chalkones, Xanthones Psilotaceae Equisetaceae Hymenophyllaceae Lycopodiaceae Ophioglossaceae Pteridaceae Selaginellaceae Marattiaceae Dennstaedtiaceae lsoetaceae Osmundaceae Dryopteridaceae Schizaeaceae Aspleniaceae Loxomataceae Marsileaceae Stromatopteridaceae Gleicheniaceae Cyatheaceae The xanthones, because of the ease of identification and apparent taxonomic importance have received considerable attention. The main compounds mangiferin and isomangiferin have been found in four disparate genera (Athyrium-Asplenium, Elaphoglossum, Hymenophyllum-Trichomanes and Marsilea). Their distribution here is valuable at the generic level (as in the angiosperms, Hostettman & Wagner, 1977}, but has not thrown much light on overall fern evolution or systematics. Other classes of flavonoids have continued to be particularly useful often correlating with known morphological features at the generic level as shown by recent work on Bommeria (Haufler, 1979; Haufler and Giannasi, 1982). (Table 5). This work CHEMISTRY AND FERN CLASSIFICATION 287 TABLE 5: Interspecific differences between species groups in Bommeria B. hispida/B. subpaleacea B. ehrenbergiana/B. pedata Luteolin glycosides present Luteolin glycosides absent Large scales present Large scales absent Cristate spores Reticulate spores was extended to look at intergeneric affinities between Bommeria and Hemionitis (Giannasi, 1980). Species having spores with cristate surfaces were shown to contain common flavonols whereas those having rugose surfaced spores lacked these compounds. It is expected that more extensive chemical examination of related cheilanthoid fern genera may help to untangle other relationships. The distribution of flavonoids and related compounds has proved to be particularly valuable in examining the complex relationships of many hybrids which are formed via allopolyploidy in ferns. This was earlier shown by the classical work of Smith and his coworkers (Smith and Levin, 1963; Smith and Harborne, 1971) onthe Appalachian Asp/enium hybrids. Here it was shown that it was relatively easy to demonstrate systematic relationships between the parental diploids and derived tetraploids on the basis of their flavonoids. Nephelea portoricensis compounds A,C,D) Do Fertile hybrid Fertile hybrid compounds A,C,Dj,Do compounds A,B,C,D),Do Alsophila dryopteroides Fertile hybrid Alsophila bryophila compounds A,C compounds A,B,C compounds A,B FIGURE 3. Flavonoid analysis of Puerto Rican tree ferns. In many tropical cloud forests, tree ferns are prominent members of the ecosystem and there is great difficulty in deciding relationships between the putative species. For example, in the Greater Antilles A/sophila and Nephelea form a baffling array of interrelated species groups. Detailed analysis of populations in Puerto Rico, on morphological criteria, showed that there was probable hybridization between A. bryophila, A. dryopteroides and WN. portoricensis, but the presumed hybrids could not be differentiated readily, as species and hybrids were all fertile and at the same ploidy level (Conant and Cooper-Driver, 1980). However, the parentage and presumed genetic linkages could be detected by examination of flavonoid patterns on 2-D paper chromatography (Figure 3). Other suspected cases of hybridization between species of the two genera in other islands, has also been detected using flavonoid data (Conant, 1982 pers. comm.). 288 FERN GAZETTE: VOLUME 12 PART 5 (1983) on ssp. bu/b/fera (BB) ssp.profrusa(PP) ssp. fenu/folia (TT) (chemistry I) (chemistry IZ) (chemistry IIL) . ee ssp. fennesseensi/s (BBPP) ssp.mackayii(PPTT) ssp."fragilis’(TTTT A) (chemistry IV) (chemistry IL) (chemistry IIL) Chemical patterns: I= flavonoids A,B+5 xanthones I= flavonoids A,B+ 3 xanthones -Ill= flavonoids B,C+ trace xanthones IV=flavonoids A,B+6 xanthones FIGURE 4. Relationships in the North American Cystopteris fragilis complex (after Timothy Reeves, 1981), supported by chromosomal, morphological and chemical data. One of the most formidable systematic challenges in the ferns is the genus Cystopteris whose species are distributed worldwide and show remarkable variability in cytotypes (2n - 8n) and morphological intergradations. The diploid taxa are found only in the Americas, arguing for origination there. Recent preliminary data on the distribution of xanthones and related compounds (Figure 4) suggest that chemistry may be valuable in resolving species relationships in this complex genus. In the lycopods, both flavonoids and the complex lysine-derived alkaloids have proved to be useful. Most authorities maintain that Lycopodium isa single genus, with three subgenera, Lycopodium sensu stricto, Cernuistachys (Lepidotis) and Selago (Urostachya) (Tryon and Tryon, 1982). While there is general agreement that Lycopodium s.s. is the most advanced sub-genus, there is often disagreement as to the evolutionary status of the other two subgenera (Pichi-Sermolli, 1977; Hickey, pers. comm.). TABLE 6: Some chemical characteristics of the genus Lycopodium Sub-genera Selago Cernuistachys Lycopodium Alkaloids lucidine cernuine lycopodine inundatine Syringyl absent absent present Lignin Flavonoids O-methylated C-glycosy| 5-O glycosyl flavones flavones flavones Braeckman et a/ (1980) surveyed 33 temperate species of Lycopodium for alkaloids and found that the three subgenera showed distinct differences in the principal alkaloids accumulated. Cernuistachys accumulated mainly the alkaloid lucidine. Se/ago mainly cernuine and inundatine and Lycopodium s.s. mainly CHEMISTRY AND FERN CLASSIFICATION 289 lycopdine (Table 6). He concluded, on biosynthetic grounds, that the subgenus Lycopodium, which accumulated mainly the alkaloid lycopodine, was the most advanced. This is supported by studies in the lignins (Towers and Maas, 1965) which show that only Lycopodium contains the more advanced and angiospermlike syringyl groups. Flavonoid data also confirms the advanced status of this subgenus and also suggests that Cernuistachys may be the most primitive. Flavones comprise the only group of flavonoids in the genus Lycopodium but within the basic flavone skeleton there is an astonishing diversity of substitution patterns based on C and O- glycosylation and O-methylation. C-glycosylation is generally regarded as the most primitive substitution pattern (Swain and Cooper-Driver, 1981)and the more primitive C-glycosyl derivatives are found in the subgenus Cernuistachys. Complex 5-O- glycosides have been isolated from Lycopodium s.s. (Markham and Moore, 1980) again confirming their advanced status. Protein Chemistry A technique that is providing one of the most significant modifications of the role that chemistry can play in studying fern evolution is analysis of protein variability through electrophoresis. Some of the evidence generated through this technique is similar to other chemosystematic data since it provides cryptic markers that can be coordinated with existing data in distinguishing species and identifying plants of hybrid origin (Werth et a/ 1980; Moran, 1981; Haufler and Soltis, 1980; 1983). o-diphenol O-quinone mvc | \“ melanins phenol-thiol phenol-sulfite complex complex FIGURE 5. A summary of the chemical reactions by which phenolic compounds inhibit enzyme activity. Electrophoretic data are of greatest utility, however, in analyzing variability among individuals and populations (Darrow and Gastony, 1981; Haufler and Soltis, 1983). Assaying protein variability in the ferns is not as straightforward as with many other plants. When standard grinding procedures are followed, the high concentrations of tannins and other phenolics found in nearly all ferns denature the proteins (Figure 5). Itis only recently that these problems have been overcome by using a complex extractant buffer and a procedure that inhibits tannin-protein interactions (Soltis et a/ 1980; 1983). Application of electrophoresis in work with ferns has proved particularly useful since ferns have some characteristics that can be studied best through enzyme analysis. For example, ferns are usually polyploid and it is possible to detect allopolyploid or duplicated sets of chromosomes through electrophoresis (Gottlieb, 1982). Many fern genera also have complex reticulate patterns of species relationships. Often it is possible to use chemical markers to determine the putative alloploid taxa have enzyme banding patterns which combine those of their presumed 290 FERN GAZETTE: VOLUME 12 PART 5 (1983) parents (Werth et a/ 1980). Ferns are unique among vascular land plants in having separate, independent gametophyte and sporophyte generations. Through detection of cryptic protein polymorphisms, it may be possible to demonstrate that these two generations contain differing amounts of variability. If it can be shown that some variability is being filtered out by the gametophyte generation, it may be possible to assign an evolutionary role to this ephemeral life cycle stage. Finally, since it has been shown that there is a diversity of factors influencing gametophyte sex expression, and therefore breeding systems in the ferns (Klekowski, 1969; Lloyd, 1974), electrophoresis is proving valuable in assaying the consequences. of this breeding system variability at the population level (Haufler and Soltis, 1983; Haufler, in prep.). The value of enzyme analysis in fern systematics has been demonstrated by Werth et a/(1980, 1981) in work with the Appalachian Asp/enium complex mentioned earlier. An examination of eleven different enzyme loci showed that variability within species of the complex is low, except between well separated populations. However.,. the parental diploid species showed well defined enzyme band patterns which were found to be additive in the putative allopolyploid derivative species. Further, by detecting regional differences in enzyme patterns, it was possible to demonstrate multiple origins for the hybrid-derived, allopolyploid taxa. In the genus Bommerta, electrophoretic analysis has provided valuable information on species relationships and evolutionary mechanisms (Haufler and Soltis, 1980; 1983). Of particular significance is that calculation of Nei’s genetic identity (Green, 1979) within Bommeria (Figure 6) showed that the species were B. HISPIDA B. SUBPALEACEA B. EHRENBERGIANA B. HISPIDA _B, SUBPALEACEA —_B. EHRENBERGIANA _B. PEDATA B HISPIDA © © 0) (Sass 0.207 0.059 ~ 0.183 B. SUBPALEACEA 1:575:siieah veteerra 0. 382 0. 348 B. EHRENBERGIANA 2.834 a ala tN em 0.183 B. PEDATA 1.697 1.055 1. 6ogyiiené ariyimte-2= FIGURE 6. Pictorial representation of Nei’s genetic distance among Bommeria species (from Haufler and Soltis, 1983). CHEMISTRY AND FERN CLASSIFICATION 291 remarkably different from each other. The average genetic identity of Bommeria species, based on determination of allelic frequencies at 13 polymorphic loci, isO.195. This figure contrasts markedly with similar infrageneric comparisons among angiosperm species where the average genetic identity was shown to be 0.673 and the values ranged from 0.28 to 0.99 (Gottlieb, 1981). These data suggest that Bommeria species may not have a common ancestor and that their morphological similarity may have resulted through convergent evolution to a similar dry-adapted morphotype. Another finding of evolutionary significance is that comparison of gametophyte and sporophyte enzyme banding patterns (Gastony and Gottlieb, 1982) shows that all populations of B. hispida are heterozygous at the cytosolic locus of phosphoglucoisomerase (PGI-1). This indicates that each population has been initiated by at least two genotypically different spores whose resultant gametophytes have outcrossed to produce the heterozygous sporophytes. While the fact that B. hispida populations occur as isolated, disjunct units argues against the likelihoodthat such an obligate outcrossing scenario could be occurring, these enzyme data, based on field-collected samples, correlate with laboratory analysis of the breeding system (Haufler and Gastony, 1978) to indicate that this is the mechanism by which fertilization is taking place in nature. Recent investigations of the genus Cystopteris have demonstrated a similar (outcrossing) breeding system in C. protrusa (Haufler, imprep.). Analysis of the enzyme variability and population structure (Figure 7) of this species have indicated that it is 2.0m ENZYME SYSTEMS a EST CAT MDH LAP IDH | 3,4 12,34 Zs 3,4 2 Ki h23. 25 3,4 2,2 Prdie. 2,5,4 Pi ie ee 12,3,4 1,23 a A BAND PATTERNS — Numbers indicate relative rates of migration FIGURE 7. Map of population of Cystopteris protrusa based on similarities in band patterns of six enzyme systems. PGI = phosphoglucoisomerase, EST = esterase, CAT = catalase, MDH = malate dehydrogenase, LAP = leucine aminopeptidase, and IDH = isocitrate dehydrogenase. This technique provides a method for determining the pattern of variability in populations and for assessing the distribution and extent of asexually reproduced clones. The actual genetic composition of these electrophoretic phenotypes has not yet been determined. 292 FERN GAZETTE: VOLUME 12 PART 5 (1983) quite polymorphic (preliminary study of range-wide samples show a polymorphic index value of 2.50 alleles per locus based on eight resolved enzyme systems) and that a majority (93%) of the range-wide polymorphism is represented in a single population (Haufler, in prep.). Studies of other plant groups (reviewed in Hamrick et a/ 1979) have demonstrated that these results would be expected from a species that is outcrossing. Therefore, as was shown with Bommeria, these enzyme data on Cystopteris support the outcrossing breeding system proposed through study of laboratory-grown gametophytes. With the advent of methods facilitating electrophoretic analysis of enzyme variability in ferns, chemistry takes a step beyond systematics and phylogeny toward describing the partitioning of variability in populations and toward understanding the mechanisms by which this variability is maintained in nature. Through coordination of breeding system and enzyme data it is possible to propose basic differences between fern taxa that may relate directly to the relative complexity of their patterns of evolution. It is clear, therefore, that enzyme analysis will play an increasingly important role in understanding interactions within and among fern species and, ultimately, in defining the various modes of evolution in the ferns. DNA Analysis The comparison of DNA’s (and RNA’s) from different organisms by annealing techniques and by electrophoretic methods following their splitting by restriction enzymes or other methods has proved to be of great importance in determining systematic and phylogenetic relationships for many organisms. These techniques have been only recently applied to ferns, mainly because of the difficulties in obtaining pure nucleic acid fractions free from contamination by tannins (Stein and Thompson, 1978). With regard to nuclear DNA, there is also the problem of polyploidy, which often makes it difficult to detect homologies. Nevertheless, much progress has been made in this field and the phylogenetic relationships of three New England Osmunda species determined. Comparisons of their nuclear DNA showed that the three species most likely arose more or less simultaneously from a common ancestor (Stein et a/ 1979). More recently an examination of fern chloropast DNA has been undertaken (Stein, pers. comm.). Here, the DNA is less complex and can be studied not only by annealing, but also by electrophoretic examination after splitting by specific restriction enzymes. Comparisons of labelled material with similar fractions from angiosperm chloroplasts has shown remarkable similarities and gives promise for more extensive future phylogenetic studies. CONCLUSION It is obvious that chemical and biochemical studies on ferns have a very positive role to play in defining phylogenetic and taxonomic relationships. It isnow quite apparentthat such studies can throw new light on variation within and between populations of a single species and on genetic interrelationships which are not discernable by morphological or other studies. Obviously the variations found demand that more careful sampling is carried out and one should not rely on results obtained from a single herbarium specimen. Fortunately, new developments of HPLC, gel electro- phoresis, GC-MS and so on will enable such data to be obtained more easily. Improved methods of computerized numerical analyses have also been developed which will allow such data to be more competently utilized (Bisby et a/ 1980). Newer approaches to chemosystematics are constantly being explored and will widen the use made of chemical compounds for systematic research (Harborne, 1980). Fortunately, investigations on ferns are in the forefront of these exciting developments and it is to be expected that it will not be long before we are able to decipher new relationships in some of the more problematical taxa in this long evolved group of plants. CHEMISTRY AND FERN CLASSIFICATION 293 REFERENCES ALSTON, R.E. & TURNER, B.L., 1963. Biochemical Systematics. Prentice Hall, New Jersey. AYALA, F.J. Ed., 1977. Molecular Evolution. Sinauer, Sunderland, Mass. BISBY, F.A., VAUGHAN, J.G. & WRIGHT, C.A., Eds., 1980. Chemosystematics: Principles and Practice. System. Assoc. 76. Academic Press, London. BRAEKMAN, J.C., NYEMBO, L. & SYMOENS, J.J., 1980. Chimiotaxonimie des Lycopodiales: distribution des alcaloides. Phytochem. 19: 803-807. CONANT, D.S. & COOPER-DRIVER, G., 1980. Autogamous allohomoploidy in A/sophila and Nephelea (Cyatheaceae): a new hypothesis for speciation in homosporous ferns. Amer. J. Bot. 67: 1269-1288. COOPER-DRIVER, G., 1980. The role of flavonoids and related compounds in fern systematics. Bull. Torrey Bot. Club 197: 116-127. DARROW, D.C. & GASTONY, G.J., 1981. Electrophoretic genetic variation within populations of Athyrium filix-femina. Bot. Soc. Amer. Misc. Publ. 160: 57. DeCANDOLLE, A.P., 1804, 1816. Essa/ sur /es proprietes medicales des plantes, comparees avec leurs formes exterieures et leur classification naturelle. Eds. 1 & 2. Paris. GASTONY, G.J. & GOTTLIEB, L.D., 1982. Evidence for genetic heterozygosity in a homosporous fern. Amer. J. Bot. 69: 634-637. GIBBS, R.D., 1963. History of plant taxonomy. In (SWAIN, T., Ed.) Chemical Plant Taxonomy. 41- 88. Academic Press, London. GIANNASI, D.E., 1974. Phytochemical aspects of fern systematics. Ann. Missouri Bot. Gard. 67: a7 GIANNASI, D.E., 1980. Flavonoid evidence for generic lines in selected gymnogrammoid ferns. Bull. Torrey Bot. Club 107: 128-133. GORNALL, R.J. & BOHM, B.A., 1978. Angiosperm flavonoid evolution: a reappraisal. Syst. Bot. 3: 353-368. GOTTLIEB, L.D., 1981. Electrophoretic evidence and plant populations. Prog. Phytochem. 7: 1-46. GOTTLIEB, L.D., 1982. Conservation and duplication of isozymes in plants. Science 276: 373-380. GREEN, D.M., 1979. A BASIC computer program for calculating indices of genetic distance and similarity. Jour. Hered. 70: 429-430. GUTFREUND, H., 1981. Biochemical Evolution. Cambridge Univ. Press, London. HAMRICK, J.L., LINHART, Y.B. & MITTON, J.B., 1979. Relationships between life history characteristics and electrophoretically detectable genetic variation in plants. Ann. Rev. Ecol. Syst. 10: 173-200. HARBORNE, J.B., 1980. New experimental approaches to plant chemosystematics. In (BISBY, F.A., VAUGHAN, J.S. & WRIGHT, C.A. Eds.) System. Assoc. 76: 39-70. Chemosystematics Principles and Practice. Academic Press, London. HAUFLER, C.H., 1979. A biosystematic revision of Bommeria. J. Arn. Arb. 60: 445-476. HAUFLER, C.H. & GASTONY, G.J., 1978. Antheridiogen and the breeding system in the fern genus Bommeria. Can. J. Bot. 56: 1594-1601. HAUFLER, C.H. & GIANNASI, D.E., 1982. Achemosystematic survey of the fern genus Bommeria. Biochem. Syst. Ecol. 10: 107-110. HAUFLER, C.H. & SOLTIS, D.E., 1980. Detection and analysis of enzyme variation in the fern genus Bommeria. Bot. Soc. Amer. Misc. Pub. 158: 49. HAUFLER, C.H. & SOLTIS, D.E., 1983. Analysis of enzyme variability inthe fern genus Bommeria. Syst. Bot. (in prep.). HEGNAUER, R., 1962. Chemotaxonomie der Pflanzen |. Birkhauser, Verlag. HOSTETTMAN, K. & WAGNER, H., 1977. Xanthone glycosides. Phytochem. 16: 821-829. KLEKOWSKI, E.J., 1969. Reproductive biology of the Pteridophyta. II. Theoretical considerations. Bot. J. Linn. Soc. 62: 347-359. LLOYD, R.M., 1974. Reproductive biology and evolution in the Pteridophyta. Ann. Missouri Bot. Gard. 67: 318-331. MARKHAM, K.R. & MOORE, N.A., 1980. Comparative flavonoid glycoside biochemistry as a chemotaxonomic tool in the subdivision of the classical ‘‘genus’’ Lycopodium. Biochem. Syst. Ecol. 8: 17-20. MORAN, R.C., 1981: X Asplenosorus shawneensis, a new natural fern hybrid between Asplenium trichomanes and Camptosorus rhizophyllus. Amer. Fern J. 71: 85-89. PICHI-SERMOLI, R.E., 1977. Tentamen Pteridophytorum genera in taxonomicum ordinem redigendi. Webbia 37: 313-512. SOLTIS, D.E., HAUFLER, C.H. & GASTONY, G.J., 1980. Detecting enzyme variation in Bommeria: An analysis of methodology. Syst. Bot. 5: 30-38. SOLTIS, D.E., HAUFLER, C.H., DARROW, D.C. & GASTONY, GJ., 1983. Starch gel electrophoresis of ferns: a compilation of grinding buffers, gel and electrode buffers, and staining schedules. Amer. Fern J. 73: 9-27. SMITH, D.M. & LEVIN, D.A., 1963. A chromatographic study of reticulate evolution in the Appalachian Asp/lenium complex. Am. J. Bot. 50: 952-958. SMITH, D.M. & HARBORNE, J.B., 1971. Xanthones in the Appalachian Asp/enium complex. Phytochem. 10: 2117-2119. SMITH, P.A., 1976. The Chemotaxonomy of Plants. Elsevier, New York. STEIN, D.B. & THOMPSON, W.F., 1978. Isolation of DNAfrom tannin containing plants. Plant Sci. Letters 11: 323-328. ji 294 FERN GAZETTE: VOLUME 12 PART 5 (1983) STEIN, D.B., THOMPSON, W.F. & BELFORD, H.S., 1979. Studies on DNA sequences in the Osmundaceae. J. Mol. Evol. 13: 215-232. SWAIN, T., Ed. Chemical Plant Taxonomy. Academic Press, London. SWAIN, T., & COOPER-DRIVER, G., 1973. Biochemical systematics in the Filicopsida Suppl. 1. Bot. J. Linn. Soc. 67: 111-134. SWAIN, T. & COOPER-DRIVER, G., 1981. Biochemical evolution in early land plants. In NIKLAS, K. (Ed.) Paleobotany, Paleoecology and Evolution. Vol. 1, Praeger, New York. 103- 134. TOWERS, G.H.N. & MAAS, W.S., 1965. Phenolic acids and lignins in the Lycopodiales. Phytochem. 4: 57-66. TRYON, R. & TRYON, A.F., 1982. Ferns and Allied Plants with Special Reference to Tropical America. Springer-Verlag, New York. WALLACE, J.W., MARKHAM, K.R., GIANNASI, D.E., MICKEL, J.T., YOPP, D.L., GOMEZ, L.D., PATTILO, J.D. & SOEDER, R., 1982. A survey for 1,3,6,7-tetrahydroxy-C- glycosyxanthones emphasizing the ‘primitive’ leptosporangiate ferns and their allies. Amer. J. Bot. 69: 356-362. WERTH, C.R., GUTTMAN, S.I. & ESHBAUGH, W.H., 1980. Allozyme variation in the Asplenium complex of eastern North America: A progress report. /ntern. Cong. Syst. Evol. Biol. Abstr. 390. WERTH, C.R., GUTTMAN, S.l. & ESHBAUGH, W.H., 1981. Multiple origins of allopolyploid species of Asplenium. Bot. Soc. Amer. Misc. Publ. 160: 60. REVIEW NEW FLORA OF JAPAN: PTERIDOPHYTA by T. Nakaike, 808 pp. 1982. Published by Shibundo Co. Ltd., Tokyo. 260 x 195mm. Price not given. This is another beautifully printed and bound Japanese fern book, again in Japanese except for the Latin names. It contains however 849 photographs of herbarium specimens of almost all taxa described (including hybrids); in many cases the specimens shown are types. The photographs could have been of higher quality and in some cases larger to show more detail but it does give a very good idea of the general appearance, size and habit. Usually the outline of the frond is clear except in a few cases of larger ferns where the herbarium specimen photographed appears as a mass of overlapping leafy herbage. It is interesting to see the number of hybrids especially in Dryopteris and Athyrium. The filmy ferns are nicely treated according to Copeland's concepts of genera, with close-ups of fertile parts. Nakaike’s concept of Blechnum excludes that group which contains our species, 8B. spicant, for which he uses the generic name Struthiopteris. For those that collect world fern floras this is a must; for the grower of hardy ferns it is also immensely useful. All we need with it is a list of those that may be hardy — and hopefully this is not far off. A.C. JERMY FERN GAZ. 12(5) 1983 295 THE FLAVONOIDS OF THE OSMUNDACEAE GAIL L. SOBEL The New York Botanical Garden, Bronx, New York 10458, USA and MICHAEL D. WHALEN L.H. Bailey Hortorium, Cornell University, Ithaca, New York 14853, USA ABSTRACT Flavonoids from fronds of all three genera of the Osmundaceae were studied. Twelve flavonoid compounds were found. Osmunda cinnamomea contains glycosides of the flavonols kampferol and quercetin. Leptopteris superba contains flavone glycosides which are absent from O. cinnamomea and both types of compounds are found in Jodea barbara. These results are in accord with current concepts of delimitation of the genera. 7odea forms a connecting link between the other two genera. INTRODUCTION All members of the fern family Osmundaceae share thick, erect stems with unique stelar structure, uniseriate hairs, fronds free of stomata on adaxial surface and large, vertically dehiscent sporangia with rudimentary, lateral annulus (Hewitson, 1962). Three extant genera are recognized in the family. Osmunda, with ca. 10 spp., has a cosmopolitan, discontinuous distribution. Leptopteris, a genus of 6 spp., occurs in New Guinea, New Zealand, Australia and Polynesia. The monotypic 7odea is a southern hemisphere disjunct found in South Africa, Australia and New Zealand. An extensive fossil record existssfor Osmundaceae (Miller, 1971) and suggests that the family was once more widespread and diverse than at present. Form genera, Thamnopteris (leaves) and Osmundacaulis (stems), are assignable to the family and date back to the Permian. Osmundites, an Osmunda-like plant, is abundant in Mesozoic rocks. The genus Osmunda itself has a fossil record extending back 70 million years. Fossils are insufficient, however, to elucidate evolutionary relationships of extant genera. Todea and Leptopteris seem closely related, and Hewitson (1962), studying morphology and anatomy, concluded that they should be considered sections of a single genus. The two are distinguished primarily by filmy fronds only two cell-layers thick in Leptopteris vs. thick coriaceous fronds in Jodea. Miller (1971) has maintained these genera as distinct, in spite of Hewitson’s interpretation. The relationship of Todea and Leptopteris to Osmunda is less clear. Wagner, et a/ (1978) chromatographed a flavonoid extract of Osmunda cinnamomea in conjunction with a chemical study of hybridization between O. claytoniana and O. regalis. The chromatographic profile found by them differs substantially from the one presented here. They did not attempt to identify the compounds found, so the extent of differences in flavonoid structures between their collection and ours cannot at this time be assessed. This present paper reports results of a preliminary comparative survey of flavonoids from fronds of all three genera of the Osmundaceae and discusses their bearing on the taxonomy of the family. Particular questions addressed are generic delimitation of 7odea and Leptopteris and the appropriateness of their inclusion in the same family with Osmunda. Species investigated were Osmunda cinnamomea L.., Todea barbara Moore and Leptopteris superba (Col.) Presl. 296 FERN GAZETTE: VOLUME 12 PART 5 (1983) TABLE 1. UV spectral data of Osmundaceae flavonoids Absorption maxima (nm) Compound MeOH NaOMe A1C1, A1C1./HC1 NaOAc NaOAc/H,BO, 1. Kaempferol 3-0 348 400 399 396 386 350 glucoside 302sh 324 350 348 307 303sh 266 DIS 303 302 215 266 275 275 2. Kaempferol 3-0- 350 400 397 395 387 353 rhamnoglucoside 301sh 328 351 349 304 296sh 267 275 303 302 273 267 273 273 3. Quercetin 3-0- 360 410 434 404 390 378 glucoside 301sh 328 334sh 369sh 325 259 267sh 271 304sh 301 PM Pe 256 273 270 4. Quercetin 3-0- 355 402 429 397 394 370 rhamnoglucoside 304sh 326 339sh 358 318 299 266sh Payee 302sh 301sh 272 261 255 274 269 5. Unidentified 340 400 340 340 355 350 flavone 280sh 320 300 300 268 265 glycoside 265 275 275 275 6. Unidentified 328 380 340 340 330 350 flavone 385sh 320 300 330 275 285 glycoside 270 275 DTS 275 265 TABLE 2. Chromatographic properties of Osmundaceae flavonoids Rf Value Colour Toren TBA —- HOA UV UV + NH, 1 0.64 0.54 Purple Green 2 0.55 0.65 Purple Green 3 0.46 0.41 Purple Yellow Green 4 0.42 0.58 Purple Yellow Green 5 0.49 0.48 Dull Purple Light Green 6 0.59 0.53 Purple Green 7/ 0.20 0.80 Dull Purple Light Yellow 8 0.63 0.56 Dull Purple Dull Purple 9 0.36 0:27 Dull Purple Light Yellow 10 0.20 0.66 Dull Purple Dull Purple 11 0.87 0.66 Dull Purple Dull Purple —_ NO © rep) To) = | To) Dull Purple Green ee a aeerernnnsan TE EIT EEE EERE FLAVONOIDS OF OSMUNDACEAE 297 TABLE 3. Taxonomic distribution of Osmundacaea flavonoids Flavonols Flavones Unidentified 1 2 3 4 5 6 7 8 9 10 11 12 Osmunda cinnamomea +++ +++ +++ +++ Todea barbara +++ ++ ++ +++ + + ee Leptopteris superba ++ + + fe +++ = highest concentration ++ = medium concentration + = low concentration HOAc:H,0 TBA: HOAc:H,0O FIGURE 1. Distribution of Osmundaceae flavonoids on chromatogram MATERIALS AND METHODS Extraction and purification 5g of dried leaf material was extracted overnight in 50ml of 85% aq MeOH. The extract was filtered and evaporated to a small volume so most MeOH was removed causing chlorophyll to precipitate from solution. The concentrated flavonoid-containing extract was applied to the lower right corner of a sheet of Whatman 3MM chromatography paper, which was developed descending with 3:1:1 TBA:HOAc:H,0O as the solvent in the first (long) dimension and 15% HOAc in the second (short) dimension. The dried chromatogram was examined for fluorescent spots in UV light before and after fuming with NH;. To isolate flavonoids for structural identification, 60-150 replicate chromatograms were run for each collection. Resulting spots were cut out and extracted briefly from the paper with spectral MeOH. For some collections, a second PC run was required for final purification of compounds. 298 FERN GAZETTE: VOLUME 12 PART 5 (1983) Spectral analyses Structural identifications of purified flavonoids relied heavily on UV spectro- photometry of their methanol solutions, employing the diagnostic reagents of Mabry, et a/(1970). The only departure from their procedures was the use of fused rather than unfused NaOAc for reading the NaOAc and NaOAc/HBO, spectra. Sugar analysis of O-glycosides Acid hydrolyses were carried out in 2N HC1 for 2 hr. Flavonoid aglycones were removed from the resulting sugar-flavonoid mixtures by extraction with EtOAc. The Sugar containing aq fractions were chromatographed by ascent with standard sugars on cellose thin-layers with 12:5:4 EtOAc:Py:H,0O as solvent. Sugar spots were detected with analine pthalate spray. Plant materials Voucher specimens for plant materials used in this study are on deposit at the Bailey Hortorium (BH), Cornell University: Osmunda cinnamomea L., G.L. Sobel 1715; Todea barbara Moore, J.J. Strudwick 2170 (BH 370609); Leptopteris superba (Col.) Presl. M. Howard, (BH 370610). RESULTS Twelve flavonoid compounds were found in fronds of members of the Osmundaceae studied. Their spectral and chromatographic properties are given in Tables 1 and2 and in Figure 1. The taxonomic distributions of these substances are summarized in Table 3. Osmunda cinnamomea and Leptopteris superba have different foliar flavonoid profiles. Glycosides of the flavonols kampferol and quercetin prevail in the former but are absent from the latter. L. superba contains flavone glycosides which are absent from O. cinnamomea. Both types of compounds are found in 7Jodea barbara. These results are in accord with current concepts of delimitation of the genera in the Osmundaceae. Although chemical profiles of Osmunda and Leptopteris are very different, 7odea forms a connecting link between them. The flavonoid structural types found in Osmundaceae are those typical of the leptosporangiate ferns in general (Voirin, 1970) and support the inclusion of the family in that group. The absence of flavonols from Leptopter/s supports the separation of that genus from Jodea. Further species of Leptopteris need to be examined to be certain that this substantial chemical distinction is consistent. ACKNOWLEDGEMENTS The senior author was supported for the majority of this study by the School of Agriculture and Life Sciences, Cornell University Honors Degree Program. | wish to thank P. Mick Richardson for his comments on the manuscript. REFERENCES HEWITSON, W., 1962. Comparative morphology of the Osmundaceae. Ann. Missouri Bot. Gard. 49:57-93. MABRY, T.J., MARKHAM, K.R. & THOMAS, M.B., 1970. The Systematic identification of flavonoids. Springer-Verlag, New York. MILLER, C.N. Jr., 1971. Evolution of the fern family Osmundaceae based on anatomical studies. Contr. Mus. Paleont. Univ. Mich. 23: 105-169. VOIRIN, B., 1970. Recherches chimiques, taxinomiques et physiologiques sur les flavonoids des Pteridophytes. These, Docteur-Science. L’Universite de Lyon. WAGNER, W.H. Jr., WAGNER, F.S., MILLER, C.N. Jr. & WAGNER, D.H. 1978. New observations on the royal fern hybrid Osmunda x ruggii. Rhodora 80: 92-106. SHORT NOTES 299 SHORT NOTES CULCITA MACROCARPA — A NEW LOCALITY IN SPAIN Culcita macrocarpa Presl. belongs to a small genus, recently placed variously within the family Cyatheaceae in the wide sense, as a tribe, Thyrsopterideae (Holttum, 1963) or asasubfamily, Thyrsopteridoideae (Lovis, 1977), or within the order Dicksoniales as a family in its own right, Culcitaceae (Ching, 1940; Pichi Sermolli, 1977). It is therefore related to the tree-ferns (and to Jhyrsopteris in particular) and is the only naturally occurring member of that group in the European flora. According to Copeland (1947) it is somewhat primitive within the Cyatheaceae. The genus Culcita is divided into two subgenera, of which the subgenus Cu/cita contains two species, C. macrocarpa, from Macaronesia, Spain and Portugal, and the closely related C. coniifolia (Hook.) Maxon from Mexico, the West Indies and S. America. Culcita macrocarpa is confined to Macaronesia (Canary Isles: N.E. Tenerife; Madeira; Azores: Sao Miguel, Terceira, Sao Jorge, Pico, Faial, Flores and Corvo) where it has long been known, and to a few localities in Atlantic Spain and Portugal, and is thus an interesting example of a Macaronesian element within the European mainland flora where it was virtually overlooked, until Allorge (1934) published on it. It was first collected from south-west Spain by Col. White in 1869 (Diels, 1899) (specimens in K! and B) not far from Gibraltar and presumably in Cadiz province near Algeciras (see Allorge, 1934). Subsequently it was collected but not published in 1929 by Ceballos (see Molesworth-Allen, 1971), and then collected in 1933 by Alorge (1934), and later by Nieschalk & Nieschalk (1965), Molesworth-Allen (1971 and 1977). In the Atlantic southern tip of Spain it is known, due mainly to Molesworth-Allen’s searches, to grow in Cadiz province in the Sierra de la Luna, Miel valley, Sierra del Algarrobo, Sierra La Paloma and Sierra de Ojen (all shortly south-west of Algeciras), in the Sierra del Nino, and almost certainly in the Sierra Blanquilla, further east. It is not common, the largest populations being up to c 100 plants in 1969 and 1977, and it is therefore important that plants should not be collected, especially as this species has already been decimated in the past for medicinal extracts, and toa certain extent more recently by unscrupulous botanists selling sets of herbarium specimens, a practice which, if happening to any large extent with threatened species, should be most strictly and severely discouraged by the Universities concerned. It should be borne in mind that particularly with the disastrous droughts affecting south Spain recently, aggravated, or in the long term perhaps even caused, to a considerable degree by the widespread felling and almost uncontrolled burning of what remains of the tree cover, the fate of these few populations hangs in the balance. Culcita macrocarpa is also known from the north of Portugal, where it was discovered by Rezende-Pinto (1940 and 1943) at Valongo, just to the N E of Porto, Douro Litoral. However this was a naturalised population of cultivated origin, as pointed out by Franco (1971 and 1974). (See also Greuter, Burdet & Long, 1981). More recently, Cu/cita was discovered in a second natural mainland locality by Lainz (1968 and 1973), at Caaveiro, Capela, La Coruna, in the north-west corner of Spain. This part of Spain, though much cooler than Cadiz province, receives considerably more rainfall, and the whole north coast of Spain is in general the main centre for the occurrence of Atlantic species in the Iberian peninsula, depending on their tolerance of the cold temperatures prevalent in winter. The natural occurrence of Cu/cita here is therefore a further indication of the richness of Macaronesian and Atlantic species in the area. On a recent visit to the central part of the coast of North Spain, occupying only four days of June 1976, in order to search for Dryopteris guanchica Gibby & Jermy, one of us (CRFJ) found C. macrocarpa by chance in Oviedo province and made a herbarium 300 SHORT NOTES specimen (CRFJ 4884, 4th June 1976; BM). But it was not until a longer visit in 1981, that in discussions with the second author the significance of this third, natural population, a considerable distance from the others, was recognised and revisited. The plants there were very small, immature, though fertile, and the same size in both 1976 and 1981, the fronds reaching a maximum of c 10cm in length. The locality is below and north east of the Col de la Cruz de Llames, between Arriondas and Colunga, E of Gijon, Oviedo province. Approximately 20 plants were found altogether, growing at the edge of apath, now slightly widened along part of its length since 1976, but with no detrimental effect on the population of Cul/cita. The path is in an open area of Erica heath on a steep north-facing slope at 500-550m alt.; the substrate is white sandstone, and the Cu/cita plants were growing among mosses and grasses at the edge of where the dark peaty soil was exposed on the upper bank of the path, which is often a suitable area for ferns. Along with Culcita there were plants of Dryopteris aemula (Ait.) O. Ktze., D. affinis (Lowe) Fras.