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FERN GAZ. 15(6) 1998 197

TRICHOMANES SPECIOSUM WILLD. (HYMENOPHYLLACEAE:
PTERIDOPHYTA) IN SOUTHERN SPAIN

FJ. RUMSEY & J.C. VOGEL
Conservation Biology Laboratory, Department of Botany,
Natural History Museum, Cromwell Road, London, SW7 5BD, U.K.

Key words: Macaronesian enclave, refugium, Tertiary relict flora, conservation.

ABSTRACT

The history of the discovery, present distribution and status of Trichomanes speciosum in
southern Spain is given. A new site in the Sierra de Montecoche, supporting the gametophyte
generation only, is reported. The sporophyte generation has apparently declined in outlying
areas; the extent to which this is a natural cyclical process in drier periods is uncertain. Habitat
degradation through human activity has undoubtedly exacerbated climatic effects. The
species array present supports the hypothesis that this area has acted as a European mainland
refugium for Tertiary relicts otherwise largely restricted to the Atlantic Islands. Protection of
the sites supporting Macaronesian floral elements is urgently required.

INTRODUCTION

Trichomanes speciosum Willd. (syn. Vandenboschia speciosa (Willd.) Kunkel) is the sole native
European species of a widespread genus (Tutin ef al., 1993). A Macaronesian-European
endemic, it has been considered one of the continent’s most threatened pteridophytes. Excessive
past collection for horticulture (Allen, 1969), and the herbarium, in combination with the
restriction of small sporophyte populations to few, scattered and vulnerable sites has
necessitated legal protection throughout the species range under the Bern Convention (Anon,
1979) and the Habitats Directive (Anon, 1992). Trichomanes speciosum has also been listed as
globally rare in the IUCN Red List (Walter & Gillet, 1998).

Recent studies, summarised in Rumsey et al. (1998), have revealed that the gametophyte
(sexual phase) of the life cycle, first described from cultivated material (Goebel, 1888), but not
reported in the wild until over a century later (Rumsey et al., 1990), is scattered across Britain
and Ireland and parts of continental Europe. Similar ‘independent’ Trichomanes gametophyte
populations have been reported in northern temperate areas of the U.S.A. (Farrar, 1967, 1985,
1992) but 7: speciosum would appear unique in that both populations with gametophytes only
and those with both gametophyte and sporophyte generations are widespread.

PAST RECORDS OF TRICHOMANES SPECIOSUM IN SOUTHERN SPAIN
Knowledge of the presence of Macaronesian floristic elements in the hills in the Algeciras area
dates from the report of the discovery of Culcita macrocarpa C. Presl in the Miel Valley, a
few kilometers W.S.W. of Algeciras (Allorge, 1934). However, both C. macrocarpa and the
now lost Dryopteris guanchica Gibby & Jermy had been collected in this area by English
botanists in the nineteenth century, who either failed to appreciate their significance or
misidentified the material (Fraser-Jenkins, 1982). Curiously Allorge (1934) did not record
Trichomanes speciosum, or Diplazium caudatum (Cav.) Jermy, both of which later authors
report as growing close to the Culcita in this area. This omission led Nieschalk and Nieschalk
(1965) to suspect that the Culcita site they subsequently discovered in the Miel Valley might
not be the same as Allorge’s, as they found T. speciosum to be a frequent associate and indeed
to be scattered elsewhere by the streambed lower in the valley. Contrary to the statement in
Salvo Tierra (1982), this was not the first discovery of T. speciosum in this region, that
distinction falling to the distinguished tropical botanist and bryologist P.W. Richards
(Richards, 1934). He had previously drawn attention to the Atlantic bryophyte interest of the

198 FERN GAZETTE: VOLUME 15 PART 6 (1998)

“waterfall valley” [= Miel Valley] which he first visited in the spring of 1931 (Richards, 1932).
Trichomanes was discovered on a return visit in April 1934, growing “very sparingly on wet
sandstone rocks in a small cave near a waterfall.”” He makes no mention of Culcita in his short
account. It seems most unlikely that this area should have been visited independently by
Richards and Allorge. The date of Allorge’s visit is not given in his account (Allorge, 1934) but
both he and Richards collected specimens of the Macaronesian moss Lepidopilum virens Card.
(as Plagiothecium argyrophyllum Geh.) here in April of that year. These were the only known
finds of this species on the European mainland until its recent rediscovery in this area by
Hakelier (Sérgio et al., 1995) and it seems probable, although not stated anywhere, that the two
botanised together.

Richards’ (1934) report and the oblique reference in Nieschalk and Nieschalk (1965) were
overlooked by many, including Molesworth-Allen (1971). It was this paper, in which she
reported T. speciosum from two unspecified localities, that brought to wider attention the
presence not just of Trichomanes but also other pteridophytes with Macaronesian distributions,
namely Diplazium caudatum, Pteris serrulata Forskal (= P. incompleta Cav.) and Cyclosorus
dentatus (Forskal) Ching (= Christella dentata (Forskal) Brownsey & Jermy), in Cadiz
province. Her careful researches in this region over the next few years revealed further sites for
T. speciosum in the Algeciras, Los Barrios and Tarifa districts, i.e. on the Sierra de Luna, Sierra
del Nino and Sierra de Ojén ranges (Molesworth-Allen, 1977). Her investigations then
effectively defined the species’ known range, with no subsequent published reports of additional
localities, although the fern may additionally have occurred a little further west, on the Sierra de
Saladavieja, the lower, westward continuation of the Ojén (Salvo, pers. comm.). It was searched
for in this area during the 1995 Group of European Pteridologists (G.E.P.) excursion, but
without success.

The distribution of Trichomanes speciosum was mapped by Salvo Tierra (1982) in his
comprehensive study of the pteridophyte flora of Andalucia, in two hectads, 60TFOO and
7OTFOO. The total number of sites and their location are not given in the published accounts
(Molesworth-Allen, 1971, 1977), which understandably sought to protect this species from
over-collection, a fate which had befallen another rare disjunct pteridophyte discovered by the
same author in this area, Psilotum nudum (L.) Beauv. (Molesworth-Allen, 1966). While she
regarded T! speciosum as “locally abundant in a number of arroyos” (Molesworth-Allen, 1977),
the three areas from which she then knew the plant supported only “40-50 plants or patches”,
of which few had been fertile.

AIMS

Two decades have passed since the above records from this phytogeographically important area,
during which time patterns of land use have changed, levels of damaging human activity have
increased, and higher local pollution loads and increasing aridity may have all affected these
vulnerable, unique, refugial areas. The distinct species assemblage which characterises the
Atlantic fringe of Europe and Macaronesia is particularly rich in cryptogamic species, many of
which are rare and endangered. Since 1994 research has been carried out at the Natural History
Museum into the distribution, ecology and historical biogeography of this community and its
constituents. As part of this wider study these isolated sites in southern Spain were visited to
determine the current extent of both generations of Trichomanes speciosum, a model organism
which had been well studied elsewhere (see Rumsey et al., 1998, in press).

MATERIALS AND METHODS
Where possible all previously reported sites of T. speciosum were visited. In addition, sites
known to support, or to have supported, other disjunct Macaronesian elements were visited to
ascertain whether the sporophyte, or the less conspicuous gametophyte generation was present.

TRICHOMANES SPECIOSUM IN SOUTHERN SPAIN 199

We were fortunate to have access to the field notes made by Mrs Molesworth-Allen and her help
and encouragement in the field.

RESULTS

Trichomanes speciosum was refound in all of the areas from which it had previously been
reported. However, the species is now present only as the gametophyte generation in the Sierra
del Nino, and on the eastern slope of the Sierra de Luna. One new site, well to the north of the
previous discoveries, was detected while investigating the distribution of other Macaronesian
taxa. Thus, we were fortunate to be guided to a recently discovered site for Culcita macrocarpa,
an arroyo draining into the Embalse del Guadarranque on the N.E. side of the Sierra de
Montecoche. The finder, Luis Federico Sanchez Tundidor, accompanied by Mrs.Molesworth
Allen, a friend, and the authors were delighted to discover that the damp rock ledges by an
impressive waterfall that supported the population of c. 15-20 Culcita plants also had extensive
mats of 7: speciosum gametophytes (Fig. 1). This extends the known range of both these species
by c. 13km to the north-east (the nearest known sites for each are on the Sierra del Nino). In the
area of the Sierra del Nifio, where 7: speciosum had previously been found, above La Granja,
only a single site with a very small quantity of gametophyte was discovered in 1997. The habitat
appeared generally unsuitable and had been further disturbed by a serious land-slip the previous
winter. Only a single sporophyte colony had ever been known to Mrs Molesworth-Allen in this
area (Molesworth-Allen, pers. comm.). No gametophytes were found in the extant site for
C. macrocarpa elsewhere in the Sierra del Nino. Very small quantities of gametophyte were also
found in a small streambed above the minor road between Tajo del Espino and Puerto de la
Zarza, on the east side of the Sierra de Luna, high above the head of the Miel Valley at c. 550m
altitude. A single large plant of C. macrocarpa grew by a small waterfall nearby.

