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FERN GAZ. 18(1):1-26. 2007 MV
THE SPECIES AND SUBSPECIES IN THE DRYOPTERIS AFFINIS
GROUP

C.R. FRASER-JENKINS

Student Guest House, Thamel, P.O. Box no. 5555, Kathmandu, Nepal
(chrisophilus @ yahoo.co.uk)

Key words: Dryopteris affinis, Europe, taxonomy, specific rank, subspecies, hybrids

ABSTRACT

Following discussion of the different levels of variation and the ranking of the
taxa, “morphotypes” have been replaced with formal names. The main taxa
within the Dryopteris affinis aggregate in Europe, formerly treated as subspecies
of D. affinis, are treated as six species, while the former geographical varieties
are treated as nine subspecies belonging to three of the species. Seven new
combinations are made at the subspecific rank and two new subspecies are
described. A new species-name, D. iranica Fras.-Jenk. replaces D. wallichiana
subsp. coriacea (Fras.-Jenk.) Fras.-Jenk. Four hybrids between the D. affinis
agg. and D. filix-mas are treated at the nothospecific rank, two of them new and
a third a new combination. Two of the hybrids are divided into six
nothosubspecies, three of them new. A detailed key and comparative diagnostic
descriptions are given, but it is not intended here to discuss the proposed
different origins for the taxa and the evidence for those origins.

“es

INTRODUCTION

The European, Macaronesian and W. Asian Dryopteris affinis (R.Lowe) Fras.-Jenk.
(“D. paleacea” auct., D. borreri Newm., D. pseudomas (Wollast.) Holub & Pouz.) has
long been known as the third member of the Dryopteris filix-mas (L.) Schott or Male
Fern group, along with two other species, D. filix-mas, D. oreades Fomin (“D.
abbreviata Newm.” sensu Manton 1950) and, later, a fourth species, D. caucasica
(A.Braun) Fras.-Jenk. & Corley. Dryopteris affinis is an apomictic complex, several of
its members presumably derived by hybridisation of various taxa, but in all cases
thought to contain a genome in common with or related to the pan-subtropical species,
D. wallichiana (Spreng.) Hyl. They belong together with the other species with parallel-
sided, + truncate segments in Sect. Fibrillosae Ching, rather than in Sect. Dryopteris,
where the rest of the Male Ferns belong.

As is well known, apomixis in D. affinis means that hybridisation with a sexual
species can give rise to a new fertile, apomictic taxon of a different cytotype. Such
hybridogenous taxa almost certainly correspond with and are the origin of most of the
species within the group (Fraser-Jenkins 1982, as subspecies), though further longer-
term study remains to be carried out to add to the known evidence for their proposed
origins. However three of the species recognised here (from continental Europe and W.
Asia) are insufficiently known as to their origin, but being well known taxa which
appear to be of major distinction and having oa been treated as subspecies, they
are now raised to specific rank along with the

The other important result of apomixis in D. one is that any variants that arise
within members of the group are preserved by the cloning type of reproduction through

Zz FERN GAZ. 18(1): 1-26. 2007

spores. The more major variants have a discrete morphological difference and a
partially distinct geographical range and are recognised here as subspecies, which may
be of quite ancient origin. They are generally of a similar morphology to the species
concerned, but are readily recognisable by their special morphological characteristics.

In addition, more minor variants also occur frequently within and between populations,

which differ only slightly from the general pattern of the subspecies and can be found
here and there in various populations throughout the range of the subspecies. Although
they may be partially distinguishable, they are connected by intermediates occurring in
other localities and while they may be more-or-less recognised from place to place they
merge into one another when all the intermediates from other populations are
considered. These more minor variants are therefore understood to be merely part of the
variation within the species and are not recognised here as being worthy of
nomenclatural distinction. If some of them were needed to be recognised they could be
more appropriately treated at the lower ranks of variety or forma, which would not have
to be included in general floristic works, including County recording etc.

Detailed study of D. affinis throughout its range in Macaronesia, Europe and W.
Asia resulted in the present author (Fraser-Jenkins 1980, 1982, 1983, 1987, 1996a and
b, Widén et al. 1996, Fraser-Jenkins & Trewren in prep.) separating a number of entities
within it. These were initially treated as infraspecific taxa at the rank of subspecies
(corresponding to the present species), with varieties (corresponding to the present
subspecies) placed within them. This ranking of the group was then considered
appropriate because D. affinis sens. lat. was widely considered to be of taxonomic
equivalence to the other species recognised within the genus and some reluctance to
accept further critical taxa within it was brought to the attention of the author at the
time. In the author’s previous and present treatments the taxa were placed hierarchically
in order to reflect their morphological and cytological relationships to each other.

Although following the present author in the taxa recognised, Jermy in Jermy &
Camus (1991), chose not to accept his hierarchical taxonomic scheme and placed all the
taxa he recognised into the category of “morphotypes”. These were intended to be
temporary unranked names as a convenient handle to use for any recognisable variant
in the complex since the authors had not themselves decided at what rank they would
place taxa and did not propose an alternative taxonomy. Morphotypes were thus invalid

ames outside the standard and universally used hierarchy and Articles of the
International Code of Botanical Nomenclature (ICBN 2005) and the authors also took
the term morphotype in a different sense from that originally defined by Danser (1950),
or Davis & Heywood (1970). In addition the relationships of taxa as reflected in the
present author’s taxonomic scheme were obscured and no longer recognised. Jermy was
followed by Pigott (1997), who added three new morphotype names, two for more
minor morphological variants, not recognised here, and the third (“morphotype
arranensis”) being an additional name for a new taxon recognised by the present author
and cited by Pigott. The new morphotypes were published as Latin names, but although
without a type or type-concept, they have been placed here in the synonymy of the
relevant species and subspecies from material kindly made available to the author by
Pigott. Morphotypes were advocated again, as the accepted treatment, by Jermy, Pigott
& Merryweather (1998) and by Merryweather (2002), though suggesting that up to 70%
of specimens were of doubtful identity and that there might be up to 11 possible species
in Britain.

Thus until now the group has remained in a state of confusion with the existence of

FRASER-JENKINS.: THE DRYOPTERIS AFFINIS GROUP 3

two alternative types of treatment and the suggestion that it is much more complex than
hitherto published or than indicated in Nature and is thus almost intractable to botanists
in general. However the relationships and relative placement of the known varieties and
subspecies in a formal taxonomic framework have continued to be appraised carefully
in the field by the present author and as the present taxonomy has been found to work
well, this scheme is again recognised here, as by Fraser-Jenkins (1996a). While it is
acknowledged that the taxa in this group, like many others, require further sophisticated
study of their origin and that a few of the apparently more minor variants within D.
borreri require study as to whether they could be of greater significance, it is hoped that
recent confusion can now be laid to rest and the established taxa can be treated in a
normal botanical framework.

Part of the reason for recourse to morphotypes was that the authors did not wish to
comment on possible relationships and the relative taxonomic significance of the taxa
and were studying locally without seeing the larger picture of the taxa throughout their
range. Although the vast majority of species have long been described without detailed
molecular evidence and continue to be so, the use of morphotypes was also felt
appropriate by some authors who sought further genetic, cytological, chemical or
molecular proof of the present taxonomic scheme. However no suggestion was made as
to where it might be mistaken, or contradicting it, and most workers today have come
to accept the main taxa recognised by the present author. Further problems have been
caused by the presence of the many minor variants within the subspecies, as following
recognition of such variant forms locally, some workers have found it difficult to accept
their placement within the species or subspecies. This has also been compounded by
prevailing cladistic concepts that a species has to be a single homogeneous entity. But
where this has been adhered to with insufficient use of subsidiary and lower ranks to
allow for different levels of variation, it has already created enormous practical
problems. Thus several now much confused and notorious Angiosperm genera have
suffered from excessive separation of unwarranted microspecies unbalanced by
practical considerations and have reached a state where almost no one can put a name
on a specimen any more. The present scheme has undergone continuing reappraisal for
more than 25 years in both field and herbarium and has been successfully used by a
number of workers both in Britain and abroad and it is therefore hoped that its
clarification here and will help to avoid some of the above difficulties. In particular it
is hoped that the temporary and rather unsatisfactory usage of non-standard morphotype
names, which have inevitably become the equivalent of formal names, may now be
superceded by reverting to standard botanical nomenclature. Doing so will not preclude
the recognition of new taxa, should that occur, nor adjustment of rank for individual
taxa, should it become necessary, but it is emphasised that the rank of the species and
subspecies, particularly in Britain now appears to have reached some stability and
general acceptance.

Arelatively major and important change in the taxonomy is effected here in altering
the rank of the main taxa from subspecies to species. Taxonomic understanding of the
group has undoubtedly changed considerably from 25 years ago, when the existence
and delimitation of its constituent taxa were little known. Given the fact that the
important taxa, the former subspecies, recognised within D. affinis by the author are
sufficiently distinct and are now generally and widely able to be recognised, they are
treated here as species in their own right. The choice of species versus subspecies is
always an arbitrary one, and originally the rank of subspecies was chosen as much the

- FERN GAZ. 18(1): 1-26. 2007

better “middle ground”, with good grounds to support it at the time. But as time has
gone on, their distinctiveness and ranges of variation have become much better known
to botanists than they were a decade ago, as have their apparent relationships. In general
they can be considered to be of roughly the same degree of distinction as the other
species of the Male Fern group — ee readily Race RIN from their gross
morphology alone, it is o recognise them as ies. This also reflects
the biology of the taxa since it i is known that most of them ae a different individual
biological (cytological and genomic) basis, though the mere fact of the existence o
such a difference, had it been in the absence of relatively major and readily recognisable

orphological separability would in the present author’s opinion be an inappropriate
as undesirable basis on its own for recognising species.

As there are few great practical difficulties in the D. affinis group, the previous
subspecies are recognised in the present paper as species, and in tandem with this the
previous varieties are treated in a standard way as geographical subspecies within them.
Variant forms below these ranks are not considered to be of sufficient taxonomic
significance to be recognised here, though many have already been named in the past
as varieties or forms. They are obviously morphologically close to the subspecies
concerned and it can readily be recognised as to where they belong; these include two
of the new morphotypes of Pigott, one of which has been found (Fraser-Jenkins &
Trewren in prep.) to have similar 16 spore-mother-celled chromosome-pairing
behaviour and to be chemically similar (Widén et al. 1996) to D. borreri, though further
study is required into variation within the latter.

TAXONOMIC TREATMENT

The three best known species occurring throughout most of the range of the group, and
the only three known so far from Britain, are D. affinis, D. cambrensis and D. borreri.
Fortunately all three names, now well known as subspecies and two of them also as
species names, appear to be usable at the specific rank, despite the involved and
complicated synonymy of the taxa within the group.

The currently accepted species and subspecies in the D. affinis group are as follows
(those with an asterisk inserted after the names are present in Britain and/or Ireland):

1. Dryopteris affinis* (R.Lowe) Fras.-Jenk., Fern Gaz. 12(1): 56 (1979), non Newm.
(1854) [nom. inval. for Lastrea filix-mas var. affinis Newm.]. Lectotype (Fraser-Jenkins
1980): from Madeira (Ribeiro Frio), R.T. Lowe, 1 Nov. 1828 in K!, paratypes: in K!
BM! M! B! etc. Cytotype: Diploid apomict.

A. subsp. affinis*. Synonyms: D. affinis [subsp. affinis] var. affinis (in Fraser-
Jenkins 1980), Dryopteris borreri var. subintegra (Aschers & Graebn.) Tavel (1937)
[based on Aspidium filix-mas var. subintegrum Dé6ll (1857), non Boreau (1840)], D.
pseudomas (Woll.) Holub & Pouzar (1967), D. mediterranea Fomin [= D. wallichiana]
f. disjuncta Fomin, D. affinis [subsp. affinis] var. disjuncta (Fomin) Fras.-Jenk. (1980),
Dryopteris resendeana Rezend.Pinto (1969), Dryopteris borreri subsp. resendeana
(Rezend.Pinto) Malagar. (1975), Dryopteris affinis [subsp. affinis] var. azorica Fras.-
Jenk. (1980). Cytotype: Diploid apomict, the rare 16-smc sporangia showing almost all
univalents (Manton 1950, Dopp 1955). Range: Can, Mad, Az, Mor, Lu, Hs, Andorra,
Ga, Co, Be, Lux, Ho, Br (Scot., Wales, Engl. [VC. S, 1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 13,
14, 16,17, 20, 21, 22,2723, 24, 27, 29, 30,.32; 33, 34, 35, 36, 37,38, 40, 41, 42, 43,
44, 45, 46, 47, 48, 49, 50, 51, 52, 55, 56, 57, 58, 59, 60, 61, 62, 63, 64, 66, 67, 69, 70,

FRASER-JENKINS.: THE DRYOPTERIS AFFINIS GROUP 5

71, 72, 74, 75, 77, 78, 79, 83, 85, 86, 87, 89, 90, 92, 95, 96, 97, 98, 99, 100, 101, 103,
104, 105, 106, 107, 108, 109, 110, 77111, 112), Hb [VC H1, H2, H3, H4, HS, H6, H7,
H8, H9, H10, H11, H12, H13, H14, H16, H17, H20, H21, H22, H27, H28, H29, H31,
H32, H35, H36, H38, H39, H40], No, Ge (W and EB), He, Liecht, It, Elba, Si, Sa, Au, Ju
(Slov., Hrvat.), Ro, An, Rs (S) (Caucasus (Karach.-Cherk.), W. Transcaucasus (Gruz.,
Abkhaz., Adzhar.)). Specimens of this subspecies with more acute teeth and slightly
convex pinnae were invalidly named by Pigott (1997) as his ““morphotype Convexa”.
This taxon is understood here to be one of the minor variant forms within subsp. affinis,
occurring with intermediates from place to place and not requiring nomenclatural
recognition. Some other specimens named as “Convexa” appear to be less lobed or
convoluted specimens of subsp. paleaceolobata and some are flatter-pinnuled
specimens of D. cambrensis. In its toothing it is slightly towards subsp. punctata and
subsp. paleaceolobata, but is obviously not as distinct as they are.

. subsp. paleaceolobata* (T.Moore) Fras.-Jenk., comb. nov., basionym: Lastrea
filix-mas var. paleaceo-lobata T.Moore, Handb. British Ferns, ed. 3: 112 (1857).
Lectotype, here designated: from Scotland, Dumbartonshire, “Lastrea filix-mas
paleaceo-lobata. Tarbet, Dumbartonshire, 1855, T.-M.” (K-Moore!), isolectotypes (x 3):
Ditto: “var. paleacea (M) crispy and incised form”, and same loc. and date, “Herbarium
Hookerianum 1867” (K! one specimen). Synonym: D. affinis [subsp. affinis] var.
napa (T.Moore) Fras.-Jenk., comb. inval. (1996). Cytotype: Diploid apomict
(Manton 1950, identified more precisely in Fraser-Jenkins & Trewren in prep.). Range:
Br oe: Wales, Engl. [VC. 1, 2, 3, 5, 35, 36, 38, 41, 42, 43, 46, 47, 48, 49, 55, 61, 62,
69, 70, 78, 85, 88, 89, 96, 98, 99, 101, 103, 105), Hb [VC. H1], [Be, adventive], ?Ge
(J. Freigang, det. CRFJ, Dec. 2002).