-Jenk. subsp. affinis, D. dilatata(Hoffm.) A. Gray, Oreopteris limbosperma (All.) Holub, Blechnum spicant (L.) Roth, Pteridium aquilinum (L.) Kuhn and Athyrium filix-femina(L.) Roth. Not far above and to the west at c. 600m alt., among rocks at the Mirador de El Fito, where the road passes through the Col de la Cruz de Llames, are Dryopteris guanchica discovered there by CRFJ in 1976 and the new species D. corley/ Fras.-Jenk., discovered there by ML in 1981 (see Fraser-Jenkins, 1982). A brief search on the hillside and in the dense forest c 100- 200m lower down failed to reveal any further plants and no mature plants were found, very probably because of the grazing and occasional burning of the locality, of which there were noticeable signs. It is emphasised and implored that in view of the small number of plants present, no further collections are made from this population. It is | also unlikely that plants would survive if transplanted into cultivation. If any further record is needed it should be a photographic one. The present discovery opens up the possibility that Cu/cita could occur in scattered localities on sandstone further west between Oviedo and La Corufa; further search is indicated. REFERENCES ALLORGE, P., 1934. La Culcita macrocarpa Presl. [=Ba/antium Culcita (L'Herit.) Kaulf.] dans les montagnes d Algesiras, Bull. Soc. Bot. France 81: 592-593. CHING, R.C., 1940. On natural classification of the family ‘‘Polypodiaceae’’, Sunyatsenia 5: 201- 268. COPELAND, E.B., 1947. Genera Filicum, the genera of ferns. Waltham, Mass. DIELS, L., 1899. Pteridophyta, in Engler, A. (Ed.), die Naturrlichen Pflazenfamilien. Leipzig. FRANCO, J. do A., 1971. Nova Flora de Portugal (Continente e Acores) 1: Lycopodiaceae - Umbelliferae. Lisbon. FRANCO, J. doA., 1974. Phytogeographical survey of Portugal. Bol. Soc. Broteriana, ser. 2a,47 (Suppl.): 89-90. FRASER-JENKINS, C.R., 1982. Dryopteris in Spain, Portugal and Macaronesia, Bol Soc. Broteriana. ser. 2a, 55: 175-336. GREUTER, W., BURDET, H.M. & LONG, G. (Eds.) 1981. Med-Checklist 1, Pteridophyta: 12-13. Geneve & Berlin. HOLTTUM, R.E.H., 1963. Cyatheaceae, in Flora Malesiana, ser. 2, Pteridophyta 1 (2): 65-176. LAINZ, M., 1968. Aportaciones al conocimiento de la flora gallega VI, An. del /nst. Forestal/invest Exp.. 36-37 LAINZ, M., 1973. Aportaciones al conocimiento de la flora Cantabra-astur X. Bo/ /nst. Estud. Asturianos 16. 163 LOVIS, J.D., 1977. Evolutionary patterns and processes in ferns. Advances in Bot. Research 4: 229-415. MOLESWORTH-ALLEN, E.A., 1971. Observations on Spanish ferns, Brit. Fern Gaz. 10: 200-202. MOLESWORTH-ALLEN, E.A., 1977. Observations on some rare Spanish ferns in Cadiz Province, Spain, Fern Gaz. 17: 271-275. NIESCHALK, A. & NIESCHALK, C., 1965. Ein Stammfarn (Culcita macrocarpa) auf europaischem Boden, Natur u. Museum 95: 495-498. SHORT NOTES 301 PICH! SERMOLLI, R.E.G., 1977. Tentamen pteridophytorum genera in taxonomicum ordinem redigendi, Webbia 31: 313-512. REZENDE-PINTO, M.C., 1940. Contribuicao para a flora criptogamica do Norte de Portugal, 4. Broteria, ser. cienc. nat., 9: 129. REZENDE-PINTO, M.C., 1943. Culcita macrocarpa Pr., contribuitao para o seu estudo monografico. Bol. Soc. Broteriana, ser. 2a, 17: 93-146. C.R. FRASER-JENKINS, c/o British Museum (Natural History), London. and M. LAINZ, S.J. Apartado 425, Colegio de la Immaculada, Gijon, Spain. AZOLLA FILICULOIDES IN EDINBURGH The Bawsinch Nature Reserve which borders Duddingston Loch and Bird Sanctuary was acquired by the Scottish Wildlife Trust in 1971, chiefly to act as a buffer zone for the Sanctuary. Since then extensive tree planting has taken place and in 1975 four ponds were excavated close to the Duddingston drainage ditch. In the ensuing 3-4 years the ponds were extensively planted as well as being colonised naturally, and it seems probable that Azo//a spores were introduced along with plants brought in from England. The presence of Azolla filiculoides was first noted in May 1980 in the smallest of the ponds, and by November 1980 it had so proliferated tha by then half the pond surface was covered. This was roughly the state in the spring of 1981 when the author started a plant survey of Bawsinch, the Azo//a having survived the relatively mild winter of 1980/81. Other surface plants in the pond included considerable quantities of Lemna trisulca (natural), some Lemna minor (natural) and Ranunculus aquatilis (planted). The Azolla fruited well in May-July, and by August a number of patches of new growth were apparent. Further excavation was undertaken in August to double the size of the pond but drought prevented the final stages until September. Meanwhile the Azo//la had expanded to cover the entire surface, and when the pond was finally allowed to double in size the Azo//a followed suit. It has not so far spread to any of the other ponds, but fronds appeared in a small hole (an abortive digging) close to Mark Pond following the autumn gales — probably wind-blown. Subsequent observations showed the Azo//a to eventually have succumbed in the severe winter of 1981/82, and it has not been seen here again since. J. MUSCOTT, 69 Warrender Park Road, Edinburgh. THE REDISCOVERY OF ASPLENIUM X CONFLUENS During a week’s holiday in Eire, in August 1982, | had the good fortune to find Asplenium x confluens (T. Moore ex Lowe) Lawalrée growing in a roadside wall in County Kerry (v.c. H1). Both parent species, A. scolopendrium L. andA. trichomanes L. subsp. guadrivalens D.E. Meyer emend. Lovis, were present, growing in the wall and otherwise in the vicinity, though neither in great numbers. The hybrid was robust and, | would guess, long established, the stock having branched to form side crowns. Collected fronds were subsequently sent to A.C. Jermy (British Museum (Natural History) ) who agreed with the determination which was further confirmed by 302 SHORT NOTES Professor T. Reichstein of Basel. A frond has been sent also to the Irish National Herbarium at Glasnevin. That, as a decidedly fallible amateur, | recognised the hybrid for what it was when | found it is indicative less of acuity on my part than of its distinctiveness: it exactly answered the description in Jermy et al. (1978). It can be presumed to be exceedingly rare since plants are unlikely to be overlooked, if seen, by those interested in ferns, and since both parent species are abundant in many parts of Britain, often together, commonly in easily accessible places, and are, besides, very popular with amateurs who scrutinise them for variants. FIGURE 1. Frond of Asplenium x confluens from Kerry. Apart from one record from Jugoslavia mentioned in Lovis (1975), and disregarding the accidental resynthesising of the hybrid in cultivation circa 1916 in the U.S.A. (Lovis and Vida, 1969), there appear to be only three accepted records, all from the British Isles. These were at Levens Park (now in Cumbria; v.c. 69) by Stabler in 1865; at Whitby (N. Yorkshire; v.c. 62) by W. Willson; and at Killarney (County Kerry; v.c. H2) by P.N. Fraser circa 1875 (Jermy et a/, Druery 1901). It is curious that these — three finds were made seemingly within a period of ten years, yet more than one hundred years elapsed before it was found again. No less curious is that | found it approximately fifteen miles from Killarney, where the last accepted previous find was made. Moreover | found A. sco/opendrium only fairly infrequently, and nowhere in abundance, in South Kerry, while A. trichomanes, though somewhat more numerous, seemed to occur in relative quantity only sporadically. Incidentally, | failed to find A. ruta-muraria and A. ceterach in South Kerry, though both have been recorded there. Druery (1901) refers to two further finds of what he listed as ‘Confluens Stabler’ under A. trichomanes, both in Ireland, by W. Forster and J.J. Smithies (no further details): these are omitted from Jermy et a/, presumably because these records cannot be validated. Druery (1910), which has an illustration of the hybrid and suggests A. scolopendrium x trichomanes as more credible parentage than A. marinum x trichomanes as some had suggested, still refers to five finds. If, notwithstanding the absence of evidence, Forster and Smithies did find A. x confluens, it would mean that of six finds in the British Isles, four were made in Ireland. ACKNOWLEDGEMENTS | thank the staff of the Fern Section, BM (NH), and Profesor T. Reichstein for confirming my determination. REFERENCES DRUERY, C.T., 1901. The book of British Ferns. London. DRUERY, C.T., 1910. British ferns and their varieties. London. JERMY, A.C., ARNOLD, H.R., FARRELL, L. & PERRING, F.H., 1978. At/as of ferns of the British Isles. London. LOVIS, J.D., 1975. Asplenium in STACE, C.A. (Ed.) Hybridisation and the Flora of the British Isles. London. LOVIS, J.D. & VIDA, G., 1969. The resynthesis and cytogenetic investigation of x Asplenophyliitis microdon and x A. jacksonii. Br. Fern Gaz. 10: 53-67. B.J. RUSH, 17 Toronto Road, Ilford, Essex. SHORT NOTES 303 AN ABERRANT FORM OF EQUISETUM TELMATEIA FROM THE ISLE OF WIGHT Whilst carrying out a botanical survey of Luccombe Chine Ledge (40-583794) on the south coast of the Isle of Wight on 5th July 1981, | was surprised to find several striking specimens of Equisetum te/mateia Ehrh. growing amongst an abundance of typical vegetative shoots of the species. The specimens in question were some 500 mm high, had moribund tips to the shoots and were producing a proliferation of side shoots, each terminating in a small cone (Fig. 1). Four or five such specimens were detected in relatively open ground over an area of some five square metres. FIGURE 1. Aberrant shoot of Equisetum te/mateia, Luccombe Chine Ledge, Isle of Wight, with numerous small cones. 304 SHORT NOTES The site is a coastal ledge produced by gault clay slipping over lower greensand deposits. The habitat is kept open by virtue of the shifting nature of the substrate. EF. telmateia is abundant and is the only Equisetum present in the immediate area. Because of the presence of gault clay and greensand, the associated species include both calcicoles and calcifuge species. The main associates are Phragmites communis, Tussilago farfara, Eupatorium cannabinum, Prunella vulgaris, Blackstonia perfoliata, Betula pubescens, Pulicaria dysenterica, Carex flacca and Lotus pedunculatus. ACKNOWLEDGEMENT | am grateful to Dr. C.N. Page for identifying the specimen and for drawing my attention to the interest of the find. C.R. POPE, Haverstreet, Isle of Wight. C-GLYCOSYLXANTHONES IN TECTARIA C-glycosylxanthones are phenolic compounds which occur sporadically in ferns. They have previously been reported from Asp/enium, Athyrium, Cardiomanes, Ctenitis, Elaphoglossum, Hymenophyllum, Marsilea and Trichomanes (see Richardson, 1982). A survey at New York Botanical Garden has discovered the compounds in the following genera: Davallia, Humata; Nephrolepis (Richardson, 1982); Acystopteris, | Gymnocarpium and Woodsia (Richardson & Thaddeus, 1983). C-glycosylxanthones have now been found in a species of Jectaria. The compounds were not detected in 21 other species of 7ectaria and in 13 related genera. Tectaria decurrens (Presl.) Copel. (NYBG number 329/76) was collected in the Enid A. Haupt Conservatory at NYBG. A voucher specimen was deposited in the NYBG herbarium. The extraction and detection process has been previously described in detail (Richardson, 1982). The two-dimensional chromatograms of the extract and its hydrolysis products indicated the presence of three C-glycosylxanthones. Twoof these were identified as mangiferin and isomangiferin by co-chromatography with authentic compounds from Asp/lenium montanum Willd. A survey of herbarium material of Po/ystichum and related genera failed to reveal the presence of C-glycosylxanthones in further taxa. The following genera and numbérs of species were examined: Jectaria (21); Hemigramma (2); Quercifilix (1); Cionidium (1); Tectaridium (11); Fadyenia (1); Pleuroderris (1); Hypoderris (1); Dictyoxiphium (1); Camptodium (1); Stenosemia (1); Heterogonium (2); Cyclopeltis (2) and Didymechlaena (1). The rare occurrence of C-glycosylxanthones in 7ectaria and related genera is typical of ferns. The only exception occurs in the Davalliaceae where almost half the examined species were found to contain the compounds (Richardson, 1982). On the basis of this evidence, it seems unlikely that C-glycosylxanthones will prove of much taxonomic value in the ferns. — REFERENCES RICHARDSON, P.M., 1982. C-glycosylxanthones in the fern genera Davallia, Humata and Nephrolepis. Phytochemistry. RICHARDSON, P.M. & THADDEUS, E. C-glycosyixanthones in the fern genera Acystopteris, Gymnocarpium and Woodsia. Phytochemistry (Submitted). P.M. RICHARDSON & E. LORENZ-LIBURNAU, New York Botanical Garden, Bronx, NY 10458, U.S.A. SHORT NOTES 305 BRITISH PTERIDOPHYTE RECORDS Compiled by A.R. Busby Since the appearance of the ‘Atlas of Ferns’, | have compiled an annual list of additions and corrections that have been notified to me by the Biological Records Centre, BSBI vice-county recorders and members of the BPS and the BSBI. Past lists have been published in the BPS ‘Bulletin’ but from now will appear in the Fern Gazette. The records are presented thus: 100km square (letters are used for Irish grid Squares to avoid confusion)/10km square followed by the recorders name. Nomenclature follows the At/as. The following additions and corrections have been received up to the 1st November 1982. PRE 1950 8.1 Botrychium lunaria 63/04 Unknown, 63/11 W. G. Clarke. 9.1 Ophioglossum vulgatum 12/90 S. B. Evans, 36/77 |. Martin. 21.8 Asplenium viride 17/53 not post-1950 as indicated. POST 1950 1.1. Lycopodium annotinum 18/62 C.W. Murray. 1.2 L.clavatum 18/32 .G. Halliday, 18/33, 53, 72 C.W. Murray. 18/60E. Vickers, 34/30 D.S. Lambert, 35/07, 16 M. Martin, 43/36 M. Robson, ; Lycopodiella inundata 18/61 M. & E. Vickers, 29/50 M.J. Marshall. Huperzia selago 18/14 M. Coulson, 18/33, 34, 36, 44, 50, 54, 63 C.W. Murray, 18/60 E. Vickers, 21/94 K. Hearn, HOO/77, 88, 97 A. Willmot, H23/78 J. McNeil. Diphasiastrum alpinum 18/24 M. Coulson. Selaginella selaginoides 17/04 J. Clark, 18/56 C.W. Murray. S. kraussiana 22/50 I.K. Morgan, 31/03 J.G. Keylock, 51/25 J. Byatt. Isoetes lacustris 18/13, 33 G Halliday, 18/55, 63, 71 C.W. Murray, 18/62 A. McG. Stirling, HOO/77, 97 A. Willmot. Isoetes echinospora 07/94 A. Slack, 17/74 A.C. Jermy, 22/86 B. Seddon, 25/97 O.M. Stewart. Equisetum hyemale 18/35, 45 C.W. Murray; H34/30 D.S. Lambert. E. x trachyodon 18/45 C.W. Murray, 35/78 G.A. Swan. E. variegatum H33/43 P. Hackney. E. fluviatile 07/94 and 17/04 J. Clark, 18/41 H.J. Birks, 18/32, 37, 43, 54, 60 C.W. Murray, 52/35 Unknown, 63/02 P.W. Lambley, 63/11 E.T. Daniels; HOO/97 A. Willmot. E. arvense 07/93, 94 and 17/04 J. Clark, 18/50, 54 C.W. Murray, 18/72 M. McCallum Webster; HOO/78 A. Willmot. 7.4x3 E. x litorale 22/51 A.M. Pell, 25/64 O.M. Stewart, 51/27 J. Byatt. 7.5 €. pratense 18/35 C.N. Page. 7.6 €E. sylvaticum 18/50 C.W. Murray, 18/72 M. McCallum Webster, 22/13 S.B. Evans, 34/84 B. Oddie. 7.7 €E. palustre 07/93, 94 and 17/04 J. Clark, 18/54, 60 C.W. Murray, 25/98, 35/39 M. Martin, 63/00 E.A. Ellis; HOO/98 A. Willmot. 7.8 E. telmateia 18/25, 26, 36 C.W. Murray. 8.1 Botrychium lunaria 41/86 W.E. Warner, 43/08 A. Willmot, 62/28 Miss C. Forrest; H24/73 D. Riley, H24/81, 84 D.S. Lambert. 9.1 Ophioglossum vulgatum 22/72 J. Humphrey, 25/46 P. Hopkins, 25/67 O.M. Stewart, 34/74 B. Oddie; H24/81 D.S. Lambert. 10.1 Osmunda regalis 18/24, 35, 41, 43, 63 C.W. Murray, 18/47 B. Philp, 24/45, 55 D. Glyn Jones, 41/96 T.R. Price, 51/44 J.E. Smith, 52/99 E.T. Daniels, 63/11 M. Wiggington. 11.1 Cryptogramma crispa 17/42 A. Kelham, 18/41 C.W. Murray, 18/71 H.J.B. Birks. 14.2. Hymenophyilum wilsonii 18/41 H.J.B. Birks; H10/07 A. Willmot. 16 ~=P. vulgare agg. 42/56 R. Lamb. 16.1 Polypodium vulgare 18/33, 42, 43, 53 G. Ramsey, 25/89 J. Martin, 32/46, 57 M.H. Rickard, 32/47 A.R. Busby, 42/44 J.G. Sampson, 45/06, 15, 16 J. Durkin; HOO/ 78, 77, 97, H10/07 A. Willmot. 16.2 P. interjectum 07/93, 94 and 17/04, 05 J. Clark, 18/44, 54G. Ramsey, 51/06 J.E. Smith, 54/41 F.E. Crackles, 63/10 P. Kirby. _— — NO eB —_ = N NNNN ® OOP WN =< No -& WN 306 SHORT NOTES P. x mantoniae 18/33 G. Ramsey, 25/97 M. Martin. P. australe 32/25, 29 P.M. Benoit, 32/39 A.R. Busby, 32/46 M.H. Rickard. P. x shivasie 25/97 M. Martin. Pteridium aquilinum 07/93, 94 and 17/04,05 J. Clark, 18/37, 41 C.W. Murray, 45/36 J. Durkin, 53/34 N.J. Hards. Phegopteris connectilis 18/26, 53 H.J.B. Birks, 18/54, 62, 63 & 72 C.W. Murray, 21/83 P. Bullard, 22/73 |. Vaughan, 32/46 M.H. Rickard, 38/73 D. Welch; HOO/77 A. Willmot. Oreopteris limbosperma 18/36, 41, 56 C.W. Murray, 41/95, 96 Surrey FC, 43/34 A. Willmot, 45/15, 16, 25 J. Durkin; HOO/77, 88, 97, 10/07 A. Willmot. Asplenium scolopendrium 17/04 J. Clark, 18/32 C.W. Murray, 18/60 M.McCallum Webster, 36/92 J. Durkin, 43/08 A. Willmot, 45/24, 26 J. Durkin, 53/34 N.J. Hard, 62/19, 63/01, 02, 10, 23, 32, 40, 42, 50 E.T. Daniels. A. adiantum-nigrum 07/93, 94 and 17/04 J. Clark, 18/41 C.W. Murray, 18/72 H.J.B. Birks, 35/18 M. Martin, 53/51 R. Stevenson, 53/70, 71 P.W. Lambley, 53/93, 63/00.01, 04,11, 12, 14, 41 E.T. Daniels, 62/01, 23 Nature in N E Essex; H34/13 H.D. Megaw. A. onopteris HOO/88, 98 A. Willmot. A. marinum 07/94 J. Clark, 18/32 C.W. Murray, 18/52 M. Coulson. A. trichomanes agg. 18/71 M. McCallum Webster, 34/79 J.G. Sampson, 41/93 J.E. Smith, 52/98, 53/93, 63/00, 03, 10, 11, 13, 20, 22, 33, 50 E.T. Daniels. 21.7a A. trichomanes, subsp. trichomanes 18/33, 45 G. Ramsey, 22/98 P.M. Benoit. 21.7b A. trichomanes, subsp. quadrivalens 17/04, 64 J.W. Clark, 18/33, 43, 44, 54, 62, 72 G. 21.8 29 21541 22.1 23:1 23:2 24.1 26.2 2x3 26.3 ZUM 212 203 27.5 27.8 27 27. 9x8 Ramsey, 35/95, 96, 97, 45/05, 06, 15, 19, 25, 26, 46/21 J. Durkin. A. viride 18/26 H.J.B. Birks; HOO/88 A. Willmot. A. ruta-muraria 22/25 A.O. Chater, 35/89 J.G. Sampson, 45/15 J. Durkin, 53/34 N.J. Hards, 53/93 P.W. Lambley, 62/19, 63/00, 11 E.T. Daniels. A. ceterach 17/48 C.W. Murray, 18/62 M. Gregory, 22/56, 67 A.O. Chater, 42/55 R. © Lamb, 43/37 L. Storer, 45/16 J. Durkin, 53/42 R. Stevenson, 62/39 J. Silverwood; H24/72 D.S. Lambert, H34/02, 03, 04, H.D. Megaw Athyrium filix-femina 07/93, 94 and 17/04 J. Clark, 18/37 C.W. Murray, 22/25 A.O. Chater, 51/05, 34 R. Clarke, 52/65 C.D. Preston, 63/02 A.L. Bull, 63/33 E.T. Daniels. Gymnocarpium dryopteris 18/34, 71 H.J.B. Birks, 18/54, A. McG. Stirling, 22/53 R. Woods, 33/25 Unknown. G. robertianum 31/03 Miss C.J. Biddens, 34/61 P. Jepson. Cystopteris fragilis 18/71 C.W. Murray, 32/57 M. Henshall, 35/96, 45/16 J. Durkin; HOO/88 A. Willmot, H33/18 D.S. Lambert, H34/24 H.D. Megaw. Polystichum aculeatum 32/17 A.C. Powell, 32/46 A.R. Busby, 34/74 B. Oddie, 36/01 N.F. Stewart, 43/08 A.Willmot, 42/25 J. Durkin, 63/10, 20, 21 E.T. Daniels. P. x bicknellii 18/52 M. Barron. P. setiferum 22/56 A.O. Chater, 41/84 J.E. Smith, 41/85 J. Knight, 41/93, 51/06 K. Page, 43/23, 32 A. Willmot, 45/04 J. Durkin, 51/27 A.C. Leslie, 53/92, 62/19, 63/01, 33 E.T. Daniels, 62/18 P.W. Lambley. Dryopteris oreades 18/42 C.W. Murray, 18/51 M. Coulson, 18/62, 71 H.J.B. Birks, 25/86 O.M. Stewart; HOO/97 A. Willmot. D. filix-mas 07/93, 94 and 17/04 J. Clark, 18/33, 43, 46, 61 C.W. Murray, 18/71 M. McCallum Webster, 35/36 M. Martin. D. affinis 17/04 J. Clark, 22/35 A.O. Chater, 32/57 M.H. Rickard, 41/24 F. Rose, 41/94 Surrey FC, 45/14, 24, 26 J. Durkin, 43/17 N.J. Hards. D. aemula 18/43, 46 C.W. Murray, 18/60 H.J.B. Birks 22/04, 32/46 M.H. Rickard. D. carthusiana 12/72 F.H. Perring, 18/25, 37 H.J.B. Birks, 41/80 J.G. Sampson; 41/83 D.C. Kelly, 43/37A. Willmot, 45/16, 15 J. Durkin, 51/05 R. Clarke, 52/89, 62/29, 63/13, 14/33 E.T. Daniels. D. austriaca 07/94 and 17/04 J. Clark, 18/43 C.W. Murray, 22/25, 35 A.O. Chater, 32/46 M.H. Rickard, 53/34 N.J. Hards, 53/94 E.T. Daniels. D. x deweveri 63/31 E.T. Daniels. 27.10 D. expansa 22/87 A.O. Chater, 28.1 Blechnum spicant 07/94 and 17/04 J. Clark, 18/37, 41, 54 C.W. Murray, 18/71 M. McCallum Webster, 43/33 A. Willmot, 63/11, 31 E.T. Daniels. REVIEW 307 Pilularia globulifera 17/04 A. McG. Stirling — a new for V.-C. 103, 63/11 E.A. Ellis. Azolla filiculoides 33/49 |.D. Wallace, 51/16, 17 J. Montgomery, 51/26 J. Byatt, 63/20 E.T. Daniels, 63/10, 11, 32 P.W. Lambley. Corrections to previous records 1.2. Lycopodium clavatum delete 63/01, 22, 23, 42, 51 fide P_\W. Lambley. 7.1. Equisetum hyemale delete 17/32 fide A.C. Jermy. 29.1 Pilularia globulifera delete 63/01 fide P.W. Lambley. REVIEWS THE FERNS OF BRITAIN AND IRELAND by C.N. Page, pp xxi, 447; 102 figures, 1982. 228 x 152mm. Cambridge University Press. Price: paperback £15 (SBN O 521 29872 5), hardback £40 (SBN O 527 23213 9). This is the first book to illustrate and describe every known species of British fern, fern ally and hybrid. As such it is an important addition to British botanical literature and will no doubt compete with We/sh Ferns as the principal work on British pteridophytes. It is written to a high scientific standard and yet is very readable at the amateur level. In the introductory sections there are several unusual features. The standard dichotomous key has been replaced by two innovative keys — achart key using easily sketched characters, and a multi-access key based on other distinctive features. Both are designed to help the beginner run a specimen down to a small group of genera. A diagram giving altitude ranges of species will be very useful, and a series of maps of environmental factors are fascinating, although that describing ‘Oceanity and Continentality’ is difficult to understand. The main text runs to about one and a half pages for each taxon and consists of a full guide to identification, notes on the natural range of variation, sources of possible confusion, technical confirmation and lengthy field notes. Taxa are arranged in five groups — ‘primitive’ ferns; modern frond forming ferns; clubmosses; quillworts; and finally horsetails. Within each group arrangement is alphabetical with hybrids again arranged alphabetically at the end of a genus. While this is an unusual arrangement | very quickly found it easy to use. Afull systematic list is included atthe beginning of the book. The main feature of this section, and indeed of the book, is the illustrations. A silhouette of a typical frond, or more usually a range of fronds, is given for every taxa. | was particularly pleased to see the illustrations of immature fronds — a traditional cause of confusion. Unfortunately two of the subspecies of Dryopteris affinis (Lowe) Fras.-Jenk. are represented only by part fronds and one is not illustrated at all. Another minor criticism is the choice of deceptively small fronds of D. oreades Fomin. These illustrations are reproduced from actual pressed material, they are therefore technically accurate — in effect modern versions of the nineteenth century nature prints. They are however, unlike their predecessors, not attractive, the black silhouette is often washed out to the point that in places it merges with the white background. In most cases the main illustration give no idea of growth form. To rectify this, habit sketches are included for many species. These are very useful for species like the crispy D. aemula (Ait.) Kuntze, while others leave something to be desired, for example the one of D. dilatata (Hoffm.) A. Gray looks little like that species. Most latin names used here have become familiar in recent years; where names are in dispute | am pleased to see that Dr Page has erred on the conservative side, e.g. Thelypteris palustris Schott is maintained and Po/ypodium australe Fée is retained for the Southern Polypody, despite pressure for achange toP. cambricumL. It is however, 308 REVIEW perhaps more surprising that Phy/litis and Ceterach have not been amalgamated in Asplenium. Problems of taxonomy within the Cystopter/s fragilis (L.) Bernh. aggregate have not been resolved: C. dickieana Sim is treated in the narrow sense in the main entry, i.e. to include only those plants from sea caves in Kincardine, but under the section on variation, plants possessing non-spiny spores but more closely resembling C. fragilis in frond form are also considered C. dickieana. Every taxon has been given a common name — in many cases for the first time, while some of the old familiar names have been changed. These changes will unfortunately cause some confusion but in most cases they are probably justified, particularly where names are inappropriate, however some losses, e.g. Intermediate Polypody for Western Polypody, achieve little in my view. The text entry for each species begins with short sections on preliminary identification, guides to habit and occurrence, and a map, which, though only small in size, shows distribution together with an indication of frequency. Unfortunately there is no indication of distribution outside the British Isles, except for one comment | noticed alongside Athyrium flexile (Newm.) Druce — ‘endemic’. The next section is a full identification guide where key characters are given in italics. The paragraph on variation only covers the range of form normally found in the wild — it is not a guide to garden varieties. Another section gives technical details e.g. chromosome number and other microscopical ways of separating some of the more difficult taxa. ‘Field Notes’ terminate the account of each species and are a highlight of the book allowing the author to share some of his observations collected during many year’s field work. Particularly useful in this section are lists of commonly associated ~ species. Perhaps inevitably coming from the pen of Dr. Page, the genus Equ/setum gets very full coverage with the text enhanced by numerous diagnostic sketches. These will be very welcome to those who find this a difficult genus. At the end of the book there are sections on growing ferns from spores, conservation, further studies needed and a bibliography very full on modern references, although most major Victorian works are omitted. Overall this is an excellent book, innovative in many ways. It is a book which achieves its principal objective, that of being a field guide. It is not a book aimed at the grower although there will be much in it to interest him. There are afew typographical errors, most are unimportant, but Cryptogramma is consistently misspelt as Cryptogamma, and the illustrations must have a ‘black mark’ for their lack of appeal. . At £15 for the paperback edition (£40 hardback) the book is expensive, but the factual quality of the contents more than make up for the shortcomings of the illustrations and the presentation inevitable in a soft cover. M.H. RICKARD REVIEW 309 ATLAS DE DISTRIBUTION DES PTERIDOPHYTES ET DES PHANEROGAMES DE LA SUISSE by M. Welten & H.C.R. Sutter, Vols 1 and 2, pp. 716 and 698 with 2572 maps + 1:500,000 Landeskarte der Schweiz and 17 transparency overlays in end pocket of vol. 2. 