Figure 1. Site of Trichomanes speciosum gametophyte population in the Sierra de Montecoche,
May 1997. (Photo FJ. Rumsey)

200 FERN GAZETTE: VOLUME 15 PART 6 (1998)

Figure 2. Trichomanes speciosum flourishing by small streams on the Dehesa de Ojén, May
1997. Also present Athyrium filix-femina (L.) Roth and, middle right, Pteris incompleta Cav.
(Photo J.C. Vogel)

TRICHOMANES SPECIOSUM IN SOUTHERN SPAIN 201

ee
Hook P
ee ae

BGS.

paren . Dis of Trichomanes she Willd. in southern eles ealed by field
urve ty ies 1997. Be oth generations recorded ® eee ameto eee eration on ae ©. The 500m
con

3b, The distri a the Sierra de Ojén, Sierra de Luna and Sierra del Nifto mapped on a
lkm grid basis. Contour lines at 100m intervals.

202 FERN GAZETTE: VOLUME 15 PART 6 (1998)

In the Miel Valley only a single sporophytic plant of Trichomanes speciosum was seen in
1995 and refound in 1997. This was growing adjacent to the only population of Culcita
macrocarpa seen on these visits. No Diplazium caudatum or Pteris incompleta were seen in the
valley in either year. The only area within which T. speciosum had maintained sporophytic
numbers was the northern flank of the Sierra de Ojén, the Dehesa de Ojén up to the Collado
Puerto del Viento on the S. E. flank of the Sierra de Luna. This population was apparently
thriving; many fertile plants were seen and juvenile sporophytes were common on gametophyte
mats (Fig. 2). The extent and condition of the sporophytes in this area would appear to depend
on the condition of the mixed evergreen-deciduous woodland in which they occur, itself
determined by activities such as cork-cutting and, more harmfully, clearance or burning. Indeed,
Molesworth-Allen (pers. comm.) had noted that in a previous visit (post her optimistic 1977
account) few sporophytes could then be found in this area. Diplazium caudatum, Pteris
incompleta and Polystichum setiferum (Forskal) Woynar were all seen in reasonable quantity
and health, the last, in number and location almost exactly as reported by Molesworth-Allen
(1977). Of concern was the apparent absence of C. macrocarpa, the loss of which could only
realistically be explained by its selective removal. The terrain is however, difficult to search and
this species and other interesting finds might still be made in the more remote areas.

The distribution of Trichomanes speciosum in southern Spain as revealed by our survey is
given in Figure 3.

DISCUSSION

The detection of the gametophyte generation of T. speciosum was to be expected given the
frequence of its occurrence in more northerly and continental areas of Europe (Vogel et al.,
1993; Rumsey ef al., 1998). What is, perhaps, surprising is that more sites were not located
fringing those that currently maintain both generations. Evidence from elsewhere and the finds
reported here from high on the Sierra de Luna and in the Sierra del Nino, demonstrate the ability
of the gametophyte to persist in areas which once supported the sporophyte. The lack of
additional gametophyte sites must reflect the general unsuitability of the habitat for both
generations; harsh environmental and climatic changes occur over such short distances that little
habitat exists that could be considered suitable for gametophytes outside of the very restricted
areas suitable for sporophyte growth.

When compared with earlier accounts (e.g. Nieschalk & Nieschalk, 1965; Molesworth-
Allen, 1971, 1977) the population present in the best known and most accessible site, the Miel
Valley, has suffered a severe and perhaps catastrophic decline. Trichomanes speciosum is not the
only taxon to have suffered in this way. Diplazium caudatum, first reported on the European
mainland from this site in 1969 (Molesworth-Allen, 1971) and suggested to have suffered from
collection (Molesworth-Allen, 1977), may now be absent from the valley. Culcita macrocarpa
has vanished from the more accessible of its past sites lower in the valley, but persists higher up
where most casual visitors do not reach. Increased rambling and other outdoor pursuits, opening
up of the undergrowth, felling in the wider area, higher incidence of fires, when coupled with a
run of drier than average years is steadily reducing the conservation interest of this area. Only
the inaccessibility of the still comparatively unspoilt and now unique vegetation on the Dehesa
de Ojen has acted to protect it. These very restricted woodland areas with their Tertiary relict
Macaronesian cryptogamic flora, the sole Iberian sites for a range of disjunct flowering plants,
e.g. Arisarum proboscideum (L.) Savi, are a national treasure and of international significance.
Their active conservation must be addressed.

ACKNOWLEDGEMENTS
We would like to thank the NERC who funded this research under grant GR3/09451. We also
must record our deep gratitude to Mrs Betty Molesworth-Allen for information, arranging
access with landowners, spritely company in the field and her optimistic navigation on the

TRICHOMANES SPECIOSUM IN SOUTHERN SPAIN 203

“road to Hell”! We also thank Fede Tundidor for sharing with us his exciting discovery of the
new Culcita locality.

REFERENCES

ALLEN, D.E. 1969. The Victorian fern craze. Hutchinson, London.

ALLORGE, P. 1934. Le Culcita macrocarpa Pres| [Balantium Culcita (L Herit.) Kauffm.] dans
les montagnes d’ Algesiras. Bull. Soc. Bot. France 81: 592-593.

ANON. 1979. The Convention on the Conservation of European Wildlife and Natural Habitats.
Council of Europe, Strasbourg.

ANON. 1992. Council directive 92/43/EEC of 21 May 1992 on the conservation of natural
habitats and of wild faunas and floras. Official journal of the European Communities 206:
7-49.

FARRAR, D.R. 1967. Gametophytes of four tropical fern genera reproducing independently of
their gametophytes in the southern Appalachians. Science 155: 1266-1267.

FARRAR, D.R. 1985. Independent fern gametophytes in the wild. Proc. R. Soc. Edinb. 86B:
361-369.

FARRAR, D.R. 1992. Trichomanes intricatum: the independent Trichomanes gametophyte in
the Eastern United States. Amer. Fern J. 82: 68-74.

FRASER- JENKINS, C.R. 1982. Dryopteris in Spain, Portugal and Macaronesia. Bol. Soc.
Brot., Sér. 2, 55: 175-336.

GOEBEL, K. 1888. Morphologische und biologische Studien: II. Zur Keimungsgeschichte
einiger Farne. Ann. Jard. Buitenz. 7: 74-119.

MOLESWORTH-ALLEN, B. 1966. Psilotum nudum in Europe. Brit. Fern Gaz. 9: 249-251.

MOLESWORTH-ALLEN, B. 1971. Observations on Spanish ferns. Brit. Fern Gaz. 10: 200-
202.

MOLESWORTH-ALLEN, B. 1977. Observations on some rare Spanish ferns in Cadiz
province, Spain. Fern Gaz. 11: 271-275.

NIESCHALK, A. & NIESCHALK, C. 1965. Ein Stammfarn (Culcita macrocarpa) auf
europadischem Boden. Natur und Museum 95: 495-498.

RICHARDS, P.W. 1932. Note on the bryophytes of the “Waterfall Valley” near Algeciras: an
outpost of the Atlantic flora. Rev. Bryol. et Lichén. 5: 1-9.

RICHARDS, P.W. 1934. Trichomanes radicans Sw. in the south of Spain. J. Bot. (London) 72:
B51

RUMSEY, F.J., SHEFFIELD, E. & FARRAR, D.R. 1990. British filmy-fern gametophytes.
Pteridol. 2: 40-42.

RUMSEY, F.J., JERMY, A.C. & SHEFFIELD, E. 1998. The independent gametophytic stage of
Trichomanes speciosum Willd. (Hymenophyllaceae), the Killarney Fern and its distribution
in the British Isles. Watsonia 22: 1-19.

RUMSEY, F.J., VOGEL, J.C., RUSSELL, S.J.. BARRETT, J.A. & GIBBY, M. in press.
Climate, colonisation and celibacy: population structure in Central European Trichomanes
speciosum (Pteridophyta). Bot. Acta.

SALVO TIERRA, A.E. 1982. Flora pteridofitica de Andalucia. Unpublished PhD. Thesis,
Universidad de Malaga, Spain.

