Full Contents - the British Pteridological Society

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Fern Gazette V18 P1 v7.qxd
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1
THE SPECIES AND SUBSPECIES IN THE DRYOPTERIS AFFINIS
GROUP
C.R. FRASER-JENKINS
Student Guest House, Thamel, P.O. Box no. 5555, Kathmandu, Nepal
([email protected])
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.
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 parallelsided, ± 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 longerterm 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 formerly been treated as subspecies, they
are now raised to specific rank along with the others.
The other important result of apomixis in D. affinis is that any variants that arise
within members of the group are preserved by the cloning type of reproduction through
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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
names 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
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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.
A relatively 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
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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 and, being readily recognisable from their gross
morphology alone, it is more appropriate to recognise them as species. This also reflects
the biology of the taxa since it is known that most of them have a different individual
biological (cytological and genomic) basis, though the mere fact of the existence of
such a difference, had it been in the absence of relatively major and readily recognisable
morphological separability would in the present author’s opinion be an inappropriate
and 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 FraserJenkins 1980), Dryopteris borreri var. subintegra (Aschers & Graebn.) Tavel (1937)
[based on Aspidium filix-mas var. subintegrum Döll (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, Döpp 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, ??23, 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,
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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, ??111, 112), Hb [VC H1, H2, H3, H4, H5, 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 E), 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.
B. 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.
paleaceolobata (T.Moore) Fras.-Jenk., comb. inval. (1996). Cytotype: Diploid apomict
(Manton 1950, identified more precisely in Fraser-Jenkins & Trewren in prep.). Range:
Br (Scot., 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. Zürich, 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, Eschelmüller & 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. H1, H2, H3,
H4, H5, H6, H7, H16].
E. subsp. jessenii (Fras.-Jenk.) Fras.-Jenk., comb. nov., basionym: Dryopteris
affinis [subsp. affinis] var. jessenii Fras.-Jenk., in Widén, Fraser-Jenkins, Reichstein,
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Gibby & Sarvela, Ann. Bot. Fennici 33: 73 (1996). Holotype: from Romania, Judeþ.
Oltenia (road above Bãile 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. 1986, 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] (1987). Cytotype:
Triploid apomict.
A. subsp. cambrensis*. Synonym: D. affinis [subsp. cambrensis] var.
paleaceocrispa (T.Moore) Fras.-Jenk., nom. inval. (1996). Cytotype: Triploid apomict,
with the 16-smc sporangia showing 41 bivalents and 41 univalents (Gibby in FraserJenkins 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-smc 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.
D. subsp. 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!
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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, FraserJenkins, 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 Dostál, Fraser-Jenkins & Reichstein, Hegi’s Illustr. 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
(1996a); 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 (Döpp 1963, Schneller in Eschelmüller &
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., ser.
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), I. 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
(Harçbeli Pass, Kirtibeº, Gölköy), 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.
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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. Zürich (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. (Hámor,
Köszeg (Güns) [= Göszbachtal, 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 1018 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 voucherspecimens, 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, ??112]), 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
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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
recognisable 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
European-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 Siâdûn, 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 morphologically
distinguishable from it in its rounded, flabellately toothed pinnule-apices etc. and
deserves specific rank. Other subspecies and unnamed variants of D. wallichiana 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.G.Lu
(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
distinguishable by their intermediate frond-morpology combined with mostly abortive
spores and by not usually forming large populations, but occurring as a few isolated
individuals in a population. However the author has seen some considerable
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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.
cambrensis, 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. oreades x D. affinis would presumably be genomically identical and probably
the same as D. cambrensis, while D. oreades x D. borreri would presumably be
genomically identical to D. x complexa and the two origins 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: xxiii (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.
B. nothosubsp. contorta* Fras.-Jenk., in Derrick, Jermy & Paul, Sommerfeltia 6: xxiii (1987) (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,
Scotland (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 FraserJenkins & Trewren in prep.). Range: Br (Wales, Engl. [VC. 47, 96]).
C. nothosubsp. eschelmuelleri Fras.-Jenk., nothosubsp. nov. planta similis ad
nothosubsp. complexam sed dentibus longioris flabellulatis et facie superiora
pinnularum valde depressa super sorum differt. Sporae plerumque abortivae. Holotype,
here designated: Germany [W.], Bayern, south Schwaben, Allgäuer Alpen, Grunten süd,
giffelstein, A. Eschelmüller [klon 11] 85/11 (Herb. T. Reichstein no. 6282, Basel, now
in Z!); isotypes: A. Eschelmüller [klon 10 and 11] 84/23, 13/8/1984, 84/36 and 84/88,
13/9/1984, 84/87, 13/9/1984 and 85/73, 19/8/1985 (Herb. A. Eschelmüller, Sulzbergim-Allgäu, now in M!). D. affinis subsp. punctata x D. filix-mas. Cytotype: Tetraploid
semi-sterile hybrid (Bär & Eschelmüller 1985). Range: Ge (W.), He, Ju (Hrvat.), An.
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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,
Südwest seite des Grünten, von Burgberg aufwarts, A. & H. Eschelmüller 85/40,
24/5/1985 (Herb. A. Eschelmüller, Sulzberg-im-Allgäu, now in M!). Plant then grown
from spores by A. Eschelmüller, 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 (Döpp 1941, Bär & Eschelmüller 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 eglandulosa. Sporae plerumque abortivae. Holotype, here designated:
Britain, Scotland (Inverness-shire, road-cutting near top of pass, Shlochd, north of
Carrbridge, south of Inverness), CRFJ 11187, 18/9/1984, 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 cuneatoacutis 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 Gätzi (1961: 44) and
Döpp, Gätzi & 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. nov., basionym: Dryopteris x
complexa nothosubsp. critica Fras.-Jenk., in Derrick, Jermy & Paul, Sommerfeltia 6: xxii (1987). Holotype: from Germany (former E. Germany), Gera
(Rudolstadt/Thüringen, zw. Partschefeld und Ühlstädt), I. & S. Jessen 1291, 22 June
1985, in Herb. S. Jessen, Chemnitz [formerly Karl-Marx-Stadt]!, isotype: in JE! D.
12
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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 FraserJenkins & 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.
Mature 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 sporesize 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.
1a. 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*.
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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
the 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 ¼ - ½ 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 ¼ adnate;
lamina not, or only slightly glossy .............................................................................. 12.
4a. Pinnules markedly crowded, some with sloping apices ..............................................
........................................................................................... D. 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 very 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
……………………………..........…………….…............... D. affinis subsp. punctata.
6b. Lamina without, or only with shallow and insignificant “punctate” indentations
above …………............................................................................................................. 7.
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 lobed
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
wide………………………………............................................................................... 8.
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*.
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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 downrolling 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 ..................................................................................... 11.
11a. 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 .......................................................................... 13.
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 long, 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 .................................. 15.
15a. Lower pinnules with a wedge-shaped base, pinnules usually not markedly lobed,
laminar upper surface ± glossy; indusia with well curved-down sides and a deep,
darkish centre ................................................................................... D. pseudodisjuncta.
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.
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16b. Pinnule-teeth ± obtuse at the tips of the pinnules ................................................ 23.
................................................................................................. D. affinis subsp. punctata.
17b. 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 pinnules on each pinna lobed with numerous small, neat lobes; stipe and
rachis scales all narrow …………………...........… D. affinis subsp. paleaceolobata*.
19b. The lower pinnules 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
..................................................................................................................................... 21.
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 ........................... 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.
................................................................................................. 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 numerous 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.
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25a. Lower pinnule-apices narrowly rounded-pointed; scales pale- to mid-brown ..........
……………………………………………………….….............. D. 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, frondaxes highly glandular .................................................... D. 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, persistent, 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
pointed .............................................................................................. 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 ……………............. 33.
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.
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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 darkgreen 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 ¼ and ½ 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 being 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 parallelsided 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
darker and narrower; the pinnule-lobes are also smaller, narrower and neater and the
spores are smaller. Appears to be the nearest Atlantic equivalent to subsp. punctata but
is more lobed and without the “punctate” indentations (apart from very small ones)
above the sori. Also similar to subsp. affinis but differs somewhat markedly in its
pinnules all being lobed at the sides and 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 developed and becomes the longest and most lobed in the frond,
and is fully stipitate. However it retains the dense, dark and narrow stipe- and rachisscales and markedly glossy lamina of D. affinis, as well as the eglandular frond and tall
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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 pinnuleapices; they also bear characteristic longer, more prominent and narrower, but obtusetipped 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) stipeand 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
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with more russet-brown scales, slightly approaching subsp. distans. The scales range
from pale, to, more usually, mid- to dark-brown, 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 short, except when growing in woods,
or between boulders. Lamina slightly thinner than in D. affinis, smooth, with the
veinlets hardly impressed above, glossy above when compared with D. borreri, though
not as glossy as in D. affinis, usually darker-green (darker than in subsp. insubrica)
though paler or yellowish when in the open, characteristically narrow, ± tapering
towards the base. Pinnae short compared to D. affinis, tapering throughout from their
wider bases, seldom flat as at least some of the ± rounded or narrowly-rounded pinnuleapices, 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 somewhat darker brown in colour. Fronds turning brown and
dying down in mid Autumn.
2B. subsp. distans. Generally intermediate in morphology between subsp.
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 D.
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
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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, Vshaped 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 basiscopic one on
each pinna may sometimes be slightly basiscopically deflexed so as to lie partly over
the rachis, the lowest basiscopic pinnule of the lowest pinna is usually fully stalked, or
stipitate. The lowest opposite-pair of pinnules on each pinna is usually well lobed, with
characteristic wedge-shaped, pointed side-lobes. Pinnule-apices rounded to rounded
truncate, as in D. affinis, but often becoming square-truncate above, bearing prominent,
long, ± narrow, but wide-based, usually somewhat acute-tipped teeth, which are
flabellate, i.e. 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.
Sori 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
long, markedly flat; pinnules ± regular in length, or the first pair slightly the longest in
each pinna, usually separated from each other above their bases by a distinctive, quite
wide, V-shaped gap or notch, and in mid and upper pinnae somewhat widely attached
to each other at their bases, often narrowed to their bases in the lowest pair of pinnules
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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; pinnuleapices 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, somewhat 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,
either sitting ± flat over the top of the ripe sporangia until dropping off, or shrivelling
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-apices of D.
borreri. Stipe-base scales with dark bases and prominent, separating, dark, glossy
streaks extending upwards into the pale area of the scale. Lamina not markedly glossy.
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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.
Usually 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 sidelobes, 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 loweror mid-frond pinnule-apices usually remain truncate, rounded-truncate, or obovate, as
opposed to their all being more rounded as in D. affinis and in most D. cambrensis
subsp. cambrensis etc.; in normal plants the pinnule-apices bear characteristically long,
mostly acute teeth, similar to those in D. filix-mas, but some plants are occasionally
almost toothless, when they can become difficult to separate from D. cambrensis subsp.
cambrensis; 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 head” outline
- the corner teeth representing the cat’s ears). Fronds eglandular. Sori smaller than in D.
affinis or D. cambrensis, and indusia noticeably relatively thin and less tall, whiteish or
pale- to mid-brown, eglandular, beginning curved down at the sides (but 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, wrinkled cone, or funnel on top of the fully exposed sporangia, and soon drop
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
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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, darker 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 russet-coloured 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 pinnuleapices 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
occasional unseparated diads or tetrads amongst them and usually with some perisporial
(sporangial) debris attached to their surfaces, also with dark debris remaining in the
sporangia (not to be confused 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 sporadically, but fairly
commonly, as occasional plants among populations of the parents, not forming
populations of their own, except some scattered rather small local, probably 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
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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 Eschelmüller, Sulzberg; the late Mr. Hugh Corley, Faringdon; Prof. Gabor
Vida, Budapest; the late Dr. Walter Gätzi, St. Gallen; M. Michel Boudrie, Clermont
Ferrand; Mr. Mike Taylor, Isle of Skye; Mr. Martin Rickard, Kyre Park; Dr. Jacob
Schneller, Zürich; 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
BÄR, A. & ESCHELMÜLLER, A. 1985. Tetraploide und pentaploide Dryopteris x
tavelii - - jetzt im Allgäu bestätigt. Mitt. Naturwiss. Arbetskr. Kempten (Allgäu) 27
(1): 57-68.
BÄR, A. & ESCHELMÜLLER, A. 1989. Beitrag zur Kenntnis von Dryopteris affinis
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25
(Lowe) Fraser-Jenkins und von Bastarden mit Dryopteris filix-mas. Mitt. Naturwiss.
Arbetskr. Kempten (Allgäu) 29 (1): 25-48.
BÄR, A. & ESCHELMÜLLER, A. 1990. Dryopteris x complexa nssp. contorta FraserJenkins - 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.
DÖPP, W. 1941. Über Dryopteris paleacea Christensen (D. borreri Newm.). Ber.
Deutsch. Bot. Ges. 59 (9): 423-426.
DÖPP, W. 1955. Experimentell erzeugte Bastarde zwischen Dryopteris filix-mas (L.)
Schott und D. paleacea (Sw.) C.Chr. Planta, Arch. Wissensch. Bot. 46: 70-91.
DÖPP, W., GÄTZI, W. & OBERHOLZER, E. 1963. Dryopteris borreri Newman (D.
paleacea Hand.-Mazz.) var. pseudodisjuncta v. Tavel. Ber. Deutsch. Bot. Ges. 76
(4): 99-111.
ESCHELMÜLLER, A. & SCHNELLER, J.J. 1980. Beitrag zur Kenntnis der
Variebilität von Dryopteris affinis im Allgäu. Mitt. Naturwiss. Arbetskr. Kempten
(Allgäu) 24 (1): 1-12.
FRASER-JENKINS, C.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 DOSTÁL, J., FRASERJENKINS, 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 Nordica 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.
GÄTZI, W. 1961. Über den heutigen Stand der Dryopterisforschung unter besonderer
Berücksichtigung von Dryopteris Borreri Newman, zugleich ein Beitrag zur
Farnflora des Südabhanges des Tannenbergs. Ber. üb. Tätigkeit St. Gallisch. Naturw.
Ges. 77: 1-73.
ICBN: GREUTER, W. et al. 2000. International Code of Botanical Nomenclature (Saint
Louis Code). Regnum Vegetabile 138. Königstein.
JERMY, A.C. & CAMUS, J.M. 1991. The illustrated Field Guide to Ferns and Allied
Plants of the British Isles. London.
JERMY, A.C., PIGOTT, A.C. & MERRYWEATHER, 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.
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FERN GAZ. 18(1): 1-26. 2007
JESSEN, S. 1991. Neue Angaben zur Pteridophytenflora Osteuropas. Farnblätter 23:
14-47.
JOHNS, R.J., EDWARDS, P.J. & PICHI SERMOLLI, R.E.G. 1996. Index Filicum
Supplementum sextum. Kew.
MANTON, I. 1950. Problems of Cytology and Evolution in the Pteridophyta.
Cambridge.
MERRYWEATHER, J.W. 2002. Identification – British male ferns. Pteridologist 4 (1):
12-14.
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(2).
RASBACH, H., RASBACH, K., REICHSTEIN, T. & SCHNELLER, J. 1983.
Tetraploide Dryopteris x tavelii Rothm. im nördlichen Schwarzwald. Farnblätter 10:
1-13.
REICHSTEIN, T. & SCHNELLER, J. 1983. Dryopteris affinis var. punctata im
Hüllerich-Wald ob Pfaffikon (SZ). Farnblätter 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 Nördosthessen. Beiträge 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. Ökologische Untersuchungen. Ber. Schweiz.
Bot. Ges. 85 (2): 110-159.
WIDÉN, 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.
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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.
Strobilus 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 µm, spinulosae. Microsporae fortiter
ourantiaceae, 23-33 µm, verrucoidae.
Holotypus: Agastyamala, Thiruvananthapuram District, Kerala, South India, 1,750
m. alt.; in evergreen forest; 9 January 1996; Raju Antony 25580 (TBGT).
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,
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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, ovateoblong, 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 µm in diameter, spinulose. Microspores deep orange,
23-33 µm in diameter, verrucoid.
Other specimens examined: Agastyamala, Thiruvananthapuram District, Kerala,
10 December 1998; Raju Antony 29880 (TBGT).
Note: This species is allied to Selaginella miniatospora (Dalz.) Bak. But can be easily
distinguished 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.
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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
Dun.
MADHUSOODANAN, P.V. & SANTHOSH NAMPY, 1994. Selaginella dixitii, a new
species of Selaginellaceae from India. Nordic J. Bot. 14: 527-529.
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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 (Preston 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
urbanisation, 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 resurvey 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).
The 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.
What 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
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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 WC1N 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 1 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 longrange 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
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FERN GAZ. 18(1): 31-38. 2007
900
Metropolitan Middlesex
(National Grid square TQ)
Northing
850
800
750
200
250
300
350
400
Easting
Figure 1. Sites for Asplenium adiantum-nigrum in the Metropolitan district of
Middlesex; the River Thames (shown grey) is its southern boundary.
900
Northing
850
800
750
200
250
300
350
400
Easting
Figure 2. Sites for Asplenium trichomanes subsp. quadrivalens in the Metropolitan
district of Middlesex; the River Thames (shown grey) is its southern boundary.
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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.,
A. 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 London1 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.
1
There is a solitary record of A. trichomanes at Chelsea, by Robert Nicholls before 1746.
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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. rutamuraria (Figure 3) is essentially random (0.1<P<0.9) in both directions.
The 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
900
Northing
850
800
750
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.
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Table 1. Annual summary of new records in the Metropolitan district.
Year
A. adiantum-nigrum
A. trichomanes
Total
1998
1
1999
2
5
7
2000
6
8
14
2001
6
10
16
2002
6
10
16
2003
8
4
12
2004
4
1
5
2005
5
1
6
2006
2
4
6
1
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, ë, of deposition per unit distance
travelled. The number of spores deposited at distance d from their source thus follows
an exponential distribution, N ˜ e-ëd, 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
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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/ë as Dm , the mean value of D. This procedure relies on each nearestneighbour 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 (which
for a true exponential distribution would be equal) are Dm = 0.875 km, = ± 0.872 km
for A. adiantum-nigrum and Dm = 0.721 km, = ± 0.891 km for A. trichomanes. 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 ë, 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 – eëD) where NO = 83 is the total number of sites (hence, nearest-neighbours) and 1/ë =
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 dispersal
In totally still air, even the most energetic process can scarcely propel spores more than
a very short distance from a dehiscing sporangium. Measurements 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)
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37
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
70
60
N>D
50
40
30
20
Cumulative sum, N>D, of nearest-neighbour distances D for
Asplenium adiantum-nigrum and A. trichomanes ssp.quadrivalens
10
0
0
0.5
1
1.5
2
2.5
D/km
3
3.5
4
4.5
5
Figure 4. Data have been evaluated separately and combined. The solid line is the
exponential fit to the joint data-set.
38
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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” (loc. 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 the
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: 5973
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
SUTER, M., SCHNELLER, J.J. & VOGEL, J.C. 2000. Investigations into the genetic
variation, population structure and breeding systems of the fern Asplenium
trichomanes subsp. quadrivalens. Int. 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 Ceterach (Pteridophyta: Aspleniaceae) based on plastid and
nuclear ribosomal ITS DNA sequences. Am J. Botany 90: 481-495
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TRICHOMANES SPECIOSUM (HYMENOPHYLLACEAE:
PTERIDOPHYTA) IN SOUTHERN SPAIN
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REFERENCES: should follow the style of a recent issue of The Fern Gazette, e.g.:HOOKER, W.J. 1864. Species Filicum, 5. Dulau & Co., London.
MORTON, C.V. 1947. The American species of Hymenophyllum, section
Sphaeroconium. Contr. U.S. Natl. Herb. 29(3): 139-201.
STEVENSON, D.W. & LOCONTE, H. 1996. Ordinal and familial relationships of
pteridophyte genera. In: CAMUS, J.M., GIBBY, M. & JOHNS, R.J. (Eds)
Pteridology in perspective, pp. 435-467. Royal Botanic Gardens, Kew.
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FERN GAZ. 18(2):41-52. 2007
41
TWO NEW SPECIES OF SELAGINELLA SUBGENUS
HETEROSTACHYS (SELAGINELLACEAE) FROM THE GUIANAS
G. CREMERS1 & M. BOUDRIE2
1
I.R.D., Muséum national d’Histoire naturelle, Département Systématique et Évolution,
USM 0602, case postale 39, 37 rue Cuvier, F-75231 Paris Cedex 05, France.
(Email : [email protected])
2
16, rue des Arènes, 87000 Limoges, France. (Email : [email protected])
Key words: Selaginella, Heterostachys, Guyana, Suriname, French Guiana, South
America.
ABSTRACT
During our study of the biodiversity of the Guianas and the preparation of the
flora of this region, we discovered that the I. Valdespino’s monograph of the
genus Selaginella subgen. Heterostachys was never published. Two taxa, S.
gynostachya and S. karowtipuensis, presented in this monograph and new to the
Guianas, are published herein.
Mots clés: Selaginella, Heterostachys, Guyana, Surinam, Guyane, Amérique du Sud.
RÉSUMÉ
Dans le cadre de notre étude de la biodiversité des Guyanes et de la rédaction de
la flore de cette région, nous avons constaté que la monographie de I. Valdespino
sur le genre Selaginella subgen. Heterostachys n’a jamais été publiée. Deux
taxons, S. gynostachya et S. karowtipuensis, présentés dans cette monographie et
nouveaux pour les Guyanes, sont publiés ici.
INTRODUCTION
In 1995 Ivan A. Valdespino Quintero completed a monographic study to obtain his
Doctor of Philosophy degree in Botany from the Graduate School of The City
University of New York. This work was supervised by J.T. Mickel as Chair of the
Examining Committee. Certain elements of the dissertation, such as the
lectotypifications of S. flagellata Spring and of S. minima Spring, are cited by Mickel
& Smith (2004), but unfortunately Valdespino’s monograph was never entirely
published.
During our study of the biodiversity of the flora of French Guiana and in relation to
the international programme of the ‘ Flora of the Guianas’, we had to consult this work.
For instance, S. minima, of which the type is from French Guiana, is treated in this
work. This species was known only from a few specimens collected before 1850 but has
been recently found again in French Guiana (Cremers & Boudrie, 2006). We are also
preparing a paper dealing with ‘Pteridophytes having at least a syntype collected in the
Guianas’ and fascicle 9 of the Pteridophytes of the Flora of the Guianas, both works
which enhance the interest of such a monograph in which two new species have not
been officially published.
We present here the descriptions of these two new species to the Guianas, with type
specimens from Guyana. What follows has been extracted verbatim from Valdespino’s
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FERN GAZ. 18(2): 41-52. 2007
monograph. Valdespino, who is currently unavailable, deserves full credit for
recognizing and distinguishing these two new species. Here we intend to validate and
to pay tribute to his work.
NEW SPECIES
Selaginella gynostachya Valdespino ex Cremers & Boudrie, sp. nov. (Figure 1 a, b, c;
Figure 2 a, b, c)
A S. hyalogramma Valdespino et S. karowtipuensis Valdespino foliis intermediis basi
subtruncatis vel rotundatis nec non foliis lateralibus supra idioblastis destitutis diversa;
ulterius a prima foliis intermediis ellipticis, ovato-ellipticis vel ovatis, foliis lateralibus
oblongis vel oblongo-ovatis necnon megasporis pallide luteis; ulterius a secunda foliis
intermediis apice acutis vel breviter acuminatis.
Type: GUYANA. Upper Potaro River, vicinity of Kopinang village, 4°58’N, 59°50’W,
670 m, 29 Jul. 1989, Boom & Samuels 8939 (holotype NY; isotype BRG).
Etymology –Gr., gyne, female or pertaining to female organs, and stachys, spike;
referring to the mostly megasporangiate strobili found on specimens of this species.
Plants terrestrial. Stems creeping, stramineous, 5-14 cm long, 0.4-0.8 mm diam., not
articulate, not flagelliform, not stoloniferous, 1- or 2-branched. Rhizophores axillary,
restricted to basal ½ of the stem, stout, (0.28-) 0.34-0.56 mm diam. Leaves dimorphic
throughout, membranaceous, the upper surface green, composed of elongate cells with
sinuate walls. Lateral leaves distant, slightly ascending, oblong to oblong-ovate, 2.8-4
x 1-1.9 mm; base subtruncate to rounded, the acroscopic base strongly overlapping the
stem, the basiscopic base not overlapping the stem; margins slightly hyaline, or pale
green, the acroscopic margin serrate to ciliate-denticulate along basal 1/3, serrulate
apically, the basiscopic margin entire to serrulate apically; apex obtuse to broadly acute,
dentate at tip; both surfaces glabrous, the upper surface without idioblasts, the lower
surface frequently with conspicuous or obscure idioblasts, the stomata present only on
the lower surface. Median leaves distant, ascending, elliptic to ovate-elliptic, 1.7-2.3 x
0.7-1.4 mm; base asymmetric with the inner base truncate and the outer base with a
slightly developed auricle; margins slightly hyaline, serrate to serrulate; apex acute to
shortly acuminate, the acumen, when present, less than 1/3 the length of the lamina,
0.1-0.26 mm long, puberulent and dentate at tip; both surfaces glabrous, the upper
surface frequently with conspicuous idioblats and stomata, the lower surface with
obscure to conspicuous idioblasts, and without stomata. Axillary leaves similar to
lateral leaves or more broadly ovate, except the base frequently rounded. Strobili
terminal on branch tips and main stem apex, lax, flattened, and dorsiventral, 5-15 mm
long. Sporophylls dimorphic, basally attached to the axis; dorsal sporophylls green,
spreading, basally attached to the axis, with an adaxial flap extending along ca. ½ to ¾
the sporophyll length, asymmetric or lanceolate to lance-ovate, 1.6-2.36 x 0.6-0.98 mm,
with a strongly developed and slightly denticulate keel along midrib, the base +
rounded, the margins slightly hyaline to hyaline greenish, serrate to shortly ciliate, the
apex acute to shortly acuminate, the upper surface green, composed of + rounded cells,
except for the portion that folds to form the flap where the cells hyaline and elongate
with sinuate walls, glabrous, frequently with conspicuous idioblasts and stomata, the
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CREMERS & BOUDRIE: TWO NEW SPECIES OF SELAGINELLA
43
Figure 1a. Scanned photograph of Selaginella gynostachya Valdespino ex Cremers &
Boudrie (Cremers 5379, CAY): Herbarium sheet.
44
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Figure 1b. Scanned photograph of Selaginella gynostachya.
Detail of the adaxial face.
2 cm
Figure 1c. Scanned photograph of Selaginella gynostachya.
Detail of the abaxial face.
FERN GAZ. 18(2): 41-52. 2007
2 cm
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45
a
1 mm
c
b
Figure 2. Drawings of the leaves of Selaginella gynostachya Valdespino ex Cremers &
Boudrie (Cremers 5379, CAY).
a = axillary leaf (abaxial face); b = lateral leaf (abaxial face); c = median leaf (adaxial
face);
The small lines on the leaves represent the idioblasts (M. Boudrie’s drawings).
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FERN GAZ. 18(2): 41-52. 2007
lower surface silvery to silvery green, composed of elongate cells with sinuate walls;
ventral sporophylls almost colourless to pale green, ascending, with a slightly
developed adaxial flap, ovate, 1.48-1.9 x 0.56-0.84 mm, with a well developed and
slightly denticulate to entire keel along midrib, the base attenuate to rounded, the
margins hyaline to pale green, serrate to very shortly ciliate, the apex acute to shortly
acuminate, both surfaces made of elongate and hyaline cells with sinuate walls,
glabrous, with or without conspicuous idioblasts, the stomata present only along midrib
on the lower surface. Megasporangia in two ventral rows; megaspores marguerite
yellow (light yellow), with a well developed equatorial flange, the proximal face
striate-reticulate to reticulate, the distal face reticulate with open reticulum of high
ridges, 300-320 µm diam. Microsporangia mostly absent from the two dorsal rows, or
few mature near the apex on dorsal rows; microspores not studied.
HABITAT AND DISTRIBUTION: On stream or river banks; 30-1000 m; known from
Venezuela, Guyana, and French Guiana.
PARATYPES: VENEZUELA. Bolivar: Gran Sabana, ca. 10 km SW of Karauin Tepui at
junction of Rio Karaurin Tepui and Rio Asadon (Rio Sanpa), 5°19’N, 61°03’W, 9001000 m, Liesner 23798 (UC).
GUYANA. Upper Mazaruni: East bank of Waruma River, 20 km S of confluence with
Kako River, 5°19’N, 60°46’W, 1000 m, Feb 1985, Renz 14188 (U); Essequibo River of
Porto Bartica, Jenman s.n. (K); Macouria Riv., Nov 1886, Jenman 2324 (BM, K, US);
Macouria Creek, Essequibo River, Jenman s.n. (NY); Upper Potaro River: vicinity of
Kopinang village, 4°58’N, 59°50’W, 670 m, 29 Jul 1989, Boom & Samuels 8939 (BRG,
NY).
FRENCH GUIANA. Rivière Sinnamary à Petit Saut, 30m, 12 Feb 1979, Cremers 5379
(CAY!, Z).
TAXONOMIC DISCUSSION: Selaginella gynostachya seems most closely related to S.
hyalogramma and S. karowtipuensis but differs by median leaves with a subtruncate to
rounded base (vs. rounded to semicordate as in S. hyalogramma or subcordate to
rounded as in S. karowtipuensis) and by lacking (vs. having) idioblasts on the upper
surface of the lateral leaves. Selaginella gynostachya differs further from S.
hyalogramma by elliptic to ovate-elliptic or broadly ovate (vs. lanceolate) median
leaves, by oblong to oblong-ovate (vs. broadly ovate to broadly ovate-lanceolate) lateral
leaves, and by light yellow (vs lemon yellow) megaspores. S. gynostachya differs from
S. karowtipuensis also by its acute to shortly acuminate (vs. long-acuminate to short
aristate) apex.
On the paratype (Cremers 5379, CAY) illustrated here, we noticed that the median
leaves do not have idioblasts on their upper surface. Therefore, this character is variable
and should be used with caution in the key. In addition, the lateral leaves are evidently
cilio-denticulate.
Selaginella karowtipuensis Valdespino ex Cremers & Boudrie, sp. nov. (Figure 3 a, b,
c; Fig. 4 a, b, c).
Selaginella seemannii Baker affinis, sed foliis lateralibus, intermediis et sporophyllis
superne idioblastis provisis diversa. A S. stenophylla A. Braun foliis intermediis basi
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Figure 3a. Scanned photograph of Selaginella karowtipuensis Valdespino ex Cremers
& Boudrie (Boom & Gopaul 7415, NY): Herbarium sheet.
48
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Figure 3b. Scanned photograph of Selaginella karowtipuensis.
Detail of the adaxial face.
2 cm
Figure 3c. Scanned photograph of Selaginella karowtipuensis.
Detail of the abaxial face.
FERN GAZ. 18(2): 41-52. 2007
2 cm
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a
1 mm
c
b
Figure 4. Drawings of the leaves of Selaginella karowtipuensis Valdespino ex
Cremers & Boudrie (Boom & Gopaul 7415, NY).
a = axillary leaf (abaxial face); b = lateral leaf (abaxial face); c = median leaf (adaxial face);
the small lines on the leaves represent the idioblasts (M. Boudrie's drawings).
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FERN GAZ. 18(2): 41-52. 2007
non nisi auriculo exteriori vel serrulato provisis diversa.
Type: GUYANA. Upper Mazaruni: Karowtipu Mountain, between camp and peak on
western side of mountain, 5°45’N, 60°35’W, 920-1080 m, 24 Apr 1987, Boom &
Gopaul 7698 (holotype NY; isotypes BM, BRG, PMA, UC, US).
Etymology—Named after the type locality.
Plants terrestrial. Stems creeping, stramineous, 20-34 (or more) cm long, 0.9-1.6 mm
diam., not articulate, not flagelliform, not stoloniferous, 2- or 3(-4)- branched.
Rhizophores axillary or occasionally with both axillary and dorsal rhizophores,
restricted to basal ½ of the stem, stout, (0.44)0.54-0.88 mm diam. Leaves dimorphic
throughout, membranaceous, the upper surface green, composed of + rounded cells
with slightly sinuate walls. Lateral leaves distant, perpendicular to the stem,
oblong-ovate to oblong, 3-5.3 x 1.5-2.6 mm; base subcordate to rounded, the acroscopic
base strongly overlapping the stem, the basiscopic base not overlapping the stem;
margins hyaline to pale green, the acroscopic margin serrate, the basiscopic margin
entire to serrulate apically; apex rounded to obtuse, dentate at tip; both surfaces
glabrous, the upper surface with conspicuous idioblasts and without stomata, the lower
surface with obscure idioblasts and with stomata along midrib. Median leaves distant,
ascending, ovate to broadly ovate, 2-3.1 x 1-1.7 mm; base subcordate, the inner base
rounded and the outer base with a well developed auricle; margins hyaline, serrate; apex
long-acuminate to short-aristate, the acumen or arista ¼ or less of the length of the
lamina, 0.28-0.54 mm long, puberulent and dentate at tip; both surfaces glabrous, the
upper surface with conspicuous idioblasts, without stomata. Axillary leaves similar to
lateral leaves or more cordiform. Strobili terminal on branch tips and main stem apex,
slightly lax and flattened, dorsiventral, 3-13 mm long. Sporophylls dimorphic, basally
attached to the axis; dorsal sporophylls green, spreading, with an adaxial laminal flap
extending along ca. ½ to 1/3 the sporophyll length, asymmetric or lanceolate to
lance-ovate, 1.54-2.08 x 0.6-0.92 mm, with a strongly developed and denticulate keel
along midrib, the base rounded, the margins slightly hyaline to hyaline-greenish,
serrate, the apex acute to shortly acuminate, the upper surface green, composed of +
rounded cells, except for the portion that folds to form the flap where the cells hyaline
and elongate with sinuate walls, glabrous, with conspicuous idioblasts, the stomata
along midrib, the lower surface silvery to silvery green, composed of elongate cell with
sinuate walls, and without idioblasts; ventral sporophylls pale green, ascending to
spreading, without an adaxial flap, ovate, 1.32-1.76 x 0.6-0.9 mm, with a well
developed and slightly denticulate keel along midrib, the base rounded, the margins
hyaline to pale green, serrate, the apex acute to shortly acuminate, the surface made of
elongate cells with sinuate walls, glabrous, with idioblasts present only on the lower
surface, the stomata present only along midrib of the lower surface. Megasporangia in
two ventral rows; megaspores maize yellow, with a prominent equatorial flange, the
proximal face striate to reticulate with open reticulum of low ridges, the distal face
reticulate with open reticulum of high ridges, 320-340 µm diam. Microsporangia
mostly absent from the two dorsal rows, or few mature near the apex on dorsal rows;
microspores not studied.
HABITAT
AND
DISTRIBUTION: On sandy or rocky soil; 920-1080 m; known only from
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Karowtipu Mountain in Guyana.
PARATYPES: GUYANA. Upper Mazaruni: Karowtipu Mountain, 5°45’N, 60°35’W, 920
m, 18 Apr 1987, Boom & Gopaul 7415 (BRG, MO, NY!, PMA, U, UC, VEN),; 9201080 m, 24 Apr 1987, Boom & Gopaul 7698 (BM, BRG, NY, PMA, UC, US).
TAXONOMIC DISCUSSION: Selaginella karowtipuensis resembles S. seemannii in its
median leaves with a well developed outer auricle and acuminate apex, and typically
oblong to oblong-ovate lateral leaves with subcordate to rounded base. It differs from
S. seemannii by conspicuous (vs. without) idioblasts on the upper surface of the lateral
and median leaves and sporophylls. In addition, S. karowtipuensis is a more robust plant
with stouter rhizophores (0.54-0.88 mm diam., compared with 0.25-0.65 mm diam. for
S. seemannii).
DISCUSSION – CONCLUSION
Of the 31 species mentioned by Valdespino (1995) within this subgenus, eight occur in
the Guianas. A key, extracted from his monograph, is given below.
Key to the species of Selaginella P. Beauv. subg. Heterostachys Baker in the Guianas.
1. Acroscopic margin of the lateral leaves ciliate to ciliate-denticulate at least along
basal ¼ ........................................................................................................................... 2
1’. Acroscopic margin of the lateral leaves serrate to serrulate or entire near the base or
throughout ...................................................................................................................... 6
2. Upper surface of the lateral leaves with idioblasts ...................................S. muscosa
2’. Upper surface of the lateral leaves without idioblasts ............................................ 3
3. Lateral leaves semicordiform to cordiform; median leaves cordiform to
ovate-deltate ............................................................................................... S. porelloides
3’. Lateral leaves ovate, ovate-orbicular, ovate-lanceolate, lanceolate, ovate oblong, or
ovate-elliptic; median leaves ovate, ovate-lanceolate, lanceolate, elliptic,
elliptic-orbicular, or, if ovate-deltate, then the apex broadly acute and ciliate at tip ... 4
4. Median leaves elliptic or ovate-elliptic ................................................................... 5
4’. Median leaves ovate, ovate-lanceolate, or lanceolate ............................... S. minima
5. Lateral leaves with an acuminate apex; upper surface of the median leaves without
idioblasts; megaspores lemon yellow ......................................................... S. sobolifera
5’. Lateral leaves with a broadly acute to obtuse apex; upper surface of the median
leaves with conspicuous to obscure idioblasts; megaspores light to maize yellow,
cream, pale yellow-orange, or pale orange-yellow ................................. S. gynostachya
6. Upper surface of lateral and median leaves with idioblasts, or idioblasts restricted to
the upper surface of the lateral leaves only, or idioblasts present on the upper surface of
the median leaves and/or on the lower surface of the lateral leaves ............................ 7
6’. Upper surface of the lateral and median leaves without idioblasts, or idioblasts
present only on the upper surface of the median leaves ............................................... 9
7. Median leaves with at least the outer base well developed into an auricle free from
the stem; rhizophores 0.3-0.86 mm diam ............................................ S. karowtipuensis
7’. Median leaves with the outer base asymmetric, rounded, oblique, or subcordate, if
with an outer auricle then this not distinctly free from the stem; rhizophores 0.08-0.56
mm diam. ....................................................................................................................... 8
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8. Apex of the lateral leaves obtuse to broadly acute; apex of the median leaves acute
to shortly acuminate, the acumen 1/8 or less the length of the lamina ... S. gynostachya
8’. Apex of the lateral leaves acute to shortly acuminate; apex of the median leaves
long acuminate to long-aristate, the acumen or arista ¼ or usually more the length of
the lamina ....................................................................................................... S. muscosa
9. Median leaves with the outer base well developed into an auricle ..... S. seemannii
9’. Median leaves with the outer base asymmetric, oblique or semicordate, without a
well developed auricle ................................................................................................. 10
10. Median leaves with the apex acute, acuminate, or short-aristate, the acumen or
arista 1/3 or less the length of the lamina ............................................... S. gynostachya
10’. Median leaves with the apex long-aristate, the arista more than 1/3 the length of the
lamina ........................................................................................................... S. flagellata
DISTRIBUTION
When comparing the distribution of the 35 species of subgenus Heterostachys from
Central and South America, it is noticed that Venezuela has the highest diversity with
17 species. The Guianas have fewer: four for Guyana, two for Suriname and five for
French Guiana. This is unusual because French Guiana typically has fewer species than
Guyana. In terms of distribution, a few species occur over widespread areas (such as S.
flagellata), whereas others are limited to the Plateau of the Guianas (S. gynostachya),
or simply known only from Guyana (S. karowtipuensis).