296 x 210 x 50mm. Birkhauser, Basel, 1982. ISBN 3-7643-1307-2 and 1308-0. Price SFr. 158 each volume. These two massive tomes show the detailed distribution of the vascular species found in Switzerland. Base maps are shaded and show the topography very effectively; two maps are given to a page. Four symbols are used to indicate frequency and altitudinal range: circles for lowland, triangles for highland, solid symbols for abundant to frequent, open symbols occasional to rare. Where additional data have been obtained from herbarium specimens or literature records these are indicated on the maps by H or L respectively. Seventeen overlays are given, some to habitats such as broad-leaved or coniferous forest, bogs and natural grassland, others to geological features and rock types, altitude and, most usefully, a key to the Swiss national maps at the 1:100,000, 1:50,000 and 1:25,000 scales. A map of the country at 1:500,000 scale is folded in the end pocket. Volume 1 contains some 30 pages of introduction, each repeated in three languages, German, French and Italian. The fern distributions are shown on 82 maps (79 species and two hybrids, Equisetum xtrachydon and Asplenium x alternifolium (as breynii)). Names follow those used in the British Atlas except Diphasium is used instead of Diphasiastrum and Thelypteris includes 7. /imbosperma and T. phegopteris. It is interesting to note the rarity of Woodsia i/vensis in this country of many potential habitats, emphasising its low level and southerly requirement. There are only five extant sites for W. pu/chel//a Bertol, a species not seen in Britain. The importance of Ticino Canton in the southern foothills of the Alps as an area where more Atlantic species can overwinter is reflected in the distribution of Po/ystichum setiferum, Matteuccia_ struthiopteris, Asplenium onopteris, Adiantum capillus-veneris and Osmunda regalis. Verified records for Cystopteris dickieana are scarce and only three extant sites are known. A very large percentage of the British flora is found in Switzerland and much ecological data can be extrapolated from the distribution maps given. Although an expensive book it is one that all institutes with an interest in European botany should try to obtain. K.P. KAVANAGH FERNS AND ALLIED PLANTS WITH SPECIAL REFERENCE TO TROPICAL AMERICA by Rolla M. and Alice F. Tryon, pp. xii, 857, 2028 figures, Springer- Verlag, New York, 1982. Price: This splendidly produced tome provides a storehouse of information about the genera of pteridophytes occurring in the New World. It could perhaps best be described as a generic ‘flora’ of American ferns since the content and organization of the text follows the pattern of a regional flora. For each genus there is provided a description (which seems rather meagre for a work on this scale) with synonymy followed by sections on ‘systematics’, ecology, geography, spores, cytology (if known) and a bibliography. The descriptions are supplemented by abundant and excellent figures and close-up 310 REVIEW photographs showing details of sori, indumentum and venation. Dot maps for each genus show its distribution in the New World and the range worldwide is also enumerated. The ecological notes on the American species are particularly valuable in view of the widely scattered floristic literature. The excellent habitat photographs mostly taken by W.H. Hodge help to provide a living picture of the wide variation in habit and form existing among modern ferns. The section entitled ‘systematics’ gives an overview of each genus, usually with a synopsis and/or a key tothe species, which should again be of considerable practical value. Generic limits tend to be widely drawn but most larger genera are subdivided and the synopses are essential for comprehension of the variation within them. The final sections on spores and cytology are also covered in portions of the introduction (as is biogeography) and the cytological information is conveniently summarized there with brief comments on each family. | would support the hypothesis put forward that many present day chromosome numbers are polyploids on a base number of 9 or 10. : For each family the description classifies the stele as proto-,soleno- or dictyostelic and includes the gametophyte but more details of both would have been valuable. A generic key is provided and also a synopsis when the family is subdivided. It is in the description and illustration of the spores that the most substantial amount of new information has been introduced. For every genus there are beautiful SEM micrographs of the spores showing the external appearance and often also the complex structure of the spore wall. The study of fern spores at a magnification of x 1000 to x 10000 has literally opened up a new dimension to pteridologists and it is to be hoped that Dr. Alice Tryon will be able to publish more of her work in this field. There is no discussion of the classification adopted and the arrangement is conservative in that among the homosporous leptosporangiate ferns all families comprising ‘Polypodiaceae’ sensu lato have been kept together. The more primitive indusiate families are thus separated from the more advanced by the very distinct exindusiate gymnogrammoid ferns (Pteridaceae and Vittariaceae). Within the group 24 families are recognized and if the subfamilies and tribes are also taken into account | find myself in almost complete agreement with their circumscription, if not with their arrangement. This general accord among pteridologists on the identification of Suprageneric taxa shows the progress that has been made since the first ‘modern’ scheme was published by Christensen as recently as 1937. FRANCES M. JARRETT FERNS AND FERN-ALLIES OF MEGHALAYA STATE, INDIA by A.K. Baishya and R.R. Rao x +161 pp. 190 x 246mm, 1962. Scientific Publishers, Jodhpur. Price US $20.00, from United Book Traders, Ratanda, Jodhpur 342001 India. Meghalaya State, formerly part of Assam abutts onto the north-eastern border of Bangladesh south of the expansive Brahmaputra river and Bhutan. It contains a low range of hills, the Khasi which are a spur from the high Nagi Hills on the Burmese border. Interestingly 133 species belonging to 62 genera recorded there (i.e. some REVIEW 311 53%) are found in the South Indian hills. Similarly some 140 species (56%) are common to the hills in and adjacent to the western Himalaya. This local flora traditional in its approach is a very good example for others to follow. Accounts on the geology, physiography soil and climate are given with a general account of the vegetation and role of pteridophytes in the State. Useful synonyms are given, as are:-keys to family, genus arid species. The book is better produced than many from India. A.C. JERMY REVIEW ASPECTS OF PLANT SCIENCES (VOLUME 6), PTERIDOPHYTES: their morphology cytology, taxonomy and phylogeny, edited by S.S. Bir. 253pp., 240 x 415mm, 1983. Today and Tomorrow’s Printers and Publishers, New Dethi, India 110005. Price Rs 95.00 : $79.00. This book is part of a serial on plant sciences and is in continuation of work discussed in Volume III (1980). We are told in an introductory note that “it reflects some aspects of the work done in Pteridology in India during the recent years’’. The contributions deal with the ecology, distribution, morphology, taxonomy and cytology of Indian members. Its authors and in particular its hard working editor have in fact reviewed papers on a world basis over the last 20 years at least, and often even older, e.g. ‘“The work on the (sic) Pteridologic geography has been discussed by Christ (1910), Winkler (1938) and Tryon (1969, 1970a)’’. The works selected are however relevant to India. The review by D.S. Loyal on ‘Some recent advances in phylogeny and life-cycle of pteridophytes, particularly ferns’, is detailed. Similarly, ‘Chromosome numbers and evolutionary status of pteridophytes of Pachmarhi Hills (Central India)’ by S.M. Vasudeva & S.S. Bir, gives chromosome diagrams for 33 species; only 15 are supported by photographs and of these the plate reproduction is very poor standard. It is interesting to note an octoploid being recorded for Se/aginella. Whilst some of the papers are superficial, the book as a whole will be useful for undergraduate (or specialist graduate) courses. A lot of references are given to papers and books available to in most university libraries. A.C. JERMY 312 THE FERN GAZETTE Original papers, articles, or notes of any length on any aspect of pteridology will be considered for publication. Contributions will be particularly welcomed on floristic, biogeographical and ecological aspects of ferns and their allies, worldwide. Members are especially encouraged to make use of short notes for reporting pteridophyte field observations and records. All may be illustrated with black and white photographs of good technical quality and line drawings. Short papers can usually by incorporated more quickly than longer ones. Contributions should be sent to: C.N. Page, Royal Botanic Garden, Edinburgh, U.K. The /ast date for receiving notes and articles to make the following early summer number is: 30th November each year Authors should follow the general style of this number. Close adherence to the following notes will help to speed the publication of contributions. NOTES FOR CONTRIBUTORS Manuscripts: Copy should be in English and submitted in double-spaced type with adequate. margins, on one side of the paper only. Abstract. All papers, other than short notes, should include a short abstract, to be set at the head of the main text, indicating the scope of the topic and the main conclusions. Headings and sub-headings: These should follow the style of this number. (Primary sub- headings are centred capitals. Secondary sub-headings U and L case side roman. Tertiary sub- headings, if necessary are U and L case italic, ranged to the left). Numbering of sub-heading should be avoided. Latin names. quote the authority at (usually) the first mention only, in the main text but, unless unavoidable for clarity, not in the title. All latin names should be underlined throughout the typed copy. Mlustrations: Any number and combination of line and half-tone illustrations (original drawings or diagrams in ink, or photographs which must be black and white, and of good technical quality) can be included with a manuscript where these help to augment or amplify the text. Photographs should be of the required magnification or larger and need not be made up to full page plates. Each drawing or photograph should be marked on the back with details of author and figure number, and the top edge clearly marked “‘top”’. Illustrations will not be returned to the author unless specifically requested. Figure numbering: Grouped illustrations should follow the numbering system, fig 1a, fig 1b, fig 2a, fig 3, etc. Figure numbers should be applied to illustrations in pencil only or ona transparent overlay. Final lettering will be added by the editor in a style and size consistent with the journal. Figure captions: Type on a separate sheet from the manuscript — include any necessary details of magnification as submitted (the editor will apply any correction or reduction). Reference lists: Please follow closely the style of this number to speed publication. Lists in other styles may have to be returned to authors for re-typing. Reprints: Twenty-five reprints are supplied free of charge to authors, who may order in advance further reprints which will be supplied at cost (plus postage) if requested at time of returning the first proofs. BOOKS FOR REVIEW Books for review in the Fern Gazette or B.P.S. Bulletin should be sent to A.C. Jermy, Botany Department, British Museum (Natural History), Cromwell Road, London, SW7 5BD. BRITISH FERNS AND THEIR CULTIVARS A very comprehensive collection is stocked by REGINALD KAYE LTD SILVERDALE, LANCASHIRE CATALOGUE ON REQUEST FIBREX NURSERIES LTD. Harvey Road, Evesham, Worcestershire Hardy and tender ferns Begonias, Gloxinias, Hederas, Hydrangeas, Primroses, Arum Lilies and Plants for the cool greenhouse Catalogue on request JOIN THE EXPERTS Los Angeles International Fern Society 28 page colour, illustrated journal plus fern lesson monthly international spore store, educational materials and books. $10 domestic, $12 foreign annual dues. = Send to a 3 = LAI ES, 14895 Gardenhill Dr., LaMirada, CA 90638, USA a eee _ THE NIPPON FERNIST CLUB a = —- _ ‘There exists in Japan this large and active os devoted both amateur and edna. For further information write to: _ Dept of Forest Botany, Faculty of Agriculture, Tokyo University, Hongo, _ Bunkyo-ku, Tokyo, Japan 113. INTERNATIONAL GARDEN FESTIVAL LIVERPOOL ‘84 A spectacular event of exhibition gardens i Bs at an international level coe Par art OF a 2 major project to regenerate Merseyside’‘s derelict sites - = From April to October 1984 ae The Society will have a stand and need plants ~ volunteers and.general support Any who can help should contact the Secretary VOLUME 12 PART 5 CONTENTS | MAIN ARTICLES HN ea wien? An ecological survey of the ferns of the dash leade District. Co. Kerry - A. Willmot ; Three new Dryopteris hybrids from Spain and the Canary Islands — Mary Gibby & C.-J. Widén ies Pah ge ee Asplenium bourgaei - a new addition to the flora of Europe ks be pete - W. Greuter et al RD SECA Contributions to the Morphology | of the Tectaroid fom Stenosemia - Subhash Chandra e. The changing role of chemistry i in fin: classifi cation se e SY - Gillian A. Cooper-Driver & C. Haufler fy en Si ie Oe ae The flavonoids of the Osmundaceae j= wn = - Gail L. Sobel & M.D. Whalen ae ie iM ies SHORT NOTES . ee | ee Culcita macrocarpa - a new locaiity i in Spain © oA a Wa - CR. Fraser-Jenkins & M. Lainz MRM ceed Nm he ts ee Azolla filiculoides in Edinburgh a riksi seer -J. Muscott | SS tai SONA hha The rediscovery of Asp/enium x confluens iat ees rate a - R.J. Rush its she Ne i An aberrant form of laine telmateia from the Isle of f Wight - C.R. Pope ba Sera C-glycosylxanthones in Jectaria aehe, in ae - P.M. Richardson & E. Lorenz-Liburnau $5 BRITISH PTERIDOPHYTE RECORDS sts REVIEWS = = ISSN 0308-0838 aa - F F Saft gl ae “7 Eat +e Ga Metloc Printers Ltd., Old Station Road, Loughton, Essex, her ae . \ eer i i, —_ . att atk LOGICAL SOCIETY xin 10 > ‘. J ‘ P 7 “ , a v4 Ms * ny hie a m is i¥ mi $ ; ; % ¥ , Z sc be t ? | THE BRITISH PTERIDOLOGICAL SOCIETY © Officers and Committee for 1984 President: A. E. Jermy Vice-Presidents J. W. Piste Dr R. E. Holttum, F. oe Ka Prof I. Manton, Dr : Honorary General Secretary and x R. Busby, 42 Lewisham Road, Editor of the Bulletin Warley, West Midlands B66 2BS. Tel: 021. Treasurer Dr B. A. Thomas, Biological Sciences Department, Goldsmit ) Rachael McMillan Building, Creek Road, Londo Meetings Secretary K. Kavanagh, c/o British Museum (Natural Cromwell Road, London S' Editors of the Fern Gazette Dr M. Gibby, Dr C. N. Page, Dr Material for publication should be sent to Dr M. Gibby, Briti (Natural History), Cromwell Road, London Editor of the Pteridologist M. H. Rickard, The Old Rectory, Leinth Ludlow, Shropshir Committee M. Barker, J. M. Camus, J. K. Cramp, P. J. Edwards, Re A. Pigott, R. Rush, G. Tonge, Dr A. Willmot, J. R. Membership Secretary _Lt.Col. P. G. Coke, Robin Hill, Stinchcomt Gloucestershire Fern Distribution Recorder a A. J. Worland, 102 Queens Clos : Camb Spore Exchange Organiser — R. F. Cartwright, 13 ek Mill Road, Pe Archivist N. A. Hall, 15 Mostyn Road, Hazel Grove, Swen Cheshire s 7 — The BRITISH PTERIDOLOGICAL SOCIETY was founded in 1891 and today continues as a foct fern enthusiasts. It provides a wide range of information about ferns through the medium publications and available literature. It also organises formal talks, informal discussions, meetings, garden visits, plant exchanges and a spore exchange scheme. The Society has membership which includes gardeners, nurserymen and botanists, both amateur and prof The Society's journals the Fern Gazette, Pteridologist and Bulletin are published annually. Gazette publishes matter chiefly of specialist interest on international pteridology, the Pte topics of more general appeal, and the Bulletin Society business and meetings reports. Membership is open to all interested in fern and fern-allies. Subscription rates (due on IstJa each year) are Full Personal Members £7; Personal Members not receiving the Fern Gazet: Student Members £5; Subscribing Institutions £8. Applications for membership should be s the Membership Secretary (address above) from whom further details can be obtained. (Remi ittan made in currencies other than Sterling are £0.50 extra to cover bank conversion charges). Back numbers of the Gazette, Pteridologist and Bulletin are available for purchase from t Treasurer (address above), from whom further details can be obtained. : ; FERN GAZ. 12(6) 1984 ots STUDIES IN THE FERN-GENERA ALLIED TO 7JECTAARIA, |. A COMMENTARY ON RECENT SCHEMES OF CLASSIFICATION R. E. HOLTTUM Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, England. During the past three years | have made a world-wide survey of the genus Jectaria and other genera which | believe to be allied to it, including a detailed study of all the species of SE Asia, Malesia and the Pacific. In the course of this work it has become evident to me that some of the genera of this group in the Old World have never been clearly distinguished from each other. | have tried to define them more clearly, and in the course of this process have discovered that a few new genera need to be distinguished. | therefore intend to prepare a series of papers on these genera in the region above specified. My work began with a detailed study of the species in the Mascarene Islands (Holttum, 1983); most of these species belong to Ctenitis as first defined (as a subgenus) by Christensen in his Monograph of Dryopteris (1913 and 1920), and within Ctenitis to two species-groups which are mainly distributed in the Neotropics. The species of Ctenitis in SE Asia and Malesia all belong to a different group which has not yet been defined. Those of SE Asia were first brought together by R.C. Ching (1938). The only considerable attempt to write a systematic account of Ctenitis in any part of Malesia is Copeland's on the Philippine species (1960). As M.G. Price (1972: 36) has pointed out, Copeland there included species of several other genera. He also wrongly transferred to Ctenitis other species native in New Guinea. Some of the Philippine species included in Ctenitis belong, in my judgement, to Tectaria, and they show the difficulty of making definitions which will distinguish the two genera clearly. | have a list of 48 species wrongly transferred to Ctenitis; they properly belong to Dryopteris, Tectaria, Dryoathyrium, Stenolepia, Acystopteris, Heterogonium, Lastreopsis, Oreopteris, and to Ctenitis subgenus Dryopsis Ching which in my opinion should rank as a separate genus, more nearly allied to Dryopteris. In 1940 Ching divided the old comprehensive family Polypodiaceae into smaller families, one being Aspidiaceae. Within this family he placed 7ectaria and its allies ina tribe Aspidieae, Dryopteris and genera related to it comprising a second tribe Dryopterideae. When | was preparing my book on the ferns of the Malay Peninsula (1955). | was impressed by the differences between Ching’s two tribes, and in a preliminary paper (1947) ranked them as subfamilies Tectarioideae and Dryopteri- doideae in a larger family Dennstaedtiaceae (| chose this name as a taxonomic indicator, rather than Polypodiaceae, because it them seemed to me improbable that Polypodium could have a place in such a family; | would now accord family status to both subfamilies). | used the distinction ‘costae and smaller rachises grooved, grooves with raised edges’’ for the Dryopteris subfamily, as against “‘costae raised, not grooved, usually hairy” for 7ectaria subfamily. | published drawings (1955: fig. 282 0n p.479, fig.311 on p.529 and fig.3140n p.533) to indicate the differences; |also showed the similarity of the two latter to the condition of Daval/ia in fig.203 on p.350. | was led to observe these differences by Ching’s observations when in 1938 he first distinguished the species of Ctenitis in Asia. | added further observations in a paper published in 1960. In 1965 Ching expressed agreement with me by recognizing a family Dryopteridaceae, distinct from Aspidiaceae (7ectaria and its allies), but he placed Peranema and a few other genera in a separate family. Since 1964 four different systems of classification of the families and genera of ferns have been published, namely those of Nayar (1974), Crabbe, Jermy & Mickel (1975), Pichi Sermolli (1977b) and Tryon & Tryon (1982). Nayar placed the Jectaria group of genera in subfamily Tectarioideae in a family named Dryopteridaceae, 314 FERN GAZETTE: VOLUME 12 PART 6 (1984) Dryopteris and its allies being in a separate subfamily; but he also included the Onocleoid and Athyrioid ferns in other subfamilies. He did not comment on distinctions between genera. Crabbe, Jermy and Mickel placed 7ectaria and its allies as a subfamily in a family named Aspleniaceae; other subfamilies are Asplenioideae, Athyrioideae and Lomariopsidoideae. In 1947 | expressed in some detail my reasons for regarding Asplenioid and Athyrioid ferns as developments on different evolutionary lines; Crabbe, Jermy and Mickel offered no reasons for rejecting this view, with which the other schemes here considered are in accord. Within their subfamily Tectarioideae | accept their list of genera except for Dryopolystichum and Didymochlaena, which | think belong with Dryopteris. Pichi Sermolli includes the genera allied to 7ectaria and also those allied to Dryopteris in one family Aspidiaceae, and discusses the relationships between the genera; | do not agree with some of his ideas on this subject, and will discuss them in some detail below. The Tryons, like Nayar, recognize a family Dryopteridaceae which includes both the 7ectaria and Dryopteris groups of genera and also other groups in several tribes. | cannot accept their allocation of genera to their tribes Dryopterideae and Peranemeae, and will adda note of this after the following discussion which is based mainly on the concepts of Pichi Sermolli. Pichi Sermolli states that if his family Aspidiaceae is to be subdivided (he makes no formal subdivision) the division must be into more than two parts. Part of his evidence for this is that there are four types of rachis structure in the family (they are distinguished in an earlier paper, 1977a: 238). But my distinction between the Dryopteris and Tectaria groups is based on the costae (or costules) of pinnules, and their relationship to the rachis which bears them, not solely on the rachis itself; the shape of the rachis is irrelevant (see Tryon & Tryon 1982: 458). In my judgement, Pichi Sermolli’s statement about four rachis types is neither comprehensive nor fully accurate. He thus defines the ctenitoid rachis: ““when the upper surface of the main rachis is grooved with the groove not open to admit the shallower groove of the secondary rachises or pinna-midribs, the raised edges of the groove of the main rachis being in any case fully fused with the secondary rachises or pinna-midribs also in their median part, where sometimes a junction is swollen”. |judge from this that Pichi Sermolli is describing the condition of Ctenitis apiciflora (Wall. ex Mett.) Ching, which belongs to Ctenitis subgenus Dryopsis Ching and is quite unlike the type of Ctenitis. | regard C. apiciflora as a member of a different genus, still to be described (see below). The distinctive feature of true Ctenitis is that the costae or costules of the ultimate leaflets are more or less prominent and always covered with ctenitoid hairs; and the distal part of pinna-rachises to which the leaflets are attached are usually not grooved. The base of the lamina of the leaflets is sometimes narrowly decurrent, and then the only real difference from Lastreopsis is that the decurrent base is not thickened and does not extend downwards to the next leaflet. My judgement still is that the genera of Pichi Sermolli’s family can be divided into two groups, to both of which (with Ching) | would accord family status. The groups are thus distinguished: Dryopteris group: midribs of ultimate leaflets grooved, the groove of the rachis bearing the leaflets being open to admit the leaflet-groove, the margin of the lamina of the leaflet being decurrent (but not prominent) down the side of the rachis; hairs of ctenitoid type lacking. Tectaria group: midribs of ultimate leaflets more or less prominent (in 7ectaria sometimes slightly grooved) and bearing ctenitoid hairs, usually many. As Pichi Sermolli points out, there are two base chromosome numbers in his family, 41 and 40. The genera which are obviously allied to 7ectar/a all have 40. But it is also evident that some which have 41 are also much more nearly related to 7ectaria FERN-GENERA ALLIED TO TECTARIA 315 than to Dryopteris. A striking example is a species from southern China first named Tectaria sinii by Ching. In frond-form it is indistinguishable from 7ectaria, and its venation is exactly that of the other species which | regard as free-veined Jectaria; but its scales, which are abundant on all axes of the frond, are closely similar to those of Ctenitis rhodolepis (Clarke) Ching and to those of the type species of Ctenitis. Ching later transferred 7. sinii to his genus Ctenitopsis (which he established to accommodate what | would call free-veined species of Jectaria). Ohwi later transferred it to Ctenitis. It should perhaps be regarded as constituting a new mono- typic genus; in any case it presents strong evidence of the close relationship between Ctenitis and Tectaria, which Christensen already recognized in 1913. The upper surface of the midribs of pinnae may appear to lack ctenitoid hairs in Pteridrys and in some species of 7ectar/a, but there are nearly always some such hairs at the bases of pinnae or of the main rachis. In some species of 7ectaria the upper surface of the pinna-midrib is shallowly grooved, never deeply so. Non-ctenitoid hairs occur on the upper surface of midribs of pinnules in species of Christensen’s group of Ctenitis subincisa (in America and Africa: Monograph part 2: 59) and in some species of Lastreopsis. These hairs are longer than normal ctenitoid hairs, acicular (apical cell acute) and do not collapse on drying, as do typical ctenitoid hairs. They are normally accompanied by some true ctenitoid hairs in C. sub/ncisa and its allies, but | cannot distinguish true ctenitoid hairs in some species of Lastreopsis, within which genus there is much variety of dermal appendages. So far as | have observed, the hairs on the upper surface of midribs in pinnae or pinnules of the Dryopteris group are never ctenitoid. In some genera (eg: Po/ystichopsis) slender hairs occur in the grooves. In Stenolepia, Nothoperanema, Acrophorus, Peranema and Didymochlaena there are complex outgrowths along the margins of grooves of the upper surface of leaflets; these structures have widened thick bases but distally are hair-like, though not like ctenitoid hairs. Pichi Sermolli does not define his genera; he only cites the original publication and the type species, and he provides no keys. Thus his generic concepts are not always clear. An example of this is his concept of the genus Heterogonium. The species of 7ectaria which have anastomosing veins can be divided into those which have ample anastomosis with many branched veinlets in their areoles (typical Tectaria) and those which have narrow costal areoles and a variable number of other rather small ones which rarely have free veinlets within them, the few such being short and excurrent (Sagenia of Presl 1836: 86, t.ll, fig.23-25). But there is every gradation from the latter condition to species which have all veins free. This situation is further complicated by the fact that some species with the venation of Sagen/a, or with free veins, have basal pinnae with reduced basal basiscopic lobes, thus differing from the form typical in 7ectaria in which such lobes are much elongate; these species also have more or less dimorphic fronds and sori of various form. One species of this group has anastomosing veins and elongate exindusiate sori; it was made the type of a new genus Heterogonium by Presl (1851: 142). Copeland added another species in 1929 (p.61) and in so doing noted its resemblance to the species originally named Aspidium sagenioides by Mettenius, which has indusiate sori, free veins, and the basal pinnae like those of Heterogonium (in which genus Ihave placed it), not 7ectaria. Ching (1938: 304) established a new genus Cteni/tops/s for species allied to 7ectaria sect. Sagenia, with free veins. In describing Cten/topsis he carefully noted its difference in venation from Ctenitis, but he did not refer to the shape of the basal pinnae and included Asp/dium sagenioides which he cited as type-species. The latter species, and also another placed in Ctenitopsis by Ching, are reported to have 40 chromosomes, as in Jectaria. But Copeland (1947: 123) included Ching’s Ctenitopsis in Ctenitis (41 chromosomes), not noting the difference in venation specified by Ching; 316 FERN GAZETTE: VOLUME 12 PART 6 (1984) at the same time (p.127) he recognized a genus Heterogonium comprising two species as in 1929. In 1949 | pointed out that these two species are not clearly distinct generically from some others which have elongate sori, and that the shape of the basal pinnae is an important common character. | was subsequently able to study a wider range of species and wrote a fuller account of Heterogonium in 1975. This genus, as thus arranged, still seems to me valid. The species have Sagenia venation (even when pinnae are quite broad), or free veins, and their fronds are almost always somewhat dimorphous (in some species much so) and show a range of soral form from round and indusiate to a fully acrostichoid condition, the form of basal pinnae being constant in all of them. This contrasts with the Sagenia section of 7ectaria in which sori are uniform and the basal pinnae always of 7ectaria form. Pichi Sermolli (1977b: 341) rejects my interpretation of Heterogonium and includes Aspidium sagenioides in Ctenitopsis, but does not explain how he would define Heterogonium, a genus which he retains in his list. Copeland later (1960: 285-299) added more Philippine species to Ctenitis. Some of them belong, in my opinion, to 7ectaria, but their venation pattern does not so clearly conform to that specified by Ching for Cten/topsis, and one of them (C. dubia Copel.) is so different from both Ctenitis and Tectaria that | think it should be regarded as representing a genus distinct from both of them. In Africa andthe West Indies also there is a group of species (Christensen‘s group of Dryopteris protensa) some of which in Africa have anastomosing and some free veins (I think that Christensen did not know of the former). Alston (1969: 70) included these species in Ctenit/s, but Kramer (1978: 104) thought them to be closer to 7ectaria. Three species of this group in West Africa are reported to have the base chromosome number 41 (Manton’s Appendix in Alston 1959: 80), an indication that Christensen was right in regarding them as allied to Ctenitis. Alston's key to the genera (p.69) does not distinguish clearly between species which he ascribes to the two genera. Thus in both the Old World and the New there are species which are not clearly assignable to well-defined genera. It is evident that more kinds of characters need to be taken into account in specifying distinctions between genera. Such characters in Ctenitis are the scales and glands. Christensen in 1913 described the scales of tropical American species of Ctenitis, but no-one has made a comparable study of the scales of species in the Old World. Cylindric glands are a constant character of Ctenitis in Malesia and SE Asia and are absent from Jectar/a and its immediate allies. The species of Christensen’'s group of C. sub/ncisa lack the characteristic glands of typical Ctenitis and have very different spores (Tryon & Tryon 1982: fig. 67.23 on p.466). They have also very large fronds and distinctive scales, and might constitute another new genus. The species of Christensen’s group of Dryopteris protensa are characterized by a peculiar frond-form and by a venation distinct from that of 7ectar/a but these have not been clearly specified. The genera which | believe Pichi Sermolli has misplaced in his scheme for the family Aspidiaceae are Didymoch/aena, Dryopolystichum and Rumohra. Didymochl/aena is placed close to Cten/tis with a statement that it has arachis of ctenitoid type. Its rachis is however extremely unlike that of any Ctenitis, being very deeply grooved and lacking ctenitoid hairs. The pinnules are very asymmetric at their bases with a thickened basiscopic margin which is not distinguishable from the costule. The relationship of pinnules to pinna-rachis is obscured by a partial articulation of the pinnules. Near the apex of a pinna, where the pinnules are not articulated, it appears that the pinna-rachis groove is interrupted at the junction witha pinnule. The thick appendages on the margins of the groove near the bases of pinnules (not mentioned by Pichi Sermolli) are comparable with those in a similar position in Peranema and allied genera (see Ching 1966: 25). | believe that Didymochlaena is allied to these genera; it is however pantropic in distribution whereas Peranema and FERN-GENERA ALLIED TO TECTARIA 317 its allies are only in SE Asia. The difference in distribution is perhaps to be correlated with the fact that Didymochl/laena occurs only a medium altitudes in tropical forest, whereas Peranema and its allies are ferns of temperate or subtropic distribution, occurring in Malesia only on high mountains. No-one has ever made a careful morphological study of the pinnae and pinnules of Didymochlaena. Dryopolystichum has grooved pinnae-midribs which near their bases bear some thick hairs on the margins of the grooves. These hairs are not well shown by herbarium specimens and need careful examination from fresh material; | think they may be comparable to the small scales on the margins of grooves in the genera allied to Peranema. | believe that Dryopo/ystichum belongs to the Dryopteris group of genera; its chromosomes have never been examined. Rumohra is the one genus which | believe does not belong either to the Dryopteris or the Jectaria alliance. As | pointed out in 1947, it differs from Daval/lia and allied genera in little except its indusia. Kato (1974) also noted that its stipes are not jointed at their bases. In the very precise dorsiventral anatomy of its rhizome (as admitted by Kato) it is exactly like Dava//ia. Also the two larger vascular strands on the adaxial side of the stipe and rachis unite distally in the rachis to produce a strand X- shaped in section as in Davall/ia (this was not observed by Kato, who does not note that the union of the strands occurs in the rachis in some species of Davallia). Kato publishes a photograph of a young leaflet showing sori almost reniform. In the genus Davallodes, closely allied to Davallia, the indusia are very varied in form in the various species, some being almost reniform @. borneensis (Hook.) Copel.). Pichi Sermolli compares the rhizome anatomy of Rumohra to that of Maxonia, a near ally of Dryopteris, quite unrelated to Jectaria and its allies; but even Maxonia is not comparable in its anatomy to the very specialized dorsiventral structure of Daval/ia. The genera Pteridrys and Pleocnemia, near which Pichi Sermolli places Rumohra, are very different from it in anatomy, scales and frond-form. | adhere to my opinion that Rumohra is related to Daval/lia and has no place in a family named Aspidiaceae. | have not attempted a detailed study of the genera which | would place in a family Dryopteridaceae. So far as | have observed, | would place Peranema, Diacalpe, Stenolepia, Nothoperanema, Acrophorus and Didymochlaena as a natural group allied to Dryopteris but distinct in the peculiar scales described by Ching (1966) for Nothoperanema; | believe that Ctenitis subgenus Dryopsi/s Ching, excluding C. eatonii (Bak.) Ching and allied species, should form an additional genus belonging less nearly to this alliance. As above noted, | also suggest that Dryopolystichum belongs to this group rather than to the 7ectaria group. | believe that Lithosteg/a is allied rather to Polystichum than to the Peranema group where Pichi Sermolli places it. Pichi Sermolli considers that the peculiar indusium of Peranema and Diacalpe is the primitive form in his Aspidiaceae. But | suggest that the indusia of almost all terrestrial ferns, except those included in Schizaeidae by Pichi Sermolli, are all homologous (ultimately derivable from the inner indusium of such ancient ferns as Dicksonia), and accordingly | regard the indusia of Peranema as a late and specialized development. The type species of Steno/epia (S. tristis (Bl.) v.A.v.R.) seems to me to differ only from the later-named Nothoperanema in the prominent receptacle to which its sporangia are attached, the base of the indusium being attached tothe receptacle. The indusium of S. tristis is cuneate, not cordate, at its base but is otherwise not distinctive. There are additional species in New Guinea which appear to belong to Steno/epia. Two of them have been placed in Ctenitis: C. speciosissima Copel. and C. hypolepioides (Rosenst.) Copel. | judge that Lastrea rubiginosa Brack. of Hawaii also belongs to Stenolepia; it has been transferred to Cteni/tis by Copeland. Amonograph of all species of Stenolepia and Nothoperanema, taking into account recent collections from New Guinea, is necessary for an understanding of these genera. 