C. subsp. punctata Fras.-Jenk., subsp. nov., facies superiora laminae laevis et
paleae stipitis rhachidis angustae ut in subsp. affine; sed facie superiora pinnularum
valde depressa super sorum et dentibus acutioribus longioribus flabellatis differt.
Holotype, here designated: Switzerland, Kt. Ziirich, above Hutten [N. side of Hohen
Rohn mountain], c. 950 m. alt., Leg. E. Oberholzer, 3 Oct. 1962; frond pressed by T.
Reichstein 740, 12 Aug. 1974, in BM! (sheet with the lower half of the frond is here
designated holotype), isotypes: in BM! Z! ZT! NY! GRU! DUB! Fronds collected by
the present author from the type-plant cult. by T. Reichstein in Basel: in BM! FR! BR!
G! NMW! D. affinis [subsp. affinis] var. punctata Oberh. & Tavel ex Fras.-Jenk. (1980)
was probably invalid as it was not indicated which specimen of TR 740, in which
herbarium, was the holotype. Thus a new subspecies has been described here, rather
than making a new combination. Synonym: D. affinis [subsp. borreri, err.| var.
splendens (Ehrler 1950, as a subvar.) Fras.-Jenk. (1980). Cytotype: Diploid apomict
(Manton 1950, Eschelmiiller & Schneller 1980, Reichstein & Schneller 1983 etc.).
Range: Ga, Ge (W.), He, Au, Ju (Slov., Hrvat.), An

D. subsp. kerryensis* (Fras.-Jenk.) Fras.-Jenk., comb. nov., basionym: Dryopteris
affinis {subsp. affinis] var. kerryensis Fras.-Jenk. in Widén, Fraser-Jenkins, Reichstein,
Gibby & Sarvela, Ann. Bot. Fennici 33: 73 (1996). Holotype: from Ireland, Co. Kerry
(Tahilla to Rossdohan, Kenmare), CRFJ 15174, 6 June 1988, in BM}, isotype: ditto, in
G! Cytotype unknown, but has relatively small, regular spores similar to those of subsp.
affinis, which almost certainly indicate that it is diploid. Range: Hb [VC. HI, 2; B33,
H4, H5, H6, H7, H16].

E. subsp. jessenii (Fras.-Jenk.) Fras.-Jenk., comb. noy., basionym: Dryopteris
affinis [subsp. affinis] var. jessenii Fras.-Jenk., in Widén, Fraser-Jenkins, Reichstein,

6 FERN GAZ. 18(1): 1-26. 2007

Gibby & Sarvela, Ann. Bot. Fennici 33: 73 (1996). Holotype: from Romania, Judep.
Oltenia (road above Baile Herculane), CRFJ 3503, 27 July 1971, in BM!, isotypes:
ditto, in G! H!, paratypes: in Herb. S. Jessen, Chemnitz! BR! G! Z! PRC! Cytotype:
Diploid apomict (Jessen 1991). Range: Ju (Srb.), Ro.

2. Dryopteris cambrensis* (Fras.-Jenk.) Beitel & W. Buck, Fiddlehead Forum, Bull.
Amer. Fern Soc. 15(2): 15-16 (1988). Holotype: Britain, Wales, Merionethshire [=
Gwynedd] (Bala, below Arennig Fawr), CRFJ 12748, 18 Sept. a in BM!, isotypes:
B! BP! DUB! E! G! H! K! NMW! NY! P! Z! Synonym: D. affinis subsp. cambrensis
Fras.-Jenk., in Derrick, Jermy & Paul, Sommerfeltia 6: xi [-xiii] dani Cytotype:
Triploid apomict.

- subsp. cambrensis*. Synonym: D. affinis [subsp. cambrensis] var.
paleaceocrispa (T.Moore) Fras.-Jenk., nom. inval. (1996). Cytotype: Triploid apomict,
with the 16-sme sporangia showing 41 bivalents and 41 univalents (Gibby in Fraser-
Jenkins 1982). Range: Lu, Hs, Ga, Br (Scot., Wales, Engl. [VC. 1, 2, 3, 35, 36, 40, 41,
42, 43, 44, 46, 47, 48, 49, 55, 57, 62, 63, 65, 66, 67, 69, 70, 72, 73, 74, 75, 76, 77, 78,
81, 85, 86, 87, 88, 89, 90, 91, 93, 94, 95, 96, 97, 98, 99, 100, 101, 102, 103, 104, 105,
106, 107, 108, 109, 110, 111, 112), Hb [Vc. H2, H7, H8, H9, H10, H16, H20, H39,
H40], No, An

B. subsp. distans (Viv.) Fras.-Jenk., comb. nov., basionym: Aspidium distans Viv.,
App. ad Flor. Cors. Prodr.: 9 (1825), non Dryopteris distans (Hook.) Kuntze. Lectotype
(Pichi Sermolli 1960): from Corsica (Ospedale), S. Serafini, 1827 in BOLO!,
isolectotype: in GE (lost). Synonym: Dryopteris affinis [subsp. cambrensis] var. distans
(Viv. [“Fiori”]) Fras.-Jenk. (1987). Cytotype: Unknown. Range: Ga, Co, It. The author
(Fraser-Jenkins 1987) initially misidentified the precise identity of the immature type of
this name until making a detailed study of the type and type-locality. This subspecies,
with a distinct range centred on western France, is consistently intermediate between
the eastern French, Mediterranean etc. subsp. insubrica and the more northerly subsp.
cambrensis, but is nearer to the latter, yet has reddish scales.

C. subsp. insubrica (Oberh. & Tavel ex Fras.-Jenk.) Fras.-Jenk., comb. nov.,
basionym: D. affinis [subsp. cambrensis] var. insubrica Oberh. & Tavel ex Fras.-Jenk.,
in Widén, Fraser-Jenkins, Reichstein, Gibby & Sarvela, Ann. Bot. Fennici 33: 73
(1996). Holotype: from Switzerland, Kt. Ticino (Lugano, Breganzona to Muzzano), F.
von Tavel, 14 Oct. 1916, in BERN!, isotypes (x 3): in BERN! Cytotype: Triploid
apomict with the 16-sme sporangia showing approximately equal numbers of bivalents
and univalents, or sometimes slightly fewer bivalents (Vida in Fraser-Jenkins 1980,
1982 and in Fraser-Jenkins & Trewren in prep.). Range: Ga, Co, Be, Lux, Ho, Da, No,
Ge (W. and E.), He, It, Elba, Si, Sa, Au, Hu, Cz (Bohem., Morav.), Po, Ju (Slov., Hrvat.),
Bu. A further, more minor local variant of subsp. insubrica from the Armorian region
of S.W. France has more acute pinnule-apices and long teeth, but is not considered
sufficiently distinct to be worthy of nomenclatural recognition here.

p- pseudocomplexa* Fras.-Jenk., subsp. nov., planta similis ad D.
cambrensi subsp. cambrensi a qua differt paleis stipitis pallidioris, lamina latiora,
pinnuli laxioris minoris confertis, apicibus earum acutioribus-rotundis, dentes plus
minusve absentis vel brevibus acutos saepe ferrentibus. Holotype, here designated:
Britain, Scotland: Woods in from gate, just south of Dunvegan Castle car-park, N.W.
Skye, Inner Hebrides, Scotland, C.R. Fraser-Jenkins 12635, 9 Sept. 1986, in BM!;
isotypes: Ditto, in H!; paratypes: ditto, nos. 12634, 12636-12640, 12644-12648, in BM!

FRASER-JENKINS.: THE DRYOPTERIS AFFINIS GROUP a

NMW! H!; woods in from gate just south of Dunvegan Castle car-park, N.W. Skye,
Inner Hebrides, Scotland. CRFJ 11130-11134, 11136-11141, 11143-11154, 16 Sept.
1984, in BM! NMW! Synonym: D. affinis [subsp. cambrensis] var. pseudocomplexa
Fras.-Jenk., nom. nud. (1996b) [form imitating D. x complexa” in Widén, Fraser-
Jenkins, Reichstein, Gibby & Sarvela: 75 (1996)]. Cytotype unknown, but spores large
and a fairly high percentage (up to c. 30%) of abortive spores, similar to those of the
triploid apomict subsp. cambrensis. Range: Ga, Br (Scot. [VC. 100, 104), Hb [VC. H2,
H6], No, ?Au. Also discovered independently by Dr. A. Church on the Isle of Arran, W.
Scotland in 1992 and thence published as “morphotype Arranensis” by Pigott (1996),
with the comment that it might be the same as the unattributed var. pseudocomplexa
[from the author’s manuscript]. This is now placed within subsp. pseudocomplexa.

3. Dryopteris pseudodisjuncta (Tavel ex Fras.-Jenk.) Fras.-Jenk., comb. nov.,
basionym: Dryopteris affinis |subsp. borreri, err.] var. pseudodisjuncta Tavel ex Fras.-
Jenk. in Dostal, Fraser-Jenkins & Reichstein, Hegi’s //lustr. Flor. Mitteleurop., ed. 2
(Kramer), 1: 147 (1983 [“1984”]). Holotype: from Switzerland, Kt. Zug (Finstersee,
Hohen Ron [mountain]), E. Oberholzer, in BERN! Synonyms: Dryopteris affinis subsp.
pseudodisjuncta (Oberholz. & Tavel ex Fras.-Jenk.) Fras.-Jenk., Nordic J. Bot. 16(1): 4

); Dryopteris affinis [subsp. cambrensis, err.] var. setosa (Christ 1900) Fras.-
Jenk. (1987). Cytotype: triploid apomict, with the 16-smc sporangia showing c. 20-24
bivalents and the remainder univalents (Dépp 1963, Schneller in Eschelmiiller &
Schneller 1980). Its proposed genomic origin is unclear, but is apparently different from
that of D. cambrensis, from which it shows slightly different pairing behaviour at
meiosis, suggesting segmental allopolyploidy, though its phloroglucinol chemistry is
similar to subsp. cambrensis. Range: Ga, Be, Lux, No, Ge (W.), He, Au.

4. Dryopteris schorapanensis Askerov, Isvest. Akad. Nauk. Azerbaidzhan S.S.R., set.
Biol. Med. Nauk, 1978(4): 4 (1979), pro hybr. [omitted by Johns et al. 1996] (described
as a proposed hybrid of what is now D. affinis with D. caucasica Fomin). Holotype:
[former] U.S.S.R., Gruzhia [Georgia] (Schorapani, Mesketskii Khrebet [range], Kavkaz
[Caucasus] Mts., N. Georgia), J. Medvedev, 18 July 1968, in TBI! Synonym: D. affinis
subsp. persica Fras.-Jenk. (1980). Cytotype: Triploid apomict (Vida in Fraser-Jenkins
1980). Its proposed genomic origin is unclear. Range: ?Ro, ?An, Caucasus (Rs (S)
(Krasnodar)) and Transcaucasia (Abkhaz., Georg., Karabakh., Azerbay., Dagest.), Ir. At
specific rank what had been called subsp. persica must be called D. schorapanensis,
one of several taxa described as hybrids by Askerov.

5. Dryopteris pontica (Fras.-Jenk.) Fras.-Jenk., comb. nov., basionym: Dryopteris
affinis subsp. pontica Fras.-Jenk. in Widén, Fraser-Jenkins, Reichstein, Gibby &
Sarvela, Ann. Bot. Fennici 33: 73 (1996). Holotype: Turkey (Anatolia), Ordu Vilhayet
(Hargbeli Pass, Kirtibe®, Gélkéy), CRFJ 14065, 15 Nov. 1987, in BM!, isotypes: ditto,
in G! H!, paratypes: ditto, CRFJ 14058-14064, 14066-14068, in BM! NMW! Cytotype:
Triploid apomict (Gibby in Fraser-Jenkins 1980, Vida in Fraser-Jenkins & Trewren in
prep.). Its proposed genomic origin is unclear, but it has a quite distinct morphology
with some characteristics in common with D. schorapanensis and rectangular lobes
similar to D. borreri. Range: An, Transcaucasus, Ir.

6. Dryopteris borreri* (Newm.) Newm. ex Oberh. & Tavel in Tavel, Verh. Schweiz.

8 FERN GAZ. 18(1): 1-26. 2007

Naturf. Ges. 118: 153 (1937). Lectotype (Fraser-Jenkins 1980): Britain, England, E.
Newman, 1879, in BM! Synonyms: Dryopteris affinis subsp. borreri (Newm.) Fras.-
Jenk. (1980), Lastrea filix-mas var. abbreviata (DC.) Newm (1854), Dryopteris
abbreviata (DC.) Manton (1950), non (Schrad.) Kuntze, Aspidium blackwellianum
Ten., nom. provis. inval., ??Polystichum induratum Schur (1866) (said to be D. filix-mas
by Christensen (1905); type not found by the author, ?in LW), Dryopteris affinis subsp.
stilluppensis (Sabr.) Fras.-Jenk. (1980), non sensu Fras.-Jenk., D. stilluppensis (Sabr.)
Holub (1986), non sensu Holub, D. affinis subsp. robusta Oberh. & Tavel ex Fras-Jenk.
(1980) (holotype, isotype and paratypes: from Switzerland, Kt. Ziirich (Hohen Ron
[mountain]), E. Oberholzer, 28 Sept. 1940, in BERN!), D. pseudomas subsp. robusta
(Oberh. & Tavel ex Fras.-Jenk.) Holub (1984), D. mediterranea Fomin [= D.
wallichiana] var. robusta (Oberh. & Tavel ex Fras.-Jenk.) Holub (1986), D. x tavelii
Rothm. (1945) (lectotype (Fraser-Jenkins 1987): from Hungary, Vas Comit. (H4mor,
K6szeg (Giins) [= Gészbachtal, Aradopatak valley], A. Waisbecker, 10 Aug. 1899, in
B!). Many other varietal names also apply to this species. Cytotype: Triploid apomict,
with the 16-smc sporangia showing c. 123 single chromosomes, or sometimes from 10-
18 bivalents (Manton 1950, Schneller 1974, 1975a-b, Manton, Vida and Rasbach in
Rube & Heise 1975, in Fraser-Jenkins 1980 and in Fraser-Jenkins & Trewren in prep.).
Fraser-Jenkins’ (1980) reports of some plants showing approximately equal numbers of
bivalents and univalents was made prior to his reidentification of Manton’s voucher-
specimens, and referred to D. cambrensis. Range: Lu, Hs, Andorra, Ga, Co, Be, Lux,
Ho, Br (Scot., Wales, Engl. [VC. S, 1, 2, 3, 4, 5, 6, 8, 9, 10, 11, 12, 13, 14, 15, 16, 17,
19, 20, 22, 23, 24, 25, 26, 27, 29, 30, 33, 34, 35, 36, 37, 39, 40, 41, 42, 43, 44, 45, 46,
47, 48, 49, 50, 52, 53, 54, 55, 57, 58, 59, 60, 62, 63, 64, 65, 66, 67, 68, 69, 70, 71, 72,
73, 74, 75, 76, 77, 78, 79, 81, 82, 83, 84, 85, 86, 87, 88, 89, 90, 91, 92, 93, 94, 95, 96,
97, 98, 99, 100, 101, 102, 103, 104, 105, 106, 108, 109, 110, 111, ?7112]), Hb [VC. H1,
H2, H3, H4, H5, H6, H7, H8, H9, H10, H13, H15, H16, H20, H21, H23, H25, ?H26,
H27, H28, H29, H30, H31, H32, H33, H35, H36, H37, H38, H39, H40], Da, No, [S],
Ge (W. and E.), He, It, Elba, Si, Au, Hu, Cz (Bohem., Morav., Slovak.), Po, Ju (Slov.,
Hrvat., Bosn.-Herc., Crna Gora, Srb., Makedon.), Ro, Bu, Tu, An, Rs (W) (Kalinin.,
Ukr.), ??Rs (K), Rs (S) (Caucasus (Krasnod., Sev.-Osset.), Transcauc. (Abkhaz.,
Georg., Dagest., Adzhar., Azerbayd.)), Ir.