SERGIO, C., BRUGUES, M. & CROS, R.M. 1995. Threatened bryophytes in the Iberian
Peninsula - Some results and comments. Crypt. Helv. 18: 51-56.

TUTIN, T.G., BURGES, N.A., CHATER, A.O., EDMONDSON, J.R., HEYWOOD, V.H.,
MOORE, D.M., VALENTINE, D.H., WALTERS, S.M. & WEBB, D.A. 1993. Flora
Europaea Vol. 1, 2nd ed. Cambridge University Press, Cambridge.

VOGEL, J.C., JESSEN, S., GIBBY, M., JERMY, A.C. & ELLIS, L. 1993. Gametophytes of
Trichomanes speciosum (Hymenophyllaceae: Pteridophyta) in Central Europe. Fern Gaz.
14: 227-232.

WALTER, K.S. & GILLETT, H.J. 1998. IUCN Red List of threatened plants of the world. New
York Botanical Garden, New York.

04 FERN GAZETTE: VOLUME 15 PART 6 (1998)

BOOK REVIEW

THE FERNS OF BRITAIN AND IRELAND. C.N. Page. 1997 (2nd edition).
Cambridge University Press, Cambridge. Hardback £95, ISBN 0 521 58380 2; paperback £40,
ISBN 0 521 58658 5.

The British Isles are not only blessed with a rich fern flora, they are also blessed with a wealth
of books about pteridophytes. One of the most outstanding and innovative contributions has
been the first edition of The ferns of Britain and Ireland by Dr C.N. Page, published in 1982.
This book has now appeared in its second edition, but at a considerably increased price. At such
a price, Page has to stand the test against the standard in European fern Floras, the //lustrierte
Flora fiir Mitteleuropa, in short ‘Hegi’, published in its third edition by Prof. K. Kramer in
1984.

Page starts his book with a general introduction to ferns/fern allies and the factors,
geography, soils and climate, which influence the distribution and occurrence of pteridophytes
in the British Isles. Fern taxa thereafter are organised in an alphabetical order, not reflecting the
phylogenetic or systematic order. For each species at least three visual aids are provided:
distribution maps (hand-drawn), a chart of the phenology and silhouettes/pictures. The text for
each taxon covers at least four parts: identification, variation, possible confusion, technical
confirmation and field notes. The distribution maps appear coarse and hand-drawn and are of a
standard which, I fear, is no longer acceptable taking into account progress in modern computer
technology. The parts on ‘identification, variation, confusion with other taxa’ and the
illustrations of the ferns (both as silhouettes and photos) are still some of the best available. Here
are the unquestionable strong parts of this book.

My appreciation of the ‘field notes’ provided for each taxon, is mixed. In a way, the field
notes are one of the highlights of the book, but they also contain some of the weakest parts,
especially when subjected to scientific scrutiny. On the one hand Chris Page lets the reader
participate in his enormous knowledge and field experience about the ecology and biology of
pteridophytes in the British Isles. I have read these chapters with great interest and I have gained
from it. On the other hand, the author can fall, in parts, into what I would describe as
‘unsubstantiated rumour’. While this has perhaps an entertainment value for the fortunate few
(like the reviewer), who can devote their life to the scientific study of ferns, it might cause
confusion and could lead to a misinformation of the general reader. We can read on page 380 of
the perpetuation of the myth of inbreeding depression being responsible for the absence of
Trichomanes speciosum sporophytes in the British Isles. Fortunately this problem is soon to be
rectified by Rumsey et al. (1999) in a forthcoming paper in the Biological Journal of the
Linnean Society. But there is worse to come on page 134: Page’s considerations about the status
of Athyrium flexile as an endemic Scottish taxon and as an “incipient insular species in
evolutionary terms”. Dr Chris Page has been in employment for many years at the Royal
Botanic Gardens of Edinburgh, a well-respected Scientific Institute. To proclaim his ‘gut
feelings’ on the status of A. flexile as a valued scientific opinion is irritating. It is well known
that scientific questions in this day and age are addressed by a well designed experiment and its
sometimes strenuous execution, and that is exactly what Heather McHaffie from Edinburgh has
been doing for the last 3-4 years. Therefore, the statement of Page that Athyrium flexile is a
Scottish endemic, whatever its genetic status, is not acceptable from a scientific point of view;
an answer to this question depends on the outcome of the experiments. Taxonomy has always
accommodated new scientific evidence, and will do so in the future. ...

Continued on page 214

FERN GAZ. 15(6) 1998 205

POPULATION ANALYSIS OF THE TYPE LOCALITIES OF SOME
RECENTLY RECOGNISED TAXA OF BRITISH PTERIDIUM
(DENNSTAEDTIACEAE: PTERIDOPHYTA)

K.M. BRIDGES, C.J. ASHCROFT & E. SHEFFIELD
School of Biological Sciences, University of Manchester,
3.614 Stopford Building, Oxford Road, Manchester, M13 9PT, U.K.

Key words: bracken, clonal growth, isozyme analyses, Pteridium

ABSTRACT
Isozyme analyses show that the type localities for plants recently ascribed to Pteridium
pinetorum subsp. osmundaceum and subsp. pinetorum contain several genetically distinct
individuals. Two geographically separated samples of P. aquilinum subsp. fulvum share
isozyme fingerprints which differ from any of the other plants analysed here, but which have
been found elsewhere in Scotland within Pteridium aquilinum subsp. aquilinum.

Although our data show that there is more than one Preridium genotype at two type
localities, they do not negate the possibility that the differences in morphology of these plants
from neighbouring P. aquilinum subsp. aquilinum are due to the influence of some
environmental factor(s).

INTRODUCTION
Prior to 1989 it was commonly accepted that all bracken (Pteridium) in the British Isles
belonged to one taxon - P. aquilinum (L.) Kuhn subsp. aquilinum. In 1989, and later in 1995, it
was proposed (Page, 1989; Page & Mill, 1995 a & b) that this was not the case.
Page’s 1989 paper introduced two taxa novel to the UK, and Page and Mill’s publications
in 1995 brought the total of Pteridium taxa reported in the UK to five, see Table 1 for summary.

Table 1: Changes in Pteridium taxonomy in the British Isles according to Page (1989)
and Page and Mill (1995a & b).

Date Pteridium taxa recognised

Up to 1989 ~— Pteridium aquilinum (L.) Kuhn subsp. aquilinum was the only widely accepted
taxon
1989 P. aquilinum subsp. aquilinum
subsp. atlanticum C.N. Page (new subsp.) Page 1989.
subsp. latiusculum C.N. Page (new to UK), Page 1989.
1995 P. aquilinum subsp. aquilinum
subsp. atlanticum
subsp. fulvum C.N. Page (new subsp.), Page and Mill 1995a
Pteridium pinetorum C.N. Page & R.R. Mill (new species), Page and Mill 1995a
subsp. pinetorum C.N. Page (new subsp.)
subsp. osmundaceum (sensu Christ 1900) (Christ) C.N. Page (new
subsp.)

These recently proposed taxa were defined solely on the basis of gross morphological and
phenological characters in field specimens. The areas of type localities described as colonised
by these taxa are small: the largest single patch encountered in this study (P. pinetorum C.N.
Page & R.R. Mill subsp. pinetorum) extends for only 150m in maximum dimension, see Figure
1. Individuals of Pteridium are, however, well known to be capable of covering large areas by
means of vegetative spread. Sheffield et al. (1989), for example, detected single genotypes

206 FERN GAZETTE: VOLUME 15 PART 6 (1998)

woodland
| pinetorum
se ‘ype locality! som

(P1-15)

oS samples taken
S ] along

transect line

possible
pinetorum
site, (217)

a patch of
x aquilinum, (P16)

Loch an Eilean PB

Figure 1: Schematic diagram of P. pinetorum subsp. pinetorum type locality, GRNH
896092, Inverness-shire, Scotland.

The shaded patch is at the cited grid reference for the type locality; the fronds in the patch on
the left of the figure had a morphology similar to that of those at the published type locality. PA-
PE are the different isozyme genotypes identified at the site. P16 is a patch of Pteridium
aquilinum subsp. aquilinum.

BRITISH PTERIDIUM 207

extending for at least 390m in the UK, and Parks & Werth (1993) mapped individual North
American genotypes up to 1015m in extent. This raises the possibility that the localities
described for the new taxa contain only single genotypes and/or individuals (see also Rumsey,
1998) rather than populations (colonies) of individuals.

The aim of this study was therefore to sample within the type areas cited as containing these
novel taxa of Pteridium in order to determine whether such localities comprise one or many
genetically distinct individuals, the morphology of which is consistent with published
descriptions. The prime focus was on localities occupied by the new species (P. pinetorum) and
the method used to detect individuals was the allozyme multi-locus genotyping technique
established by Sheffield et al. (1989). This technique allows an unequivocal statement of
whether two or more plant samples are genetically different, but it cannot provide an
unequivocal demonstration of genetic identity.