318 FERN GAZETTE: VOLUME 12 PART 6 (1984) R.M. and A.F. Tryon (1982), dealing mainly with Neotropic genera, have recognized a family Dryopteridaceae very different from Ching’s; itincludes the genera placed in Aspidiaceae by Pichi Sermolli and also many others. The Tryons divide their family into six tribes, one of which (p.455) is Peranemeae (including Peranema, Acrophorus, Nothoperanema and Steno/epia but not Didymochlaena) and another is Dryopterideae which includes Dryopteris, Ctenitis and Tectaria. Their tribes Peranemeae and Dryopterideae thus together correspond to the Aspidiaceae of Pichi Sermolli. Other tribes are Physematieae (including Athyrium and Diplazium), Onocleeae, Oleandreae and Bolbitideae; Pichi Sermolli has placed the genera of these groups in other families, and in general | agree with him. As above indicated, | believe that the genera placed by the Tryons in their tribe Peranemeae are closely allied to Dryopteris, and to place Dryopteris together with Ctenitis and Jectaria in another tribe appears to me quite unnatural. The Tryons however give a better description than Pichi Sermolli of the upper surface of costules of leaflets in relation to the axes which bear them, and provide excellent photographs of these features. Like Pichi Sermolli, they place Rumohra near Lastreopsis but do not mention the rhizome anatomy of the former. Thus a taxon corresponding approximately to my concept of the Jectaria group of genera is recognized by Ching and Nayar, also by Crabbe, Jermy and Mickel, though only Ching accords family status to it, and | have strong objections to its being placed in a family Aspleniaceae by Crabbe, Jermy and Mickel. Pichi Sermolli’s Aspidiaceae (excluding Rumohra) corresponds exactly to my concepts of the Dryopteris and Tectaria groups of genera, but he does r.ot think that a natural separation of the family into two parts is possible. The Tryons’ arrangement differs from all the others andis to me unacceptable. lt also appears evident to me that the genera within the Jectaria group in Asia, Malesia and the Pacific are not all well defined, and that to distinguish them more clearly monographic studies involving all species of all genera are necessary. The whole range of leptosporangiate forms is so great and so varied in detail that no-one can have a critical knowledge of all of them. Thus the family concepts in the schemes above mentioned are all based on partial knowledge, and they are all different. The result is that the name Aspidiaceae has no definite meaning unless one refers to a particular author and date of publication. In practice the only way in which the different families named Aspidiaceae can be described is to list the genera in each; the name Aspidiaceae merely means a family typified by the genus Aspidjum Sw., this genus being typified by the species A. trifoliatum (L.) Sw. (Tectaria Cav. is typified by the same Linnaean species, so the names Aspidiaceae and Tectariaceae are equivalent). Until there is general agreement as to the determination of families of ferns, it seems to me irrational to argue about their legitimacy; to suppose that a family name is defined by the publication of a description in words is a delusion. REFERENCES ALSTON, A.H.G. 1959. The ferns and fern-allies of West Tropical Africa. London, Crown Agents. CHING, R.C. 1938. A revision of the Chinese and Sikkim-Himalayan Dryopteris with reference to some species from neighbouring regions. Bu//. Fan Mem. Inst. Biol. Bot. 8: 275-334. CHING, R.C. 1940. A natural classification of the family “‘Polypodiaceae’’. Sunyatsenia 5: 201- 268. CHING, R.C. 1965. Dryopteridaceae, a new fern family. Acta Phytotax. Sinica 10: 1-5. CHING, R.C. 1966. Three new fern genera. Acta Phytotax. Sinica 11: 17-30. CHRISTENSEN, C. 1913. A monograph of the genus Dryopteris, part 1. K.Dansk. Vid. Selsk. Skr. Vi. 10: 55-282. CHRISTENSEN, C. 1920. Ibid., part 2. K. Dansk Vid. Selsk. Skr. Vill, 6: 1-132. COPELAND, E.B. 1929. The oriental genera of Polypodiaceae. Univ. Ca/. Pub/. Bot. 16: 45-128. COPELAND, E.B. 1947. Genera Filicum. Waltham, Mass., Chronica Botanica. COPELAND, E.B. 1960. Fern Flora of the Philippines, part 2. Manila, Bureau of Printing. FERN-GENERA ALLIED TO TECTARIA 319 CRABBE, J.A., JERMY, A.C. & MICKEL, J.T. 1975. A new arrangement for the pteridophyte herbarium. Fern Gazette 17: 141-163. ee Ea Mee: A revised classification:of Leptosporangiate Ferns. Journ. Linn. Soc. Bot. - 123- HOLTTUM, R.E. 1949. The fern-genus Heterogonium. Sarawak Mus. Journ. 5: 156-166. HOLTTUM, R.E. 1955. A Revised Flora of Malaya, Il, Ferns of Malaya. Singapore, Government Printing Office. HOLTTUM, R.E. 1960. Vegetative characters distinguishing the various groups of ferns included in Dryopteris in Christensen’s Index Filicum and other ferns of similar habit. Gard. Bull. Singapore 17: 361-367. HOLTTUM, R.E. 1975. The genus Heterogonium Presl. Kalikasan 4: 205-231. HOLTTUM, R.E. 1983. The fern-genera Jectaria, Heterogonium and Ctenitis in the Mascarene Islands. Kew Bull. 38: 107-130. KATO, M. 1974. A note on the systematic position of Rumohra adiantiformis. Acta Phytotax. Geobot. 26: 52-57. KRAMER, K.U. 1978. The Pteridophytes of Surinam. Utrecht, Naturwet. Studiekring voor Surinam en de Ned. Antillen. NAYAR, B.K. 1974. A classification of Leptosporangiate Ferns, pp. 111-201 in Nayar & Kaur, Companion to Beddome’s Handbook to the Ferns of British India. New Delhi, Chronica Botanica. PICH| SERMOLLI, R.E.G. 1977a. Fragmenta Pteridologiae, VI. Webbia 31: 237-259. PICH| SERMOLLI, R.E.G. 1977b. Tentamen pteridophytorum genera in taxonomicum ordinem redigendi. Webbia 37: 313-512. PRESL, K.B. 1836. Tentamen Pteridographiae. Pragae, Haase. PRESL, K.B. 1851. Epimeliae Botanicae. Pragae, Haase. . PRICE, pie oi Asummary of our present knowledge of the ferns of the Philippines. Ka/ikasan i: 17-58. TRYON, R.M. and TRYON, A.F. 1982. Ferns and allied plants with special reference to tropical America. New York and Berlin, Springer Verlag. 320 FERN GAZ. 12(6) 1984 A NEW SPECIES OF C7EN/7T/IS FROM BORNEO AND A NEW COMBINATION IN THE GENUS R. E. HOLTTUM Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, England. ABSTRACT Ctenitis muluensis, a new species from the Gunung Mulu National Park, Sarawak, is described, and the new combination Ctenitis subobscura (Christ) Holttum made. INTRODUCTION | have in preparation a monograph of the fern genus Ctenitis C.Chr. in SE Asia, Malesia and the Western Pacific. Material from this monograph relevant to two species from the Gunung Mulu National Park is here published in advance, so that their names may be included in the list of pteridophytes in the Park. One is a new species, Ctenitis muluensis, found on limestone; it has very small leaflets densely covered with glands and hairs. The other is widely distributed in Western Malesia and has been given two names in other genera; it is nere transferred to Ctenitis and a statement of its synonymy is presented. Ctenitis muluensis Holttum, sp. nov. Inter species malesianas generis Ctenitidis propter foliola minuta supra pilis glandulisque densissime vestita peculiaris; stipes usque 12cm longus, basi paleis angustis c. 10mm longis vestitus; lamina usque 23cm longa; pinnae usque 10-jugatae; pinnae infimae usque 7cm longae, pluripinnatae, pinnulis infimis basiscopicis et acroscopicis fere aequalibus; foliola tertiaria 3mm longa; sori mediani, indusiis parvis pilis glandularibus minutis vestitis praediti; sporae c. 55mm, saccato-verrucatae, sporodermis granulis bacilliformibus. Type: Borneo, Sarawak: Fourth Division, Gunung Mulu National Park, Gunung Api, between Camp 4 and summit, 1600-1700m, on roots, stems or mossy limestone rocks in montane forest with stunted Leptospermum, 15 April 1978,A.C. Jermy 14156 (holotype, BM; isotype K, SAR). Caudex erect, covered with shiny castaneous scales less than 1mm wide at base, gradually tapering to a filiform apex and up to 13mm long, margins+ entire with a few short glands. Fronds up to 32cm long; stipes c. 12cm long, dark brown, densely covered with glands, short hairs and scattered scales, those near the base similar to the scales on the caudex, those above up to 5mm long, more ovate-triangular with a long attenuate apex, clathrate, red brown in centre and with paler margins, toothed towards base and with short glands above; rachis with similar indumentum; /amina triangular-lanceolate, bipinnate- pinnatifid, pinnae c. 10 pairs grading into apex of frond, lowermost 5-7cm long; acroscopic and basiscopic pinnules of similar length, pinnatisect at base, pinnatifid above with rounded lobes; lamina surface and veins covered with dense glandular stiff hairs above, underneath sparsely covered with appressed hairs; lower surface of costae and costules densely covered with glandular hairs and scales similar to those on the upper part of the stipe, upper surface with stiff short hairs and scattered scales at least on the costae. Sor/ median, indusium reniform covered with glandular hairs; spores saccate-verrucate, sporoderm with small rod-shaped granules c. 55m. The following new combination is validated here, as this species has also been recorded in Gunung Mulu National Park. Ctenitis subobscura (Christ) Holttum comb. nov. Phegopteris subobscura Christ, Bull Herb. Boiss. 6. (1898) 836. Type: G. Schneider 35, Sumatra (P). Synonyms: Dryopteris squamulifera v.A.v.R., Bull. Jard. Bot. Btzg Il, 16: (1914) 9. Type: C.G. Matthew 669, Sumatra, Padang Panjang (BO). Ctenitis mannii sensu Holttum, Revised Fl. Malaya 2: (1955) 497. quoad pl. males. FERN GAZ. 12(6) 1984 321 CHROMOSOME NUMBERS AND ECOLOGICAL OBSERVATIONS OF FERNS FROM EL TIROL, PARAGUAY A.R. SMITH The Herbarium, University of California, Berkeley 94720, U.S.A. and MERCEDES S. FOSTER Museum Section, U.S. Fish and Wildlife Service, National Museum of Natural History, Washington DC 20560, U.S.A. ABSTRACT Chromosome numbers are given for 32 species of ferns from Paraguay, mostly from El Tirol. Included is the first report for the genusAntigramma, n= 144. Collections of Polypodium singeri, P. truncorum, Thelypteris dentata, and T. torresiana are the first reported of these species from Paraguay. About half of the 47 fern species from El Tirol are widespread in tropical America; the remaining species are more narrowly distributed in southern Brazil, Uruguay, Paraguay, and northeastern Argentina. Most of the ferns at El Tirol occur terrestrially in the forest, especially along creeks. Ten species are epiphytes, including three that are restricted to trunks of the tree fern, Nephelea setosa. Vegetative and reproductive growth appear to be concentrated in the months of heaviest rainfall, October to December. INTRODUCTION To date, there are chromosomal counts for very few South American pteridophytes. Reports are available only for plants from the Galapagos (Jarrett et al. 1968) and Amazonian Brazil (Tryon et al. 1975), or for a few species in particular families (eg: Polypodiaceae; Evans, 1963). Herein we report counts for 32 species (in 18 genera) from Paraguay. Our data represent the first counts for 13 of the species (including first counts for the genus Antigramma). For another 12 species, our counts agree with previous reports for the species in other parts of their ranges. Counts for seven species differ from previous reports for the same species in other localities. This last finding emphasizes the need for repeated sampling over the range of a fern species before a complete understanding of its chromosomal complexity can be achieved. Equally poorly known are most aspects of fern ecology in tropical America. We are unable to locate a single report that provides information on habitat, phenology (including dormancy and time of year when meiosis and spore release occur), host specificity, abundance, and revivability for a community of ferns in this region. MATERIALS AND METHODS For all counts, material was fixed in the field in Carnoy’s solution (6:3:1, ethyl alcohol: chloroform:acetic acid) by Foster and was stained with acetocarmine. Chromosome counts (by Smith) are observations of the configuration at metaphase | of meiosis. Photographs of chromosomes were made with a Zeiss photomicroscope. Collections were made by Foster primarily from an ornithological study site at El Tirol (ca. 55°47’'W, 27°11'S; elevation, 170-260m), 19.5km by road + NNE of Encarnacion, Dpto Itapua, Paraguay. Afew additional counts were made from material collected at Parque Nacional Ybycui (ca. 56°59’W, 26°1'S), ca. 30km NE of Ybycui, Dpto Paraguari (abbreviated Y), 26-27 Mar 1980, and at Parque Nacional Cerro Cora (ca. 56°3’W, 22°39’S), Dpto Amamby (abbreviated C), 1-19 Sept 1980 and 30 Oct-4 Nov 1983. El Tirol was visited eight times: 29 Sept-17 Nov 1976; 4Dec 1977-14 Jan 1978; 31 Aug-13 Oct 1978; 1-17 Oct 1979; 17-28 Aug and 23 Sept-3 Nov 1980; 21 Sept-19 Oct 1981; 19 Sept-3 Oct 1982; and 7-23 May 1983. These repeated visits allowed observations in different seasons and years. Voucher specimens are deposited in the Herbarium of the University of California (UC), with duplicates and a few unicates of many collections in MO, LIL, and AAU. A 322 FERN GAZETTE: VOLUME 12 -PART 6 (1984) representative set will also be deposited in the herbarium of the Servicio Forestal Nacional in Asuncion. Rain data for El Tirol were obtained from records maintained at the site by Armando Reynaers. Temperature data, from the Meteorological Service of the Ministry of Defense, are for Capitan Miranda, a town ca. 4km by road SW of El Tirol. STUDY SITES The three collecting localities lie in a zone of temperate moist forest, though Ybycui and Cerro Cord average 15-20% drier and a few degrees warmer than El Tirol (Farina Sanchez 1973). Days with rain per month are fairly constant throughout the year at El Tirol (Fig. 1), though rainfall may fluctuate greatly within months between years. July is the driest month; October, November, and December usually are the wettest. Temperatures vary seasonally; the warmest period occurs from November through February and the coolest from June through August, when temperatures occasionally may drop below freezing (Fig. 2). 38 36 34 32 130 42-340 28 -278 = 170 OS Baila 2 —~ 160 Cr) 12 E = E 150 = 10 5 140 8 c = 130 6 rv 120 | 17-225 18-285 4 I< 69.8 78.6 110 2 100 O J EMA Mie ln te Ava Si Oe Nice URE IM)! VA Mig Jy A S) OS UNe SD (7.6) (6.3) (7.3) (56) (64) (74) (6.4) (7.0) (68) (8.0) (7.1) (6.6) Month Month FIGURE 1. Mean monthly rainfall at El Tirol, Paraguay, January, 1971 — September, 1982. Range and one standard deviation adjacent to dots. Mean number of days with rain for each month in parentheses. FIGURE 2. Mean monthly maximum and minimum temperatures at Capitan Miranda (4km by road SW EI Tirol), Paraguay, January, 1964 — August, 1982. The habitats at El Tirol are extremely varied. The main Tirol site includes about 50ha of forest adjacent to Hotel El Tirol de Paraguay. This forest, which is situated on moderately steep hillsides as well as flat areas, is cut by a series of northeast- and eastward-flowing creeks. Several are permanent; others flow intermittently following heavy rains. The forest is relatively low, with a canopy height in level areas of ca. 15-18 metres; the understory is open. This forest has not been extensively logged, though selected trees have been cut. In the last several years, however, the area has been increasingly disturbed and reduced in size by the cutting of trails for visitors, clearing of land for gardens, electric lines, and a road, and the diversion of a creek for irrigation. Similar areas of forest, some on hillsides, others on flat or only gently sloping areas, are connected by corridors along creeks, though these zones are also rapidly disappearing. Adjacent to the Tirol forest are large areas of tall second growth originally planted in coffee or other crops. Also common (the zone is primarily FERNS FROM EL TIROL, PARAGUAY 323 agricultural) are tung orchards, cattle pastures, and fields cultivated with corn, manioc, or other vegetables. The entire area is characterized by a number of natural springs. These feed into permanent creeks or, in some flat sites, cause the formation of permanently boggy areas that flood with heavy rains. Natural seeps are found irregularly through the forest; these occasionally give rise to small pools or marshy areas. Collections and observations at Ybycui National Park were made either in forest areas that have experienced minimal human disturbance (Zona Primitiva of Comision de Parques Nacionales de Paraguay, 1973) or in areas of extreme disturbance along roads and in park recreation zones. Collections at Cerro Cord National Park were made in areas of moderately disturbed forest, secondary growth, and scrubby pastures. FERNS FROM EL TIROL Below is a complete list of ferns known from El Tirol, plus a few additional Paraguayan species for which we have chromosomal counts. Collections cited (Foster numbers) are from El Tirol unless otherwise indicated. In the list, superscript ‘‘a’’ denotes the first chromosomal count for the species; superscript ‘‘b’’ denotes a count that differs from that previously reported; superscript ‘‘c’’ denotes a count agreeing with previous reports; (C) denotes Cerra Cora National Park and (Y) denotes Ybycui National Park. FIGURE 3. Meiosis in Antigramma brasiliensis showing 144 bivalents (Foster 87-23). FIGURE 4. Chromosomes of Antigramma plantaginea, showing irregular meiosis, with pairs and univalents (Foster 80-2-67). Adiantopsis radiata (L.) Fee © 30 bivalents (79-16). Additional collections: 76-8; 78-2-26; 80-14 (Y); 80-2-26 (C). Adiantum pseudotinctum Hieron:@ c. 58 bivalents (81-5). Additional collections: 76-20; 78-2- 24; 79-12; 80-2-5; 81-4; 80-2-29 (C). Adiantum tetraphyllum Humb. & Bonpl. ex Willd» 30bivalents (80-2-17, C); 29 bivalents (80-12, Y). Walker (1966) reported n = 58-60 for a plant from Jamaica. Anemia phyliitidis (L.) Sw.© 76 bivalents (79-3). Additional collections: 76-12; 76-21; 78-2-10; 80-15 (Y); 80-2-37 (C). Antigramma brasiliensis (Sw.) Moore? 144 bivalents (81-23), Fig. 3; c. 144 bivalents (79-19; 81-35). Additional collections: 76-4; 78-2-18; 81-39; 80-27 (Y). Spores are well-formed, reniform, and 64 per sporangium. This represents the first count for the genus, placed by nearly all recent pteridologists in the asplenioid alliance (Aspelniaceae). The chromosome number indicates that the species is octoploid, based on x = 36. Antigramma plantaginea (Schrad.) Presl|? Meiosis irregular, with many outlying univalents, bivalents, and perhaps a few trivalents (Fig. 4); the best cellshowed c. 731II+ 113lat meiotic 324 FERN GAZETTE: VOLUME 12 PART 6 (1984) metaphase (80-2-67), so the total is perhaps 2n = 288 (octoploid). Spores are 16 per sporangium and are globose, indicating that this fern has unreduced spores and possibly an apogamous life cycle. Additional collections: 76-42; 78-2-59; 81-41; 81-42. Asplenium claussenii Hieron? 36 bivalents (78-2-66; 79-14); c. 36 bivalents (79-13, MO; 81- 44). Additional collections: 78-2-19; 78-2-41; 80-6 (Y). Asplenium hostmannii Hieron. (76-11). Growing with A. claussenii, but differing from that species in the much lower number of lateral pinnae, pinnae broader and longer, with the lowermost the largest or only slightly shorter than the next pair. Also similar to A. /Jaetum Sw., which differs in having a creeping rhizome and more atropurpureous Stipes. Asplenium mucronatum Presl (76-43: 78-2-31; 79-25; 81-15). Asplenium serratum LO ©. 144 bivalents (79-18). Additional collections: 78-2-66; 80-2-6; 80-2- 7; 80-2-45 (C). The only previous count for this species was n= 72 (Wagner, 1963), based on material from Florida. One additional population (81-37) of this species from Paraguay was also examined cytologically, but an exact count could not be obtained. It showed a large number of chromosomal pairs, certainly more than 72. Spores are well-formed, 64 per sporangium. Blechnum australe L. subsp. auriculatum (Cav.) Sota (80-2-53; 80-2-55). Blechnum brasiliense Desv. (79-64; 81-19; 81-22; 80-16, Y; 80-2-44, C). Blechnum ensiforme (Liebm.) C.Chr.4 29 bivalents (80-2-47; 81-18); c. 29 bivalents (79-42). Additional collections: 78-2-51; 78-2-73. In South America, this species has often gone under the name B. meridense (Klotzsch) Mett., but that name is a later homonym of B. meridense Klotzsch [B. fraxineum Willd.]. We are unable to find reliable characters by which Paraguayan material differs from Mexican material; the type of B. ens/forme was collected in Mexico. Blechnum occidentale L© c. 31 bivalents+ 31 univalents [30-31 Il + 29-331, 3 cells examined] (80-2-32, C). Another collection (80-18, Y) also showed irregular meiosis with many univalents and strongly malformed spores; exact counts could not be obtained. This species was previously recorded as n= 62 (eg, Walker, 1973). Hybrids of B. occidentale with several other species are known, but the Paraguayan specimen counted as triploid seems typical for the species. As Walker (1973) discussed, B. occidentale reproduces readily by stolons, so the triploids may be able to maintain themselves indefinitely without producing viable spores. Additional collections: 79-29 (spores malformed); 80-2-50. Campyloneurum lapathifolium (Poir.) Ching? 37 bivalents (80-2-71). Additional collection: 81- 31. Campyloneurum phyliitidis (L.) Presl b 37 bivalents (80- 2-1). Additional collections: 78- 2-16; 78- 2-17; 79-57. Previously counted as n = 72, for example by Wagner (1963) from Florida, Walker (1966) from Jamaica, and Jarrett et al. (1968) from the Galapagos. Paraguayan material generally has a narrower rhizome, with fronds arising at longer intervals, than material from North America and northern South America. Blades of Paraguayan specimens are also narrower. Monographic revision is needed. Cheilanthes chlorophylla Sw.4 30 bivalents (79-39; 79-54; 79-55, MO; 81-12; 81-33; 80-2- 10). One collection (79-55) showed some cells with 27-29 Il+ 2-61. Additional collections: 76-14; 78-2-77; 80-2-64. Cheilanthes concolor (Langsd. & Fisch.) R. & A. Tryon) 30 bivalents (79-1; 79-10). Additional collections: 76-7; 80-24 (Y); 83-2-26 (C). This species previously counted as n = 60, from Taiwan (Mitui, 1968). It has often been placed in Doryopteris [=Doryopteris concolor (Langsd. & Fisch.) Kuhn]; the base chromosome number may lend additional support for placement in Cheilanthes (x = 30, a few 29) over Doryopteris (mostly x = 29). Ctenitis connexa (Kaulf.) Copel.2 41 bivalents (81-27). Additional collections: 78-2-34; 79-46. Closely related to C. subinc/sa (Willd.) Ching, widespread in tropical America; the differences seem rather slight. Ctenitis submarginalis (Langsd. & Fisch.) Ching® c. 41 bivalents (81-13). Additional collections: 78-2-11; 80-2-31 (C). Dennstaedtia globulifera (Poir.) Hieron.© 47 bivalents (81-21; 80-2-70). Additional collections: 79-66; 80-2-69; 83-2-33 (C). Dennstaedtia obtusifolia (Willd.) Moore (78-2-83; 79-47). Didymochlaena truncatula (Sw.) J. Smith® c. 41 bivalents (79-24). Additional collections: 78-2- 33; 80-2-65. Diplazium ambiguum Raddi. No count, but undoubtedly a high polyploid, possibly octoploid (n = 164) (81-14). Additional collections: 78-2-74; 79-49. Most of the tropical American species of Diplazium that are bipinnate-pinnatifid or more divided seem to be octoploid (see Walker, 1966, for counts on D. expansum, D. fuertesii, D. striatum, all Jamaican species of this alliance.) FERNS FROM EL TIROL, PARAGUAY 325 Diplazium cristatum (Desr.) Alston©® c. 82 bivalents (80-2-66). Additional collections: 78-2-37; 78-2-42; 79-27. Doryopteris nobilis (Moore) C.Chr. 4 ¢. 6Obivalents (80-2-48). Additional collections: 76-40; 78- 2-9; 80-23 (Y); 80-2-27 (C). Doryopteris nobilis x pedata var. multipartita 4 c. 9Ounivalents (79-31 ); Meiosis irregular, spores malformed. ¥ b Doryopteris pedata (L.) Fée var. multipartita (Fee) Tryon~ 29 bivalents (79-5; 79-32; 79-33, MO). Additional collections: 76-9; 78-2-7; 80-5 (Y); 80-2-28(C). The species previously counted as n = 60 from the Galapagos (Jarrett et al. 1968), n= 30 from Jamaica (Walker, 1966), and n= 58 (Kurita, 1971). Gymnopteris tomentosa (Lam.) Underw. (83-5; 80-2-43, C). Lastreopsis effusa (Sw.) Tindale® 41 bivalents (79-23; 79-34; 81-28). Additional collections: 78- 2-36; 80-3 (Y). Microgramma lindbergii (Mett.) Sota? 21 bivalents + 30 univalents; 21 II+ 291; 33 11+ 61; 3411+ 41 (80-13, Y), many malformed spores and meiotic abnormalities, but some spores appear well- formed. Additional collection: 80-2-15 (C). Microgramma squamulosa (Kaulf.) Sota (76-6; 78-2-56; 80-11, Y). Microgramma vacciniifolia (Langsd. & Fisch.) Copel.© 36 bivalents (80-2-16, C); ca. 20 II + 341: 34 Il + 6 | (80-25, Y). Evans (1963) also reported n = 36 for a plant from Brazil. Additional counts of species of Microgramma are desirable. Most other species have been reported as being based on x = 37; however, M. vacciniifolia and M. lindbergii seem to be x = 36, with some populations showing strong meiotic abnormalities. Nephelea setosa (Kaulf.) Tryon (78-2-35; 79-61; 81-29). Pityrogramma trifoliata (L.) Tryon (78-2-46; 79-4; 80-2-13; 81-40; 83-2-23, C). Polypodium filicula Kaulf. a 37 bivalents (79-37; 79-38; 79-60). Additional collection: 80-2-34 (C). Polypodium hirsutissimum Raddi (78-2-75). Polypodium pleopeltifolium Raddi@ 35 bivalents (80-2-8); 37 bivalents (81-6). Additional collections: 78-2-15; 79-59; 80-2-35 (C). Foster 80-2-8 showed incomplete pairing in all five meiotic figures examined, from 30-34 II + 2-91(sum, 2n= 70); nevertheless, spores appeared well-formed, 64 per sporangium. Evidence is accumulating that this group of Po/ypodium, probably closely allied to Pleopeltis, has a base chromosome number(s) other than x = 37. Smith and Mickel (1977) reported n = 34 for Pleopeltis angusta, n= 35 or 36 for Polypodium pleolepis and n = 35 for P. friedricsthalianum. Several Old World species of Pleopeltis (Lepisorus) are known to be based on x = 35. Polypodium polypodioides (L.) Watt var. minus (Fée) Weatherby (78-2-14; 79-68; 80-2-19, C.). Sometimes treated as a distinct species, Po/ypodium squalidum Velloso. Polypodium recurvatum Kaulf.© 37 bivalents (80-17, Y). Polypodium siccum Lindm. (78-2-32; 79-36). Polypodium singeri Sota (76-10; 80- 2-49). First reports of the species from Paraguay; listed only from southern Brazil and northeast Argentina (Misiones) by Evans (1969). Polypodium truncorum Lindm.2 37 bivalents (81-43). Additional collections: 78-39; 81-16. First reports of the species from Paraguay; listed only from southern Brazil and northeastern Argentina by Evans (1969). Polystichum platyphyl/um (Willd.) Pres! D ¢. 82 bivalents (81-20). Additional collections: 79-22; 81-24. Previously reported as n = 41 from Mexico (Smith and Mickel, 1977). Pteridium aquilinum (L.) Kuhn var. arachnoideum (Kaulf.) Brade 79-62, MO; 79-63; 80-22 (Y). Pteris deflexa Link (78-2-71; 81-26). Pteris denticulata Sw.2 116 bivalents (81-30, octoploid); 174 bivalents (79-8, MO, dodecaploid); ca. 174 bivalents (79-9). Additional collections: 78- 2-20; 80-2-24, C; 80-2-51; 80-20(Y). 12x is the highest reported chromosome number in Pteris. Tectaria incisa Cav. 78-2-76; 80-2-42 (C). Thelypteris dentata (Forssk.) E.St.John© 72 bivalents (79-51). Additional collections: 78-2-12; 83-2-31 (C). First reports of the species from Paraguay (see Smith, 1971); it is introduced from the Old World and now widely naturalized in tropical America. Thelypteris hispidula (Decne) Reed© 36 bivalents (79-7). Additional collection: 81-25. Thelypteris interrupta (Willd.) lwatsuki 79-65. Thelypteris torresiana (Gaud.) Alston® cc. 62 bivalents (79-6; 79-43; 81-32). Additional collection: 80-2-39 (C). First reports of the species from Paraguay; introduced into the New World and rapidly expanding its range. 326 FERN GAZETTE: VOLUME 12 PART 6 (1984) DISTRIBUTION AND BIOGEOGRAPHY Four species occurring at El Tirol represent the first published records for Paraguay. Polypodium singeri and P. truncorum are recorded from the adjacent Argentine province of Misiones as well as from southern Brazil (Evans, 1969). Two species of Thelypteris, both introduced and widely naturalized in the Neotropics, are also new to Paraguay: Thelypteris dentata (see Smith, 1971) and 7. torresiana, also known previously from Misiones, Argentina, and southern Brazil. Nearly half of the species known from El Tirol are widespread in tropical and subtropical America, with distributions ranging from Paraguay and north-eastern Argentina to southern Mexico and the Antilles. Two species, Thelypteris hispidula and T. interrupta, are even pantropical. Seventeen species have a more restricted range, occurring in Paraguay, northeastern Argentina and southern Brazil, with some extensions into Uruguay and Bolivia. Included in this category are Adiantum pseudotinctum, Antigramma spp., Asplenium mucronatum, Blechnum australe subsp. auriculatum, Ctenitis connexa, Microgramma squamulosa, Nephelea setosa, Polypodium hirsutissimum, P. pleopeltifolium, P. siccum, P. singeri, P. truncorum, Gymnopteris tomentosa (also Peru), Doryopteris nobilis (disjunct in Colombia), and Pteris denticulata (disjunct in Hispaniola and Cuba). Many of the species known from El Tirol are likely to be encountered in other areas of primary vegetation in Paraguay. Although Ybycui and Cerro Cora National Parks were visited only briefly, 22 (of the 47 total species at El Tirol) were collected at one or the other (or both) of these two sites. ECOLOGICAL OBSERVATIONS Habitats and Abundance Of the 47 species and one hybrid recorded from El Tirol, 36 were terrestrial, 10 were epiphytic, one hemi-epiphytic, and one was both terrestrial and epiphytic. The terrestrial species occurred in a variety of different habitats that can be categorised relative to light and moisture conditions. Blechnum brasiliense, Ctenitis connexa, Dennstaedtia obtusifolia, Didymochlaena, Diplazium spp., Nephelea, Polystichum, Pteris deflexa, and Tectaria were found only along banks of permanent creeks in the forest. Dip/azium cristatum occasionally grew on rocks in the creek as well. In most instances, plants of these species were in full shade. Some of the larger tree ferns (Nephel/ea) leaning into the creek beds received filtered sunlight, and the B/echnum also was often in somewhat sunnier areas. Nephelea lined the permanent creeks in a zone extending 1 to 3m from the stream edge at high water. In a few areas it also occurred on high banks 3 to 5m above the stream bed and, in one area of forest, pure stands of 0.5 to 1 hectare were growing. Although stands were bordered by permanent water along one side, most of the ferns were on a moderately steep slope. Whether this colonial habit is indicative of vegetative reproduction, as discussed by Gastony (1973) for Nephelea sternbergii var. acanthomelas in southern Brazil, is not known. Of streambank species, perhaps the most common was Didymochlaena; insome areas it formed pure stands 5to 10m wide to the exclusion of nearly all other vascular plants. Several other terrestrial species also showed a preference for moister areas. Thelypteris hispidula and T. torresiana were found along both creek banks and growing in the forest. The latter thrived in more open areas at the forest edge. Antigramma plantaginea was a locally common species often found in rocky areas around natural seeps and also in dense stands, often tens of metres across, in areas where the soil seemed particularly moist, but where no seeps were apparent. A large number of terrestrial species grew only in the forest. Asp/enium serratum was found only occasionally, usually in clumps of several plants. Antigramma brasiliensis was one of the most common ferns in undisturbed forest; in second FERNS FROM EL TIROL PARAGUAY 327 growth areas it was much less abundant. Isolated plants of Campyloneurum lapathifolium and Doryopteris nobilis were found in deep forest under low light intensities. Although both were relatively rare, the latter was locally abundant in patches several metres across. Doryopteris pedata was moderately common within the forest but also grew in shady areas at the forest edge. Po/ypodium singeri and Asplenium hostmanii grew only within the forest. Lastreopsis effusa, Pteris denticulata, and Ctenitis submarginalis were somewhat less specialized in their habitat preferences. Lastreopsis was found along permanent creeks but also carpeted large expanses of the forest floor in areas of filtered sunlight and thick leaf litter. ln some areas it grew with the Ctenitis which, along with the Pteris, also covered large areas of forest. All three species were especially common along well used paths. In addition, the Cten/tis occurred along forest and second growth edges; this may reflect its greater tolerance of sunny, dry conditions. Several other species occupied a variety of habitats. Adjantum pseudotinctum was scattered in areas of deep shade but grew more commonly in areas of filtered sun at edges of second growth, where it formed carpets. Che/lanthes concolor and Blechnum occidentale similarly occupied both sites. Ad/antopsis was extremely common everywhere in the forest and at the forest edge. Anemia was scattered in the forest, but became much more common in more open areas, such as along paths and at forest edge. Dennstaedtia globulifera grew in large clumps along forest creeks, at forest edge, and in open areas. Though occasionally found in heavy second growth, Cheilanthes chlorophylla was more characteristic of the forest edge, especially where areas had been cleared for pasture. Pinnules of this species were extremely soft and nearly always appeared wilted; however, fronds revived early in the morning, late in the evening, and during rains. Blechnum australe and Pteridium aquilinum were characteristic of the second growth forest/pasture interface. Their blades were extremely leathery and transient wilting was not observed. 