D. borreri is morphologically very close indeed to the triploid apomict D.
pseudofilixmas Rothm. (1945), from Mexico, which may be due to chance convergence
following the combination of their ancestors; their chemistry and presumably their
origin is different, but further study is required to investigate its relationship to D.
borreri. There are numerous minor variants of D. borreri, which all appear to merge
into each other and are probably merely due to less significant variation being preserved
through apomictic cloning, as also occurs within the subspecies of the other species.
Being obviously below the rank of subspecies these are not recognised here even
though many of them have received varietal names, albeit uncritically, in the past. The
botanical varietal names have been detailed in Fraser-Jenkins & Trewren (in prep.), but
a great many more varietal names given to abnormal cultivars are also valid at varietal
rank and apply to the basic more normal plant from which the monstrosities were
derived. Any attempt to apply lower ranking names to the minor variants within D.
borreri or within the subspecies of the other species would therefore be extremely
difficult. Applying new names would be still less advisable unless they were for more
fundamental entities at the subspecific rank, which the author has not recognised in

FRASER-JENKINS.: THE DRYOPTERIS AFFINIS GROUP 9

Nature so far. It would probably also be of dubious value because whereas individual
minor variants can recur frequently in many populations and be more-or-less
prnesere from place to place, intermediates occur in other populations and in
different regions and it appears doubtful whether any of them represent discrete entities.
They have therefore not been recognised nomenclaturally by the present author, even at
the rank of forma, though it is evident that further research is required. One such form
that occurs frequently with rather small, squarely truncate pinnules similar to the type
of D. borreri, has been separated by Pigott (1997) as “morphotype Insolens”. It may be
directly synonymous with D. borreri itself, or may turn out to be a distinct entity on
further investigation. It occurs commonly throughout Britain and in parts of the western

uropean-continental range of the species, whereas in N.E. Europe D. borreri usually
has wider and often more lobed pinnules (these forms also occuring in Britain and W.
Europe). The present author mistakenly validated one of von Tavel & Oberholzer’s
many varietal names within this species as D. affinis subsp. robusta, which was taken
up by Holub, Jermy and then Pigott in various slightly different senses, the two latter as
“morphotype Robusta”. But this name simply referred to plants of D. borreri with
slightly long, developed, but hardly lobed lowest basiscopic pinnules, with more
rounded pinule-apices, which can occur in several of its minor variants and perhaps
varies continuously in degree of development, though requiring further study. Subsp.
robusta has already been sunk by Fraser-Jenkins (1987, 1996b) but the name is still
being used in various senses by some authors.

7. Dryopteris iranica Fras.-Jenk., nom et stat. nov. for Dryopteris affinis subsp.
coriacea Fras.-Jenk., Willdenowia 10: 12-113 (1980), non Dryopteris coriacea (Brause)
Ching [= Thelypteris (Amphineuron) coriacea (Brause) Ching]. Holotype: Iran, Talesh,
Gilan (above Assalem, S. of Siadiin, Astara to Bandar-e Pahlavi), CRFJ 5933, 30 June
1977, in BM!, isotype: ditto, in BM!; paratypes: ditto, 5926-5932, 5934-5939, BM!
NMW! H! G! FR! Synonyms: D. wallichiana subsp. coriacea (Fras.-Jenk.) Fras.-Jenk.
(1996b), D. affinis subsp. coriacea var. gigantea Fras.-Jenk. (1980). Cytotype: triploid
apomict (Vida in Fraser-Jenkins 1980). This subspecies, though more closely related to
D. wallichiana in its long scales and very coriaceous lamina is readily nae sos
distinguishable from it in its rounded, flabellately toothed pinnule-apices etc.
deserves specific rank. Other subspecies and unnamed variants of D. wa Shien in
Asia are generally closer to subsp. wallichiana itself and are maintained by the author
as subspecies, with the exception of Dryopteris himalaica (Ching & S.K.Wu) S.GLu
(syn.: D. wallichiana var. himalaica Ching & S.K.Wu, D. wallichiana subsp. himalaica
Fras.-Jenk. (1997), the latter fortuitously using the same epithet as Ching’s, despite
having inadvertently lost track of the earlier name at varietal rank). D. wallichiana
subsp. coriacea var. gigantea Fras.-Jenk. (1980) from N. Turkey and N. Iran appears
very similar morphologically to D. himalaica but was reported as triploid by Gibby,
while D. himalaica was reported as diploid (Gibby 1985, CRFJ 10364).

HYBRIDS
The cytotypes of the D. affinis x D. filix-mas hybrids detailed here are tetraploid or
pentaploid, whereas the species are diploid or triploid. They are normally
—— by their intermediate frond-morpology combined with mostly abortive
s and by not usually forming large populations, but occurring as a few isolated
indbviduals in a population. However the author has seen some considerable

10 FERN GAZ. 18(1): 1-26. 2007

populations of D. x complexa nothosubsp. complexa in S.W. Ireland, which may have
something to do with spore-germination being favoured by the luxuriant, mild and
consistently moist climate there, as elsewhere in the west. So far no definite D. affinis
agg. x D. oreades Fomin has been discovered, previous records all being of D.
dombrensi which almost certainly had such an origin. Some further candidates have
recently been suggested, but are at present of uncertain identity. A further complication
is that D. oe x D. affinis would presumably ws nn identical and probably
the same as D. cambrensis, while D. oreades . borreri would presumably be
genomic ae identical to D. x complexa and min two abana for both would probably
give indistinguishable results. Only D. cambrensis x D. oreades should be expected to
be a distinctive hybrid worth looking out for, though like any other hypothetical taxa
within the group which may or may not exist, it need not be mentioned further until
actually discovered in Nature.

The nomenclature of several of the hybrids in the group requires revision now that the
subspecies of the D. affinis group have been raised to the specific rank. Hybrids within
the group known to the author are as follows:-

1. Dryopteris x complexa* Fras.-Jenk., in Derrick, Jermy & Paul, Sommerfeltia 6: x-
xiii (1987). D. affinis x D. filix-mas. Holotype: from Germany [former W. Germany]
(Gaisbrunnen, c. 2 km S.E. of Herrenalb, S. of Karlsruhe), G Philippi, H. & K.
Rasbach, T. Reichstein & A. Sleep TR 4020, 14 Aug. 1976, in Z!, isotype: in M! Fronds
from type-plant cult. by T. Reichstein in Basel in herb. CRFJ, in BM! NMW! Cytotype:
Tetraploid semi-sterile hybrid.

A. nothosubsp. complexa*. D. affinis subsp. affinis x D. filix-mas. Cytotype:
Tetraploid semi-sterile hybrid, with the 16-smc sporangia showing c. 41 bivalents and
c. 82 univalents (Manton 1950 and in Fraser-Jenkins & Trewren in prep., Rasbach in
Rasbach, Rasbach, Reichstein & Schneller 1983 and in Fraser-Jenkins & Trewren in
prep.). Range: Lu, Hs, Ga, Be, Br (Scot., Wales, Engl. [VC. S, 1, 2, 3, 5, 11, 32, 35, 36,
41, 42, 44, 51, 55, 62, 63, 69, 70, 82, 85, 87, 99, 103, 105]), Hb [H1, H2, H3, H6, H8,
H10, H20, H22, H28, H29, H32, H39], Ge (W.), He, It, Au, Ro.

nothosubsp. contorta* Fras.-Jenk., in Derrick, Jermy & Paul, Sommerfeltia 6: x-
Xiii (8 (erroneously thought at first to be D. cambrensis subsp. cambrensis x D.
filix-mas). D. affinis subsp. paleaceolobata x D. filix-mas. Holotype: from Britain,
bape (Glen Affric, Inverness-shire, CDFJ and ABJ, 1967 [erroneously thought to
have been from Leicestershire, at The Brand, Woodhouse Eaves, CRFJ, Oct. 1968]),
CRFJ 11494, ex hort. Newcastle House, Bridgend, 25 Aug. 1985, in BM!:; isotypes:
ditto, in G! H! NMW! Cytotype: Tetraploid semi-sterile hybrid (Gibby in Fraser-
Jenkins & Trewren in prep.). Range: Br (Wales, Engl. [VC. 47, 96]).

C. nothosubsp. eschelmuelleri Fras.-Jenk., nothosubsp. nov. planta similis ad

nothosubsp. complexam sed dentibus ee Fflabellulatis et facie superiora
pinnularum valde depressa super sorum diffe abortivae. Holotype,
here designated: Germany [W.], Bayern, south Schwaben, Allgauer Alpen, Grunten siid,
eines A. paper [klon 11] 85/11 (Herb. T. Reichstein no. 6282, Basel, now
Z!); isotypes: schelmiiller [klon 10 and 11] 84/23, 13/8/1984, 84/36 and 84/88,
13/9) 1984, ie Gone and 85/73, 19/8/1985 (Herb. A. Eschelmiiller, Sulzberg-
im-Allgau, now in M!). D. affinis subsp. punctata x D. filix-mas. Cytotype: Tetraploid
semi-sterile hybrid (Bar & Eschelmiiller 1985). Range: Ge (W.), He, Ju (Hrvat.), An.

FRASER-JENKINS.: THE DRYOPTERIS AFFINIS GROUP 11

2. Dryopteris x convoluta* Fras.-Jenk., nothosubsp. nov., frons angustiora quam in D.
x complexa segmenta parviora et pinnulae basiscopicae ad superam curvatae. Par
inferum pinnularum quoquae pinnae valde longiorum. Paleae stipitis rubricantae.
Indusia glandulosa. Sporae plerumque abortivae, sed sporae bene formatae maximae.
Holotype, here designated: Germany [W.], Bayern [= Bavaria], south Schwaben,
Siidwest seite des Griinten, von Burgberg aufwarts, A. & H. Eschelmiiller 85/40,
24/5/1985 (Herb. A. Eschelmiiller, Sulzberg-im-Allgau, now in M!). Plant then grown
from spores by A. Eschelmiiller, frond coll. AE 88/72, 9/1988 (photo!). D. cambrensis
x D. filix-mas. Cytotype: Pentaploid semi-sterile hybrid.

A. nothosubsp. convoluta. D. cambrensis subsp. insubrica x D. filix-mas.
Cytotype: Pentaploid semi-sterile hybrid (Dépp 1941, Bar & Eschelmiiller 1989).
Range: Co, Ge (W.), He.

B. nothosubsp. occidentalis* Fras.-Jenk., nothosubp. nov., planta similis ad
nothosubsp. convolutam sed paleae stipitis flavo-fuscae, dentes pinnularum minori.
Indusia fere smal a Sporae tyes eee Holotype, here designated:
Britain, Scotland (Inverness-shire, road-cutt top of pass, Shlochd, north of
Carrbridge, south of Inverness), CRFJ 11187, 1e/071984, in NMW. D. cambrensis
subsp. cambrensis x D. filix-mas. Cytotype: unknown. Range: Ga, Co, Br (Scot., Wales,
Engl. [VC. 47, 49, 62, 69, 74, 87, 96, 104]), Hb [VC. ?H6].

C. nothosubsp. inconspicua Fras.-Jenk., nothosubsp. nov., planta similis ad
nothosubsp. occidentalem sed segmenta minore lobata et dentes insignificanti, paleae
stipitis pallidae vel flavae. Holotype, here designated: France, Haut-Rhin (east-facing

woodland scree, c. 1 km along Sentier des Roches foot-path, south-east side of Col de
la Schlucht, west of Colmar on Remiremont road, Vosges mountains), CRFJ 13688,
9/8/1987, in NMW (with D. x sarvelae. CRFJ 13876, 17 Sept. 1987, NMW). Plant
cultivated in hort. M.H. Rickard, Leinthall Starkes, Herefordshire, later transferred to
Kyre Park, Ludlow, Shropshire, where it subsequently died. D. cambrensis subsp.
pseudocomplexa x D. filix-mas. Cytotype: unknown. Range: Ga, Br (Scot. [VC. 100])
It should be searched for further in Scotland and Ireland, where it is likely to occur more
commonly.

3. Dryopteris x complanata Fras.-Jenk., hybr. nov., frondes similes ad D. x criticam sed
lamina valde complanata et pinnulis minoris lobatis, apicibus pinnularum cuneato-
acutis similis ad D. pseudodisjunctam sed dentes plures. Sporae plerumque abortivae.
Holotype, here designated: Germany [W.], Nordrhein-Westfalen (east Arnsberg, below
Ramsbeck waterfall, south of Meschede, north Rothaargebirge, Dortmund to Kassel,
wooded slopes across stream), C.R. Fraser-Jenkins 13748, 19/8/1987, in BM!, isotypes
(x 8): FR! BR! G! H! Herb. T. Reichstein (no. TR 6862), Basel, now in Z! NMW! K!,
paratypes (x 3): Ditto, CRFJ 13749 in NMW. D. pseudodisjuncta x D. filix-mas.
Cytotype: unknown. Range: Ge (W.). Reports of this hybrid by Gatzi (1961: 44) and
Dopp, Giitzi & Oberholzer (1963) have been investigated by the author and were in
error for D. filix-mas and D. pseudodisjuncta respectively.

4. Dryopteris x critica* (Fras.-Jenk.) Fras.-Jenk., comb. noy., basionym: Dryopteris x
complexa nothosubsp. critica Fras.-Jenk., in tte Jermy & Paul, Sommerfeltia 6: x-
xii (1987). Holotype: from Germ (former E. Germany), Gera
(Rudolstadt/Thiiringen, zw. Partschefeld et “Ohstiat), I. & S. Jessen 1291, 22 June
1985, in Herb. S. Jessen, Chemnitz [formerly Karl-Marx-Stadt]!, isotype: in JE! D.

12 FERN GAZ. 18(1): 1-26. 2007

borreri x D. filix-mas. Cytotype: Pentaploid partly sterile hybrid, with the 16-smc
sporangia showing a range from nearly all univalents to c. 68-81 bivalents and the rest
univalents (Manton 1950, Vida in Rasbach, Rasbach, Reichstein & Schneller 1983,
Schneller 1974, 1975a-b, Rasbach in Jessen 1985). Range: ?Hs, Ga, Be, Lux, Br (Scot.,
Wales, Engl. [VC. 27, 35, 36, 40, 47, 48, 49, 52, 55, 57, 60, 62, 63, 64, 69, 88, 92, 97,
104, 105], Hb [VC. H4, H23, ?H39 and collected by R.L. Praeger], No, Ge (W. and E.),
He, It, Au, Hu, Ju (Hrvat.), Cz (Moravia), Ro, Bu, Tu. Some plants of D. x complexa
were erroneously distributed to various British gardens as being this hybrid from the
collection of the late Prof. I. Manton, Leeds, by the late Dr. A. Sleep, the two having
become transposed at Leeds University Botanical Garden.