MATERIALS AND METHODS

All of the study sites listed herein have been pinpointed by Dr C.N. Page whilst in the field with
one of the authors. If collections were not made in the company of Dr Page, the same site was
revisited after precise notation of the exact location. At the type locality for P. pinetorum subsp.
pinetorum (grid ref. NH 869092) there is a continuous patch of Preridium; fifteen samples (one
pinna, basal or near basal) were taken (11.8.96) at approximately 5-10m intervals along a
transect (samples numbered P1-15). A sample was also taken from Pteridium adjacent to the
type locality which did not match the published description of P. pinetorum subsp. pinetorum
and was taken to be P. aquilinum subsp. aquilinum (sample P16) and from across a nearby
stream which appeared close to the published description of P. pinetorum subsp. pinetorum
(sample P17), see Figure 1.

At the P. pinetorum subsp. osmundaceum (Christ) C.N. Page type locality (grid ref. NN
778578) four discrete patches of Preridium were present; all were consistent with the published
description of P. pinetorum subsp. osmundaceum. Samples of basal/near basal pinnae were
taken at each patch (samples Os1-28), see Figure 2. At Patch 1, samples were taken across a
transect and a sample from every 4m was used (Os1-7); at Patch 2, samples were taken along a
transect through the patch and samples from every 4m were used (Os8-12); Patch 3 was very
narrow, so random samples were taken throughout, (Os13-22) and at Patch 4, samples were
taken on a transect and samples from every 8-10m were used (Os23-28).

A collection (Os29) was also made of Pteridium near the P. pinetorum subsp. osmundaceum
type locality, but in a different vegetation type (woodland) and also from the opposite side of
the road, but not within the woodland, see Figure 2. The sample from the opposite side of the
road resembled the Preridium plants in the P. pinetorum subsp. osmundaceum type locality.

Not all collected material was analysed isozymically. Both ends of the patches, and samples
along the transect (where applicable) were included in the current analyses, the remainder is in
storage at -80°C pending further biochemical and molecular analysis.

For P. aquilinum subsp. fulvum C.N. Page, one collection was made, (grid ref. NN 887603).
This site is on a steep slope near Loch Faskally, Perthshire and is one continuous patch. The
fronds in this patch are clearly distinct in terms of their morphology from the surrounding
Pteridium fronds, which were presumed to be P. aquilinum subsp. aquilinum. Material was
collected from an individual frond (matching the P. aquilinum subsp. fulvum description) at this
locality. Another (single) collection was made of Pteridium on the shores of Loch Rannoch (grid
ref. NN 589586). Plants from both sites were identified by Dr C.N. Page as P. aquilinum subsp.
fulvum. See Figure 3.

All frond samples were sealed in ziplock bags and stored at 4°C overnight before being sent
to Manchester where portions were put into storage at -80°C.

P. aquilinum (L.) Kuhn subsp. atlanticum C.N. Page has been the subject of previous
investigations following Page’s 1989 publication (see Rumsey ef al., 1991; Wolf et al., 1994),

208 FERN GAZETTE: VOLUME 15 PART 6 (1998)

Tummel Bridge

Patch 4 (Os23-28)

60m OA, OE,OF
<___>
30m Heather
moorland © ~
2 | &
Patch 1 101m S 0329
(Os1-7) 24m ros
OA, OB a
<_\_—__—___>
29m 60m
Patch 2 Patch 3
(Os8-12) (Os13-22)
OC D
Aberfeldy

|

Figure 2: P. pinetorum subsp. osmundaceum type locality, GR 778578, Perthshire,
Scotland.

OA-OF represent the different isozyme genotypes identified, and the measurements are the
greatest extent in the indicated orientation. All four patches are part of the type locality.

‘ulvumn

ee Loch Rannoch

Dunalastair Water

X '
(fd type locality
a {not sampled]
(osmundaceum type locality)

Figure 3: The location of sample sites for P. aquilinum subsp. fulvum and P. pinetorum
subsp. osmundaceum included in this study.

BRITISH PTERIDIUM 209

and is the subject of other publications (Ashcroft & Sheffield in press) so will not be considered
further here.

Isozyme analyses were carried out as per Soltis et al. (1983) with only minor modifications.
All samples were stained for: aconitase (ACON) EC 4.2.1.3; aldolase (ALD) EC 4.1.2.13;
aspartate aminotransferase (AAT) EC 2.6.1.1; glucose-6-phosphate dehydrogenase (G6PDH)
EC 1.1.1.49; glyceraldehyde-3-phosphate dehydrogenase (G3PDH) EC 1.2.1.12; hexokinase
(HK) EC 2.7.1.1; isocitrate dehydrogenase (IDH) EC 1.1.1.42; leucine aminopeptidase (LAP)
EC 3.4.11; malate dehydrogenase (MDH) EC 1.1.1.37; malic enzyme (ME) EC 1.1.1.40;
phosphoglucoisomerase (PGI) EC 5.3.1.9; phosphoglucomutase (PGM) EC 2.7.5.1;
6-phosphogluconate dehydrogenase (6PGD) EC 1.1.1.44; shikimate dehydrogenase (SKDH)
EC 1.1.1.25; triosephosphate isomerase (TPI) EC 5.3.11.

All gels were run with marker wicks of S48 (British P. aquilinum subsp. aquilinum). This
Pteridium plant grows at the University of Manchester’s Experimental Grounds in Fallowfield,
Manchester.

Herbarium specimens of material used in this study have been deposited at the Manchester
Museum Herbarium (MANCH).

RESULTS
No variation was detected for: AAT, ACON, ALD, F16D, G3PDH, G6PDH, HK, LAP, MDH-
2, ME, PGI-1, PGM, 6PGD, TPI-1 in any of the samples analysed in this study. Variability was
detected for IDH, MDH-1, MDH-3, PGI-2, SKDH, TPI-2.

Five isozyme genotypes were found at the P. pinetorum subsp. pinetorum site: (PA-E), see
Tables 2 and 3, Figure 1. Five were for samples taken as P. pinetorum subsp. pinetorum (P1-15,
17); the frond sampled as being P. aquilinum subsp. aquilinum (P16) matches the isozyme
genotype of P14 and P15.

Table 2: Genotypes found at the P. pinetorum subsp. pinetorum site

Sample number Genotype
P1-7 PA

P8 PB (=OsA)
P9 PC

P10-11 PD (=OsD)
P12-16 PB (=OsA)
Ply PE

At the P. pinetorum subsp. pinetorum site P17 (the sample taken from across the stream,
morphologically similar to P. pinetorum subsp. pinetorum) does not have the same genotype as
any of the other samples, but P16 (sample from frond of standard P. aquilinum subsp.
aquilinum appearance) shares the same isozyme profile as P12-15. Four different genotypes
were found in the continuous patch of bracken at the P. pinetorum subsp. pinetorum type locality
(samples P1-15).

Four isozyme genotypes were found at the P. pinetorum subsp. osmundaceum site: (OsA-
D), see Tables 4 and 5 (Figure 2). Three were for fronds sampled in patches 1-4 (Os1-28) and
one was for the sample taken from across the road (Os29).

The different P. pinetorum subsp. osmundaceum genotypes do not directly correspond to the
four patches sampled. That is, there is some overlap of genotypes between patches. (See Figure
2). Samples Os24 and Os26 (within Patch 4) have the same genotype as samples Os1-6 (within
Patch 1). Patch 1 has two genotypes; Patch 2 has one; Patch 3 has one, and Patch 4 has three
genotype. Os29 is the sample from across the road, this sample has the same genotype as Os14-
22 (Patch 3).

210 FERN GAZETTE: VOLUME 15 PART 6 (1998)

The two P. aquilinum subsp. fulvum samples (from near Loch Faskally and Loch Rannoch) were
the same as each other and different from every other sample analysed here.

Table 3: Zymogram of P. pinetorum subsp. pinetorum samples, for variable isozymes
only. Asterisks denote heterozygous individuals.

Table 4: Genotypes found at the P. pinetorum subsp. osmundaceum site

Sample number Patch number Genotype
Os1-6 1 OsA (=PB)
Os7 I OsB
Os8-12 253 OsC
Os13-22 8) OsD (=PD)
Os23 4 OsE
Os24-28 4 OsA

Os29 : OsD (=PD)

Comparison of isozyme profiles between type sites.