7helypteris dentata was especially common along the forest edge; it also was found in more open areas. Thelypteris interrupta and Pityrogramma trifoliata were the only species found primarily in full sun. Though occasionally growing at the forest edge, they occurred most often in old fields and pastures where they sometimes formed dense carpets. The Thelypteris was most characteristic of marshy areas. The epiphytic ferns all grew in shady sites. The most common microhabitat was in the axils of old stipe bases of Nephelea. Asplenium mucronatum, Polypodium siccum, and P. truncorum were found only in this niche. Polypodium truncorum generally formed a carpet of tiny plants along the length of the Nephel/ea trunk. The Asplenium, with pendant fronds to 75cm long, was rarely found below 2m and was distributed irregularly among the tree ferns, which bore zero to several individuals. Two species were epiphytic both on Nephelea and on other trees. Campyloneurum phyllitidis was found on Nephelea, on rocks along the creek, and on fallen trees in the forest. Asp/enium claussenii, a common component of the forest floor, occasionally grew on Nephelea. Microgramma squamulosa and four species of Polypodium (P. filicula, P. hirsutissimum, P. pleopeltifolium, and P. polypodioides) were epiphytic on trunks (not Nephelea) and large limbs (diameter greater than 6cm) of forest trees. The Po/ypodium spp. were generally several metres up on the tree trunk; the last two species were especially common 5 to 6m up; P. Airsutissimum was common even higher. In contrast, P. filicula grew low down, forming large carpeted areas on tree trunks, often within a metre of the ground. Rhizomes of Microgramma squamulosa formed dense mats most commonly on horizontal branches. All of these species seemed to continue 328 FERN GAZETTE: VOLUME 12 PART 6 (1984) growing well when the host tree or limb fell to the ground. This is true also of Campyloneurum phyliitidis. Blechnum ens‘forme, the only hemi-epiphyte, was observed only along permanent creeks. The largest, reproductive plants grew up the trunks of Nephelea. Small, young sporophytes were moderately common in crevices of rocks bordering the creek. Phenology Because observations were not made throughout the year, it is difficult to describe adequately the phenology of the ferns at El Tirol. However, areview of the information available for the months of observation suggests several patterns of activity. In many species reproduction appeared to be correlated with the periods of greatest rainfall, even though the number of days with rain are fairly constant from month to month (Fig. 1). In at least 29 of the 47 species, meiosis occurred from October to December. Some meiotic activity was evident as early as August in Anemia, Antigramma brasiliensis, Campyloneurum phyllitidis, Cheilanthes spp., Diplazium cristatum, Polypodium truncorum, Pteris denticulata, Tectaria, and Thelypteris dentata. In most of these, spores were in earliest stages of development; spore release may not occur until the later, wetter months. Polystichum platyphy/lum sampled in October had fertile fronds bearing sporangia with mature spores as well as sporangia with spores undergoing meiosis; this suggests that reproduction was initiated at least in September. In contrast, Adiantum pseudotinctum and Asplenium serratum were entirely non-reproductive in August and September. Indirect evidence suggests that some species reproduce earlier in the year, during the cold winter months of June to August. Asp/enium mucronatum fronds bore old sori from October through January; sori produced in September were somewhat younger, though still post-meiotic. During the one fall visit, in May, young fertile fronds were present on some Adiantum pseudotinctum, on nearly all Anemia phyllitidis and Thelypteris torresiana, and on most Pityrogramma trifoliate and Polypodium pleopeltifolium. Large numbers of plants of 26 additional Tirol species had no fertile fronds or only old, dried ones. Fertile fronds were found only on a few of nearly 100 plants of Pityrogramma trifoliata examined during the last half of August; these fronds were old and dried. In general, the epiphytes, which are particularly sensitive to moisture conditions, underwent meiosis and produced spores in the wetter months. However, no fertile fronds of Microgramma squamulosa were located. For a few species, flushes of new growth were obvious. In late August and September, vegetative growth in Pityrogramma was pronounced. In September, new growth was abundant in WNephelea, Blechnum occidentale, and Adiantum pseudotinctum. Generally, those species showing growth flushes were non- reproductive. Active new growth was evident in Antigramma spp., Asplenium claussenii, Blechnum brasiliense, B. ensiforme, Cheilanthes concolor, Didymochlaena, Polypodium truncorum, and Pteris denticulata in October. Finally, all species examined in May, 1983 showed active new growth. This was, however, one of the rainiest Mays on record. Biological Interactions Herbivore damage was noted consistently on only four species of ferns. Diplazium ambiguum suffered what appeared to be heavy insect damage on most plants. On many of these the costules were completely stripped of pinnules. This coincides with observations by Smith (unpubl.) on Dip/azium spp. in Venezuela and southern Mexico. Fronds of Nephe/ea also were heavily eaten or otherwise damaged, at least in September and May when few pinnules are intact. Insect damage was noted occasionally on Antigramma brasiliensis and old fronds of Thelypteris torresiana. FERNS FROM EL TIROL, PARAGUAY 329 ACKNOWLEDGEMENTS We acknowledge the assistance of John Strother with photography. Minister of Agriculture Hernando Bertoni and Ing. Hilario Moreno of the Servicio Forestal Nacional (SFN) of Paraguay arranged for the plant collecting permits, and the SFN and the U.S. Peace Corps for logistical support in Ybycui and Cerro Cord National Parks. A.L. Gardner, D.B. Lellinger, and S.F. Smith commented on an early draft of the manuscript. Partial support of this research was provided by Grant No. 7699 from the Penrose Fund of the American Philosophical Society and a grant from the National Geographical Society to Foster, and by funds from the Department of Zoology and Museum of Vertebrate Zoology, University of California, Berkeley, for Foster. To all these individuals and institutions we extend our sincerest thanks. REFERENCES COMISION DE PARQUES NACIONALES DE PARAQUAY, 1973. Plan de manejo y desarrollo conceptual del parque nacional Ybycui. Doc. Tec. Trabajo No. 5 (Proyecto FAO-RLAT/TF/ 199). EVANS, A.M., 1963. New chromosome observations in the Polypodiaceae and Grammitidaceae. Caryologia 16: 671-677. EVANS, A.M., 1969. Interspecific relationships in the Po/ypodium pectinatum-plumula complex. _ Ann. Missouri Bot. Gard. 55: 193-293. FARINA SANCHEZ, T., 1973. The climate of Paraguay. Pp. 33-38 /n J.R. Gorham (ed.), Paraguay: Ecological Essays. Acad. Arts Sci. Americas. Miami, Florida. GASTONY, G.J., 1973. A revision of the fern genus Nephelea. Contr. Gray Herb. 203: 81-148. JARRETT, F.M., MANTON, I., and ROY, S.K., 1968. Cytological and taxonomic notes on a small collection of living ferns from Galapagos. Kew Bull. 22: 475-480. KURITA, S., 1971. Chromosome study of four species of leptosporangiate ferns. Ann. Rep. Foreign Students’ Coll. Chiba Univ. 3: 47-52. MITUI, K., 1968. Chromosomes and speciation in ferns. Sci. Rep. Tokyo Kyoiku Daigaku, B. 13: 285-333. SMITH, A.R., 1971. Systematics of the Neotropical species of The/ypteris section Cyclosorus. Univ. Calif. Publ. Bot. 59: 1-143. SMITH, A.R. and MICKEL, J.T., 1977. Chromosome counts for Mexican ferns. Brittonia 29: 391- 398. TRYON, A.R., BAUTISTA, H.P., and DE SILVA ARAUJO, I., 1975. Chromosome studies of Brazilian ferns. Acta Amazonica 5: 35-43. WAGNER, W.H., JR., 1963. A biosystematic study of United States ferns. Preliminary abstract. Am. Fern J. 53: 1-16. WALKER, T., 1966. A cytotaxonomic survey of the pteridophytes of Jamaica. 7rans. Roy. Soc. Edinburgh 66: 169-237. WALKER, T., 1973. Additional cytotaxonomic notes on the pteridophytes of Jamaica. 7rans. Roy. Soc. Edinburgh 69: 109-135. 330 FERN GAZ. 12(6) 1984. A NEW OPHIOGLOSSUM FROM INDIA SHARDA KHANDELWAL School of Studies in Botany, Jiwaji University, Gwalior, India. H. K. GOSWAMI Department of Genetics, School of Biological Sciences, Bhopal University, Bhopal, India. ABSTRACT A new species of Ophioglossum, from India, O. eliminatum Khandelwal & Goswami, is described. INTRODUCTION This paper presents observations on a new Ophioglossum species with features hitherto unknown in the genus. The plants described here were first collected in 1973 from Kanharjhir (near Gwalior) and at Chandpata (Shivpuri). Further collections during the rainy season (July — September) have been very extensive, and morphological, anatomical, biochemical and chromosomal studies have already been published (Khandelwal, 1978; Khandelwal and Goswami, 1977; Khandelwal, Goswami and Chaterjee, 1980; Goswami and Khandelwal, 1980; Das, Khandelwal and Dubey, 1981), incorporating other species occurring in the close vicinity (viz. O. costatum R. Br., O. nudicaule L., O. /usitanicum L., O. gramineum Willd. andO. thermale Kumarov). DESCRIPTION Ophioglossum eliminatum Khandelwal & Goswami sp. nov. Holotypus. Kanharjhir, Gwalior, Sharda Khandelwal, 9 August 1973 (National Botanic Garden, Botanical Survey of India, Howrah). lsotypes: Forest Research Institute, Dehra Dun and Royal Botanic Gardens, Kew. Plantae atro-virides, 1.8-15.0cm altae; rhizoma subglobosum; surculus communis 0.5- 4.0cm longus; trophophyllum 0.5-3.5 x 0.3-2.5cm, ovatum, mucronatum, sine vitta mediana pallida, venatio non duplex et cum areolis longis in parte inferiore laminae sterilis. Surculus fertilis 1.0-8.0cm longus, spica O.3-2.2cm longa; sporae triletae, interdum biletae, aletae et monoletae, liberae vel conjunctae, 18.0 (34.5) 45.Oum in diam. Plants dark green in colour, 1.8-15.0cm high; rhizome subglobose; common stalk 0.5- 4.0cm long, trophophyll 0.5-3.5 x 0.3-2.5cm, ovate in shape, mucronate, without pale median band, venation not double and with long stretched areoles at the lower part of the sterile blade, tip of the trophophyll with a pointed end; fertile stalk 1.0-8.0cm long; spike 0.3-2.2cm long; spores trilete, bilete, monolete and alete in one andthe same sporangium, occasionally jointed, 18.0-45.Oum in diameter (average 34.5); exine scubrate and with or without any outgrowth; spore output in a sporangium 4190-6651. Meiosis shows 90 bivalents, 2n = 180. The name of this species derives from the suggestion that it has originated through chromosome eliminations (Goswami and Khandelwal, 1980). ACKNOWLEDGEMENTS We are very grateful to Dr N.C. Majumdar, Botanical Survey of India, Howrah, for the Latin diagnosis. Thanks are also due toA.C. Jermy, British Museum (Natural History), London, for suggestions. REFERENCES DAS, R.R., KHANDELWAL, S. and DUBEY, S.N. 1981. Activity of certain oxido-reductases in different species of Ophioglossum L. Bionature 1, 1 & 2: 43-44. GOSWAMI, H.K. and KHANDELWAL, S. 1980. Chromosomal elimination in natural population of Ophioglossum L. Cytologia, 45: 77-86. KHANDELWAL, S. 1978. Cytological and physiological studies of Ophioglossum L. Ph D thesis. Jiwaji University, Gwalior, India. KHANDELWAL, S. and GOSWAMI, H.K. 1977. Stomata in Ophioglossum palmatum L. Curr. Sci. 46, 2: 51-54. KHANDELWAL, S., GOSWAMI, H.K., and CHATERJEE, A.K. 1980. Nitrogen and oil contents in Ophioglossum L. J. Ind. Bot. Soc. 59, 1: 78-80. FERN GAZ. 12(6) 1984 331 ASPLENIUM OBOVATUM IN BRITTANY, NW FRANCE A. LABATUT Puypezac, Rosette, 24100 Bergerac, France R. PRELLI 32 rue des Salles-Plestan, 22400 Lamballe, France and J. SCHNELLER Institut fur systematische Botanik, Universitat Zurich, 107 Zollikerstrasse, 8008 Zurich, Switzerland ABSTRACT The Asp/enium obovatum group consists of two closely related species which may be difficult to distinguish on gross morphology. A. obovatum Viv. is a diploid Mediterranean species whereas A. bi//otii F.W. Schultz is a tetraploid with a much wider, more Atlantic, distribution. The occurrence of A. obovatum in Brittany has recently been verified by chromosome counts, a discovery which throws new light on the distribution of this species. Further field investigations, however, will be necessary to determine whether these finds represent isolated relict localities or an hitherto overlooked Atlantic extension of A. obovatum. INTRODUCTION The Asp/enium obovatum group consists of two closely related species, A. obovatum Viv. and A. billotii F.W. Schultz. The former is a diploid species confined to the Mediterranean region but occurring more commonly in the eastern part. A. bi/lotii, on the other hand, is a tetraploid species having a more Atlantic distribution; in France it occurs in scattered localities in Lorraine, Alsace, the Bassin Parisien region, and the Massif Central, as well as along the Atlantic seaboard of the Iberian peninsula andthe western Mediterranean coast (Sleep, 1966). O. obovatum andaA. billotii are of similar appearance and may at times be difficult to distinguish (Aymonin, 1974). The first diagnostic feature of the gross morphology which may be used to separate them is pinnule shape. InA. obovatum the pinnules are markedly rounded with very wide, obtuse, often inconspicuous teeth, whereas in A. billotii they are more acute, their teeth being more serrate and markedly distinct one from the other. The pinna apex is another good diagnostic character. It is rather wide androunded inA. obovatum, being formed by the decidedly confluent apical pinnules. InA. billotii it is more acuminate, nearly all the pinnules remaining distinct (Badré et al., 1981). The occurrence in France of a third similar species, namely A. foreziense Héribaud, does not make matters easier. This allotetraploid, derived by hybridisation between A. fontanum (L.) Bernh. and A. obovatum, bears a morphological resemblance to the two taxa of the A. obovatum group. As a result, identification is more difficult and the respective geographic distribution of the three species cannot be clearly defined (Jalas & Suominen, 1972). Finally, it should also be borne in mind that the nameA. obovatum was at one time frequently used to describe what is known today as A. bi//otii. DETAILS OF THE BRITTANY PLANT In July 1981, in crevices of coastal rocks at Pointe de Brezellec, NW of Cléden-Cap- Sizun, Finistére, Brittany, A. Labatut noticed a few plants of Asp/enium which on morphological grounds he suspected to be A. obovatum (Fig. 1). 332 FERN GAZETTE: VOLUME 12 PART 6 (1984) | b FIGURE 1.Asp/enium obovatum Viv. from Pointe de Brezellec, 1 May 1982; the two cytologically confirmed plants. 1a, Prelli 1322; 1b, Prelli 1323. Scale = 1cm (silhouette by A. Zuppiger). Spore measurements often provide a useful micro-morphological character for distinguishing between diploid and tetraploid species. Such measurements were accordingly made on this material by R. Prelli and they confirmed the original supposition; the spore length of our specimens proved to be 25-32y1m, compared with 32-38um for the tetraploid A. bi/fotii. In order to confirm our identification of A. obovatum a cytological study was undertaken by J. Schneller on material fixed in the field by R. Prellion 1 May 1982. Chromosome counts (made according to the technique of Manton, 1950) from spore mother cells at meiosis in young sporangia from two different plants (Prelli 1322 and 1323) gave the same result; both plants were diploid with c. 36 bivalents. The occurrence of the diploid species, Asp/enium obovatum, in Brittany has thus been clearly established. ASPLENIUM OBOVATUM IN BRITTANY 333 The habitat of A. obovatum at Pointe de Brezellec is reminiscent of its ecology on the Mediterranean coast. About 10 plants have been observed in shady crevices of well-sheltered siliceous coastal rocks facing due east. These rocky outcrops bear an open vegetation composed of Umbilicus rupestris, Armeria maritima, Silene maritima, Sedum anglicum, Daucus carota spp. gummifer, Cochlearia danica, Hedera helix, together with the lichens Ramalina sp. and Rocella sp. The grassy ledges carry a heathland type of vegetation with Euphorbia portlandica, Dactylis glomerata, Chrysanthemum leucanthemum, Jasione montana, Endymion non-scriptus, Scilla verna, Pteridium aquilinum, Violariviniana, Ulex gallii, Erica cinerea, Calluna vulgaris, Teucrium scorodonia, Ruscus aculeatus, Lonicera periclymenum, and Rubus sp. This is not the only locality. Specimens collected on 18 August 1891 by E. Gadeceau at Pointe de Kerharo (1km E of Pointe de Brezellec) were identified as A. obovatum by G. Denizot (P!). Although the plants are small, measurement of spores taken from these specimens confirms the identification. G. Aymonin, in a handwritten note, indicates that he was unable to locate these plants in the field in spite of repeated searches over a number of years. What is to be made of such Atlantic sites of a species up to now regarded as Mediterranean? They might be just a few relict localities, testifying to aprevious, much wider, distribution stretching over the oceanic regions. On the other hand, these sites might be an hitherto overlooked extension of the distribution of this species which, in that case, would have to be regarded as Mediterranean-Atlantic. Only further discovery of other localities after a thorough investigation of the Armorican coast will help resolve this question. REFERENCES AYMONIN, G., 1974. L’Asplenium obovatum Viv., plante de Corse. Observations sur la définition de l’espéce. Bull. Soc. Bot. Fr. 121, 95e sess. extr.: 61-65. BADRE, F., BOUDRIE, M., PRELLI, R., and SCHNELLER, J., 1981.Asplenium x sleepiae (A. billotii x A. foresiense) et Asplenium x bouharmontii (A. obovatum x A. onopteris), hybr. nov. Bull. Mus. natn. Hist. nat. Paris, 4e ser., 3, sect. B: 473-481. JALAS, J., and SUOMINEN, J., 1972. At/as florae europaeae. | Pteridophyta. Helsinki. MANTON, I., 1950. Problems of cytology and evolution in the Pteridophyta. Cambridge University Press. SLEEP, A., 1966. Some cytotaxonomic problems in the fern genera Asplenium and Polystichum. PhD thesis, Leeds. 334 SHORT NOTES SHORT NOTES SPIKE AND SPORANGIAL ABNORMALITIES IN OPH/IOGLOSSUM OF RAJASTHAN, INDIA Teratological conditions relating to spike, Sporangium and leaf are well known in Ophioglossaceae. A survey has provided some undescribed abnormalities in Ophioglossum which are reported here. A variety of branching patterns of spike were observed in natural populations of O. costatum R.Br. in SW Rajasthan, including a dichotomously branched spike, a twice branched spike with branches arising at different levels and oriented in different planes simulating the compound spike of Botrychium, and a branching of the spike with the lower part unbranched in aspecimen from SE Rajasthan, acondition reported earlier in O. pendulum L. (Bower, 1926), O. pa/matum L. and O. pedunculosum sensu Prantl (Mahabale, 1962) and recently in O. nudicaule L. (Goswami & Khandelwal, 1973). A similar type of branching of spike was observed in O. petio/atum Hook. from. Central Rajasthan, where a small protuberance of the spike was perpendicular to the main spike. In O. costatum from SE Rajasthan, dichotomously branched spike stalks bearing normal unbranched spikes were frequently observed. In O. reticulatum L. from the same locality the branching of stalk was conspicuously unequal. Sporangia as arule are paired and lateral in Ophioglossum. However, terminal unpaired sporangia were found in O. nudicaule from SE Rajasthan and O. petiolatum from Central Rajasthan as recorded earlier in O. costatum and O. gramineum Willd. (Goswami & Khandelwal, 1973). In the same spike of O. petio/atum containing the unpaired sporangium, three successive sporangia were also found fused forming a linear compound sporangium or a synangium on one side of the spike only, similar to the fusion reported in O. gramineum but involving only two sporangia (Goswami, 1978). A plant of O. petio/atum cultivated in the Botanical Garden, Government College, Ajmer (Central Rajasthan) has a leaf with an emarginate apex. Unlike Goswami & Khandelwal (1973), we could not locate any specimen with sporangia on the tropophyll in O. nudicaule or any other species of the genus. It is significant, however, that in plants with sporangial or spike abnormalities we could not observe any well marked difference in the spore types (normal, irregular, enucleate or deformed spores) in normal and/or abnormal spikes, nor was there any relationship between the twisting of spike and occurrence of enucleate spores in the twisted portion. Spike twisting is a regular feature in O. petio/atum in Rajasthan and was not observed in the other species. Twisting of the spike seems to be relatedto an enhanced period of humidity affording a prolonged period of development to the spike and does not indicate that a genetic mechanism leading to sterility is operative as suggested by Goswami & Khandelwal (1973). Grateful thanks are due to Principal N.P. Vadehra, and Dr P.L. Mital, Head, Department of Botany, Government College, Ajmer. REFERENCES BOWER, F.O. 1926. The ferns Vol. //. Camb. Univ. Press. GOSWAMI, H.K. 1978. A morphogenetic study of sporangia-partitioning complex. Acta Soc. Bot. Poloniae. 47: 307-315. GOSWAMI, H.K. & KHANDELWAL, S. 1973. Abnormalities in Ophioglossum. Brit. Fern Gaz. 10: 311-314. MAHABALE, T.S. 1962. Species of Ophiog/ossum in India. Their taxonomy and phylogeny. Bull. Bot. Surv. India. 4: 71-84. B.L. YADAV and T.N. BHARDWAJA, Pter(dophyte Biology Lab., Department of Botany, Government College, Ajmer — 305001, India. FERN GAZ. 12(6) 1984 335 A NEW SPECIES OF M/CROLEPIA FROM SOUTH INDIA B.K. NAYAR and P.V. MADHUSOODANAN Botany Department, Calicut University, 673 635, Kerala, India. ABSTRACT A new species of Microlepia, M. manohara, similar to M. spe/uncae, is described from Wynad, South India. INTRODUCTION The Wynad plateau (South India) is c. 700m above sea level and juts into the Malabar plains, ending rather precipitously on the western side. Lying on the windward side of the Western Ghats, Wynad receives an average annual rainfall of c. 380cm, mostly between May and August. It is criss-crossed by streams and supports a dense tropical forest which until recently was nearly impenetrable and hence floristically little known. Destruction of the forests for establishing tea, coffee and cardamom plantations has now opened up most regions for botanical exploration. While studying the fern flora of Wynad we came across a new species of Microlepia which is described here. Microlepia C. Presl. (Dennstaedtiaceae) contains 47 species, 10 of which occur in India (Beddome, 1892; Nayar & Kaur, 1963, 1964). Microlepia manohara* Nayar & Madhusoodanan sp. nov. Species Microlepia speluncae similis; sed magnitudine grandior, folia magna ovato- deltata cum Sporangiis et sporis minoribus. Rhizoma ramosum, c. 2cm crassum, pilis uniseriatis lanea marcidaque ornatum. Folium erectum, c 3m longum. Lamina foliorum late ovato-deltata, c. 170x150cm, quadripinnata. Omnes divisiones acroscopicae basales ampliores quam divisiones ceterae eiusdem ordinis. Pinnae primariae oblongo-lanceolatae, 75x25cm cum apicibus longe acuminatis. Pinnae secundariae late lanceolatae, 12x3cm cum apicibus longe acuminatis. Pinnae tertiariae acroscopicae basales pinnatim divisae; ceterae pinnatifidae, 2.0 x 0.75cm, rhombicae, cum apicibus obtusis. Pinnae quaternariae rhombicae, 6 x 3mm, cum marginibus serratis et apicibus obtusis. Rachis etomnes rami et primae venae laminae pilis uniseriatis glandulosis etiam pilis uniseriatis acicularibus praeditae. Lamina pilis parvis acicularibus praeter illas duas species pinnarum praedita. Sporangia cum paraphysibus mixta. Annulus sporangii 20-26 cellulis longus. Sporae triletae, leves, 20x30pm, in forma trianguli cum lateribus concavis. Holotypus.: B.K. Nayar & P.V. Madhusoodanan CU 7265, Chembra Peak, Wynad, India Meridionalis, 1000m. Feb. 1976, in Central National Herbarium, Calcutta. Isotype in the herbarium of Calicut University, Kerala. Paratypes: B.K. Nayar & P.V. Madhusoodanan CU 7280, Chembra Peak; P.V. Madhusoodanan CU 21306, Chembra Peak, both in Calicut University Herbarium. Rhizome long-creeping, branched. Fronds to 3m tall; stipe c. 125cm x more than 10mm at base, erect, glabrous and shining when mature, purplish-brown basally, with a shallow broad adaxial median groove. Young stipes covered with hairs which are shed except in adaxial groove when mature. Lamina c. 175 x 150cm, broadly ovate-deltate in outline, broadest slightly above base, quadripinnate. All lamina divisions oblique at base, acroscopic side markedly longer than basiscopic; acroscopic basal division of each series conspicuously larger than others, resembling divisions of previous order. Primary pinnae up to 75 x 25cm, oblong-lanceolate, with c. 22 pairs of 2° pinnae, acuminate at apex. Secondary pinnae c. 12 x 3cm, broadly lanceolate, distinctly stalked, long-caudate at apex. Basal acroscopic 3° pinnae distinctly pinnate, other 3° pinnae deeply pinnatifid, c. 20 x 7mm, rhombic, obtuse at apex, broadly rounded at acroscopic base. Ultimate lobes (Fig. 1) ovate-rhombic, crenate at margin and obtuse at apex. Quaternary pinnae c. 6 x 3mm, serrate at margin and with pinnate venation. Rachis, its branches and midribs of ultimate pinnae straight, adaxially grooved, groove not continuous on branches of different orders. Lamina with hairs throughout, most of which are shed from the main rachis during maturity. Sori submarginal at apex of veinlets (mostly on acroscopic basal veinlets of ultimate lobes). Indusium broader than long (0.5 x 0.8mm), with entire glabrous margin. *The specific epithet ‘manohara’ is derived from Sanskrit and means ‘elegant’. 336 FERN GAZETTE: VOLUME 12 PART 6 (1984) FIGURE 1. Microlepia manohara, tertiary pinna showing sori and venation (scale = 3mm). Sporangia mixed with uniseriate glandular paraphyses. Annulus of (20-)22(- 26) indurated cells. Spores 20 x 32um (Px E), trilete-tetrahedral, with triangular amb, broadly rounded corners and concave sides, flat proximal face and tenui-marginate laesura, with a few large yellowish-brown oil globules and many plastids when fresh. Microlepia manohara is a \arge and elegant fern forming clumps near margins of forests. Except for its markedly larger size, it resembles MVM. speluncae, which is widespread in South India (Sledge, 1956). The broadly deltate lamina of M. manohara is nearly three times as broad with the primary and secondary pinnae more than twice the size of those of V/. spe/uncae, while the ultimate segments are much smaller. Asin M. haflangensis (Nayar & Kaur, 1964) the margin of the indusium is naked. The annulus of the sporangium consists of 20-26 cells in contrast to 15-20 cells in M. speluncae. Despite the markedly larger size of the plant, the spores of VM. manohara are the smallest in the genus (Nayar & Devi, 1968) being only 20x 32um in contrast to those of MV. spe/uncae (28 x 34um) and M. hance: (25 x 40um). The exine is smooth in M. manohara whereas it is granulose in the other species mentioned. ACKNOWLEDGEMENTS It is a pleasure for us to thank Mgsr Rev. Fr Thomas Moothedan for correcting the Latin diagnosis. REFERENCES BEDDOME, R.H. 1892. Handbook to the ferns of British India, Ceylon and Malay Peninsula (with supplement). Thacker, Spink & Co., Calcutta. NAYAR, B.K., and DEVI, S. 1968. Spore morphology the the Pteridaceae — II: the Dicksonioid, Dennsta 2dtioid and Lindsaeoid ferns. Grana Palyn. 7: 185-203. NAYAR, B.K., and KAUR, S. 1963. Ferns of India — VIII: Micro/epia Presl. Bull. nat/ bot. Gdns 79: 1-25. NAYAR, B.K., and KAUR, S. 1964. Ferns of India — XII: some new taxa of Indian ferns. Bu//. nat/ bot. Gdns 94: 1-15. SLEDGE, W.A. 1956. Microlepia speluncae (L.) Moore, M. trapeziformis (Roxb.) Kuhn and ™. firma Mett. in Kuhn. Kew Bull. 3: 523-531. FERN GAZ. 12(6) 1984 337 ANOTHER INTERGENERIC HYBRID IN GRAMMITIDACEAE: CTENOPTERIS LONGICEPS x GRAMMITIS SUMATRANA BARBARA S. PARRIS Botany School, Downing Street, Cambridge, CB2 3EA, England.* ABSTRACT The naturally occurring hybrid between Ctenopteris longiceps (Rosenst.) Copel. and Grammitis sumatrana (Baker) Copel. is reported for the first time. It is found in Papua New Guinea. INTRODUCTION The family Grammitidaceae is represented in New Guinea by approximately 150 species. The largest genera are Grammitis and Ctenopteris with 64 and c. 50 species respectively. As with many other groups of tropical plants, a large number of species belonging to the same family may be found in one area, theoretically providing opportunities for hybridisation, but hybrids even within a genus appear to be extremely uncommon. It is of great interest, therefore, to find plants that are intermediate between two species from different genera and which are apparently of hybrid origin. In 1980 | collected ferns on Mt Hagen (Western Highlands Province) and Mt Giluwe (Southern Highlands Province) in Papua New Guinea. Ctenopteris longiceps (Rosenst.) Copel. and Grammitis sumatrana (Baker) Copel. occur on both mountains, and plants intermediate between the two species were occasionally found in both places. | was unable to find any intermediates during extensive collecting in other years (Mt Hagen in 1971 and Mt Giluwe in 1977 and 1981) although both species were found during these expeditions. DESCRIPTION Species of Grammitis have simple + entire fronds; those of Ctenopteris usually have fronds pinnately divided to pinnate, although some may be bipinnate. Grammitis sumatrana has slehder fronds entire to crenately lobed to 5- 12mm from the midvein in the centre of the frond, while Ctenopteris longiceps has broader fronds which are pinnate or very deeply pinnately divided to c. 0.5mm from the midvein in the centre of the frond. Plants intermediate between these two species have fronds of intermediate proportion, rather irregularly pinnately divided to 2-4mm from the midvein in the centre of the frond. Figure 1 shows the two species and the intermediate. All three taxa have the following characters in common: short to long-creeping rhizome, with + lanceolate + acute pale to medium red-brown glabrous scales, rather dense + patent red-brown simple eglandular stipe hairs c. 0.1-0.3mm, scattered to occasional + patent medium to dark red-brown simple eglandular hairs c. 0.2-0.7mm on margin and midvein or rachis above and below, the lateral veins forked several times, and the sori + circular in outline. The microscopic characters given in Table 1 show no significant differences in sporangial size, length and number of sporangial setae nor in spore size between the two species. The spores of both species are + globose and chlorophyllous, while those of the intermediate plants are + globose, usually pale and without chlorophyll, or occasionally blackish. *Present address: Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE, England. 338 FERN GAZETTE: VOLUME 12 PART 6 (1984) TABLE 1. Microscopic characters of Ctenopteris longiceps, Grammitis sumatrana and intermediate plants, based upon material from Mt Hagen and Mt Giluwe (10 measure- ments of each character from each plant: range in brackets, other figures represent one standard deviation each side of the mean). C. longiceps intermediates G. sumatrana (5 plants) (3 plants) (7 plants) sporangia (170-)203 - 235(-250) um (170-)200 - 252(-260) um (180-)207 - 257(-310) um setae 1-5, medium to dark red- 1-4, medium to dark red- 1-6, medium to dark red- brown, 100-180 yum brown, 100-170 um brown, 110-200 um spores (29-)32 - 40(-45) yum (23-)26 - 31(-33) um (28-)30 - 40(-44) yim DISTRIBUTION AND ECOLOGY C. longiceps is endemic to Papua New Guinea and has a rather limited distribution in the Western, Southern and Eastern Highlands and Morobe provinces between 2300 and 3000m alt. G. sumatrana occurs elsewhere in Malesia (Sumatra, Borneo, Java and the Philippines), and is one of the commonest and most wide-spread species of the family in New Guinea, occurring between 950 and 3350m alt. On Mts Giluwe and Hagen both species are rather common between 2300 and 3000m alt. in midmontane forest which is often dominated by Nothofagus. C. longiceps is usually a low epiphyte from + ground level to c. 3m up on tree trunks of 3cm to more than 1m in diameter in primary montane forest, but occasionally grows on the downstream side of large mossy boulders in the beds of small mountain streams in forest. G. sumatrana is usually epiphytic from + ground level to at least 8m up on tree- ferns, lianes and tree trunks of 1¢cm to more than 1m in diameter in primary montane forest, young secondary forest and rather open recently logged forest, but occasionally grows on the down-stream side of large mossy boulders in streams in exactly the same habitat as C. /Jongiceps and sometimes with it. The intermediate plants were found between 2300 and 2820m alt. as epiphytes from O.7to 2m up on lianes and tree trunks of 1cm to 20cm in diameter in primary montane forest. In each of the three localities where the intermediate was found it was as a solitary plant, with neither species present in the immediate vicinity, but with both species present in the same area. No intermediate plants were found in areas where only one species occurred. DISCUSSION It is obvious from the above descriptions that not only are the intermediate plants probably hybrids between C. /ongiceps and G. sumatrana (although artificial synthesis of this hybrid and study of its chromosome behaviour at meiosis would provide useful confirmation that hybridisation is possible), but also that these two species are e closely related in spite of their superficially very different appearance. The genus Grammitis contains many species groups each of which contains a number of closely related species, but the relationships between the species groups are sometimes unclear. Ctenopteris as presently understood also contains a diversity of species groups, some of which appear to be more closely allied to species groups currently treated in other genera than to other groups within Ctenopteris, eg: C. longiceps with the group of G. sumatrana, C. heterophylla (Labill.) Tindale with the group of G. billardieri Willd. (Parris, 1977) and C. curtisii (Baker) Copel. (syn. Polypodium decrescens Christ) with the group of G. fasciata Blume (Copeland, 1952) which should probably also include X/phopteris conjunctisora (Baker) Copel. (Parris, 339 CTENOPTERIS LONGICEPS x GRAMMITIS SUMATRANA sdaaibuo] sisajdouajy) 1614 BUBIJEUINS SIJIWWeID x Sdazibuo sisaj]douajy 91\1Ua0 BUBIIEUINS SIIWWweslD 149] ‘L AYNDIS 340 FERN GAZETTE: VOLUME 12 PART 6 (1984) 1983). C. heterophylla and G. billardieri hybridise, as do C. curtisii and G. fasciata. G. sumatrana, together with the only other member of its species group, G. torricelliana (Brause) Parris of the Philippines and New Guinea, has a rather isolated taxonomic position with the genus Grammitis because of the lateral veins being several times forked and bearing several rows of sori on each side of the midvein in well-developed fronds. C. Jongiceps appears at first sight to be a typical member of Ctenopteris because of its frond dissection, but has no near relatives in that genus; apart from the size of the frond and the depth of the lobes it is nearly identical with G. sumatrana. The smaller geographical and ecological range of C. /Jongiceps compared to G. sumatrana may indicate that the former has originated from the species group of the latter in Papua New Guinea. Although both grow in primary montane forest there appears to be some ecological separation between them as they very rarely actually grow together (only on stream boulders). Reproductive isolation between the two species may be indicated by the absence of normal spores in the hybrid (although it would be of interest to attempt the germination and growth of both the pale and the dark spores of the hybrid to test whether they are viable in spite of their appearance). Alternatively G. sumatrana may be regarded as a stabilised hybrid between an unknown Grammitis species or G. torricelliana and C. longiceps; the artificial synthesis of a hybrid between these two species would be interesting. There is no evidence at present to indicate which possibility is more likely. Some of the problems in supra-specific classification within the Grammitidaceae are outlined above. Both the variation and the relationships of many species in the family are poorly known and it is premature to tamper with the existing generic classification until these problems are resolved. It must be borne in mind, however, that Ctenopteris as generally understood is an artificial assemblage of Grammitids which have a certain type of frond dissection and lack the distinctive characters of Adenophorus and Prosaptia; Grammitis may also be artificial in part, and Xiphopteris is certainly so. | am currently working on a more natural circumscription of the Grammitid genera. ACKNOWLEDGEMENTS | am indebted to the trustees of the Percy Sladen Memorial Fund and the Eileen and Phyllis Gibbs Travelling Fellowship, Newnham College, Cambridge for the funds to enable me to carry out this research. | am also grateful to the staff of the Office of Forests, Western Highlands Province for providing transport to Mt Hagen, to Mr & Mrs D.J. King for hospitality in Mt Hagen town, to Mr T. Payne for hospitality in Mendi and transport to Mt Giluwe, to Mr R.J. Johns for transport in the Highlands Provinces, to Beechwood Company for access along their logging roads on Mt Giluwe, to the Director of the herbarium of Division of Botany, Department of Forests at Lae for the use of drying facilities and permission to examine material, to Dr J.P. Croxall for assistance in the field and to Miss J.A. Hulyer and Mr C.R. Chalk for taking the photograph. REFERENCES COPELAND, E.B. 1952. Grammitis. Philippine Journal of Science 80: 214. PARRIS, B.S. 1977. A naturally occurring intergeneric hybrid in Grammitidaceae (Filicales): Ctenopteris heterophylla x Grammitis billardieri. New Zealand Journal of Botany 15: 597-599. PARRIS, B.S. 1983. A taxonomic revision of the genus Grammitis Swartz (Grammitidaceae: Filicales) in New Guinea. B/umea 29: 13-222. FERN GAZ. 12(6) 1984 341 HYBRIDIZATION IN ELAPHOGLOSSUM IN THE MASCARENE ISLANDS D.H. LORENCE Instituto de Biologia, Apartado postal 70-233, U.N.A.M., Mexico 20, D. F., Mexico. ABSTRACT Five new interspecific hybrids in the fern genus E/aphog/ossum (Lomariopsidaceae) from the Mascarene Islands of Mauritius and Réunion are described: F. x adulterinum D. Lorence, E. x cadetii D. Lorence, £. x heterophlebium D. Lorence, E. = revaughanii D. Lorence and E. x setaceum D. Lorence. Habitat preferences and probable parentage of the hybrids is discussed together with the possible role of hybridization in the evolution of the genus. INTRODUCTION The importance of the role played by hybridization in the evolution of angiosperms has been well documented (Raven, 1980; Stebbins, 1969, 1977), and hybridization is thought to be particularly significant in the evolution of island floras (Carlquist, 1965, 1974; Gillett, 1972; Raven, 1972). Likewise, hybridization among ferns appears to bea common phenomenon (see Knobloch, 1973 and 1976 for compilations and an extensive bibliography) and the evolution of discrete taxa w/a hybridization, at times involving a reticulate pattern of phylogeny, has been well documented in temperate ferns (Manton, 1950; Wagner, 1954, 1969, 1973; S. Walker, 1961). However, in spite of some excellent work done in the tropics (eg: Manton and Sledge, 1954; Wagner and Wagner, 1975; Wagner, Wagner and Gomez, 1978; T. Walker, 1958, 1973), relatively less attention has been focused on interspecific hybridization in tropical ferns, particularly on islands, with only a single previous report involving hybridization in Elaphoglossum (Lomariopsidaceae) (Knobloch, 1976), a large and complex genus of c. 400 species distributed in tropical montane forests of both hemispheres. METHODS AND CRITERIA EMPLOYED While undertaking a systematic revision of the species of Elaphoglossum occurring in the Mascarene Islands (SW Indian Ocean) for ‘Flore des Mascareignes”’ (Lorence, in press), | collected a number of specimens in Mauritius differing morphologically from well known taxa. Similar irregularities were observed in a collection from neighbouring Réunion Island. As pointed out by Wagner (1962), morphological and structural irregularities are usually strong indicators of hybridism in pteridophytes. It is further stated (Wagner and Lim Chen, 1965) that a fern hybrid may be detected by: (1) possession of morphological characters intermediate between two species; (2) deficiencies in the reproductive apparatus, ie: a large percentage of underdeveloped or aborted spores or sporangia resulting from abnormal chromosomal pairing behaviour at meiosis (may also be observed microscopically in meiotic squash preparations); (3) apomixis, which often occurs in the case of sterile triploid hybrids; (4) artificial synthesis of the hybrids by crossing parental gametophytes (see Lovis, 1968 for the experimental procedures involved). Of the five hybrid combinations described here, which most likely represent F, segregates, four display abnormal sporogenesis in addition to being morphologically intermediate. The fifth hybrid, E. x setaceum, was not fertile but the fact that it was growing intimately associated with its putative parents, in addition to being morphologically intermediate, leaves little doubt as to its probable parentage. Elaphoglossum gametophytes are slow-growing and long-lived (Stokey and Atkinson, 1957), as are the sporophytes which may take several years to achieve fertility and therefore no attempt was made to recreate the hybrids described here. A study of the gametophytes in culture to detect the presence of apomixis was not undertaken for the same reason. 