KEY TO THE D. AFFINIS GROUP.

Comparative diagnostic descriptions of the various taxa have been provided by Fraser-
Jenkins & Trewren (in prep.) and these and the key from that work are provided here.
Unfortunately, as with many complex groups in pteridophytes, apparently “diagnostic”
characteristics can on occasion vary sufficiently in some individuals so as to be
misleading and may have to overridden in order to place a specimen in its species or
subspecies correctly. The key allows for this to some degree concerning the main
exceptions likely to be encountered, but it may still be necessary to balance and
evaluate a combination of characteristics simultaneously when examining a specimen.

ature specimens are dealt with in this key and both immature ones and occasional
extremes may have to be abandoned, at least as far as the key and diagnostic
descriptions are concerned (though with experience they can usually be correctly
recognised identified more intuitively). Garden-grown specimens can also be
particularly difficult when, as often occurs, rather exposed, or not well developed. It is
also pertinent to point out that specimens which have been pressed and mounted in such
a way that the very important upper (adaxial) surface of the lower part of the frond is
covered over or scrumpled up may also be very difficult to identify. The smaller spore-
size of D. affinis has not been used here as a key-characteristic, partly because many
people will not have easy access to a microscope. However it should be pointed out that
it can be useful in cases of doubt, as long as the spores are not mostly shed and as long
as they are fully mature. Youngish spores in the D. affinis group, even after they have
become brown and developed their perispore, are normally smaller than mature ones. It
takes a little experience to recognise their very slightly paler colour, less wrinkled
perispore, greater irregularity in size and partial clumping as being typical of an
immature spore-sample, and not of a hybrid, and thus not to rely on their small size in
such cases. However it is not uncommon to find specimens of D. borreri (which usually
has the smallest spores of any of the triploids) with small spores almost approaching
those of D. affinis, though they are less regular. A chromosome-count is not considered
here to be a necessary or normal part of the identification process.

la. Spores (when ripe and not mostly shed) almost entirely and highly irregularly
abortive, with a few very large good ones (c. 5% - 20%) present, incuding + spherical
ones. [This characteristic has to be observed in combination with the occurrence of
intermediate-type frond-morphology, as on rare occasions spore-abortion alone is not
always absolutely reliable in this group.] Plants of sporadic occurrence and not
normally forming populations
D. x complexa*, D. x convoluta*, D. x complanata and D. x critica*.

FRASER-JENKINS.: THE DRYOPTERIS AFFINIS GROUP 13

1b. Spores (when ripe and not mostly shed) mostly good, or with a majority, or high
proportion of good ones mixed with abortives, so that the good spores predominate in
e sample 2.

2a. Rachis-scales markedly long and narrow, strongly exserted from the rachis. Laminar
texture very stiffly coriaceous; pinnules or pinna-lobes deeply joined together at their
bases (i.e. sinus between the pinnules not extending as deeply as in D. affinis) in the
mid-upper parts of the lamina and pinnae D. wallichiana agg.
2b. Rachis-scales shorter and usually wider, clothing the rachis more tightly. Laminar
texture often coriaceous, but hardly stiffly coriaceous; pinnules or pinna-lobes not so
deeply joined together at their bases (i.e. sinus between the pinnules very nearly
reaching the costa) in the mid-upper parts of the lamina and _ pinnae
D. affinis* agg.

3a. Sori tall, indusium + thick, only slightly lifting and usually splitting on ripening and
not, or only slightly shrivelling. The lowest basiscopic pinnule of the lowest pinna
usually 4 - % adnate to the pinna-costa; lamina considerably glossy above ................ 4.
3b. Sori not tall, indusium + thin, lifting on or after ripening and shrivelling, not
splitting [except for D. affinis, included under 28a. for comparison]. The lowest
basiscopic pinnule of the lowest pinna usually fully stalked, or sometimes % adnate;
lamina not, or only slightly glossy 12.

4a. Pinnules markedly crowded, some with sloping apices

. affinis subsp. kerryensis*.
4b. Pinnules not markedly crowded, with symmetrical apices 5

5a. Scales black, pinnule-apices squarely truncate . .. D. affinis subsp. jessenii.
5b. Scales pale, reddish, or brown (rarely with: 4 very y dark bases), pinnule-apices
rounded-truncate to rounded-pointed, or pointed 6.

6a. Lamina with marked “punctate” indentations on the top surface above each sorus
s subsp. punctata.
6b. Lamina without, or only with shallow and staigiihicken:! ‘punctate” indentations
above a.

7a. Lamina with a wide base, the lowest basiscopic pinnules of the lowest pinna
developed and longer than the rest, the lower pinnules on each pinnule markedly lo
with small, neat lobes; stipe and rachis scales all narrow (indusium shrivels more than
in other D. affinis subspecies) D. affinis subsp. paleaceolobata*.
7b. Lamina more-or-less tapering to the narrowed base, the lowest basiscopic pinnules
of the lowest pinna not usually the longest, lower pinnules on each pinna unlobed, or +
shallowly lobed with somewhat coarse lobes; stipe and rachis scales narrow or
i 8.

wide

8a. Lamina flat, thickly coriaceous, with veinlets markedly impressed above. Pinnae
long; pinnules markedly regular in size, with parallel sides, seldom lobed at the sides
apart from a rounded basal auricle on the lowest pair of pinnules on each pinna. Sori
large, spores small mostly regular (i.e. nearly all good)
D. affinis subsp. affinis*.

14 FERN GAZ. 18(1): 1-26. 2007

8b. Lamina not flat as at least some pinnules are normally curved up at their tips,
coriaceous, but not thickly so, veinlets not markedly impressed above. Pinnae short;
pinnules often irregular in size as the first opposite-pair of pinnules on each pinna is
frequently somewhat longer than the rest, not completely parallel at the sides as they
are often slightly spathulate and wider at their apices (sometimes due to the down-
rolling of the lower-mid sides), usually lobed at the sides, at least on the lowest pair of
pinnules on each pinna. Sori relatively small, spores large with a considerable
proportion of abortive spores (sometimes nearly equal to the number of good spores)

9.

9a. Pinnule side-lobes wedge-shaped and pointed, pinnule-apices with prominent, +
markedly long, acute, markedly flabellate teeth; frond-axes highly glandular, scales
markedly reddish-yellow or red in colour. D. cambrensis subsp. insubrica.
9b. Pinnule side-lobes not wedge-shaped or pointed, but rounded, pinnule-apices with
smallish, + obtuse to slightly acute, weakly flabellate or non flabellate teeth (or teeth
absent); frond-axes mostly eglandular, scales varying from pale or whiteish to red or
dark brown in colour 10

10a. Scales red coloured D. cambrensis subsp. distans.
10b. Scales whiteish or brown

lla. Scales all pale, lamina pale-green, pinnae long with well separated pinnules and
teeth often absent, or, when present, narrowly acute
D. cambrensis subsp. pseudocomplexa*.
11b. Scales not all pale (apart from in some exceptional plants), but brown or slightly
yellowish-russet brown, lamina mid- to dark-green, pinnae short, teeth often absent, or,
when present, somewhat obtuse, though often with a pointed apex ............
D. cambrensis subsp. cambrensis*.

12a. Pinnule-teeth obviously present 43.
12b. Pinnule-teeth + absent 28b
13a. Pinnule-apices squarely truncate 14.
13b. Pinnule-apices + rounded, or pointed 16.

14a. Scales black, lamina glossy and thick, pinnule-teeth — but wide and irregular
in length on each pinnule-apex D. affinis subsp. jessenii.

14b. Scales pale or brown, lamina + matt and thin, pinnule-teeth long or short, with a +
narrowly acute apex, + regular in length on each pinnule- -apex

15a. Lower pinnules with a wedge-shaped base, pinnules usually not markedly lobed,
laminar upper surface + glossy; indusia with well curved-down — and a dee

darkish centre D. pse endbalitieleté:
15b. Lower pinnules with a rectangular base, pinnules usually not markedly lobed,
laminar upper surface matt; indusia with less markedly curved down sides and a
shallow, concolorous centre D. borreri*.

16a. Pinnule-teeth + narrowly acute at the tips of the pinnules 17.

FRASER-JENKINS.: THE DRYOPTERIS AFFINIS GROUP 15

16b. Pinnule-teeth + obtuse at the tips of the pinnules 23.

17a. Lamina with marked “punctate” indentations on the top surface above each sorus
D. affinis subsp. punctata.
7b. Lamina without, or only with very shallow and insignificant “punctate”
indentations 18.

18a. Lower pinnules with markedly rectangular side-lobes with pointed corners
D. borreri*.
18b. Lower pinnules + without rectangular side-lobes (though often with rounded, or
pointed ones) 19.

19a. The lower oe on each pinna lobed with numerous small, neat lobes; stipe and
rachis scales all narrow . affinis subsp. paleaceolobata*.

19b. The lower ace on each pinna unlobed or lobed with few, often somewhat
coarse lobes; stipe and rachis scales include many wide ones 20

20a. Pinnules long and narrow, widely separated
D. cambrensis subsp. pseudocomplexa*.
20b. Pinnules not noticeably long and narrow, crowded, or + narrowly separated

21a. Lamina pale-green and matt above, pinnule-teeth wide-based, narrowing +

abruptly to a long, acute apex; only a weak dark spot present at pinna-axils, or absent
altogether; indusium rapidly lifting and shrivelling on sporangial ripening ................
D. schorapanensis.
21b. Lamina + darker green and + glossy above, pinnule-teeth not abruptly narrowed
from a wide base, long- or shortly acute; an obvious dark spot present at the pinna-axils;
indusium remaining inflected at the sides until shrivelling soon after sporangial
ripening 22.

22a. Scales paler- or brighter reddish in colour, pinnae short, lamina well tapered so that
the lowest pinnae are usually the shortest, frond-axes highly glandular; pinnule-apices
rounded to somewhat acutely pointed ..............c::ceeee D. cambrensis subsp. insubrica.
22b. Scales pale, or slightly smokey-yellow in colour, pinnae long, the lowest pinna +
as long as those above, frond-axes + eglandular; pinnule-apices varying from narrowly
truncate to narrowly pointed, but frequently lop-sided so that one apical corner is taller
than the other D. pseudodisjuncta.

23a. Lamina with marked “punctate” indentations on the top surface above each sorus
D. affinis subsp. punctata.
23b. Lamina without, or only with very shallow and insignificant “punctate”
indentations 24

24a. The lower pinnules on each pinna lobed with eee small, neat lobes; stipe and
rachis scales all narrow D. affinis subsp. paleaceolobata*.

24b. The lower pinnules on each pinna unlobed, or lobed with few, somewhat coarse
lobes; stipe and rachis scales include many wide ones 25

16 FERN GAZ. 18(1): 1-26. 2007

25a. Lower pinnule-apices narrowly rounded-pointed; scales pale- to mid-brown ..........
. pseudodisjuncta.
25b. Lower pinnule-apices obtusely rounded-crenate; scales mid-brown or reddish

26.

26a. Scales brown D. cambrensis subsp. cambrensis*.
26b. Scales reddish or red 27

27a. Pinnule side-lobes wedge-shaped and pointed, pinnule-teeth markedly long, frond-

axes highly glandular . cambrensis subsp. insubrica.
27b. Pinnule side-lobes rounded or rounded-truncate, pinnule-teeth short, frond-axes
hardly or not glandular D. cambrensis subsp. distans.

[28a. Indusia large, markedly tall, thick, se eri splitting, scales mostly to almost all
narrow, lamina very glossy D. affinis* (included here for comparison)].
28b. Indusia small, not markedly tall, relatively thin, not splitting, shrivelling later,
scales include many wide ones, lamina matt or somewhat glossy 29.

29a. Pinnule-apices squarely truncate, though often with rounded corners, so becoming
truncately rounded 30.
29b. Pinnule-apices rounded to rounded-pointed 32.

30a. Indusia + flat to shallowly curved, but begin slightly inflected at their edges, then
shrivel rapidly; pinnule side-lobes (usually present on lower pinnules) square or

rectangular with pointed corners 31.
30b. Indusia markedly curved and begin well inflected at their edges with a central
depression before shrivelling later; pinnule side-lobes absent, or wedge-shaped and
pointe D. pseudodisjuncta.

31a. Stipe-base scales varying from + concolorous to having vaguely defined darker
bases and centres, or if well defined, the dark colour not extending upwards in narrow

streaks D. borreri*.
31b. Stipe-base scales bicolorous, pale, with glossy blackish streaks extending upwards
from a clearly contrasting dark base D. pontica.

32a. Pinnules well separated, pinnae long; scales very pale or whiteish ...................
D. cambrensis subsp. pseudocomplexa*.
32b. Pinnules crowded, pinnae short; scales reddish or bown

33a. Scales reddish or red D. cambrensis subsp. distans.
33b. Scales yellowish brown to brown D. cambrensis subsp. cambrensis*.

COMPARATIVE DIAGNOSTIC DESCRIPTIONS
1. D. affinis*.
1A. subsp. affinis*. The “purest” D. affinis morphology after D. wallichiana and
the most reminiscent of D. wallichiana in its flat frond, “plasticy” (like a moulded

plastic) upper surface. smooth laminar texture and regular, rounded-truncate pinnules.

FRASER-JENKINS.: THE DRYOPTERIS AFFINIS GROUP 17

Frond usually + wide (except when exposed on screes etc.), with a relatively short stipe,
except in large, luxuriant plants in woods; lamina generally tapering somewhat towards
the base. Stipe and rachis densely clothed with somewhat narrow scales which stick out
from the rachis, but not as much as in D. wallichiana; scales variable in colour, most
commonly mid- to deep golden-brown, with darker centres and bases, but in some
localised clones deep-brown to blackish, in others pale. Lamina thicker, more highly
coriaceous and markedly more glossy above than in the other taxa, of a noticeably
plastic-like texture above, with the veinlets impressed in the upper surface, somewhat
glaucous below with the veinlets darkened. Variable in colour above, but usually dark-
green except when exposed, when becoming yellower. Pinnae not, or hardly tapering
throughout most of their length, but extending to a long caudate apex, flat; pinnules
more regular in size and shape than in the other taxa, sometimes becoming + crowded
when growing in open places, though not overlapping, but usually + separated, or well
separated by a U-shaped gap (or “disjunction”), especially when growing in woods,
though when not markedly “disjunct” the pinnules may be slightly more widely
attached to each other at their bases than in the other taxa; usually + unlobed except for
a + rounded basal auricle on the lowest-opposite pair of pinnules on each pinna (though
occasional plants may have prominently lobed and/or even sharply toothed pinnules),
the lowest opposite-pair of pinnules on each pinna usually + the same length as the rest,
basiscopic pinnules at the bases of the lowest pinnae not usually becoming developed
or longer, with the lowest basiscopic pinnule of the lowest pinna either the same size
as, or often somewhat smaller than the rest and usually between 4% and % attached (or
adnate) to the pinna-costa, usually nearly all along the acroscopic side of its stalk with
most of the base on the other side free; pinnule-apices rounded to rounded-truncate
(rarely more pointed further up the frond in large plants), bearing rather few, obtuse
pinnule-teeth. Fronds eglandular. Sori the largest in the group, indusia thick, large. +
tall, somewhat pale until older, eglandular, markedly curved down and inwardly
inflected at the margins until the spores ripen, when the indusia lift slightly at the edges
and usually split open radially in one or more places as the sporangia turn black (the
splits ei difficult to see once the sporangia have dehisced), but hardly shrink and do
not normally shrivel up or lift completely (except in a few exceptional specimens),
persisting until the fronds begin to die. Spores relatively small and regular, ripening
later than in the other taxa. Fronds + persistent throughout most of winter.