ACKNOWLEDGEMENTS
We would like to express our gratitude and thanks to Drs. T. Lobova, J.T. Mickel, R.C.
Moran and S.A. Mori (NYBG) who have kindly provided access to the specimens and
a scan, to the technical staff of the National Museum of Natural History in Paris (P) who
carried out one scan, and to Dr. J.J. de Granville (CAY Herbarium, French Guiana) who
has made the detailed photographs of the specimens.
REFERENCES
CREMERS, G. & BOUDRIE, M. 2006. Ptéridophytes de Guyane française non
récoltées depuis plus d’un siècle ou récemment retrouvées. Acta Bot. Gallica 153
(1): 3-48.
MICKEL, J.T. & SMITH, A.R. 2004. The Pteridophytes of Mexico. Mem. New York
Bot. Gard. 88: 1-1054, t.1-328.
VALDESPINO QUINTERO, I.A. 1995. A monographic revision of Selaginella P.
Beauv. subgenus Heterostachys Baker in Central and South America. Degree of
Doctor of Philosophy, The City University of New York 1- 405.
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53
SYSTEMATICS OF TRICHOMANES (HYMENOPHYLLACEAE:
PTERIDOPHYTA), PROGRESS AND FUTURE INTERESTS
A. EBIHARA1, J.-Y. DUBUISSON2, K. IWATSUKI3 & M. ITO1
1
Department of System Sciences, Graduate School of Arts and Sciences, the
University of Tokyo, 3-8-1 Komaba, Tokyo 153-8902, Japan,
2
Université Pierre et Marie Curie, 12 rue Cuvier, F-75005 Paris, France,
3
The Museum of Nature and Human Activities, Hyogo, Yayoigaoka 6-chome, Sanda
669-1546, Japan
(1Email: [email protected]; present address: Department of Botany, 4-1-1
Amakubo, Tsukuba 305-0005, Japan)
Key words: filmy ferns, rbcL, Trichomanes
ABSTRACT
Trichomanes L. sensu lato (s.l.), is a large group of Hymenophyllaceae to which
ca. 250 species are attributed, distributed from the tropics to temperate regions
around the world,. Their life forms and morphology are more diversified than
those of the other large filmy-fern genus Hymenophyllum. Phylogenetic analyses
were performed based on the rbcL sequences of 81 Trichomanes taxa, covering
most of the major groups within the genus, in addition to morphological,
anatomical and cytological investigations, that offer a number of insights
concerning evolution of the genus. Eight robustly supported clades are
recognized within Trichomanes, while some traditional trichomanoid taxa (e.g.,
Pleuromanes) are transferred to the Hymenophyllum clade.
INTRODUCTION
Because of their simplified morphology among pteridophytes, filmy ferns
(Hymenophyllaceae) have attracted the attention of many researchers, especially those
interested in evolution and phylogeny. The family’s basal placement among
leptosporangiate ferns was already suggested by morphological evidence (oblique
annuli of sporangia; Bower, 1926) and supported by recent molecular phylogenetic
work (Pryer et al., 2004). In contrast, it is difficult to reconstruct its intrafamiliar
lineages from its morphological characters, probably as a result of numerous parallel
evolutions (Dubuisson, 1997a). Several different classification systems (e.g., Copeland,
1938; Morton, 1968; Iwatsuki, 1984) are currently in use for this family.
Dubuisson (1997b) first adopted molecular phylogeny to infer intrafamiliar
relationships among the Hymenophyllaceae, specifically targeting one of the two
largest groups, Trichomanes sensu lato (s.l.), to explore the reliability of the chloroplast
rbcL marker at the infrageneric level. The result showed better resolution than for the
other group, Hymenophyllum s.l., in which less genetic variation was found for rbcL
(Pryer et al., 2001; Ebihara et al., 2002; Hennequin et al., 2003). Trichomanes,
however, is a large genus comprising around 250 species (Iwatsuki, 1990) occurring
nearly throughout the tropics and extending into the temperate zone, especially the
southern latitudes. Despite the study by Dubuisson et al. (2003a) focusing on
Neotropical Trichomanes, many distinctive Palaeotropical taxa remained unsampled.
Recently, Ebihara et al. have added considerable new rbcL data, making a global
revision of the genus possible.
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COVERAGE OF SAMPLING
In total, 81 species of Trichomanes, approximately a third of the estimated number of
extant species, were included in this study, though some traditional trichomanoid taxa
(Microtrichomanes Copel. pro parte and Cardiomanes reniforme (G.Forst.) C.Presl)
whose affiliations to the Hymenophyllum lineage have already been suggested (Pryer et
al., 2001; Ebihara et al., 2004) were not counted as Trichomanes. This sampling
covered all of Copeland’s trichomanoid genera (Abrodictyum C.Presl, Callistopteris
Copel., Cephalomanes C.Presl, Crepidomanes (C.Presl) C.Presl, Crepidopteris Copel.
[= Reediella Pic.Serm.], Davalliopsis Bosch, Didymoglossum Desv., Feea Bory,
Gonocormus Bosch, Lecanium C.Presl, Macroglena (C.Presl) Copel., Microgonium
C.Presl, Nesopteris Copel., Pleuromanes (C.Presl) C.Presl, Polyphlebium Copel.,
Selenodesmium (Prantl) Copel., Trichomanes sensu stricto (s.s.) and Vandenboschia
Copel.) and most of Morton’s (1968) sections under Trichomanes, except for four small
sections of the subgenus Achomanes C.Presl (sections Odontomanes (C.Presl) C.Chr.,
Trigonophyllum (Prantl) C.Chr., Homoeotes (C.Presl) C.Chr. and Ragatelus (C.Presl)
C.Chr.).
A data matrix consisting of 1206 bp fragments of the rbcL sequences from 81
Trichomanes species as well as 12 other Hymenophyllaceae and 4 nonHymenophyllaceae (Polypodium glycyrrhiza D.C.Eaton, Matonia pectinata R.Br.,
Osmunda cinnnamomea L. and Angiopteris evecta Hoffm.; the last taxon was treated as
an outgroup) was analyzed using the maximum-parsimony (MP) method with
PAUP*4.0 software (detailed in Ebihara et al.).
EVOLUTIONARY RELATIONSHIP
In the resulting consensus MP tree (Ebihara et al., in prep.; Figure 1), both the
Trichomanes and Hymenophyllum clades are strongly supported, as in Pryer et al.
(2001) and Ebihara et al. (2004). Pleuromanes, a group which has always been
considered a member of Trichomanes, is indeed embedded in the Hymenophyllum clade
and eight robustly-supported subclades (BS > 90) are recognizable in the Trichomanes
lineage. These groupings do not match any of the existing classifications, but seem to
be closely related to the plants’ morphological characters, chromosome base numbers
and geographical distributions.
The phylogenetic framework suggests three evolutionary scenarios for the genus:
(1) Cytological data reveals that the chromosome base numbers are constant within
each “clade,” so the observed diversity in number may originate from aneuploid
reduction from the ancestral x = 36. (2) Life form data indicate that four monophyletic
clades (Cr, Di, Pa and Va in Fig. 1), consisting mostly of epiphytic and epipetric taxa
displaying evolutionary traits that tend towards morphological and anatomical
reductions, are supported with fairly high reliability (Ebihara et al.). Assuming that the
common ancestor of Trichomanes was terrestrial, the occurrence of epiphytism related
to evolutionary regression was apparently quite a significant event in the genus, and at
least four independent evolutionary transitions led to the epiphytic habit. (3)
Geographical distribution data show that four of the eight clades are subcosmopolitan;
the NT clade is nearly confined to the Neotropics, while the Ca, Ce and Cr clades are
confined to the Palaeotropics. Peculiar monotypic trichomanoid “genera” distributed in
the southern hemisphere, which formed the basis of the Antarctic origin theory of
Hymenophyllaceae (Copeland, 1938, 1939), are all placed in derivative positions in the
present phylogeny.
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EBIHARA et al.: SYSTEMATICS OF TRICHOMANES
55
76
96
98
97
100
Cr
100
91
99
100
66
98
88
Va
100
73
50
99
98
100
80
88
66
57
90
Di
100
80
100
64
98
87
98
100
Po
100
100
100
Ca
100
97
100
93
99
77
62
NT
97
99
100
96
65
53
93
97
79
100
Pa
64
98
75
52
100
100
Ce
82
99
50
99
68
100
100
100
T. (Vandenboschia) fallax
T. (Vandenboschia) melanotrichum
T. (Crepidomanes) walleri
T. (Vandenboschia) schmidtianum
T. (Crepidomanes) latealatum
T. (Crepidomanes) christii
T. (Crepidomanes) bipunctatum
T. (Crepidomanes) kurzii
T. (Microtrichomanes) vitiense
T. (Crepidopteris) humile
T. (Gonocormus) mannii
T. (Gonocormus) minutum
T. (Nesopteirs) intermedium
T. (Nesopteris) grande
T. (Nesopteris) thysanostomum
T. (Vandenboschia) birmanicum
T. (Vandenboschia) speciosum
T. (Vandenboschia) davallioides
T. (Vandenboschia) maximum
T. (Vandenboschia) johnstonense
T. (Vandenboschia) radicans
T. (Vandenboschia) auriculatum
T. (Vandenboschia) rupestre
T. (Didymoglossum) krausii
T. (Didymoglossum) aff. angustifrons
T. (Didymoglossum) pinnatinervium
T. (Didymoglossum) exiguum
T. (Didymoglossum) gourlianum
T. (Microgonium) hildebrandtii
T. (Microgonium) tahitense
T. (Lecanium) membranaceum
T. (Microgonium) cuspidatum
T. (Microgonium) bimarginatum
T. (Microgonium) ekmanii
T. (Crepidopteris) endlicherianum
T. (Crepidopteris) vieillardii
T. (Polyphlebium) venosum
T. (Vandenboschia) colensoi
T. (Vandenboschia) angustatum
T. (Vandenboschia) capillaceum
T. (Vandenboschia) exsectum
T. (Vandenboschia) ingae
T. (Vandenboschia) diaphanum
T. (Vandenboschia) borbonicum
T. (Callistopteris) apiifolium Fiji
T. (Callistopteris) apiifolium Thai
T. (Trichomanes) crispum
T. (Trichomanes) egleri
T. (Trichomanes) robustum
T. (Trichomanes) pilosum
T. (Trichomanes) galeottii
T. (Trichomanes) pinnatum
T. (Trichomanes) arbuscula
T. (Trichomanes) alatum
T. (Trichomanes) trigonum
T. (Trichomanes) polypodioides
T. (Vandenboschia) scandens
T. (Trichomanes) lucens
T. (Feea) osmundoides
T. (Feea) mougeotii
T. (Feea) diversifrons
T. (Davalliopsis) elegans
T. (Trichomanes) ankersii
T. (Macroglena) laetum
T. (Selenodesmium) dentatum
T. (Selenodesmium) obscurum
T. (Selenodesmium) elongatum
T. (Selenodesmium) rigidum
T. (Macroglena) tamarisciforme
T. (Macroglena) meifolium
T. (Macroglena) flavofuscum
T. (Macroglena) caudatum
T. (Abrodictyum) boninense
T. (Macroglena) schlechteri
T. (Macroglena) asae-grayi
T. (Macroglena) strictum
T. (Macroglena) brassii
T. (Cephalomanes) atrovirens
T. (Cephalomanes) boryanum
T. (Cephalomanes) javanicum
T. (Pleuromanes) pallidum
Cardiomanes reniforme
Hymenoglossum cruentum
T. (Microtrichomanes) palmatifidum
T. (Microtrichomanes) nitidulum
H. hirsutum
H. polyanthos
Serpyllopsis caespitosa
H. tunbrigense
H. fucoides
Matonia pectinata
Polypodium glycyrrhiza
Osmunda cinnamomea
Angiopteris evecta (OG)
Figure 1. Evolutionary relationship of Trichomanes: a strict consensus of seven most
parsimonious trees retrieved from an unequally weighted rbcL data matrix (3717.96
steps, CI=0.32 and RI=0.74). Bootstrap values ≥50% are shown. Taxon names
generally follow the Morton’s (1968) system, and the trichomanoid genera defined by
Copeland (1938) are indicated in parentheses.
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FUTURE CHALLENGES
Usefulness of rbcL
In the case of Trichomanes, rbcL phylogeny is a useful tool for inferring the
“macroevolution” or global relationships among sublineages of the genus. Indeed, eight
principal clades have recently been defined as eight distinct genera (Ebihara et al.).
Although our results show that there is some useful genetic variation for discussing
relationships among species with closely-related rbcL sequences, further consideration
is necessary before adopting chloroplast-coding markers for this purpose. If
hybridization or reticulate evolution (e.g., Wagner 1954; van den Heede et al., 2003;
Ebihara et al., 2005) occurred in a clade, the relationship between the members as
reconstructed from chloroplast DNA, which is maternally inherited in ferns (Gastony &
Yatskievych, 1992), would not be accurate.
For example, Copeland’s genus Gonocormus, ranging from Africa to the Pacific
region, consists of several taxa segregated by leaf shape and proliferation (e.g.,
Trichomanes saxifragoides C.Presl, T. minutum Blume and T. proliferum Blume);
however, Yoroi and Iwatsuki (1977) argued that it should be difficult to recognize such
taxa morphologically and tentatively clustered them into a single species, T. minutum
(Iwatsuki, 1984). Although much genetic variation is also found in rbcL sequences
taken from several specimens of the Gonocormus species collected from various
localities, there does not seem to be a clear relationship between their morphological
and genetic variation (Ebihara, unpublished data). Considering the fact that diploid,
triploid and tetraploid series are reported for Gonocormus (Braithwaite, 1969, 1975;
Yoroi & Iwatsuki, 1977), reticulate evolution has probably occurred in this complex.
Another example is the Trichomanes (Vandenboschia) radicans complex (the Va clade),
which includes the American T. radicans Sw., the European T. speciosum Willd., the
African T. giganteum Willd., the Japanese T. orientale C.Chr. and the Asian T.
birmanicum Bedd. Our study, utilizing the biparently-inherited nuclear GapCp marker,
suggests that hybridization involving at least three biological units occurred in Japan,
despite an observed difference in rbcL of up to 1.66% (20/1206 bp; Ebihara et al., 2005)
among them. In addition, it is likely that both T. speciosum and T. giganteum are of
hybrid origin (Ebihara, unpublished data). Considering the high incidence of
allopolyploid hybridization in ferns (Soltis & Soltis, 1999), such examples may be
ubiquitous rather than exceptional. These results strongly indicate the importance of
biparently-inherited markers in species-level analyses. Analyses using sequences from
multiple gene regions are also necessary to clarify relationships among the clades that
are unresolved in the present rbcL phylogeny.
Importance of field observations
The evolution of life forms is one of the most interesting topics concerning
Hymenophyllaceae. Their habits are usually divided into several types, such as
terrestrial, epiphytic, epipetric, hemi-epiphytic and true lianas (Dubuisson et al.,
2003b), but in fact their ecological classification is routinely complex and requires
careful and precise in situ observations. For example, we have never seen herbarium
specimens of T. auriculatum with underground parts (roots). Rather, most herbarium
specimens consist only of epiphytic parts, i.e., leaves with climbing rhizomes. Our field
observations, however, reveal that the plant germinates on the ground and the rhizomes
climb up tree trunks. Field investigations are therefore essential for evolutionary
studies.
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57
Diversification history and conservation
Extant filmy ferns exclusively prefer moist and shady environments, but the habitat
where their common ancestor acquired its unique one-cell thick lamina in the Triassic
or earlier (AxSmith et al., 2001; Pryer et al., 2004) remains unknown for certain. The
fact that roughly half of the Hymenophyllaceae species are attributed to the
hymenophylloids, which are mostly epiphytic on tree trunks and diversified relatively
recently compared to trichomanoids (Hennequin et al., 2003; Schuettpeltz & Pryer,
2006), is consistent with the trend of extant fern diversification in angiosperm forests
(Schneider et al., 2004). Our phylogeny also suggests that a few species in Trichomanes
with an epiphytic habitat on trees may have recently acquired this habit.
Filmy ferns are one of the fern groups most sensitive to environmental changes,
particularly decreased humidity caused by deforestation. Habitat conservation and
conservation biology studies of already-endangered species (e.g., Rumsey et al., 1998)
are crucial for the future study of this family and to maintain its present diversity.
ACKNOWLEDGEMENTS
The authors are grateful to Dr. H. Schneider for valuable comments on the manuscript.
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species. Taxon 53: 935-948.
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59
SPATIAL PATTERN OF INDIVIDUAL GENETIC SIMILARITIES
IN POPULATION OF ASPLENIUM CETERACH
(ASPLENIACEAE: PTERIDOPHYTA)
VIRÁG KRÍZSIK,1 ISTVÁN PINTÉR2, ÁGNES MAJOR3
& GÁBOR VIDA1, 2
1
2
Department of Genetics, Eötvös Loránd University, Budapest, Hungary
HAS – ELTE Research Group for Evolutionary Genetics Budapest, Hungary
3
The Hungarian Natural History Museum
Keywords: intrapopulational structure, Aspleniaceae, RAPD, UPGMA
ABSTRACT
The genetic relatedness of individuals can give a population finer scale spatial
pedigree structure. The relation of genetic similarity and spatial distance refers
to the dispersal characters and reproductive relations existing among
individuals. Our main purpose here was to obtain information on the genetic
spatial pattern before a more profound spatial autocorrelation analysis of
Asplenium ceterach individuals. Three physically isolated subpopulation
patches of the tetraploid A. ceterach subs. ceterach were identified on the
southern rocky faces of the St. György Hill in Hungary. The genetic properties
were scored and cluster analysis, UPGMA, was carried out in three steps using
progressively larger samples: 42, 85 and 320 individuals were chosen. Cluster
analysis revealed a minimum of 70% genetic similarities among individuals
indicating intensive gene flow between subpopulations, but there was also
detectable correspondence between individual genetic similarities and spatial
position.
INTRODUCTION
The homosporous, autotetraploid, xerophytic rock fern, Asplenium ceterach L. subsp.
ceterach is one of three cytotypes of the species Asplenium ceterach (Reichstein 1981,
Vida 1965, 1973). Asplenium ceterach and related taxa are circumscribed by special
morphological characters within Asplenium. The taxonomic treatment of these ferns
have been the subject of debate; some authors have recognised Ceterach as a separate
genus within Aspleniaceae (Reichsten, 1981), but others consider it as a clade or
subgenus within the genus Asplenium (Pintér et al. 2002, Viane et al. 1993).
The study of geographical distribution and patterns of genetic diversity within
Asplenium cetercah has revealed its postglacial colonisation (Trewick et al. 2002). The
European pteridophyte flora has an ancient origin with the Mediterranean areas having
a major role as refugia during the glaciation periods (Vogel et al. 1999). Several
experimental studies demonstrate that autopolyploids are derived from diploids
(Manton, 1950; Reichstein, 1981; Vida, 1973). Although the ancestral (diploid) and
derived (polyploid) taxa have the same substrate preference, they have distinct
geographical distributions. During the glaciation periods the Balkans and Italy acted as
refugia for Asplenium ceterach. The diploids are mostly restricted to the Mediterranean
basin (Pintér et al. 2002.), whereas the tetraploids occupy areas further north in Europe.
The wide occurrence of the tetraploid Asplenium ceterach subsp. ceterach across
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Europe can be explained by its better dispersal capacity owing to single spore
colonisation in this taxon (Vogel et al. 1999). Analysing the haplotype patterns of
European Asplenium ceterach has elucidated recurrent polyploid formation and
colonisation from different population sources throughout Europe (Trewick et al.
2002). The tetraploid subspecies, Asplenium ceterach subsp. ceterach, is common and
widespread in Europe bit in Hungary, diploid Asplenium ceterach subsp. bivalens
predominates in natural habitats and the occurrence of tetraploid populations has been
reported only from the St. György Hill and the Buda Hills parts of the Transdanubian
Mountains (Vida 1973, Vida 1965). On the bases of chloroplast DNA analyses two
different haplotype sequences were identified in Hungary (‘red’ and ‘green’ according
to Trewick et al. 2002.). The haplotype sequence ‘red’ is found both in diploids and
tetraploids, whereas the ‘green’ haplotype is found only in tetraploids. Diploid
populations with the ‘red’ haplotype are described mainly from the Balkans, while
tetraploid populations of ‘red’ and ‘green’ haplotypes are rather frequent in SouthWestern Europe (Trewick et al. 2002). The tetraploid Asplenium ceterach populations
on the St. György Hill studied by us have the ‘green’ haplotype (Morgan-Richards,
Vogel unpublished). While the volcanic soil of the hill has been utilized for vinegrowing, the steep rocky slopes are natural habitats. These dry rocks may have
colonized after the last glaciation. Here, six physically isolated subpopulation patches
of A. ceterach subsp. ceterach were identified on the southern rocky faces of the hill
(Figure 1).
The genetic relatedness of individuals can reveal a fine-scale spatial pedigree
structure in a population (Vekemans & Hardy 2004). The relationship of genetic
similarity and spatial distance reflects the dispersal characters and reproductive
relationships among individuals (Hardy 2003). The aim of this study was to examine
the genetic constitution of this particular tetraploid population in the context of the past
colonisation events before a more profound spatial autocorrelation analysis. The
analysis of the population’s genetic constitution and its spatial structure as revealed by
molecular markers can provide information about microevolutionary processes
occurring when the individuals are linked by bonds of mating and parenthood (Hardy
& Vekemans 2002).
Genetic data derived from allozyme studies indicate that tetraploid Asplenium
ceterach shows high genetic variation between populations and low within population
variation, unlike diploids (Vogel et al. 1999). Genetic differences between diploid and
tetraploid populations result from differences in their breeding systems. The high
degree of population differentiation in the tetraploid taxon is caused by low level of
outcrossing. (Vogel, pers. com.). There was no evidence of vegetative propagation
among our cultivated individuals, nor are there any reported cases in the literature.
MATERIAL AND METHODS
In this study we present detailed information about the genetic relatedness of
individuals in a tetraploid population. For measuring spatial distances the exact position
of individuals was mapped (see below). This provides a 3D coordinate system for the
finer scale spatial analysis that follows. Use of a large number of RAPD products gives
a powerful tool for documenting offspring-parent relationships and dispersal distances
(Levitan and Grosberg 1993, Hadrys 1992) especially at lower taxonomic levels
(Rodrigues et al. 2002.).
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61
Sampling strategy
Individuals of Asplenium ceterach were collected from the southern rocky faces of St.
György Hill near Lake Balaton in Hungary. Here, six physically isolated subpopulation
patches were identified from where we collected 440 leaf samples (1 leaf per
individual). To obtain information on the genetic spatial pattern we surveyed three out
of six patches with progressively larger samples. The sample size was increased by
adding new individuals to the previously selected ones. These samples included 42, 85
and 320 individuals.
Constructing the spatial position map of the individuals
The task was to score the genetic properties of individuals and to give their exact
position in space in order to construct the 3-dimensional population genetic structure
before embarking on the fine scale analysis. For describing individual plants’ location
on the hill, GPS localized reference points were used. The WGS84 GPS-based
coordinates were converted to the Hungarian National Grid (EOV) (Timár et al. 2002,
Molnár & Timár 2002, Timár & Molnár 2002) (Figure 2). Reference coordinates were
the starting points of a grid, and before calculating the coordinates of each individual in
3D space. This coordinate system can be matched to a geographical map. The positional
data formed the basis for the future analysis of the genealogical connections.
Figure 1. St. György Hill and the subpopulation patches of A. ceterach subsp. ceterach
(1a, 1b, 2) on the southern rocky faces of the hill.
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Genetic studies
Genetic studies on the reliability of RAPD markers prove that this method is as suitable
for determinating genetic identity as the AFLP techniques (Albert et al. 2003). Bias of
polymorphic band selection is the weak point of this method. Many polymorphic bands
can be generated by only a few RAPD primers (Stewart & Porter 1995), but a few are
reliable. The applied bands have to be selected carefully. In our case, the decreased
amount of primers might influence precise indication. However, polymorphic bands
obtained from several primers give a more reliable result.
For total plant DNA isolation we used DNeasy Plant Mini Kit (Qiagen). The
extraction protocol was slightly modified for pteridophyte material. A dry mass of
leaves, 100mg of tissue was ground in liquid nitrogen the presence of Polyclar AT. A
volume of 600µl extraction buffer and 6-8µl Rnase (100mg/ml), and 16-17µl
β-mercaptoethanol were added. This suspension was homogenized and incubated at
65ºC for more than 1 hour with frequent and careful mixing. After adding 195-200µl
precipitation buffer, the mixing was repeated. This was followed by 10 minutes
incubation on ice and centrifugation at 12,000 rpm for 5-8 mins. The supernatant was
removed and applied to the QIAshredder column, combined in a 2ml eppendorf and
was centrifuged at 8000 rpm for 2 mins. Taking 1.5:1 proportion of the AP3/E and the
liquid that came down the column and it was mixed carefully again. This suspension
was centrifuged through the DNeasy mini column at 8000rpm for 1 min. Next a
washing buffer was added, followed by centrifugation at 8000rpm for 1min. This
process was repeated and an additional centrifugation step was applied for 3 mins.
The converted EOV coordinates of the subpopulational patches
167320
1b1
167300
1a
167280
1b1
1b2-1b3
1b4
1b5
167260
SÆN
167240
167220
23
167200
167180
22-24-25
167160
21
167140
167120
527940
527960
527980
528000
528020
528040
528060
528080
528100
528120
WÆE
Figure 2. To describe the location of individual plants on the hill, GPS localized
reference points were used. The WGS84 GPS-based coordinates were converted to the
Hungarian National Grid (EOV) (Timár et al. 2002, Molnár & Timár 2002, Timár &
Molnár 2002).
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Finally the DNS was eluted twice in 100µl and 50µl. The DNA samples were then
either stored at +4ºC for immediate use or stored at -20ºC.
For the DNA concentration quantification a self-made DNA standard was used. Its
concentration was measured by lambda 35 spectrophotometer and UV winlab program.
The DNA samples were diluted to 1ng/ µl.
RAPD procedure
For genetic identification the multilocus genotype (phenotype) of RAPD polymorphic
loci was used. The RAPD method uses a large set of oligonucleotide primers annealing
with various parts of the genome. RAPD markers produce good Mendelian characters,
typically, but not always dominant (Hillis et al. 1996). For the generation of genetic
data 60 oligonucleotid primers 10 basepairs long were screened. The best 8 were: OPA7, OPA-19, OPB-18, OPK-9, OPK-19, OPP-4, OPP-5, OPP-14. These primers
generated strong, clear and repeatable polymorphic bands.
For PCR reaction we applied 5µl (1ng/ µl) DNA template. The PCR reactions were
carried out in a PerkinElmer thermocycler. The amplification cycles ran: 60 sec at 94°C,
two cycle of 30 sec at 94°C, 30 sec at 41°C, 120 sec at 72°C, 20 cycles of 30 sec at
94°C, 15 sec at 39°C, 15 sec at 45°C, 90 sec at 72°C, 18 cycles of 30 sec at 94°C, 15
sec at 39°C, 15 sec at 45°C, 120 sec at 72°C and the final cycle 72°C 5min (Schneller
1998.).
A 4-5 hour long electrophoresis at 180-200V was used for the separation of the
fragments.
Data Analysis
The polymorphic bands of RAPD markers were scored 1 for presence and 0 for
absence. We applied simple matching coefficient for the dissimilarity matrix (Sokal &
Michener 1958). SM=a+d / a+b+c+d; (a, d: the numbers of the binaric variable
coincide, b, c: the numbers of the binaric variable do not coincide.)
For cluster analysis the UPGMA (unweighted pair group method with arithmetic
mean) method was used. The complete multivariate analysis and its presentation were
managed with the NTSYS-pc program Version 2. (Rohlf 2000, Podani 1997). The
results were illustrated with dendrograms reflecting the genotypic (phenotypic)
similarity between the objects.
The programmes SAHN module offers two options to deal with ties. The FIND
option constructed each alternative clustering. As there were several ties in the
dissimilarity matrix we had maximum 50 output trees. It was possible to compute their
cophenetic correlation to check how well the different tied trees represent the original
distance matrix (Rogrigues et al. 2002.). First the COPH module computed a
cophenetic (ultrametric) value matrix from each dendrogram. These matrices are used
to test the goodness of fit of clustering analysis by using MXCOMP module (Rohlf and
Sokal 1981). The module evaluates the cophenetic correlation (product moment
correlation: r) between the cophenetic value and the original distance matrices. We
selected trees with the highest evaluated correlation coefficient from among the tied
tries. The normalized Mantel test was also performed. The Mantel test statistic Z is
frequently used to measure the correspondence between matrices (Mantel 1967, Podani
1997).
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RESULTS
Our main purpose was to obtain information on the genetic spatial pattern before a more
profound spatial autocorrelation analysis of 440 positioned Asplenium ceterach
individuals. Based on the scored genetic properties and their spatial position the
individuals were clustered in three steps using progressively larger samples: 42, 85 and
320 individuals were chosen. The progressive increase of the size of the samples may
indicate the sensitivity of the spatial pattern analysis to the sample size.
The multivariate analysis of RAPD data revealed a high level of genetic relatedness
among either individuals or subpopulations. In all samples the assessment of the
minimum genetic similarity between individuals was approximately 70%, and at least
half of the plants showed 80-90% genetic similarity. The high level of genetic
relatedness between individuals either from the same or different subpopulations
suggested a high level of gene flow between the different subpopulations. Since
individuals show close genetic relatedness the UPGMA produced several alternative
clusters. The cophenetic correlations did not detect important difference in goodness of
fit among the dendrograms. Similar moderate correlation coefficient values were
computed for each alternative dendrogram. The matrix correlations (= normalized
Mantel stat Z) for different dendrograms were between r=0.62 –0.65 for 42 individuals,
r=0.71–0.73 for 85 individuals and r=0.618-0.625 for 320 individuals. The Mantel test
gave p (probability random Z< observed Z) = 1 for 1000 permutations for each tested
dendrogram. We selected one tree with the highest evaluated cophenetic correlation
coefficient (product moment correlation: r) (Rohlf and Sokal 1981) (Figures 3 & 4).
The first two sample sets (42, 85) showed a similar range of SM values for the
clusters (Figures 3, 4). The SM index varied from 70% up to 100%. In half of the
sampled individuals a similarity of more than 90% was detected. The genetic similarity
of the common individuals in the first two sample set was practically the same. Among
the first 42 individuals two homogenous subclusters were recognizable with 80-83%
similarity level (‘subpopulations 1a2’ and ‘2’). In the sample of 85 individuals another
homogenous ‘subpopulation 2’ cluster appeared with approximately 80% similarity.
The individuals of these two ‘subpopulation 1a2’ and ‘2’ also displayed their
cohesiveness )Figure 5). The rest of the dendrogram did not show any distinguishable
cluster.
The first two samples with 42 and 85 individuals were relatively small compared
with the total number of individuals. The doubling of the sample size did not alter the
intrapopulational spatial genetic pattern. The next step was the analysis of 320
individuals. The dendrogram constructed with 320 individuals showed not only
quantitative but also qualitative change in the picture of the spatial pattern. Increasing
the sample size gave a more detailed spatial pedigree structure along the dendrogram.
Subpopulations appeared as slightly separated subclusters of the tree. The minimum
similarity value for these characterized subclusters was around 77-80% (The minimum
genetic similarity between any two individuals is 70%). These distinguishable
substructures may demonstrate a significant number of non-random events in the
reproductive processes and propagation. This tree represents pedigree structure of these
ferns, and it differs from random. Large number of individuals appear to be outside their
characteristic subpopulational clusters scattered along the dendrogram.
The cluster analysis revealed minimum 70% genetic similarities among individuals.
The spatial genetic pattern indicates intensive gene flow between subpopulations, but
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also there was detectable correspondence between genetic similarities and spatial
position. This dual spatial pattern can be derived from the two step process of
pteridophyte reproductive biology and haploid-diploid life cycle, as the spores and
gametes have different dispersal capacities.
DISCUSSION
Asplenium ceterach is a homosporous fern with hermaphroditic gametophytes. The
spatial pattern of the genetically related individuals within the population is influenced
by both reproductive biology and dispersal characteristics. According to Klekowski’s
terminology two different kinds of mating types might produce different levels of
heterozygosity: intragametophytic selfing, yielding completely homozygotic
sporophytes (with the theoretical possibility of homeologous heterozygosity), and
intergametophytic mating between two sib or non-sib gametophytes resulting in
varying degree of heterozygosity (Klekowski 1979). The tetraploid Asplenium ceterach
is reported as a highly successful in-breeder infered from allozyme data (Trewick et al.
2002.).
The alternating haploid-diploid life-cycle is an important characteristic of ferns
(Page 2002). Spores and gametes have different dispersal capacity. The distance
between two gametophytes accomplishing mating is very much limited in space (Suter
2000). However, the distance over which spores travel is in comparatively unlimited.
This kind of haplotype transport can reorganize the population structure. Yet some
authers propose that 95% of the spore dispersal is within 1-10m from the parent plant
(Vogel et al. 1999.) The question is how the mating system and spore dispersal affect
Figure 3. Cluster analysis of 42 fern individuals. The individuals in the dendrogram are
labelled according to their subpopulational (1a, 1b, 2) and intra-subpopulational (1a1,
1a2, 1b1, 1b2, 1b3, 1b4, 1b5, 21, 22, 23, 24, 25) positions:
Subpopulations and their symbols:
1a1: 1a2: 1b1: 1b2: 1b3: 1b4: 1b5: 21:
22:
23:
24:
25:
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intrapopulation genetic pattern. The genetic similarity of neighbouring individuals may
help elucidate the nature of breeding system. Cluster analysis of the RAPD multilocus
phenotype suggests high levels of gene flow between these distant subpopulations and,
likewise, a correspondence between individual genetic similarities and spatial position.
Since the subpopulations are further than 10m apart, this suggests that spore dispersal
should exceed 10 m. But cluster analysis is not reliably informative to dissect the
influence of spore travel from the influence of mating type in the determination of
individuals genetic relatedness. This can be carried out by further, more profound
spatial autocorrelation analysis.
Figure. 4. Cluster analysis of 85 fern individuals. The individuals in the dendrogram
are labelled according to their subpopulational (1a, 1b, 2) and intra-subpopulational
(1a1, 1a2, 1b1, 1b2, 1b3, 1b4, 1b5, 21, 22, 23, 24, 25) positions:
1a1: 1a2: 1b1: 1b2: 1b3: 1b4: 1b5: 21: 22:
23:
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Figure. 5a. Cluster analysis of 320 fern individuals. The individuals in the dendrogram are labelled according to their subpopulational (1a, 1b,
2) and intra-subpopulational (1a1, 1a2, 1b1, 1b2, 1b3, 1b4, 1b5, 21, 22, 23, 24, 25) positions:
1a1: 1a2: 1b1: 1b2: 1b3: 1b4: 1b5: 21:
22:
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25:
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Figure. 5b. Cluster analysis of 320 fern individuals. The individuals in the dendrogram are labelled according to their subpopulational (1a, 1b,
2) and intra-subpopulational (1a1, 1a2, 1b1, 1b2, 1b3, 1b4, 1b5, 21, 22, 23, 24, 25) positions:
1a1: 1a2: 1b1: 1b2: 1b3: 1b4: 1b5: 21:
22:
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ACKNOWLEDGEMENTS
I would like to express my thanks to Maria Takács and Maria Tuschek for their kind
help, and many thanks for Gábor Timár and Bálint Halpern for their help in the GPS
work.
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HARDY, O. & VEKEMANS, X. 2002. SPAGeDi: a versatile computer program to
analyse spatial genetic structure at individual or population levels. Molecular
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KLEKOWSKI, E. 1979. The genetics and reproductive biology of ferns. In: The
Experimental Biology of Ferns. DYER, A.F. (Ed.) Academic Press London.
LEVITAN, D.R. & GROSBERG, R.K. 1993. The analysis of paternity and maternity in
the marine hydrozoan Hydractinia symbiolongicarpus using random amplified
polymorphic DNA(RAPD)markers. Molecular Ecology 2:315-328.
MANTON, I. 1950. Problems of cytology and evolution in the Pteridophyta.
Cambridge University Press.
MANTEL, N. A. 1967. The detection of disease of clustering on generalized regression
approach. Cancer Researches 27:209-220.
MOLNÁR, G. & TIMÁR, G. 2002. Az EOV – koordináták nagypontosságú közelítése
Hotine-féle ferdetengelyû Mercator-vetülettel (in Hungarian with English
summary). Geodézia és Kartográfia [Budapest] 54(3) 18-22.
PAGE,C.N. 2002. Ecological strategies in fern evolution: a neopteriodological
overview. Review of Palaeobotany and Palynology 119 (2002) 1-33.
PINTÉR, I; BAKKER, F.; BARRETT, J.; COX, C.; GIBBY, M.; HENDERSON, S.;
MORGAN-RICHARDS, M.; RUMSEY, F.; RUSSELL, S.; TREWICK,
S.;SCHNEIDER, H.;VOGEL, J. 2002. Phylogenetic and biosystematic
relationships in four disjunct polyploid complexes in the subgenera Ceterach and
Phyllitis in Asplenium (Aspleniaceae). Organism. Diversity and Evolution 2, 299311.
PLUMBI, S. R. 1996. Nucleic Acids II: The Polymerase Chain Reaction. In: HILLIS,
D. M.; MORITZ, C.; MABLE B.K. (Eds.) Molecular Systematics, pp. 205-247.
Sinauer Associates, Inc.