Each type locality contained some genotypes identical for all isozymes tested to genotypes
found elsewhere. Genotype OsA (P. pinetorum subsp. osmundaceum samples Os1-6 and Os 24-
28) is the same as PB (P. pinetorum subsp. pinetorum samples P8 and P14-16). OsD
(P. pinetorum subsp. osmundaceum samples Os 13-22 and Os29) has the same isozyme genotype
as PD (P. pinetorum subsp. pinetorum samples P 10-11). See Tables 2 & 3. The isozyme profile
of plants referred to as P. aquilinum subsp. fulvum is identical to that of a plant sampled
elsewhere in Scotland (GR 972511), not the location of a Pteridium type locality, analysed in a
parallel study (Ashcroft, unpublished data).

DISCUSSION
Clonal pteridophytes present special problems for population analysis because individuals
(genets) generate many morphological units (ramets). The identification of Pteridium genets
within populations relied until recently upon morphological differences between individuals
(e.g. Webster & Steeves, 1958; Oinonen, 1967). Such identification methods resulted in

BRITISH PTERIDIUM 11

Table 5: Zymogram of P. pinetorum subsp. osmundaceum samples, for variable
isozymes only.
Asterisks indicate heterozygous individuals.

underestimates of clonal size (Parks & Werth, 1993), and have been superseded by methods
including multilocus allozyme genotyping (Sheffield et al., 1989; Parks & Werth, 1993). The
latter method reveals differences in the structure of only a limited number of proteins and cannot
necessarily detect every genet, but genotypes that are revealed as distinct by the method do most
certainly represent individuals that are genetically different from each other. It is particularly
important to employ such methods where forms of a pteridophyte are reported to be correlated
with edaphic parameters, as reported for the new Pteridium taxa (Page, 1997). The influence of
a particular set of conditions, for example, soil, on a single genotype could generate atypical
morphology of an otherwise standard form.

Pteridium has a ‘guerrilla-type’ clone-forming growth strategy, which results in intergrowth
amongst genets. Patches cannot be assumed to consist of single genets, nor are boundaries
between genets sharp (Parks & Werth, 1993). As forms of Pteridium are thought to be associated
with particular sets of environmental parameters, it is crucial to determine whether such forms
represent ramets of neighbouring genets of the common taxon (P. aquilinum subsp. aquilinum)
which appear different only because of environmental pressures, or distinct genets.

272 FERN GAZETTE: VOLUME 15 PART 6 (1998)

The data presented herein do not support the hypothesis that the new taxa of bracken are
based upon environmentally-modified extensions of neighbouring individuals fitting the
descriptions of the common taxon of the genus. Each area occupied by plants ascribed to new
taxa was found to contain one or more individuals genetically distinct from the surrounding
bracken. This study has used more enzyme loci than many studies, and it is highly unlikely that
the genotypes found with one or more heterozygous loci (the majority of the plants sampled)
would be repeated, as bracken is known to be an outcrossing species (Sheffield, Wolf & Haufler
1989; Wolf et al., 1988). Thus, their distinctive morphology cannot merely reflect perturbations
of the morphology of clones surrounding these type areas which have grown into the habitats in
question. These plants might of course, share an environmentally-induced phenotype. The areas
identified as typifying the new taxa of the species P. pinetorum investigated in the present study
do contain more than one individual matching the published description of the taxa. This means
that the correct description of the P. pinetorum-inhabited areas described herein, and in Page
1997, is indeed “colonies” of individuals, 1.e. populations, and not large single clones.

Spatial analysis of bracken genets based on reliable data is still in its infancy, and few
studies have been published, but the current study is in broad agreement with others in finding
that Pteridium populations comprise both small and large individuals. Growth rates for the
species are similarly under-researched, but range from 25cm to over 100 cm per year for the
P. aquilinum subsp. aquilinum taxon in one report (Sheffield et al., 1989). Given the northerly
locations of the site localities and a growth rate which 1s likely to be lower than that of the weedy
form, it does seems reasonable to conclude that some of the individuals detected herein
represent clones of considerable antiquity. Estimates of age of clones have been based on very
different models of plant growth (see Sheffield et al., 1993 for full account) but however
conservative the estimate, it seems possible that the individuals detected in this study have
survived for a substantial time in the face of competition from neighbouring P. aquilinum subsp.
aquilinum individuals (the most aggressive of the Pteridium taxa; Page, 1997). It could also of
course indicate that the P. aquilinum subsp. aquilinum taxon has only recently ventured into the
neighbourhood. Only time, and further sampling in decades to come, would determine the
correct interpretation of the current situation at these sites.

Finally, it should be noted that although the type areas of the new P. pinetorum taxa
comprise populations and not merely single individuals, the isozyme profiles detected were not
unique for each taxon. In each population there were individuals with isozyme profiles identical
to individuals in populations ascribed to a different taxon. For P. pinetorum subsp. fulvum, no
other individuals were detected with the same profile as the profiles recorded during the present
study, but a parallel study has located individuals with this genotype that matched the
description of P. aquilinum subsp. aquilinum. This does not necessarily question the validity of
taxonomic recognition for these taxa, as there are organisms for which isozymes do not reflect
taxonomic distinctions, but taxonomic considerations were not the focus of the current study
and are the subject of ongoing research which includes common-garden cultivation of
specimens collected at the study sites listed here.

ACKNOWLEDGEMENTS
The authors would like to thank Dr J. Bridges, Dr R.S. Callow, C. Ellis, Dr C.N. Page, S. Ross
and J. Usherwood for help collecting material, and S. Challinor for technical assistance.
Funding was provided by a BBSRC studentship to KMB and a NERC studentship to CJA.

BRITISH PTERIDIUM 213

REFERENCES

ASHCROFT, C.J. & SHEFFIELD, E. In press. Rejection of Pteridium aquilinum subspecies
atlanticum (C.N. Page). Bot. J. Linn.Soc.

OINONEN, E. 1967. Sporal regeneration of Bracken (Preridium aquilinum (L.) Kuhn) in Finland
in the light of the dimensions and the age of its clones. Acta Forrest. Fenn. 83: 1-96.

PAGE, C.N. 1989. Three sub-species of Bracken, Pteridium aquilinum (L.) Kuhn, in Britain.
Watsonia 17: 429-434.

PAGE, C.N. 1997. The ferns of Britain and Ireland. 2nd ed. Cambridge University Press,
Cambridge.

PAGE, C.N. & MILL, R. 1995a. Scottish Bracken (Pteridium): new taxa and a new
combination. Bot. J. Scot. 47: 139-140.

PAGE, C.N. & MILL, R. 1995b. The taxa of Scottish Bracken in a European perspective. Bot.
J. Scot. 47: 229-247.

PARKS, J. & WERTH, C. 1993. A study of spatial features of clones in a population of bracken
fern, Pteridium aquilinum (Dennstaedtiaceae). Amer. J. Bot. 80: 537-544.

RUMSEY, FJ. 1998. Review of Page, C.N. Ferns of Britain and Ireland 2nd ed., Cambridge
University Press. Ann. Bot.(Oxford) 81: 297-301.

RUMSEY, FJ., SHEFFIELD, E. & HAUFLER, C.H. 1991. A re-assessment of Pteridium
aquilinum (L.) Kuhn in Britain. Watsonia 18: 297-301.

SHEFFIELD, E., WOLF, P.G. & HAUFLER, C.H. 1989. How big is a bracken plant? Weed Res.
29: 455-460.

See. ES WOLF, P-G., RUMSEY, FJ., ROBSON, D., RANKER, T.A. &
CHALLINOR, S.M. 1993. Spatial distribution and reproductive behaviour of a triploid
bracken (Pteridium aquilinum) in Britain. Ann. Bot.(Oxford) 72: 231-237.

SOLTIS, D., HAUFLER, C.H., DARROW, D. & GASTONY, G. 1983. Starch gel
electrophoresis of ferns: a compilation of grinding buffers, gel and electrode buffers, and
staining schedules. Amer. Fern J. 73: 9-27.

WEBSTER, B., & STEEVES, T.A. 1958. Morphogenesis in Pteridium aquilinum (L.) Kuhn -
general morphology and growth habit. Phytomorphology 8: 30-41.

WOLF, P.G., HAUFLER, C.H. & SHEFFIELD, E. 1988. Genetic variation and the mating
system of the clonal weed Pteridium aquilinum (L.) Kuhn. Evolution 42: 1350-1355.

WOLF, P.G., SHEFFIELD, E., THOMSON, J.A. & SINCLAIR, R.B. 1994. Bracken taxa in
Britain: a molecular analysis. In: Smith, R.T. & Taylor, J.A., (eds). Bracken: an
environmental issue, pp 16-20. International Bracken Group, Aberystwyth.