1B. subsp. paleaceolobata*. Can be confused with D. cambrensis subsp.
cambrensis because of its lobed pinnules, but the pinnae are longer and more parallel-
sided than in D. cambrensis and the lamina is more widely lanceolate with a somewhat
wide base base or just above it, tapering from there to the apex, more glossy and with
more twisted pinnules including their side-lobes, and the stipe-scales are consistently
a ra the pinnule-lobes are also smaller. narrower and neater and the

ores are ler. Appears to be the nearest Atlantic equivalent to subsp. punctata but
i more lobed and without the * - tate” indentations (apart from very small ones)
above the sori. Also similar to p. affinis but differs somewhat markedly in its

pinnules all being lobed at the ae rst usually slightly irregularly twisted or tilted up
from side-to-side (seldom curved up at the apices as in D. cambrensis). The lowest

basiscopic pinnule is devel and becomes the longest and most lobed in = frond,
and _is fully stipitate. However it retains the dense, dark and narrow sti chis-

scales and markedly glossy lamina of D. affinis, as well as the selaideineds frond ee tall

18 FERN GAZ. 18(1): 1-26. 2007

though not as thick, eglandular indusium, which splits, but then, unlike in subsp. affinis,
shrivels somewhat on ripening. It also has similarly small spores and usually more
obtuse teeth, though these often become somewhat acutely deltate (acuminate) and
flabellately spread out around the pinnule-apex. The pinnule side-lobes are rounded and
not as rectangular as in D. borreri.

1C. subsp. punctata. Differs from subsp. affinis in having larger, more crowded,
often somewhat crinkled pinnules, bearing more side-lobes, including a pair of auricles
at their bases, and with less regularly parallel margins and more rounded pinnule-
apices; they also bear characteristic longer, more prominent and narrower, but obtuse-
tipped teeth splayed out around the pinnule-apices in a fan-like arrangement. The upper
surfaces of the pinnules have a marked circular, “punctate” indentation above each
sorus, which is highly characteristic, and deeper than the small ones that sometimes
occur in subsp. affinis etc. The lowest basiscopic pinnule on the lowest pinna is more
narrowly attached to the costa than is usual in subsp. affinis, normally being fully
stipitate, and the lower pinnules are usually markedly more lobed. Fronds eglandular.
The indusium is eglandular and usually (but not always) splits on sporangial ripening,
but is thinner than in subsp. affinis and lifts up and normally completely shrivels, except
in plants from exposed places, which have thicker and less shrivelling indusia. In its
scales, glossy, thick, usually dark-green, flat, regular lamina and large sori, it obviously
belongs to D. affinis.

1D. subsp. kerryensis*. Similar to D. cambrensis subsp. cambrensis, but with a
smaller, somewhat diminutive frond, and markedly flatter _and dark-green, glossy
lamina. Stipe- and rachis-scales narrow, dark-brown to blackish; pinnae flat, or slightly
convex from above, with the pinnules often slightly curved down at their edges and
apical halves. Pinnules markedly crowded, + rectangular, the lowest opposite-pair on
each pinna often bearing a few distinctive + wedge-shaped, pointed side-lobes and a
similar basal auricle. Pinnule-apices varying from rounded to more usually rounded- to

Square-truncate, often slightly sloping obliquely from one side to the other, bearing

somewhat long, acute teeth, but which are slightly wider up to their apices than in D.
borreri.

1E. subsp. jessenii. Perhaps distantly related to subsp. punctata, and like it, has
lobed pinnules and usually a fully stalked lowest basiscopic pinnule on the lowest pinna
(but sometimes when + unlobed, becomes half adnate to the costa in exposed plants, or
in dry situations), it also has smaller punctate depressions above the sori and somewhat
small and thin, but inflected indusia, some of which split before shrivelling; but it
differs in its usually markedly darker (usually almost black, at least at their bases) stipe-
and rachis-scales, its smaller, more disjunct, and irregularly margined pinnules, and
especially in at least the lower pinnule-apices being markedly squarely truncate, while
the upper ones in the frond may become pointed, but are not as rounded as in subsp.
punctata. The pinnule-teeth are characteristically longer and narrower than in the other
varieties of D. affinis, becoming somewhat finger-like, but not acuminate, with slightly
obtuse. if narrow apices, and are curiously irregular in length, slightly reminiscent of a
monstrosity or abnormality. Axes, lamina and indusia eglandular.
2. D. cambrensis*.

2A. subsp. cambrensis*. Intermediate between D. affinis and D. oreades, and the
nearest triploid to D. affinis in morphology. The British plants are usually the least
toothed, with the brownest (or occasionally pale) scales, though in places in north Wales
and the Cairngorm Highlands of Scotland they may become slightly more toothed, and

FRASER-JENKINS.: THE DRYOPTERIS AFFINIS GROUP 19

with more russet-brown scales, slightly approaching subsp. distans. The scales range
from pale, to, more usually, mid- to —— usually with darker bases, and are
glossy and usually slightly twisted; the larger ones are wider than in D. affinis and
sometimes more glossy. Stipe usually relatively ate except when growing in woods,
or between boulders. Lamina slightly thinner than in D. affinis, smooth, with the
— hardly im mr above, glossy above when compared with D. borreri, thou
as glossy as in D. affinis, usually darker-green (darker than in — insubrica)

eve paler or cawah when in the open, characteristically narrow, + ta g
towards the base. Pinnae short compared to D. affinis, tapering ae from their
wider bases, seldom flat as at least some of the + rounded or narrowly-rounded pinnule-
apices, especially the lowest basiscopic ones and lowest opposite-pair on each pinna,
are usually curved upwards out of the plane of the frond, and the pinnules sometimes
twist irregularly laterally, resulting in a slightly crisped appearance to the frond (but not
as markedly as in D. affinis subsp. paleaceolobata). Lowest opposite-pair of pinnules
of each pinna (i.e. the lowest acroscopic (upper) and basiscopic (lower) pinnule of a
pinna) the longest, and usually overlapping the rachis, the lowest one often deflexed
basiscopically (i.e. bent slightly towards the base of the pinna) so as to lie slightly over
the rachis, the lowest basiscopic pinnule of the lowest pinna is usually fully stalked, or
stipitate. The pinnules vary from well lobed with rounded lobes to almost unlobed, the
pinnule-apices are rounded and vary from fairly prominently toothed to more frequently
small-toothed or + untoothed and in either case have shorter and more obtuse teeth than
in subsp. insubrica. In some specimens there are a few (up to c. 30) glands on the edge
of the indusium, or the frond-axes are sparsely glandular, but most plants are +
eglandular. Sori smaller, or nearly the same size as in D. affinis, the indusia are
somewhat thick, thicker than in D. borreri, though with a thinner margin than in D.
affinis, + tall, somewhat pale. or greyish, until old, markedly curved down and inflected
as in D. affinis, but when the spores ripen the indusia frequently split, then lift up. to lie
over the top of the sorus, finally shrivelling considerably and after some time, dropping
off, but not shrivelling as much as in D. borreri and persisting longer, becoming brown.
Spores markedly larger than in D. affinis and containing a markedly higher proportion
of abortive ones; also — darker brown in colour. Fronds turning brown and
dying down in mid Autu

2B. subsp. distans. Pi glossy: between. saben,
cambrensis and subsp. insubrica, but closer to the former except in scale-colour. Fronds
are often taller than in subsp. insubrica; pinnules with small, rather insignificant teeth,
occasionally becoming longer and slightly flabellate; pinnule-lobes when present
rounded, not cuneate. Axes and indusia eglandular or bearing only few, scattered, small
glands. Stipe-scales markedly reddish or reddish-brown, thin, often rather twisted.
Some British and Norwegian populations of subsp. cambrensis approach it, but the
distinction between the two is less clear in parts of those areas.

2C. subsp. insubrica. Strongly reminiscent of the pale russet-scaled, glandular and
somewhat narrowly toothed forms of D. oreades of Corsica, Sardinia, Elba and northern
Italy. More acute-toothed, pointed-lobed and with more reddish-russet coloured scales
than in subsp. cambrensis. The larger stipe- and rachis-scales are wider than in
affinis, more glossy and slightly thicker, often + concolorous, ranging from pale yellow
(particularly in eastern Europe), to pale-russet, to a more common and characteristic
reddish or russet-brown, often with darker, castaneous bases and some dark flecks or
streaks in the scales; some of the narrower ones characteristically exserted and

20 FERN GAZ. 18(1): 1-26. 2007

somewhat twisted spirally, particularly on the upper stipe and lower rachis. Stipe
usually relatively short except in large, luxuriant plants in woods, or when growing
between boulders. Lamina narrow, + tapering towards the base, slightly thinner and less
highly coriaceous than in D. affinis, but of a slightly crispaceous texture, + matt above
veinlets usually slightly impressed in the upper surface. Variable in colour above, but
often dark- to mid-green, except when exposed, when yellowish. Pinnae usually short,
tapering from their wider bases, + flat. Pinnules rather irregular in length, crowded and
sometimes slightly overlapping, though they may become separated by a + narrow, V-
shaped gap; the lowest opposite-pair on each pinna usually longer than the rest, and
usually close to the rachis or somewhat overlapping it, the lowest oe pint one on
each pinna may sometimes be slightly basiscopically deflexed so as to lie partly o
the rachis, the lowest basiscopic pinnule of the lowest pinna is sulfa
stipitate. The lowest opposite-pair of pinnules on each pinna is usually well lobed, w
characteristic wedge-shaped, pointed side-lobes. Pinnule-apices rounded to Smee
acne as in D. affinis, but often becoming square-truncate above, bearing prominent,
ong, + narrow, but _wide-based, usually somewhat acute-tipped teeth, which are
Pars i.é. splayed out around the apex in the manner of D. oreades. Many small,
fairly scattered, or dense, white glands on the frond-axes and on the indusia as well.
ori usually smaller than in D. affinis, usually also slightly thinner, though thicker than
in D. borreri, + tall, somewhat pale, or greyish, until old, usually glandular near their
margins, markedly curved down and inflected as in D. affinis, but when the spores ripen
the indusia frequently split, then lift up to lie over the top of the sorus, finally shrivelling
considerably and after some time dropping off, but not shrivelling as much as in D.
borreri_and_ persisting longer, usually remaining paleish to pale-brown until
considerably later, when becoming brown. Fronds turning brown and dying down early
in Autumn.
2E. subsp. pseudocomplexa*. Somewhat similar to D. x complexa, but with a paler
coloured lamina and smaller, narrower, less lobed pinnules, and with a high proportion
of good spores. It also differs in forming extensive populations. Generally similar to
subsp. cambrensis, but with pale to very pale-russet. thin, but slightly glossy. ovate
stipe-base scales; the lamina thin. pale- to mid-green, wider and more lax than in subsp.
cambrensis; pinnules usually rather widely separate, or at least not as crowded, long and
narrow, with the side-lobes smaller and neater when present. Pinnule- -apices narrowly
rounded, often becoming somewhat pointed or rounded-pointed in larger plants,
varying from being + toothless to bearing small, somewhat narrow, often acute teeth,
more acute than in subsp. cambrensis. Indusia somewhat thin, inflected at first, not as
tall as in subsp. cambrensis and not, or seldom splitting, soon lifting and shrivelling
markedly.

3. D. pseudodisjuncta.

Stipe- and rachis-scales + pale, usually with darker-brown bases; somewhat abundant
narrow, hair-like, pale scales present on the costae on the underside of the pinnae. Frond
wider than in D. cambrensis; lamina slightly glossy; pinna-axils very occasionally
without the dark spot (darkened area of the petiole at the junction beneath) ; pinnae
ong, markedly flat — + regular in length, or ~ first pair slightly the longest i in
each pinna, usually from bases

V-sh ap mi upper pi what widely attache

oe other at their bases, on fa) o their in owest pai innul:

FRASER-JENKINS.: THE DRYOPTERIS AFFINIS GROUP 21

in the lower pinnae which thus become + wedge-shaped below, + unlobed (including
the lowest basiscopic pinnules of the lowest pinnae) apart from + rounded (occasionally
more rhombic) basal auricles in the basal pinnules of mid and upper pinnae in some
plants, the lowest opposite-pair of pinnules of each pinna usually slightly longer than
the rest and often somewhat deflexed towards the rachis (which rarely occurs also in D.
borreri); the lowest basiscopic pinnule of the lowest pinna (i.e. the lowest one of the
frond) sometimes becomes longer and developed, but is not usually the longest one,
fully stalked or stipitate, pinnules usually sloping towards their narrow apices; pinnule-
apices narrow or wedge-shaped, varying in the upper part of the frond from rounded to
rounded-truncate (or often somewhat markedly truncate), but characteristically
rounded-pointed in the lower pinnae. Fronds eglandular. Indusium tall, eglandular,
small, + thin, mid- to dark-brown, but greyish when younger, matt, markedly inflected
at the margins, with a + obvious depression in the centre that becomes noticeable when
the indusium begins to lift slightly at the edges on ripening; often splitting before lifting
slightly on ripening, though the edges remain downturned and enclosing much of the
top of the sorus; only later, after the spores have been shed, do the edges lift up
markedly before the indusium shrivels. Fronds turning brown and dying down in mid
or late Autumn.

4. D. schorapanensis.
Close to D. filix-mas and often mistaken for it. Frond + wide with a long stipe, tapering
slightly to the lamina-base. Stipe and rachis not very densely clothed with scales; scales
thin, + wide, pale to pale-brown, concolorous, or with a dark base, or sometimes a few
dark streaks in the centre. Lamina as thin as in D. borreri, or thinner, hardly coriaceous,
+ matt above, slightly glaucous below, lighter green than in D. borreri, veinlets slightly
impressed above, not darkened. Pinna-axils usually without the dark spot normally
present in the living state in all the other species and subspecies (but see under D.
pseudodisjuncta); pinnae flat; pinnules + regular in length, or the lowest opposite-pair
of each pinna slightly the longest, seldom much separated, often somewhat narrowed
towards their bases near the bottom of each pinna, + unlobed, or with insignificant
shallow, + square lobes, the lowest basiscopic pinnule of the lowest pinna not usually
becoming longer and developed, and not the longest one, fully stalked or stipitate;
pinnule-apices truncate to rounded- truncate in the lower part of the frond, often
becoming more pointed and wedge-shaped in the upper part of the frond, bearing
characteristic subulate, wide-based teeth, cnr abruptly narrowed to + and often
rather irregularly long-acute points and spread out around the apex in a flabellate. or
fan-like arrangement. Fronds eglandular. Sori similar to D. borreri, the indusium
eglandular, pale, starting only slightly curved down and not much inflected at the
margins (less so than in D. borreri), until lifting, when it shrinks but does not split,
ither sitting + flat over the top of the ripe sporangia until dropping off, or shrivellin
considerably, though more persistent than in D. borreri. Spores large and long, with
much abortive material present. Fronds turning brown and dying down early in

Autumn.