PODANI, J. 1997. Bevezetés a többváltozós biológiai adatfeltárás rejtelmeibe. Scientia
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REICHSTEIN, T. 1981. Hybrids in European Aspleniacea (Pteridophyta). Botanica
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RODRIGUES, F. M., DINIZ-FILHO, J. A. F., BATAUS, L. A. M. & BATOS, R. P.
2002. Hypothesis testing of genetic similarity based on RAPD data using Mantel
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SCHNELLER, J.J. 1998. Patterns of genetic variation detected by RAPDs suggest a
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71
ASSESSING THE CONSERVATION STATUS OF PTERIDOPHYTES,
A CHALLENGE FOR THE GLOBAL STRATEGY FOR PLANT
CONSERVATION
S. BLACKMORE & K. WALTER
Royal Botanic Garden Edinburgh, 20a Inverleith Row, Edinburgh, EH3 5LR,
Scotland, UK
Key words: conservation status, Global Strategy for Plant Conservation,
Pteridophytes, Red Data Lists.
ABSTRACT
The Global Strategy for Plant Conservation (GSPC) sets out a series of activities
with targets intended to halt the decline in plant biodiversity by 2010. This
article examines the current state of knowledge concerning the conservation
status of ferns in relation to Target 2 of the GSPC. The change in criteria used
by the World Conservation Union (IUCN) for assessing conservation status has
led to data generated prior to 1997 being marginalised. The latest information,
using the revised criteria and published in 2003, refers to a smaller number of
pteridophyte species, with only ten species being common to both the 1997 and
2003 assessments. There is an urgent need to capture the knowledge that
pteridologists and other specialists undoubtedly have, relating to the
conservation status of ferns and fern allies, and to incorporate this into Red Data
Lists to provide firmer foundations for the GSPC.
INTRODUCTION
The biodiversity crisis affecting our world is undoubtedly a real problem (Pimm et al.,
1995; Heywood & Watson, 1995; Gomez-Pompa, 2004) and, like global warming with
which it is intimately connected (Thomas et al., 2004), it poses great challenges to the
future of humanity (Wilson, 2002). The biodiversity crisis might, however, be much
more amenable to solution than global climate change if appropriate practical
conservation policies are implemented at the local level around the world. The Global
Strategy for Plant Conservation (GSPC) adopted in 2002 under the Convention on
Biological Diversity sets out measurable targets as milestones towards halting the loss
of plant biodiversity by 2010 (Anon, 2003). At present, however, the rate of progress on
global conservation programmes is severely constrained by the lack of investment in
biodiversity science (see for example, Blackmore, 2002; House of Lords, 2002;
Wheeler et al., 2004).
The global status and conservation of pteridophytes has been the focus of a recent
conference, the proceedings of which provide an excellent overview (Dyer et al. (eds),
2002) and there are many published case studies for particular taxa. The purpose of this
contribution is to review the current state of knowledge about pteridophytes in relation
to selected targets of the GSPC. The point we wish to emphasise is that whilst the extant
pteridophytes are less species rich than angiosperms and might therefore seem an easier
group to conserve, much of the basic information needed to do this effectively is
lacking. In particular we highlight the paucity of information concerning the
conservation status of pteridophytes.
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CONSERVATION STATUS OF PTERIDOPHYTES
The first three targets of the GSPC are all concerned with understanding and
documenting plant diversity and concern the underpinning knowledge needed for
conservation efforts to be demonstrably effective. Whilst most plant taxonomists are
aware of the extent to which the initial exploration and documentation of biodiversity
is incomplete, it continues to surprise many people, including policy makers, that many
new species are discovered each year and how incomplete the documentation of life on
earth is. Perhaps the most widely authoritative estimate, derived from the global
biodiversity assessment (Heywood & Watson, 1995), is that 1.7 million species have so
far been described out of an estimated total of 13 million. However, current estimates
are approximate at best and the question of how many flowering plant species there are,
for example, has seen much recent discussion. For pteridophytes, Roos (1996)
estimated that between 10,500 and 11,300 species have been described from a total of
between 12,000 and 15,000. In other words somewhere between 6% to as many as 30%
of pteridophytes species have not yet been discovered. This has an important bearing on
Target 1 of the GSPC: the preparation of “a widely accessible list of all plants, as a step
towards a complete world flora.” The importance of the target derives from the widely
acknowledge fact that one cannot be certain of conserving species without even
knowing that they exist, although one important aspect of protected areas, especially in
biodiversity hotspots is that, hopefully, they will harbour a good proportion of the
unknown taxa. The technical rationale accompanying Target 1 makes it clear that a
working list rather than a definitive list of species is the objective for 2010. Any such
list can, by definition, only be a list of the known species and thus for pteridophytes our
current state of knowledge would permit a list that contains somewhere between 70%
and 94% of the estimated world total. It is encouraging to note that good progress has
already been made towards the achievement of Target 1. The international plant names
index (www.ipni.org) provides a highly accessible database of names and associated
bibliographic information and now incorporates the data from the published volumes of
Index Filicum (Johns, 1996) on the names of ferns and fern-allies. Web-based resources
are not only the most accessible means of delivering such information but, in many
ways, also the most appropriate, given that they can be updated much more readily than
printed
literature.
The
world
of
ferns
website
(http://homepages.caverock.net.nz/~bj/fern/) is another highly accessible source of
information on names and synonymy in pteridophytes.
A much less favourable situation applies to Target 2: “a preliminary assessment of
the conservation status of all known plant species, at national, regional and international
levels”. As the technical rationale for Target 2 makes clear, the conservation status has
so far been assessed for over 60,000 plant species, of which 34,000 are classified as
globally threatened with extinction (Anon, 2003) and, given the vast amount of working
still to be done, the intention is to undertake a preliminary assessment for the “datadeficient” species by 2010. The scale of this challenge emphasises just how little
information we have to hand on the conservation status of plants in general. The
situation for pteridophytes will be explored in more detail.
Information derived from the IUCN website (www.iucnredlist.org) in June 2004
indicated that there are 13,025 known species of pteridophytes. Of these, conservation
status has only been determined for 180 of these species and as a result 111 species were
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BLACKMORE et al.: CONSERVATION STATUS OF PTERIDOPHYTES
73
considered to be threatened in 2003. Although, on this information, only 1% of the total
number of species of ferns and fern allies are known to be threatened, with two species
being extinct, 62% of those that have actually been evaluated are considered to be
threatened. More complete information on the status of 186 species of pteridophytes in
2003 from the IUCN web site is summarised in Table 1. Clearly we are in a state of
relative ignorance about the true scale of the threats to pteridophytes species. If 62% of
all pteridophytes were threatened, this would constitute some 8,075 species. There is,
of course, no sound scientific basis for making such a projection.
It is interesting to compare this most recent summary of knowledge with earlier
information available on the web. The web site of the World Conservation Monitoring
Centre (www.unep-wcmc.org) presents information from the earlier assessment
contained in Walter & Gillett (1997). According to this source, a more comprehensive
survey of the conservation status of pteridophytes had already been previously
undertaken (Table 2). In 1997, using the IUCN’s earlier system for evaluating threats,
768 species of pteridophytes had been assessed (compared to just 111 under the criteria
in use in 2003), of which nine were extinct in the wild and a further 13 were suspected
of being extinct. The 1997 Red Data List (Walter & Gillett, 1997) did not list species
that were of indeterminate status or were not threatened.
Although good reasons were advanced for changing the criteria and categories for
threatened plants between the 1997 and the 2003 assessments, the fact remains that
hard-earned information recorded prior to 1997 was marginalised as a result. We
estimate that preparation of the 1997 Red List required the equivalent of about 250
person years of work. Furthermore, very little re-assessment of previously studied
species, to bring the pre-1997 data into the new system, has been undertaken. Only ten
species out of the 768 species assessed as threatened in 1997 have been reassessed in
the 2003 list (Table 3). In general it is easy to see, from the list of those species that
occur in both the 1997 and the 2003 list, that reassessment is either not taking place is
or is restricted to assessments of particular groups of endemic species, such as the
Table 1. Data on pteridophytes from the 2003 Red Data List. EX = extinct, EW =
extinct in the wild, CR – critically endangered, EN = endangered, VU = vulnerable, NT
= near threatened, LC = least concern, DD = data deficient. Species listed as NT, LC
or DD are not considered to be threatened and therefore the total number of threatened
pteridophytes detailed in Table 1 is 111, comprising those listed as CR, EN, or VU.
EX
EW
CR
EN
VU
NT
LC
DD
Total
LYCOPSIDA
0
0
1
2
8
1
1
0
13
SELAGINELLOSIDA
0
0
0
0
1
1
0
0
2
ISOETOPSIDA
0
0
0
0
1
0
0
0
1
POLPYPODIOPSIDA
2
0
26
22
50
12
7
45
164
Totals
2
0
27
24
60
14
14
45
186
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FERN GAZ. 18(2): 71-76. 2007
pteridophytes of Ascension Island.
As a consequence of the extremely low rate of re-assessment and because of the
change in criteria it is almost impossible to extract evidence of trends in the
conservation status of pteridophytes, or other plants. We consider that this poses one of
the greatest challenges to the GSPC. Although we strongly support efforts to move
ahead with data gathering under the new system we are concerned that this is not
happening rapidly enough. The reasons for this are difficult to determine but we suspect
that for many species, especially for endemics, more information is known, informally
by specialists on particular taxa or localities, than has been recorded. We therefore urge
pteridologists, and other botanists, to enter such information as they do have into the
IUCN system.
The importance of a better state of knowledge on the conservation status of ferns
impacts on other targets within the GSPC. For example, Target 8 aims to have 60% of
the world’s threatened species conserved in situ and 10% of them to be included in
species recovery programmes. On the basis of the Red List data 2003, this would
require only 66 threatened species of fern to be conserved in situ, with 6 being the
subject of recovery programmes. That has probably already been achieved but this is
hardly a satisfying observation in that it reflects ignorance rather than progress.
CONCLUSIONS
Current syntheses of the documented taxonomic diversity of pteridophytes provide a
sound basis for the achievement of a preliminary checklist of the world’s ferns and fern
allies in the context of Target 1 of the GSPC. This is not to deny the need for significant
further revisionary work to define taxonomic concepts, particularly fro large and
complex genera. Syntheses of information on the conservation status of known
pteridophytes are, in contrast, woefully inadequate. This results in part from changes of
Table 2. Summary of the status of pteridophytes recorded in the 1997 Red List (Walter
& Gillett, 1997). Abbreviations: Ex = extinct, Ex/E = possibly extinct, E = endangered,
R = rare, V = vulnerable, I = indeterminate. Total threatened species is the number of
species assessed as Ex/E, E, V, R or I (in other words, it excludes extinct species). %
threatened is the total threatened species as a percentage of the number of known
species.
Ex Ex/E E V R I
Total
Number
%
Threatened
of
Threatened
Species known
species
LYCOPODIOPSIDA
1
0
3
6 11
3
23
519
4.4
SELAGINELLOPSIDA
1
0
3
5
9
6
23
713
3.2
ISOETOPSIDA
2
0 11
4 22
2
39
79
49.7
POLYPODIOPSIDA
5
13 85 97 356 133
683
9053
7.5
Totals
9
13 102 112 398 144
768 10,364
7.4
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BLACKMORE et al.: CONSERVATION STATUS OF PTERIDOPHYTES
75
criteria used for evaluating conservation status but even more fundamentally from the
low proportion of known species for which assessments of status have been recorded.
There is now an urgent need to harness the expertise of people who know about
pteridophytes and their status and to complete the recording of conservation status if we
are to understand the conservation needs of ferns and fern allies properly. This case
study focuses on pteridophytes but it is likely that a similar situation applies to plants
in general. We end by urging pteridologists and others to familiarise themselves with
the procedures for assessing conservation status
(see: www.iucn.org/themes/ssc/redlists/RLcats2001booklet.html).
Table 3. Summary information on the ten species of pteridophytes that feature in both
the 1997 and the 2003 Red Lists with an assessment of whether their status in the wild
appears to have improved or worsened and their geographical distribution.
1997 Red
List
2003 Red
List
Change in
Status
Distribution
Anogramma ascensionis
Ex/E
Ex
Unchanged Ascension
Asplenium ascensionis
R
NT
Improved
Ctenitis squamigera
E
CR
Unchanged Hawaii
Cyathea bipinnata
R
VU
Worse
Ecuador
Cyathea heliophila
R
EN
Worse
Ecuador
Dryopteris ascensionis
Ex/E
Ex
Unchanged Ascension
Elaphoglossum pellucidum
R
CR
Worse
Hawaii
Marattia purpurascens
R
NT
Improved
Ascension
Pteris adscensionis
E
CR
Unchanged Ascension
Xiphopteris ascensionense
R
NT
Improved
Ascension
Ascension
76
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REFERENCES
ANON. 2003. The Global Strategy for Plant Conservation. Botanic Gardens
Conservation International, UK.
BLACKMORE, S. 2002. Biodiversity update: progress in taxonomy. Science 298: 365.
DYER, A.F., SHEFFIELD, E. & WARDLAW, A.C. (eds). 2002. Fern floras worldwide:
threats and responses. Fern Gazette 16: (6,7,8).
GÓMEZ-POMPA, A. 2004. The role of biodiversity scientists in a troubled world.
Biosicience: 54: 219 – 227.
HEYWOOD, V.H., & WATSON, R.T. 1995. Global Biodiversity Assessment.
Cambridge University Press, Cambridge.
HOUSE OF LORDS. 2002. What on Earth? The threat to the science underpinning
conservation. HL Papers 118(i) and 118 (ii).
JOHNS, R.I. 1996. Index Filicum: Supplementum Sextum pro annis 1976 – 1990.
Royal Botanic Gardens, Kew.
PIMM, S.L., RUSSELL, G.J., GITTLEMAN, J.L. & BROOKS, T.M., 1995. The future
of biodiversity. Science 269, 347 - 350.
ROOS, M.C. 1996. Charting tropical plant diversity: Europe’s contribution and
potential. In: BLACKMORE, S. & CUTLER, D. (Eds) Systematic Agenda 2000:
The Challenge for Europe. Pp. 55 – 88. Linnean Society Occasional Publication –
Number 1. Samara Publishing, Tresaith.
THOMAS, C.D., CAMERON, A., GREEN, R.E, BAKKENES, M., BEAUMONT,
L.J., COLLINGHAM, Y.C., ERASMUS, B.F.N., FERREIRA DE SIQUEIRA, M.,
GRAINGER, A., HANNAH, L., HUGHES, L.; HUNTLEY, B., VAN
JAARSVELD, A.S., MIDGLEY, G.F., MILES, L., ORTEGA-HUERTA, M.A.,
PETERSON, A.T., PHILLIPS, O.L. & WILLIAMS, S.E. 2004. Extinction risk from
climate change. Nature 427: 145 148.
WALTER, K.S. & GILLETT, H.J. (Eds) 1997. IUCN Red list of threatened plants.
IUCN, Gland.
WHEELER, Q.D., RAVEN, P.H. & WILSON, E.O. 2004. Taxonomy: impediment or
expedient. Science 303: 285.
WILSON, E.O. 2002. The future of Life. Random House Inc. New York.
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BOOK REVIEW
ENCYCLOPEDIA OF GARDEN FERNS. Olsen, S. 2007. Hardback, 444pp., 692
colour photographs, 2 line drawings. ISBN 13: 978-0-88192-819-8. Timber Press,
Portland. US$59.95; £40.00. 11” x 8” (28cm x 22cm).
This is a highly informative and enjoyable book that most fern enthusiasts will wish to
own. It is intended primarily for fern collectors and growers, amateur and commercial
(see the review in Pteridologist, 2007), but there is much to interest fern biologists. Its
considerable size and weight, and large number of colour photos, suggest a ‘coffee table
book’ but these usually receive superficial attention and Sue Olsen’s book will be
regularly consulted for detailed information on identification, cultivation and, in many
cases, natural habitats of nearly 1000 species. Despite its all embracing title, there is a
strong North American emphasis and the book is similar in scope to John Mickel’s
Ferns for American Gardens (second edition 2003, first edition 1994, reviewed
American Fern Journal 84 (3), p.104), but contains almost twice as many species. There
is an overlap also with Rickard’s book The Plantfinder’s Guide to Garden Ferns (2000,
published in UK by David & Charles, Newton Abbot, reviewed Fern Gazette 16(3), pp.
146, 167) but Rickard writes from a British perspective and gives more attention to the
horticultural cultivars popular during the British fern craze. A strong case can be made
for acquiring all three books because of their complementary strengths.
Olsen’s Encyclopedia contains 5 chapters, 8 Appendices, a Glossary, a Reference
List and an Index. The ‘ferns’ of the title are taken to include not only horsetails, as we
are now required to do, but also Psilotum and the lycophytes Diphasiatrum, Huperzia,
Lycopodiella, Lycopodium and Selaginella. Isoetes and Tmesipteris are not, however,
discussed. An initial preamble includes acknowledgement of the owners of gardens
where she has taken photographs. Amongst these are several past and present BPS
members including ex-Presidents Martin Rickard and Alastair Wardlaw. Then four short
chapters deal with, in turn, “Ferns through the ages”, “Cultivating ferns”, with guidance
on topics from garden habitats to companion planting, “Propagating ferns”, with
descriptions of propagation techniques together with an outline of the typical life cycle,
and “Fern structure and basic diagnostics” with a brief introduction to the main
characters, including frond morphology, used in fern recognition.
Chapter 5, “Ferns from around the world”, is the heart of the book and, with 320
pages, the major part of it. Nearly 1000 fern species are arranged in alphabetical order
of genera; about 700 are accompanied by colour photographs. For most species the
entry begins with the Latin name, the common name, the translation of the Latin
specific epithet, the height, whether it is “evergreen” or “deciduous”, and the plant
hardiness zone(s) to which it is adapted. Then follows a concise but informative
description of diagnostic features, an indication of its natural geographic range and
habitat, and guidance on cultivation, together with additional helpful comments when
appropriate. Some less frequently cultivated species are represented by a brief entry
under “Shorter Notes”.
In the Appendices are a Plant Hardiness Zone map for the USA, a comparable map
for Europe (unfortunately with a different colour coding), a list of ferns that have been
awarded an RHS Garden Merit Award, and a list of ferns designated “Great Plants” by
the Elisabeth C. Miller Botanical Garden, Seattle. Appendix 4 consists of “Top twenty
ferns” lists provided by 19 enthusiasts from USA, UK and Germany and representing
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hardiness zones from 4 to 11. Appendices 5 to 8 provide recommendations for ferns for
special situations, a brief introduction to fern societies (are the BPS, AFS and the Hardy
Fern Foundation really the only fern societies?), a list of gardens noted for ferns, and a
list of commercial growers. The book finishes with a glossary of about 120 essential
terms including “discombobulated: the feeling an uninitiated fern amateur might have
when encountering technical terms”, a somewhat idiosyncratic list of books and papers
referred to in the text, and an index of Latin and common plant names.
The list of featured species is impressive but nevertheless represents less than 10%
of the world flora. The criteria for selecting the chosen species are not explicitly stated
but most of the ferns mentioned are temperate species hardy at temperatures below
freezing during winter (Hardiness Zone 9 or below), thereby eliminating most ferns,
which are tropical. Many of those chosen are familiar garden subjects but I cannot
believe that all of them were listed because they are commercially available. Ferns from
all over the world are represented, but not every fern in cultivation is included. Even for
Britain, the list is not quite complete. For example, Cystopteris dickieana is recognised
in the text as a distinct species and shown in a photograph (p.168) but there is no
descriptive entry for it even though this attractive dwarf fern is easy to grow, and plants
propagated from spores collected at the type locality in a Scottish seaside cave have
been widely distributed, at least among British fern enthusiasts. I suspect that the list
simply represents those species which can be grown somewhere in the USA or are
otherwise known to the author. Nevertheless, it will be possible to identify most garden
ferns using the descriptions and illustrations provided.
With such an eclectic list of ferns, it is important to provide guidance to the readers
about which ferns will grow in their area. Guidance on suitable climatic conditions is
given by means of Plant Hardiness Zones, based on the “average annual minimum
temperature” and thus a guide to how cold it can get in winter. This guidance is widely
used in the USA but it is less familiar in the British Isles, perhaps because it is not
sufficiently discriminatory. Most of Britain away from the coast, from Wick to
Winchester, is in Zone 8. In continental Europe, Zone 8 extends from Stavanger to
Salamanca. In eastern USA, Zone 8 extends from southern North Carolina to northern
Florida. Clearly, few if any ferns will grow equally well throughout Zone 8 because of
other environmental factors. Hardiness Zones are, at best, broad indicators of which
ferns are worth trying.
In the introductory chapter on propagation (p.74), there is a brief account of hybrid
formation, but the treatment of fertile allopolyploids is inconsistent and potentially
confusing to the uninitiated. For example, there is no indication that research has
revealed that Polystichum aculeatum is the allotetraploid hybrid of two ancestral
species, P. setiferum and P. lonchitis, formed by ancient cross-fertilisation followed by
chromosome doubling. However, tetraploid Dryopteris filix-mas, with a similar type of
history thought to involve D. caucasica (or something similar) and D. oreades, is stated
to be “Dryopteris caucasica x D. oreades” and within the description is mention of the
fact that it is a fertile hybrid. Woodsia alpina, another allotetraploid species, is defined
as Woodsia glabella x W.ilvensis” and this time, its status as “a fertile hybrid” is
emphasised in the first sentence of the description. Some readers may not realise that
all three species have arisen in a similar way, and that indeed many fern species are
fertile polyploids derived from two or more diploid ancestors. In the absence of any
mention of ploidy or chromosome number in the species entries, they might also find it
difficult to distinguish these fertile polyploids of ancient hybrid origin from recent,
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sterile, hybrids with no chromosome doubling. There is also an inconsistent treatment
of these recent hybrids. For example, Dryopteris x australis (p.214) and D. x boottii
(p.216) are correctly presented as recent sterile natural hybrids but a similar hybrid, D.
x complexa, is described as fertile without explaining that the apparent fertility is due
to inherited apomixis (apogamy plus diplospory) rather than to the compatibility of the
parental genomes. To compound the confusion, Dryopteris x remota, a recent rare
hybrid of D.affinis and D. expansa, is listed as “D. remota”, implying a species, but then
described as a “fertile hybrid”, spore production being again due to inherited apomixis.
A consistent treatment of hybrids, and some explanation of the role of hybridisation and
polyploidy in species formation and evolution in ferns, perhaps in Chapter 1 “Ferns
through the Ages”, would have been helpful.
Whilst acknowledging that fern gardening is all about the decorative sporophytes,
in a book called an Encyclopedia I would like to have seen a little more about the vital
gametophyte generation. There is a very brief mention in the outline of the fern life
cycle (p.69) and in the account of propagation (p.72), and on p.151 the author
comments that Blechnum spicant is “the only species where I have seen prothalli in
nature”. The ecology of gametophytes in natural habitats is largely unknown and cannot
therefore inform the choice of conditions for propagation from spores. However an
indication of the diversity of form of gametophytes, with photographs, would have been
welcome. I would also expect a mention of the perennial gametophytes that can be
cultivated, but Anogramma is not included, even as a sporophyte, and under
Trichomanes (p.389) the statement that filmy ferns are “sometimes only seen in a
permanent state without sporophytes” will not be easily understood by readers who are
not familiar with examples of independent gametophytes.
There are commendably few mistakes or typographical errors, though on p.69, “one
pound [2kg] of spores” leaves the reader wondering whether Cyathea medularis
produces 454gm or 2000gm of spores each year.
To the grower, I would suggest that you use the horticultural information in this
book as a useful guide but don’t be inhibited by indications that a species might not be
successful in your conditions; you might be pleasantly surprised. Not knowing that
Ceterach officinarum is “wretchedly difficult” (p.49), I wedged my only plant between
small rocks at the foot of a south-east facing slope and, some ten years later and with
no further assistance other than some lime chippings, it is still growing, slowly but
steadily. Less than 2m away I have three plants of the calcifuge Cryptogramma crispa
of similar age and also slowly increasing even though this species is said by Olsen to
be “almost impossible to cultivate” (p.180). Although these successes are more due to
a lucky combination of garden conditions than to horticultural prowess, it does
demonstrate that whatever the advice, it is always worth trying a fern you wish to grow,
preferably with some attempt to imitate its natural habitat, and in several different
conditions of aspect, shade and moisture.
To the fern biologist, I would say don’t dismiss this book as solely for gardeners. It
is a goldmine of interesting information on the ecology and growth requirements of a
wide selection of species.
A.F. Dyer
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will be refereed
aligned.
aligned.
a legible form.
Supplements.
FERN GAZ. 18(3):81-90. 2008
81
THE CYTOTAXONOMY OF THE ASPLENIUM STUHLMANNII
COMPLEX (ASPLENIACEAE, PTERIDOPHYTA) IN AFRICA
A. F. BRAITHWAITE
4 Kendal Drive, Beeston, Nottingham, NG9 3AW
(Email: [email protected])
Key words: Asplenium mantoniae, Asplenium jaundeense, new species, chromosome
numbers.
ABSTRACT
A new octoploid species, Asplenium mantoniae, intermediate between two
tetraploids species, A. stuhlmannii and A. jaundeense, is described from West
Africa. The morphology and distribution of members of the complex are
consistent with the view that the octoploid has probably arisen by hybridization
of the two tetraploids and chromosome duplication
INTRODUCTION
Asplenium stuhlmannii Hieron. was first described from tropical East Africa and is
widely distributed from northern Tanzania to the southern Sudan. Similar, though not
identical, material occurs in West Africa and was placed in A. stuhlmannii by TardieuBlot (1953) and Alston (1959) and reported by Manton (1959) to be octoploid with n =
144 chromosomes, based on meiotic analysis. Subsequent cytological examination of
East African material has shown the genuine A. stuhlmannii to be tetraploid with n = 72
chromosomes (Appendix in Braithwaite 1964; Fig.1). Further studies of the two
cytotypes suggest that they are sufficiently distinct morphologically and geographically
to merit recognition as two separate species and the octoploid material from West Africa
is described here as a new species, A. mantoniae. The new species also bears some
resemblance to A. jaundeense Hieron., a tetraploid species (Fig.1) from West Africa,
and its possible relationships to the two tetraploids is discussed.
Figure 1. Acetocarmine meiotic squash preparations. A, Asplenium stuhlmannii n = 72,
Somalia (Brittish Somaliland; B, A. jaundeense n = 72, arrow points to an out of focus
bivalent,Yaunde, Cameroon. Scales = 10µm
Locality and origin
Chromosome number
Spore size*
Length
A. stuhlmannii Hieron
A. mantoniae sp. nov.
Breadth
Somalia, collected by Mr Desmond Kelsall (O. S Dept, n = 72
St Andrews) on a hill walking expedition in 1949, ex
University Botanic Gardens, St Andrews.
40·9 ± 2·49
30·3 ± 2·06
Kenya, living plant sent to Kew with a consignment of n = 72
orchids
39·7 ± 2·0
26·4 ± 1·0
Nigeria, plant raised in Leeds from spores taken from n = 144
Hambler 513 (BM)
47·1 ± 3·57
31·7 ± 2·36
Ghana, locality not known, collected by Adams (see n = 144
Manton 1959), plant in this study raised from spores
taken from herb. specimen of Manton’s plant
45·7 ± 2·13
30·4 ± 2·13
Cameroon, Yaunde, collected for the author as a living n = 72
plant and first established at Kirstenbosch in 1961 then
sent to Kew and subsequently transferred to Leeds
34·1 ± 2·85
23·4 ± 1·69
* means ± SD (n = 100)
Table 1. Locality, source, chromosome numbers and spore sizes of living plants of the Asplenium stuhlmanii complex.
FERN GAZ. 18(3): 81-90. 2008
A. jaundeense Hieron.
82
Species
BRAITHWAITE: THE ASPLENIUM STUHLMANNII COMPLEX
83
Live plants for cytological investigation were accumulated from a variety of sources
(see Table 1) and finally established in the Botanic Garden, University of Leeds. In
addition, all the relevant herbarium material at BM and K was examined. Voucher
specimens will be deposited in BM.
Chromosome counts were made on acetocarmine squash preparations of meiosis
using the standard method for ferns (Manton 1950).
Spore samples for measurement were mounted in either gum chloral or Depex and
measured using a calibrated eyepiece and ×40 objective.
THE SPECIES
Asplenium stuhlmannii Hieron.
In Engl. Pflanzenw. Ost.-Afr. C: 83, 1895; type: Kassesse, Towalio in Western des Vict.Njansa, Stuhlman 936 (B!—lectotype), 953!, 1179, 3246, 3981a (B—paratypes); non
Tardieu-Blot, in Mem. Inst. fr. Afr. noire 28: 190, t. 36, fig. 3, 1953; Alston in The Ferns
and Fern Allies of West Tropical Africa, suppl. 2 ed. Flora of West Tropical Africa, 59,
1959.
Asplenium stuhlmannii var. laciniata Hieron., in Engl. Pflanzen. Ost-Afr. C: 83,
1895; type: Gumango, Lande der Niamniam, Schweinfurth 3915 (B!—holotype;
BM!, K!—isotypes).
Asplenium amoenum C. H. Wright, in Johnstone, The Uganda Prot. 1: 326, 1902,
non Presl 1836; type: Uganda, E. Toro, R. Kiawiume, alt. 5000´. Doggett s.n. (K!—
holotype).
Rhizome creeping with rather stiff loosely tufted fronds. Rhizome scales 3–5mm long,
0·5–0·75mm wide at the base, linear subulate with a rounded base and ending in a short
hair point, clathrate with smooth cell walls; median cells dark thick-walled and
compressed with narrow, sometimes obscured, lumens, becoming thinner walled and
pellucid towards the margins. Fronds up to 24–45cm long, pinnate to deeply
bipinnatifid; stipes black becoming green near apex, sparsely scaly. Lamina bright
green, linear lanceolate, 15–37 × 2·5–9·5cm with 10–17 pairs of opposite or
subopposite spreading pinnae. Pinnae 1·8–5·0 × 1·0–2·3cm, coriaceous, broadly trullate
to trullate, base broadly cuneate; basal pinnae deeply divided, upper pinnae less so, into
3–5 lobes; largest lobes (pinnules) cuneate-obovate and further cut into 3 or 4 segments
with truncate lacerate apices, smaller lobes rectangular with truncate lacerate-crenate
apices. Veins and sori subflabelliform. Spores plano-convex, (32–) 36–44 (–50) × (20–)
24–30 (–35) µm perispore costate with loosely anastomosing low ridges or folds.
Reproduction sexual, chromosome number n = 72 (2n = 144).
Ecology and distribution. Asplenium stuhlmannii is a fern of rocky outcrops often
in grassland at altitudes from 850 – 2188m. It grows in sheltered rock crevices or in
shade around the base of rocks or on top of rocks in the shade or less often in full sun.
It is widely distributed in East Africa: Tanzania, Rwanda, Uganda, Kenya, E Zaire,
Sudan and E Central African Republic (Figure 1).
Notes. The name A. stuhlmannii is here restricted to the East African material with
creeping rhizomes bearing scales made up of smooth walled cells, pinnate to deeply
bipinnatifid fronds with broadly trullate to trullate pinnae and spores with costate-alate
spores with rather low smooth folds or wings (Figure 3). All the material is tetraploid
in so far as can be judged from comparisons of spore measurements from specimens in
84
FERN GAZ. 18(3): 81-90. 2008
the herbaria with those from the cytologically authenticated material.
Asplenium stuhlmannii var laciniata with larger pinnae, more deeply dissected into
laciniate lobes probably represents a larger luxuriant shade form. The rhizome scales
and spore ornamentation and size match those of typical A. stuhlmannii.
Specimens examined. ZAIRE: Haut Zaire: Zande Dist.: entre Doruma et Niangoro,
1931, J. Lebrun 3180 (K, BM). SUDAN: Western Equatoria: Yambio Dist.: Mt
Bangenze, in rocky crevice, in shade on hillside, local, 900m, G. A.. Prowse 146 (BM);
ibid. on stone outcrop near Mt Bangenze, F. W. Andrews A1497 (BM); Rest House, on
granite outcrop near Borago, 1939, F. W. Andrews A1612 (BM); Mt Ledua near Ibba, J.
G. Myers 6570 (K); Baragu-marshes, 1939, J. W. G. Wyld 542 (BM). Maridi Dist..: Azza
Forest, in crevice of rocks in depression forest, tufted frond, local, 850m, 1953, Prowse
153 (BM); near Azza Forest, 1939, F. W. Andrews A1347 (BM). Eastern Equatoria: Yei
Dist.: Lado, Yei R., F. Sillitoe 110 (K); Kapigo, in pockets of earth on open rocky
hillside especially near bushes, common, 1273m, 1953, G. A. Prowse 33 (BM); Mt
Gumbiri, in Selaginella mats on open rock surface, common, 1500m, 1953, G. A.
Prowse 17 (BM); Mt Kala, rocky crevice and under Euphorbia sp., locally common,
1200m, 1953, G. A. Prowse 191 (BM); Imatong Mts, Talanga, 4° 01´ N, 32° 45´ E,
950m, 1980, I. Friis & K. Vollesen 577, 578 (K). UGANDA: Northern Prov.: West Nile
Dist.: Koboko Hill, damp crevices in granite rocks, 4400ft, R. J. Chancellor 111 (K);
Mt Otzi,, 500ft below summit, 5000 feet, fairly common, 1951, E. M. Cherry 5 (K,
BM). Acholi Dist.: Chua, fern on rocky outcrop at Ukuti, 1935, W. J. Eggeling 2411,
2413 (K); Lotuturu, Chua, Maxwell-Forbes 94 (PRE). Lango Dist.: Maruzi, ground fern
growing in interstices of rocks of Kibuji Hill, 1937, R. G. Sangster 321 (BM). Buganda
Prov.: Mubende Dist., Bugangadzi, grows on granitic rocks, 4200ft, Dawe 137 (K).
Mengo Dist., Entebbe, Loe, 1909, R. Fyffe 71 (K). Western Prov.: Toro Dist.: Kyaka,
growing on tree trunk, 4500ft, 1913, J. D. Snowden 150 (K), 1914 (BM); Mwenge,
Oruha Hill, fern in crack of gneiss rock, 1951, H. A. Osmaston 1221 (BM); Kyegegwa,
granitic outcrop, 4200ft, 1933, A. S. Thomas (K). Ankole Dist.: Igara, rocky outcrop in
1000 Km
Figure 2. Distributions of Asplenium stuhlmannii (z), A. jaundeense () and A.
mantoniae (S). The locality of A. jaundeense in Gabon not placed.
85
short grassland, 5500ft, 1939, Purseglove 577 (K); Nyakalakye-Buyenja, Kashari, 0°
22´ S, 30° 31´ E, growing in and around rock crevices, 1680m, 1986, P. K.
Rwaburindove 2235 (K). Eastern Prov.: Teso Dist.: Kyere, crevices among rocks
forming rocky outcrops, 3650ft, P. Chandler 971 (K). Busoga Dist..: Bulimba Banga,
hill N of Mpumuda, Kibili Rd, growing in shade under large round boulders on hill top,
1950, G. A. T. Wood Y2 (K). KENYA: Rift Valley Prov.: Trans-Nzoia Dist.: NE Elgon
foothills, Simpsons rocks, rock crevice in outcrop rock in savannah, 6200ft, Tweedie
1832 (K). Eldoret Dist.: Oldan Sapuk, c. 7000ft, 1951, P. J. Greenway 8538 (BM).
Yanza Prov.: North Kavirondo Dist.: S foothills of Elgon above Kimilili, growing in
rock crevices, 5900´, Tweedie 2921 (K). Uasin-Gishu Dist.: Oldan Sapuk, c. 7000ft,
1951, P. J. Greenway 8538 (K). Nandi Dist.: Kaimosi Farm, 5800ft, G. R. Williams 571
(K). Central Prov.: Fort Hall Dist..: Fort Hall, 1903, R. Meinertzhagen s.n. (BM).
Coastal Prov.: Teita Dist..: Teita Hills, Mwatata R. Valley 5.8km Mwatata-Wundanye
along new alignment, 870–1000m, R. B. Faden 70/443 (K). ? Masai Prov.: Masai Dist..:
Emoli Hill ft, 5900ft, Opiko 326 (K). TANZANIA: Lake Prov.: Mwanza Dist.: coast of
Speke Gulf near Mwanza, moist cracks and hollows below great granite boulders,
3740ft, 1931, B. D. Burtt 2474 (K, BM); Mwanga, fern common on rocky hills near
Lake Mwanga, 19385 (?8), P. E. Glover 303 (K). Ngara Dist.: Bugarama, Bushubi, on
rock face in crevice, 5000ft, 1961, R. Tanner 5829 (K). RWANDA: Biumba Territory,
Region de Mutara, env. de Mimuli, colline Nyakagenge, anfractuosités dans
affleurements granitiques, 1975, Troupin 3726 (K); ibid., creux ombragé sous roche
surplomb, 1975, Troupin 3267; ibid., affleurements rocheux, 1450 – 1500m, 1975,
Troupin 4212 (K).
Asplenium mantoniae A. Braithw. sp. nov.
Asplenium stuhlmannii et A. jaundeense similis sed differt chromosomate numero (n =
144) et sporis majoribus; a A. stuhlmannii pinnis trapezio-trullatis, basaleis acutis et
lobis paucioribus; a A. jaundeense lobatis profunde trapezioid-trullatis pinnis.
Holotypus: Nigeria, Ondo Province, insel-berg nr Ado Aiye, 7° 54´ N, 3° 30´ E, with
creeping rhizome growing under overhanging boulders, Hambler 513 (BM).
Asplenium stuhlmannii sensu Tardieu-Blot, in Mem. Inst. fr. Afr. noire 28: 190, t. 36,
fig. 3, 1953; Alston in The Ferns and Fern Allies of West Tropical Africa, suppl. 2nd
ed., Fl. W. Trop. Afr., 59, 1959.
A
B
C
Figure 3. Morphology of the Asplenium stuhlmannii complex, spores. A, A
stuhlmannii, Somalia; B, A. mantoniae, Hambler 513; C, A. jaundeense, Yaunde,
Cameroon. Scales: A – C = 10 µm.