214 FERN GAZETTE: VOLUME 15 PART 6 (1998)

BOOK REVIEWS

AUTHORS OF SCIENTIFIC NAMES IN PTERIDOPHYTA R. E. G. Pichi Sermolli. 1996.
xxx + 78 pp. Royal Botanic Gardens, Kew. Price £12. ISBN 0 947643 90 7. Softback.

Every pteridophyte taxonomist should have their own copy of this book! For starters it is more
manageable than that fine source of author abbreviations Authors of Plant Names by Brummitt
and Powell (1992), to which Professor Pichi Sermolli contributed the main list of pteridophyte
authors, and for seconds it contains an additional 401 names. It consists of a revised and updated
list of pteridophyte authors and their abbreviations in the standard forms recommended by
Authors of Plant Names, and an appendix of author abbreviations adopted in recent publications
which differ from those used in APN and in this book together with their standard form
equivalent. Needless to say it has been compiled with the accuracy and painstaking attention to
detail we have come to expect from Professor Pichi Sermolli. The introduction is a fascinating
read in its own right. In the section dealing with the citation and spelling of authors’ names I
particularly enjoyed the long discussion on Chinese names and the Wade-Giles and Pin-yin
conventions for their romanisation. This slender volume is an absolutely indispensible tool for
the pteridologist monographer and Flora writer.

B.S.Parris

THE FERNS OF BRITAIN AND IRELAND C.N. Page. Continued from page 204.

... If the general reader wants to learn about pteridophytes in the British Isles, Page’s ‘field
notes’ provides a lot of information, but how does one judge what to take as an authoritative
statement, and what with a pinch of salt. A good solution to this problem would be to test his
observations. One can only encourage the many fern enthusiasts in Britain and Ireland to take
the ‘field notes’ as an inspiration to go out for one’s self and to test Page’s hypotheses by
observing and experimenting. It might help to raise awareness out in the field and foster your
scientific urges to prove Page right or wrong, but please do not forget to communicate your
findings to the wider pteridological community by publishing in the Fern Gazette and other
journals. It might help Chris Page with his third edition.

This book is a good and valuable read and it is the most comprehensive book on British and
Irish ferns. As these Isles harbour a wealth of species the book is of great interest to fern
enthusiasts from abroad. If you do not have the first edition, this book is a must. However,
concerning biosystematics, progress in pteridology, and even additions to the British fern flora,
the book rather reflects the state of knowledge of perhaps 1990, but certainly not that of 1997,
the year it was published. The hardbound copy is £95, and this is a lot of money for a book on
ferns (paperback £40). Despite all the good qualities this book certainly has, its second edition
strikes me as a missed opportunity. The first edition was an outstanding piece of work but
suffered from the poor reproduction of the fern specimen pictures, and this certainly has been
rectified in the second edition. However, there were other areas where improvements would
have to be made in order to stay ahead. The claim that the book has been fully updated is not
supported by the contents. Sorry Chris, if I want to hold THE authoritative book of ferns in
Europe in my hands, I will grab the Hegi.

J.C. Vogel

FERN GAZ. 15(6) 1998 Zn

THE ADDITION OF ACROSORUS STREPTOPHYLLUS
(BAKER) COPEL. (GRAMMITIDACEAE: PTERIDOPHYTA) TO
THE FLORA OF THAILAND

B.S. PARRIS
Fern Research Foundation, 21 James Kemp Place,
Kerikeri, Bay of Islands, New Zealand

Key words: Acrosorus streptophyllus, Grammitidaceae, Thailand.

ABSTRACT
Acrosorus streptophyllus (Grammitidaceae: Pteridophyta) is reported for the first time from
Thailand.
INTRODUCTION

Tagawa & Iwatsuki (1989) report only one species of the fern genus Acrosorus
(Grammitidaceae) for Thailand. They record A. triangularis (Scort. ex Bedd.) Copel. for three
regions, Surat Thani (Khao Nong), Nakhon Si Thammarat (Khao Luang) and Yala (Gunong
Ina), in Peninsular Thailand. The correct name for this species is A. friderici-et-pauli (H. Christ)
Copel., syn. A. triangularis Copel., a name based on Polypodium triangulare Scort. ex Bedd.
(1887) non J.F. Gmel. (1791) nec Dulac (1867). Tagawa & Iwatsuki (1989) mention that another
species, A. streptophyllus (Baker) Copel., is common in Malaya and Borneo and distinguish it
from A. friderici-et-pauli by size of fronds, texture and soral construction without including it
in the Flora.

Material of Acrosorus from Peninsular Thailand held in the herbaria of the Botany School,
University of Cambridge (CGE) and the Royal Botanic Gardens, Kew (K) has been examined
and found to contain both A. friderici-et-pauli and A. streptophyllus.

MATERIALS
Acrosorus streptophyllus specimens examined: Thailand: Peninsular; Yala (Gunung Ina) [Malay
Peninsula, Perak, Gunung Inas], epiphytic on jungle trees, 3900 ft, 6 Dec. 1899, R.H. Yapp 405
(CGE p.p., K) and 405 bis (CGE). Yapp 405 at CGE is a mixture of A. friderici-et-pauli and
A. streptophyllus.

KEY TO ACROSORUS IN THAILAND
Lamina 80-480 x (4-)6-9mm, texture thickly coriaceous, sori deeply sunken in pouches in
lamina A. friderici-et-pauli
Lamina 40-200 x 3-6mm, texture thinly coriaceous, sori protected by folded lamina
A. streptophyllus

DISCUSSION
Holttum (1968) describes and illustrates both Acrosorus streptophyllus and A. friderici-et-pauli
(as A. triangularis).

Tagawa & Iwatsuki (1989) report the ecology of A. friderici-et-pauli as ‘on mossy tree
trunks in dense evergreen forests on ridges at medium or high altitudes’. Evidently both species
occur in similar habitats in Thailand, as in Peninsular Malaysia (Gunung Beremban), Sabah (Mt
Kinabalu) and Sarawak (Gunung Mulu), where I have collected both growing as low epiphytes
in low or open ridge crest lower montane and mid-montane forest between 1500 and 1900 m alt.
Acrosorus streptophyllus should be searched for in Peninsular Thailand in the localities where
A. friderici-et-pauli has been found.

216 FERN GAZETTE: VOLUME 15 PART 6 (1998)

Gunung Ina (Inas) was formerly included in the state of Perak in northern Peninsular
Malaysia, but is now part of southern Peninsular Thailand. It is possible that old collections
from areas formerly included in northern Perak, like these of Yapp, are now wrongly filed under
Malaysia in some herbaria and that further new records for the flora of Thailand remain to be
discovered.

REFERENCES
HOLTTUM, R.E. 1968. Revised Flora of Malaya Vol. 2 Ferns. 2nd ed. Government Printing
Office, Singapore.
TAGAWA, M. & IWATSUKI, K. 1989. Flora of Thailand Vol. 3 Part 4. Chutima Press,
Bangkok.

BOOK REVIEWS

DISTRIBUTION, ECOLOGY AND CONSERVATION STATUS OF
CLUBMOSSES (DIPHASIASTRUM SPECIES) IN LOWER SAXONY AND BREMEN
(in German). Karsten Horn. 1997. Naturschutz und Landschaftspflege in Niedersachsen, Heft
38, 83 pages. Erlangen. Available from the Niedersachisches Landesamt fiir Oekologie, Abt.2,
An der Scharlake 39, D-31135 Hildesheim.

This book is a beautifully illustrated and well researched study on the distribution, ecology and
conservation status of Diphasiastrum taxa in the north western parts of Germany. It is, as stated
on the inside cover, a slightly altered version of the Master’s thesis by Karsten Horn submitted
in 1997 and supervised by Professors H.W. Bennert (Bochum) and W. Nezadal (Erlangen).

The book has six major parts: morphology and systematics of clubmosses, introduction to
the area, material and methods, results, discussion and recommendation for the conservation of
clubmosses. It starts off with comprehensive introductions to the geography, climate, geology,
soil types and natural vegetation of the area, as well as a collation of many interesting facts
about the ecology and biology of Diphasiastrum.