5. D. pontica.

Similar to D. schorapanensis, but with the truncate lobes and _pinnule-

borreri. Stipe-base scales with dark bases and prominent, separati

streaks extending upwards into the pale area of the scale. Lamina not markedly glossy.

22 FERN GAZ. 18(1): 1-26. 2007

Pinnules becoming wider towards their bases and somewhat remote towards their
apices, with obliquely sloping, truncate-tipped basal lobes on each side and often a few
smaller lobes above; pinnule apices varying from obtuse to more usually narrowly
truncate, with rather prominent teeth. Indusia thin, beginning well turned down, but
lifting and partly shrivelling on ripening. Spores larger than in D. borreri, with a similar
proportion of abortive spores to that in D. cambrensis. Fronds turning brown and dying
down early in Autumn.

6. D. borreri.

sually nearer in morphology to D. filix-mas than are the other taxa (except D.
schorapanensis and D. cambrensis subsp. pseudocomplexa). Frond usually + wide and,
except when exposed, with a relatively long stipe, the lamina not usually tapering much
below, or not as much as in the other taxa (excluding D. schorapanensis), except in
more exposed plants and rarely in some localised clones. Stipe and rachis sometimes
less densely clothed in scales than in the other taxa (except D. schorapanensis and D.
cambrensis subsp. pseudocomplexa) and the scales often paler (sometimes with dark
bases) and wider, though very variable. Lamina relatively thinner (except in exposed
plants) than the other taxa (except D. schorapanensis and D. cambrensis subsp.
pseudocomplexa), less coriaceous, hardly glossy, but more matt above, and usually
lighter green than in D. affinis, though the colour varies considerably, veinlets not
impressed above or darkened beneath, lamina not glaucous beneath. Pinnae + flat;
pinnules not as regular in length as in D. affinis, adjacent, but not crowded, though
seldom much separated (though occasionally so, by V- or rarely U-shaped gaps), the
lower pinnules on lower pinnae, particularly the lowest basiscopic one of the frond.
usually lobed with characteristic + rectangular side-lobes, and a larger, rectangular
basal auricle (though they can be absent), the lowest opposite-pair of pinnules of each
pinna often curved slightly away from the rachis, usually + the same length as the rest,
except that the basiscopic one frequently becomes a little longer, at least in the lower
pinnae, the lowest, or second lowest basiscopic pinnule of the lowest pinna is usually
the longest in the frond, is often markedly developed and lobed with rectangular side-
lobes, and is fully stalked or stipitate; pinnule-apices characteristically markedly
squarely truncate in the lower parts of the frond, but becoming rounded or pointed,
particularly in the upper frond, in many larger specimens, but at least some of the won
or mid-frond pinnule-apices usually remain truncate, rounded-truncate, or obovate, a
opposed to their all being more rounded as in D. affinis and in most uD. cambrensis
subsp. cambrensis etc.; in normal plants the pinnule-apices bear lon
pein acute teeth, similar to those in D. filix-mas, but some plants are occasionally

t toothless, when they can become difficult to separate from D. “8

poreogene in the truncate-pinnuled plants the teeth may be longer above each corner
of the pinnule than in the centre of the apex (the late H.V. Corley’s “cat’s “Soaee outline
- the corner teeth representing the cat’s ears). Fronds eglandular. Sori oncom than in D.
affinis or D. cambrensis, and indusia noticeably relatively thin and 1, whiteish or
pale- to mid-brown, eglandular, beginning curved down at the sides a not inflected,
or turned in, as in D. affinis and D. cambrensis) until the spores ripen, when they lift

right up. without splitting. shrivel and shrink markedly to become a very small,
inverted, wri cone, or el on top of the fully exposed i soon dro
off. Spores relatively larger and with more abortive material than in D. affinis and quite
often with as many abortive spores as in D. cambrensis, but they are usually more

FRASER-JENKINS.: THE DRYOPTERIS AFFINIS GROUP 23

regular_and often noticeably smaller than in the other triploid taxa, and can be
considerably smaller than expected. Spores ripen from two weeks to up to nearly a
month earlier than in D. affinis. Fronds mostly turn brown and die down early in winter,
though some persist longer.

The most widespread and commonest member of the group, though absent from
Macaronesia, north-west Africa and southernmost mainland Italy and less common in
Portugal and Spain than subsp. affinis.

7. D. affinis agg. hybrids with D. filix-mas.
The frond-morphology does not fit in with any of the other taxa in the D. affinis agg.
and_is intermediate between the parents, i.e. markedly towards D. filix-mas when
compared with that of any other members of the D. affinis agg., though the lamina
clearly has more of the glossiness and darker colour (especially in D. x complexa) of D.
affinis than is normal in D. filix-mas. Fronds and segments often very large (though this
can also occur sometimes in the rest of the D. affinis agg., especially in D. affinis subsp.
affinis and in D. borreri and occasionally also in D. cambrensis subsp. cambrensis), the
lamina is sometimes rather narrow in the lower part of the frond (particularly in D. x
complexa and D. x convoluta). The stipe bears denser, ae and narrower scales than
in D. filix-mas (but it should be noted that occasionally D. filix-mas can also be very
densely scaly), which often become more comand and slightly towards those of
D. cambrensis subsp. insubrica; the pinnules of the lower pinnae have more parallel
sides than in D. filix-mas and sometimes slightly more rectangular side-lobes, if present,
though not as parallel and rectangular as in D. borreri, but they become much more like
D. filix-mas with more curved and sloping sides and more pointed lobes and pinnule-
apices further up the frond. The pinnule-teeth are + acute, though less so in D. x
complexa. The sori (except in D. x complexa) have less inflected indusia than in D.
affinis etc. The spores, which are most important [examined dry at c. 100-150 x
magnification when ripe, yet not mostly shed and lost], are _characteristically highly
abortive, without which feature being examinable it is difficult to confirm D. x
complexa except in very obvious specimens. Apart from the mass of abortive spores
there is also a small and variable percentage (c. 5-20%. occasionally up to c. 50%) of
mostly very large, or exceptionally large, sometimes nearly spherical, apparently good
spores present. The all-good, very small spores of D. filix-mas, by contrast, are instantly
separable (and much smaller than in any species or subspecies of the D. affinis agg.),
while those of the species and subspecies of the D. affinis agg. are usually at least 50%,
to mainly good, unless nearly all shed. The abortive spores of the hybrids may be many
small fragments (possibly confusable with slightly immature spores, but smaller and
with more developed and darker perispores), or misshapen spores with an irregular,
often darker perispore (not to be confused with shapeless, dark pieces of earth), with
casional tient a or tetrads amongst them and usually with some perisporial
az angial) debris attached to their surfaces, also with dark debris remaining in the
sporangia (not to be eee with earth or dust). They are usually more irregular and
often with wider wings of perispore than in the less numerous abortive spores of any
other members of the D. affinis agg. The hybrids occur patent but fairly
commonly, as occasional plants among populations of the parents. not fo g
populations of their own, except some scattered rather small local, seers mostly
vegetative ones, and seldom more than 1-5 clumps per local wood, except in a few
favoured habitats where the parents have long been together in quantity, or conditions

24 FERN GAZ. 18(1): 1-26. 2007

are very luxuriant, damp and favourable for spore-germination, when the hybrid can
become more numerous. Rarely large-sized populations of hybrids may occur, for
example, in places in south-western Ireland, and are likely to be discovered
occasionally elsewhere in Atlantic Britain etc. Hybrids can occur throughout the range
of the D. affinis agg. as D. filix-mas occurs in all the range of the D. affinis agg. as well
as in its own, wider range. Without being properly familiar with all the taxa of the D.
affinis agg. it is difficult to recognise, or even accurately guess the presence of genuine
hybrids. In the past D. borreri was widely misidentified as being such hybrids and was
often reported under the name D. x tavelii Rothm.

8. D. wallichiana subsp. wallichiana.

Morphology like an extreme D. affinis. Frond persistent in winter, tall and usually
tapering towards the base. Stipe short, except in occasional luxuriant wide-fronded
specimens with large segments; stipe and rachis densely clothed with longer, narrower
scales than in any of the D. affinis agg.. which stick out markedly from the rachis.
Lamina more highly coriaceous and stiff. more glossy above and glaucous below and
more crispaceous than in any of the D. affinis agg. Pinnae flat and regular; pinnules +
crowded, usually markedly squarely truncate at their apices, those in the upper parts of
the frond more widely joined together at their bases than in D. affinis; pinnule-teeth few,
more obtuse and short, though usually with an acute tip at their apex. Fronds eglandular.
Sori smaller than in D. affinis, with thick, inflected, eglandular indusia that split on
ripening and persist for a long time. Spores relatively small and regular (diploid
apomict).

A pan subtropical montane taxon. Not in the European (sens. lat.) flora, where it is
replaced by the W. Asian D. iranica (usually with a longer stipe, wider lamina-base,
larger segments; triploid apomict, and with larger and less regular spores).

ACKNOWLEDGEMENTS
The author wishes to thank the following for their various help and contributions to this
study:

Prof. Mary Gibby, Edinburgh; the late Dr. Trevor Walker, Newcastle; Miss Alison
Paul, London; Miss Josephine Camus, London; Mr. Peter Edwards, Kew; the late Prof.
I. Manton, Leeds; Mr. Ken Trewren, Egton Bridge; Mr. Anthony Pigott, Mendlesham;
Mr. Alfred Eschelmiiller, Sulzberg; the late Mr. Hugh Corley, Faringdon; Prof. Gabor
Vida, eeu the late Dr. Walter Gatzi, St. Gallen; M. Michel Boudrie, Clermont

ran . Mike Taylor, Isle of Skye; Mr. Martin Rickard, Kyre Park; Dr. Jacob
Schneller, Zirich; Dr. Stefan Jessen, Chemnitz; Herr Jens Freigang, Germany; Herr
Walter Bujnoch, Germany, and above all his colleague of 25 years, the late Prof. Tadeus
Reichstein, Basel and Agarone.

The great many correspondents and the staff of nearly 200 herbaria who have
assisted in his inquiries have been acknowledged in full in the forthcoming study by
Fraser-Jenkins & Trewren (in prep.).

REFERENCES
BAR, A. & ESCHELMULLER, A. 1985. Tetraploide und pentaploide Dryopteris x
tavelii - - jetzt im Allgau bestatigt. Mitt. Naturwiss. Arbetskr. Kempten (Allgiu) 27
(1): 57-68.
BAR, A. & ESCHELMULLER, A. 1989. Beitrag zur Kenntnis von Dryopteris affinis

FRASER-JENKINS.: THE DRYOPTERIS AFFINIS GROUP 25

(Lowe) Fraser-Jenkins und von Bastarden mit Dryopteris filix-mas. Mitt. Naturwiss.
Arbetskr. Kempten (Allgiu) 29 (1): 25-48.

BAR, A. & ESCHELMULLER, A. 1990. Dryopteris x complexa nssp. contorta Fraser-
Jenkins - ein seltener Farnbastard in Bayern. Ber. Bayer. Bot. Ges. 61: 91-97.

DANSER, B.H. 1950. A theory of systematics. Bibl. Biotheor., ser. D, 4: 117-180.

DAVIS, P.H. & HEYWOOD, V.H. 1970. Principles of Angiosperm Taxonomy: 10.
Cambridge.

DOPP, W. 1941. Uber Dryopteris paleacea Christensen (D. borreri Newm.). Ber.
Deutsch. Bot. Ges. 59 (9): 423-426.

DOPP, W. 1955. Experimentell erzeugte Bastarde zwischen Dryopteris filix-mas (L.)
Schott und D. paleacea (Sw.) C.Chr. Planta, Arch. Wissensch. Bot. 46: 70-91.

DOPP, W., GATZI, W. & OBERHOLZER, E. 1963. Dryopteris borreri Newman (D.
paleacea Hand.-Mazz.) var. pseudodisjuncta v. Tavel. Ber. Deutsch. Bot. Ges. 76
(4): 99-111.

ESCHELMULLER, A. & SCHNELLER, J.J. 1980. Beitrag zur Kenntnis der
Variebilitét von pera affinis im Allgau. Mitt. Naturwiss. Arbetskr. Kempten
(Allgau) 24 (1): 1-1

FRASER-JENKINS, 2 R. 1980. Dryopteris affinis: a new treatment for a complex
species in the European Pteridophyte flora. Willdenowia 10 (1): 107-115.

FRASER-JENKINS, C.R. 1982. Dryopteris in Spain, Portugal and Macaronesia. Bolm.
Soc. Broteriana, ser. 2a, 55: 175-335.

FRASER-JENKINS, C.R. 1983. Dryopteris affinis, in DOSTAL, J., FRASER-
JENKINS, C.R. & REICHSTEIN, T., in HEGI, G, Illustrierte Flora von
Mitteleuropa, ed. 3, 1, pp. 136-169. Berlin.

FRASER-JENKINS, C.R. 1987. Taxonomic and Nomenclatural notes 5, Dryopteris, in
DERRICK, L.N., JERMY, A.C. & PAUL, A.M., Checklist of European
Pteridophytes. Sommerfeltia 6: x-xiii.

FRASER-JENKINS, C.R. 1996a. Dryopteris affinis subsp. pseudodisjuncta (Oberh. &
Tavel ex Fras.-Jenk.) Fras.-Jenk., comb. nov., in JONSELL, B. (ed.),
Lectotypifications and new combinations for Flora iced Vol. 1 (Lycopodiaceae
— Papaveraceae). Nordic J. Bot. 16 (1): [3-] 4 [-8]. [not published in Nordic J. Bot.
15 (1995), as stated in Flora Nordica]

FRASER-JENKINS, C.R. 1996b. A reaffirmation of the taxonomic treatment of
Dryopteris affinis (Dryopteridaceae: Pteridophyta). Fern Gaz. 15 (3): 77-81.

FRASER-JENKINS, C.R. & TREWREN, K. in prep. The Dryopteris affinis complex

and its species, subspecies and hybrids.

GATZI, W. 1961. Uber den heutigen Stand der Dryopterisforschung unter besonderer
Beriicksichtigung von Dryopteris Borreri Newman, zugleich ein Beitrag zur
Farnflora des Siidabhanges des Tannenbergs. Ber. iib. Tatigkeit St. Gallisch. Naturw.
Ges. 77: 1-73.

ICBN: GREUTER, W. et al. 2000. International te of Botanical Nomenclature (Saint
Louis Code). Regnum Vegetabile 138. KG6nigste

JERMY, A.C. & CAMUS, J.M. 1991. The PP Field Guide to Ferns and Allied
Plants of the British Isles. London.

JERMY, A.C., PIGOTT, A.C. & MERRY WEATHER, J.W. 1998. Dryopteris, in RICH,
T.C.G. & JERMY, A.C. (eds.), The Plant Crib, pp. 29-33. London.