86
FERN GAZ. 18(3): 81-90. 2008
Rhizome creeping with stiff tufted fronds; rhizome scales dark brown, up to 6mm long,
1mm wide, linear-subulate, clathrate, cell walls thick and opaque in the middle,
becoming thinner and pellucid towards the margin, walls facing lumens often
irregularly minutely denticulate (Figure 3). Fronds 15–44cm long, pinnate to deeply
pinnatifid; stipe dark purplish brown 4·5–20cm long with small scales similar to those
of the rhizome at the base; rachis dark purplish brown, for the most part becoming green
towards the apex, sparsely scaly. Lamina 10–24cm long, 3·5–7·5cm wide, narrowly
ovate with 8–13 pairs of subopposite spreading pinnae merging into a lobed apex.
Pinnae 2·0–6·5 × 0·75–2·5cm, narrowly parallelogram-shaped to trapezoid-trullate,
base narrowly cuneate then divided into 3–4 lobes merging into lacerate-serrate,
sometimes attenuate, acute apex (Figure 2); largest basal acroscopic lobe obtriangular
to rectangular and further cut into 3 lobes with irregularly crenate apices, remaining
lobes rectangular with irregularly crenate apices, becoming progressively smaller
towards apex of pinna. Veins and sori subflabellate. Spores plano-convex, (40–) 46 – 48
(–55) µm × (28–) 31–32 (–37) µm, costate-alate with much anastomosing high wavy
(undulating) ridges or wings. Reproduction sexual, chromosome number n = 144 (2n =
288).
Ecology and distribution. This new species is generally associated with rocky
outcrops, usually but not exclusively granite, e.g. the inselbergs of Nigeria, where it is
found in the shade on rock faces, in damp rock crevices or growing under overhanging
boulders from 100 – 400m. It has a predominantly West Africa distribution (Guinea,
Sierra Leone, Ivory Coast, Ghana, Nigeria, Cameroon) with outliers in the Central
African Republic (Figure 2).
Notes. The new species resembles A. stuhlmannii in its general growth habit, frond
architecture, colour and texture, but is distinguished by its slightly larger rhizome scales
made up of cells with thicker and often dentate cell walls, generally larger but less
divided and more trapezioid or parallelogram-shaped pinnae with narrower cuneate
bases and by its larger costate-alate spores with broader more frequently anastomosing
wavy (undulating) perispore folds or wings. In these respects it resembles the following
species, A. jaundeense, despite being traditionally confused with A. stuhlmannii. (see
Figures 2 & 3).
Etymology. Dedicated to the late Professor Irene Manton F. R. S. in recognition of
her contribution to knowledge of the cytology of the fern flora of West Tropical Africa.
specimens examined. GUINEA: Haut Niger (Fouta Djallon): Mt Bambaya, vers 100m,
1945, P. Jaeger 2113 (K); Macenta, Adams 5513 (P); Gueckedou, Adams 5581 (P).
SIERRA LEONE: Northern Prov., Bumba (?Bumban), 1200ft,1932, R. Glanville 436
(BM). IVORY COAST: Séguéla, rocher granitique a 15km E. sur la route de Beoul,
sous un surplombe humide, 1948, H. des Abbayes 619 (BM); entre Seguela et
Maukono, fissure rocheuse, 1954, R. Schell (K); Rocher d’Issia, in groove on granitic
rock, 250m, 1962, A. J. M. Leeuwenberg 4140 (K); Pays des Ayoles, sommet du Mont
Kouan prés Aanamé (manelog granitique), 400m, 1909, A. Chevalier 21269 (K,P); Mt
Dou, Man, Porteres s.n. (P); Mt Semelebou, A. Chevalier 22095 (P). GHANA:
Bhongo-Ahafo Prov.: Ebaw Rock near Mim, growing in moist crevices on granite
outcrop near thicket margin, 1973, Hall & Abbiw 44559 (K). NIGERIA: Oyo Prov.: hill
3 km N of Iseyin, 7° 58´ N, 3° 34´ E, 350m, rather shady crevice in granitic rocks, 196?,
J. B. Gillett 15399 (K); Okeho, group of low hills about 3 miles from Iseyin road, in
savannah country, 1959, R. W. J. Keay FHI 37748 (K); Ibadan, Oje Rock, sheltered rock
crevices near summit, 1967, D. P. M. Guile 3018 (K). Ondo Prov.: Erio near Aramoko-
87
Figure 3. Morphology of the Asplenium stuhlmannii complex showing silhouettes of
fresh juvenile fronds (top, scale = 5cm), pinnae from lower part of mature dried fronds
(middle, scale = 2 cm) and sections (middle to margin, scale = 0·1mm) of the rhizome
scales (bottom). A, A. stuhlmannii, Somalia; B, A. mantoniae, Hambler 513; C, A.
jaundeense, Yaunde, Cameroon.
88
FERN GAZ. 18(3): 81-90. 2008
Ekiti, base of Hildegardia on steep rock faces, 1968, J. B. Hall 67 (K); ibid., shade of
rock on inselberg, 1968, J. B. Hall 71 (K); Idanre, Jones s.n. (FHI 14846) (BM); among
granite boulders on the lower more or less forested slopes of granite hills, 1946, A. P.
D. Jones 14844 (BM); Idanre, behind Rest House, in rock fractures, texture coriaceous,
dark green, 1968, D. Gledhill 957 (K); inselberg NE of Ado Rock, 7° 51´ N, 3° 30´ E,
amongst rubble under overhanging ledge – small quantity only growing with Pellia
doniana, 1958, D. J. Hambler 419 (BM); inselberg near Tapa, 7° 34´ N, 3° 30´ E, under
overhanging boulders growing with Pellia doniana and Sanseviera sp., 1958, D. J.
Hambler 526 (BM). Plateau Province: near Farm Rua between Willsani Camp and
Marhai, soil filled crevices in shady side of rock, 1968, J. B. Hall 659 (K); Sha, 3600ft,
rock crevice in deep shade, scarp slope, dark green erect fern, D. W. Lawlor & J. B. Hall
570 (K). Taraba Prov.: Gembu, Mambilla Plateau, fern up to 25 cm high with grooved
rachis, found growing up between rocks on a wet rocky outcrop behind the Vet. Rest
House, c. 5500ft, 1973, H. M. Chapman 102 (K). CAMEROON: Nord Prov.: 7km S of
Poli, Mango, 1974, G. Fotius 2190 (K). CENTRAL AFRICAN REPUBLIC: BaminguiBangoran Prov., Manova-Gounda- St Floris National Park, WWF Inter. Elephant Cons.
Project, 2 km w of Camp Koumbala on Koumbala Creek, 8° 30´ N, 21° 12´ E, 570m
sandstone plants with some outcropping of sandstone ‘buttes’, under shaded overhang
in sandstone, 1983, J. M. Fay 5615 (K). Haut Kotto(Obangui-Chari): Wadda, sur les
rocheurs near Wadda, 1922, G. le Testu 4106 (BM).
A. jaundeense Hieron.
Engl. Bot. Jahrb. Syst. 46: 369, 1911; Tardieu. Mem. Inst. fr. Afr. noire 28: 192, t. 36,
fig. 4, 1953; type: Cameroon, an Felsen und Baustammen des Urwaldes bei Jaunde in
800 – 1000 Hohe ü. M., in dem Jahren 1890 – 1892, Zenker 214 (B!—lectotype,
selected here), 1894 – 1895, Zenker 214a (K!, BM!— isolectotypes); 23 Okt 1894,
Zenker & Staudt 526 (B! K!—paratypes); 31 Juli 1897, Zenker 1492 (B, K!—
paratypes); in lichten Waldungen an steinigen abhangen der Lagoo-Berge in 300 Hohe
ü. M., 25 Juni 1909, Ledermann 4396 (B–paratype).
Asplenium dimidiatum var. zenkeri Hieron., in Engler Veg. der Erde 9: 28, fig. 24,
1908.
Rhizome creeping giving rise to fronds a few millimetres apart; rhizome scales dark
brown, up to 6 × 1mm, subulate, clathrate with median cells dark thick-walled and
compressed, often with narrow occluded lumens, marginal cells thin-walled and
pellucid, walls facing lumens distinctly minutely dentate. Fronds up to 66 cm long;
stipes up to 26cm, matt black with scales at the base similar to those of the rhizome.
Lamina up to 41 × 8cm, oblong to linear-oblong with 10 – 20 opposite-subopposite
pairs of spreading pinnae merging progressively into a lobed apex; rachises similar to
stipe but glabrous and becoming green towards the apex. Pinnae up to 6 × 3cm,
trapezoid-rhomboid to cuneiform hastate, base narrowly cuneate, inaequilateral,
acroscopic side up to 1·5cm long, basiscopic side up to 3cm long; largest pinnae
subtrilobed to trilobed, lateral lobes truncate irregularly inciso-dentate, terminal lobe
long, elongate-deltoid, sparsely alternately inciso-dentate, ending in an acute point;
trilobing tending to disappear and pinnae becoming trapezoid-rhomboid towards apex
and in juvenile fronds. Costae not evident, veins and sori subflabellate. Spores planoconvex, (28–) 33–36 (–40) µm × (19–) 22–24 (–27) µm, perispore costate-alate with
anastomosing high wavy ridges or folds (wings). Reproduction sexual, chromosome
89
number n = 72 (2n = 144).
Ecology and distribution. The limited ecological information available suggests that
this species is associated with rocky outcrops or boulders at altitudes ranging from 300
– 1000m. It is known only from West Africa: Gabon (locality not placed), Cameroon
and eastern Nigeria (Figure 2).
Notes. Distinguished from Asplenium mantoniae by its trapezioid- rhomboid three
lobed pinnae with often long attenuate apical lobe and smaller spores, (28–) 33 – 36
(–40) µm × (19–) 22 – 24 (–27) µm.
The morphologically similar A. megalura Hieron. differs by its erect rhizome, rather
brittle wiry fronds with fewer and often long-attenuate pinnae, a large 3-lobed apical
pinna and spores with sparsely anastomosing shallow ridges or wings. It also differs
ecologically being usually epiphytic in forest and is widely distributed in tropical and
subtropical Africa.
Specimens examined. NIGERIA: Ogoja Prov.: Ikom and Obudu Divs, nr top of 2500ft
hill about pillar no. 30 on the E boundary of Boje enclave, Jones & Onochie s.n. (FHI
No. 18757) (BM). CAMEROON: Milbraed, III Reise nach Afrika 1913-14. Ubergangs
–und Kampfgebiet gegen die Savanna an der Nord-grenze der Hylea südlich des Sanaga
zwischen Yaunde und Dangdeng umwelt der Vereinigung von Lom (Sanaga) und
Djerem Etwa 68 Km N. O. Jaunde, Feb 1914, Sommer 8718 (K). GABON: Andoun,
rocheres de tete, Testu 238 (locality not placed) (BM).
DISCUSSION
The three species of the Asplenium stuhlmannii complex are united by the presence of
a creeping rhizome clothed in clathrate scales with dark median band and translucent
margins, and bearing rather stiff once- pinnate narrowly lanceolate or narrowly ovate
fronds. The rhizome scales, pinnae and spores of the three species are shown in Figures
2 & 3.
The clathrate rhizome scales of all three species are made up of dark thick-walled
cells with compressed lumens in the middle parts becoming thinner-walled and
translucent towards the margins. Apart from differences in the size of the cells related
to ploidy, there are subtle differences in the inner surface of the cell walls bordering the
lumens. Those of A. stuhlmanni are smooth while those of A. jaundeense are minutely
denticulate. Those of A. mantoniae are often, but not always, sparsely denticulate
(Figure 2), and thus somewhat intermediate.
The pinnae and spore ornamentation of A. mantoniae are also intermediate between
those of the two tetraploids. The pinnae possess a narrower acute base and are more
trapezoid-rhomboid and less dissected than the broadly trullate pinnae of A. stuhlmannii
and possess a broader cuneate base and are more trulllate with deeper lobes than the
trapezio-rhomboid almost entire pinnae of A. jaundeense. The spores of A. mantoniae
are larger than those of the two tetraploids on account of their higher level of
polyploidy. Their higher more anastomosing wavy perispore folds or wings contrast
with the rather low sparingly anastomosing straight perispore wings of A. stuhlmannii.
In this respect they resemble the spores of A. jaundeense.
The morphological analysis suggests a close relationship between the three
members of the complex and is consistent with the view that A. mantoniae has most
likely arisen by hybridisation between the two tetraploids and subsequent doubling of
the chromosome number of the hybrid to produce the octoploid. The present
90
FERN GAZ. 18(3): 81-90. 2008
distributions of A. jaundeense and A. stuhlmannii are widely separated in West and East
Africa respectively so that such a hypothesis would imply that in the past their
distributions must have either been sympatric or in close contact. The distribution of A.
mantoniae partly falls between those of the two tetraploids, but has also extended the
distribution of the complex particularly in West Africa where it may now be found as
far west as Sierra Leone and Guinea.
REFERENCES
ALSTON, A. H. G. 1959. The ferns and fern allies of West Tropical Africa. Suppl. to
ed. 2 of Flora of West Tropical Africa. Crown Agents, London.
BRAITHWAITE, A. F. 1964. A cytotaxonomic investigation on the Asplenium
aethiopicum complex in Africa. Unpublished Ph. D. Thesis, University of Leeds.
MANTON, I. 1950. Problems of cytology and evolution in the pteridophyta.
Cambridge University Press, Cambridge.
MANTON, I. 1959. Cytological information on the ferns of West Tropical Africa. In
ALSTON, A H. G. The ferns and fern allies of West Tropical Africa. Suppl to 2nd
ed. of Flora of West Tropical Africa, pp. 75–81. Crown Agents, London.
TARDIEU-BLOT, M. L. 1953. Les Pteridophytes de l’Afrique Intertropicale Francaise.
Mem. Inst. Fr. Afr. Noire 28: 1–241.
FERN GAZ. 18(3):91-97. 2008
91
ASPLENIUM DELAVAYI (PTERIDOPHYTA, ASPLENIACEAE)
A FERN SPECIES NEW TO NEPAL
ARUN RIJAL
P.O. Box 4326, Kathmandu, Nepal
(E-mail: [email protected])
Key words: Asplenium delavayi, Pteridophyta, west Nepal, disjunct
ABSTRACT
Asplenium delavayi (Franch.) Copel., a Sino-Himalayan species, described from
S.W. China and previously known only as far west as Sikkim, has been
discovered in the Baitadi District of west Nepal. This is much further west than
might be expected and emphasises the possibility of more species being
discovered in Nepal with increased collection and study.
INTRODUCTION
Although Nepal is only c. 885 km from west to east and 145 to 241 km from north to
south, it contains extreme differences in climatic and geographic situation, as a result of
the great range of altitude, from c. 60 m. in the southern terai zone to more than 8000
m. in the high Himalaya. Plant diversity ranges from stunted alpines surviving in harsh
environments in the frozen mountains, to mighty trees of the steamy lowland jungles.
This diverse environment provides habitats for a surprisingly large number of plant
species and more than 6000 species of flowering plants have been recorded so far from
Nepal. It has been estimated that over 6600 species will be listed from the country when
the poorly known remote regions are fully explored (RBGE 2007).
Nepal also has a rich fern-flora with 532 species of pteridophytes reported so far
(Thapa 2002; Fraser-Jenkins & Thapa in prep.). Nepalese pteridophytes belong to two
main phytogeographical elements, with a smaller third element. The dominant group of
species are Sino-Himalayan species, with their centre of diversity in S.W. China, which
have spread westwards along the Himalayan chain and are typically plants of the main
and inner ranges, occurring from mid to higher altitudes, though some may occur in the
lower, outer ranges nearer the plains. Such species usually occur more-or-less
throughout Nepal, though the number of species is higher in the higher rainfall areas of
the east. The second group of species are the S.E. Asian elements, which require more
tropical conditions and high rainfall. They occur predominantly in E. Nepal from lowermid to mid altitudes, and although they also extend westwards through Nepal, their
numbers decrease rapidly further westward, particularly towards far-west Nepal. There
is a small second enclave of S.E. Asian species in the locally high-rainfall area around
Pokhara in W.C. Nepal. The third element is a small group of either European or W.
Himalayan species, the latter a special part of the Sino-Himalayan group. These extend
eastwards from the W. Indo-Himalaya into the far-west part of Nepal, with several
occurring mainly behind the Himalayan line in C. Nepal.
Asplenium is one of the larger genera in the flora and contains 32-34 species and an
additional subspecies in Nepal (Thapa 2002, Fraser-Jenkins & Thapa in prep.) of which
19-21 species may be considered Sino-Himalayan elements, eight S.E. Asian elements
and five European elements (Fraser-Jenkins pers. comm., Dec. 2007). Most new
92
FERN GAZ. 18(3):91-97. 2008
records for Nepal concern what is presumably the richest part of the flora in eastern
Nepal. But the central region is by far the best collected due to its greater accessibility
and, as currently known, more species are recorded from central Nepal than further east.
This is likely to be an artificial anomaly resulting from the greater attentions of
collectors there. The least known area is western Nepal, especially the far west. There
is a higher proportion of semi-arid and European and W. Himalayan elements here, but
it is clear that fewer species occur in W. Nepal due to the lower rainfall. However
because the area is so under-collected some surprises occasionally turn up there, with
some species extending further west than might be expected.
In 1997, the author came across an unusual species in far W. Nepal which was not
familiar to him. It had the long, indusiate sori of an Asplenium, but a uniquely rounded,
entire and simple lamina of small size, and a black stipe. This was subsequently
identified in Dec. 2007 by C.R. Fraser-Jenkins as the very rare and little known species,
Asplenium delavayi (Franch.) Copel., which was not previously known from Nepal,
even in the east.
TAXONOMIC TREATMENT
Asplenium delavayi (Franch.) Copel., Genera Filicum: 165 (1947).
Basionym: Scolopendrium delavayi Franch., Bull. Soc. Bot. France, sér. 2, 32: 28
(1885); synonyms: Phyllitis delavayi (Franch.) C.Chr., Index Fil. 2: 492 (1906);
Schaffneria delavayi (Franch.) Tardieu, Naturaliste Malgache 9(1): 30 (1957);
Sinephropteris delavayi (Franch.) Mickel, Brittonia 28(3): 327 (1976).
Type: China, Yunnan: “in montibus, ad Chouang-che-teou, supra Ta-pin-tze, prope
Tali [now Dali]”. Abbé P.J.M. Delavay 67, 19 Aug. 1883 (P, K).
Distribution: W. Nepal, Sikkim, Bhutan, N.E. India (Manipur), N. Myanmar, S.W.
and S.C. China (Yunnan, Sichuan, Guizhou and Guangxi).
Asplenium delavayi was illustrated by Clarke (1888), who first reported it from
India, Ching (1930), Mickel (1976) and Wu (1999) and is probably related to the
similar, but larger, Chinese and Japanese species, A. cardiophyllum (Hance) Baker
(syn.: Boniniella cardiophylla (Baker) Tagawa). Although several splinter-genera,
including Sinephropteris, were raised at various times, these are now generally sunk
N
TIBET
Study area
INDIA
Figure 1: Map of Nepal showing the location of A. delavayi in Baitadi District.
RIJAL: ASPLENIUM DELAVAYI - A FERN SPECIES NEW TO NEPAL
93
into Asplenium as they all have a recognisably similar “asplenioid” morphology and are
connected by intermediate species to the main-stream of the genus.
Asplenium delavayi was collected by the author in Nepal at Nwaghargad, Baitadi
District, Far-West Development Region (Mahakali Zone); in crevices of damp rock in
open places, 1258 m. alt. (4125 ft.). A. Rijal 3251, 15 January 1997 (KATH). The
distribution of A. delavayi is that of a Sino-Himalayan species, though occurring at
lower altitude.
Description: Rhizome short, unbranched, up to c. 1cm. long, erect, bearing a tuft of
branched, dark-brown roots below, scales on the apex similar to those on the stipes but
slightly smaller, and a tuft of semi-erect, radiating stipes. Stipes cylindrical with an
adaxial groove, blackish-brown to black, glossy, 2.5-5cm. long, erect, bearing a tuft of
small, greyish, narrowly lanceolate, acuminate, slightly toothed scales towards their
bases, c. 2-3mm. long, 0.4-0.7mm. wide towards their bases. Lamina persistent in
Winter, simple, 2-2.6 x 2.1-2.6 cm., glabrous above, bearing a few scattered small
scales beneath, bright green, paler below, entire to slightly wavy at the margin,
markedly rounded-ovate with a shallow cordate base and sometimes a slightly roundedpointed apex, a vague darkish midrib present towards the base of the lamina, veins
immersed, inconspicuous, radiating from the midrib, several times dichotomously
forked, with the smallest forks towards the margin, but not reaching it, with occasional
anastomoses near to the margin. Sori c. 20-28 per frond, elongated, straight to slightly
curved, placed along the veins, not forked, obliquely radiating, starting near the midrib
and ending below the margin, larger main sori c. 1cm. long, occasional smaller sori
borne between the larger sori from about half way along their length, indusiate;
indusium pale, very thin, becoming brown, lifting and shrivelling somewhat on soral
ripening, opening towards the adaxial (midrib) side, Spores good, regular, bilateral,
with short wings of perispore, c. 28 x 20 µm.
The general habit of the fern is similar to a small, more delicate Adiantum reniforme
L. (from Macaronesia and S.W.China), but with the lamina not horizontal as in that
species and the sori elongated and radiating on the lower surface of the frond, not short
and around the edge. It is also similar to a very small Asplenium scolopendrium L.
(from N. America, Europe, Macaronesia, W. Asia, China and Japan), but with round,
not elongated ovate fronds and thin, black stipes.
CONCLUSION
The discovery of this rare species in far-west Nepal where it has been overlooked so far
is probably connected with that area being under-collected. The considerable
disjunction of this population from those further east is not likely to be an actual
botanical reality. It is also likely that further unrecorded species may be found if more
intensive collecting were made, particularly if carried out by specialists familiar with
all the Nepalese pteridophytes who could easily pick out any unfamiliar taxa. Some
other species also show similar disjunctions which may be expected to be only
temporary until further study has been made. At one time there was a considerable gap
in recording between Darjeeling/Sikkim and the W. Himalaya in Kumaon
(Uttarakhand) for the majority of species extending that far west. But since the
pteridophyte-flora of Nepal has become largely known over the latter half of the 20th
Century, to date, the “central Himalayan gap” has largely been filled and can be seen as
an artificial one. A few large disjunctions still remain, these are:
Acrophorus paleolulatus Pic.Serm. - gap from C. Nepal to Kumaon (Chamoli).
94
FERN GAZ. 18(3):91-97. 2008
2-3 mm
1 cm
Single scale
Soral arrangement
1 cm
Structure
Figure 2: Sketch of the general habit, a stipe-base scale and the abaxial surface of
the frond with sori.
95
Actiniopteris radiata (Sw.) Link - gap from E. Nepal to Lumaon (Garhwal).
Acystopteris tenuisecta (Blume) Tagawa - gap from E. Nepal to Kumaon (Nainital).
Anogramma reichsteinii Fras.-Jenk. - gap from C. Nepal (Chapagaon, Kathmandu;
locality perhaps doubtful) to Kumaon (Nainital).
Asplenium delavayi (Franch.) Copel. - gap from Sikkim to far-west Nepal (Baitadi).
Athyrium otophorum (Miq.) Koidz. - gap from N.E. India (Meghalaya) to Kumaon
(Pithoragarh).
Athyrium roseum Christ - China; Darjeeling (Fraser-Jenkins) and gap to W.C. Nepal
(below Annapurna Base-Camp. Fraser-Jenkins).
Cornopteris quadripinnatifida M.Kato - gap from C. Nepal to Kumaon (Chamoli).
Deparia lancea (Thunb.) Fras.-Jenk. - gap from C. Nepal (Sankhu) to ?Kumaon
(Pithoragarh. Punetha & Kholia).
Plate 1 & 2: Asplenium delavayi from Baitadi, general view.
96
FERN GAZ. 18(3):91-97. 2008
Dryopteris pulvinulifera (Bedd.) Kuntze - gap from E. Nepal to Kumaon (Nainital).
Lepisorus sublinearis (Baker ex Takeda) Ching - gap from W.C. Nepal (Pokhara) to
Kumaon (Nainital).
Lindsaea commixta Tagawa - gap from N.E. India (Assam) to W.C. Nepal (Andheri
Khola, N. of Tansen; locality sometimes given on label as Mussoorie in error).
Matteuccia intermedia C.Chr. - gap from W.C. Nepal (Annapurna Base Camp. FraserJenkins) to Kumaon (Pithoragarh).
Microlepia hallbergii (d’Almeida) C.Chr. - gap from C. Nepal (Sankhu) to Kumaon
(Garhwal).
Microsorum zippelii (Blume) Ching - gap from far E. Nepal to Kumaon (Chamoli).
Pteris kathmanduensis Fras.-Jenk., in prep. - gap from C. Nepal (Kathmandu) to
Kumaon (Pithoragarh).
Pteris vittata L. subsp. vermae Fras.-Jenk. - gap from China, Tibet and ?Bhutan
(Thimphu. Fraser-Jenkins) to Kumaon (Nainital).
Thelypteris (Christella) lebeufii (Baker) Panigrahi - gap from N.E. India (Assam State)
to Kumaon (Tanakpur. Fraser-Jenkins).
Trichomanes parvifolium (Baker) Copel. - gap from Myanmar to W.C. Nepal (Gorkha).
Trichomanes saxifragoides C.Presl - gap from Bhutan to W.C. Nepal (Pokhara).
It is to be hoped that future work, concentrating on Nepal, may continue to make the
full distribution patterns of Himalayan pteridophytes clear and other new records may
be discovered.
Plate 3: Single frond (abaxial surface) showing the arrangement of the sori.
97
ACKNOWLEDGEMENTS
The author wishes to thank Mr. C.R. Fraser-Jenkins, of Naya Bazaar, Kathmandu,
Research Associate of the Royal Botanic Garden Edinburgh, for helping in
identification of the collection, revision of this paper and providing information on
disjunct species.
REFERENCES
CHING, R.C. 1930. Icones Plantarum Sinicarum 1: t. 26.
CLARKE, C.B. 1888. On the Plants of Kohima and Muneypore. J. Linn. Soc. (Lond.),
Bot., 25: [1-] 93 [-101] et tt., t. 41.
MICKEL, J.T. 1976. Sinephropteris, a new genus of scolopendrioid ferns. Brittonia
28(3): 326-328.
ROYAL BOTANIC GARDEN EDINBURGH (RBGE) 2007. Plants of Nepal: Nepal
and its plants. Webpage of Royal Botanic Garden Edinburgh
(http://www.floraofnepal.org/, June 17 2007).
THAPA, N. 2002. Pteridophytes of Nepal. Department of Plant Resources (DPR),
Ministry of Forest and Soil Conservation (MFSC), Thapathali, Kathmandu, Nepal.
WU, S.H. (1999). Aspleniaceae, in Flora Reipublicae Popularis Sinicae etc. 4(2): [1-]
143-145 [-265].
98
FERN GAZ. 18(3). 2008
SHORT NOTE
CORRECTION TO A REVISED HANDBOOK TO THE FLORA OF
CEYLON, VOLUME XV, PARTS A & B, FERNS AND FERN-ALLIES,
2006. M.D. Dassanayake (General Editor), Monika Shaffer-Fehre (Editor).
ISBN, SET 1-57808-411-3
Monika Shaffer-Fehre, The Herbarium, Royal Botanic Gardens , Kew,
Richmond, Surrey
([email protected])
These volumes devoted to ferns and fern allies contribute to the Flora of Ceylon.
Unfortunately some errors have been identified.
The following corrections are required to the contents: the family Dryopteridaceae
(Vol. A) is treated in two chapters rather than one only, and Polypodiaceae begins
Volume B. Regarding the Woodsiaceae, the name of the senior author is missing in
‘Contents’; three of the four genera treated, viz.: Hypodematium, Deparia and Athyrium
have been described by C.R. Fraser-Jenkins and one, Diplazium, has been contributed
by Dr M. Zink. There is one further omission: Vol. B, page 356: PTERIDACEAE, B.
Verdcourt. The following text is missing from Hemionitis tomentosa at the very end of
the treatment:
Specimens Examined. KANDY DISTRICT: Above Kandy: Roseneath Valley, The
Heremitage, Sledge 1139 (BM, K); Abatage on Gampola-Pusselawa road, Faden &
Faden 76/246 (K); Peradeniya, Matthews s.n.; cultivated at Kew from Ceylon material
in 1954 (K).
The editor apologises for these mishaps, and a correct list of contents is given on the
next page. The editor would appreciate hearing from readers of any other mistakes. A
list of additional corrections will then be published at a later date.
99
FERN GAZ. 18(3). 2008
FOREWORD
ACKNOWLEDGEMENTS
INTRODUCTION AND SURVEY OF TAXA WITH REFERENCES AND
GENERAL LITERATURE by M. Shaffer-Fehre and P.J. Edwards
ii—-xx
KEY to the FERNS and FERN-ALLIES of SRI LANKA by P.J. Edwards xxi-xxix
VOLUME 15 PART A
ASPLENIACEAE by D.Philcox
AZOLLACEAE by Jennifer M. Ide
BLECHNACEAE by P. Jayasekara
CYATHEACEAE by D. Philcox
DAVALLIACEAE by H. P. Nooteboom
DENNSTAEDTIACEAE by M. J. Zink
DRYOPTERIDACEAE 1 by C. R. Fraser-Jenkins
DRYOPTERIDACEAE 2 by M. J. Zink
EQUISETACEAE by B. Wadhwa
GLEICHENIACEAE by Monika Shaffer-Fehre
GRAMMITIDACEAE by B. S. Parris
HYMENOPHYLLACEAE by P. Jayasekara
ISOETACEAE by B. Wadhwa
LOMARIOPSIDACEAE by M. J. Zink
LOXOGRAMMACEAE by M. G. Price
LYCOPODIACEAE by D. Philcox
MARATTIACEAE by D.M.U.B. Dhanasekara
MARSILEACEAE by Jennifer M. Ide
OLEANDRACEAE by B. Verdcourt
OPHIOGLOSSACEAE by D. Philcox
OSMUNDACEAE by P. Jayasekara
PARKERIACAE by B. Wadhwa
1—35
36—39
40—47
48—55
56—66
67—99
100-155
156-170
171-172
173-175
176-213
214 235
236-237
238-248
249-253
254-270
271-274
275-283
284-294
295-304
305-307
308-310
VOLUME 15 PART B
POLYPODIACEAE by P. H. Hovenkamp
PSILOTACEAE by B. Wadhwa
PTERIDACEAE by C. R. Fraser-Jenkins, B. Verdcourt & T. G. Walker
SALVINIACEAE by Jennifer M. Ide
SCHIZAEACEAE by D. Philcox
SELAGINELLACEAE by D. Philcox
THELYPTERIDACEAE by Monika Shaffer-Fehre
VITTARIACEAE by Monika Shaffer-Fehre
WOODSIACEAE by C.R. Fraser-Jenkins and M. J. Zink
New Names published in this Volume
List of endemic species
Selective Glossary of Terms
Index to Scientific Names
311-348
349-351
352-427
428-432
433-439
440-454
455-520
521-531
532-576
577
578-579
580-590
591-616
100
FERN GAZ. 18(3). 2008
SHORT NOTE
TWO NEW SPECIES OF SELAGINELLA SUBGENUS HETEROSTACHYS
(SELAGINELLACEAE) FROM THE GUIANAS - A CORRECTION
In the recent paper by Cremers & Boudrie (2007), the two names are ascribed to
Valdespino ex Cremers & Boudrie. However, in referring to the descriptions, Cremers
& Boudrie state that “what follows” was extracted verbatim from Valdespino’s
monograph. As both the names and the descriptions are therefore ascribed to
Valdespino, under Art. 46.2 the correct attribution of the names should be:
Selaginella gynostachya Valdespino in Fern Gaz. 18: 42[-46].
Selaginella karowtipuensis Valdespino in Fern Gaz. 18: 46[-51]. 2007.
2007
I thank Professor John McNeill for advice.
CREMERS, G. & BOUDRIE, M. 2007. Two new species of Selaginella sugbenus
Heterostachys (Selaginellaceae) from the Guianas. Fern Gazette 18(2):41-52.
Mary Gibby
FERN GAZ. 18(3): 101-109. 2008
101
MORPHOTYPE AND CONFLICTING TAXONOMIES IN PTERIDIUM
(DENNSTAEDTIACEAE: PTERIDOPHYTA)
JOHN A. THOMSON
National Herbarium of NSW, Botanic Gardens Trust, Mrs Macquaries Road, Sydney,
NSW 2000, Australia.
Key words: Dennstaedtiaceae, bracken, genetics, morphology, morphotype,
polymorphism, Pteridium, subspecies, taxonomy.
ABSTRACT
Conflicting taxonomic treatments of the diploid bracken ferns of the Laurasian
lineage within Pteridium are briefly reviewed. The infraspecific entities
recognised in this section of the genus can be separated on morphological,
phenological and physiological criteria into two main groups referred to
respectively as ‘aquilinum’ and ‘latiusculum’ morphotypes. Evidence of gene
flow between morphotypes is discussed, including the finding from DNA studies
that taxa of the ‘aquilinum’ and ‘latiusculum’ morphotypes in each major
geographic region are more closely related to the contrasting morphotype from
the same region than to the corresponding morphotype from another region. It is
proposed here that the ‘aquilinum’ and ‘latiusculum’ morphotypes may be
regarded as mediated by a simple genetic polymorphism involving alternative
pleiotropic alleles at a single major morphogenetic locus comparable to those
previously described in Athyrium and postulated in Asplenium and Pteridium,
but other possible models are discussed.
It is concluded that P. aquilinum is best treated as a single species containing at
present 11 subspecies.
INTRODUCTION
A vigorous ongoing polemic continues to cloud taxonomic treatment of the diploid (2n
= 104; Wolf et al., 1987,1988) bracken ferns [Pteridium aquilinum (L.) Kuhn]
comprising a Laurasian lineage within the genus. Thomson and colleagues (Thomson,
2004; Thomson et al., 2005; Thomson et al., 2008; contra Thomson, 2000) regard P.
aquilinum as a grouping of 11 subspecies (Table 1) that in general correspond to
varieties within Tryon’s (1941) P. aquilinum subsp. aquilinum. In contrast, certain other
contemporary classifications are based on separation of these taxa into two phenotypic
groups designated here as the ‘aquilinum’ and ‘latiusculum’ morphotypes (Table 1).
Page (1997) and coauthors (Page & Mill, 1995; Gureyeva & Page 2005) regard these
‘aquilinum’ and ‘latiusculum’ morphotypic groups as multi-species complexes
represented in Europe by P. aquilinum and by P. pinetorum C.N. Page & R.R. Mill
respectively, each with several subspecies. Fraser-Jenkins (1997), on the other hand,
recognises P. latiusculum (Desv.) Hieron. ex R.E.Fr. at species level and accords
subspecific rank to P. latiusculum subsp. pinetorum (C.N. Page & R.R. Mill)
Fraser-Jenk. Stace (1997) and Karlsson (2000) group the European ‘aquilinum’
phenotypes as P. aquilinum subsp. aquilinum and those of ‘latiusculum’ as P. aquilinum
subsp. latiusculum (Desv.) Hultén. Shorina & Perestoronina (2000) distinguish the
European ‘latiusculum’ morphotype as P. aquilinum var. pinetorum (C.N. Page & R.R.
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FERN GAZ. 18(3): 101-109. 2008
Table 1: Geographic distribution and current subspecific nomenclature of northern
hemisphere diploid (2n = 104) bracken taxa indicating ‘aquilinum’ or ‘latiusculum’
morphotype, together with Tryon’s (1941) varietal names for reference.
Geographic region/
morphotype
Europe
‘aquilinum’
‘latiusculum’
Asia
‘aquilinum’
‘latiusculum’
Africa
‘aquilinum’
‘latiusculum’
North America
‘latiusculum’
Hawaiian Islands
‘intermediate’
P. aquilinum subspecies
P. aquilinum variety
aquilinum
pinetorum (C.N. Page & R.R.
Mill) J.A. Thomson1,2
aquilinum
latiusculum (Desv.) Underw.
ex Heller
wightianum (Wall. ex J.
Agardh) Shieh3
japonicum (Nakai) A. Löve &
D. Löve1
wightianum (J. Agardh) R.
Tryon
capense (Thunb.) C. Chr.4
centrali-africanum Hieron. ex
R.E. Fr.4
aquilinum
feei (W. Schaffn. ex Fée) J.A.
Thomson, Mickel & K.
Mehltreter5
feei (W.Schaffn. ex Fée)
Maxon ex Yuncker
latiusculum (Desv.) Hultén
auct. Thomson (2004)1
pseudocaudatum (Clute)
Hultén5
ex Heller
pubescens (Underw.) J.A.
Thomson, Mickel& K.
Mehltreter5
pubescens Underw.
decompositum (Gaudich.)
Lamoureux ex J.A. Thomson1
decompositum (Gaudich.) R.
Tryon
ex Heller
africanum Bonap.
pseudocaudatum (Clute)
Heller
1
Thomson, 2004
2
Also referred to as P. aquilinum subsp. latiusculum (Desv.) Hultén; P. aquilinum
var. pinetorum (C.N. Page & R.R. Mill) Perestor.; P. pinetorum C.N. Page &
R.R. Mill; P. latiusculum subsp. pinetorum (C.N. Page & R.R. Mill) Fraser-Jenk.
3
Also referred to as P. revolutum (Blume) Nakai
4
Thomson et al., 2005
5
Thomson et al., 2008
THOMSON: CONFLICTING TAXONOMIES IN PTERIDIUM
103
Mill) Perestor.
A dearth of useful discriminatory characters often leads to difficulties in
pteridophyte taxonomy, especially in the case of low order taxa that may be separated
on as few as one single character (Perrie & Brownsey, 2005). In many fern species the
segmental units represented at successive levels by frond, pinna, pinnule, and even
pinnulet, form a repetitive hierarchical series for which a feature expressed at one level
may not be independent of a corresponding feature scored at another segmental level.
In general, such attributes should be treated as relating to a single character, and may
well share a common genetic basis. Examples are evident in the morphometric
documentation of the relative number, size, shape and spacing of pinnae, pinnules and
pinnulets in subspecies of Pteridium aquilinum (L.) Kuhn (Thomson et. al., 2005;
Thomson et. al., 2008). Further, many commonly used features of indumentum, false
indusium and laminal segmentation are extremely labile, being affected by age,
shading, water-logging and other environmental conditions (Ashcroft & Sheffield,
1999; Karlsson, 2000; Thomson et al., 2005; Thomson et al., 2008) and must be
employed quantitatively and with caution.