Large parts of Lower Saxony are relatively flat with sandy soils prevailing (good for
D. tristachyum and D. zeilleri), but it also includes mountainous areas rising up to over 1,000
metres, in the Harz mountains. The latter have the greatest diversity, with all five taxa
(D. alpinum, D. issleri, D. complanatum, D. tristachyum, D. zeilleri) still present. The
geographical variation of the overall area makes for a rich clubmoss flora. The current and
historic distribution of all taxa is meticulously documented and is based on several years of field
work and herbarium studies. These detailed studies allow the author to investigate the decline
in number and abundance of clubmosses in Lower Saxony and to determine some of the causes
of this decline. The colonisation of new sites by clubmosses normally follows disturbance. As
poor competitors they are forced out when other vegetation becomes established in the area.
Change of land use, e.g. heathland, which no longer include regular disturbance, have
contributed to the decline. Therefore, many well stocked clubmoss assemblies are nowadays
confined to ski slopes or military areas. In addition, collecting for commerical purposes or
herbaria, as well as liming of forests, may have contributed to the loss of populations.

From this overview it becomes clear that Diphasiastrum has been, and still is, benefitting
from or even depending on, human interference. This in turn must raise the question of where
Diphasiastrum may have had natural sites in the area investigated, and whether protection
measures are actually desirable for an organism following Man’s activities? It is stated by the
author that four taxa of Diphasiastrum are at the edge of their distribution in Lower Saxony, but
to conclude from this there is a need for special protection is not the only possible inference

FERN GAZETTE: VOLUME 15 PART 6 (1998) ily

especially since their survival depends on wintersports activities. While the causes for decline
are well documented by Horn, a forward-looking conservation strategy remains only
- rudimentary.

The section about the biosystematics of Diphasiastrum was a disappointment for me. Horn
enthusiastically states “that the biosystematics of Diphasiastrum in Europe has finally been
resolved” with the publication of a recent study, referring to Stoor et al. (1996), which he co-
authored. However, the “evidence” on which this claim is based cannot be accepted. Horn
(1997) and Stoor et al. (1996) present the reader with a triangle of three ancestral taxa and their
three “hybrids”. Their scheme is based on biochemical evidence derived from allozyme
eletrophoresis of only two enzyme systems. The evidence for the hybrid origin of some taxa is
based on “additive” allozyme banding patterns. While the use of “additive”, and fixed, banding
patterns has been applied for determining the diploid ancestry of allopolyploid taxa (Werth,
1989), its application in biosystematic studies cannot be extended to homoploid taxa. If a
Diphasiastrum plant is the result of outbreeding and cross-fertilisation, either within or between
taxa, the Fi Diphasiastrum plant will show additive banding patterns if the parental plants have
different allelic forms in the locus investigated. However, taken the homoploid status and the
fertility of putative hybrid Diphasiastrum into consideration, allozyme patterns will segregate
in the F2 in Mendelian ratios. Therefore additive bands will not be fixed in hybrids between taxa
and the use of allozymes as a biosystematic marker in Diphasiastrum might be limited.
Furthermore, it is stated that “esterase is the most useful enzyme for unequivocally determining
the identity of Diphasiastrum species” (Horn - in Stoor et al., 1996, p.152; emphasis added).
Esterases are plentiful in plants with two to ten loci, and rarely is a genetic basis available to aid
interpretation (Weeden & Wendel, 1989). Therefore, it is not, or very rarely, used in systematic
studies. The esterase picture published by Stoor et al. (1996) shows two gels pasted together and
therefore do not represent the desirable side-by-side within-gel comparison. Furthermore, no
genetic interpretation for the picture is given. It is well known that esterases can be tissue
specific, that their activity may be environmentally induced, and that the absence of breeding
and hence segregation data “can lead to grossly misleading conclusions” (Oxford, 1973).
Taking the arguments above, I fear that we are a long way off a satisfactory answer to the
puzzling biosystematics of Diphasiastrum. Molecular methods can certainly play a part in
solving the riddle but caution should prevail in the light of insufficient data sets and data

analysis.
Despite the criticism on a small, but important, part of the book, I congratulate the author

on his work about the ecology and distribution of Diphasiastrum in northern Germany. There is
much need for such comprehensive studies on European pteridophytes. If you can, try to get a
copy. The detail and effort presented in this book shows that Karsten Horn loves “his
organisms”, but such love should not impair his vision.

J.C. Vogel

REFERENCES:

OXFORD, G.S. (1973) The genetics of Cepaea esterases. 1. Cepaea nemoralis. Heredity
30:127-139.

STOOR, A.M., BOUDRIE, M., JEROME, C., HORN, K, & BENNERT, H.W. 1996.
Diphasiastrum oellgardii (Lycopodiaceae, Pteridophyta), a new lycopod species from
central Europe and France. Feddes Repert. 107: 149-157.

WEEDEN, N.F. & WENDEL, J.F. 1989. Genetics of plant isozymes. In: Soltis, D.E. & Soltis,
P.S. (eds). Isozymes in Plant Biology, pp. 46-72. Dioscorides Press, Portland.

WERTH, C.R. (1989). The use of isozyme data for inferring ancestry of polyploid
pteridophytes. Biochem. Syst. Ecol. 17: 117-130.

218 FERN GAZETTE: VOLUME 15 PART 6 (1998)

INDEX FILICUM SUPPLEMENTUM SEXTUM PRO ANNIS_ 1976-1990.
R. J. Johns. 1996. x + 414 pp. Royal Botanic Gardens, Kew. Price £30. ISBN 1 900347 11 3.
Softback.

Index Filicum and Supplements 1-3 were prepared by Carl Christensen and incorporated his
opinions on the taxonomic status and generic and specific synonymy of ferns. Each contained
additional material at the end of the volume in sections termed Addenda and/or Corrigenda
which were inconvenient to use and sometimes ignored by consulting pteridologists unaware of
their existence. Supplement 4 presented the names as published, without taxonomic judgment,
as did Supplement 5, which included fern allies for the first time. Supplements 4 and 5 dealt
explicitly only with validly published names, included the details of publication of infraspecific
basionyms and, to the delight of users, did not include Addenda/Corrigenda sections.
Supplement 6, like its two predecessors presents the names as published, without taxonomic
judgment and, like Supplement 5, includes names of fern allies. Entries for Supplement 6 were
compiled by the Index Kewensis staff, with associated nomenclatural problems from 1976 to
1980 solved by Frances Jarrett and those from 1981 to 1989 dealt with by Barbara Parris. A
useful innovation in the latest supplement is the inclusion of new infra-specific taxa for the first
time. Some infra-specific taxa published before 1976 are also included in the main part, e.g.
Polystichum munitum subsp. imbricans, while many more are included in the Addenda etc. A
useful additional inclusion would have been the card index held at Kew of infra-specific
pteridophyte taxa compiled for the later part of the years covered by Supplement 5. Infra-
specific basionyms have been included without their place of publication, however, contrary to
the policy of Supplements 4 and 5, e.g. Aspidium munitum var. imbricans, Dryopteris sciaphila
vars rapaensis and raivavensis, Lycopodium miyoshianum var. coreanum and Polystichum
lobatum var. ruwenzoriense.

Unfortunately Supplement 6 differs in some other ways from the two preceding
supplements and appears rather muddled in concept and confused in execution, particularly the
Addenda etc. Addenda etc. were a feature of Index Filicum and Supplements 1-3, but were
dropped from Supplements 4 and 5. In these last two supplements names dating from before the
starting dates (1934 and 1961 respectively) which were not included in the original Index
Filicum nor in preceding supplements were included in the body of the work, but preceded by
an asterisk to distinguish them from names published in the years covered by the supplements.
Thus they are clearly visible to the reader without the need to consult a separate section. It is
disappointing to find an Addenda etc. tacked on to the end of Supplement 6 behind the
Catalogus literaturae almost as an afterthought. I do not understand why validly published
names are included here rather than in the body of the text. Unfortunately the new names in the
Addenda have no distribution, unlike those elsewhere in the volume. Additional data and
omissions from Supplement 5 are not all in the Addenda by any means. A sample of pre-1976
names in the main part of the Supplement (without asterisk) includes Aspidium lobatum (later
homonym), Ctenopteris khasyana (earlier combination), and Alsophila and Sphaeropteris spp.
(omitted from Supplement 5). Carpogymnia dryopteris, Huperzia perrieriana and Lunathyrium
lasiostipes are included in Supplement 5 and repeated in Addenda to Supplement 6 with the
same reference. Goniophlebium manmiense and Loxogramme tibetica are in both the main body
of text to Supplement 6 and in the Addenda.

The inclusion of invalidly published names, both in the body of the text and in the Addenda
etc., differs from previous supplements and is a source of confusion. If this is a standardisation
of Index Filicum to the Index Kewensis format and will be a feature of future supplements a note
to this effect would have been useful. The Introduction notes that Ching published some of his
species several times in different publications and these have been listed so that the
monographer has access to all these published names (even if the species had already been
validly published). As one of that rare breed, a fern monographer, who has consulted /ndex

FERN GAZETTE: VOLUME 15 PART 6 (1998) 219

Filicum for more than thirty years, I am not interested in anything other than validly published
names and am not convinced that multiple citation adds anything other than padding.