JESSEN, S. 1985. A reappraisal of Dryopteris affinis subsp. borreri var. robusta and
new records of D. affinis subspecies in eastern Europe. Fern Gaz. 13 (1): 1-6.

26 FERN GAZ. 18(1): 1-26. 2007

JESSEN, S. 1991. Neue Angaben zur Pteridophytenflora Osteuropas. Farnblatter 23:

JOHNS, R.J., EDWARDS, & PICHI SERMOLLI, R.E.G. 1996. Index Filicum
Supplementam sextum.

MANTON, I. 1950. Pawan of Cytology and Evolution in the Pteridophyta.
Cambridge.

MERRY WEATHER, J.W. 2002. Identification — British male ferns. Pteridologist 4 (1):
12-1

PICHI SERMOLLI, R.E.G. 1960. The identification of Aspidium distans Viv. American
Fern J. 50 (1): 133-138.

PIGOTT, A.C. 1997. Affinis Watch Newsletter, Special Issue British Pteridological
Society, insert in Pteridologist 3(

RASBACH, H., RASBACH, K., REICHSTEIN, T. & SCHNELLER, J. 1983.
Tetraploide Dryopteris x tavelii Rothm. im nérdlichen Schwarzwald. Farnblatter 10:
1-13.

REICHSTEIN, T. & SCHNELLER, J. 1983. Dryopteris affinis var. punctata im
Hiillerich-Wald ob Pfaffikon (SZ). Farnblatter 9: 9-21.

ROTHMALER, W. 1945. Der Formenkreis von Dryopteris paleacea (Sw.) Hand.-Mazz.
Candollea 10: 91-101.

RUBE, G. & HEISE, K. 1975. Neufund von Dryopteris borreri Newm. (Dreiwertiger
Speuschuppenfarn) in N6rdosthessen. Beitrige zur Naturkunde in Osthessen 9-10:
169-173.

SCHNELLER, J.J. 1974. Untersuchungen an einheimischen Farnen, insbesondere der
Dryopteris filix-mas-Gruppe 1.Teil. Ber. Schweiz. Bot. Ges. 84 (3): 195-217.

SCHNELLER, J.J. 1975a. Untersuchungen an einheimischen Farnen, insbesondere der
Dryopteris filix-mas-Gruppe 2.Teil. Cytologische Untersuchungen. Ber. Schweiz.
Bot. Ges. 85 (1): 1-17.

SCHNELLER, J.J. 1975b. Untersuchungen an einheimischen Farnen, insbesondere der
Dryopteris filix-mas-Gruppe 3.Teil. Okologische Untersuchungen. Ber. Schweiz.
Bot. Ges. 85 (2): 110-159.

WIDEN, C.-J., FRASER-JENKINS, C.R., REICHSTEIN, T., GIBBY, M. &
SARVELA, J. 1996. Phloroglucinol derivatives in Dryopteris sect. Fibrillosae and
related taxa (Pteridophyta, Dryopteridaceae). Ann. Bot. Fennici 33: 69-100.

FERN GAZ. 18(1):27-29. 2007 27

SELAGINELLA AGASTYAMALAYANA (SELAGINELLACEAE:
PTERIDOPHYTA), A NEW SPECIES FROM SOUTH INDIA

RAJU ANTONY, A.E. SHANAVAS KHAN & G SREEKANDAN NAIR
Tropical Botanic Garden and Research Institute,
Palode, Thiruvananthapuram — 695 562,
Kerala, India

Key words: Fern ally, Selaginella agastyamalayana, India

ABSTRACT
Selaginella agastyamalayana, a new species (Selaginellaceae: Pteridophyta) is
described from Kerala, South India.

INTRODUCTION
The genus Selaginella P. Beauv. comprises about 700 species distributed throughout the
world, but mainly concentrated in tropical and subtropical areas. Dixit (1992) recorded
62 species in India. Later Madhusoodanan and Nampy (1994) added one more species
from Kerala.

As part of our floristic survey and ex-situ conservation of pteridophytes in the
Western Ghats, South India, we made repeated plant exploration trips to Agastyamala,
one of the well-known ‘hot spots’ of South India, which resulted in the collection of
interesting specimens of Selaginella growing in wet rock crevices (1,750 m alt.) in
evergreen forests. On careful examination it proved to be a species new to science. The
new species is described here and named after the type locality. Selaginella
agastyamalayana is known so far only from the type locality.

Selaginella agastyamalayana Raju, Khan & Nair sp. nov. Figure 1.
Caulis gracilis, 3.5-7 cm, prostratus, sursum a ramosum, ramus brevis. Rhizophora
longa, tenuis, filo metallico similis, omnio plant praesentia. Folia heteromorpha, pallide
virens, remote posita, membranacea; folia lateralis 1-2 x 0.8-1.5 mm, suborbicularia,
obtusa ad apicem, cordata ad basim, margine denticulata; folia axillaris suborbicularia,
obtusa ad apicem, cordata ad basim, margine denticulata; folia medianus 0.7-1 x 0.2-0.3
mm, ovata, accuminata vel aristata ad apicem, cordata ad basim, margine denticulatus.
. 2.5-4 x 1.8-2 mm, terminalis, singularis vel duplex ad ramus et ramulus;
sporophyllum dimorphus; sporophyllum magnus 1-1.5 x 0.4-0.8 mm, ovato-oblongus,
subacutus ad apicem, cordatus ad basim, margine denticulatus; sporophyllum parvus
1-1.2 x 0.6-0.7 mm, late ovatus, acuminatus ad apicem, cordatus ad basim, margine
denticulatus. Megasporae pallid luteae, 173-313 um, spinulosae. Microsporae fortiter
ourantiaceae, 23-33 um, verrucoidae

Holotypus: Agastyamala, Tidavanaithoguiain District, Kerala, South India, 1,750
m. alt.; in evergreen forest; 9 January 1996; Raju Antony 25580 ( 7).

Stems slender, 3.5-7 cm, prostrate, branched from the base, branches short.
Rhizophores long, thin, wiry, present throughout the plant. Leaves heteromorphic
throughout, pale green, membranous; lateral leaves distant on main stem and branches,
1-2 x 0.8-1.5 mm, suborbicular, obtuse at apex, cordate at base, margin denticulate;
axillary leaves suborbicular, obtuse at apex, cordate at base, margin denticulate; median
leaves distant on main stem but subcontiguous on branches, 0.7-1 x 0.2-0.3 mm, ovate,

28 FERN GAZ. 18(1):27-29. 2007

acuminate to aristate at apex, cordate at base, margin denticulate. Strobili 2.5-4 x 1.8-2
mm, terminal, single to double at the branches and branchlets; sporophylls dimorphic;
larger sporophylls in the same plane as the median leaves, 1-1.5 x 0.4-0.8 mm, ovate-
oblong, subacute at apex, cordate at base, margin denticulate; smaller sporophylls 1-1.2
x 0.6-0.7 mm broadly ovate, acuminate at apex, cordate at base, margin denticulate.
Megaspores dull yellow, 173-313 um in diameter, spinulose. Microspores deep orange,
23-33 um in diameter, verrucoid.
Other specimens examined: Agastyamala, Thiruvananthapuram District, Kerala,
= December 1998; Raju Antony 29880 (TBGT).
e: This species is allied to Selaginella miniatospora (Dalz.) Bak. But can be easily
aes from it by prostrate stems, rhizophores present throughout the plant,
suborbicular lateral leaves with obtuse apices, median leaves acuminate to aristate at

Figure 1. Selaginella agastyamalayana
A - habit; B - lateral leaf; C - axillary leaf; D — median leaf; E — strobilus; F — larger
sporophyll; G — smaller sporophyll.

RAJU ANTONY et al.: SELAGINELLA AGASTYAMALAYANA 29

apex, denticulate sporophylls, spinulose megaspores as against stems erect, rhizophores
restricted to basal one fourth, ovate-oblong lateral leaves, subacute at apex, median
leaves cuspidate at apex, dentate sporophylls and papillate megaspores.

ACKNOWLEDGEMENTS
The authors thank Dr V.B. Hosa Goudar, Tropical Botanic Garden and Research
Institute, Palode, for the latin diagnosis. We also thank E.S. Santhosh for his help and
S. Suresh Kumar for the illustration.

REFERENCES
DIXIT, R.D. 1992. Selaginellaceae of India. Bishen Singh Mahendra Pal Singh, Dehra

un.
MADHUSOODANAN, P.V. & SANTHOSH NAMPY, 1994. Selaginella dixitii, a new
species of Selaginellaceae from India. Nordic J. Bot. 14: 527-529.

30 FERN GAZ. 18(1). 2007

BOOK REVIEW

CHANGE IN THE BRITISH FLORA 1987-2004 (A report on the BSBI Local
Change Survey). Braithwaite, M.E., Ellis, R.S. & Preston, C.D. 2006. Softback,
382pp., ISBN 0 901158 34 8. Botanical Society of the British Isles, London. £12.00.

Superficial comparison of the distribution maps in the New Atlas of the British and Irish
Flora sai et al 2002) with those recorded in the Atlas of the British Flora (Perring
& Walters 1962) show many changes resulting from a variety of causes — increased
seteniiuisots changes in farming practices, spread of non-native invasive species, or
climate change. A survey of species from 811 tetrads in a regular grid across Britain,
carried out in 1987 by BSBI recorders, provided a firm basis for further research to re-
survey these sites and measure distributional changes for individual species. This
publication is the result of the excellent collaboration between the valuable network of
BSBI recorders and staff of the Biological Records Centre (BRC).
range of species covered include native and long-established introductions, and
more recent introductions as well as some rare or very widespread species. In reporting
the results, the species are grouped under broad habitats — coniferous woodland,
calcareous grassland etc., to try to identify patterns of change within a habitat.
at is happening to ferns? Their inclusion is clear from the front cover, with an
illustration of Hart’s Tongue Fern, Asplenium scolopendrium (referred to here as
Phyllitis scolopendrium). This species is included in two broad habitats, ‘Broad-leaved,
mixed and yew woodland’, and ‘Inland rock’ that includes cliffs, screes, quarries and
limestone pavement. In both habitats it seems to be increasing its range, and in Scotland
appears to be colonising walls and open woodland, perhaps in response to a reduction
in frosts. Similarly, Polystichum setiferum, another woodland species, also shows
significant increase and may also be benefitting from milder winters. In contrast
Cystopteris fragilis, more commonly found in rock outcrops or walls in the north and
west, shows decline, particularly in lowland sites that may no longer provide damp,
shaded conditions. Large decline is also recorded for Dryopteris expansa and D.
oreades, although the results of their analyses are based on small samples.
This book raises a lot of questions. It also highlights the importance of good and
accurate recording and, I hope, stimulates further activity in BPS field meetings.

REFERENCES
PERRING F.H. & WALTER, S.M., eds. 1962. Atlas of the British Flora. London:
Thomas Nelson & Sons.
PRESTON, C.D., PEARMAN, D.A. & DINES, T.D. 2002. New Atlas of the British and
Irish Vascualar Plants. Oxford: Oxford University Press.

M. Gibby

FERN GAZ. 18(1): 31-38. 2007 31

DYNAMICS OF LONG-DISTANCE DISPERSAL: THE SPREAD OF
ASPLENIUM ADIANTUM-NIGRUM AND ASPLENIUM TRICHOMANES
(ASPLENIACEAE: PTERIDOPHYTA) ON LONDON WALLS

J.A. EDGINGTON
19 Mecklenburgh Square, London WCIN 2AD

ABSTRACT

Asplenium adiantum-nigrum L. and A. trichomanes L. have spread widely
across inner London since about 1995 but have not yet colonised the district
uniformly. The spacing between sites, and evidence that at most a few founding
populations are involved, suggests that a sequence of long-distance dispersal
events is responsible. A simple model of dispersal is proposed which predicts an
exponential distribution of nearest-neighbour distances. The combined data for
both species yield a good fit to such a distribution and a value of approximately
0.75 km for the mean distance between a newly-established sporophyte and the
nearest potential parent plant. This leptokurtic distribution accounts for some
offspring growing 5 km or more from their parent while the populated zone
expands at up to | km per year. Assumptions underlying these conclusions are
examined.

INTRODUCTION

It is well-known that plants can colonise fresh sites, such as oceanic islands, at great
distances from existing populations but there are few experimental studies of long-
range dispersal — in the case of seed plants, virtually none (see Cain, Milligan and
Strand, 2000, for a review). The likelihood of establishing spore-producing plants as a
function of distance from the source sporophyte is likewise poorly understood. Though
measurements have been reported of atmospheric concentrations of spores at varying
distance from their apparent source, the complex relationship between spore dispersal
and viability on the one hand, and subsequent germination and development on the
other, means that these data cannot be taken to be representative of the likelihood of
sporophyte establishment as a function of distance from the parent plant. Knowledge of
this would provide both reference data for the phytogeography of spore plants and
constraints on models of migration and expansion.

There are few instances in which the spread of bryophytes and pteridophytes into a
virgin habitat has been followed closely in time and space. A recent example is a survey
(Miller and McDaniel, 2004) of the bryophyte flora of a road driven 65 years ago
through the Adirondack Mountains in New York State. By observing the presence, in an
otherwise acidic natural terrain supporting only a calcifuge flora, of a large number of
calcicole bryophytes on man-made mortared structures, Miller and McDaniel
concluded that some bryophytes may be capable of routine dispersal over distances of
at least 5 km, and that long-distance colonisation had occurred at the rate of at least one
species per year.

The present paper describes an analogous case involving ferns observed in the
process of occupying a habitat which has recently become able to support a flourishing
pteridophyte flora. The ferns concerned are calcicole epiliths and the habitat is mortared
brick walls in central London. Colonisation seems to have occurred within the last

FERN GAZ. 18(1): 31-38. 2007

400

900
Metropolitan Middlesex
(National Grid square TQ)
850 -

jo2)

=

<=

5

C)

=

800 +
750 — ;
200 250 300 350
Easting

Figure 1. Sites for Asplenium adiantum-nigrum in the Metropolitan district of

Middlesex; the River Thames (shown grey) is its southern boundary.
900

400

Metropolitan Middlesex
(National Grid square TQ)
850
ao
£
<£
4
ic]
z
800 -
750 : Ss
200 250 300 350
Easting

Figure 2. Sites for Asplenium trichomanes subsp. quadrivalens in the Metropolitan
dary.

district of Middlesex; the River Thames (shown grey) is its southern boundary.

EDGINGTON: DYNAMICS OF LONG-DISTANCE DISPERSAL 33

fifteen years and the distribution of sites, dense near the first recorded occurrence,
sparser further away, suggests that is not yet complete. A model is proposed for the
spatial distribution of long-distance dispersal events, which when fitted to the data
yields an estimate of the mean dispersal distance and the likelihood of colonisation as
a function of distance.

COLONISATION OF LONDON BY SPECIES OF ASPLENIUM

Since 1998 the author and others have recorded ferns growing on walls in the built-up
area of inner London, specifically the so-called Metropolitan district (Kent, 1975)
whose boundaries are essentially those of the pre-1965 London County Council north
of the River Thames. The area of this district is about 140 km2. A recent paper
(Edgington, 2003), comparing current and historic records, showed there had been a
significant increase in the number and variety of mural ferns since about 1990, probably
associated with a contemporaneous decline in SO2 levels. More than twenty different
species have been recorded. Excluding casual aliens, and a few native ferns represented
by only one or two records, they fall in one of three groups.