Three recent re-evaluations of disputed taxonomic treatments in ferns highlight the
contribution that genetic evidence, either directly from breeding experiments or
indirectly from molecular data, may make to more consistent and objective assignment
of taxonomic rank. In Athyrium distentifolium Tausch ex Opiz (Woodsiaceae),
morphotype ‘flexile’ is endemic to Scotland and is found only at localities where
‘distentifolium’ is also present (McHaffie et al., 2001). Sporophytes of the ‘flexile’
phenotype are distinguished from ‘distentifolium’ by frond morphology, position of sori
and response to nutrients. The two phenotypes are controlled by alleles of a single major
gene with pleiotropic effects expressed in both gametophyte and sporophyte, the
‘flexile’ morphotype being recessive to ‘distentifolium’. In one natural population the
frequency of the recessive allele was about 0.4. The two morphotypes were originally
described as separate species but McHaffie and colleagues recommend a taxonomic
status not higher than variety.
In Asplenium hookerianum Colenso (Aspleniaceae), morphotype ‘hookerianum’
occurs in Australia and New Zealand and is distinguished from the endemic New
Zealand morphotype ‘colensoi’ which has consistently narrower pinnules. Stands in
New Zealand commonly comprise morphotype ‘hookerianum’ alone or co-occurring
with ‘colensoi’: ‘colensoi’ by itself is rare (Perrie & Brownsey, 2005). The two forms
were originally described as separate species. Neither nuclear genomic fingerprinting
by AFLP nor the chloroplast sequence results separated the specimens according to
morphotype. Geographic source accounts for much more of the total DNA-sequence
variation than pinnule morphology in both the AFLP data (29% of genetic variation
versus 0%) and the chloroplast haplotype data (58% of genetic variation versus 3%;
Perrie & Brownsey, 2005). Plants of contrasting morphotype originating from close
sites are more similar to each other genetically than to plants of the their own
morphotype from distant sites. The phenotypic stability of plants of the two
morphotypes even when growing intertwined in the field argues against a major role for
environmental variables, and ‘colensoi’ is purported to breed true from spores (Perrie &
Brownsey, 2005). It appears probable that morphotypic variation in Asplenium, like that
in Athyrium, results from a simple genetic polymorphism based on allelic variation at a
single locus. Perrie & Brownsey (2005) conclude that there is no significant genetic
discontinuity between the two morphotypes, and no justification for separating them at
104
FERN GAZ. 18(3): 101-109. 2008
higher than varietal or form rank.
Within P. aquilinum in North America, the status of the two eastern taxa now
designated subsp. latiusculum and subsp. pseudocaudatum (Table 1) proved
contentious until detailed analyses of morphology and isozyme complements were
made by Speer & Hilu (1999) and Speer et al. (1999). These taxa are clearly distinct on
morphological and distributional grounds (Speer & Hilu, 1999; Thomson et al., 2008).
Over their main east-coast range, ‘latiusculum’ is more abundant in the northern states,
‘pseudocaudatum’ in the south (Speer & Hilu, 1999), with only very few narrow zones
of overlap where intermediates suggesting introgression are seen. Detailed isozyme
analyses showed unrestricted gene flow between the two morphotypes at one such zone
of co-occurrence (Speer et al., 1999). Speer and colleagues postulate that the
‘latiusculum’ and ‘pseudocaudatum’ phenotypes may be controlled by alternative
alleles at a single genetic locus with pleiotropic effects. If the ‘latiusculum’ phenotype
is dominant and determined by an allele with high frequency in the north of its range,
while the recessive allele for the ‘pseudocaudatum’ phenotype is at high frequency or
fixed in the south, the roughly equal frequencies of the two forms observed in the North
Carolina piedmont would correspond to a frequency of about 0.7 for the recessive allele
(Speer et al., 1999). Strong selective pressure in relation to environmental features and
habitat preference are indicated, with a taxonomic ranking of subspecies for the two
morphotypes (Thomson et al., 2008).
CHARACTERISATION OF GENERALISED ‘AQUILINUM’ AND
‘LATIUSCULUM’ MORPHOTYPES IN PTERIDIUM
Characters that reflect the relative number, relative size, spacing and shape of laminal
segments at blade, pinna, pinnule and pinnulet levels (Thomson, 2000; Thomson et al.,
2005; Thomson et al., 2008), are particularly effective in discriminating a
‘latiusculum’-like group of taxa from an ‘aquilinum’-like phenotypic group.
The ‘aquilinum’ morphotype: Blade long, ovate to linear, typically 2-3 times the
length of the longest pinna which is generally one of the third to the fifth pair from the
lamina base towards the tip. Pinnae, pinnules (and less markedly pinnulets), relatively
long and narrow; typically 13-18 pinnules in the basal half of the longest pinna. Pinnae
and pinnules commonly inserted on rachis and costa at 70-90o. Frond expansion
gradual, progressing from base to apex. Basal 1-2 pinna pairs often marcescent before
expansion of the distal pinnae is complete. Frond axes relatively thick and fibrous;
dense stands leave heavy, often partially upright, litter in winter that is slow to collapse
next season.
The ‘latiusculum’ morphotype: Blade short, broadly triangular, typically 1-1.5 times
the length of the longest pinna which is generally one of the first or second pair from
the base towards the tip. Pinnae, pinnules (and less markedly, pinnulets) relatively short
and broad; typically 6-9 pinnules in the basal half of the longest pinna. Basal segment(s)
of each order sometimes much reduced, even vestigial. Pinnae and pinnules commonly
inserted on rachis and costa at 45-70o. Frond expansion rapid, often almost
synchronous, although progressive from base to apex. Basal pinna pairs not marcescent
before frond expansion is complete. Frond axes relatively thin, not heavily fibrous;
dense stands leave thin litter in winter that collapses readily next season.
The ovate to linear form of the expanded blade and pinna in ‘aquilinum’
morphotypes results from both the shorter length of the lowest pinna and pinnule pair(s)
compared with those placed more centrally and the relatively higher number of these
105
segments on the frond axis and costa respectively. The broadly triangular frond blade
and pinna of the ‘latiusculum’ morphotype could be envisaged as derived from the
‘aquilinum’ type by loss of pinna and pinnule pairs from the base of blade and pinna
upwards, a process which would at the same time increase the apparent length, relative
to more distal segments, of the pinnae/pinnules which thus become the basal or near
basal components. Overall the ‘latiusculum’ morphotype appears to reflect adaptation
to conditions providing a short growing season.
Where zones of contact or parapatry occur locally, as in Europe (Rumsey et al.,
1991; Karlsson, 2000), NE Asia (Tryon, 1941), and Africa (Verdcourt, 2000), recurrent
field observations of morphological intermediates provide evidence of recent, probably
ongoing, interbreeding between ‘aquilinum’ and ‘latiusculum’ morphotypes.
Multi-locus isozyme marker studies also reveal gene flow between multiple genets
(Bridges et al., 1998) of subsp. pinetorum in Scotland and local populations of subsp.
aquilinum (Rumsey et al., 1991).
DNA fingerprinting using the Arbitrarily Primed Polymerase Chain Reaction (APPCR) and Inter Simple Sequence Repeat (ISSR) procedures show that genomic
similarity is higher between ‘aquilinum’ and ‘latiusculum’ morphotypes from the same
geographic region than either is with morphotypes of its own group from other regions
(Thomson et al., 2005; Thomson et al., 2008). Thus subspp. capense and centraliafricanum from sub-Saharan Africa are more similar genomically than capense is to
aquilinum of Europe or centrali-africanum is to the N American latiusculum (Thomson
et al., 2005; Thomson et al., 2008). Similarly, subspp. pinetorum and aquilinum of
Europe share more genomic markers than pinetorum shares with latiusculum of North
America (Thomson, 2000; Thomson et al., 2005). Chloroplast genome haplotype for
the rps4-trnS region based on presence or absence of one or both of two 5-base direct
repeats is not concordant with the ‘aquilinum’ or ‘latiusculum morphotypic groupings
(Thomson et al., 2005; Thomson et al., 2008). For example, the haplotypes (Type A or
B, Thomson et al., 2005) of three geographically contiguous ‘latiusculum’ and
‘aquilinum’ morphotype pairs are: subspp. pinetorum/aquilinum (Europe) respectively
Types A and B, subspp. centrali-africanum/capense (Africa) both Type B, and subspp.
japonicum/wightianum (NE Asia) both Type A.
These findings provide strong evidence against a taxonomy based on separation of
‘latiusculum’ and ‘aquilinum’ clades (Thomson et al., 2008), and in particular argue
against recognition of ‘latiusculum’ or ‘aquilinum’ as reproductively-separated
groupings that individually merit full species status. Within Pteridium aquilinum as
recognised here (Table 1), reticulate phylogenetic relationships overlie evolutionary
trends presumably based on diversification and local adaptation.
IS THE ‘AQUILINUM’ VERSUS ‘LATIUSCULUM’ DISTINCTION A SIMPLE
GENETIC POLYMORPHISM?
Descriptively, the morphological polymorphisms analysed in Athyrium (McHaffie et
al., 2001) and postulated in both Asplenium (Perrie & Brownsey, 2005) and Pteridium
(Speer et al., 1999) behave as if based on variant alleles of major genes, with allele
frequencies being maintained by balancing selection pressures in particular
environments and/or differentially in the gametophyte or sporophyte stage. The
contention that the ‘aquilinum’ versus ‘latiusculum’ contrast also represents a genetic
polymorphism is supported by distributional data (Table 1), the genetic relationships of
the contrasting morphotypes in areas of overlap, and evidence that laminal segment
106
FERN GAZ. 18(3): 101-109. 2008
morphology at frond, pinna, pinnule and even pinnulet levels is hierarchically repetitive
and likely to be under control of a common gene or genes with alternative, apparently
pleiotropic, alleles. Definitive characterisation of the genetic basis of morphotype in
bracken must await integrated breeding and molecular analyses. The loci involved are,
however, clearly distinct from those responsible for the small-scale morphological
variants often collected and cultivated by growers (see, for example, Andersson, 1927)
that are rare in natural populations, with very low allele frequencies maintained by
recurrent mutation opposing negative selection (McHaffie et al., 2001).
While for descriptive purposes morphotypic variation in the ferns discussed here
may be adequately treated in terms of single locus polymorphisms with pleiotropy,
functional and structural molecular analyses of the loci responsible are now required to
reveal how the underlying developmental process are controlled. In general, the more
numerous and diverse the multiple phenotypic effects involving disparate
morphological and physiological characteristics that appear to be under single gene
control, the more an explanation invoking pleiotropy becomes questionable. In the
present cases we therefore need to consider other possible models of gene structure such
as that exemplified by the complex S locus responsible for self-incompatibility
(SI)/compatibility (SC) in a number of families of flowering plants such as
Brassicaceae. This super-locus provides co-ordinated unitary genetic control over
multiple aspects of SI mediated by tightly linked co-adapted sequence segments
collectively forming an S haplotype (Charlesworth et al., 2005). Multiple
developmental, cellular, physiological and biochemical processes in different S
phenotypes are mediated by the alternative S haplotypes represented in polymorphic
populations or in different taxa (Bechsgaard et al., 2006; Sherman-Broyles et al., 2007).
The several features characterising each S phenotype are controlled by distinct
sequence elements and do not involve pleiotropy. Where aspects of phenotype are
controlled by a complex super-locus or through a stepwise multigene pathway, multiple
mutations may appear as a single event if a mutation acting at an early step blocks
expression of accumulated changes effective later in the pathway. If features
distinguishing allopatric morphotypes are subject to selection, the allelic variants
responsible will be expected to change in frequency more rapidly than ‘neutral’
genomic elements. Phenotypic features under positive selection are likely to become
fixed faster than other genomic segments so that the probability of finding DNA
sequence markers correlating with morphotype is expected to be low, increasing with
the time of separation of the contrasting forms. In Asplenium, Perrie & Brownsey
(2005) found that neither AFLP fingerprinting of the nuclear genome nor chloroplast
sequence data grouped the plants analysed according to their morphotype. In Pteridium,
Thomson (2000) identified markers detected by AP-PCR that are specific to subsp.
wightianum (as P. revolutum) and other markers that are specific to subsp. capense
[misnamed by Thomson (2000) as P. aquilinum var, africanum; see Thomson et al.,
2005). These major markers are in both cases restricted in occurrence to geographically
localised taxa here regarded as of subspecific status, and neither is shared by other
‘aquilinum’ morphotypes. Distinctive alleles, supergene haplotypes and/or modified
dominance relations may account for the ‘intermediate’ morphotype of subsp
decompositum (Table 1) in which genomic elements of both subsp. japonicum
(‘latiusculum’) and subsp. wightianum (‘aquilinum’) are present (Thomson, 2000).
107
CONCLUSIONS
In most taxonomic investigations of ferns, especially where multiple taxa must be
considered, controlled breeding experiments of the kind so admirably applied in
Athyrium (McHaffie et al., 2001) are not practicable. More widely feasible are genomic
comparisons based on methodologies such as AP-PCR (RAPD), AFLP, ISSR, or
microsatellite analyses. If plants of taxon A from locality 1 are more closely related
genetically to plants of taxon B from locality 1 than to taxon A plants from localities
2...n, and/or a comparable situation holds for the relationships of taxon B, there is prima
facie evidence of recent and/or ongoing gene flow between the taxa that is likely to be
inconsistent with the degree of reproductive isolation normally associated with full
specific status. Rather, such a situation strongly suggests that the taxa concerned should
be accorded infraspecific rank, for example as subspecies, varieties, or forms (Perrie &
Brownsey, 2005; Thomson et al., 2008).
The work of McHaffie and colleagues on the ‘distentifolium’/‘flexile’
polymorphism in Athyrium represents an extraordinary contribution to our
understanding of fern morphogenetics by showing how morphological and
physiological studies of the gametophyte as well as the sporophyte may reveal at these
two life stages ‘antagonistic pleiotropy’ (Shaw, 2001) that determines allele frequencies
in natural populations through otherwise unsuspected opposing positive and negative
selective pressures. However, as pointed out above, questions remain open regarding
the structure of the genetic loci involved in the control of fern morphotype, and thus on
the strict applicability of the term pleiotropy in these cases.
P. aquilinum appears at present to be best treated as a single species containing 11
subspecies as listed in Table 1. Further morphometric and genetic studies are needed to
clarify the status of a number of named taxa described from many parts of Eurasia
which on present evidence should, if named, be ascribed varietal, form or ecotype
rankings (Stace, 1997).
ACKNOWLEDGEMENTS
I thank Elizabeth Brown (NSW) and Barbara Parris (Fern Research Foundation,
Kerikeri) for helpful discussion and comments. Miguel Garcia (NSW) provided
generous library assistance. I am indebted to Paul Wolf (Utah State University) for a
constructive critical review leading to substantial improvement of the final manuscript.
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inheritance. Hereditas 9: 157-179.
ASHCROFT, C.J. & SHEFFIELD, E. 1999 Rejection of Pteridium aquilinum
subspecies atlanticum (C. N. Page). Bot. J. Linn. Soc. 130: 157-170.
BECHSGAARD, J.S., CASTRIC, V., CHARLESWORTH, D., VEKEMANS, X. &
SCHIERUP, M.H. 2006. The transition to self-compatibility in Arabidopsis thaliana
and evolution within S-Haplotypes over 10 Myr. Mol. Biol. Evol. 23: 1741-1750.
BRIDGES, K.M., ASHCROFT, C.J. & SHEFFIELD, E. 1998. Population analysis of
the type localities of some recently recognised taxa of British Pteridium
(Dennstaedtiaceae: Pteridophyta). Fern Gaz. 15 :205-213.
CHARLESWORTH, D., VEKEMANS, X., CASTRIC, V. & GLÉMIN, S. 2005. Plant
self-incompatibility systems: an evolutionary perspective. New Phytol. 168: 61-69.
FRASER-JENKINS, C. R. 1997. New species syndrome in Indian pteridology and the
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ferns of Nepal. International Book Distributors, Dehra Dun.
GUREYEVA, I.I. & PAGE, C.N. 2005. Towards the problem of bracken taxonomy in
Siberia. Taxonomy Notes of the Krylov Herbarium of Tomsk State University 95:
18-26.
KARLSSON, T. 2000. Pteridium. In: JONSELL, B. (Ed.) Flora Nordica, vol. 1, pp.
43-47. Royal Swedish Academy of Sciences, Stockholm.
MCHAFFIE, H.S., LEGG, C.J. & ENNOS, R.A. 2001. A single gene with pleiotropic
effects accounts for the Scottish endemic taxon Athyrium distentifolium var. flexile.
New Phytol. 152: 491-500.
PAGE, C.N. 1997. The ferns of Great Britain and Ireland, 2nd ed. Cambridge
University Press, Cambridge.
PAGE, C.N. & MILL, R.R. (1995) The taxa of Scottish bracken in a European
perspective. Bot. J. Scotl. 47: 229-247.
PERRIE, L.R., & BROWNSEY, P.J. 2005. Genetic variation is not concordant with
morphological variation in the fern Asplenium hookerianum sensu lato
(Aspleniaceae). Amer. J. Bot. 92: 1559-1564.
RUMSEY, F.J., SHEFFIELD, E. & HAUFLER, C.H. 1991. A re-assessment of
Pteridium aquilinum (L.) Kuhn in Britain. Watsonia 18: 297-301.
SHAW, A.J. 2001. Antagonistic pleiotropy and the evolution of alternate generations.
New Phytol. 152: 365-368.
SHERMAN-BROYLES, S., BOGGS, N., FARKAS, A., LIU, P., VREBALOV, J.,
NASRALLAH, M.E. & NASRALLAH, J.B. 2007. S Locus genes and the evolution
of self-fertility in Arabidopsis thaliana. Plant Cell 19: 94-106.
SHORINA, N.I. & PERESTORONINA, O.N. 2000. Taxonomic studies of Russian
bracken I. Taxonomy of Pteridium in territories of European Russia, Crimea and
Caucasus. In: TAYLOR, J.A. & SMITH, R.T. (Eds.) Bracken fern: toxicity, biology
and control, pp. 48-51. International Bracken Group, Aberystwyth.
SPEER, W.D. & HILU, K.W. 1999. Relationships between two infraspecific taxa of
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305-312.
SPEER, W.D., WERTH, C.R. & HILU, K.W. 1999. Relationships between two
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Syst. Bot. 23: 313-325.
STACE, C.A. 1997. Field flora of the British Isles, 2nd ed. Cambridge University
Press, Cambridge.
THOMSON, J.A. 2000. Morphological and genomic diversity in the genus Pteridium
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THOMSON, J.A. 2004. Towards a taxonomic revision of Pteridium
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THOMSON, J.A., CHIKUNI, A.C. & MCMASTER, C.S. 2005. The taxonomic status
and relationships of bracken ferns (Pteridium: Dennstaedtiaceae) from sub-Saharan
Africa. Bot. J. Linn. Soc. 148: 311-321.
THOMSON, J.A., MICKEL, J.T. & MEHLTRETER, K. 2008. Taxonomic status and
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Central and North America. Bot. J. Linn. Soc. 157: 1-17.
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VERDCOURT, B. 2000. Pteridium. In: BEENTJE, H.J. (Ed.) Flora of tropical East
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WOLF, P.G., HAUFLER, C.H. & SHEFFIELD, E. 1987. Electrophoretic evidence for
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WOLF, P.G., HAUFLER, C.H. & SHEFFIELD, E. 1988. Electrophoretic variation and
mating system of the clonal weed Pteridium aquilinum (L.) Kuhn (bracken).
Evolution 42: 1350-1355.
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BOOK REVIEW
THE LIVERWORTS, MOSSES AND FERNS OF EUROPE, BY
WOLFGANG FREY (HORNWORTS AND LIVERWORTS),
JAN-PETER FRAHM (MOSSES), EBERHARD FISCHER AND
WOLFRAM LOBIN (FERNS AND FERN-ALLIES). Translated and
updated by the authors with illustrations prepared by H. Lunser
(Bryophytes) and E. Fischer (Ferns). Harley Books. English edition revised
and edited by T.L.Blockeel .2006. 512pp.( ferns and former fern allies pp.363455), 166 figs, each incorporating numerous line drawings (ferns and former
fern allies figs. 121-166). 218 X 155mm, hardback. ISBN 0 946589 70 4. £45.
German edition first published 1995.
Wide ranging titles such as this usually mean a book will be a ‘Jack of all subjects’ and
master of none, so you can imagine the pleasant surprise I had on dipping into it. Ferns
are indeed a relatively minor part filling fewer than 100 pages but the coverage is more
comprehensive than given in many field handbooks.
Apart from a couple of prefaces there are no introductory chapters. The books start
straight away with the dichotomous keys. At the end there is a glossary and
bibliography.
None but the most well read specialist will be familiar with all the taxa described here.
Recently discovered species and subspecies are included. There are no lengthy field
notes, details are precise and to the point. Most species are covered with fewer than four
lines of text. Distribution is given by country or island within Europe and more briefly
outside of Europe. As with the rest of the book the entire fern section is a generously
annotated dichotomous key. The key is supported by many line drawings which work
well for the well defined species but by the time you get down to the really critical taxa
they are not so helpful. I looked long and hard at one sketch of the base of the
lowermost pinna of Gymnocarpium robertianum but I still cannot work it out!
The fern tourist venturing anywhere in Europe, be it the Azores, Crete, Cape Verde,
Madiera, Russia or closer to home will find all the local fern species covered. A few
hybrids are also mentioned although the criteria for their inclusion is obscure to me. For
example the very rare Aspelnium x heufleri is in the key but many others, such as A. x
sarniense, are not.
I can think of no other up to date book covering all the ferns of Europe. Remy Prelli`s
Les fougeres et plantes allies de France et d’Europe occidentale has a more restricted
range. Where the two overlap I suggest the Prelli book will be the more user friendly
even though it is in French. For full coverage of Europe in one go, however, this guide
is a useful volume.
Martin Rickard
111
FERN GAZ. 18(3). 2008
BOOK REVIEW
BIOLOGY AND EVOLUTION OF FERNS AND LYCOPHYTES. Tom A.
Ranker & Christopher H. Haufler (eds). 2008. 480pp., 66 line figures 45
halftones. ISBN 978-0521-87411-3 hardback; ISBN 978-0521-69689-0
paperback. Cambridge University Press. £75.00, £35.00.
The editors of this new book have taken up the challenge of presenting the reader with
the major research developments that have taken place over the last 10 years that
contribute to the current understanding of the biology of ferns and lycophytes. To
achieve this goal they have brought together the work of some 28 contributors all active
in research, mainly from USA but also from Canada, Europe and Japan.
The subject areas are diverse. The first section on development and morphogenesis
starts with the gametophyte and alternation of generations, before exploring the
structure of meristems and subsequent diversification into the various organs – root,
stems and leaves. The next section deals with genetics and reproduction, including
chapters on the role of antheridogens, breeding systems, and the chloroplast and nuclear
genomes. Ecology is a complex discipline in itself, and the focus here is on phenology
and habitat specificity in tropical ferns, the ecology of gametophytes and a review of
conservation biology. The final section, systematics and evolutionary biology, includes
a review of the evolution of ferns and fern-like plants from the fossil record, a chapter
on species and speciation, exploration of diversity, biogeography and floristics, and
ending with phylogeny and classification.
Each chapter presents a personal review of the subject by it author(s), with summaries
of discoveries and developments, explanations of concepts and ending with a section on
prospects for the future - all supported by an extensive list of references.
This is a scholarly work, clearly aimed at the undergraduate student market, and I can
see it being added to the essential reading list for many courses in plant evolutionary
biology. But it is more than being a standard undergraduate textbook and with such a
wide variety of topics covered, there will be sections to interest a variety of general
readers and particularly for those with a fascination for fern biology. It is good to see
that it available as a more modestly priced paperback as well in hardback format.
Mary Gibby
112
FERN GAZ. 17(3). 2005
will be refereed
aligned.
aligned.
a legible form.
Supplements.
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FERN GAZ. 18(4):120-215. 2008
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AN ANNOTATED CHECKLIST OF THE MONILOPHYTES (FERNS)
AND LYCOPHYTES OF EL SALVADOR
J. MONTERROSA1 & A. K. MONRO2
1
Herbario LAGU, Asociación Jardín Botánico La Laguna, Urbanización Industrial
Plan de La Laguna, Antiguo Cuscatlán, apartado postal 1197CG
Email: [email protected]
2
Botany Department, The Natural History Museum, London SW7 5BD, U.K.
Email: [email protected]
Key words: checklist, El Salvador, pteridophytes, monilophytes, lycophytes, ferns,
Central America
ABSTRACT
We report a total of 397 species, 89 genera and 28 monilophyte and lycophyte
familes for El Salvador. 29 of these are new records for the country and we have
excluded 28 previous records. We summarise the extent of collecting within El
Salvador and list the major collectors for the country. For each record, the local
name, herbarium collections (verifiable biodiversity data), distribution within El
Salvador and conservation status (where assessed) are given. This study is the
result of a literature review and a review of 6397 herbarium specimens at BM,
F, ITIC, LAGU, MHES and NY.
RESUMEN
Se reportan un total de 397 especies, 89 generos y 28 familias de monilophyte y
lycophyte para El Salvador. 29 de las especies son nuevos registros para el país
y hemos excluido 28 registros anteriores. Se explican como el alcance de las
colectas en El Salvador y se listan los principales colectores para el país. Para
cada registro, nombre común, muestras de herabrio (datos de biodiversidad
verificable), Distribución dentro de El Salvador y estado de conservación (dónde
se valora) son dados. Este estudio es el resultado de una revisión de la literatura
y 6397 muestras de herabrio de BM, F, ITIC, LAGU, MHES and NY.
INTRODUCTION
El Salvador is located on the Pacific coast of Central America and covers 21,000 km2.
It has a population of 6.9 million (The World Bank Group, 2007), making it the most
densely populated country in Latin America. It is also, by far, the most deforested
country in continental Central and South America. Holdridge (1975) estimated that
99.5% of El Salvador would have been forested prior to human occupation. The vast
majority of the country’s biodiversity including ferns, would have been associated with
that forest cover. In 2001 a forest survey of the country based on sattelite data estimated
undisturbed natural forest cover to be 7% (Ventura & Villacorta, 2001). The loss of so
much natural forest suggests that a huge proportion of El Salvador’s fern diversity is at
risk. Conservation tools such as checklists and identification guides are therefore of
critical importance since they enable the accurate identification and monitoring of
diversity. A fact recognised by El Salvador in the Estrategía Nacional de Biodiversidad
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FERN GAZ. 18(4): 120-215. 2008
(Ministerio de Medio Ambiente y Recursos Naturales, 2000) and the Global Strategy
for Plant Conservation (Secretariat of the Convention on Biological Diversity and
Botanic Gardens Conservation International, 2002).
Past studies of the Salvadoran Pteridophyte flora
Ferns represent one of the most studied groups of organisms in El Salvador. The first
study of Salvadoran ferns was undertaken in 1930 by Maxon & Standley, Ferns of the
Republic of Salvador . This was based on the collections of Standley and listed 87
species for the country. In 1941 Calderón & Standley published the Lista Preliminar de
Plantas de El Salvador (Preliminary list of the plants of El Salvador) which included
97 species. Thirteen years later Löetschert (1954), based at the ITIC herbarium of the
University of El Salvador, published Nuevas Pteridofitas para El Salvador (New
pteridophytes for El Salvador) in which 75 new records were included, bringing the
total unmber of species to 172. In 1982, Ralph Seiler, a US Peace Corp worker based at
the Museo de Historia Natural de El Salvador (MHES) from 1980 to 1982, published
Contribuciones a la Pteridología Centroamericana (Contributions to Central American
Pteridology) in which he recognised 316 species for El Salvador. This was based on a
combination of his own collections (denoted by a ‘*’ in Seiler, 1982) and those that he
studied at ITIC. Towards the end of the 1980’s Pfeiffer-Berendsohn, published Listado
Básico de la Flora Salvadorensis. Pteridophyta. Cuscatlania 1(1). This was based on the
PLANTAS collections database of the research section of the Jardín Botánico La
Laguna and listed 346 species for the country. This number was reduced with the
publication of Flora Mesoamericana (Volume 1: Psilotaceae to Salviniaceae, Davidse
et al.) in 1995. Flora Mesoamericana recorded 304 species for El Salvador although the
collections at LAGU and ITIC were not consulted.
MATERIALS AND METHODS
The checklist is based on an inventory of the fern collections at BM, F, ITIC, LAGU,
MHES and NY. All material was critically studied using the following reference works:
Listado Básico de la Flora Salvadorensis. Pteridophyta. Cuscatlania 1(1) (Berendsohn,
W. G., 1989); ‘The Pteridophytes of Mexico’ (Mickel & Smith, 2004); and Flora
Mesoamericana Volume 1 (Davidse et al., 1995). In additions colleagues in Costa Rica
and Mexico (Alexander Rojas Alvarado, Jardín Botánico de Lankaster, Costa Rica y
Daniel Tejero-Diez, Universidad Nacional Autónoma de Mexico, Iztacala, Mexico)
were contacted where there were doubts over the indentifion of a species. In total 6397
collections were examined: 1329 at BM, 931 at F, 523 at ITIC, 1523 at LAGU, 992 at
MHES and 1099 at NY.
Species were considered as being part of the Salvadoran flora where a herbarium
collection of known Salvadoran origin was seen by the authors, or where a species was
cited in one of the publications listed in Table 1 and referenced to a herbarium
collection. Species records published elsewhere, or where cited in a publication in Table
1 but not referenced to a herbarium collection (or where the identification resulting in
a citation was incorrect) were excluded. Of the species that met the above criteria for
inclusion in the checklist some were excluded where a herbarium collection was based
on cultivated material. Examples of other species that were excluded from the checklist
are: Athyrium dombei Desv., a species described from Peru and whose citation for El
Salvador is known only from Seiler (1982) but was not based on any herbarium material
from El Salvador; and Asplenium castaneum Schlecht. & Cham. (C.D. Adams, 1995)
MONTERROSA & MONRO: CHECKLIST OF THE FERNS OF EL SALVADOR 122
whose citation was based on the misidentification of a Salvadoran collection (Seiler
160, F) of Asplenium polyphyllum Bertol.
Where species were cited in one of the publications but no specimens were seen by
these authors, this is indicated in the checklist by the symbol: ‘X’. Species were
considered new records where that species, or its basionym / synonym had not been
cited as occurring in El Salvador in a previous floristic publication encompassing the
♦’.
country (see Table 1). Such species are indicated in the checklist by the symbol ‘♦
The classification of ferns used in this study follows Smith et al. (2006).
RESULTS
A total of 397 species, 89 genera and 28 pteridophyte familes are reported for El
Salvador (summarized in Table 3). This compares with previous estimates ranging from
87 to 346 (Maxon & Standley, 1930; Calderón & Standley, 1940; Löetschert, 1954;
Seiler, 1982; Berendsohn, 1989; Davidse et al., 1995). The records are listed below. 25
of these are new records for the country and we have excluded 28 previous records.
Only two species are endemic to El Salvador: Elaphoglossum microproductum A. Rojas
(known only from the type collection) and Anemia salvadorensis Mickel.
New records for El Salvador
29 new records were encountered during the course of this work. They are indicated by
♦’ preceding the taxon name in Appendix 1.
the symbol ‘♦
Asplenium cirrhatum Rich. ex Willd.
Asplenium cristatum Lam.
Asplenium monodon Liebm.
Asplenium theciferum (Kunth) Mett.
Bolbitis bernoullii (Kuhn ex Christ) Ching
Ctenitis equestris var. erosa Stolze
Dennstaedtia globulifera (Poir.) Hieron.
Dennstaedtia spinosa Mickel
Diplazium drepanolobium A.R. Sm.
Elaphoglossum lonchophyllum (Fée) T. Moore
Elaphoglossum minutum (Pohl ex Fée) T. Moore
Hymenophyllum crispum Kunth
Lastreopsis effusa subsp. divergens (Willd.) Tindale
Megalastrum pulverulentum (Poir.) A. R. Sm. & R. C. Moran
Olfersia cervina (L.) Kunze
Pityrogramma ochracea (C. Presl) Domin
Polypodium diplotrichum Mickel & Beitel
Polypodium thyssanolepis A. Braun ex Klotzsch
Polystichum muenchii (Christ) C. Chr.
Psilotum nudum (L.) P. Beauv.
Pteris erosa Mickel & Beitel
Terpsichore delicatula (M. Martens & Galeotti) A. R. Sm.
Thelypteris angustifolia (Willd.) Proctor
Thelypteris linkiana (C. Presl) R.M. Tryon
Thelypteris obliterata (Sw.) Proctor
Thelypteris serrata (Cav.) Alston
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FERN GAZ. 18(4): 120-215. 2008
Thelypteris standleyi (Maxon & C. V. Morton) R. M. Tryon
Trichomanes hymenophylloides Bosch
Vittaria bradeorum Rosenst.
Species excluded from the checklist
Asplenium auritum Sw. var. auritum
The only record of this species (Seiler, 1982) is an incorrectly identified collection of
Asplenium monodon Liebm. (Seiler 466, MUHNES).
Asplenium castaneum Schlecht. & Cham.
The only record of this species (Adams, 1995), is an incorrectly identified collection of
Asplenium polyphyllum Bertol. (Seiler 160, F).
Asplenium hastatum Klotzsch ex Kunze
Seiler (1982) referred to the presence of this species in El Salvador, but referred to its
presence as dubious because he had not seen a collection from the country. No
collections of this species were located by Monterrosa & Monro.
Asplenium otites Link
Seiler (1982) reported this species for El Salvador. No specimens or additional citations
associating this species to El Salvador were encountered and reports of this species
from Mesoamerica as a whole are uncertain (Adams, 1995; Mickel & Smith, 2004).
Asplenium radicans L.
Seiler (1982) reported Asplenium radicans var. partitum for El Salvador. This variety
has since been placed in synonymy with A. flabellulatum. (Adams, 1995).
Asplenium rigidum Sw.
This species was reported from El Salvador (Adams, 1995) based on misidentified
material (see Mickel & Smith 2004: 90, under discussion for Asplenium cuspidatum
Lam.). No collections of this species from El Salvador have been located.
Athyrium dombei Desv.
This species was reported from El Salvador (Adams, 1995) based on misidentified
material (see Mickel & Smith 2004: 137 under discussion for Athyrium bourgeaui E.
Fourn.). No collections of this species from El Salvador have been located.
Dennstaedtia dissecta (Sw.) T. Moore
This species is restricted to the Greater Antilles, Costa Rica, Panama and the Andes
(Navarrete & Øllgaard, 2000). It is likely that Seiler (1982) mistakenly identified
material of Dennstaedtia cornuta (Kaulf.) Mett., which was previously treated as
conspecific to D. dissecta (see discussion of D. cornuta in Mickel & Smith, 2004).
Elaphoglossum firmum (Mett.) Urb.
Seiler (1982) referred to the presence of this species in El Salvador as dubious because
he had not seen a collection from the country. No collections of this species were
located by Monterrosa & Monro.
Elaphoglossum formosum (M. Martens & Galeotti) Urb.
Elaphoglossum longifolium (Jacq.) J. Sm.
Elaphoglossum pilosum (Humb. & Bonpl. ex Willd) T. Moore
Elaphoglossum revolvens (Kunze) C. Chr.
Elaphoglossum rigidum (Aubl.) Urban
Although Stolze (1981) reported this species (as a synomym of E. flaccidum) from El
Salvador, Guatemala and Mexico, Mickel & Smith 2004 do not recognize this species
from Mexico and Mickel in Davidse et al. (1995) treats E. flaccidum as native to South
America.
Grammitis heteromorpha (Hook. & Grev.) C. V. Morton
he did not see a collection from the country. No collections of this species were located
by Monterrosa & Monro. In addition, Mickel & Smith (2004) state that G.
heteromorpha is restricted to the Andes.
Grammitis meridensis (Klotzsch) F. Seym.
Huperzia mexicana (Herter) Holub
Huperzia verticillata (L.f) Trevis.
Lellingeria delitescens (Maxon) A. R. Sm. & R. C. Moran
Notholaena affinis (Mett.) Hook. ex T. Moore
The only record of this species (Seiler 88, MHES, NY; Seiler, 1982) is an incorrectly
identified collection of Notholaena meridionalis Mickel.
Polypodium arcanum Maxon
Polypodium montigenum Maxon
Seiler (1982) referred to the presence of this species in El Salvador but did not cite a
collection. No collections of this species were located by Monterrosa & Monro and it
is possible that Seiler incorrectly identified a collection of Polypodium alansmithii R.
C. Moran as P. montigenum.
Polypodium ptilorrhizon Christ
This species was reported in El Salvador by Seiler (1982) and Berendsohn (1989) and
is based on a misidentification by Seiler (1982).
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FERN GAZ. 18(4): 120-215. 2008
Polystichum ordinatum (Kunze) Liebm.
Maxon & Standley (1930) and later Calderón & Standley (1941) reported this species
from El Salvador. Seiler (1982) referred to the presence of this species in El Salvador
as dubious because he did not see a collection from the country. No collections of this
species were located by Monterrosa & Monro.
Pteris pulchra Schltdl. & Cham.
Maxon & Standley (1930) reported this species as present in El Salvador. Seiler (1982)
referred to the presence of this species in El Salvador as dubious because he did not see
a collection from the country. No collections of this species were located by Monterrosa
& Monro.
Pteris polita Link.
Seiler (1982) cites Pteris polita as present in El Salvador based on the misidentification
of material of Pteris orizabae.
Thelypteris opposita (Vahl) Ching
Seiler (1982) cites Thelypteris opposita as present in El Salvador on the
misidentification of material of material of Thelypteris balbisii (Spreng.) Ching and T.
resinifera (Desv.) Proctor.
Vittaria lineata (L.) J. Sm.
Calderón & Standley (1941) reported this species as present in El Salvador. Seiler
(1982) referred to the presence of this species in El Salvador as dubious because he did
not see a collection from the country. No collections of this species were located by
Monterrosa & Monro.
Uses
Marattia and Equisetum have documented (Pers. Obs.) medicinal uses: Marattia
excavate as a contraceptive; Equisetum giganteum and E. myriochaetum as diuretics
and hypoglycaemics and Equisetum species “cola de caballo” are commonly sold in
local markets for these properties (Pers. Obs.). Polypodium pleurosorum “calaguala” is
also sold as a medicinal plant, however the purported properties are unknown. Cyathea
divergens, Nephrolepis spp. “cola de ardilla” and Polypodium pleurosorum are used as
ornamentals in gardens and landscaping and the stems of Cyathea species are sold as a
growing medium for the cultivation of orchids. No other uses are known for Salvadoran
ferns.
Table 1. State-wise distribution of fern species records based on collections at LAGU.