342 FERN GAZETTE: VOLUME 12 PART 6 (1984) Fern scales provide excellent diagnostic features in groups where most taxa exhibit strikingly constant and distinctive types of palea on their rhizome, stipe and lamina. The scales of Elaphoglossum species have long played an important role in their taxonomy and many keys rely to a large extent on scale morphology; scales also appear to be an extremely reliable indicator of hybridism, at least among Mascarene members of the genus. Venation in the vast majority of E/aphoglossum species is dichotomous-pinnate, although inter-vein reticulations occur in a few members (eg: E. reticu/atum (Kaulf.) Gaud. from the Hawaiian Islands). Among the dichotomous-pinnate members vein endings in most species are free and often thickened near the margin, sometimes into hydathodes. A number of species, however, possess veins which reunite into an intra- marginal commissure or series of arcs around the periphery of the lamina. Species displaying this latter character were formerly placed in the genera Aconiopteris and Olfersia. Some Mascarene Island species displaying this feature are £. macropodium (Fée) Moore, £. richardii (Bory) Christ, —. sieberi (Hook & Grev.) Moore and E&. tomentosum (Bory ex Willd.) Christ. Apart from two species to be discussed later /(E. angulatum (BI.) Moore and E. /anatum (Bojer ex Baker) D. Lorence), | found venation type to be extremely constant for all species and also a feature useful in detecting hybrids. Other useful morphological characters employed in this investigation were shape of the lamina and characters of the rhizome, including spacing of the fronds and number of rows in which they are produced. Finally, spore size and percentage of aborted spores were employed as criteria for detecting reproductive irregularities. DESCRIPTION OF NEW HYBRIDS Elaphoglossum x revaughanii* D. Lorence, hybr. nov. (E. sieberi (Hook. & Grev.) Moore x £. /epervanchii (Fée) Moore). Hybrida E/laphoglosso sieberi similis sed differt: habitatio terrestris (haud numquam epiphytica) in clones densos vegetatos aggregata; frondes steriles c. duplo quam eis E. sieberi grandiores apice breviter acuminatae; venis laminis sterilis pro parte liberis, pro parte in arcus intramarginales conjunctis; paleae rhizomatis sparsiores et multo breviores angusto-ovatae ad subulatae, 3.0-6.0x 0.5-0.8mm, nigrae nitentes opacae; sporae fuscae irregulares, c. 65% abortae. Fig. 1. Type: Mauritius, Mare Longue Plateau, Lorence 14.3 sub MAU 15878 (holotype, MAU; isotypes, K, MAU, MEXU, MO, P, REU). Other specimens examined: Mauritius, Mare Longue Plateau, Lorence 77.7 (MAU, MEXU, MO), Lorence 8.6 sub MAU 155317 (K, MAU, MO, P, REU), Lorence 1176 (MO); Pétrin Nature Reserve, Lorence 1497 (MAU, MO). Rhizome short-creeping, dorsiventral, occasionally branching, 5-6mm diam., densely paleaceous at apex, fronds in 2 ranks, 6-15mm distant; rhizome scales narrowly ovate to subulate, 3-6 x 0.5-0.8mm, black, shiny, rigid, apex long acute, base truncate to cordate or auriculate, paler, with short glandular cilia, margins entire or with a few long, glandular cilia, cells small, seriate, rectangular, black and opaque; phyllobase scaly, grayish to yellowish-brown, 15-20mm long x 3.0-3.5mm diam., slightly swollen, abcission distinct. Sterile frond (300-) 440-530mm long; stipe 140-160 (-190) x 2.0-2.6mm, canaliculate, stramineate, scaly when young, scales dark brown to black, ovate to subulate, 0.5-4.0 x 0.2-0.5mm, base cordate to auriculate, fringed by short, glandular cilia, apex sinuate, margins sinuate with scattered glandular cilia, cells dark brown; lamina narrowly elliptic to very narrowly elliptic, (190-)300-350 x (43-) 55-70mm, apex acute to acuminate, base acutely cuneate, decurrent along top 1/4-1/3 of stipe then continuing as dark lines, texture subcoriaceous, margin revolute, rounded, veins visible, slightly raised, forking 1-2 times, making a 75-80° angle with costa, in part free and thickened, in part reuniting into intramarginal arcs, costa canaliculate adaxially, bearing abaxially scattered brown scales 0.5mm diam., with glandular cilia. Fertile frond (300-) 350-420mm long; stipe (150-) 1 70- 220mm long; lamina 150-230x15-26mm, very narrowly elliptic or lorate, apex acute to acuminate, tip rounded, base acutely cuneate, adaxial surface pallid with minute, *Named in honour of Dr R.E. Vaughan, former curator of the Mauritius Herbarium, who called my attention to the hybrid and whose numerous contributions to the botany of Mauritius include initiation of the ‘‘Flore des Mascareignes”’ project. HYBRIDIZATION IN MASCARENE ELAPHOGLOSSUM 343 scattered scales, veins not visible, texture thin, flexible. Spores: 4-5% very large (75-88 x 50-60um, excluding perine); 65-67% aborted; 29-30% normal (40-50 x 30-35um, excluding perine), perine bearing large, ruffled crests 10-15um high. Distribution: Mauritius, known only from Mare Longue Plateau (630m elevation, precipitation 3400mm per year) and Pétrin Nature Reserve (660m elevation, precipitation 4000m per year). Elaphoglossum x revaughanii was first collected from a single large and subcircular clone c. 13 x 17m on Mare Longue Plateau growing in the light shade of low, scrubby vegetation over lava with skeletal lateritic soil. The hybrid apparently reproduces vegetatively by rhizome branching, as young plants and sporelings were not observed in the clone or in the vicinity. Wagner (1963) states, ‘sterile hybrids of most other species of ferns are likewise capable of such vegetative reproduction, usually by rhizome “‘‘twinning” and dying off of the older stem parts’’. The area covered by this clone consists of a fairly dense network of rhizomes and mass of fibrous roots which have formed a thick layer of humus. This massive humus deposit, the clone’s large size, and the relatively slow growth rate of the hybrid suggest that it is quite old. Rhizomes of plants in cultivation were observed to elongate at the rate of c. 30mm per year. As the radius of the clone is c. 7.5m, it could conceivably be approximately 250 years old. Asecond much smaller clone was subsequently discovered growing in open Philippia heath vegetation at Pétrin Nature Reserve 2km distant. E. x revaughani superficially resembles £. s/eberi, for which it was first mistaken. Closer examination revealed its morphology to be intermediate between E£. sieberi and E. lepervanchii, although the hybrid is more robust than either parent. Both putative parents occur in this same biotope, usually as depauperate individuals growing at the bases of stunted trees and shrubs because it is a marginal, suboptimal habitat for these shade-loving, wet forest epiphytes. —. x revaughanii, however, behaves like a terrestrial heliophilous or hemisciophilous species and is obviously much more successful than either parent in this particular ecological setting of low scrub vegetation. The hybrid’s most distinctive morphological irregularities are apparent in the venation and scales. £. x revaughanii has veins which are partly free (as in E. /eper- vanchii) and partly reunited into an intramarginal commissure (as in £. s/eber/). Scales on the lower portion of the stipe and phyllobase of the hybrid are intermediate between those of both parents. Most spores are completely aborted (65-67%), about a third are relatively normal (c. 30%), and a few are exceptionally large (4-5%). All spores are strikingly dark in colour with highly cristate perines. In addition, the internal sporangial walls are lined with a dark coloured deposit, findings that agree with those reported by Wagner and Lim Chen (1965) for Dryopteris hybrids. Furthermore, the number of spores per sporangium in £. x revaughanii is extremely variable (16, 17, 18, 32 or 48), probably as a result of pairing abnormalities at meiosis. These phenomona are suggestive of observations reported by Morzenti (1962) and Hickok and Klekowski (1973) regarding non-reduction meiosis in fern hybrids. Further research on spore viability and chromosomal behaviour in £. x revaughanii is obviously desirable. Elaphoglossum < heterophlebium* D. Lorence, hybr. nov. (E. heterolepis (Fée) Moore x E. tomentosum (Bory ex Willd.) Christ). Hybrida Elaphog/osso heterolepe similis, sed differt: paleae rhizomatis castaneae margine sparse et breviter dentato ad ciliato; paleae stipitis magis pallido-brunneae marginibus *The specific epithet refers to the hybrid’s mixed venation, as the veins from the middle portion of the lamina unite into an intramarginal commissure (as in £. tomentosum) while those at the extremities of the lamina are free with thickened tips (as in £. hetero/epis). All examples of E. x heterophlebium that | encountered were robust. 344 FERN GAZETTE: VOLUME 12 PART 6 (1984) tHE MAUOTTUS BERR RIM Proxa Frea nf thn Mascarene tlands Se ya DEER OL ELD n> EEA Le By ene EE SEARIR ET TS AE ER TE Scnpane cise Pere Ne Bese nee Demi Shae ee Sek: Rion ge RUS en DO RORSR RR a: BR Beis BEG BS ae Ee BIRR TN BIEN SSRIS me Be resexee ox 8) SORES: Berens Pas SS Ri Sa ectas nc BOK Ree: aS Fee me Beka tote Skee Rees Se, peat % FOG AUST a = Hora af: Maanitus Rg SEO Sekine BS ESAT GS Roar aa SeRAoye kia. Fant Seqguegaseanen So eee ee KERRY NON | ON RESTS ew E Kea ne DASE Mens 9& Rader tele ( Genie we. meee Sea Nae ah Bee Moe Cy: SQ Se: Mo ARNe WK yee Cle eee: awh Aeon Sane Coney ARES CHEARAE SPCR Ce as SeaR Nes, ERPS BRAS See GS Ae Sooke eatin: Ry CORY SENS OT Rake Mextne a be corn BERNER wok. goat SoA UNIS SHORES NS Re OD ee eee Afecmce Ve \ERS2 wives serene Epler ane My ye SES ERE THE: BALSTIRSS BERBARIDM hercan Of CHOKE BE ISEES ES Shane Pensa Cn ern cee aaa hii crete? Rigetntn w€ hye: ; ® Ehepes Tih BAKA: HERB Flota of Mauritius THE MAURTORS HEMAABI Rey {HE MAURTNUS RERRARIOM Bug eestertemn K £eXwcen in, 3 ® es Pablarree HYBRIDIZATION IN MASCARENE ELAPHOGLOSSUM 345 3 FIGURE 1. Holotype collection of Elaphoglossum * revaughanii from Mare Longue Plateau, Mauritius, Lorence 14.3 sub MAU 158178 (MAU). FIGURE 2. Holotype collection of Elaphoglossum = heterophlebium from Pétrin Nature Reserve, Mauritius, Lorence 14.2 sub MAU 15827 (MAU). FIGURE 3. Holotype collection of Elaphoglossum ~* cadetii from Plaine des Sables, Réunion, Cadet 2063 (REU). FIGURE 4. Holotype collection of Elaphoglossum x adulterinum from Cascade 500 Pieds (Cascade Alexandra), Mauritius, Lorence 874 sub MAU 16309 (MAU). FIGURE 5. Holotype collection of Elaphoglossum xX setaceum and _ its putative parents, all from Tamarind Falls Reservoir, Mauritius: a) &. hydridum var. hybridum, Lorence sub MAU 715823a (MAU); b) E xX setaceum, Lorence sub MAU 15823b (MAU); c) E. lanatum, Lorence sub MAU 15823c (MAU). scarioso-fimbriatis et centris castaneis; lamina sterilis apice acuta ad vix acuminata, superficiebus ambabus indumento homogeneo squamarum uniformium bubalinarum peltatarum orbicularium ad ovatarum praeditis; venae laminae sterilis apicibus incrassatis pro parte liberis ornatae, pro parte in arcus intramarginales conjunctae; sporae 52-85% abortae. Fig. 2. Type: Mauritius, Pétrin Nature Reserve, Lorence 14.2 sub MAU 158217 (holotype, MAU). Other specimens examined: Mauritius, Brise Fer Road, Lorence 16.2 sub MAU 15819 (MAU); Plaine Champagne, Lorence sub MAU 15099 (MAU), Lorence 996 sub MAU 16518 (MAU), Lorence 1544 (MAU, MO). Rhizome short-creeping, 5mm diam, occasionally branching, densely paleaceous, fronds caespitose, 2-ranked; rhizome scales thick, opaque, castaneous, narrowly ovate to subulate, 5-7 x 0.5-0.7mm, slightly falcate, apex acute, setaceous, base cordate, margins thin, bearing short sparse acicular teeth or rare glandular cilia; phyllobase dull brown, 10- 20mm long, unthickened, abcission irregular. Sterile frond 330-450mm long; stipe 80- 130x 1.5-2.0mm, stramineate, flattened adaxially, densely scaly when young; stipe scales narrowly ovate, 2-5 x 0.8-1.5mm, falcate, centres dark, castaneous, opaque, margins pale brown, scarious, fimbriate to ciliate-dentate, with acicular or glandular-tipped teeth, base peltate and obtuse, or basifixed and cordate to auriculate; lamina very narrowly elliptic to lanceolate, 225-320 x 26-34mm, base acutely cuneate, apex acute to slightly acuminate, costa raised and slightly caniculate adaxially, margin rounded, slightly revolute, texture chartaceous, veins making a 70-75° angle with costa, in part free with thickened tips, in part reuniting into intramarginal arcs; both laminar surfaces bearing a homogeneous covering of thin, appressed and matted scales, soon caducous; laminar scales buff coloured, hyaline, peltate, orbicular to ovate, 0.5-1.0 x 0.5mm, base and margins prolonged into numerous arachnoid cilia 0.1-0.3mm long with acicular or inflated tips, cells thin walled. Fertile fronds 325-360mm long; stipe 165-195x 1.2mm; lamina linear, 346 FFRN GAZETTE: VOLUME 12-PART 6 (1984) 160-165 x 10-15mm, adaxial laminar surface and abaxial surface of costa covered by a thin layer of scales. Normal spores 35-38 x 25-27um (excluding perine), perine cristate, crests 2-3um high, minutely sptnulose. In Lorence 14.2 52% of the spores were aborted, whereas 85% of the spores were aborted in Lorence sub MAU 15099. Distribution: endemic to Mauritius. All four gatherings of Elaphoglossum x heterophlebium were made on the central plateau of Mauritius (elevation 600-700m, precipitation 2400-4200mm annually) in the following evergreen formations: Philippia heath formation at Pétrin; low, stunted scrub over laterite at Plaine Champagne; open lower montane wet forest of Sideroxylon puberulum A.DC. at Pétrin; mature lower montane wet forest at Brise Fer in a light gap. In all cases the hybrids were growing in fairly close proximity to or in association with the parental species. Although both £. hetero/epis and E. tomentosum are generally epiphytic or only casually terrestrial and prefer moderate to dense shade, E. x heterophlebium almost always occurs terrestrially and prefers light shade or partial exposure to sunlight. As was the case for the previous hybrid, £. x heteroph/lebium is able to occupy a niche that the parental species are unable to exploit. Segregates of £. x heterophlebium are remarkably uniform morphologically and very nearly intermediate between both parents, although tending to resemble E. heterolepis somewhat more than £. tomentosum. Elaphoglossum ~ cadetii* D. Lorence, hybr. nov. (E. acrostichoides (Hook. & Grev.) Schelpe x £. coursi/ Tard). Hybrida Elaphoglosso coursii similis sed differt: paleae rhizomatis duplo longiores sed tenuiores, 2-4 x 0.5-1.0mm, marginibus subintegris vel sparsim ciliatis; frondes steriles breviores, 60-140mm longae, apice obtuse-acuminatae; sporae c. 90% abortae. Fig. 3. Type: Reunion, Plaine des Sables, Cadet 2063 (holotype, REU; isotype, P). Rhizome short-creeping, 2-3mm diam., densely paleaceous; rhizome scales shiny black, discolorous, ovate, 2-4 x 0.5-1.0mm, apex black, opaque, acute to acuminate, base pale brown, cordate to auriculate, margins bearing rare, glandular cilia, fronds in two dorsal ranks, 2-4mm distant; phyllobase 5.0-7.5mm long, swollen, dark brown to black, colour change and abcission distinct, bearing at base scales similar to those on rhizome. Sterile frond (60-)80-140mm long; stipe (22-)36-65 x 0.5-1.0mm, stramineate, adaxially canaliculate, bearing scattered dark brown, discolorous, sparsely ciliate, ovate scales 2-3x 0.5- 1.0mm; lamina narrowly elliptic, 40-100 x 7- 15mm, apex acute with rounded tip, base acutely cuneate, decurrent as low ridges along top half of stipe, margin thin, cartilaginous, revolute, costa slightly canaliculate adaxially, veins immersed, indistinct, forking once, making a 50-60° angle with costa, tips free and thickened, texture coriaceous, lamina glabrous adaxially, abaxially bearing scattered, minute, brown, substellate scales 0.5mm diam., with a few larger scales along abaxial surface of costa. Fertile frond 140-185mm long; stipe 75-140mm long, about three times longer than sterile stipe; lamina narrowly elliptic, 60-85 x 9-10mm, glabrous adaxially, apex obtuse, base acutely cuneate, sides slightly decurrent, costa stramineate, veins indistinct, texture thin, flexible. Distribution: known from Réunion Island (La Réunion). A collection from Mt Ankarotra, Madagascar (Humbert & Cours 17521, P) may possibly represent this hybrid, as both parents also occur in Madagascar. The type of Elaphoglossum * cadeti/ is from Plaine des Sables on the Piton de la Fournaise massif, Réunion’s active volcano, where it was found growing in lava fissures. The hybrid is morphologically intermediate between E. acrostichoides and E. coursii which both occur in the wet and cloud forest zones of Réunion. Although Cadet makes no mention of associated species of E/aphoglossum on the type specimen, E. acrostichoides occurs in the Piton de la Fournaise region (D. Lorence, pers. obs.). The Plaine des Sables (elevation 2300m, precipitation c. 3500mm per year) supports *Named in honour of its collector, Dr T. Cadet who has contributed eminently to the study of the flora of Réunion and that of the Mascarene Islands in general. HYBRIDIZATION IN MASCARENE ELAPHOGLOSSUM 347 scattered patches of high altitude heath vegetation and herbaceous plants over porous rocky and sandy substrates. No forest occurs in the area and the few epiphytes able to grow here are restricted to sheltered rock crevices. This appears tobe yet another case where the hybrid is able to occupy a habitat unfavourable to either parent. Scale morphology of E£. x cadeti/ is intermediate between that of E. acrostichoides and £. coursii. Venation is free in all three taxa. The hybrid’s somewhat irregular frond morphology and the extremely high percentage of aborted spores (about 90%) leave little doubt as to its probable hybrid origin. Hybridization in E/aphoglossum lanatum Elaphoglossum /anatum is unique in being the only Mascarene representative of the genus to grow exclusively on vertical faces of cliffs and stream banks composed of decaying lava, a fact noted by Bojer on the type collection at Kew (‘ad rivulorum ripas’’). Certain morphological features of this species are extremely variable, notably frond size and shape, the degree to which the veins are free or reunite into an intramarginal commissure, and percentage of aborted spores (Lorence, 1976). For example, some individuals possess linear laminas 250-300mm long with acute to acuminate apices, while others have narrowly obovate, obtuse laminas only 65mm long in fertile plants. Certain other species of Elaphoglossum are quite polymorphic and the range of variation encountered in laminar size and shape for F. Janatum certainly lies within the range of normal phenotypic variability. As previously mentioned, venation type is almost invariably constant in most taxa, being either free or united at the margin. However, in £. /Janatum intermediates and extremes of both conditions may occur, even on the same individual. A similar situation was found in plants of £. angu/atum from Réunion. In addition, both species produce a significant percentage of aborted spores, (4-40% in £. angu/atum and 15- 32% in E. Janatum), much higher than the 5-10% observed for most other species. These morphological and reproductive irregularities suggest that —&. angu/atum and E. lanatum may be of hybrid origin. Although of possible hybrid origin, E. /‘anatum is certainly a viable species that exploits a niche which the other members of the genus in Mauritius are unable to. In addition, it is able to reproduce sexually and appears to hybridize with two other species of the genus forming two hybrid combinations described below. Elaphoglossum * adulterinum D. Lorence, hybr. nov. (E. lanatum (Bojer ex Baker) D. Lorence x £. tomentosum (Bory ex Willd.) Christ). Hybrida Elaphoglosso tomentoso similis sed differt: rhizoma resiniferum; paleae rhizomatis in margine glanduloso-dentatae; stipites c. duplo longioribus; paleae laminae sterils peltatae vel basifixae, margo ciliis longis arachnoideis implicatis ferens; venae laminarum sterilium pro parte liberae ad apicem incrassatae sed pro parte majore in arcus intramarginales conjunctae; sporae c. 82% abortae. Fig. 4. Type: Mauritius, Valley of Cascade 500 Pieds (Cascade, Alexandra), Lorence 874 sub MAU 16309 (holotype, MAU). Other specimens examined: Mauritius, Valley of Cascade 500 Pieds (Cascade Alexandra), Lorence sub MAU 16307 (MAU, MO), Lorence 1609 (MO), Lorence 1670 (MO). Rhizome prostrate, short-creeping, rarely branching, resiniferous, 2-3mm diam., densely paleaceous apically, fronds in two ranks, 3-5mm distant; rhizome scales opaque, castaneous, rigid, narrowly ovate to subulate, 4-6 x 0.4-0.6mm, apex filiform, sinuate, base cordate to auriculate, entire or bearing clear, globose marginal glands, margin sparesely glandular dentate-ciliate; phyllobase dark brown, 6-8mm long, thickened, colour change abrupt, abcission distinct, bearing basally scales like those of rhizome. Sterile frond (270-)350-485mm long; stipe (80-)100-150x 1.0-1.5mm, stramineate, canaliculate adaxially, bearing scattered, appressed, brown, caducous, ovate scales 1-2 x 0.5mm, margins scarious, glandular-ciliate, these intermixed with smaller, matted scales: sterile lamina (190-)250-320 x (14-)18-25mm, very narrowly elliptic to linear, base acutely cuneate to decurrent, apex acute to acuminate, both surfaces of lamina covered with a homogenous layer of thin, matted + appressed, pale brown to buff coloured scales; laminar 348 FERN GAZETTE: VOLUME 12 PART 6 (1984) scales orbicular to ovate, 1.0-1.5x 1.0mm, peltate or basifixed, base rounded to cordate or Sagittate, base and margins bearing long, spreading arachnoid cilia, pedicel and centre darker brown, subtended by cluster of globose glands, cells hyaline, thin-walled, scales caducous adaxially, costa stramineate, rounded abaxially, canaliculate adaxially, veins visible, slightly raised, making a 60-65° angle with costa, simple or forking once, in part free with thickened tips, in part reuniting intramarginally, texture subcoriaceous. Fertile frond 200-250mm long; stipe 100-115x1.0mm; lamina linear, 110-140x7-10mm, base acutely cuneate, apex acute, adaxial surface with a dense layer of matted scales as in sterile frond, margin revolute, texture thin, flexible. Spores 82% aborted; normal spores 35-38 x 25-27um (excluding perine), perine cristate, crests low, 1.5-3.0um high, sinuate to finely erose-dentate. Distribution: endemic to Mauritius. Elaphoglossum * adulterinum is known only from the type locality at Cascade Alexandra in the Valley of Cascade 500 Pieds located at the southern extremity of the central plateau, altitude ca. 600m, precipitation 4800mm per year. Habitats of both E. lanatum, which grows on cliff faces, and E. tomentosum, epiphytic or epipetric, are adjacent along the margins of this particular valley, thus providing a unique opportunity for hybridization between the species. Lorence 874 was found growing on a mossy boulder with £. tomentosum atthe base of a cliff supporting large populations of £. /anatum; the other collections of E. = adulterinum were also growing in the vicinity. The hybrid resembles £. tomentosum in habitat prefence (epiphytic or epipetric), in its erect fronds (pendulous in £. /anatum) and in the shape of the lamina. Laminar scales of E. x adu/terinum are intermediate between the parental types. Rhizome scales, however, are glandular and resiniferous as in E. lanatum. Venation is also intermediate between that of the parents, being in part free, and in part united at the margin as in £. tomentosum. Both conditions occur in E. lanatum, as previously noted. The high percentage of aborted spores (c. 82%) is also suggestive of hybridity. Elaphoglossum + setaceum D. Lorence, hybr. nov. (E. hybridum (Bory) Brack. * E. Janatum (Bojer ex Baker) D. Lorence). Hybrida Elaphoglosso hybrido var. hybrido similis, sed differt: rhizoma gracile 3mm diametro; paleae rhizomatis breviter dentatae dentibus glandulosis; laminae frondium sterilium paleis minoribus pallidioribus in Ssuperficiebus ambabus laminae dispersis; paleae laminae margine soium basi involuto, basi paribus aliquot ciliorum elongatorum glandulosorum praedita; marginibus ciliis aut dentibus acicularibus ascendentibus brevibus praeditis. Fig. 5. Type: Mauritius, Tamarind Falls Reservoir, Lorence sub MAU 15823b (Holotype, MAU). Rhizome dorsiventral, short-creeping, 3mm _ diam., densely paleaceous, fronds caespitose, in 3ranks, 2mm distant; rhizome scales light castaneous, subulate to narrowly ovate, 3-4 x 0.5mm, base cordate, the apex filiform, margins bearing short teeth each terminated by a globose gland, cells rectangular-fusiform; phyllobase 2-4mm long, dark brown, scaly abscission distinct. Sterile frond 80-140mm long; stipe 30-35 x 0.5-0.7mm, yellowish-brown, adaxially grooved, bearing numerous simple capitate hairs and scales; stipe scales brown, squarrose, bristle-like, subulate 1.5-2.5 x 0.2mm, apex filiform, base cochleariform with involute margins bearing several long cilia and short glandular teeth or cilia, cells fusiform; lamina narrowly elliptic to narrowly ovate, (50-)90-105:x (10-)15- 20mm, apex acuminate, base acutely cuneate, decurrent as low ridges along stipe, costa pale yellow-brown, raised on both surfaces, veins slightly raised and visible, forking once, tips free and thickened, margin slightly revolute, rounded, ciliate with fringe of pale brown bristle-like scales; marginal scales 2-3 x 0.2-0.3mm, apex filiform, base cochleariform, margins involute basally and bearing several long, spreading glandular-tipped cilia and sparse acicular teeth; both surfaces of lamina villous with bristle-like scales intermixed with smaller flat, deltoid scales, each with a single pair of long, spreading basal cilia and scattered simple hairs. Distribution: endemic to Mauritius. HYBRIDIZATION IN MASCARENE ELAPHOGLOSSUM 349 A single sterile individual of Elaphoglossum x setaceum was found growing ina population of £. /anatum on a shady stream bank in secondary forest at Tamarind Falls Reservoir (altitude c. 500m, precipitation 2600mm) at the western edge of the central plateau of Mauritius. A vigorous fertile clone of &. hybridum var. hybridum was growing at the base of the tree directly overhead but no other members of the genus occurred in the vicinity. E. x setaceum possesses morphological characters which are almost exactly intermediate between those of the putative parents (Fig. 5), notably shape of the lamina and scale architecture. Furthermore, its rhizome bears three ranks of fronds, whereas E£. /anatum is a two-ranked species and £. hybridum var. hybridum has three to four ranks. Unfortunately, the only specimen is sterile, thus precluding an examination of the spores. The hybrid’s association with its congeners and its intermediate appearance, however, leave little doubt as to its putative parentage. DISCUSSION AND CONCLUSIONS Elaphoglossum is a large and taxonomically complex pan-tropical genus of over 400 species with its greatest concentration in the mountains of Central and South America. Seventeen species are reported from the Mascarenes (nine in Mauritius, 14 in Réunion), of which seven are endemic to the islands (three in Mauritius, two in Réunion, two shared) (Lorence, in press). The genus is in an active state of evolution in the Mascarenes and hybridization appears to play a significant role in speciation of the members. Although up to four or five E/aphog/ossum species may occur sympatrically in the montane wet and cloud forests of Mauritius and Réunion, hybrids are virtually absent here. This may be due to the fact that hybridization in stable habitats is adaptively disadvantageous (Lewontin and Birch, 1966). Indeed, all five Mascarene Elaphoglossum hybrids occur in either of two situations: (1) at the abrupt interface of two distinct habitats, thus bringing together species with differing ecological require- ments and greatly enhancing their chances for hybridization, (eg: £. x adu/terinum, E. = setaceum); (2) in habitats generally unfavourable to either parent (eg: £. x cadetii, E. x heterophlebium, E. x revaughanii). In the first instance hybrids must occupy a habitat not differing from that of one of the parental species. Intermediate hybrids would be adaptively disadvantageous here and the hybrid would be expected to persist only if its ecological requirements matched those of either parent. This appears to be the case for £. x adulterinum and E. x setaceum, each of which grows together with one parent. The remaining three hybrids occupy habitats apparently not greatly utilized by either parent. £. x revaughanii has been able to thrive in a habitat where both parents are uncommon and only marginally successful. Similarly, £. x heterophlebium was collected four times in more highly exposed and isolated habitats than those in which the parental species occur. This likewise appears to be the case for E. x cadetii, although more information is required regarding distribution of its putative parents. Finally, hybrid origin can be suspected for successful and widespread but often ecologically specialized species such as £. /anatum and E. angulatum which display certain morphological and reproductive irregularities. In conclusion, the findings presented here suggest that hybridization in the large genus Elaphoglossum may provide an evolutionary stimulus capable of generating novel genetic combinations enabling certain hybrid segregates to survive better in habitats unfavourable to either parent. A successful hybrid could thus maintain itself either by vegetative propagation or by apogamy. ACKNOWLEDGEMENTS | am indebted to Dr John Dwyer of the Missouri Botanical Garden for his valuable assistance in preparing the Latin diagnoses. Dr Alwyn Gentry and Dr Warren G. 350 FERN GAZETTE: VOLUME 12 PART 6 (1984) Wagner of the Missouri Botanical Garden, and Dr T.P. Ramamoorthy of the Universidad Nacional Autonoma de México provided many useful suggestions and critical comments onthe manuscript. Finally, |am gratefulto Messrs L.S. De Relan and J. Forget of the Mauritius Sugar Industry Research Institute for preparing the photographs. REFERENCES CARLQUIST, S. 1965. /s/and life. Natural History Press, New York. CARLQUIST, S. 1974. /sland biology. Columbia University Press, New York. GILLETT, G.W. 1972. The role of hybridization in the evolution of the Hawaiian flora. /n, D.H. Valentine (ed.), Taxonomy, Phytogeography, and Evolution, 205-219. Academic Press Inc., London. HICKOK, L.G. and KLEKOWSKI, E.J., JR., 1973. Abnormal reductional and nonreductional meiosis in Ceratopteris: alternatives to homozygosity and hybrid sterility in homosporous ferns. Amer. J. Bot. 60: 1010-1022. KNOBLOCH, I.W. 1973. The present status of hybridity among the pteridophytes. 7Ja/wania 18/1): 29-34. KNOBLOCH, |.W. 1976. Pteridophyte hybrids. Pub/. Mus. Michigan State Univ., Biol. Ser. 5(4): 277-352. LEWONTIN, R.C. and BIRCH, L.C. 1966. Hybridization as source of variation for adaptations to new environments. Evo/ution 20: 315-336. LORENCE, D. 1976. Notes on some Mascarene species of E/aphoglossum (Lomariopsidoideae sensu Holttum). Fern Gaz. 717. 199-205. LORENCE, D. Lomariopsidacées. /n, J. Bosser et al. (eds), Flore des Mascareignes. L. Carl Achille, Imprimeur du Gouvernment, Port Louis, Ile Maurice (in press). LOVIS, J.D. 1968. Fern hybridising at the University of Leeds. Brit. Fern Gaz. 10: 13-20. MANTON, |. 