There are some careless errors in Supplement 6. Some names have been placed in the wrong
genera e.g. Selenodesmium bellisquamata, S. plebiscopa, S. werneri should be under Selliguea.
Thelypteris remotipinna is correctly spelt in the main body of text, but appears in Addenda etc.
with the same authors and place of publication as T. remotifinna, likewise Mecodium coreanum,
included in Supplement 5, is misspelt as Mecodium doreanum in Addenda etc. the details of
author, place and date of publication are identical. Compounded epithets with hyphen, contrary
to the /nternational Code of Botanical Nomenclature Article 60.9, have been included e.g.
Crypsinopsis albido-paleata and Hypodematium squamuloso-pilosum. One might have
expected a publication from Kew to have authors’ names consistent with the excellent Authors
of Plant Names by Brummitt and Powell, but Hermann Christ is consistently cited as ‘Christ’
rather than H.Christ as recommended by Brummitt and Powell and Lev. has been used instead
of H.Lév. There are also inconsistencies in the citation of place of publication. Proctor’s Ferns
of Puerto Rico and the Virgin Islands is cited just as Mem. New York Bot. Gard. 53 (1989) with
the journal reference included in the Catalogus liteaturae, but Mickel and Beitel’s Pteridophyte
flora of Oaxaca, Mexico is cited as ‘Pterid. Fl. Oaxaca (Mem. New York Bot. Gard. 46 (1988))’
and the journal title is not mentioned in the Catalogus literaturae, while Brownlie’s
Pteridophyte flora of Fiji is cited as “Nova Hedwigia Beth. 55 (Pterid. Fl. Fiji) (1977). Hybrids
are indicated by x in front of their specific epithet, but not for Leptopteris x intermedia (André)
Brownsey. The paper where this combination is made is missing from the Catalogus literaturae;
it is: Brownsey, P. J. (1981). A biosystematic study of a wild population of Leptopteris hybrids
in New Zealand. New Zealand J. Bot. 18: 343-352. The mistakes listed above were found during
an hour’s browse and doubtless many others await discovery.

Although it is not obvious at first sight, Supplement 6 differs from its predecessors in a very
important point, which has changed the way in which I use Index Filicum. The original and
previous five supplements included references to names when they were first combined, so
looking up a particular combination was quite straight-forward one started with the most recent
supplement and worked back until finding it (or not). Supplement 6 includes the same
combinations but made by different authors or in different references, and thus introduces an
unexpected level of confusion. Some names are included both in Supplement 5 and Supplement
6 with different authors and/or references, e.g. Grammitis cavisora, Plesioneuron fulgens,
Pronephrium lakhimpurense, P. nudatum, P. penangianum, P. triphyllum, Pseudocyclosorus
ochthodes; those in Supplement 6 are superfluous and should not have been included. Again,
there is no explanation for the change of policy in Supplement 6. Perhaps the editor was
unaware of the careful checking carried out by editors of earlier supplements of Index Filicum
to ensure that combinations were not duplicated unnecessarily. Do not expect to browse back
through the /ndex Filicum supplements to find the name you are seeking. Now you will have to
begin at the original, with its Addenda and Corrigenda (before the Catalogus literaturae), and
progress to Supplement 1 and Corrigenda (after the Catalogus literaturae), Supplement 2 and
Corrigenda (after the Catalogus literaturae), Supplement 3 and Addenda et Corrigenda (before
the Catalogus literaturae), Supplements 4 and 5 (mercifully without Addenda and/or
Corrigenda) and Supplement 6 with Addenda etc. There are numerous names cited twice in
Supplement 6, e.g. Ctenitis tonduzii, Deparia polyrhizon, Grammitis poeppigiana, G. rigescens,
Huperzia kamaensis, H. nylamensis, Palhinhaea cernua var. sikkimensis, Xiphopteris
bryophylla. Only the earliest citation should be included as all others are nomenclaturally
superfluous. At least in Emendata the later combinations are marked ‘non’!

All taxonomic pteridologists will need Index Filicum Supplement 6, which contains much
useful information, but I am sorry to see that the high standards of earlier supplements have not
been maintained and look forward to the day when the /ndex Filicum data is available on CD-

rom and more easily accessible than at present.
B.S. Parris

220 FERN GAZETTE: VOLUME 15 PART 6 (1998)

INSTRUCTIONS FOR AUTHORS

Manuscripts on all subjects of pteridology are welcome and should be sent to the editor:
Miss J.M. Camus, Department of Botany, The Natural History Museum, Cromwell Road,
London SW7 5BD, UK [fax +44 - (0)171- 9260, e-mail j.camus@nhm.ac.uk].

PLEASE NOTE that papers should not usually exceed 20 printed pages and are generally
expected to be considerably shorter. Subject review articles will be considered. Short notes are
acceptable e.g. new records. The senior author should supply fax and e-mail numbers if possible
to facilitate correspondence.

MANUSCRIPTS should be written in English (British), double spaced, preferably on A4 paper.

Three hard copies are required in the first instance; a version on disc will be required after

revision (WORD in WINDOWS 95, DOS preferred). All manuscripts will be refereed. The title

should reflect the content of the paper and be in BOLD CAPITALS and centrally aligned.

Generic and specific names should be in italics and any title containing a generic or specific

name must be followed by the family and Pteridophyta in brackets e.g.

TRICHOMANES SPECIOSUM WILLD. (HYMENOPHYLLACEAE:

PTERIDOPHYTA) IN SOUTHERN SPAIN

Author abbreviations should follow Pichi Sermolli’s (1996) Authors of scientific names in

Pteridophyta, Royal Botanic Gardens, Kew.

MAIN HEADINGS: in BOLD CAPITALS and centrally aligned.

SUBSIDIARY HEADINGS: in italics and left aligned.

AUTHORS’ NAMES AND FULL ADDRESSES: following the title and centrally aligned.
KEY WORDS: up to five.

ABSTRACT: should reflect the content of the paper.

FIGURES: there is no distinction between photographs, line drawings and maps in numbering
and all should be presented in a form ready for reproduction. They should be of the required
magnification or larger with a scale bar where appropriate. Lettering or numbers (Arabic) should
be sufficiently large to be legible if reduction is necessary during printing. The number of
photographs allowed in any one issue is limited by cost. Figure captions should be on a separate
sheet.

TABLES: can be printed in either portrait or landscape format. Authors should consider this
when they are preparing tables. If possible tables should be presented in a form that can be
reproduced (like a figure).

MEASUREMENTS: should follow the metric system.

REFERENCES: should follow the style of this issue of The Fern Gazette, e.g.:-

HOOKER, W.J. 1864. Species Filicum, 5. Dulau & Co., London.

PICHI SERMOLLI, R.E.G. 1977. Fragmenta pteridologiae VI (Crypsinopsis). Webbia 31: 240-
242.

STEVENSON, D.W. & LOCONTE, H. 1996. Ordinal and familial relationships of
pteridophyte genera. In: CAMUS, J.M., GIBBY, M. & JOHNS, R.J. (eds) Pteridology in
perspective, pp. 435-467. Royal Botanic Gardens, Kew.

Journal abbreviations should follow Botanico Periodicum Huntianum & Supplements.

Alterations from the original text at proof stage will be charged for unless they are minor points
of detail. Twenty-five offprints will be supplied free to the senior author. Additional copies may
be ordered at the page proof stage.

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should be sent to:

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_ Department of Botany

The Natural History Museum
Cromwell Road

London SW7 5BD

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The British Pteridolog :

THE FERN GAZETTE

VOLUME 15 PART 6

CONTENTS

MAIN ARTICLES

Trichomanes speciosum Willd. (Hymenophyllaceae: Pteridophyta)
in southern Spain
FJ. Rumsey & J.C. Vogel

Population analysis of the type localities of some recently recognised taxa
of British Pteridium (Dennstaedtiaceae: Pteridophyta)

K.M. Bridges, C.J. Ashcroft & E.Sheffield
The addition of Acrosorus streptopnyllus (Baker) Copel to the

flora of Thailand
B.S. Parris

BOOK REVIEWS

INSTRUCTIONS FOR AUTHORS

1998

Page

197

205

215

204, 214,
216, 218

220

THE FERN GAZETTE Volume 15 Part 5 was published on 15th December, 1997

Published by THE BRITISH PTERIDOLOGICAL SOCIETY, c/o Department of Botany,

The Natural History Museum, London SW7 5BD

ISSN 0308-0838

Printed by Metloc Printers Ltd, Caxton House, Old Station Road, Loughton, Essex IG10 4PE