State
# State records
State
# State records
Ahuachapán
141
San Miguel
34
Cabañas
24
San Salvador
107
Chalatenango
142
San Vicente
46
Cuscatlán
10
Santa Ana
236
La Libertad
71
Sonsonate
87
La Paz
12
Usulután
47
Morazán
122
Total
952
Also of economic significance is the invasive species Pteridium aquilinum var. feei
(bracken) that has been responsible for the poisoning of cattle throughout the country
(Ministerio de Agricultura y Ganaderia, Pers. Comm., 2005).
Locality distribution data within El Salvador
Species records for a locality are the product of species distribution and sampling effort,
with differences in the number of species records from one state to the next not
necessarily reflecting differences in fern diversity or sampling effort. Combined with
knowledge of the geography and vegetation of a country and collecting effort as
indicated by herbarium records, however, they can be used to give an indication of both
variation in sampling effort and diversity.
Analysis of the 1523 collections held at LAGU indicates that sampling effort has
been very uneven in El Salvador - Santa Ana, Ahuachapán, Chalatenango and Morazán
accounting for ca 75% of species-records whilst Cuscatlan and La Paz account for just
1.7% of records. In part this correlates with the expected distribution of fern diversity
based on altitudinal gradients in these states. The unevenness of the records is, however,
much greater than would be expected from altitudinal gradients and it appears that there
has been collecting bias. Most likely collection effort has been focused on protected
areas: the cloud forest of reserve of Montecristo (Santa Ana), the lowland tropical moist
forest reserve of El Impossible (Ahuachapán), El Pital (Chalatenango) and the Río Sapo
oak and pine forest reserve (Morazán). This also corresponds with the authors’ own
knowledge of collecting effort. A future collection program focussing on the states with
the fewest species records (Cuscatlan, La Paz, Cabañas, San Vicente and San Miguel)
and agricultural (e.g. shade coffee farms) and private land would be expected to provide
valuable baseline data essential for the conservation of ferns in El Salvador.
Principal collectors and collections
The principle collectors of ferns in El Salvador have been William Lötscher (also
known as Loetschert, first set of collections at ITIC), William Maxon (first set of
collections at US), Paul Standley (main sets of collections at US and F), Calderón (main
sets of collections at US), John Tucker (first set of collections at DAV), Antonio Molina
R. (first set of collections at EAP), Norman Fasset (first set of collections at WIS),
Louis Williams (first set of collections at F) and Margery Carlson (first set of
collections at F) and Ralph Seiler (first set of collections at MHES and F)
DISCUSSION
El Salvador’s fern diversity in a national context
Ferns are one the best studied groups of organisms in El Salvador, alongside birds and
reptiles. The first checklist to the ferns of the country dates to the 1920s and several
have followed (see introduction). This comparitively strong knowledge of the fern flora
of El Salvador makes ferns a good candidate for developing national inventory
protocols and conservation strategies that will contribute to El Salvador’s commitment
to the Global Strategy for Plant Conservation (2002). A Darwin Initiative project
initiated in 2006 with the Ministerio de Medio Ambiente y Recursos Naturales
(Ministry of the Environment and Natural Resources), ‘Tools and training for fern
conservation and monitoring (El Salvador)’, seeks to do exactly this and use ferns to
develop a country-wide monitoring strategy that can subsequently be applied to other
groups of organisms.
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FERN GAZ. 18(4): 120-215. 2008
Fern species-records for Salvadoran ferns are very unevenly distributed between
states, indicating an apparent sampling bias towards the country’s national parks and
against private land and ‘disturbed’ vegetation (secondary forest, agroforest, small
forest fragments). The relatively poorly sampled disturbed vegetation represents the
greater part of the El Salvador’s forest cover (19% versus 7%) and further sampling in
this areas should generate many new species-records at the state and possibly national
level. As an example of this potential, an inventory of the ferns growing within 27 shade
coffee farms enumerated 38 fern species (Monro et al., 2002) and accounts for a
significant proportion of the fern records for the state of Cabañas. It should therefore be
seen as a priority to sample such areas.
The economic importance of ferns to El Salvador although yet to be evaluated, is
likely to be minimal and restricted to medicinal and ornamental uses and the toxicity of
the invasive Pteridium aquilinum.
El Salvador’s fern diversity in a regional context
El Salvador has relatively high species diversity per km2 compared with the
neighbouring countries of Central America. This is possibly a sampling artifact, El
Salvador having been more intensively collected for ferns than other countries in
Central America. Although the most active collectors of ferns in El Salvador expended
greater collecting effort in Guatemala, Guatemala has a greater area - over five times
the size of El Salvador. El Salvador’s high fern diversity could also be ascribed to its
large altitudinal variation (2418 m) and variation in rainfall across the country (11001700 mm pa), for its size. The transition from the dry coastal lowlands to the wetter
highlands inland, combined with the presence of a number of isolated volcanic peaks
that reach altitudes greater than 1700 m (Santa Ana, Coatepeque, Apaneca, Izalco, San
Salvador, Ilopango, San Miguel, San Vicente, Tecapa and Conchagua) has resulted in a
great diversity of forest and non-forest habitat types (Ventura et al., 2001) at altitudes
associated with high fern diversity elsewhere (Kluge et al., 2006), many of which are
isolated from each-other. Kluge et al. (2006), studying fern diversity along an altitudinal
gradient in Costa Rica found that fern diversity peaked at mid-elevations (ca 1700 m)
and that the mid-domain position together with humidity and temperature were the most
powerful explanatory variables species diversity. It may be that a combination of the
relative frequency of mid-domain positions in volcanoe-dominated landscapes is high
and that this, combined with a relatively steep rainfall gradient across the landscapes
contributes to the relatively high diversity of ferns in El Slavador.
As is the case for Neotropical ferns in general the level of endemism is very low at
ca 0.5% of the species.
Table 2. Current estimates of fern diversity for Central America.
Country
Fern diversity (source)
Guatemala
652 (Stolze, R. G., 1976, 1981, 1983)
Belize
234 (Balick et al. 2000)
S u r f a c e Altitudinal Spp. per
area km2 range (m) 1000 km2
108,900
4220
6.0
22,960
1124
11.9
El Salvador 397 (this article)
21,000
2418
18.9
Costa Rica
1112 (Hammel et al., 2004)
51,100
3810
21.7
Panama
938 (Correa et al., 2004)
75,520
3475
12.4
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TEJERO-DÍEZ, J. D. & PACHECO, L., 2004. Notas Taxonómicas y de Distribución en
Polypodium plesiosorum y P. rhodopleuron (Polypodiaceae: Pteridophyta). Anales
Inst. Biol. Univ. Nac. Auton. Mexico, Bot. 75(1): 11-37.
TEJERO-DÍEZ, J. D. & PACHECO, L., 2004. Taxa Nuevos, Nomenclatura,
Redefinición y Distribución de las Especies Relacionadas con Polypodium colpodes
Kunze (Polypodiaceae, Pteridophyta). Acta Bot. Mex. 67: 75-115.
THE WORLD BANK GROUP, 2007. El Salvador Data Profile.
http://devdata.worldbank.org/external/CPProfile.asp?SelectedCountry=SLV&CCO
DE=SLV&CNAME=El+Salvador&PTYPE=CP
VENTURA, N. E., VILLACORTA, R. F., 2001. Mapeo de Vegetación Natural de
Ecosistemas Terrestres y Acuáticos de El Salvador (CD Rom). San Salvador:
Ministerio de Medio Ambiente y Recursos Naturales.
ACKNOWLEDGEMENTS
We would like to thanks the curators at BM, F, ITIC, LAGU, MHES and NY for
providing access to the collections. In particular we would like to thank Alison Paul
(BM) for help with the identification of some of the material at BM, Robbin Moran and
John Mickel (NY) for their hospitality and kind help in the identification of material
whilst at NY, José Daniel Tejero-Díez of the Faculty of Iztacala (UNAM) for the
revision of the checklist and help with the nomenclature of a number of Polypodium
names, Alan Smith (UC) for his kind help in the identification of material of
Thelypteris, David Barrington (VT) for his kind help in the identification of material of
Polystichum and Alexander Rojas Alvarado (Lankester Botanic Garden) for his help
with the identifaction of Cyathea and Elaphoglossum collections. We would also like to
thank the Association La Laguna Botanic Garden for permission to undertake this
research and access to the PLANTAS database and Gerrit Davidse and Jim Solomon at
MO for provding a dump of Salvadoran fern records in TROPICOS. Jorge Monterrosa
benefited from a Darwin Scholar Programme Award (No. EIDPS7), was able to visit F
as part of Darwin Initiative project 640/15/015 and was able to visit NY as part of the
Norwegian Agency for Development Cooperation project “Desarrollando Capacidades
y Compartiendo Tecnología para la Gestión de la Biodiversidad en Centroamérica”
through The Natural History Museum of El Salvador (MHES). We would also like to
thank Harald Schneider and Bob Press (BM) for their comments on earlier drafts of this
manuscript.
APPENDIX 1. CHECKLIST OF THE MONILOPHYTES (FERNS) AND
LYCOPHYTES OF EL SALVADOR
Ferns or monilophytes are arranged by family in alphabetical order following Smith et
al. (2006) asignment of genera to family. Lycophytes are arranged by family in
alphabetical order following the asignment of genera to family of Flora
Mesoamericana (Davidse et al., 1995).
Conspectus of the monilophytes and lycophytes of El Salvador (see pages 131-132)
♦”. Endemic species are indicated by
New records for El Salvador are indicated by “♦
“z”. Species for which no collections have been seen but where we are confident of
their inclusion by previous authors (see Table 1) are indicated by “X”.
DIVISION LYCOPHYTA
LYCOPODIACEAE
Huperzia cuernavacensis (Underw. & F. E. Lloyd) Holub, Folia Geobot. Phytotax. 20:
72. 1985.
Lycopodium cuernavacensis Underw. & F. E. Lloyd, Urostachys cuernavacensis
(Underw. & F. E. Lloyd) Hetter ex Nessel
Global distribution: Mexico, Guatemala.
Salvadoran distribution: Chalatenango: R. Seiler 157 (MHES).
Huperzia dichotoma (Jacq.) Trevis., Atti Soc. Ital. Sci. Nat. 17: 248. 1874.
Lycopodium dichotomum Jacq., Urostachys dichotomus (Jacq.) Herter . Local name:
candelillo.
Global distribution: USA, Mexico, Guatemala, Belize, Honduras, Nicaragua, Costa
Rica, Panama, Antilles, Colombia, Venezuela, Trinidad and Tobago, French Guyana,
Peru, Brazil, Bolivia, Paraguay.
Salvadoran distribution: Ahuachapán: Arana s.n. (JBL04315) (LAGU); R. Villacorta
et al. 908 (B, LAGU, MO); W. Berendsohn & M. Sandoval 1634 (LAGU).
Chalatenango: H. Sipman et al. s.n. (JBL02486) (LAGU).
131
FERN GAZ. 18(4): 120-215. 2008
TABLE 3. Conspectus of the monilophytes and lycophytes of El Salvador.
S
p
p.
Huperzia
11
DIVISION
Lycopodiales
Lycopodiaceae
Lycopodiella
3
LYCOPSIDA
LYCOPHYTA
Lycopodium
2
Selaginellales
Selaginellaceae
Selaginella
13
Botrychium
3
Ophioglosales
Ophioglosaceae
PSILOTOPSIDA
Ophioglossum
1
Psilotales
Psilotaceae
Psilotum
2
EQUISETOPSIDA Equisetales
Equisetaceae
Equisetum
4
MARATTIOPSIDA Marattiales
Marattiaceae
Marattia
2
Osmundales
Osmundaceae
Osmunda
2
Hymenophyllum 9
Hymenophyllales Hymenophyllaceae
Trichomanes
9
Dicranopteris
1
Diplopterygium 1
Gleicheniales
Gleicheniaceae
Gleichenella
1
Sticherus
4
Lygodiaceae
Lygodium
1
Schizaeales
Anemiaceae
Anemia
6
Marsileaceae
Marsilea
1
Salviniales
Azolla
2
Salviniaceae
Salvinia
2
Culcitaceae
Culcita
1
Cibotiaceae
Cibotium
1
DIVISION
Alsophila
3
EUPHYLLOPHYTA
Cyatheales
Cyatheaceae
Cyathea
7
SUB-DIVISION
Sphaeropteris
1
MONILOPHYTA
Dicksonia
1
POLYPODIOPSIDA
Dicksoniaceae
Lophosoria
1
Lindsaea
1
Lindsaeaceae
Lonchitis
1
Dennstaedtia
5
Histiopteris
1
Dennstaedtiaceae
Hypolepis
4
Pteridium
3
Acrostichum
2
Adiantum
18
Polypodiales
Ananthacorus
1
Anogramma
2
Bommeria
1
Pteridaceae
Ceratopteris
2
Cheilanthes
11
Cheiloplecton
1
Doryopteris
1
Eriosorus
1
Hemionitis
3
Division
Class
Order
Family
Genus
TABLE 3. (continued from previous page)
Division
Class
Order
Family
Genus
S
p
p.
Mildella
1
Notholaena
2
Pellaea
1
Pityrogramma
6
Pteridaceae
Polytaenium
1
Pteris
11
Scoliosorus
1
Vittaria
2
Aspleniaceae
Asplenium
29
Macrothelypteris
1
Thelypteridaceae
Thelypteris
29
Athyrium
3
Cystopteris
1
Woodsiaceae
Diplazium
10
Woodsia
1
Blechnum
15
Blechnaceae
Woodwardia
2
Arachniodes
1
Bolbitis
3
Ctenitis
8
DIVISION
Dryopteris
5
EUPHYLLOPHYTA
Elaphoglossum
29
POLYPODIOPSIDA Polypodiales Dryopteridaceae
SUB-DIVISION
Lastreopsis
1
Megalastrum
1
MONILOPHYTA
Olfersia
1
Phanerophlebia
2
Polystichum
8
Cyclopeltis
1
Lomariopsidaceae
Nephrolepis
4
Tectariaceae
Tectaria
5
Ceradenia
2
Cochlidium
2
Lellingeria
2
Melpomene
6
Micropolypodium
1
Terpsichore
7
Campyloneurum
6
Polypodiaceae
Loxogramme
1
Niphidium
1
Pecluma
2
Phlebodium
1
Pleopeltis
4
Polypodium
22
Serpocaulon
2
Total
397
133
FERN GAZ. 18(4): 120-215. 2008
Huperzia hippuridea (Christ) Holub, Folia Geobot. Phytotax. 20: 73. 1985.
Lycopodium hippurideum Christ in Pittier, Urostachys hippurideus (Christ) Herter ex
Nessel
Global distribution: Mexico, Guatemala, Honduras, Costa Rica, Panama, Antilles,
Colombia, Venezuela, Guayana, Ecuador, Peru, Bolivia.
Salvadoran distribution: Santa Ana: R. Villacorta et al. 1284 (B, LAGU, MO).
Chalatenango: R. Villacorta et al. 2052 (B, LAGU, MO); R. Seiler 158 (F, MHES).
Huperzia linifolia (L.) Trevis., Atti Soc. Ital. Sci. Nat. 17: 248. 1874.
Lycopodium linifolium L., Urostachys linifolius (L.) Herter
Global distribution: Mexico, Guatemala, Belize, Honduras, Nicaragua, Costa Rica,
Panama, Antilles, Colombia, Venezuela, Trinidad and Tobago, Suriname, Ecuador,
Brazil, Bolivia.
Salvadoran distribution: Santa Ana: H. Weber 277 (ITIC); R. Seiler 247 (MHES); S.
Winkler 8b (ITIC).
Huperzia myrsinites (Lam.) Trevis., Atti Soc. Ital. Sci. Nat. 17: 248. 1874.
Lycopodium myrsinites Lam., L. skutchii Maxon, Plananthus myrsinites (Lam.) P.
Beauv., Urostachys myrsinites (Lam.) Herter
Global distribution: Mexico, Guatemala, Honduras, Nicaragua, Costa Rica, Antilles,
Colombia, Venezuela, Trinidad and Tobago, Guayana, Ecuador, Peru.
Salvadoran distribution: Santa Ana: R. Seiler 235 (F, ITIC, MHES); 288 (MHES); R.
Villacorta et al. 1278, 1280 (B, LAGU, MO). Chalatenango: R. Villacorta et al. 2057
(B, LAGU, MO); R. Seiler 401 (JBL00110) (F, LAGU, MHES); 747 (F); S. Winkler 1a
(ITIC). Morazán: R. Seiler 1021 (F, MHES).
Huperzia orizabae (Underw. & F. E. Lloyd) Holub, Folia Geobot. Phytotax. 20: 75.
1985.
Lycopodium orizabae Underw. & F. E. Lloyd, Urostachys orizabae (Underw. & F. E.
Lloyd) Herter ex Nessel
Global distribution: Mexico, Guatemala, Honduras.
Salvadoran distribution: Kovar 2594, US (Øllgaard in Davidse et al. 1995). Santa Ana:
R. Seiler 764 (MHES).
Huperzia pithyoides (Schldtl. & Cham.) Holub, Folia Geobot. Phytotax. 20: 76. 1985.
Lycopodium pithyoides Schltdl. & Cham., L. gigas Herter, Urostachys pithyoides
(Schltdl. & Cham.) Herter, U. gigas (Herter) Herter
Global distribution: Mexico, Guatemala, Honduras, Nicaragua, Costa Rica, Antilles,
Colombia, Venezuela.
Salvadoran distribution: Santa Ana: R.A. Carballo et al. 90 (B, LAGU); R. Seiler 310
(MHES); S. Winkler 6b (ITIC). Chalatenango: R. Seiler 420 (JBL00111) (LAGU); R.
Seiler 746 (MHES). Morazán: R. Seiler 982 (F, ITIC, MHES).
Huperzia pringlei (Underw. et F. E. Lloyd) Holub, Folia Geobot. Phytotax. 20: 76.
1985.
Lycopodium pringlei Underw. & F. E. Lloyd, L. tuerckheimii Maxon, Urostachys
pringlei (Underw. & F. E. Lloyd) Herter ex Nessel, U. tuerckheimii (Maxon) Herter,
Huperzia tuerckheimii (Maxon) Holub
Salvadoran distribution: Santa Ana: H. Weber 294 (ITIC); J. Lagos 12 (ITIC); R.
Seiler 180, 767 (MHES); R. Villacorta & H. Sipman 1275 (B, LAGU, MO); S. Winkler
3 (ITIC). Chalatenango: R. Seiler 159 (ITIC, MHES); 743 (MHES). Morazán: E.A.
Montalvo s.n. (ITIC); R. Seiler 1001 (F, MHES); 1022 (MHES).
Note: Seiler (1982) and Berendsohn (1989) treat L. pringlei and L. tuerckheimii as
distinct species.
Huperzia reflexa (Lam.) Trevis., Atti Soc. Ital. Sci. Nat. 17: 248. 1874.
Lycopodium reflexum Lam., Urostachys reflexus (Lam.) Herter
Panama, Antilles, Colombia, Venezuela, Trinidad and Tobago, Ecuador, Peru, Brazil,
Bolivia, Paraguay.
Salvadoran distribution: Sonsonate: R. Villacorta & M.L. Reyna de Aguilar 716 (B,
ITIC, LAGU, MO). Santa Ana: A. Molina et al. 21712 (F, ITIC); M.C. Carlson 979
(F); R. Seiler 267 (F, MHES); 883 (F, ITIC, MHES); S. Winkler 7a (ITIC). La
Libertad: M.C. Carlson 249 (F); R. Villacorta & E.A. Montalvo 2314 (B, LAGU, MO).
San Salvador: A. Molina et al. 21862 (F). San Vicente: E.A. Montalvo s.n. (ITIC).
Morazán: R. Seiler 1013, 1085 (F, MHES); 1052 (MHES); 1077 (F, ITIC, MHES).
Huperzia taxifolia (Sw.) Trevis., Atti Soc. Ital. Sci. Nat. 17: 248. 1874.
Lycopodium taxifolium Sw., Urostachys taxifolius (Sw.) Herter
Panama, Antilles, Colombia, Venezuela, Trinidad and Tobago, Ecuador, Peru, Bolivia.
Salvadoran distribution: Ahuachapán: A. Molina et al. 21760 (ITIC); E.A. Montalvo
5463 (ITIC); R. Seiler 959 (F, MHES). Santa Ana: G. Davidse et al. 37244 (B, BM,
ITIC, LAGU, MO, L). González 1981 (ITIC); R. Seiler 307 (F, MHES); S. Winkler 4
(ITIC). San Vicente: J. Lagos 2381 (ITIC).
Huperzia watsoniana (Maxon) B. Øllg., Ann. Missouri Bot. Gard. 79: 701. 1992.
Lycopodium watsonianum Maxon
Global distribution: Costa Rica, Panama.
Salvadoran distribution: Molina et al. 16823, F (Øllgaard in Davidse et al. 1995).
Santa Ana: R. Seiler 232 (MHES); 289 (F, MHES). Chalatenango: R. Seiler 744
(MHES).
Lycopodiella alopecuroides (L.) Cranfill, Amer. Fern J. 71: 97. 1981.
Lycopodium alopecuroides L.
Global distribution: USA, Mexico, Guatemala, Antilles, Colombia, Venezuela,
Ecuador, Peru, Brazil, Bolivia, Paraguay, Uruguay.
Salvadoran distribution: Morazán: R. Seiler 1078 (F, ITIC, MHES).
Lycopodiella caroliniana (L.) Pic. Serm., Webbia 23: 165. 1968.
Lycopodium carolinianum L., L. meridionale Underw. & F. E. Lloyd, L. carolinianum
L. var. meridionale (Underw. & F. E. Lloyd) Nessel, Pseudolycopodiella meridionalis
(Underw. & F. E. Lloyd) Holub, Lycopodiella caroliniana (L.) Pic. Serm. var.
meridionalis (Underw. & F. E. Lloyd) B. Øllg. & P. G. Windisch.
Global distribution: USA, Mexico, Guatemala, Belize, Honduras, Nicaragua, Panama,
135
FERN GAZ. 18(4): 120-215. 2008
Antilles, Colombia, Venezuela, Guayana, Suriname, French Guyana, Ecuador, Peru.
Salvadoran distribution: Morazán: R. Seiler 1014 (F, MHES).
Lycopodiella cernua (L.) Pic. Serm., Webbia 23: 166. 1968.
Lycopodium cernuum L., Lepidotis cernua (L.) P. Beauv., Palhinhaea cernua (L.) Vasc.
& Franco.
Rica, Panama, Antilles, Colombia, Venezuela, Trinidad and Tobago, Guayana,
Suriname, French Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Trópicos y
Subtrópicos del Viejo Mundo.
Salvadoran distribution: Santa Ana: B. Bechyne 3908 (ITIC); E. Peñate s.n. (ITIC); R.
Seiler 39 (F, MHES); 1204 (F); Thomson 55 (ITIC). San Salvador: N.C. Fassett 28601
(ITIC); W. Eggler 682 (F). Chalatenango: A. Díaz s.n. (ITIC); M.C. Carlson 592 (F);
R. Villacorta & A. Araniva 980 (LAGU, MO, B, ITIC). Usulután: J. Lagos 2 (ITIC).
Morazán: R.A. Carballo et al. 1032 (LAGU, MO); R. Seiler 978 (F, ITIC, MHES);
1012, 1043 (F, MHES).
Lycopodium clavatum L., Sp. Pl. 2: 1101. 1753.
Lycopodium aristatum Humb. & Bonpl. ex Willd., L. piliferum Raddi, L. trichophyllum
Desv., L. aristatum Humb. & Bonpl. ex Willd. var. incurvum Grev. & Hook., L.
aristatum Humb. & Bonpl. ex Willd. var. robustius Grev. & Hook., L. clavatum L. var.
raddianum Spring, L. clavatum L. var. aristatum (Humb. & Bonpl. ex Willd.) Spring,
L. clavatum L. var. desvauxianum Spring, L. clavatum L. var. jamaicense Spring, L.
aristatum Humb. & Bonpl. ex Willd. var. desvauxianum Spring, L. trichiatum Bory var.
desvauxianum Spring, L. eriostachys Fée, L. clavatum L. var. eriostachys (Fée) Nessel
& Hoehne, L. clavatum L. var. piliferum (Raddi) Nessel & Hoehne, L. clavatum L. var.
equisetoides Schwacke ex Nessel & Hoehne, L. clavatum L. var. trichophyllum (Desv.)
Nessel & Hoehne.
Local name: guía de roble.
Global distribution: USA, Mexico, Guatemala, Honduras, Nicaragua, Costa Rica,
Panama, Antilles, Colombia, Venezuela, Guayana, Suriname, French Guyana, Ecuador,
Peru, Brazil, Bolivia, Paraguay, Europe.
Salvadoran distribution: Santa Ana: A. Molina et al. 12538, 16715 (F; B). PfeifferBerendsohn 21 (B, LAGU, MO); E.A. Montalvo s.n., 3128, 4136 (ITIC); J. Lagos, 8,
1480 (ITIC); M.L. Reyna de Aguilar 1029 (MHES); N.C. Fassett 28310 (F, ITIC); R.
Seiler 115 (F, ITIC, MHES); 769 (F, MHES); R. Villacorta et al. 1046 (B, LAGU, MO);
W. Berendsohn et al. 1720 (B, BM, LAGU, MO). Chalatenango: J. Tucker 1051 (F);
R. Villacorta et al. 2077 (B, LAGU, MO); W. Berendsohn et al. 1565 (B, LAGU, MO).
Lycopodium thyoides Humb. & Bonpl. ex Willd., Sp. Pl., ed. 4, 5(1): 18. 1810.
Diphasiastrum thyoides (Humb. & Bonpl. ex Willd.) Holub, Lycopodium complanatum
L. var. validum Weath., L. complanatum L. var. adpressifolium Spring, L. complanatum
L. var. thyoides (Humb. & Bonpl. ex Willd.) Christ, L. complanatum L. var. tropicum
Spring . Local name: helecho guía de roble.
Global distribution: Mexico, Guatemala, Honduras, Costa Rica, Panama, Colombia,
Venezuela, Ecuador, Peru, Brazil, Bolivia, Argentina.
Salvadoran distribution: Padilla 170 (US) (Øllgaard in Davidse et al. 1995).
Chalatenango: W. Berendsohn et al. 1567 (B, LAGU, MO).
SELAGINELLACEAE
Selaginella acutifolia (Stolze) Valdespino, Mem. New York Bot. Gard. 88: 558. 2004.
Selaginella pallescens (C. Presl) Spring var. acutifolia Stolze
Global distribution: Mexico, Guatemala, Belize, Honduras, Nicaragua, Costa Rica.
Salvadoran distribution: Ahuachapán: E. Sandoval & F. Chinchilla 635 (B, LAGU);
J.M. Rosales 1062, 1169 (AAU, B, BM, LAGU, MO). Sonsonate: D. Rodríguez & M.
Trejo 77 (B, LAGU, MO). La Libertad: K. Sidwell et al. 509 (B, BM, ITIC, LAGU,
MO); M. Renderos & R. Villacorta 286 (B, EAP, LAGU, MO); R.A. Carballo et al. 48
(B, LAGU, MO). Chalatenango: R. Seiler 365, 549 (MHES). La Paz: J. Lagos 4
(ITIC). San Miguel: R. Seiler 541 (MHES); R. Villacorta et al. 2753 (B, BM, EAP,
LAGU, MO). Cabañas: R.A. Carballo et al.289 (B, LAGU, MO). Morazán: R.A.
Carballo et al. 1067 (LAGU).
Selaginella bernoullii Hieron., Hedwigia 41: 192. 1902.
Salvadoran distribution: Chalatenango: R. Seiler 733 (F, MHES).
Note: M.E. Fraile in Dvidse et al. (1995) cites this name as a synonym of S.
porphyrospora A. Braun. We have chosen to maintain the species distinct following the
criteria cited by Mickel & Smith (2004).
Selaginella convoluta (Arn.) Spring in Martius, Fl. Bras. 1(2): 131. 1840.
Lycopodium convolutum Arn., Selaginella longispicata Underw. & Millsp.
Global distribution: Mexico, Guatemala, Honduras, Antilles, Brazil, Bolivia, Paraguay,
Argentina.
Source: Löetschert (1954): 23; Seiler (1982): 382; Berendsohn (1989) 1: 24.
Selaginella estrellensis Hieron., Hedwigia 41. 200. 1902.
Global distribution: Costa Rica.
Source: Fraile in Davidse et al. (1995): 32; Alston 1955 (no visto).
Selaginella hoffmannii Hieron., Hedwigia 41. 184. 1902.
Local name: yayuco colorado.
Global distribution: Mexico, Belize, Guatemala, Honduras, Nicaragua, Costa Rica.
Salvadoran distribution: Ahuachapán: E. Sandoval & F. Chinchilla 666 (B, LAGU); J.
Lagos 5 (ITIC); J.M. Rosales 1231, 1232 (B, BM, LAGU, MO); 1257 (AAU, B, BM,
LAGU, MO); 1300 (B, BM, EAP, LAGU, MO); 1619 (B, LAGU); R. Villacorta et al.
905 (B, LAGU, MO). Santa Ana: A.K. Monro & R. Douglas 3462 (B, BM, ITIC,
LAGU, MO); R. Seiler 705 (F, MHES); R. Villacorta 2179 (B, LAGU, MO). La
Libertad: A.K. Monro et al. 3082 (B, BM, ITIC, LAGU, MO); R.A. Carballo et al. 40
(B, LAGU, MO). San Salvador: Grupo Ecológico Cantón El Zapote s.n. (JBL03316)
(LAGU, MO); R. Seiler 634 (F, MHES); 1509 (F). Chalatenango: R. Seiler 143 (F,
ITIC, MHES).
Selaginella lepidophylla (Hook. & Grev.) Spring in Martius, Fl. Bras. 1(2): 126. 1840.
Lycopodium lepidophyllum Hook. & Grev.
Global distribution: USA, Mexico.
Salvadoran distribution: San Salvador: R. Seiler 48 (F, ITIC, MHES).
137
FERN GAZ. 18(4): 120-215. 2008
Selaginella martensii Spring, Acad. Roy. Sci. Belgique 24: 129. 1849.
Selaginella solmsii Baker
Global distribution: Mexico, Guatemala, Honduras, Nicaragua, Costa Rica, Panama.
Salvadoran distribution: Santa Ana: A. Molina et al. 12633, 16866, 16883 (F); E.A.
Montalvo 4137 (ITIC); G. Davidse et al. 37242, 37304 (B, BM, ITIC, LAGU, MO); J.
Lagos 14 (ITIC); J. Tablas s.n. (ITIC); J. Toledo s.n. (ITIC); M.C. Carlson 890 (F); R.
Seiler 108 (F, ITIC); 184 (F, MHES); 472 (F, ITIC, MHES); R. Villacorta et al. 1279
(B, LAGU); 2679 (B, F, LAGU, MO). La Libertad: F. López s.n. (ITIC).
Selaginella pallescens (C. Presl) Spring in Martius, Fl. Bras. 1(2): 132. 1840.
Lycopodium pallescens C. Presl, L. cuspidatum Link, Selaginella cuspidata (Link)
Link, S. cuspidata (Link) Link var. elongata Spring
Local names: helecho tuya, resurrección.
Panama, Colombia, Venezuela, Suriname, Brazil.
Salvadoran distribution: Ahuachapán: E. Sandoval et al. 602 (B, LAGU); 1338 (K,
LAGU); J.M. Rosales 1299 (B, BM, EAP, LAGU, MO); 1403, 1586 (AAU, B, BM,
LAGU, MO); M. Sandoval & E. Sandoval 20 (B, F, LAGU); T. Croat 42064 (ITIC).
Sonsonate: R. Seiler 8 (F, MHES); T. Croat 42239 (ITIC). Santa Ana: A. Molina et al.
21881 (F, ITIC); T. Croat 42300 (ITIC); V.M. Martínez s.n. (CMC00262) (B, LAGU).
La Libertad: J.C. González & M.A. Hernández 361 (B, EAP, LAGU, MO). San
Salvador: M.C. Carlson 331 (F); R. Seiler 53 (F, ITIC, MHES); 506 (F, MHES); Tobar
s.n. (12-802) (MHES). Chalatenango: D. Sloot & A. Reina 262 (MAG00691) (B,
LAGU, U). La Paz: R. Villacorta et al. 293 (B, LAGU, MO). San Miguel: W. Eggler
663, F (ITIC). Cabañas: R. Seiler 521 (F, MHES); R. Villacorta & R.M. Rivas 2119 (B,
LAGU, MO). Cuscatlán: F. Wunderlin 3026 (ITIC). Morazán: A.K. Monro et al. 3833
(B, BM, ITIC, LAGU, MO); R. Villacorta et al. (B, BM, LAGU, MO).
Selaginella porphyrospora A. Braun, Ann. Sci. Nat. Bot., sér 5, 3: 286. 1865.
Selaginella bulbifera Baker, S. binervis Liebm. ex Baker
Global distribution: Mexico, Guatemala, Honduras, Nicaragua, Costa Rica, Panama,
Colombia, Venezuela, Ecuador.
Salvadoran distribution: Sonsonate: R. Seiler 609 (F, MHES). Santa Ana: J. Williams
30 (ITIC); R. Seiler 877 (F, ITIC, MHES). San Salvador: H. Weber 7 (ITIC).
Selaginella sertata Spring, Nouv. Mém. Acad. Roy. Sci. Bruxelles 24(1): 104. 1850.
Selaginella nicaraguensis Baker
Panama.
Salvadoran distribution: Ahuachapán: J.M. Rosales 1603 (AAU, B, BM, LAGU,
MO). Santa Ana: R. Villacorta et al. 1237 (B, LAGU, MO). La Libertad: R. Villacorta
& D.Lentz 1109 (B, LAGU, MO). Chalatenango: R. Seiler 147 (MHES). La Paz: R.
Villacorta & S. Villacorta 1154 (B, LAGU, MO). San Miguel: R. Seiler 543 (MHES).
Cabañas: R. Seiler 526 (MHES).
Selaginella silvestris Asplund, Ark. Bot. 20A(7): 30 (F). 3-5. 1926.
Colombia, Venezuela, Ecuador, Peru, Bolivia.
Salvadoran distribution: Tucker 1174 (UC) (Fraile in Davidse et al. 1995).
Source: Stolze (1983): 60; Berendsohn (1989) 1: 24; Fraile in Davidse et al. (1995): 28;
Mickel & Smith (2004): 596.
Selaginella stellata Spring, Flora 21: 194. 1838.
Selaginella galeottii Spring
Global distribution: Mexico, Guatemala, Belize, Honduras, Costa Rica, Panama.
Source: Berendsohn (1989) 1: 24.
Selaginella tenella (P. Beauv.) Spring, Bull. Acad. Roy. Sci. Bruxelles 10: 234. 1843.
Diplostachium tenellum P. Beauv., Lycopodium tenellum (P. Beauv.) Desv. ex Poir.,
Selaginella cladorrhizans A. Braun, S. subcaulescens Baker
Local name: helecho tuya.
Global distribution: Mexico, Guatemala, Belize, Honduras, Panama, Antilles,
Colombia, Venezuela, Trinidad and Tobago, Brazil.
Salvadoran distribution: Ahuachapán: E. Sandoval & O.Rivera D. 1376 (B, LAGU);
J.M. Rosales 1298 (B, BM, EAP, LAGU, MO); 1340 (B, LAGU). La Libertad: R.
Villacorta 316 (B, EAP, LAGU, MO).
DIVISION EUPHYLOPHYTA, SUB-DIVISION MONILOPHYTA
ANEMIACEAE
Anemia guatemalensis Maxon, N. Amer. Fl. 16: 46. 1909.
Local name: petatillo.
Global distribution: Guatemala, Honduras, Nicaragua.
Salvadoran distribution: Santa Ana: B. Pfeiffer-Berendsohn & W. Berendsohn 15 (F,
LAGU, MO); R. Villacorta et al. 1264 (B, ITIC, LAGU, MO). Chalatenango: D. Sloot
& A. Reina 109 (MAG00538) (B, LAGU, U); L. Lara 220 (MHES); R. Seiler 562 (F,
MHES). Morazán: J. Monterrosa & C. Rivera 712 (B, BM, LAGU, MO); R. Seiler
1050 (F, MHES).
Anemia hirsuta (L.) Sw., Syn. Fil. 156. 1806.
Osmunda hirsuta L., Ornithopteris hirsuta (L.) Bernh., Anemia hirsuta (L.) Sw. var.
achillaeifolia M. Martens & Galeotti, A. opaca Fée
Local name: helecho pluma de perico.
Antilles, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia.
Salvadoran distribution: Ahuachapán: E. Sandoval et al. 484, 1317 (B, LAGU); F.
Chinchilla s.n. (ISB00522) (AAU, B, LAGU); s.n. (ISB00501) (AAU, B, F, LAGU);
s.n. (ISB00763) (B, LAGU); J.M. Rosales 898, 972, 1404 (AAU, B, BM, LAGU, MO);
M. Sandoval & E. Sandoval 10, 39 (B, LAGU); R. Villacorta et al. 406 (B, LAGU,
MO); 952 (B, LAGU); W. Berendsohn & E. Sandoval 1481 (B, LAGU). Sonsonate: R.
Seiler 35 (F, MHES); R. Villacorta 2402 (AAU, B, LAGU). Santa Ana: B. PfeifferBerendsohn 57 (B, LAGU, MO); M.L. Reyna de Aguilar 765 (MHES); R. Villacorta &
H. Sipman 1270 (B, LAGU, MO). La Libertad: N. Ventura 5013 (ITIC); O. Rohweder
s.n. (JBL01254) (LAGU). San Salvador: J. Lagos 1076 (ITIC); R. Seiler 69 (F,
MHES); 445 (MHES); R. Villacorta & M.A. Sanabria 402 (B, LAGU, MO).
Chalatenango: R. Seiler 366 (F, MHES); 553, 569 (MHES); R. Villacorta 2362 (B,
139
FERN GAZ. 18(4): 120-215. 2008
LAGU, MO). La Paz: E.A. Montalvo 6388 (AAU, B, ITIC, LAGU, MEXU, MO). San
Vicente: N.C. Fassett 28348 (F). Usulután: D. Williams 217 (LAGU); 354 (B, LAGU,
MO). Cabañas: J. Monterrosa & R. Carballo 295 (B, BM, LAGU, MO). Morazán: J.
Monterrosa & C. Rivera 664 (B, BM, LAGU). La Unión: S. Calderón 2374 (F).
Anemia oblongifolia (Cav.) Sw., Syn. Fil. 156. 1806.