1950. Problems of cytology and evolution in the Pteridophyta. Cambridge University Press. MANTON, I. and SLEDGE, W.A. 1954. Observations on the cytology and taxonomy of the pteridophyte flora of Ceylon. Philos. Trans. Ser. B. 238: 127-185. MORZENTI, V.M. 1962. A first report on pseudomeiotic sporogenesis, a type of spore reproduction by which “‘sterile’’ ferns produce gametophytes. Amer. Fern J. 52: 69-78. RAVEN, P.H. 1972. Evolution and endemism in the New Zealand species of Epilobium. /n. D.H. Valentine (ed), Taxonomy, Phytogeography, and Evolution, 259-274. Academic Fiess Inc., London. RAVEN, P. 1980. Hybridization and the nature of species in higher plants. Canadian Botanical Association Bulletin, Suppl. vol. 13, 1: 3-10. STEBBINS, G.L. 1969. The significance of hybridization for plant taxonomy and evolution. Jaxon 18: 26-35. STEBBINS, G.L. 1977. Processes of Organic Evolution. Prentice Hall, Englewood Cliffs, New Jersey. STOKEY, A.G. and ATKINSON, L.R. 1957. The gametophyte of some American species of Elaphoglossum and Rhipidopteris. Phytomorphology 7: 275-292. WAGNER, W.H., JR. 1954. Reticulate evolution in the Appalachian Aspleniums. Evo/ution 8: 103-118. WAGNER, W.H., JR. 1962. Irregular morphological development in hybrid ferns. Phytomorphology 12. 87-100. WAGNER, W.H., JR. 1963. Biosystematics and taxonomic categories in lower vascular plants. Regnum Veg. 27: 63-71. WAGNER, W.H., JR. 1969. The role and taxonomic treatment of hybrids. BioScience 19: 785- 789. WAGNER, W.H., JR. 1973. Reticulation of Holly ferns (Po/ystichum) in the western United States and adjacent Canada. Amer. Fern J. 63: 99-115. WAGNER, W.H., JR., and LIM CHEN, K. 1965. Abortion of spores and sporangia as a tool in the detection of Dryopteris hybrids. Amer. Fern J. 55: 9-29. WAGNER, W.H., JR., and WAGNER, F.S. 1975. A hybrid polypody from the New World tropics. Brit. Fern Gaz. 17: 125-135. WAGNER, W.H., JR., WAGNER, F.S. and GOMEZ, P., L.D. 1978. The singular origin of a Central American fern, Pleuroderris michleriana. Biotropica 10(4): 254-264. WALKER, S. 1961. Cytotaxonomic studies in the Dryopteris spinulosa complex. Il. Amer. J. Bot. 48: 607-614. WALKER, T.G. 1958. Hybridization in some species of Pteris L. Evolution 12: 82-92. WALKER, T.G. 1973. Additional cytotaxonomic notes on the pteridophyta of Jamaica. 7rans. Roy. Soc. Edinburgh 69: 109-135. FERN GAZ. 12(6) 1984. 351 STOMATA IN PS/LOTUM AND 7MESIPTERIS T. G. WALKER Department of Plant Biology, The University, Newcastle upon Tyne, NE1 7RU, England. ABSTRACT Stomata in some species of Psi/otum and 7Tmesipteris show a characteristic incrustation of an unknown nature. Unlike the situation in other plants the incrusta- tion is confined to the guard cells, the other epidermal cells being naked. However, whilst the common possession of such features may be an additional argument for a close relationship between Psi/otum and TJmesipteris, the total absence of incrustation in Psilotum complanatum is a warning that absence may be of only limited value as evidence of lack of affinity. INTRODUCTION During the course of a comparative investigation of triploid and tetraploid specimens of Psilotum nudum (L.) Beauv. measurements were made of stomatal length (Walker, in press). In some cases the limits of the guard cells were not very clearly defined and it was also evident that the stomata were not lying in exactly in same focal plane as the general surface of the epidermis. In order to clarify the position the epidermis was examined in the SEM. These initial scans lead to a broadening of the investigation to include other members of the Psilotales and the results are presented below. MATERIALS AND METHODS The wild origins of the plants used are set out in Tab. 1. The samples of material of Psilotum nudum were taken from living plants which had been in cultivation at Newcastle for almost two decades, whilst samples of P. complanatum Sw. and Tmesipteris lanceolata Dang. agg. were from herbarium specimens. The herbarium material was rehydrated by immersion in hot water containing some detergent and then treated as for the living samples. This involved fixation for 3 hours in 0.1 cacodylate at pH 7 and room temperature, followed by dehydration in an ascending acetone series and critically point dried using a Tousimis Samdri 780 critical point drier with liquid carbon dioxide. The specimens were then sputter-coated with gold. OBSERVATIONS Both triploid and tetraploid forms of Psilotum nudum from Papua New Guinea and Mauritius respectively had deeply sunken stomata on their aerial shoots. These stomata were heavily incrusted by a network of material which totally obscured the guard cells, leaving only the opening free (Figs. 1-4). A noteworthy fact is that the incrustation is confined to the stomata and does not occur elsewhere on the epidermis (Fig. 1). Whilst cuticular secretion of waxes is acommon phenomenon in plants (see, for example, Martin & Juniper, 1970; Juniper & Jeffree, 1983; Brandham & Cutler, 1978) resulting in deposits of various morphology, as far as the author is aware all such deposits occur over both epidermal and stomatal cells. The present observations are still at a preliminary stage and an analysis of the nature of the incrustation has not yet been attempted. However, immersion of the material for several hours in xylene and in chloroform has no effect and presumably it is not composed of the more soluble waxes. It will be seen that the incrustation is relatively thick, the edge of the guard cell being arrowed in Fig. 3. Although accurate measurements have not been possible because of the angle of the specimen the indications are that the thickness is of the order of 8um. The investigation was extended to include the stomata of the other very different- looking member of the Psilotales, 7mesipteris. 352 FERN GAZETTE: VOLUME 12 PART 6 (1984) FIGURES 1-3. Psi/otum nudum C2142 (4x) aerial shoot. 1. Epidermis & stomata, x 300; 2. Stoma, x 1000; 3. Part of stoma, edge of guard cell arrowed, x 2000. FIGURE 4. P. nudum, T8897 (3x) aerial shoot; part of stomatal incrustations, x 4000. STOMATA IN PSILOTUM AND TMESIPTERIS 353 FIGURES 5-7. 7mesipteris lanceolata, T12399, leaf surface. 5. Epidermis & stomata, x 300; 6. Stoma, x 1000; 7. Part of stoma, x 3000. FIGURE 8. Psilotum complanatum, T9545, Stoma, x 1000. 354 FERN GAZETTE: VOLUME 12: PART 6 (1984) TABLE 1. Sources of material of Ps/lotum and Tmesipteris. Collector's Number Origin P. nudum (L.) Beauv. a) triploid Walker T8897 Papua New Guinea. Near Moro, Finisterre Mts, c. 1700m, terrestrial 1964. b) tetraploid Page 2142 Mauritius. Macabé Forest, c. 580m, epiphytic, 1968. P. complanatum Sw. Walker T9545 Papua New Guinea. Wagau, c. 1130m, epiphytic, 1964. Imesipteris Walker T12399 Sulawesi, on ridge above N. bank lanceolata Dang. agg. of River Pasui, Latimodjong Mts, c. 1750m, on trunk of tree fern, 1969. Rehydrated herbarium material of a leafy appendage of 7. /anceolata agg. gave satisfactory results (Figs. 5-7), although it was contaminated by a sparse network of fungal hyphae. In this instance the stomata do not appear to be as deeply sunken as in Psilotum nudum but they also have the incrustation which is likewise absent from other epidermal cells. Some idea of the fineness of the constituent fibrils of the incrustation may be gained by comparing the diameter of the fibrils with that of the contaminating fungal hyphae (Fig. 6 and bottom left hand corner of Fig. 7). It will also be seen that the fine structure of tne 7mes/pteris incrustation is virtually identical with that found in triploid Ps//otum (compare Figs. 4 & 7) and argues for it being of the same or very similar nature in both cases. Psilotum consists of only a few species, many authors recognizing only two, P. nudum and P. complanatum. P. complanatum contrasts with the former species in being an epiphyte with long pendulous, flat, ribbon-like shoots instead of the erect rounded or angular ones of P. nudum. A piece of herbarium specimen of P. complanatum was rehydrated and on examination showed very deeply sunken stomata as in P. nudum but contrasted strongly from other members of the Order in completely lacking any incrustation (Fig. 8). DISCUSSION The presence of incrustations on the stomata of pteridophytes appears to be a rare phenomenon, particularly when associated with a lack of similar material on the epidermis. The only other notable example of stomatal ornamentation occurs in Equisetum (Page, 1972) but here the structure and constitution are quite different, being siliceous in nature and in the form of very precisely delimited protrusions or small papillae instead of a reticulum. The presence in common of such unusual incrustations in both Ps//jotum and Tmesipteris helps to strengthen the evidence for the association of the two genera in the same Order. However, the lack of such a phenomenon in P. complanatum indicates that presence of incrustations is more important than absence as positive evidence of relationships. ACKNOWLEDGEMENTS My thanks are due to Josephine Camus, British Museum (Natural History) for the identification of 7mesipteris, to Dr C.N. Page, Royal Botanic Garden, Edinburgh, for the living Ps//otum plant from Mauritius, and also to Dr Trevor Booth, Department of Plant Biology, University of Newcastle upon Tyne, for technical assistance. REFERENCES BRANDHAM, P.E. and CUTLER, D.F., 1978. The influence of chromosome variation on the organisation of the leaf epidermis in a hybrid A/oe (Liliaceae). Bot. J. Linn. Soc. Lond. 77: 1-16. JUNIPER, B.E. and JEFFREE, C.E., 1983. Plant surfaces. Edward Arnold, London. MARTIN, J.T. and JUNIPER, B.E., 1970. The cuticles of plants. Edward Arnold, London. PAGE, C.N. 1972. An assessment of inter-specific relationships in Equisetum subgenus Equisetum. New Phytol. 77: 355-369. WALKER, T.G. Cytological observations on Psi/lotum. Indian Fern J. (in press). FERN GAZ. 12(6) 1984. 355 CYTOMORPHOLOGICAL STUDIES OF PROGENY OF ADIANTUM PERUVIANUM B.M.B. SINHA and A.K. VERMA Department of Botany, Patna University, Patna 800 005, India. ABSTRACT Adiantum peruvianum Klotz. (2n = 60), a plant of horticultural importance in India, has a gigas habit, irregular meiosis and is propagated vegetatively. Germination of spores is low, and selfed prothalli give rise to sporophytes that do not resemble the parent in morphology. The progeny show variation in form and in chromosome number, from 2n = 56 to 2n = 62; some plants show regular meiosis whereas others have mostly bivalents but a few univalents at metaphase 1. The probability that A. peruvianum is of hybrid origin, and derived from A. capi//us-veneris L. is discussed. INTRODUCTION Adiantum peruvianum Klotz. exhibits a conspicuous gigas habit of fronds as compared with other species of the genus Ad/antum. It is one of the most popular ferns in Indian gardens, and is maintained by vegetative propagation of the rhizome. Meiotic investigations of the taxon have established its diploid nature (2n = 60), with irregular meiosis and a presumed hybrid origin (Roy and Sinha, 1956). Development of the gametophyte has been studied by Kachroo and Nayar, 1953. Cytological and morphological studies of selfed progeny of this taxon are reported here. MATERIALS AND METHODS Spores were obtained from live plants in cultivation in several botanic gardens of India, and referable toAd/antum peruvianum. The procedure followed for sowing andraising the gametophytes was that of Lovis (1968). In the experimental gardens where these investigations were carried out, no other variety or form of Adiantum was grown, to prevent cross-contamination. Fully developed prothalli with archegonia were selfed in a watch-glass in a drop of tap-water containing a good swim of antherozoids from prothalli bearing mature antheridia. The young sporophytes were raised to maturity in a temperature-controlled glasshouse, the process taking a year to eighteen months. Such mature plants were selected for morphological studies. Cytological preparations were made following the schedule of Manton (1950). All micrographs have been taken from permanent slides. Live plants are growing inthe department greenhouse and herbarium specimens have been deposited in the Department of Botany, Patna University. RESULTS A dense sowing of spores produced only a few prothalli that had different shapes and sizes at maturity; some remained vegetative, whilst others bore sex-organs. Details of the selfing attempts and the results are presented in Tab. 1. Morphology: None of the selfed progeny resembles the parent stock plant, and the progeny show considerable morphological variation (Fig. 1). Using major points of difference, the progeny have been divided into five groups, andthe characters of each of these groups are set out in Tab. 2. Cytology: A few of the sporophytes showing wide morphological differences were cytologically investigated to determine the chromosome number and the pairing behaviour of chromosomes during meiosis. Formation of regular bivalents was observed in most cells, but the number of bivalents was variable, ranging from 28 to 31 in different sporophytes (see Tab. 3 and Fig. 2). In some plants a few univalents are observed; in such cases, the total chromosome number was higher than the diploid number of the parent. 356 TABLE 1. Details of gametophytes selfed, percentage of sporophytes raised and number of adult sporophytes surviving. Year 1969 | 1970 | 1971 | TOTAL Number of gametophytes selfed Number of Number of Percentage of sporophytes adult sporo- sporophytes obtained phytes survived in relation to number of gametophytes selfed 6 3 18.75 6 5 15.62 19 15 17.85 2 2 25.00 12 11 34.37 3 2 25.00 13 9 2577N 61 47 21.86 (Mean) FERN GAZETTE: VOLUME 12 PART 6 (1984) TABLE 2: Morphology of different groups of selfed plants General appearance of the frond Group number | Oblong Il Lanceolate Il Ovate IV Spathulate V Elliptic Size of frond in cms 16 14 20.8 14 4.8 Size of stipe in cms 5.5 4.6 Sh 2 1.6 Number Size of of pinnae pinnae (average) in cms 16 2.1 13 1.9 13 3 11 0.9 9 0.8 Shape of Scales pinnules Rhomboid. Margins of lower pinnules with shallow incision but terminal ones with deep incision. Pinnules distantly placed. Absent Rhomboid. Incisions deep. Pinnules rather close. Sparse Pinnules Dense dissected, incision irregular, pinnules very close giving denser appear- ance to the fronds. Somewhat orbicular, with irregular dentate margin, pinnules closer. Sparse Rhomboid with shallow incisions in margin, pinnules sparse. Dense PROGENY OF ADIANTUM PERUVIANUM 357 PV 180 PV 181 FIGURE 1. Silhouettes of fronds of the stock plant 1/5 nat. size and a few raised progeny. Each frond bears the number of the selfed plant from which the frond has been taken. 358 FERN GAZETTE: VOLUME 12 PART 6 (1984) FIGURE 2. Dividing spore mother cells at diakinesis from a few selected sporophytes showing: 28 pairs (A), 29 pairs (B), 29 pairs and 2 singles (C), 30 pairs (D), 31 pairs and 1 single (E) and 31 pairs (F), all at x1000. TABLE 3. Cytological observations in the selfed plants Number of Nature of chromosomal associations investigated Univalents Bivalents Total number of plants Range Mean Range Mean chromosomes 2 0-2 2 27-28 27 56 25 ) ) 30 30 60 10 0-2 2 29-30 29 60 8 ) 0 29 29 58 1 1-3 1 30-31 31 63 1 19-23 21 22-25 24 69 PROGENY OF ADIANTUM PERUVIANUM 359 DISCUSSION The population of Adiantum peruvianum that has been investigated is a garden sample, and belongs to a single clone since it is always propagated vegetatively. Nayar (1962) has described the taxon in India, mentioning its gigas habit. Results of the experimental investigations here are noteworthy for the total absence of progeny resembling the parent in morphology. These progeny differ not only from the parent plant, but also amongst themselves. Of the five groups into which they have been divided (Tab. 2), one is very similar to A. capi/lus-veneris whilst others resemble garden varieties of this species. It is suggested that some form of A. capi/lus-veneris and one of its varieties may have hybridized, giving rise to a plant with fronds with a gigas habit. Selfing of such a hybrid would result in segregation of genes, with morphological variation in the offspring and possible loss of the gigas habit. The selfing attempts were inter-gametophytic and this, coupled with the non- homology of a few chromosomes in A. peruvianum (Roy and Sinha, 1956) may be responsible for the presence of univalents in some of the progeny. The range in morphology and chromosome number in the progeny supports the view that A. peruvianum is of hybrid origin. ACKNOWLEDGEMENTS The authors are grateful to Prof. R.P. Roy, F.N.A., for encouragement and critical comments. We are further thankful to Dr D.B. Lellinger, Prof. G.L. Stebbins and DrT.G. Walker for their comments. Financial assistance from the U.G.C., New Delhi has been received in the form of a research grant. REFERENCES KACHROO, P. and NAYAR, B.K. 1953. Studies in Pteridaceae |. Observations on gametophytes of some species of Adiantum L. Phytomorphology, 3: 240-248. LOVIS, J.D. 1968. Fern hybridists and fern hybridising Il. Fern hybridising at the University of Leeds. Brit. Fern Gaz. 10: 13-20. MANTON, I. 1950. Problems of cytology and evolution in the Pteridophyta. University Press, Cambridge. NAYAR, B.K. 1962. Studies of Indian Ferns (Adiantum). Bull. N.B. Garden, Lucknow. ROY, R.P. and SINHA, B.M.B. 1956. Meiotic studies in Adiantum peruvianum Klotz. Curr. Sc. 25: 268-269. 360 FERN GAZ. 12(6) 1984 A CHROMOSOME COUNT FOR OSMUNDA VACHELLII FROM PENINSULAR MALAYSIA A. BIDIN Faculty of Life Sciences, Universiti Kebangsaan Malaysia Bangi, Selangor, Malaysia. ABSTRACT A chromosome count of 2n = 44 (22 bivalents) is recorded for Osmunda vachellii from peninsular Malaysia, thus providing further confirmation of the cytological uniformity of the genus. INTRODUCTION Osmunda L. is a nearly cosmopolitan genus, with about fifteen species belonging to the temperate and tropical regions of both northern and southern hemispheres (Tryon & Tryon, 1981). Ten species are present in Asia, some of which have overlapping ranges. O. vachel/ii Hk. occurs in southern China, and has been found at several localities in Vietnam, Cambodia and Thailand (Schmid, 1974; Holttum, pers. comm.). The only representative of the genus so far recorded inthe Malaysian region is O. javanica Blume, where it is mainly a species of higher elevations (Holttum, 1966). CYTOLOGY The genus Osmunda has been the subject of many studies, especially focussed on the temperate species O. rega/is L. Autotetraploid and triploid plants were experimentally produced by Manton (1950). Cytological records for wild species from all over the world are, however, uniformly 2n= 44 (Lovis, 1977). The author found O. vachellii plants on a rocky bank of the River Tembeling (elevation c. 50m.), together with Adjantum stenochlamys in Taman Negara National Park in Pahang, Malaysia, in March 1983. Only two populations were found, at different places along the riverbank. The plants differed from O. javanica, where the FIGURE 1. Meiosis in Osmunda vachellii Hook. from West Malaysia National Park in Pahang (AB | 1741). 1000 x. SHORT NOTES 361 fertile pinnae are restricted to the middle part of the frond, as in both populations large and small specimens regularly produced fertile pinnae from both the base and middle part of the frond, whilst the upper half remained sterile. Professor R.E. Holttum at the Royal Botanic Gardens, Kew, confirmed descriptions and photographs of the plants as O. vachellii, adding that he had seen a number of plants of this species growing in a similar habitat in Canton, China. Live specimens brought back to the Universiti Kebangsaan Malaysia rapidly became fertile, and a meiotic squash clearly showed that these specimens are diploid, with 22 pairs of chromosomes at metaphase 1 (Fig. 1). ACKNOWLEDGEMENT This work is partially supported by the Universiti Kebangsaan Malaysia Research Grant No. 18/82, to which | am indebted. REFERENCES HOLTTUM, R.E., 1966. Flora of Malaya, Vol. Il. Ferns of Malaya, Singapore. LOVIS, J.D., 1977. Evolutionary patterns and processes in ferns. Adv. Bot. Res., 4: 229-45. MANTON, I., 1950. Problems of cytology and evolution in the Pteridophyta. Cambridge Univ. Press, London. SCHMID, M., 1974. Vegetation du Viet-Nam. Office de La Recherche Scientifique, Paris. TRYON, R.M. & TRYON, A.F., 1981. Ferns and allied plants. Springer-Verlag, New York, Heidelberg, Berlin. SHORT NOTES PSILOTUM NUDUM: A NEW RECORD FOR ARABIA Psilotum nudum (L.) Beauv. is widely distributed in the tropics but until now has not been found in North Africa, tropical North East Africa or South West Asia. In October 1983 a single gathering was made by Mr Ken Gordon in the extreme south of the Yemen Arab Republic, in the Hujariyah region near Turbah above Aduf. This new locality fills a large gap in its distribution in the Old World Tropics, its nearest station being some 1200 miles away in southern Kenya. The Hujariyah region is particularly interesting, and forms a distinct phytogeographical unit within SW Arabia, containing many endemics. It is relatively dry with uncertain spring rain, although unfortunately no rainfall figures are available. Psilotum was found growing associated with a small plant of Ficus salicifolia (Moraceae) in a shaded crevice on sandstone cliffs at about 5600ft. Only one plant was seen but the area is botanically little explored, and other sites could well be found. |am grateful to John Wood for the following comments about the locality: ‘“The valley is very interesting; there is an abundance of the endemic species Crotalaria squamigera (Leguminosae), Vernonia bottae (Compositae) and a new species of Centaurea (Compositae). There is also plentiful Wen/andia arabica (Rubiaceae), A/oe rivierei (Liliaceae) and many other interesting plants. The outstanding plant, however, is Fuirena felicis (Cyperaceae), known only from this valley. The locality consists of a steep ‘V’-shaped valley cut through sandstone descending from the Dubhan plateau. A nearly permanent stream flows along the valley bottom with deeper pools, and there are springs emerging from the cliff side.” A voucher specimen has been deposited in the herbarium at the Royal Botanic Garden, Edinburgh. A.G. MILLER Royal Botanic Garden, Edinburgh 362 SHORT NOTES ASPLENIUM x CONFLUENS ? IN SNOWDONIA A single plant of what appears to be Asp/enium = confluens (T. Moore ex Lowe) Lawalrée (A. scolopendrium L. x A. trichomanes L. subsp. quadrivalens D.E. Meyer emend. Lovis) has been found near Llyn Idwal in Snowdonia. It was growing under a rock on a heavily grazed slope of mostly Festuca ovina L. and Nardus stricta L. Many similar rocks in the vicinity had Blechnum spicant (L.) Roth growing under them. Neither of the parents of the hybrid were evident in the immediate area, although A. trichomanes subsp. quadrivalens was seen a few hundred metres away. A. scolopendrium and A. trichomanes subsp. trichomanes are recorded in the relevant 10Km square. The underlying rock in the vicinity appears to be acidic ordovician rhyolite, but there are other lime-rich rocks within a short distance. The occurrence of the hybrids on such apparently acidic soil is surprising when the preferences of the parents are considered. FIGURE 1. The Asp/enium hybrid from Cwm Idwal (life size). The plant appears to be quite consistent with A. x confluens although there seems no reason why it might not be A. sco/opendrium ~x A. trichomanes subsp. trichomanes as this would be very difficult to distinguish morphologically. A small piece of the plant has been propagated and it is intended for this to be examined cytologically. A frond of the plant has been deposited in the herbarium of the British Museum (Natural History) (Fig. 1). A.C. PIGOTT 43 Molewood Road, Hertford, Herts., SG14 3AQ REVIEWS 363 BRITISH AND IRISH PTERIDOPHYTE RECORDS Compiled by A.R. BUSBY, BPS Recorder The following records are additions to the At/as of Ferns (1978) and are communicated to the vice-county recorder concerned. One error is reported. | would like to express my thanks to all BPS and BSBI members, to the many vice-county recorders, and especially to C.D. Preston and his staff at the Biological Records Centre, Monks Wood (ITE), for their help. As in previous years, records are presented thus; 100km square/10km square followed by the recorder’s name. Irish records are prefixed with an H. POST 1950 12 Lycopodium clavatum 36/51 J. Durkin, 31/49 A. Lewis. 5.1 Selaginella selaginoides 36/82 N.J. Hards. 6.2 Isoetes echinospora 07/94 A.A. Slack. TAX2 Equisetum *< trachydon H10/57 T. O'Mahony. 73 Equisetum fluviatile 45/22 J. Durkin. 7.3x7 Equisetum * dycei 17/39, 18/30 H. McHaffie. 7.4x7 Equisetum *< rothmaleri 18/30 H. McHaffie. 7.4x3 Equisetum ~ litorale 17/49, 27/04, 60, 36/15 H. McHaffie. 7.7x8 Equisetum ~* font-queri 33/32 A.R. Busby. 78 Equisetum telmateia 45/34 J. Durkin. 14.2 Hymenophylilum wilsonii 22/91 M. Dohrn. t6:1 Polypodium vulgare HOO/56 R. Rush, 38/80 J. Durkin, 52/94, 62/13 E.M. Hyde. 162 Polypodium interjectum HOO/56, 67, 68 R. Rush, 35/15, 45/24 J. Durkin, 42/57 R.P.H. Lamb, 62/16 E.M. Hyde. 16.2x1 Polypodium < mantoniae 36/16 H. McHaffie, 52/94, 62/03, 13, 16 E.M. Hyde. 16.3 Polypodium australe H10/67 R. Rush, 35/42 F.J. Roberts. 7.1 Pteridium aquilinum 45/46 J. Durkin, 53/24 N.J. Hards. 19.1 Phegopteris connectilis H13/69 D. Kingston. 2A Asplenium scolopendrium 36/51 J. Durkin, 53/04 N.J. Hards. 21.1x7b Asplenium x confluens HOO/86 R. Rush. Pd |e Asplenium adiantum-nigrum 35/94, 45/05 J.Durkin, 44/95, 53/04, 54/07, 16 E. Chicken. oa WY 4 Asplenium trichomanes agg. 44/84, 54/07 E. Chicken. 21.7b Asplenium trichomanes subsp. quadrivaléns 35/76, 88, 45/29, 46/02, 12, 22 J. Durkin. 21.9 Asplenium ruta-muraria 36/51, 45/18, 28 J. Durkin, 44/84, 54/05, 16, 27 E. Chicken. Ze VA Asplenium ceterach 51/50 R. Hibbs. 22.1 Athyrium filix-femina 36/51 J. Durkin, 53/34 N.J. Hards. 24.1 Cystopteris fragilis 36/51, 46/14 J. Durkin. 26.2 Polystichum aculeatum 36/51 J. Durkin. 26.3 Polystichum setiferum HOO/55 R. Rush, 36/43 BSBI Party, 43/97, 53/34 N.J. Hards, 54/14 F.E. Crackles. OTF Dryopteris filix-mas HOO/56 R. Rush, 27/09 N.J. Hards, 36/51, 80,45/46 J. Durkin. 27.3 Dryopteris pseudomas 35/15, 36/80, 90, 45/44 J. Durkin. 27.5 Dryopteris aemula H13/69 D. Kingston, 22/64 I.K. Morgan. 27:8 Dryopteris carthusiana 45/05 J. Durkin. 27.9 Dryopteris austriaca HOO/46 R. Rush, 36/51, 45/46 J. Durkin. 29.1 Pilularia globulifera 17/04 A. McG. Stirling. 30.1 Azolla filiculoides 36/27 J. Muscott, 43/97 |. Weston, 45/27 J. Durkin, 52/76 G. Heathcote. ERRORS 23.2. Gymnocarpium robertianum NOT 33/21 A.R. Busby. 364 REVIEWS REVIEWS MEMOIR OF THE LIFE AND WORK OF EDWARD NEWMANby his son Thomas Richard Newman, 82 pp. 1876. 147 x 213mm. Reprinted by E.W. Classey, 1980, (from Park Road, Faringdon, Oxon SN7 7DR). Price £2.50. This small book is an interesting sketch of the life of the most energetic of naturalists. Born on 13 May 1801 Edward Newman developed his interest in ferns when at school in Gloucestershire. Although his first love was entomology, which remained with him throughout his life, he began his first fernery in 1826 at Leominster in Herefordshire. Newman observed how ferns grew in the wild and built replicas of their habitats in his garden, the principles of which are graphically described in the Introduction to his History of British Ferns (1844). This little book that Mr. Classey has reprinted makes interesting reading. One wonders how Newman found time for botany! It emphasises, however, the need for someone to write a biography of Edward Newman — the pteridologist. A.C. JERMY THE FERNS AND FERN ALLIES OF SOUTHERN AFRICA by W.B.G. Jacobsen, 542pp. 1983. Butterworths, Durban. 225 x 290mm. Price £47. This book is more than a fern Flora in the conventional sense. The first 113 pages contain six chapters. A general introductory one on classification and morphology includes a note on the conservation of ferns. The author's discussion on the loss of habitats is realistic and frightening and emphasises the role of invading exotics (Australian Wattle and other Acacia species in particular) in destroying dry land scrub, macchia, and kloof (ravine) forest. Deprivation of forests is mainly the result of man’s clearing; indigenous forest now covers less than 0.2 per cent of the whole of South Africa and even less of Zimbabwe. A chapter on the distribution and one on phytogeography follow and two very substantial chapters on ecology, one in which the factors of topography, substrate and climate are discussed and a second where the fern flora of some 42 communities are described for this extensive area. The second part (382 pp) is the Flora proper — detailed descriptions of families, genera and some 300 species. As might be expected of this author, extensive and original field notes are given, as are keys, distribution maps and photographs (of herbarium specimens usually — unfortunately). This is litthe mention of spore characters which is a pity, especially in the discussion on Cystopteris fragilis, where we would like to know the distribution of the C. dickieana-spore type. The production is clear in two columns but the printers have not used a very fine screen for the photographs and the detail is lost. This is unfortunate as the habitat pictures in part 1 are unique. There are two appendices: A, a list of the species given and their relationships to rainfall, environment and distribution; and B, a similar list of species showing the relationship to altitude. There are few printing errors; one, which is consistent, is Hymenophyllum tunbridgense which it is interesting to see is the commonest filmy-fern in South Africa, and | noticed Widén in the bibliography misspelt as Vidén. But these are minor points. This is an excellent book and the author, by profession a geologist, has indeed contributed to our knowledge of ferns as they are seen in the field in southern Africa. This may not be the last word on the taxonomy of these ferns but it is certainly a mile-stone and an excellent example of field observation. A.C. JERMY REVIEWS 365 PTERIDOPHYTIC FLORA OF GARHWAL HIMALAYA by S.S. Bir, C.K. Satija, S.M. Vasudeva and P. Goyal. 83 pp 1983. Jugal Kishore, Dehra Dun, 248001 India. 195 x 258mm. Price Rs 95. This is an annotated catalogue of the ferns and fern allies of Mussoorie, Dehradun, Chakrata and the adjoining hills; in all, the flora consists of 157 species. The account abounds with bibliographical references but individual records are listed with useful ecological notes. There is a general chapter on the ecology of pteridophytes in the region which is interesting for the account of lithophytes and xerophytes. The reader is reminded of the food value of young Diplazium esculentum, D. polypodioides and, surprisingly, the hairy Botrychium lanuginosum. Medical uses are several: Adiantum for expectorants and diuretics, spleenworts to counteract enlarged spleens, and the siliceous stems of Equisetum debile for gonorrhoea. There is still much taxonomic work to be done on the ferns of India. Professor Bir’s little book highlights some of the problems and will hopefully stimulate comprehensive monographic studies. A.C. JERMY THE AUSTRALIAN FERN JOURNAL, published annually by the Fern Society of Victoria, Vol 1 No 7 1984. In 1979 the Fern Society of Victoria in Australia was launched under the Presidency of Chris Goudey, with an initial membership of over 200. Now it has more than 500 members and is still expanding. Right from its beginning the Society issued a monthly Newsletter which has increased much in size and in the interest of its contents since then. | enjoy receiving this publication. Now the Society has taken another step forward and has begun to publish the Australian Fern Journal under the editorship of David Jones. Acopy of Vol 1 No 1 for 1984 is before me as | write. It consists of 33 pages, containing much fern matter of interest, with 26 colour photographs (24 of them of ferns) of superb quality, most of them the work of Chris Goudey and David Jones, of the same high standard as those appearing in their book Exotic ferns of Australia. The coloured cover depicts a magnificent tree-fern in its natural habitat. Papers included are on the ferns of Victoria and New South Wales, the endemic ferns of Lord Howe and Norfolk Islands, and a trip to Borneo. | intend to offer copies of this first issue for sale through BPS Booksales and estimate that | shall be able to sell them at around £1.60. Purchasers of this first issue may find it of interest to join the Fern Society of Victoria. The annual subscription for membership, which includes the monthly News/etter, but not the Australian Fern Journal, is Australian Dollars 12.50for members outside Australia. Information on the additional cost for the new journal can be obtained from the Secretary of the Fern Society of Victoria, 14 Afton Street, Essendon, Victoria 3040, Australia. J.W. DYCE 366 REVIEWS PTERIDOPHYTES OF TAIWAN by C.M. Kuo, 1982. Published by the Taiwan Provincial Department of Education, Taiwan, Republic of China. Available from K.S. Hsu, Taiwan Forestry Research Institute, Botanical Garden, 53 Nan- Hai Road, Taipei, Taiwan. This is Volume Ill in a set of ten books prepared for high-school students. It is written in Chinese, but an English edition will likely be published within a year or two. A systematic list of the families and genera of Taiwan pteridophytes is given, which follows Pichi Sermolli’s classification of 1977. Acolour plate and a description of one species in each genus found on Taiwan is given. Family and scientific names are in Latin, common names are in Chinese. No synonyms are given. The world-wide distribution, ecological notes, and distribution in Taiwan are given for the species illustrated. The colour photos are excellent and are printed on high grade glossy paper. Nearly all photos were taken of plants in their natural habitat. Mr Kuo uses the ‘small genus’ concept so, instead of using one species to represent Lycopodium, he gives six photos and descriptions for six segregate genera recognized. Some of the other volumes in this set also deal with pteridophytes, but there is no duplication of photos or descriptions. For example, in Volume |, The Rare and Theatened Plants of Taiwan, seven species are listed and the descriptions and illustrations are not the same as those used in Volume Ill. Dr Hsu Kuo-Shih tends to use more familiar names, for example: Schizaea digitata (L). Sw. instead of Actinostachys digitata (L.) Wall. ex J. Sm. CHARLES E. DEVOL BOOK NOTES 367 BOOK NOTES FLORA OF ECUADOR No 18 (edited by G. Harling and B. Sparre) 14 (4) POLYPODIACEAE-THELYPTERIDOIDEAE by Alan R. Smith. 147 pp.1983. 168 x 234mm. Swedish Research Council, Stockholm. One hundred and one species, all treated as of 7he/ypteris Schmidel, are concisely and clearly described, with a key to their identification. Useful notes are given on their ecology reflecting the high standards of recent collections of the Aarhus school whose herbarium labels give more information than just altitude! The taxonomy is very sound and the whole work is what we have come to expect from the pen of Alan Smith. We look forward to other accounts in this series. ICONOGRAPHIA SELECTA FLORAE AZORICAE, Vol. 1, fasc. 1 & 2, edited by A. Fernandes and Rosette Batarda Fernandes, pp 285, 51 plates. 