Four species (Asplenium scolopendrium L. and three woodland ferns - Dryopteris
filix-mas (L.) Schott, Dryopteris dilatata (Hoffm.) A. Gray and Pteridium aquilinum
(L.) Kuhn) have been recorded regularly since 1869 and are essentially ubiquitous,
found on a great many walls in central London; the former is always highly fertile while
the woodland ferns, though normally sterile when growing on walls, are abundant in
surrounding countryside and are presumably recruited from fertile plants growing in
nearby woodland. Another five (Adiantum capillus-veneris L., Asplenium ceterach L.,

. ruta-muraria L., Polypodium vulgare L. and P. interjectum Shivas) were rarely
recorded in the past and are still known from rather few places. The most abundant are
P. vulgare (sensu lato) and A. ruta-muraria with over a dozen sites each; some have
been known for centuries but others are transient and there is little evidence of recent
rapid spread, simply of better recording. Colonisation is evidently a slow and erratic
process, like the occasional escape of alien ferns from cultivation.

Two species, however, A. adiantum-nigrum L. and A. trichomanes L. subsp.
quadrivalens, which were apparently absent from central London’ until recently, are
now frequent. The historical flora of Middlesex (Kent, 1975) has no modern records in
the district for either fern but the Supplement (Kent, 2000) with records up to 1997 lists
four sites for A. adiantum-nigrum (including the City of London where Mr JM
Montgomery found one plant in 1980 — the others were in 1988, 1990 and 1995) and
two for A. trichomanes (1995 and 1996, Bloomsbury). Both ferns are now widespread
and appear to be still increasing, with 40 and 43 records respectively at the time of
writing, compared with 27 and 37 records respectively in Edgington (2003). They
behave as strict calcicoles, growing only on mortared walls (or in one case, on eroded
limestone boulders used as design features in a small park), generally in shade and
often, though not exclusively, towards the top of the wall. Old crumbling mortar is
preferred but many sites are on apparently sound mortar of modern walls where there
is some permanent dampness. Figures 1 and 2 show their distributions within the
Metropolitan district. Each record includes the street and sometimes the house number
but in the Figures and what follows 6-figure National Grid references are used; sites
closer than 100 m are treated as a single colony.

'There is a solitary record of A. trichomanes at Chelsea, by Robert Nicholls before 1746.

34 FERN GAZ. 18(1): 31-38. 2007

Evidence for a small number of founding events

Both ferns form compact colonies where they occur; sporelings mature rapidly and
_ within a year or two all plants are shedding enormous numbers of sooty spores. Though
present in many districts around London they were, and still are, “thinly scattered ... and
often scarce where they occur” (Burton, 1983), unlike A. ruta-muraria, for example,
which is much more abundant on suitable sites in the surrounding countryside. Their
sudden appearance in the Metropolitan district and their rapid spread suggests in the
case of each species one or at most very few founding events followed by spore
dispersal and establishment in suitable places with the cycle then repeating. Their
distributions (clustered near their earliest recorded sites in the City and in Bloomsbury,
sparser further away) are highly non-random. Dividing the district into equal areas
along north-south and east-west axes, counting the number of sites in each sector and
evaluating 2 assuming random distributions yields significant (P<0.05) and highly
significant (P<<0.01) departures from randomness from east to west and from north to
south respectively, for both species. In contrast the distribution of sites for A. ruta-
muraria (Figure 3) is essentially random (0.1<P<0.9) in both directions.

e number of new records of A. adiantum-nigrum and A. trichomanes each year is
shown in Table 1. Coverage is necessarily incomplete — only sites accessible to the
public have been surveyed — but if recording efficiency is assumed to be constant across
the whole area, the model of dispersal described below is still applicable. The decline
in new records after 2003 is consistent with early colonisation of the most suitable sites
and slower spread subsequently. If this process were to continue these species would no

Metropolitan Middlesex
(National Grid square TQ)

850 +
ao
4
z
=
C)
z

800 -

200 250 300 350 400

Easting
Figure 3. Sites for Asplenium ruta-muraria in the Metropolitan district of Middlesex;
the River Thames (shown grey) is its southern boundary.

EDGINGTON: DYNAMICS OF LONG-DISTANCE DISPERSAL 35

Table 1. Annual summary of new records in the Metropolitan district.

Year A. adiantum-nigrum| A. trichomanes Total
1998 1 1
1999 2 5 7
2000 6 8 14
2001 6 10 16
2002 6 10 16
2003 8 + 12
2004 4 1 $
2005 5 1 6
2006 2 4 6

doubt eventually become as common as A. scolopendrium, a longer-established and
equally freely-fertile fern found throughout inner London on almost all suitable
mortared walls; its present distribution is essentially uniform and tells us nothing about
spore dispersal.

A MODEL FOR THE DISPERSAL PROCESS

The following simple model is proposed. During periods of calm, ejected spores will
fall close to the plant. In windier weather some, maybe most, will be carried away and
are capable of travelling considerable distances. As long as the atmosphere is turbulent
they are likely to remain suspended. One mechanism for bringing them to ground is
capture by (or nucleation leading to formation of) raindrops, a process that will
obviously deposit them in a wet environment and enhance the chances of germination.
The combination of an (assumed constant) wind velocity with a random process of
precipitation (probably, but not necessarily by rain) acting on an (extremely large)
population of spores results in an equal probability, €, of deposition per unit distance
travelled. The number of spores deposited at distance d from their source thus follows
an exponential distribution, N ~ e-éd, as in radioactive decay where time is the variable,
and just as the time of the first decay after a clock has been started follows the same
law, so does the distance from the source at which the first spore is deposited. If each
deposited spore is equally likely to produce a mature sporophyte, the same exponential
law will hold for the distance of these from their parent plant. Dispersal will take place
in the direction of the prevailing wind, which will be variable but is assumed not to
change significantly during each episode of spore release and travel. The predominance
of stations in Figures. 1 and 2 to the north-east of the presumed initial sites is consistent
with the prevalence of south-westerly winds.

A priori we do not know the source of spores arriving at a particular site. To
continue the analogy with radioactivity, where the interval between consecutive decays
also follows the exponential law, so too does the distance between adjacent spore

36 FERN GAZ. 18(1): 31-38. 2007

depositions; that is, by assumption, between sites of mature ferns. It is sufficient,
therefore, to identify each station’s nearest neighbour, determine their separation D and
estimate 1/6 as Dm , the mean value of D. This procedure relies on each nearest-
neighbour pair being in a direct parent-offspring relationship. The sparse distribution of
these two ferns makes this a plausible assumption. (Its breakdown is one reason why
this model cannot be applied to dense distributions like that of A. scolopendrium.) It is
immaterial which of the pair is the parent and which the offspring. If there is more than
one founding colony the exponential law will hold for each lineage separately and the
estimated value of Dm (based on the combined populations) will be a lower bound to
the true value.

Results of nearest-neighbour analysis

The data in Figures 1 and 2 have been analysed using this model. The distributions of
D for the two species turn out to be similar. The mean and standard deviations —
for a true exponential distribution would be equal) are Dm = 0.875 km, 872 km
for A. adiantum-nigrum and Dm = 0.721 km, = + 0.891 km for A. ae This
agreement between Dm and , excellent in the first case and adequate though not so
compelling in the second, gives some support for the simplest model of a single
founding event in each case.

If it is assumed that both data sets have the same true value of &, they can be
combined, yielding the estimate Dmjoint = 0.795 km with a standard error of m = 0.097
km. Since plants separated by less than 100 m are treated as a single colony this
procedure overestimates the mean separation by approximately 50 m; the corrected
value is thus Dmjoint = 0.745 km The combined data are plotted in Figure 4 as a
cumulative distribution of (N > D) versus D. The solid curve represents N = NO (1 — e-
€D) where NO = 83 is the total number of sites (hence, nearest-neighbours) and 1/é =
0.745 km is the mean dispersal distance.

The exponential distribution is strongly leptokurtic (kurtosis k = 9; compare k = 3
for a Gaussian distribution) with a long tail of improbable but not impossible events.
The two largest observed values of D are 3.5 and 4.8 km, for A. adiantum-nigrum and
A. trichomanes respectively. Values as large as this or larger would be expected in about
1 in 100, and 1 in 1500 cases respectively, for the given exponential distribution. A
separation of 3.5 km is therefore entirely consistent with the size of the data set (83
values of D). A separation of 4.8 km is less likely but not implausibly so (P = 0.05).

DISCUSSION
Two assumptions underlying this analysis are, that long-distance dispersal is not an
unusual event but a regular process undergone by large numbers of spores, and that the
breeding system allows growth of sporophytes from a small number of spores, perhaps
only one.

Experimental data on spore production and

In totally still air, even the most energetic process cam scarcely Bones spores more than
a very short distance from a dehiscing sporangium of the deposition rate
around a single fertile fern do indeed show that it falls rapidly with distance, reaching
(in the case of woodland ferns such as Pteridium) background levels beyond 5-10 m
(Dyer, 2005, in litt). However, in experiments involving the mosses Atrichum
undulatum (Hedw.) P. Beauv. and Bryum argenteum Hedw., Miles and Longton (1992)

EDGINGTON: DYNAMICS OF LONG-DISTANCE DISPERSAL a7

estimated that 85-95% of spores were dispersed more than 2 m from the parent
gametophyte, presumably for the same reasons that pollen grains drift long distances in
air currents. Such an experiment has not been carried out for pteridophytes but it does
not seem unreasonable to suppose that similar results would be obtained. The fact that
wind speeds generally increase with height is an argument favouring long-distance
dispersal for spores of ferns growing on vertical walls. Since the number of spores per
frond of A. trichomanes has been estimated to be about 750,000 (Page, 1979), a colony
of a few mature plants, with a hundred or so fronds, could release up to 108 spores each
season. If, as with the mosses in Miles and Longton’s studies, the majority spread
beyond the immediate vicinity, then hundreds of spores could travel 10 km or more,
even with a mean dispersal distance as short as 0.75 km.

Germination and development in Asplenium

Sporophyte production from a single spore requires either self-fertilisation or
apogamous development of the gametophyte. The apogamous life-cycle occurs widely
among epilithic taxa but is apparently present in only about 3% of Asplenium species
(Van den heede, Viane and Chase, 2003); neither A. adiantum-nigrum nor A.
trichomanes is known to be apogamous. In a study of germination and gametophyte
development in three Asplenium species, Pangua, Lindsay and Dyer (1993) showed that
gametophytes of A. trichomanes were protogynous with about equal numbers
developing archegonia only, and both antheridia and archegonia; rather few developed
antheridia only. Fully 90% of archegoniate gametophytes went on to develop
sporelings. These findings were obtained with rather densely sown cultures (ca 250
spores cm -2) so the likelihood of inter-gametophytic fertilisation was a priori high;
indeed the authors concluded that the populations of A. trichomanes they studied were
predominantly outbreeding. More recently, experiments on five Swiss populations of A.
trichomanes subsp. quadrivalens (Suter, Schneller and Vogel, 2000) showed that more

80 - Seeeeo

N>D

Cumulative sum, N>D, of nearest-neighbour distances D for
Asplenium adiantum-nigrum and A. trichomanes ssp.quadrivalens

0 0.5 1 1.5 2 2.5 3 3.5 4 45 5
D/km
Figure 4. Data have been evaluated separately and combined. The solid line is the
exponential fit to the joint data-set.

38 FERN GAZ. 18(1): 31-38. 2007

than half (mean 56.4%, maximum in one population 83.3%) of deliberately isolated
prothalli succeeded in forming sporophytes. Suter, Schneller and Vogel concluded that,
although there was some evidence for cross- fertilisation, the taxon is highly inbreeding
“with the capability of single-spore colonisation and subsequent founding of new
populations” (/oc. cit.). It seems safe to conclude that intra-gametophyte selfing is
common in A. trichomanes and, if the similar behaviour of the taxa in London is any
guide, in A. adiantum-nigrum too.

CONCLUSIONS
The present distributions of A. adiantum-nigrum and A. trichomanes in central London
are well described by an exponential model of dispersion. Both species appear to have
migrated about ten km from their presumed initial sites, the former in 15-20 years, the
latter in 10-15 years. Their mean distance of travel is about 0.75 km while there is
evidence for dispersal distances of up to 5 km during this period. This value is similar
to that inferred for bryophytes in a comparable situation.

In the absence of experimental proof that the various colonies of each fern form a
single lineage these conclusions must be tentative; evidence for single founding events
is mainly circumstantial. A. trichomanes in particular is a variable species. Although
most examples from central London are clearly referable to the tetraploid subsp.
quadrivalens others appear morphologically close to subsp. hastatum or even
diploid subsp. inexpectans. Lacking biological evidence of affinity between colonies,
this account must be treated as a hypothesis requiring validation by genetic studies to
confirm the lineages.

REFERENCES

BURTON, R.M. 1983. Flora of London. London Natural History Society, London.

CAIN, M.L., MILLIGAN, B.G. & STRAND, A.E. 2000. Long-distance seed dispersal
in plant populations. Am. J. Botany 87: 1217-1227

EDGINGTON, J.A. 2004. Ferns of the metropolis — a status report. Lond. Nat. 82: 59-
a3

KENT, D.H. 1975. The historical flora of Middlesex. The Ray Society, London.

KENT, D.H. 2000. Flora of Middlesex: A supplement to The historical flora of
Middlesex. The Ray Society, London.

MILES, C.J. & LONGTON, R.E. 1992. Deposition of moss spores in relation to
distance from parent gametophytes. J. Bryol. 17: 355-368

MILLER, N.G. & McDANIEL, S.F. 2004. Bryophyte dispersal inferred from
colonization of an introduced substratum on Whiteface Mountain, New York. Am.
J. Botany 91: 1173-1182

PAGE, C.N. 1979. Experimental aspects of fern ecology. In DYER, A.F. (Ed.) The
experimental biology of ferns, pp. 552-589. Academic Press, London.

PANGUA, E., LINDSAY, S. & DYER, A. 1994. Spore germination and gametophyte
development in three species of Asplenium. Ann. Bot. 73: 587-593

, M., SCHNELLER, J.J. & VOGEL, J.C. 2000. Investigations into the genetic

variation, population structure and breeding systems of the fern Asplenium
trichomanes subsp. ioe — J. Plant Sci. 161 (2): 233-244

VAN DEN HEEDE, C.J., VIANE, P.L.L. & CHASE, M.W. 2003. Phylogenetic analysis
of Asplenium subgenus peas (Pretadonlentes Aspleniaceae) based on plastid and
nuclear ribosomal ITS DNA sequences. Am J. Botany 90: 481-495

40 FERN GAZ. 18(1). 2007
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THE FERN GAZETTE

VOLUME 18 PART 1 2007

CONTENTS

C.R. Fraser-Jenkins

ginelia agasty gi Pteridophyta), a new species
Raju Antony, A.E. Shanavas Khan & G Sreekandan Nair

_ Dynamics of long-distance dispersal: the spread of Asplenium