Osmunda oblongifolia Cav., O. humilis (Cav.) Sw., Anemia oblongifolia (Cav.) Sw. β
humilis (Cav.) Hook., A. pilosa M. Martens & Galeotti α brevistipes Liebm., A. cornea
Prantl, A. seemannii Hook.
Local name: helecho rosquilla pacha.
Panama, Colombia, Venezuela, Guayana, Suriname, French Guyana, Ecuador, Peru,
Brazil, Bolivia, Argentina.
Salvadoran distribution: Ahuachapán: E. Sandoval & F. Pérez 1347 (B, LAGU, MO);
W. Berendsohn & R. Villacorta 1383 (B, LAGU). Sonsonate: R. Seiler 36, 494
(MHES); 73 (MHES). Santa Ana: O. Rohweder s.n. (JBL01255), s.n. (JBL01257),
2660 (JBL01256) (LAGU). San Salvador: E.A. Montalvo 3775 (ITIC).
Chalatenango: L. Lara s.n. (JBL02202) (B, LAGU); 77 (MHES); R. Seiler 558 (F,
MHES); S. Calderón 2474 (F). Usulután: D. Williams 140 (B, LAGU, MO). Morazán:
J. Monterrosa & M. Rivera 602 (BM, LAGU); K. Sidwell et al. 773 (B, BM, ITIC,
LAGU, MO); M. Renderos et al. 629 (AAU, B, LAGU, MO); R. Villacorta et al. (B,
BM, LAGU, MO).
Anemia pastinacaria Moritz ex Prantl, Unters. Morph. Gefasskrypt. 2: 110. 1881.
Anemia pilosa M. Martens & Galeotti β longistipes Liebm., A. longistipes (Liebm.) C.
Chr., A. helveola Fée
Local names: petatillo.
Panama, Antilles, Colombia, Venezuela, Trinidad and Tobago, Guayana, Suriname,
French Guyana, Ecuador, Peru, Brazil, Bolivia.
Salvadoran distribution: Santa Ana: B. Pfeiffer-Berendsohn 60 (LAGU, MO); R.
Seiler 134 (MHES). San Salvador: R. Seiler 205b, 782 (F, MHES). Chalatenango: D.
Sloot & A. Reina 183 (MAG00612) (B, LAGU, U); R. Seiler 145 (F, MHES); 565
(MHES). Morazán: J. Monterrosa & C. Rivera 665 (BM, LAGU).
Anemia phyllitidis (L.) Sw., Syn. Fil. 155. 1806.
Osmunda phyllitidis L., Anemidictyon phyllitidis (L.) J. Sm.
Local names: helecho cola de ratón, helecho de montaña.
Antilles, Colombia, Venezuela, Trinidad and Tobago, Guayana, Suriname, Ecuador,
Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay.
Salvadoran distribution: Ahuachapán: B. Smalla & N. Kilian 81 (B, LAGU); E.
Sandoval & H. Rivera 1319 (B, LAGU); J.M. Rosales 1005 (B, LAGU); M. Sandoval
& E. Sandoval 53 (B, LAGU); R. Villacorta et al. 876 (B, LAGU); 955 (B, ITIC,
LAGU, MO). Santa Ana: J. Monterrosa et al. 260 (B, BM, LAGU, MO); 674 (B, BM,
LAGU); 921 (LAGU); 1027 (B, LAGU, NY); R. Villacorta 2172 (LAGU). San
Salvador: M.C. Carlson 332 (F); R. Seiler 65, 781 (F, MHES). Chalatenango: R.
Seiler 566 (MHES); 570 (F, MHES). Usulután: A.K. Monro et al. 3004 (B, BM, ITIC,
LAGU, MO); D. Williams 127, 131 (LAGU); 302 (LAGU, MO). Morazán: J.
Monterrosa & C. Rivera 674 (B, BM, LAGU); R. Seiler 1087 (F); 1090 (F, MHES).
z Anemia salvadorensis Mickel, Amer. Fern. J. 74: 113. 1984.
Local names: helecho de rosa, helecho plumerillo.
Global distribution: Endemic.
F. Chinchilla s.n. (ISB00541) (B, LAGU); s.n. (ISB00542) (LAGU); W. Berendsohn et
al. 1395 (B, LAGU, MO); R. Seiler 947 (MHES, NY).
ASPLENIACEAE
Asplenium abscissum Willd., Sp. Pl., ed. 4, 5(1): 321. 1810.
Asplenium polymorphum M. Martens & Galeotti. Local name: helecho quezalío.
Rica, Panama, Antilles, French Guyana; Colombia, Venezuela, Trinidad and Tobago,
Ecuador, Peru, Brazil, Bolivia, Uruguay, Paraguay.
Salvadoran distribution: Ahuachapán: E. Sandoval & F. Chinchilla 41 (LAGU); G.
Davidse et al. 37420 (B, BM, ITIC, LAGU, MO); R. Seiler 901 (F, ITIC, MHES).
Sonsonate: R. Seiler 619, 623 (F, MHES). Santa Ana: R. Seiler 111, 182, 363 (F,
MHES).
Asplenium achilleifolium (M. Martens & Galeotti) Liebm., Kongel. Danske Vidensk.
Selsk. Skr., Naturvidensk. Math. Afd. Ser. 51: 249. 1849.
Caenopteris achilleifolia M. Martens & Galeotti, Athyrium conchatum Fée, Asplenium
conchatum (Fée) T. Moore, A. grande Fée
Global distribution: Mexico, Guatemala, Honduras, Nicaragua, Costa Rica, Peru.
Salvadoran distribution: Sonsonate: A. Molina et al. 21623 (F); R. Seiler 624 (F,
MHES). Santa Ana: J. Tucher 1248 (F); M.B. Siu s.n. (ITIC); M.L. Reyna de Aguilar
762 (MHES); R. Seiler 468 (F, MHES); R. Seiler 469 (F, ITIC, MHES); R. Seiler 269
(ITIC (F, LAGU (photo), NY)); R. Seiler 872 (ITIC); S. Winkler 37 (ITIC). San
Salvador: M.C. Carlson 417 (F); R. Seiler 346 (F, ITIC, MHES).
Asplenium alatum Humb. & Bonpl. ex Willd., Sp. Pl. ed. 4, 5(1): 319. 1810.
Global distribution: Mexico, Guatemala, Nicaragua, Costa Rica, Panama, Antilles,
Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia.
Salvadoran distribution: Santa Ana: R. Seiler 309 (MHES). R. Seiler 465 (F, LAGU
(photo), NY).
Asplenium auriculatum Sw., Kongl. Vetensk. Acad. Handl. 38: 68. 1817.
Asplenium salicifolium L. var. auriculatum (Sw.) Proctor, A. semicordatum Raddi, A.
pimpinellifolium Fée & Schaffner ex Fée
Antilles, Colombia, Venezuela, Trinidad and Tobago, Ecuador, Peru, Brazil, Bolivia,
Paraguay.
Salvadoran distribution: Ahuachapán: D. Witsberger 686 (MHES); E. Sandoval & M.
Sandoval 1423 (B, LAGU); R. Seiler 821, 823, 961 (F, MHES); 910 (F, ITIC, MHES);
S. Winkler 6a (ITIC). Sonsonate: R. Seiler 615 (F, MHES). Santa Ana: A. Molina et
141
FERN GAZ. 18(4): 120-215. 2008
al. 12650, 16760, 16802a, 16806 (F; B). López s.n. (ITIC); B. Pfeiffer-Berendsohn 32,
36 (B, MO, LAGU); E.A. Montalvo 12 (ITIC); G. Cerén & D. Rodríguez s.n.
(JBL04659) (B, LAGU, MO); G. Davidse et al. 37297 (B, BM, ITIC, LAGU, MO); J.
Monterrosa & R. Carballo 554 (B, LAGU, MO); J. Tucker 1250 (F); K. Sidwell et al.
849 (B, BM, ITIC, LAGU, MO); M.C. Carlson 995, 999 (F); M.L. Reyna de Aguilar
685 (MHES); R. Seiler 92 (F, ITIC, MHES); 361, 646 (F, MHES); R. Villacorta 1121
(B, LAGU). San Salvador: M.C. Carlson 428 (F); R. Seiler 338 F (MHES). Morazán:
R. Seiler 1037 (F, ITIC, MHES); 1061 (F, MHES).
Asplenium blepharophorum Bertol., Novi Comment. Acad. Sci. Inst. Bononiensis 4:
443. 1840.
Athyrium macrocarpon Fée, Asplenium macrocarpon (Fée) T. Moore, A. myapteron
Fée, A. commutatum Mett. ex Kuhn, A. onustum Christ, A. subvestitum Copel.
Antilles.
Salvadoran distribution: Sonsonate: R. Seiler 15 (F). Santa Ana: R. Seiler 11 (ITIC,
MHES); 85 (F, MHES).
♦ Asplenium cirrhatum Rich. ex Willd., Sp. Pl., ed. 4, 5(1): 321.1810.
Asplenium radicans L. var. cirrhatum (Rich.) Rosenst.
Global distribution: Mexico, Belize, Guatemala, Honduras, Nicaragua, Costa Rica,
Panama, Antilles, French Guyana, Colombia, Venezuela, Ecuador, Peru, Brazil,
Bolivia.
Salvadoran distribution: Santa Ana: M.L. Reyna de Aguilar 848 (LAGU, MHES).
♦ Asplenium cristatum Lam., Encycl. 2: 310.1786.
Asplenium cicutarium Sw.
Rica, Panama, Antilles, Guayanas y Galapagos, Colombia, Venezuela, Trinidad and
Tobago, Ecuador, Peru, Brazil, Bolivia, Paraguay.
Salvadoran distribution: Ahuachapán: A.K. Monro et al. 1993 (B, BM, ITIC, LAGU,
MO).
Asplenium cuspidatum Lam., Encycl. 2: 310.1786 .
Asplenium lacerum Schltdl. & Cham., A. mexicanum M. Martens & Galeotti, A.
fragrans Sw. var. mexicanum (M. Martens & Galeotti) Christ, A. auritum Sw. var.
bipinnatifidum Kunze, A. pyramidatum Liebm., A. auritum Sw. var. bipinnatisectum
Mett.
Local name: helecho trensillo de altura.
Antilles, Colombia, Venezuela, Ecuador, Peru, Bolivia. Brazil and Paraguay.
MO); D. Witsberger 687 (MHES); E.A. Montalvo 60 (ITIC); E. Sandoval & M.
Sandoval 1292 (B, LAGU); R. Seiler 827, 911 (ITIC, MHES); 962 (F, MHES); 963
(MHES); W. Berendsohn & M. Sandoval 1633 (LAGU); W. Berendsohn et al. 1519 (B,
LAGU). Sonsonate: R. Seiler 183 (F); 617 (MHES). Santa Ana: A. del Cid 1618
(ITIC); J. Monterrosa & R. Martínez 949 (B, LAGU, MO, NY); L. González 1617
(ITIC); M.L. Reyna de Aguilar 816 (MHES); R. Seiler 83, 183 (MHES); S. Winkler 47
(ITIC). San Salvador: R. Seiler 213 (ITIC, MHES). Chalatenango: R. Seiler 384
(JBL00083), 414 (JBL00084) (F, LAGU, MHES). San Vicente: R. Seiler 319 (MHES);
330 (F, MHES). Usulután: A. del Cid s.n. (ITIC); A.K. Monro, et al. 3010 (B, BM,
ITIC, LAGU, MO); D. Williams 52, 137 (B, LAGU, MO); D. Williams & R.W. Herrera
299 (LAGU); E.A. Montalvo 5033, 5045 (ITIC). Morazán: R. Seiler 975 (F, ITIC,
MHES); 1018, 1036 (MHES).
Asplenium feei Kunze ex Fée, Mém. Foug. 5: 194. 1852.
Asplenium sanguinolentum Kunze ex Mett., A. sarcodes Maxon
Antilles, Colombia, Venezuela, Suriname, Brazil, Bolivia.
Salvadoran distribution: San Salvador: M. Carlson 428a (F).
Asplenium flabellulatum Kunze, Linnea 9: 71. 1834.
Asplenium flabellulatum Kunze var. dentatum Klotzsch, A. radicans L. var dentatum
(Klotzsch) Bonap., A. flabellulatum Kunze var. partitum Klotzsch, A. partitum
(Klotzsch) C. Chr., A. radicans L. var partitum (Klotzsch) Hieron
Antilles, Colombia, Venezuela, Ecuador, Peru. Paraguay.
Salvadoran distribution: Santa Ana: M.L. Reyna de Aguilar 824 (MHES).
Chalatenango: R. Seiler 406 (JBL00086) (F, LAGU, MHES).
Asplenium formosum Willd., Sp. Pl., ed. 4, 5(1): 329.1810.
Asplenium formosum Willd. var. incultrum H. Christ, A. subalatum Hook. & Arn.
Panama, Antilles, Colombia, Venezuela, Guayana, French Guyana, Ecuador, Peru,
Brazil, Bolivia, Paraguay, Argentina, Africa, S India, Sri Lanka.
Local names: helecho peine de niño, helecho peinetero, helecho de rosquilla.
Salvadoran distribution: Ahuachapán: E. Sandoval & F. Chinchilla 99, 541 (B,
LAGU); E. Sandoval & F. Pérez 1264 (B, LAGU); R. Seiler 927 (F, MHES); R.
Villacorta et al. 939 (B, LAGU); W. Berendsohn et al. 1401, 1432, 1611 (B, LAGU);
W. Berendsohn et al. 1453 (LAGU). Sonsonate: R. Seiler 76 (F, MHES). Santa Ana:
B. Pfeiffer-Berendsohn 18 (B, LAGU, MO); M. Magaña 28 (MP-00028) (LAGU); S.
Winkler 14 (ITIC). La Libertad: E.A. Montalvo 6002 (ITIC); R. Seiler 713 (F, MHES).
San Salvador: R. Seiler 49 (MHES). Chalatenango: J. Lagos 1720 (ITIC); L. Lara 234
(MHES); R. Seiler 146 (F, MHES); 372 (F, ITIC, MHES). Usulután: A. del Cid 1031
(ITIC); A.K. Monro et al. 3014 (B, BM, ITIC, LAGU, MO); D. Williams & R.W.
Herrera 315 (LAGU, MO); D. Williams 503 (LAGU); L. González 1085 (ITIC).
Cabañas: A.K. Monro et al. 3748 (B, BM, ITIC, LAGU, MO). Morazán: J.
Monterrosa & C. Rivera 676 (B, BM, LAGU, MO); R. Seiler 1089 (F, MHES); S.
Winkler 5 (ITIC).
Asplenium fragrans Sw., Prodr. 130. 1788.
Asplenium auritum Sw. var. tripinnatum E. Fourn., A. cuspidatum Lam. var.
tripinnatum (E. Fourn.) C.V. Morton & Lellinger, A. foeniculaceum Kunth, A.
cuspidatum Lam. var. foeniculaceum (Kunth) C.V. Morton & Lellinger, A. coriifolium
Liebm.
143
FERN GAZ. 18(4): 120-215. 2008
Antilles, Colombia, Venezuela, Ecuador, Peru, Bolivia.
Salvadoran distribution: Chalatenango: R. Seiler 151 (F, MHES); 422 (JBL00087) (F,
ITIC, LAGU, MHES).
Asplenium harpeodes Kunze, Linnea 18: 329.1844.
Asplenium erectum Bory var. harpeodes (Kunze) Mett., A. pendulum Fée, A. donnellsmithii Christ . Local names: helecho llorón, quetzal silvestre.
Antilles, Colombia, Venezuela, Guayana, French Guyana, Ecuador, Peru, Brazil,
Bolivia.
Salvadoran distribution: Santa Ana: A. Molina et al. 16802 (F); E.A. Montalvo 15b,
19 (ITIC); J. Monterrosa et al. 221 (B, BM, LAGU, MO); M.L. Reyna de Aguilar 1458
(LAGU); R. Seiler 120 (F, ITIC, MHES); R. Villacorta et al. 1039, 1281 (B, LAGU,
MO); S. Winkler 2 (ITIC). Chalatenango: J. Tucker 1105 (F). Morazán: J.L. Linares
& A. Pool 2285 (JBL01930 ) (LAGU); J.L. Linares & C. Peña H. 2749 (JBL01928)
(LAGU).
Asplenium hoffmannii Hieron., Hedwigia 60: 258. 1918.
Asplenium membraniifolium Maxon
Local names: helecho de roca.
Colombia, Venezuela, Trinidad and Tobago.
Salvadoran distribution: Ahuachapán: J.M. Rosales 1104 (B, LAGU, MO); J.M.
Rosales 1233 (AAU, B, BM, LAGU, MO). Sonsonate: D. Rodríguez & M. Trejo 98
(B, BM, LAGU, MO), NY. La Libertad: R. Seiler 718 (F, MHES).
Asplenium laetum Sw., Syn. Fil. 79, 271. 1806.
Asplenium lugubre Liebm.
Panama, Antilles, Colombia, Venezuela, Trinidad and Tobago, French Guyana,
Ecuador, Peru, Brazil, Bolivia, Paraguay.
Salvadoran distribution: Ahuachapán: M. Sandoval & E. Sandoval 297 (B, F, LAGU).
Asplenium miradorense Liebm., Mexic. Bregn. 243. 1849.
Asplenium bradeorum Hieron., A. schlechtendahlianum Hieron., A. standleyi Maxon
Colombia, y Brazil.
Salvadoran distribution: Sonsonate: A. Molina & E.A. Montalvo 21777 (JBL01478)
(LAGU). Morazán: R. Seiler 1038 (F, MHES).
Asplenium monanthes L., Mant. Pl. 130.1767.
Asplenium monanthemum L., A. arcuatum Liebm., A. obtusissimum Fée, A. galeotti
Fée, A. leptophyllum Fée, A. curvatum Fée
Global distribution: S USA, Mexico, Guatemala, Honduras, Nicaragua, Costa Rica,
Panama, Antilles, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, Uruguay, Chile,
Argentina, Africa, Hawaii.
Salvadoran distribution: Ahuachapán: F. Wunderling 1226 (ITIC); Santa Ana: B.
Pfeiffer-Berendsohn 80 (LAGU, MO); G. Cerén & D. Rodríguez s.n. (JBL04646)
(LAGU); J. Monterrosa et al. 209, 908 (LAGU); J. Monterrosa et al. 931 (B, LAGU,
MO, NY); M. B. Siu s.n. (ITIC); M.C. Carlson 724 (F); M.L. Reyna de Aguilar 673
(MHES); R. Seiler 313 (F, MHES). San Salvador: R. Seiler 210 (F, MHES); 516
(MHES). Chalatenango: R. Seiler 387 (JBL00088) (F, LAGU, MHES); R. Villacorta
et al. 2073 (B, LAGU, MO); W. Berendsohn et al. 1572 (B, LAGU, MO). San Vicente:
R. Seiler 331 (JBL04807) (F, LAGU, MHES).
♦ Asplenium monodon Liebm., Mexic. Bregn. 247. 1849.
Asplenium auritum Sw. var. monodon (Liebm.) E. Fourn.
Global distribution: USA, Mexico, Guatemala, Honduras, Costa Rica, Panama,
Antilles, Venezuela, Colombia, Trinidad and Tobago, Ecuador.
Salvadoran distribution: Santa Ana: R. Seiler 466 (MHES).
Asplenium polyphyllum Bertol., Novi Comment. Acad. Sci. Inst. Bononiensis 4: 443.
1840.
Asplenium polymeris T. Moore, A. bertolonii Donn. Sm., A. trichomanes L. var.
viridissimum Christ, A. viridissimum J. Bommer ex Christ
Venezuela, Ecuador, Peru, Bolivia.
Salvadoran distribution: Santa Ana: B. Pfeiffer-Berendsohn 41 (LAGU, MO); J.
Monterrosa & R. Carballo 252 (B, BM, LAGU, MO); J. Monterrosa & R. Carballo 837
(B, LAGU, MO); M.L. Reyna de Aguilar 764 (MHES); R. Seiler 464 (F, MHES); R.
Villacorta & L. Lara 2663 (B, LAGU, MO); V.M. Martínez s.n. (CMC01046) (B,
LAGU, MO); L. Lara 238 (MHES). Chalatenango: R. Seiler 160 (F, ITIC, MHES).
Asplenium potosinum Hieron., Hedwigia 60: 247.1918.
Asplenium regulare Sw. var. latior E. Fourn., A. lunulatum Sw. var. latius E. Fourn., A.
hesperium Mickel & Beitel
Local names: helecho cola de pavo real.
Salvadoran distribution: Ahuachapán: E. Sandoval & F. Chinchilla 40 (LAGU, MO);
R. Seiler 824 (JBL04817) (F, ITIC, LAGU, MHES); R. Seiler 903 (F, ITIC, MHES);
955 (F, MHES); W. Berendsohn & E. Sandoval 1513 (B, LAGU). Usulután: A.K.
Monro et al. 3008, 3015 (B, BM, ITIC, LAGU, MO); D. Williams 53 (LAGU, MO); D.
Williams 413 (LAGU).
Asplenium praemorsum Sw., Prodr. 130. 1788.
Salvadoran distribution: Ahuachapán: M. Magaña 33 (MP-00018) (LAGU); R. Seiler
819 (F, MHES). Morazán: R. Seiler 1035 (F); 1072 (F, MHES).
Asplenium pulchellum Raddi, Opusc. Sci. 3: 291.1819.
Asplenium pulchellum Raddi var. subhorizontale Rosenst.
Global distribution: Mexico, Honduras, Nicaragua, Costa Rica, Panama, Colombia,
Venezuela, Ecuador, Peru, Brazil, Bolivia, Paraguay.
Source: Seiler (1982): 387; Berendsohn (1989): 5; Mickel & Smith (2004): 114.
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FERN GAZ. 18(4): 120-215. 2008
Asplenium pumilum Sw., Prodr. 129. 1788.
Asplenium anthriscifolium Jacq., A. leucothrix Maxon, Athyrium verapax Christ
Rica, Panama, Antilles, Colombia, Venezuela, Trinidad and Tobago, Ecuador, Peru,
Brazil, Bolivia, Africa.
Salvadoran distribution: Sonsonate: M.L. Reyna de Aguilar 1453 (LAGU); R. Seiler
74 (MHES); 495 (F, MHES). Santa Ana: R. Seiler 703 (MHES). San Salvador: R.
Seiler 582 (F, MHES). Usulután: A. del Cid 1785 (ITIC); J. Lagos 1786 (ITIC).
Asplenium seileri C.D. Adams, Novon 2: 291.1992.
Global distribution: Mexico, Guatemala, Costa Rica.
Salvadoran distribution: Ahuachapán: A. Molina et al. 21797 (ITIC); G. Davidse et al.
37372 (B, BM, ITIC, LAGU, MO); R. Seiler 957, (F, MHES). Sonsonate: R. Villacorta
751 (LAGU).
Asplenium serra Langsd. & Fischer, Pl. Voy. Russes Monde I: 16.1810.
Asplenium serra Langsd. & Fischer var. incurvatum H. Christ, A. insigne Liebm., A.
progrediens Fée
Bolivia, Paraguay, Uruguay.
Salvadoran distribution: Santa Ana: A. Molina et al. 16755 (F); J. Monterrosa & R.
Carballo 860 (B, LAGU, MEXU, MO); R. Seiler 106, 191b (F, MHES); 114 (MHES);
192 (F); V.M. Martínez s.n. (CMC01095) (B, EAP, LAGU, MO). Chalatenango: J.
Tucker 1045 (F); R. Seiler 424 (F, MHES); W. Berendsohn et al. 1574 (B, LAGU, MO).
Asplenium sessilifolium Desv. var. sessilifolium, Ges. Naturf. Freunde Berlin Mag.
Neuesten Entdeck. Gesammten Naturk. 5: 322.1811.
Asplenium sessilifolium Desv. var. guatemalense Hieron.
Salvadoran distribution: Sonsonate: R. Seiler 625 (F, MHES). Santa Ana: A. Molina
& E.A. Montalvo 21512 (JBL01516) (F, LAGU); B. Pfeiffer-Berendsohn 55 (LAGU,
MO); J. Monterrosa & R. Martínez 936 (B, LAGU, MO, NY); R. Seiler 121
(JBL04803) (F, LAGU, MHES); R. Seiler 13 (ITIC, MHES); 84 (ITIC); 354 (F, ITIC,
MHES). Chalatenango: L. Lara 233 (MHES); R. Seiler 378 (JBL00085) (F, LAGU,
MHES).
Asplenium tenerrimum Mett. ex Kuhn, Linnea 36: 97. 1869.
Asplenium cuspidatum Lam. var. tenerrimum (Mett. ex Kuhn) C.V. Morton & Lellinger,
A. fournieri Kuhn ex E. Fourn.
Global distribution: Mexico, Honduras, Nicaragua, Colombia, Venezuela, Ecuador.
Salvadoran distribution: Santa Ana: R. Seiler 545 (F).
♦Asplenium theciferum (Kunth) Mett., Ann. Sci. Nat. Bot. sér. 5, 2: 227. 1864.
Davallia thecifera Kunth, D. lindenii Hook., Darea thecifera (Kunth) E. Fourn.,
Loxoscaphe lindenii (Hook.) T. Moore, L. thecifera (Kunth) T. Moore
Global distribution: Mexico, Honduras, Nicaragua, Costa Rica, Panama, Antilles,
Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, Africa.
Salvadoran distribution: Santa Ana: J.L. Linares 391 (JBL01410) (LAGU).
Asplenium virillae Christ, Bull. Herb. Boissier, sér. 2, 4: 1090. 1904.
Asplenium alfredii Rosenst.
Venezuela.
Salvadoran distribution: Sin loc: P. Standley 19774, US (fide Adams, 1995). Santa
Ana: J. Monterrosa et al. 1457 (BM, LAGU, MO, NY): 130.
BLECHNACEAE
Blechnum appendiculatum Willd., Sp. Pl., ed. 4, 5(I): 410. 1810.
Blechnum glandulosum Kaulf. ex Link, B. meridionale C. Presl, B. occidentale L. var.
minor Hook., B. occidentale L. var. pubirhachis Rosenst.
Local names: helecho esqueleto de pescado, petatillo.
Bolivia, Paraguay, Argentina.
MO); F. Pérez s.n. (ISB00003) (B, LAGU, MO); G. Davidse et al. 37417 (B, BM, ITIC,
LAGU, MO); J.G. Sandoval et al. s.n. (ISB00027) (B, LAGU, MO), US; K. Sidwell et
al. 814 (B, BM, ITIC, LAGU, MO); S. Castillo s.n. (ISF00594) (B, F, LAGU, MO).
Sonsonate: R. Villacorta & A. González 693 (B, LAGU, MO). Santa Ana: B. PfeifferBerendsohn 65 (LAGU, MO); D. Rodríguez & G. Cerén s.n. (JBL04618) (LAGU); E.A.
Montalvo 3654 (ITIC); J. Monterrosa et al. 907 (B, LAGU, MO, NY); M.B. Siu s.n.
(ITIC); M.L. Reyna de Aguilar 684 (MHES); R. Seiler 349, 477 (F, MHES); V.M.
Martínez s.n. (CMC01096) (B, EAP, LAGU, MO). La Libertad: A. Mangandi et al.
s.n. (JBL04492) (LAGU); A.K. Monro et al. 2276, 3086 (B, BM, ITIC, LAGU, MO);
J.C. González 415 (B, LAGU); M. Renderos et al. 412 (LAGU). San Salvador: A.K.
Monro et al. 2217, 2223 (B, BM, ITIC, LAGU, MO, L).O. Williams et al. 15232 (F);
M.C. Carlson 329 (F); R. Seiler 29 (F, MHES). Chalatenango: D. Sloot & A. Reina
216 (MAG00645), 433 (MAG00862) (LAGU, U); L. Lara 218, 225 (MHES); M.
Renderos & R. Villacorta 141 (B, LAGU); R. Seiler 413 (JBL00091) (LAGU, MHES);
W. Heed 26 (ITIC). San Vicente: E.A. Montalvo 4615, 4618 (ITIC). Usulután: L.
González 1089, 1787 (ITIC). San Miguel: K. Sidwell et al. 885 (B, BM, ITIC, LAGU,
MO); M. Renderos et al. 687 (LAGU, MO). Cuscatlán: S. Dar s.n. (JBL03666)
(LAGU, MO). Morazán: J. Monterrosa & M. Rivera 615 (B, BM, LAGU); J. Tucker
608 (F).
Blechnum appendiculatum Willd. x Blechnum gracile Kaulf
Global distribution: Mexico, Honduras, Nicaragua, Colombia, Venezuela.
Salvadoran distribution: Santa Ana: R. Seiler 763 (MHES). Chalatenango: L. Lara
209 (MHES).
Blechnum x caudatum Cav., Descr. Pl. 262. 1802.
Panama, Colombia, Venezuela, Suriname, Ecuador, Peru, Brazil.
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FERN GAZ. 18(4): 120-215. 2008
Salvadoran distribution: Santa Ana: B. Pfeiffer-Berendsohn 59 (B, LAGU, MO); C.
Salazar s.n. (ITIC). Morazán: A.K. Monro et al. 3804 (B, BM, ITIC, LAGU, MO); J.
Monterrosa & R. Carballo 794 (LAGU).
Note: Blechnum x caudatum is a hybrid between Blechnum gracile y B. occidentale
(Mickel & Smith 2004: 146, Fig.: 55C).
Blechnum divergens (Kunze) Mett., Ann. Sci. Nat. Bot., sér. 5, 2: 255.1864.
Lomaria divergens Kunze, L. exaltata Fée, Struthiopteris exaltata (Fée) Broadh.
Salvadoran distribution: Santa Ana: R. Seiler 765, 892 (F, MHES); 1156, F (ITIC).
Blechnum ensiforme (Liebm.) C.Chr., Index Filic. 153: 1905.
Lomaria ensiformis Liebm., Struthiopteris ensiformis (Liebm.) Maxon
Panama, Colombia, Venezuela, Guayana, Ecuador, Peru, Bolivia.
Salvadoran distribution: Santa Ana: G. Cerén & D. Rodríguez s.n. (JBL04660)
(LAGU); R. Seiler 299 (F, MHES). Chalatenango: R. Seiler 389 (JBL00092) (F,
LAGU, MHES). Morazán: R. Seiler 1026 (F, ITIC, MHES).
Blechnum falciforme (Liebm.) C. Chr., Index Filic. 154. 1905.
Lomaria falciformis Liebm., L. deflexa Liebm., Struthiopteris falciformis (Liebm.)
Broadh.
Ecuador.
Salvadoran distribution: Sonsonate: R. Seiler 632 (F, MHES). Santa Ana: A.
Hernández s.n. (ITIC); M.C. Carlson 725, 891, 952 (F); R. Seiler 348, 478 (MHES);
863 (F, ITIC, MHES). Chalatenango: N. Ventura 5026 (ITIC); R. Seiler 409 (MHES).
Morazán: R. Seiler 990 (F, MHES).
Blechnum fragile (Liebm.) C.V. Morton & Lellinger, Amer. Fern J. 57: 68. 1967.
Lomaria fragilis Liebm., L. polypodioides (Sw.) Desv., L. onocleoides (Sw.) Spreng., L.
mexicana Fée, Blechnum binervatum (Poir.) C.V. Morton & Lellinger subsp. fragile
(Liebm.) R.M. Tryon & Stolze, B. onocleoides Sw., B. polypodioides (Sw.) Kuhn,
Osmunda polypodioides Sw.
Panama, Antilles, Colombia, Venezuela, Trinidad and Tobago, Ecuador, Peru, Bolivia.
Salvadoran distribution: Santa Ana: R. Seiler 654 (F, MHES). Morazán: R. Seiler
1041 (F, ITIC, MHES).
Blechnum gracile Kaulf., Enum. Filic. 158. 1824.
Blechnum intermedium Link., B. subdimorphum Copel., B. fraxineum auct., non Willd.
French Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina.
Salvadoran distribution: Chalatenango: M.B. Siu s.n. (ITIC); M.C. Carlson 594 (F);
R. Seiler 572 (F, ITIC); (MHES). Cabañas: A.K. Monro et al. 3746 (B, BM, ITIC,
LAGU, MO). Morazán: R. Seiler 1083 (F, ITIC, MHES).
Blechnum lehmannii Hieron., Bot. Jahsb. Syst. 34: 473. 1904.
Blechnum l´herminieri (Bory) C. Chr. subsp. lehmannii (Hieron) Lellinger,
Struthiopteris maxonii Broadh.
Salvadoran distribution: Sonsonate: J.L. Linares & C.A. Martínez 3033 (JBL01933)
(LAGU). Santa Ana: F. Wunderling 1245 (ITIC); G. Cerén & D. Rodríguez s.n.
(JBL04650) (LAGU); R. Seiler 894 (F, ITIC, MHES). Morazán: R. Seiler 979, 1039,F
(ITIC, MHES).
Blechnum occidentale L., Sp. Pl. 2: 1077. 1753.
Local names: helecho esqueleto de pescado.
Salvadoran distribution: Ahuachapán: A.K. Monro et al. 1929, 2984 (B, BM, ITIC,
LAGU, MO); A. Sermeño 98 (JBL00999) (B, LAGU, MEXU, MO); E. Sandoval & M.
Sandoval 864 (B, LAGU); J.M. Rosales 1188 (B, LAGU, MO); 1715, 1723 (B, BM,
LAGU, MO); M. Magaña 19 (MP-00032) (LAGU); M. Sandoval & E. Sandoval 136
(B, F, LAGU); R. Villacorta 622 (B, LAGU, MO); S. Castillo & B.A. Martínez s.n.
(ISF00406) (AAU, F, LAGU); W. Berendsohn et al. 1316, 1368, 1382, 1393 (B,
LAGU, MO). Sonsonate: D. Rodríguez & G. Trejo 203 (LAGU). Santa Ana: G.
Davidse et al. 37226 (B, BM, ITIC, LAGU, MO); J. Monterrosa & R. Carballo 509
(LAGU); K. Prestegard 10 (MP-00031) (LAGU); M.L. Reyna de Aguilar et al. 1439 (B,
ITIC, LAGU, MO). La Libertad: F. Wunderling 461 (ITIC); N. Ventura 5001, 5004
(ITIC); O. Tejada et al. s.n. (JBL04504) (LAGU); R.Guerra et al. s.n. (JBL04502)
(LAGU); R.Menjivar & J. Sermeño s.n. (JBL04483) (LAGU). San Salvador: M.C.
Carlson 4a (ITIC); R. Seiler 457 (F, MHES); 783 (F, ITIC, MHES). Chalatenango:
E.A. Montalvo 30, 3202, 3778 (ITIC); L. Lara 498 (MHES); M.B. Siu s.n. (ITIC).
Usulután: D. Williams 54 (LAGU, MO). San Miguel: A.K. Monro et al. 2944 (B, BM,
ITIC, LAGU, MO); A. Salgado 66 (MHES). Cuscatlán: E.A. Montalvo 3312 (ITIC).
Cabañas: A.K. Monro & K. Sidwell 2778 (B, BM, ITIC, LAGU, MO); G. Davidse et
al. 37083 (B, BM, ITIC, LAGU, MO). Morazán: J. Monterrosa et al. 45 (B, LAGU,
MO).
Blechnum polypodioides Raddi, Opusc. Sci. 3: 294.1819, non (Sw.) Kuhn, 1868.
Asplenium blechnoides Lag. ex Sw., Blechnum unilaterale Sw., B. blechnoides (Lag. ex
Sw.) C. Chr., B. scabrum Liebm., B. angustifrons Fée
Local names: helecho peineta amarilla.
Salvadoran distribution: Ahuachapán: W. Berendsohn et al. 1469 (LAGU).
Chalatenango: M.B. Siu s.n. (ITIC); L. Lara 230 (MHES); R. Seiler 564 (MHES); 571
(F, MHES). Usulután: D. Williams 55 (LAGU, MO). Morazán: J. Monterrosa & C.
Rivera 690 (B, BM, LAGU); J. Tucker 643 (F).
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FERN GAZ. 18(4): 120-215. 2008
Blechnum schiedeanum (Schltdl. ex C. Presl) Hieron., Hedwigia 47: 239. 1908.
Lomaria schiedeana Schltdl. ex C. Presl, L. spectabilis Liebm., L. longifolia Schltdl. ex
M. Martens & Galeotti, L. danaeacea Kunze, L. acrodonta Fée, L. varians E. Fourn.,
L. costaricensis H. Christ, Struthiopteris costaricensis (H. Christ) Broadh., Blechnum
danaeaceum (Kunze) Christ, B. varians (E. Fourn.) C. Chr., B. costaricensis (H. Christ)
C. Chr.
Salvadoran distribution: Ahuachapán: B. Pfeiffer-Berendsohn 51 (B, LAGU, MO); 53
(B, LAGU); J. Monterrosa & R. Carballo 262 (B, LAGU, MO); R. Seiler 196, 681 (F,
MHES). Santa Ana: A. Hernández s.n. (ITIC). Morazán: R. Seiler 1034 (F, ITIC,
Blechnum serrulatum Rich., Actes Soc. Hist. Nat. Paris I: 114. 1792.
Suriname, French Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina,
Australia.
Sources: Calderón & Standley (1941): 30; Seiler (1982): 389; Berendsohn (1989), 1: 6;
Moran in Davidse et al. (1995): 330; Mickel & Smith (2004): 151.
Blechnum stoloniferum (Mett. ex E. Fourn.) C. Chr., Index Filic. 154. 1905.
Lomaria stolonifera Mett. ex E. Fourn., L. ghiesbreghtii Baker, Struthiopteris
stolonifera (Mett. ex E. Forun.) Broadh., Blechnum ghiesbreghtii (Baker) C. Chr.
Global distribution: Mexico, Guatemala, Honduras, Costa Rica.
Salvadoran distribution: Santa Ana: R. Seiler 678 (F); 780 (MHES). Chalatenango:
R. Seiler 731 (F, MHES).
Blechnum wardiae Mickel & Beitel, Mem. New York Bot. Gard. 46: 89. 1988.
Global distribution: Mexico, Nicaragua, Costa Rica, Panama, Colombia, Ecuador.
Salvadoran distribution: Santa Ana: A. Molina et al. 16837 (F); E.A. Montalvo 17
(ITIC).
Woodwardia semicordata Mickel & Beitel, Mem. New York Bot. Gard. 46: 403.
1988, pro hybr.
Global distribution: Mexico, Costa Rica.
Source: Mickel & Smith (2004): 694.
Woodwardia spinulosa M. Martens & Galeotti, Mém. Foug. Mexique 64. 1842.