1980-1983. Coimbra. Volume One of this nicely prepared and printed work contains Pteridophyta and Gymnospermae; of the 51 plates only four however are gymnosperms. Each species account (47 in all) is individually signed and authors include the editors, Isabel Noqueira and Magarida Queiros, the latter with M. Lameiras being the principal artists. Edmund Launert and Georg Paiva describe a new Marsilea (M. azorica). All plants are described in full and although the text is in Portuguese, the full-page plates are excellent and the books are well worth obtaining for these alone. A.C. JERMY AZOLLA AS A GREEN MANURE: USE AND MANAGEMENT IN CROP PRODUCTION by Thomas A. Lumpkin and Donald L. Plucknett. Westview Tropical Agriculture series No 5 Boulder, Colorado. 230 pp. 1982. 148 x 224mm. Azolla species contain, in cavities within their leaves, bluegreen algae (Anabaena spp.) which have the ability to ‘fix’ atmospheric nitrogen. On decay (rapid in the tropics), nitrogenous compounds are released and become available to other plants growing nearby. The species is there encouraged in the rice fields and Azo//a culture is ‘big business’ in SE Asia and in China in particular. This book is about management of the crop but it contains useful chapters onthe identification, morphology and ecology of Azo//a species. There is obviously need for a definitive monograph and further field studies on this interesting genus. The book is impressive for its extensive bibliography, especially Chinese (450 entries). A.C. JERMY 368 THE FERN GAZETTE Original papers, articles, or notes of any length on any aspect of pteridology will be considered for publication. Contributions should be sent to: Dr. M. Gibby, British Museum (Natural History), Cromwell Road, London SW7 5BD The /ast date for receiving notes and articles to make the following summer number is: 31st December each year Authors should follow the general style of this number. Close adherence to the following notes will help to speed publication. NOTES FOR CONTRIBUTORS Manuscripts: Copy should be in English and submitted in double-spaced type with adequate margins, on one side of the paper only. Abstract: All papers, other than short notes, should include a short abstract, to be set at the head of the main text, indicating the scope of the topic and the main conclusions. Headings and sub-headings: These should follow the style of this number. (Primary sub- headings are centred capitals. Secondary sub-headings U and L case side roman. Tertiary sub- headings, if necessary are U and L Case italic, ranged to the left). Numbering of sub-heading should be avoided. Latin names: quote the authority at (usually) the first mention only, in the main text but, unless unavoidable for clarity, not in the title. All latin names should be underlined throughout the typed copy. /lustrations: Any number and combination of line and half-tone illustrations (original drawings or diagrams in ink, or photographs which must be black and white, and of good technical quality) can be included with a manuscript where these help to augment or amplify the text. Photographs should be of the required magnification or larger and need not be made up to full page plates. Each drawing or photograph should be marked on the back with details of author and figure number, and the top edge clearly marked “‘top”’. Illustrations will not be returned to the author unless specifically requested. Figure numbering: Grouped illustrations should follow the numbering system, fig 1a, fig 1b, fig 2a, fig 3, etc. Figure numbers should be applied to illustrations in pencil only or on a transparent overlay. Final lettering will be added by the editor in a style and size consistent with the journal. Figure captions: Type on a separate sheet from the manuscript — include any necessary details of magnification as submitted (the editor will apply any correction or reduction). Reference lists: Please follow closely the style of this number to speed publication. Lists in other styles may have to be returned to authors for re-typing. Reprints: Twenty-five reprints are supplied free of charge to authors, who may order in advance further reprints which will be supplied at cost (plus postage) if requested at time of returning the first proofs. BOOKS FOR REVIEW Books for review in the Fern Gazette or Bulletin should be sent to A.C. Jermy, Botany Department, British Museum (Natural History), Cromwell Road, London, SW7 5BD; for review in the Pteridologist books should be sent to M.H. Rickard, The Old Rectory, Leinthall Starkes, Ludlow, Shropshire SY8 2HP. BRITISH FERNS AND THEIR CULTIVARS A very comprehensive collection is stocked by REGINALD KAYE LTD SILVERDALE, LANCASHIRE CATALOGUE ON REQUES r FIBREX NURSERIES LTD Harvey Road, Evesham, Worcestershire te Hardy and tender ferns “88 ‘Begonias, Gloxinias, Hederas, Hydrangeas, Primroses, Arum Lilies and Plants for the cool greenhouse Catalogue on request JA CKAMOOR’S HARDY PLANT FARM eobalds Park Road, Crews Hill, Enfield, Middlesex EN2 9BG : Tel: 01-363 4278 Hardy ferns, unusual hardy herbaceous plants, shade and moisture loving perennials Send two ten-pence stamps for list i 5 att oe pe JOIN THE EXPERTS ee Los Angeles International Fern Society ss 28 page colour, illustrated journal plus fern lesson monthly _ international spore store, educational materials and books. ‘ ‘ * $10 domestic, $12 foreign annual dues. “ee Send to ~LAIFS, 14895 Gardenhill Dr., LaMirada, CA 90638, USA Sea THE NIPPON FERNIST CLUB . za , ae exists in Japan this large and active Society devoted - pes both amateur and professional. be For further information write to: Dept of Forest Botany, Faculty of Agriculture, Tokyo University, Hongo, Bunkyo-ku. Tokyo, Japan 113. THE FERN GAZETTE VOLUME 12 PART 6 CONTENTS MAIN ARTICLES Studies in the fern-genera allied to 7ectaria, 1. Acommentary on recent schemes of classification - R.E. Holttum A new species of Ctenitis from Borneo and a new combination in the genus — R.E. Holttum Chromosome numbers and ecological observations of ferns from El Tirol, Paraguay — A.R. Smith & Mercedes S. Foster A new Ophioglossum from India - Sharda Khandelwal & H.K. Goswami Asplenium obovatum in Brittany, NW France — A. Labatut, R. Prelli & J. Schneller A new species of Microlepia from South India -— B.K. Nayar & P.V. Madhusoodanan Another intergeneric hybrid in Grammitidaceae: Ctenopteris longiceps Grammitis sumatrana - Barbara S. Parris Hybridization in E/aphoglossum in the Mascarene Islands - D.H. Lorence Stomata in Psi/otum and Tmesipteris - T.G. Walker Cytomorphological studies of progeny of Adiantum peruvianum - B.MB. Sinha & A.K. Verma A chromosome count for Osmunda vachellii from peninsular Malaysia - A. Bidin SHORT NOTES Spike and sporangial abnormalities in Ophiog/ossum of Rajasthan, India - B.L. Yadav & T.N. Bhardwaja Psilotum nudum: a new record for Arabia - A.G. Miller Asplenium * confluens ? in Snowdonia - A.C. Pigott BRITISH AND IRISH PTERIDOPHYTE RECORDS REVIEWS BOOK NOTES 367 (THE FERN GAZETTE Volume 12 Part 5 was published on 5th September 1983) Published by THE BRITISH PTERIDOLOGICAL SOCIETY, c/o Department of Botany, British Museum (Natural History), London SW7 5BD ISSN 0308-0838 Metloc Printers Ltd., Old Station Road, Loughton, Essex. Qk Oo Regul Ba - FERN GAZETTE VOLUME 12 the ws 7 t ) 2 © - on A a 4x rs Seo 4c > ‘Le Ly is ~ \ ns : i FERN GAZETTE — INDEX VOLUME 12 Aconiopteris 342 Arthromeris wallichiana 87 Acrophorus Sia oli, ote Aspidium 247 Acropterygium 213 goeringianum 246 Acrostichum 185 sagenioides 3tS;, 36 aureum 98 trifoliatum 318 speciosum 98 Asplenium 50, 74, 80, 102, Actinostachys digitata 366 106, 115, 188, 189, Acystopteris 304, 313 246, 286, 287, 290, Adenophorus 340 304, 308, 327, 331 Adiantopsis 327 adiantum-nigrum 5—8, 16, 17, 22, 24, radiata ae 78, 80, 103—106, Adiantum 31,..90, 76, 215, 115, 446; 136, 137, 221,355, ; 365 143, 144, 149, 152, capillus-veneris Tide 1S, TE: 252, 255, 259, 306, 363 23—25, 78, 81, aethiopicum 50 84, 151, 195, 263, x alternifolium 309 264, 309, 355, 359 anceps 157, 158 caudatum 50 billetii 214 edgeworthii 84 billotii 17, 116, 264, 331, incisum 85 332 latifolium 98 bourgaei 271—274 lunulatum 85 breynii 309 malesianum ZA bulbiferum 33 peruvianum 355—359 calcicola 214 pseudotinctum 323, 326—328 ceterach 18, 19, 23-25, raddianum 195 133, 136, 144, reniforme 156 152). 252) 255. stenochlamys 360 259, 302, 306, tetraphyllum a23 363 venustum 85 claussenii 324.327, 326 Africa, Macrothelypteris new to, 117 coenobiale 214 Aglaomorpha 225—228 x confluens 252. 259. 301, pilosa 227 302, 362, 363 Aleuritopteris 85 cuneifolium 5—8, 103—106 Alsophila 287 dalhousiae 87 bryophila 287 ensiforme 87 dregei 195 exiguum 87 dryopteroides 287 fissum 81 Amauropelta 160 fontanum 81,275, Sat Ampelopteris prolifera 87 foreziense 331 Amphineuron opulentum 98 fuscipes 214 Anemia 320528 gemmiferum 193 dregeana 193 glandulosum 81 mexicana 36 glaucophyllum 98, 101 phyllitidis 323, a2e goeringianum 246 Angiopteris 51, 161, 162, 244 hostmannii 324, 327 evecta 161, 162 incisum 105 smithii 161 indicum 87 Anogramma 78, 80, 81 Jahandiezii 272—273 leptophylla 75, 76, 78—81, 85 kobayashii 105—106 264 laciniatum 87 Ant associations with Bornean laetum 324 rain forest ferns, 243 lepidum 81 Antigramma 321,.326, 328 longissimum 98, 100, 101 brasiliensis 323, 326, 328 lunulatum 194 plantaginea 323, 326 marinum 12.18. 79. Appearance and disappearance 23—25, 136, 144, of a Dryopteris carthusiana 152, 252, 259, colony, 224 302, 306 Arabia, Psilotum nudum new to, 361 montanum 304 Arachniodes 130 mucronatum 324, 326—328 Araiostegia pseudocystopter is 86 nidus 97, 98, 100, 101 FERN GAZ. 12 INDEX obovatum onopteris pellucidum petrarchae platyneuron prionitis pulcherrimum ruta-muraria rutifolium sandersonii xX sarniense scolopendrium septentrionale serratum splendens x ticinense trichomanes subsp. quadrivalens trojani unilaterale varians viride X Asplenophyllitis jacksonii angustisquamatum anisopterum drepanopterum falcatum filix-femina flexile foliolosum goeringianum var. pictum jseanum 331—333 57, 72,180, 1103, 104, 106, 115, NG 25279259) 262, 306, 309 98, 101 80, 81, 189 16S 2s—25; 136, 137, 143, 144, 149, 152, 153, 252, 255, 259, 302, 306, 363 193 193 116 12, 16, 17, 22—25, 115, 136, 137, 143, 144, 149— 15272527255; 259, 263, 301, 302, 306, 362, 363 80, 264 324, 326, 328 193 115 187119), 22—25; 74, 112, 136, 137, 143, 144, 149, 152, 157, 252200209, 274, 302, 306, 362, 363 19) 2 s6; 144, 252, 259, 301, 306, 362, 363 252, 259, 260, 305, 306 105 189, 246, 247, 286, 294, 304, 318 11, 18.19) 22) 25, 136, 137, 143, 146, 148— 150, 152, 252, 255, 260, 300, 306 latilobum megistophy/llum nigripes niponicum var. meta/licum var. Niponicum forma cristatoflabellatum var. pictum otophorum oxyphyllum pectinatum proliferum rupestre rupicola schimperi yokoscence Azolla filiculoides pinnata var. imbricata BADRE, F. & PRELLI, R. BARKER, MARION Belvisia Berwickshire, the ferns of, BHAMBIE, S. & MADAN, PARKASH BIDIN, A. BIGNALL, ERIC Blechnum attenuatum var. giganteum australe subsp. auriculatum brasiliense ensiforme fraxineum indicum meridense occidentale orientale punctulatum var. krebsii spicant sylvaticum tabulare Bolbitis quoyana Bommeria ehrenbergiana hispida pedata subpaleacea Borneo, A new species of Ctenitis trom, Borneo, Asplenium pulcherrimum new to, Botrychium lanuginosum 246 230, 232, 240, 301, 367 301, 307, 363 360 is 102, 199, 294, 326 194 327 324, 326 324, 326, 328 324, 328 a2 324, 327, 328 98 195 194 20222827 135-137, 145, 148, 150—152, 252, 255, 261, 294, 300, 306, 362 195 195 275 286, 287, 290— 292 287, 290 287, 290, 291 287, 290 287, 290 320 214 28; 215, 217; 221, 222, 334 84, 216, 218, 365 /unaria ternatum British Isles, Asp/enium x confluens rediscovered in, British Isles, Dryopteris x sarvelae new to, British pteridophyte records, British record of Asp/enium cuneifolium suspect, British record, second of Equisetum x font-queri, Burren, Ferns of the, BUSBY, A.R. Camptodium Camptosorus sibiricus Campy/oneurum lapathifolium phyllitidis Canary Islands, New Dryopteris hybrids from, Cardiomanes Ceratopteris Ceterach cordatum officinarum C-Glycosy|xanthones in Tectaria, CHANDRA, SUBHASH Cheilanthes albomarginata anceps var. brevifrons austrotenuifolia bergiana caudata chlorophylla concolor var. kirkii contigua coriacea dalhousiae distans farinosa fragrans gracillima hirsuta hispanica inaequalis var. buchananii lasiophylla marantae microphylla persica pteridioides pumilio shirleyana 15, 136, 140, oe, 202; 257; 305 84, 216, 218 301, 362 178 305, 363 103 61 9 305, 363 279, 277; 276, 280, 281, 304 74 105 160 324, 327 324, 327, 328 225; 299 74,650,121 — 129) 159,274, 324, 328 122, 124, 125, e277, 194 121 324, 327 324, 327, 326 193 N26; 127 195 27 FERN GAZ. 12 INDEX sieberi subsp. pseudovellea subrufa tenuifolia vellea viridis var. involuta var. macrophylla Chemistry in fern classification, Christella arida dentata Parasitica subpubescens China, First Pteridological Symposium in, Christensenia aesculifolia Chromosome count of Asplenium anceps, Chromosome count of Christensenia, Chromosome count of Macroglossum, Chromosome count of Osrunda vachell/ii, Chromosome count of Phanerosorus, Chromosome counts of Paraguayan Ferns, Cionidium Classification of genera allied to Tectaria, Commentary on recent schemes of, Coniogramme affinis caudata COOPER-DRIVER, GILLIAN A. & HAUFLER, CHRISTOPHER Corfu, Fern-dominated wall communities in, CORIEEY AHLVs & GIBBY, IM: Crepidomanes insigne Crypsinus stenopteris Cryptogramma crispa Ctenitis apiciflora connexa dubia eatonii hypolepioides mannii muluensis rhodolepis speciosissima subobscura subincisa submarginalis Ctenitopsis 127—-V25) eT, 128 308 136, 140, 152, 305 189; 275; 277, 278, 280, 281, 304, 313-318, 320, 327 314 324, 326 316 SH 317 320 320 3t5 ee HL 320 315, 316, 324 324, 327 S15 7390 FERN GAZ. 12 INDEX Ctenopteris barathrophylla brooksiae curtisii heterophylla longiceps rufidula Culcita coniifolia macrocarpa Cyathea latebrosa Cyclogramma auriculata Cyclopeltis Cyclophorus dimorphus stellatus Cyclosorus dentatus gongylodes interruptus Cyrtomium caryotideum Cystopteris dickieana fragilis subsp. bulbifera subsp. protrusa subsp. tenu/folia protrusa Cytomorphology of Adiantum peruvianum progeny, Cytotaxonomy of Azolla pinnata, Danaea Davallia canariensis chaerophylloides denticulata pulcherrima Davallodes borneensis Dennstaedtia flaccida globulifera obtusifolia scabra Diacalpe Dicksonia Dicranoglossum Dicranopteris linearis var. subpectinata Dictymia brownii Dictyoxiphium Didymochlaena 337, 338, 340 118 180 338, 340 338, 340 337—340 118 186, 199, 299, 119 50, 288, 291, 292 308, 309, 364 18, 19, 23—25, 74, 81, 136, 137, 143, 146, 149, 151, 152, 156, 252, 259, 260, 263, 288, 306, 308, 363, 364 288 288 288 74, 291 355 230 51, 162 53, 188, 304, 313, 317 UZ 194, 195, 196 96, 98, 100, 101 214 Sil 98 198, 199, 206 197—206 304 304, 314-318, 326, 328 truncatula Didymoglossum insigne Diphasiastrum alpinum x issleri Diphasium Diplazium ambiguum angustisquamatum asperum cristatum esculentum expansum fuertesii latilobum lobulosum megistophy/lum polypodioides spectabile striatum Diplopterygium Diplora Dipteris conjugata DIXIT, R. D. & DAS, ANJALI Doodia media Doryopteris concolor nobilis pedata var. mul/tipartita Drought tolerance in Cheilanthes with special reference to the gametophyte, Drymogtossum piloselloides Drynaria glauca mollis quercifolia sparsisora Drynarioid ferns, A new concept of, Drynariopsis Dryoathyrium Dryopolystichum Dryopteris aemula affinis subsp. borreri subsp. sti//upensis 252, 254, 305 136, 138, 152 309 189, 318 326, 98, 365 182 123, 128, 199 124, 127 324 121 98, 100, 101 225—228, 244 119 98, 100, 101, 227, 244 225 225—228 313 314, 316, 317 27, 28, 36, 50 56, 74, 153, 183, 184, 246, 247, 267, 294, 313—315, 317, 318, 343 21, 22) 135; 136, 147, 152, 245, 252, 255, 261, 267, 270, 300, 306, 307, 363 56, 156, 183, 184, 252, 254, 255, 260, 267— 270 300, 306, 307 252, 261 249, 252, 261 x ambroseae ardechensis austriaca x bootii campyloptera carthusiana x cebennae xX cedroensis chrysocoma cochleata corleyi X Geweveri dilatata expansa filix-mas x fraser-jenkinsii goeringiana X gomerica guanchica inaequalis laeta marginalis marginata odontoloma oligodonta oreades paleacea protensa Pseudomas remota xX sarvelae xX sarvelii sparsa squamulifera submontana x tavelii tyrrhena villarii wallichiana Dryostachyum 56, 178 183, 184 1.1, 20—22, 25, 74, 136, 137, 145, 147—150, 152, 254, 306, 25, 1350, 137, 145, 147, 148, 150—153, 178, 224, 252, 261, 306, 363 - 300 133, 136, 148, 150, 178, 306 115,224, 252, 254, 255, 261, 268, 269, 300, 307 178, 249, 252, 261, 263, 306 11, 20—24, 36, 37, 136, 137, 145, 147—150, 152,252, 255, 260, 264, 306, 267 300 267, 268 249, 252, 260, 262, 306, 307 156 316 20-24, 27,56, 135197, 140, 147—152, 254, 184 G21, 25; fou, 261 183, 184 35, 183, 184, 264 156 225, 226 FERN GAZ. 12 INDEX Ecology of Paraguay ferns, Ecology of Phanerosorus, Edinburgh, Azo/la filiculoides in, EDWARDS, P.J. Elaphoglossum acrostichoides x adu/terinum angulatum x cadetii coursii heterolepis x heterophlebium hybridum lanatum lepervanchei macropodium reticulatum x revaughnii richardii xX setaceum sieberi tomentosum England, Asp/enium adiantum- nigrum aggregate in, England, Asplenium cuneifolium new to, England, Equisetum x font-queri new to, England, Equisetum x trachydon new to, Equisetum arvense bogotense debile x dycei fluviatile x font-queri hyemale var. californicum x litorale 321 209 301 224, 241 160, 286, 304, 341, 342, 346, 347, 349 346, 347 341, 345 347- 349 342, 347, 349 341, 345—347 349 346, 347 343, 346 341, 343—346, 349 345, 348, 349 342, 345, 347— 349 342, 343 195, 342 342 341—345, 349 342 341, 345 348, 349 342, 343 342, 343, 346— 348 5 5, 103 61 59 27, 30, 40, 41, 59-62, 74, 115353; 179; 286, 304, 308 13, 14,.22, 36; 116,136; 137, 139, 141, 148, 150,152, 252, 255, 256, 305 137, 139, 141, 150.152; 178; 179} 252, 255, 256, 305, 363 6 62, 116, 363 57—60, 113, 13651397 152; 179, 180, 252, 256, 305, 307 36 13; 133,136, 139) 4-79; 252, 255) 256; 305, 363 FERN GAZ. 12 INDEX mackati xX moorei palustre pratense ramosissimum robustum x rothmaleri sylvaticum te/mateia x trachyodon variegatum var. wilsoni wilsoni Eschatogramme Europe, Asp/enium bourgaei new to, Fadyenia FERREIRA, R.E.C. Flavonoids in Osmundaceae, Forked vein and foliar fibres in Selaginella, France, Asplenium obovatum new to Brittany, France, New records of Asplenium and Equisetum hybrids in, FRASER-JENKINS, C.R. FRASER-JENKINS, C.R. & JERMY, A.C. FRASER-JENKINS, C.R. & LAINZ, M., S.J. FRASER-JENKINS, C.R., RUSH, R. & CHING, R.C. GERSON, URI GIBBY, MARY & WIDEN, CARL-JOHAN Glaphyropteridopsis erubescens Gleichenia polypodioides GOMEZ, LUIS D. Goniophlebium mehipitense rajaense verrucosum Goniopteris prolifera Grammitis armstrongii billardieri clemensiae 57 249, 252, 263 14, 15, 61, 62, 116, 136, 137, 139, 141, 150, 1527 1785 179) 252, 257, 305 305 56, 116, 195 36 117, 363 sh USS, USE, 139, 150—152, 2527/2577, 505 14, 15, 61, 62, 116, 136, 139, (150, 162, 2EZ, 257, 303—305, 363 13, 57—60, 113, 114, 179, 180, 252, 256, 305, 309, 363 13, 56—60, 113, 179, 180, 252, 256, 305 256 256 331 iis 56, 155, 183 56 186, 208, 211, 213 195 196) 337,338, 340 165, 166 338, 340 118 fasciata intromissa kerguelenensis magellanica maxwellii papuensis plana poeppigiana sparsipila subdichotoma suck lingiana sumatrana torricelliana viridula vittariifolia GREUTER, W., PLEGER, R., RAUS, TH., ZIMMER, B. & GREUTER, J.J. Gymnocarpium dryopteris x heterosporum robertianum Gymnopteris tomentosa vestita Helminthostachys zeylanica Hemigramma Hemionitis Hemistachyum HENNIPMAN, E., DE JONCHEERE, G.L & PRICE, M.G. Heterogonium pinnatum Hicriopteris glauca Holostachyum HOLTTUM, R.E. Humata k inabaluensis var. subvestita subvestita Huperzia selago Hymenophyllum tunbrigense wilsoni Hypodematium crenatum Hypoderris Hypolepis SParsisOora India, a new species of Microlepia from, India, a new species of Ophioglossum from, 271 245, 304 19, 74, 133, 136, 146, 150— 152, 249, 252, 260, 263, 264, 306 74 74, 75, 264, 306, 363 325, 326 85, 88 PAWS) EPA Ti 2221) 222 215-218 275, 277, 280, 281, 304 287 225 47 189, 275, 281, 304, 313, 315, 225—228 185; 337320 28, 304 118 118 13;/25;,.136; 138, 152, 252, 254, 255, 305 102, 286, 304 252, 255, 257; 364 15) 1S 2o2 255, 257, 263; 305, 363 86, 211 304 188, 236, 237 195 335 330 Intergeneric hybrid in Grammitidaceae, Iran, Cheilanthes coriacea new to, Isle of Wight, Aberrant form of Equisetum te/lmateia from, lsoetes dodgei echinospora hystrix lacustris macrospora Japan, Equisetum x rothmaleri new to, JERMY, A.C. & PAGE, C.N. Kau/fussia aesculifolia KHANDELWAL, SHARDA & GOSWAMI, H.K. Killarney, The ferns of, KUNG-SHAR, SHING LABATUT, A., PRELLI, R. & SCHNELLER, J.J. Laminal flap in Se/agine//a, Lastrea foenisecii rubiginosa Lastreopsis shepherdii effusa Lecanopteris carnosa sinuosa spinosa Lemmaphy/l/lum Lepisorus amaurolepida excavatus kashyapii kuchanensis nudus scolopendrius Leptopteris superba Light response in Marsilea crenata, Light response in Regnel/lidium diphyllum Lithostegia Lomegramma LORENCE, D.H. Loxogramme involuta LOYAL, D.S., GOLLEN, A.K. & RATRA, RAMAN Lunathyrium japonicum Lycopodiella inundata Lycopodium alpinum var. decipiens annotinum carolinianum cernuum 252, 256, 305, 363 264 74, 252, 255, 256, 264, 305 74 117 113 51 330 249 119 331 180 186 147 317 123; 126; 189; 313—515; 318, 204 1997°295),2968 295) 29773296 62 252, 254, 305 36, 74, 288, 289, 366 FERN GAZ. 12 INDEX clavatum var. inflexum complanatum var. flabelliforme gnidioides quadrangulare verticillatum Lygodium flexuosum longifolium microphyllum Macroglossum alidae smithii Macrothelypteris bukoensis torresiana Malaya, New species of Plagiogyria in, Marattia fraxinea var. salicifolia Marginaria Marginariopsis Marsilea azorica crenata macrocarpa vestita Mascarene Islands, Hybridization in Elaphoglossum in, Matonia pectinata sarmentosa Matteucia struthiopteris Maxonia Meniscium Meringium Merinthosorus Microgramma lindbergii lycopodioides squamulosa vacciniifolia Microlepia haflangensis hancei manohara speluncae Microsorium membranaceum punctatum scolopendria MILLER, A.G. WHT GHELL ak: Moffat Hills, Woodsia i/vensis in, Mohria caffrorum Monographic studies on ferns, The continuing need for, Morphology of Azo//a pinnata, 36, 136, 138, 152, 252,254, 305, 307, 363 195 182 40, 51, 52, 161, 162 194 206 206 1317 132, 286; 304, 367 209-213 160, 206, 325 325 193 325—328 325 188, 335 336 336 335, 336 98, 101, 335, 336 225 88, 90 98, 100, 101, 193, 195 FERN GAZ. 12 INDEX Morphology of Po/ypodium macaronesicum and P. australe, Morphology of Stenosemia, MUKHERJEE, R.N. & SEN, U. MUKHOPADHYAY, R. & SENAU MURRAY, C.W. MUSCOTT, J. NAIR, G.B. Nannothelypteris Nainital (Western Himalayas), Ferns of NAYAR, B.K. & MADHUSOODANAN, P.V. Nectaries of Pteridium aquilinum, Field observations of, Nephelea portoricensis setosa sternbergii var. acanthomelas Nephrodium affine Nephrol/epis acutifolia biserrata exa/tata hirsutula radicans Neurosoria New combination in Bornean Ctenitis, New combinations in Southeast Asian ferns, Niphidium Nomenclature of Athyrium goeringianum, Nomenclatural notes on Dryopteris, Notholaena marantae sinuata standleyi trichomanoides Nothoperanema Oil palm plantations in West Malaysia, Ferns of, Oleandra distenta Olfersia Onoclea Onychium continguum divaricatum lucidum melanolepis siliculosum Ophioglossum azoricum costatum eliminatum gramineum lusitanicum 83 335 233 287, 326—328 287 S21) o2 On S26 100, 304 98, 100, 101 93, 96—98, 100, 101 33, 34, 195 211, 244 86, 98, 100, 101 199 320 118, 180 160, 204 246 123 215, 2179219 — 222, 330, 334 264 216, 219-221, 330, 334 330 216, 219—221, 330, 334 216,219) 220, 222, 264, 330 nudicaule palmatum pedunculosum pendulum petiolatum reticulatum thermale vulgatum Oreogrammitis clemensiae Oreopteris limbosperma Orthiopter is Osmunda cinnamomea claytoniana javanica regalis xX ruggiil vachellii Osmundacaulis Osmundites PAGE, C.N. PAGE, C.N. & BENNELL, FRANCES M. Paesia scaberula Paltonium Paraceterach PARRIS, B.S. PATERSON, SUSAN Pellaea atropurpurea calomelanos glabella involuta ornithopus Peltate scales in Sacco/oma, Peranema Phanerosorus major sarmentosus Phegopteris connectilis subobscura Philippines, New species of Selaginelia in, 216, 219-221, 330 334 234 98, 100, 101, 334 84, 90, 216, 219-221 216, 219-221, 334 216, 219, 220, 222, 330 14, 15, 84, 90, 136, 140, 241, 242, 252, 257, 305 118 313 27, 136, 137, 141, 142, 148, 150, 152, 249, 252, 255, 258, 40, 41, 50, 292; 2958 29e, 360 295,297, 298 36, 74, 295 360 15, 16; 74, 133; 136, 140, 152, 195, 252, 255; 257, 264, 295, 305, 309, 360 56, 57, 117, 178, 233 5 237 121 204 199 11:7), 1ahS;, Hes; 180, 214, 337 313, 315—st18 209, 211-213 209 209, 210, 212 136, 142, 150, 152, 245, 252, 258, 306, 363 320 169 Phlebodium aureum Photinopteris Phyllitis scolopendrium Phy matodes longissima nigrescens oxyloba scolopendria stracheyii Phymatopteris glauca taeniata var. borneensis PIGGOTT, A. G. PIGOTT, A. C. Pilularia globulifera Pityrogramma calomelanos ‘var. aureoflava trifoliata Plagiogyria euphlebia malayensis tuberculata Platycerium coronarium grande holttumii superbum wandae Platyzoma Pleocnemia irregularis Pleopeltis angusta excavata gibbsiae kashyapii macrocarpa scolopendria Pleuroderris Pleurosorus hispanicus Pneumatopter is Polypodium amoenum angustifolium argutum australe azoricum barathrophyllum bellisquaamatum brooksiae bryophyllum cambricum 123, 160 124, 127, 128 225—228 188, 308 104, 106, 115, 252, 261, 307, 363 328 96, 99, 101 195 325, 327, 325 208 27, 36, 50, 74, 153, 160, 188, 196, 199, 206, 245, 313, 325, 327 17, 25, 69—73, 252, 258, 306, 307, 363 69 118 119 180 119 307 FERN GAZ. 12 INDEX filicula friedricsthalianum glycyrrhiza griffithianum var. borneense hirsutissimum interjectum intromissum lachnopus macaronesicum x mantoniae maxwellii mehipitense microrhizoma neriifolium papuense planum pleolepis pleopeltifolium poly podioides subsp. ecklonii var. Minus rajaense recurvatum rufescens rufidulum x shivasiae siccum singeri sparsipilum squalidum stenopteris subamoenum subdichotomum suck lingianum torricellianum truncorum virginianum viridulum vittariifolium vulgare subsp. azoricum var. serratum Polystichopsis Polystichum acanthophyllum aculeatum 325, 327 325 74 119 325-327 17, 720473, 116, 136, 142, 252, 258, 305, 363 69—73 9, 17, 25. 136, 142, 249, 252, 258, 306, 363 118 325 325—328 327 193 325 118 325 118 118 306 325—327 321, 325-327 180 321, 325—328 74, 223 118 118 15-17, 22—25, 37, 71—74, 78, 79, 136, 137, 141, 142, 148— 152, 188,223, 245, 252, 255, 258, 264, 305, 27, 50, 74, 108—110, 153, 304, 317, 326 86 19, 20, 22—24, 74, 86, 136, 137, 145, 146, 149—152, 252, 255, 260, 306, 363 FERN GAZ. POPE, C.R. Pronephrium Prosaptia 12 INDEX var. angulare alaskense andersonii bicknellii braunii falcinellum imbricans indicum lobatum lonchitis lucidum luctuosum munitum nigropaleaceum obliquum platyphyllum polyblepharum pungens setiferum var. nigropaleaceum setigerum squarrosum tsus-simense triphyllum Pseudocyclosorus tylodes repens Pseudodry naria Psilotum complanatum flaccidum nudum Psomiocarpa Pteridium aquilinum var. arachnoideum var. wightianum var. yarrabense caudatum subsp. yarrabense var. yarrabense Pteridophyte and arthropod associations, 86 109 74, 109 US), ZQ, USI, 252, 260, 306 146 136, 146, 151, 152, 156, 252, 260, 263 11, 20—25, 86, WAS, VSS, USS), 136, 146, 149— 52252259) 260, 306, 309, 351, 354, 361 351, 353, 354 98, 101 98, 101, 351, 352, 354, 361 ATS, AU, PTS), 280, 281 Bi, SS, WAN, 236, 237 Up UG, Uo 4 ZZ 26), PAS), Pal, P48), S67 lis5—1S7, 141, 142, 148— Bre WE) ZO 239), 252) 255, 258, 263, 300, SOG6S2Z7, 335, 363 Pteridrys Pteris biaurita buchananii catoptera var. horridula cretica deflexa dentata subsp. flabellata denticulata ensiformis excelsa longipinnula multifida Quadriaurita semipinnata tripartita vittata Pyrrosia africana angustata beddomeana dimorpha flocculosa longifolia stellata stictica Quercifilix QUIRK, HELEN & CHAMBERS, T.C. Regnellidium diphyllum RASBACH, H., RASBACH, K. & SCHNELLER, J.J. Reviews: AUSTRALIAN FERN JOURNAL BAISHYA, A.K. & RAO, R.R. Ferns and fern allies of Meghalaya State, India BENL, G. : The pteridophyta of Fernando Po BIR, S.S. (ed.) Aspects of plant sciences (volume 6), pteridophytes BIR, S.S., SATIJA, C.K., VASUDEVA, S.M. & GOYAL, P. Pteridophytic flora of Garhwal Himalaya CEIEEORD. leas CONSTANTINE, J. Ferns, fern allies and conifers of Australia CODY, WILLIAM, J. Ferns of the Ottawa district DE LA SOTA, ELIAS, R. Flora de la provincia de Jujuy, Republica Argentina (ed. Angel L. Cabrera) Part II Pteridopitas 315, 317 31, 101, 115, 25) 221327 194 85, 89, 114, 15 325, 326 98 100, 101 365 310 110 Sa 365 190 46 DEVI, SANTHA Spores of Indian ferns 46 BYER, A-.F. Experimental biology of ferns 107 FERNANDES, A. & FERNANDES, ROSETTE BATARDA (eds.) Iconographia selecta florae azoricae 367 HENNIPMAN, E. & ROOS, M.C. A monograph of the fern genus Platycerium (Polypodiaceae) 266 HOLTTUM, R.E. Flora Malesiana Ser. 2 Pteridophyta Vol. 1 Part 5. Thelypteridaceae 208 HUANG, TSENG-CHIENG Spore flora of Taiwan 248 INSTITUTE OF TERRES- TRIAL ECOLOGY Overlays of environmental and other factors for use with Biological Records Centre distribution maps 82 IWATSUKI, K. List of the type specimens in the herbaria of Japan. Fern families 229 JACOBSEN, W.B.G. The ferns and fern allies of Southern Africa 364 JONES, D.L. & CLEMESHA, Sa) Es Australian ferns and fern allies 240 KRAMER, K.U. The pteridophytes of Suriname 102 KUO, C.M. Pteridophytes of Taiwan 366 KURATA, S. & NAKAIKE, T. (eds.) Illustrations of pteridophytes of Japan D235 LAKELA, OLGA & LONG, ROBERT W. Ferns of Florida — an illus- trated manual and identifica- tion guide 46, 130 LAN, CHANG KIAU (ed.) The eightieth birthday of R. E. Holttum, 1975 28 LOWS J.D: Evolutionary patterns and processes in ferns 50 LUMPKIN, THOMAS A. & PLUCKNETT, DONALD L. Azolla as a green manure: use and management in crop production 367 MICKEL, JOHN T. How to know the ferns and fern allies 74 NAKAIKE, T. Selected pteridological papers 1952—1978 of Professor S. Kurata 130 FERN GAZ. 12 INDEX NAKAIKE, T. New flora of Japan: pteridophyta NEWMAN, THOMAS RICHARD Memoir of the life and work of Edward Newman PAGE, C.N. The ferns of Britain and Ireland PETRIK-OTT, ALETA JO The pteridophytes of Kansas, Nebraska, South Dakota and North Dakota ROBERTS, R.H. The flowering plants and ferns of Anglesey ROUX, J.P. Cape Peninsula ferns SEILER, RALPH Una guia taxonomica para helechas de El Salvador SMITH, ALAN R. Flora of Chiapas (ed. Dennis E. 294 364 307 282 Breedlove) Part 2 Pteridophytes 196 SMITH, ALAN R. Flora of Ecuador No. 18 (ed. G. Harling & B. Sparre) 14(4) Polypodiaceae — Thelypteridoideae SHOEZE, ROBERT “G: Ferns and fern allies of Guatemala: Part 2 Polypodiaceae TAGAWA, M. & IWATSUKI, K. Flora of Thailand Vol. 3 Part 1 Pteridophy tes TAYLOR, THOMAS N. Palaeobotany — an introduc- tion to fossil plant biology THOMAS, BARRY The evolution of plants and flowers TRYON, ROLLA M. & ALICE FE. Ferns and allied plants with special reference to tropical America WAGNER, DAVID Systematics of Po/ystichum in Western North America WELTEN, M. & SUTTER, AGarne Atlas de distribution des pteridophytes et des phanero- games de La Suisse RICHARDSON, P.M. & LORENZ-LIBURNAU, E. ROBERTS, R.H. ROBERTS, R.H. & PAGE, C.N. ROUX, J.P. Root connections in Ophioglossum vulgatum, Rumohra 367 160 ua 164 309 108 309 304 1,69 61 191 241 316—318 FERN GAZ. 12 INDEX adiantiformis RUSH, R.J. Saccoloma elegans Sadleria cyatheoides Sagenia Salvinia auriculata molesta natans Schizaea digitata pectinata Schizoloma ensifolia Scotland, Equisetum x dycei in, Scotland, Equisetum x trachyodon in, Scotland, Pteris cretica and Selaginella kraussiana naturalized in, Selaginella adunca apoensis ascendens atimonanensis bisulcata boninensis caffrorum caulescens ciliaris chrysocaulos chrysorrhizos denticulata dregei griffithii helferii helvetica heterostachys intermedia intertexta involvens kraussiana lepidophylla longiaristata martensii mittenii myosurus nummularia peltata plana pricei reticulata rupestris selaginoides 3073S 9S 301 Se Whey, 1 ZAS) 114 28, 40, 41, 78, 80, 115, 169, Ue, WHS, 12/7, 180, 181, 232, 311 175—177 169, 170, 173 169 169-171 180, 181 173 195 ZS 175 175, 176 175 75, 76, 78—80, iW Se263 195 WS, WHS 175 169, 175, 176 WOE, UA, U7 SS), 1 7AS) 114, 115, 194, 252, 254, 261, 169, 170 NESE 0) 172, 180, 181 13, 14, 25, 114, 134, 136, 138, N52) 2497252, 254, 263, 305, 363 springiana subdiaphana tenera uncinata vaginata willdenovii Selliguea bellisquamata gibbsiae SHIMWELL, D.W. Sinephropteris SINHA, B.M.B. & VERMA, A.K. SLEEP, ANNE SMITH, A.R. & FOSTER, MERCEDES S. SOBEL, GAIL L & WHALEN, MICHAEL D. Solanopteris brunei South Atlantic and South Indian Ocean Grammitis, Spain, New Dryopteris hybrids from, Spain, New locality for Culcita macrocarpa in, Sphaerostephanos Spike and sporangial abnorme- heterocarpus polycarpus lities in Ophioglossum, Stegnogramma Steiropteris Stenochlaena palustris Stenolepia tristis Stenosemia aurita dimorpha pinnata Sticherus Stigmatopteris Stomata in Psilotum and Tmesipteris, Struthiopteris Taenitis blechnoides cordata TAN, BENITO C. & JERMY, A: GLIVE Tectaria brooksii coadunata decurrens devexa fuscipes incisa vasta sinit Tectaridium Thamnopteris Thayeria 169 180, 181 175, 176, 180, 181 334 160 160 185 93, 98, 100, 101 SISA Sion on 318 Sil 7/ 189) 275—> 281; 304 275,276; 279 275 169 189) -275;277, 278, 280, 281, 304, 313-318, 326, 328 304 281 Thelypteris bergiana brunnea confluens dentata erubescens hispidula interrupta l/imbosperma palustris phegopteris repens subvillosa thelypteroides torresiana totta xylodes Thyrsopteris Tmesipteris Todea lanceolata barbara Transkei, Southern Africa, Pteridophyta of, Trichomanes pyxidiferum var. melanotrichum radicans speciosum Trigonospora Turkey, Equisetum variegatum new to, Venation patterns in Ophioglossum, Botrychium and He/lminthostachys, VERMA, S.C. & KHULLAR, S.P Vittaria elongata ensiformis isoetifolia Wales, Killarney fern in, WALKER, T.G WALKER, T.G., & JERMY, A.C. WALKER, T.G., & PAGE, C.N. WILLMOT, A. Woodsia x abbeae alpina ilvensis oregana pulchella 102, 160, 196, 208, 309, 326, 327, 367 194 ot, a2 195, 208 321, 325—328 17, 75, 309 208, 252, 254, 258, 264, 307 328 351, 354 351, 353, 354 40, 41, 199, 295, 298 1955299; 297, 298 191 102, 196, 286, 304 193 1 145252, 257 208 56 215 83 100 98, 99, 101 98, 99, 101 igs 1 51, 161, 351 209 151, 263 65—67, 74, 80, 309 46 309 FERN GAZ. 12 INDEX Xiphopteris 340 bryophylla 119 conjunctisora 338 YADAV, B.L & BHARDWAAJA, T.N. 334 Metloc Printers Limited, Old Station Road, Loughton, Essex ‘ se 4 eas 4 aie’ NA ath .. 4 ’ At A, ite! OT alle ety a + rea rs ‘NQsA> HSONIAN _ i Zz 47) oa Sd = = oO za IOSHLINS SS3IYVUGIT oe = Ra 2 S ~~ AX: = = re 22) pp THSONIAN INSTITUTION z Ne. =< WW, = Y¥ AQ: S TN. . ” as = = Ww . 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