Global distribution: Mexico, Guatemala, Honduras, Nicaragua.
Salvadoran distribution: Santa Ana: G. Davidse et al. 37216 (B, BM, ITIC, LAGU,
MO); V.M. Martínez s.n. (CMC00979) (B, LAGU, MO); s.n. (CMC01024) (B, BM,
EAP, LAGU, MO); R. Seiler 471 (F, MHES); W. Berendsohn et al. 1679 (LAGU).
Chalatenango: R. Seiler 727 (MHES). Morazán: R. Seiler 1002 (F); 1027 (F, MHES).
CIBOTIACEAE
Cibotium regale Verschaff. & Lem., Ill. Hort. 15: under pl. 548 (F). 1868.
Cibotium guatemalense Rchb. f. ex Kuhn, C. wendlandii Mett. ex Kuhn
Local names: cabeza de niño.
Global distribution: Mexico, Guatemala, Belize, Honduras.
Salvadoran distribution: R. Seiler 151, MO (fide B. Pérez-García in Davidse et al.
1995). Santa Ana: R. Seiler 186 (MHES); 304 (F, ITIC, MHES). La Libertad: K.
Prestegard 36 (MP-00025) (LAGU). Usulután: D. Williams & R.W. Herrera 471 (B,
LAGU, MEXU, MO).
Note: listed as ‘at risk’ by MARN, 2004.
CULCITACEAE
Culcita coniifolia (Hook.) Maxon, Annual Rep. Board Regents Smithsonian Inst. 1911:
488, t. 13C. 1912.
Dicksonia coniifolia Hook.
Salvadoran distribution: Molina R. 12682, GH (Pérez-García in Davidse et al. 1995).
Santa Ana: A. Molina et al. 12682 (F); R. Seiler 485 (F, ITIC, MHES); 1129 (F, ITIC);
L. Lara 279 (MHES).
Note: listed as ‘at risk’ by MARN, 2004.
CYATHEACEAE
Alsophila firma (Baker) D. S. Conant, J. Arnold Arbor. 64: 372. 1983.
Cyathea firma (Baker) Domin, C. mexicana Schltdl. & Cham., C. trejoi Christ, C.
articulata Fée, Nephelea mexicana (Schltdl. & Cham.) R. M. Tryon
Panama, Ecuador.
Local names: helecho de espina, helecho cuerpo espín
Salvadoran distribution: Ahuachapán: M. Sandoval & E. Sandoval 9 (B, LAGU); W.
Berendsohn et al. 1455 (B, LAGU).
Alsophila salvinii Hook. in Hooker & Baker, Syn. Fil. 36. 1866.
Cyathea salvinii (Hook.) Domin, Alsophila muenchii Christ
Global distribution: Mexico, Guatemala, Belize, Honduras, Nicaragua, Peru.
Salvadoran distribution: R. Seiler 899, MO (Riba in Davidse et al. 1995). Santa Ana:
A. Molina et al. 16749 (F); R. Seiler 296 (F, ITIC, MHES).
Note: Listed as ‘threatened’ by MARN, 2004.
Alsophila tryoniana (Gastony) D.S. Conant, J. Arnold Arbor. 64: 371. 1983.
Nephelea tryoniana Gastony
Salvadoran distribution: R. Seiler 1154, MO (Riba in Davidse et al. 1995). Santa Ana:
R. Seiler 236, 300 (F, ITIC, MHES). Morazán: R. Seiler 976 (F, ITIC, MHES).
Note: Listed as ‘threatened’ by MARN, 2004.
Cyathea bicrenata Liebm., Mexic. Bregn. 289 (reprint 137). 1849.
Alsophila bicrenata (Liebm.) E. Fourn., A. scabriuscula Maxon, Trichipteris bicrenata
(Liebm.) R. M. Tryon, T. scabriuscula (Maxon) R. M. Tryon
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FERN GAZ. 18(4): 120-215. 2008
Salvadoran distribution: Santa Ana: R. Seiler 301 (F, ITIC, MHES).
Note: Listed as ‘at risk’ by MARN, 2004; listed as Apendix II by CITES.
Cyathea costaricensis (Mett. ex Kuhn) Domin, Acta Bot. Bohem. 9: 107. 1930.
Hemitelia costaricensis Mett. ex Kuhn, Trichipteris costaricensis (Mett. ex Kuhn)
Barrington . Local name: helecho pescuezo de violín.
Panama.
Salvadoran distribution: Ahuachapán: K. Prestegard 18 (MP-00026) (LAGU); R.
Seiler 929 (F, MHES); W. Berendsohn et al. 1352, 1414 (B, LAGU, MO). Sonsonate:
J. Tucker 1355 (F). Santa Ana: S. Winkler s.n. (ITIC). La Libertad: A.K. Monro et al.
2307 (B, BM, ITIC, LAGU, MO). San Salvador: R. Seiler 224, 368, 535, 787 (F,
MHES).
Note: Listed as ‘at risk’ MARN, 2004; listed as Apendix II by CITES.
Cyathea divergens Kunze var. divergens, Linnaea 9: 100. 1834
Alsophila subaspera H. Christ, C. divergens var. hirta I. Losch, C. divergens var. minor
Rosenst., C. pelliculosa H. Christ, C. subaspera (H. Christ) Domin.
Global distribution: Costa Rica, Panama, Colombia, Venezuela, Guayana, Suriname,
Ecuador, Peru.
Note: This species is cultivated at Los Planes, Montecristo national park (Pers. Obs.).
Cyathea divergens var. tuerckheimii (Maxon) R.M. Tryon, Contr. Gray Herb. 206: 56.
1976.
Cyathea tuerckheimii Maxon, Cyathea juergensenii E. Fourn.
Global distribution: Mexico, Belice, Guatemala, Nicaragua.
Salvadoran distribution: Santa Ana: J.Monterrosa & D.Rodríguez 1539 (LAGU); R.
Seiler 276 (MHES). Morazán: R. Seiler 980 (F, MHES).
Note: Listed as ‘at risk’ MARN, 2004; listed as Apendix II by CITES.
Cyathea fulva (M. Martens & Galeotti) Fée, Mém. Foug. 9: 34. 1857.
Alsophila fulva M. Martens & Galeotti, Cyathea delicatula Maxon, C. compersa H.
Christ, C. furfuracea H. Christ, C. mollis Rosenst., C. molliuscula Domin, C. onusta H.
Christ, C. papyracea H. Christ, C. underwoodii H. Christ
Colombia, Venezuela, Ecuador.
Note: Listed as Apendix II by CITES.
Sources: Mickel & Smith (2004): 237.
Cyathea godmanii (Hook.) Domin, Pterid. Dominica 262. 1929.
Alsophila godmanii Hook., A. mexicana Mart., Cyathea mexicana Schldtl. & Cham., C.
valdecrenata Domin, Trichipteris mexicana (Mart.) R.M. Tryon
Global distribution: Mexico, Guatemala, Honduras, Panama.
Note: Listed as ‘threatened’ by MARN, 2004; listed as Apendix II by CITES.
Cyathea schiedeana (C. Presl) Domin, Pterid. Dominica 263. 1929.
Alsophila schiedeana C. Presl, A. crassifolia H. Christ, A. chnoödes H. Christ, Cyathea
chnoödes (H. Christ) Domin, Trichipteris schiedeana (C. Presl) R.M. Tryon, T.
crassifolia (H. Christ) Gastony
Panama, Colombia.
Salvadoran distribution: Santa Ana: J. Monterrosa & R. Carballo 848 (LAGU); R.
Seiler 172, 237 (F, MHES). Chalatenango: J. Tucker 1129 (F); R. Seiler 760 (F,
MHES). Morazán: J. Tucker 788 (F); R. Seiler 1082 (F, ITIC, MHES).
Note: Listed as ‘threatened’ by MARN, 2004; listed as Apendix II by CITES.
Sphaeropteris horrida (Liebm.) R.M. Tryon, Contr. Gray Herb. 200: 20. 1970, non
Cyathea horrida (L.) Sm.
Cibotium horridum Liebm., C. princeps Linden ex J. Smith, Cyathea princeps E.
Mayer, C. bourgeaui E. Fourn., C. glauca E. Fourn., C. muenchii Christ
Global distribution: Mexico, Guatemala, Belize, Honduras, Nicaragua.
Salvadoran distribution: Santa Ana: R. Seiler 242 (MHES), F (ITIC).
Note: Listed as ‘at risk’ by MARN, 2004.
DENNSTAEDTIACEAE
Dennstaedtia bipinnata (Cav.) Maxon, Proc. Biol. Soc. Wash. 51: 39. 1938.
Dicksonia bipinnata Cav., D. adiantoides Humb. & Bonpl. ex Willd. Local name:
helecho palma de paraiso.
Rica, Panama, Colombia, Venezuela, Trinidad and Tobago, Ecuador, Peru, Bolivia.
Salvadoran distribution: P.Standley 19760 (GH) (Moran in Davidse et al. 1995).
Ahuachapán: E. Sandoval & F. Chinchilla 123 (LAGU); S. de Moore s.n. (12-909)
(MHES). San Salvador: R. Seiler 1214 (F).
Dennstaedtia cornuta (Kaulf.) Mett., Ann. Sci. Nat. Bot,, sér. 5, 2: 260. 1864.
Dicksonia cornuta Kaulf.
Panama, Antilles, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, Argentina.
Salvadoran distribution: Santa Ana: R. Seiler 165, 271, 693, 1192 (F, MHES).
Dennstaedtia distenta (Kunze) T. Moore, Index Fil. 306. 1861.
Dicksonia distenta Kunze
Global distribution: Mexico, Guatemala, Costa Rica, Panama, Antilles.
Salvadoran distribution: Standley & Valerio 49449, GH (Moran in Davidse et al. 1995).
Morazán: Tucker 726 (F).
♦ Dennstaedtia globulifera (Poir.) Hieron., Bot. Jahrb. Syst. 34: 455. 1904.
Polypodium globuliferum Poir., Dicksonia globulifera (Poir.) Kuntze, Dennstaedtia
gracilis A. Rojas & Tejero
Antilles, Colombia, Venezuela, Ecuador, Trinidad and Tobago, Ecuador, Peru, Brazil,
Bolivia, Paraguay, Argentina, Uruguay.
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FERN GAZ. 18(4): 120-215. 2008
Salvadoran distribution: San Salvador: S. Winkler 74 (ITIC).
♦ Dennstaedtia spinosa Mickel, Amer. Fern J. 58: 90. 1968.
Salvadoran distribution: Ahuachapán: W. Berendsohn et al. 1320 (B, LAGU, MO).
Note: This collection was cited as Dennstaedtia cicutaria (Moran in Davidse et al.
1995), however it posesses spines that are diagnostic for D. spinosa.
Histiopteris incisa (Thunb.) J. Sm., Hist. Fil. 295. 1875.
Pteris incisa Thunb., Litobrochia incisa (Thunb.) C. Presl
Antilles, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, Africa, Asia.
Salvadoran distribution: R. Seiler 1195 (UC) (Moran in Davidse et al. 1995). Santa
Ana: A. Molina et al. 12534 (F); R. Seiler 253 (F, MHES). Chalatenango: J. Tucker
1122 (F).
Hypolepis blepharochlaena Mickel & Beitel, Mem. New York Bot. Gard. 46: 219.
1988.
Salvadoran distribution: Santa Ana: A. Molina et al. 12528, 16775, 16885 (F); R.
Seiler 249, 294 (F, MHES). Chalatenango: R. Seiler 427 (JBL00108) (LAGU,
MHES).
Hypolepis bogotensis H. Karst., Fl. Columb. 2: 91. 1865.
Dennstaedtia munchii H. Christ, Hypolepis flexuosa Sodiro
Global distribution: Mexico, Honduras, Costa Rica, Panama, Antilles, Colombia,
Salvadoran distribution: Chalatenango: Tucker 1121 (F).
Hypolepis nigrescens Hook., Sp. Fil. 2: 66, t. 90C. 1852.
Salvadoran distribution: Santa Ana: J.Monterrosa & D.Rodríguez 1532 (LAGU)
Hypolepis repens (L.) C. Presl, Suppl. Tent. Pterid. 162. 1836.
Lonchitis repens L., Cheilanthes repens (L.) Kaulf., Hypolepis mexicana Liebm.
Salvadoran distribution: Santa Ana: Tucker 1303 (F, LAGU (photo), NY).
Pteridium aquilinum var. feei (W. Schaffn. ex Fée) Maxon ex Yunck., Publ. Field Mus.
Nat. Hist., Bot. Ser. 17: 308. 1938.
Pteris feei W. Schaff. ex Fée, Pteridium feei (W. Schaffn. ex Fée) Faull
Salvadoran distribution: Santa Ana: R. Seiler 777 (F, MHES). Chalatenango: R.
Seiler 417 (JBL00109) (F, LAGU, MHES). Morazán: J. Tucker 696 (F); R. Seiler 1058
(F, MHES).
Note: An invasive species that has been responsible for the poisoning of cattle
throughout the country (Ministerio de Ganaderia, Pers. Comm., 2005).
Pteridium arachnoideum (Kaulf.) Maxon, J. Wash. Acad. Sci. 14: 89. 1924.
Pteris arachnoidea Kaulf., P. aquilina L. var. arachnoidea (Kaulf.) D. C. Eaton,
Pteridium aquilinum (L.) Kuhn var. arachnoideum (Kaulf.) Brade, P. caudatum (L.)
Maxon subsp. arachnoideum (Kaulf.) Lellinger .
Local name: helecho petatillo
Antilles, Colombia, Venezuela, Trinidad and Tobago, Guayana, Suriname, French
Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay.
Salvadoran distribution: Ahuachapán: D. Witsberger 665 (MHES); E. Sandoval & F.
Chinchilla 429 (B, LAGU); R. Seiler 665 (MHES). Santa Ana: R. Seiler 306 (F,
MHES). Chalatenango: D. Sloot & A. Reina 310 (MAG00739) (LAGU, U); R. Seiler
266 (F, MHES). San Vicente: R. Seiler 333 (MHES). Morazán: R. Seiler 1093 (F);
1094 (F, MHES).
Pteridium caudatum (L.) Maxon, Proc. U.S. Natl. Mus. 23: 631. 1901.
Pteris caudata L., P. aquilina L. var. caudata (L.) Link, P. aquilina L. var. mexicana
Fée, P. caudata L. var. mexicana Fée, Pteridium aquilinum (L.) Kuhn var. caudatum
(L.) Sadeb., P. aquilinum subsp. caudatum (L.) Bonap.
Suriname, French Guyana, Ecuador, Peru.
Salvadoran distribution: Santa Ana: J. Monterrosa & R. Carballo 251 (B, LAGU,
MO); J. Tucker 1298 (F). La Libertad: J.C. González 372 (B, LAGU, MO); J. Lagos
s.n. (ITIC). San Salvador: M.C. Carlson 404, 418 (F); S. Calderón 461 (F).
Chalatenango: M.C. Carlson 611 (F); R. Villacorta & A. Araniva 981 (B, LAGU, MO).
Usulután: D. Williams & R.W. Herrera 451 (LAGU). Morazán: J. Tucker 697 (F).
DICKSONIACEAE
Dicksonia sellowiana Hook., Sp. Fil. I: 67. 1844.
Dicksonia gigantea H. Karst., D. ghiesbreghtii Maxon
Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, Uruguay.
Salvadoran distribution: Santa Ana: A. Molina et al. 12523, 16757, 16915, 16917 (F);
M.C. Carlson 964 (F); R. Seiler 240 (F, ITIC, MHES); S. Winkler 100 (ITIC).
Chalatenango: J. Tucker 1127 (F, MO) (Pérez-García in Davidse et al. 1995); R. Seiler
416 (JBL00094), (JBL00095) (F, LAGU, MHES).
Note: Note: listed as ‘at risk’ by MARN, 2004; listed as Apendix II by CITES.
Lophosoria quadripinnata (J. F. Gmel.) C. Chr. in Skottsberg, Nat. Hist. Juan
Fernández I: 16. 1920.
Polypodium glaucum Sw., P. quadripinnatum J. F. Gmel., P. pruinatum Sw., Alsophila
quadripinnata (J. F. Gmel.) C. Chr., A. pruinata (Sw.) Kaulf. ex Kunze, Trichosorus
glaucescens Liebm., T. densus Liebm., T. frigidus Liebm.
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FERN GAZ. 18(4): 120-215. 2008
Antilles, Colombia, Venezuela, Trinidad and Tobago, Ecuador, Peru, Bolivia, Chile,
Argentina.
Salvadoran distribution: Santa Ana: A. Molina et al. 16773 (F); G. Cerén & D.
Rodríguez s.n. (JBL04655) (B, LAGU); s.n. (JBL04656), s.n. (JBL04657) (LAGU); R.
Seiler 292 (F, ITIC, MHES). Chalatenango: J. Tucker 1150 (F). Morazán: R. Seiler
1030 (F, MHES).
Note: Listed as ‘threatened’ by MARN.
DRYOPTERIDACEAE
Arachniodes denticulata (Sw.) Ching, Acta Bot. Sin. 10: 260. 1962.
Polypodium denticulatum Sw., Aspidium denticulatum (Sw.) Sw., A. jucundum Fée,
Polystichum denticulatum (Sw.) J. Sm., Dryopteris denticulata (Sw.) Kuntze, D.
denticulata f. jucunda (Fée) C. Chr., Rumohra denticulata (Sw.) Copel., Byrsopteris
denticulata (Sw.) C.V. Morton
Panama, Antilles, Colombia, Venezuela, Guayana, Ecuador, Peru, Brazil, Bolivia.
et al. 12497 (F); F. Wunderlin 543 (ITIC); G. Cerén & D. Rodríguez s.n. (JBL04653)
(B, LAGU, MO); J. Monterrosa et al. 204 (AAU, B, BM, INB, LAGU, MEXU, MO);
M.C. Carlson 953 (F); R. Seiler 126 (F, ITIC, MHES); 179 (MHES); 239 (F, MHES);
V.M. Martínez s.n. (CMC01027) (B, LAGU, MO). Chalatenango: J. Tucker 1060 (F);
R. Seiler 154 (F); 410 (F, ITIC, MHES); 423 (JBL00082) (LAGU, MHES); W.
Berendsohn et al. 1573 (B, LAGU, MO). Morazán: J.L. Linares & A. Pool 2291
(JBL01780) (LAGU); R. Seiler 1065 (F, MHES).
Bolbitis aliena (Sw.) Alston, Kew Bull. 1932: 310. 1932.
Acrostichum alienum Sw., Gymnopteris aliena (Sw.) C. Presl, Leptochilus alienus (Sw.)
C. Chr.
Global distribution: Mexico, Honduras, Nicaragua, Costa Rica, Panama, Antilles,
Source: Calderón & Standley (1941): 31 (as Leptochilus alienus); Seiler (1982): 389;
Berendsohn (1989) 1: 9.
♦ Bolbitis bernoullii (Kuhn ex Christ) Ching in Christensen, Index Filic., Suppl. 3: 47.
1934.
Acrostichum bernoullii Kuhn ex Christ, Gymnopteris bernoullii (Kuhn ex Christ) Diels
in Engler & Prantl, G. tuerckheimii Christ, G. donnell-smithii Christ, Leptochilus
bernoullii (Kuhn ex Christ) C. Chr., L. tuerckheimii (Christ) C. Chr., L. donnell-smithii
(Christ) C. Chr., Bolbitis donnell-smithii (Christ) Ching in C. Chr.
Local names: helecho cola de león.
Panama, Colombia, Ecuador.
Salvadoran distribution: Ahuachapán: M. Sandoval & E. Sandoval s.n. (ISB00801)
(AAU, B, F, LAGU); M. Sandoval & E. Sandoval 299 (B, F, LAGU). La Libertad: W.
Berendsohn et al. 1454 (LAGU).
Bolbitis portoricensis (Spreng.) Hennipman, Amer. Fern J. 65: 30. 1975.
Acrostichum portoricensis Spreng., A. cladorrhizans Spreng., A. irregulare Liebm.,
Gymnopteris portoricensis (Spreng.) Fée, Leptochilus cladorrhizans (Spreng.) Maxon,
Bolbitis cladorrhizans (Spreng.) Ching in Christensen .
Local names: helecho colchón de león, helecho salamandra, pescadillo.
Panama, Antilles, Colombia, Venezuela, Trinidad and Tobago, Ecuador.
LAGU, MO); A. Sermeño s.n. (JBL01118), 97 (JBL01000) (B, LAGU, MO); E.
Sandoval 1149 (B, LAGU); F. Chinchilla s.n. (ISB00314) (LAGU); J.M. Rosales 6
(AAU, B, BM, EAP, LAGU, MEXU, MO); 2450 (B, BM, LAGU, MEXU, MO); M.
Sandoval & E. Sandoval 269 (B, F, LAGU); P. Standley et al. 2604 (F); R. Seiler 923
(MHES); T. Croat 42096 (ITIC); W. Berendsohn 1317 (B, LAGU, MO); 1724 (B,
LAGU). Santa Ana: J. Monterrosa et al. 84 (B, LAGU, MO); M. Magaña 45 (MP00023) (LAGU); R. Seiler 706 (MHES). La Libertad: A.K. Monro et al. 3771 (B, BM,
ITIC, LAGU, MO); B. Pfeiffer-Berendsohn et al. 67 (B, LAGU, MO); F. Wunderlin
460 (ITIC); K. Sidwell et al. 479 (B, BM, ITIC, LAGU, MO); M.A. Hernández s.n.
(JCG00562) (B, LAGU, MO); W. Heed 27, 28, 29 (ITIC). San Salvador: K. Sidwell et
al. 569 (B, BM, ITIC, LAGU, MO); R. Seiler 314, 454, 458 (F, MHES).
Chalatenango: R. Seiler 144 (F, MHES). Cabañas: A.K. Monro et al. 3750 (B, BM,
ITIC, LAGU, MO); R.A. Carballo et al. 259 (LAGU). La Unión: A.K. Monro et al.
3537 (B, BM, ITIC, LAGU, MO).
Ctenitis equestris (Kunze) Ching var. equestris, Sunyatsenia 5: 250. 1940.
Lastrea ciliata Liebm., Polypodium alsophiloides Liebm., Aspidium bourgeaui E.
Fourn., A. scabriusculum Davenp., Dryopteris davenportii C. Chr., D. equestris
(Kunze) C. Chr. var. mentiens C. Chr.
Panama.
MO); G. Davidse et al. 37379 (B, BM, ITIC, LAGU, MO); R. Seiler 814, 906 (F,
MHES); 954 (MHES); W. Berendsohn & E. Sandoval 1510 (B, LAGU). Sonsonate: A.
Molina et al. 21630, 21763, 21775 (F, ITIC); 21770, 21773 (F); R. Seiler 612 (F,
MHES). Santa Ana: A. Molina et al. 12654, 21496 (F); 21513 (F, ITIC); D. Rodríguez
& G. Cerén s.n. (JBL04604) (LAGU); J. Tucker 1216 (F); K. Sidwell et al. 440 (B, BM,
ITIC, LAGU, MO, L). Lara 227, 228 (MHES); M. Magaña 23 (MP-00010) (LAGU);
R. Seiler 19 (MHES); 350 (F, MHES); S. Winkler 67 (ITIC); T. Croat 42232 (ITIC);
V.M. Martínez s.n. (CMC00819) (B, LAGU, MO); s.n. (CMC01000) (B, EAP, LAGU,
MO). San Salvador: M.C. Carlson 427, 477 (F); R. Seiler 208 (MHES); 209 (F,
MHES). Chalatenango: R. Seiler 435b (JBL04943) (LAGU, MHES). San Vicente: R.
Seiler 332 (F, MHES). Morazán: R. Seiler 1006 (F, MHES).
♦ Ctenitis equestris var. erosa Stolze, Amer. Fern. J. 67: 41. 1977.
Salvadoran distribution: Santa Ana: R. Seiler 484, 690 (F, MHES).
Ctenitis erinacea A.R. Sm., Mem. New York Bot. Gard. 88: 223-224. 2004.
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FERN GAZ. 18(4): 120-215. 2008
Salvadoran distribution: R. Seiler 890 (UC) (Mickel & Smith, 2004); Croat 42398
(CAS, MEXU) (Mickel & Smith, 2004). Santa Ana: B. Pfeiffer-Berendsohn 52 (B,
LAGU, MO); J. Monterrosa & R. Carballo 287 (B, BM, LAGU); R. Seiler 136
(MHES); 312 (F); 479 (F, ITIC, MHES). Chalatenango: Tucker 1140 (LL, UC, US)
(Mickel & Smith, 2004).
Ctenitis excelsa (Desv.) Proctor, Rhodora 63: 34. 1961.
Polypodium excelsum Desv., Dryopteris excelsa (Desv.) C. Chr. Local name: helecho
colchón de león.
Panama, Antilles.
LAGU, MO); E. Sandoval 1148 (LAGU, MO); E. Sandoval & F. Chinchilla 488 (B,
LAGU); R. Seiler 922 (F, MHES). Santa Ana: R. Seiler 879 (F).
Ctenitis hemsleyana (Baker) Copel., Gen. Fil. 124. 1947.
Polypodium hemsleyanum Baker ex Hemsl., Dryopteris hemsleyana (Baker ex Hemsl.)
C. Chr., Ctenitis costaricensis R.C. Moran
Salvadoran distribution: Morazán: R. Seiler 1024 (F, MHES).
Ctenitis interjecta (C. Chr.) Ching, Sunyatsenia 5: 250. 1940.
Dryopteris interjecta C. Chr.
Global distribution: Mexico, Guatemala, Belize, Honduras, Costa Rica.
Source: Löetschert (1954): 27; Seiler (1982): 388; Berendsohn (1989) 1: 9.
Ctenitis nigrovenia (Christ) Copel., Gen. Fil. 124. 1947.
Nephrodium nigrovenium Christ in Donnell Smith, Dryopteris nigrovenia (Christ) C.
Chr. Local name: helecho trecillo.
Colombia, Venezuela, Peru, Bolivia.
Salvadoran distribution: Ahuachapán: D. Witsberger 692, 655 (MHES); E. Sandoval
1135 (B, LAGU, MO); R. Seiler 817 (F, ITIC, MHES, NY).
Ctenitis sloanei (Poepp. ex Spreng.) C.V. Morton, Amer. Fern J. 59: 66. 1969.
Polypodium sloanei Poepp. ex Spreng.
Global distribution: USA, Mexico, Honduras, Nicaragua, Costa Rica, Panama, Antilles,
Colombia, Venezuela, Trinidad and Tobago, Ecuador, Peru, Bolivia.
Salvadoran distribution: Ahuachapán: N. Herrera s.n. (JBL03707) (LAGU, MO). San
Salvador: A.K. Monro et al. 3453 (B, BM, ITIC, LAGU, MO).
Dryopteris karwinskyana (Mett.) Kuntze, Revis. Gen. Pl. 2: 813. 1891.
Aspidium karwinskyanum Mett., Nephrodium karwinskyanum (Mett.) Baker in Hooker
& Baker . Local name: helecho pata de paloma.
Global distribution: Mexico, Guatemala, Honduras, Nicaragua, Costa Rica.
MO); M. Sandoval & E. Sandoval 69, 102 (B, F, LAGU); W. Berendsohn et al. 1642
(B, BM, LAGU, MO). La Libertad: N. Ventura 5010, 5024 (ITIC). San Salvador: K.
Prestegard 34 (MP-00011) (LAGU); R. Seiler 70 (F, MHES); 438 (F, ITIC, MHES).
Chalatenango: R. Seiler 522 (F); 554 (F, MHES). Cabañas: R. Seiler 520 (F, MHES).
Morazán: J. Monterrosa & C. Rivera 679 (BM, LAGU).
Dryopteris maxonii Underw. & C. Chr., Amer. Fern. J. I: 96. 1911.
Dryopteris patula (Sw.) Underw. var. moreliae Christ in Lecomte
Global distribution: Mexico, Guatemala, Costa Rica.
Salvadoran distribution: R. Seiler 1485 (MO) (Moran in Davidse et al. 1995)
Source: Moran in Davidse et al. (1995): 213.
Dryopteris nubigena Maxon & C.V. Morton, Proc. Biol. Soc. Wash. 50: 179. 1937.
Local names: helecho petatillo de montaña.
Global distribution: Mexico, Guatemala, Honduras, Costa Rica, Panama.
Salvadoran distribution: Santa Ana: R. Seiler 691 (F, MHES). Chalatenango: W.
Berendsohn et al. 1570 (B, LAGU, MO).
Dryopteris patula (Sw.) Underw., Native Ferns, ed. 4, 117. 1893.
Aspidium patulum Sw., A. paupertinum Kunze, A. mexicanum C. Presl var. obtusilobum
Kunze ex Mett., A. apertum Fée, Dryopteris aperta (Fée) C. Chr., D. simplicior Mickel
& Beitel
Local names: helecho cola de pavo real, helecho pata de comadreja, helecho pata de
paloma, helecho pluma de faisán.
Antilles, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, Paraguay.
Salvadoran distribution: Ahuachapán: E. Sandoval & M. Sandoval 1371 (B, LAGU);
F. Chinchilla s.n. (ISB00538) (AAU, B, LAGU); s.n. (ISB00659) (AAU, B, F, LAGU);
M. Sandoval & E. Sandoval 19 (B, LAGU); M. Sandoval 66 (B, F, LAGU). Sonsonate:
R. Seiler 34, 611 (F, MHES); T. Croat 42245 (ITIC). Santa Ana: K. Prestegard 1 (MP00012) (LAGU); M.B. Siu s.n. (ITIC); R. Seiler 82 (MHES); 500 (F, ITIC, MHES); 660
(F, MHES). La Libertad: E.A. Montalvo & R. Villacorta 6268 (B, LAGU). San
Salvador: R. Seiler 25, 52 (F, MHES); 437 (F, ITIC, MHES); 441 (MHES); S. Winkler
79 (ITIC). Chalatenango: R. Seiler 567 (JBL04812) (F, LAGU); R. Seiler 142, 568 (F,
MHES). San Vicente: R. Seiler 318 (JBL04806) (F, LAGU, MHES). Morazán: J.
Monterrosa & M. Rivera 643 (B, EAP, LAGU); J. Monterrosa & C. Rivera 648 (BM,
LAGU); S. Winkler 84 (ITIC).
Dryopteris wallichiana (Spreng.) Hyl., Bot. Not. 1953: 352. 1953.
Aspidium paleaceum D. Don, A. wallichianum Spreng., A. donianum Spreng., A.
paleaceum Sw., A. parallelogrammum Kunze, A. crinitum M. Martens & Galeotti,
Dryopteris paleacea (Sw.) C. Chr., D. parallelogramma (Kunze) Alston
Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, Paraguay, Africa, Asia, Hawaii.
Salvadoran distribution: Chalatenango: Hammer s.n. (12-274) (MHES); R. Seiler 163
(F, MHES). San Vicente: S. Winkler 77 (ITIC).
Elaphoglossum affine (M. Martens & Galeotti) T. Moore, Index Fil. 4. 1857.
Acrostichum affine M. Martens & Galeotti
Global distribution: Mexico, Guatemala, Costa Rica, Panama, Colombia, Venezuela.
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FERN GAZ. 18(4): 120-215. 2008
Salvadoran distribution: M.C. Carlson 730, F (Mickel, 1995: 256).
Source: Mickel in Davidse et al. (1995): 256; Mickel & Smith (2004): 285.
Elaphoglossum albomarginatum A.R. Sm., Proc. Calif. Acad. Sci. ser. 4, 40: 220 (F).
4A. 1975.
MO). Santa Ana: R. Seiler 876 (F, ITIC, MHES); S. Winkler 62 (ITIC).
Elaphoglossum baquianorum A. Rojas, Rev. Biol. Trop. 51(1): 35. 2003.
Salvadoran distribution: Santa Ana: R. Seiler 290 (F, MHES); R. Seiler 688 (F, MHES,
NY).
Elaphoglossum ellipticifolium A. Rojas, Rev. Biol. Trop. 51(1): 5. 2003.
Salvadoran distribution: Santa Ana: A. Molina et al. 16855 (F); T. Croat 42423,
holotipo MO, isotipo NY; R. Seiler 170 (F, MHES). Chalatenango: R. Seiler 397
(JBL00100) (F, LAGU, MHES); R. Seiler 429 (JBL00098), F (LAGU). Morazán: R.
Seiler 992 (F, MHES).
Elaphoglossum erinaceum (Fée) T. Moore var. erinaceum, Index Fil. 9. 1857.
Acrostichum erinaceum Fée,
Salvadoran distribution: Santa Ana: M.L. Reyna de Aguilar 839, 872 (MHES).
Elaphoglossum eximium (Mett.) Christ, Monogr. Elaph. 107. 1899.
Acrostichum eximium Mett.
Salvadoran distribution: Santa Ana: A. Molina et al. 16824; B. Pfeiffer-Berendsohn 35
(LAGU); J. Lagos s.n. (ITIC); R. Seiler 174 (F, MHES); S. Winkler 617 (ITIC).
Usulután: L. González 1942 (ITIC).
Elaphoglossum glabellum J. Sm., London J. Bot. I: 197. 1842.
Acrostichum glabellum (J. Sm.) Klotzsch
Global distribution: Mexico, Costa Rica, Panama, Antilles, Colombia, Venezuela,
Trinidad and Tobago, Guayana, Suriname, Ecuador, Peru, Bolivia.
Salvadoran distribution: Ahuachapán: P. Standley 20172 (US).
Source: Mickel & Smith (2004): 290.
Elaphoglossum glaucum T. Moore, Index. Fil. 10. 1857.
Acrostichum glaucum Fée, Elaphoglossum eucraspedum Christ
Salvadoran distribution: Santa Ana: R. Seiler 668 (MHES). Chalatenango: R. Seiler
399 (JBL00101) (F, LAGU, MHES).
Elaphoglossum guatemalense (Klotzsch) T. Moore, Parker´s Cat. 1858.
Acrostichum guatemalense Klotzsch, A. salvinii Baker ex Hemsl.
Global distribution: Mexico, Guatemala, Belize, Honduras.
Salvadoran distribution: Santa Ana: A. Molina et al. 16868 (F). Chalatenango: R.
Seiler 396 (F, MHES). San Vicente: R. Seiler 321 (F, MHES). Morazán: R. Seiler
1066 (MHES).
Elaphoglossum herrerae A. Rojas, Brenesia 45-46: 13. 1996.
Elaphoglossum lanceum Mickel, Amer. Fern J. 69: 101. 1979.
Salvadoran distribution: Santa Ana: R.A. Carballo 649 (B, LAGU, MO); R. Seiler
104, 173 (F, MHES); 684 (MHES). Chalatenango: R. Seiler 161, 739 (F, MHES).
Elaphoglossum latifolium (Sw.) J. Sm., J. Bot. 1: 197. 1842.
Acrostichum latifolium Liebm., A. sartori Liebm., A. scapellum Kunze ex Fée,
Elaphoglossum latifolium (Sw.) J. Sm. var. lutescens Rosenst., E. sartori (Liebm.)
Mickel, E. scapellum (Kunze ex Fée) T. Moore
Panama, Antilles, Colombia, Venezuela, Guayana, Ecuador, Peru, Bolivia, Brazil.
Salvadoran distribution: Santa Ana: G. Cerén & D. Rodríguez s.n. (JBL04645)
(LAGU); G. Davidse et al. 37290 (B, BM, ITIC, LAGU, MO); J.L. Linares & C.A.
Martínez 1520 (JBL01904) (LAGU); J. Monterrosa & R. Carballo 859 (LAGU); J.
Monterrosa et al. 174, 179 (LAGU); R. Seiler 103, 175, 657 (MHES); R. Villacorta et
al. 1286, 1018 (B, LAGU, MO). Chalatenango: W. Berendsohn et al. 1596 (B,
LAGU); R. Seiler 150, 749; 753 (F, MHES).
Note: Rojas-Alvarado (2003) notes that Elaphoglossum latifolium is taxonomically
difficult because of variation in rhizome habit and scales, frond scales, size, apex and
base shape. Rojas-Alvarado (2003) and Mickel & Smith (2004) suggests that E.
latifolium sensu stricto may be restricted to the Caribbean islands. It would seem
probable that the collections cited here as E. latifolium include more than one species.
♦ Elaphoglossum lonchophyllum (Fée) T. Moore, Index. Fil. 360. 1862.
Acrostichum lonchophyllum Fée, Elaphoglossum acutissimum Christ
Antilles.
Salvadoran distribution: Santa Ana: B. Pfeiffer-Berendsohn & W. Berendsohn 28
(LAGU, MO).
Elaphoglossum longistipitatum A. Rojas, Brenesia 47-48: 6. 1997.
Salvadoran distribution: Santa Ana: W. Löetschert 375 (US).
Source: Rojas-Alvarado (2002): 997.
Elaphoglossum mcvaughii Mickel, Brittonia 32: 336. 1980.
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FERN GAZ. 18(4): 120-215. 2008
Salvadoran distribution: Santa Ana: J. Monterrosa et al. 1011 (B, LAGU, MO, NY).
San Vicente: R. Seiler 322 (F, MHES).
Elaphoglossum mesoamericanum A. Rojas, Rev. Biol. Trop. 51(1): 8. 2003.
Salvadoran distribution: Santa Ana: M. Carlson 753 (F). San Salvador: M. Carlson
593 (F). Chalatenango: R. Seiler 258 (F, MHES); S. Calderón 2448 (F).
Note: Mickel & Smith (2004) cite Seiler 578 as Elaphoglossum yourkeorum (see
below), this specimen may be conspecific to E. mesoamericanum but the two species
have been maintained distinct here pending further research.
Elaphoglossum mexicanum (E. Fourn.) A. Rojas, Rev. Biol. Trop. 51(1): 42. 2003.
Acrostichum mexicanum E. Fourn.
Salvadoran distribution: Santa Ana: R. Seiler 130 (F, MHES); W. Löetschert 369, US
(Rojas-Alvarado, 2003).
z Elaphoglossum microproductum A. Rojas, Brenesia 47-48: 9. 1997.
Global distribution: Endemic.
Salvadoran distribution: Chalatenango: R. Seiler 742 (MHES, NY).
♦ Elaphoglossum minutum (Pohl ex Fée) T. Moore, Index Fil. 12. 1857.
Acrostichum minutum Pohl ex Fée, A. conforme Sw. var. alpinum J. Bommer ex H.
Christ?, A. leptophyllum Fée, Elaphoglossum leptophyllum (Fée) T. Moore
Global distribution: Costa Rica, Panama, Antillas, Colombia, Venezuela, Ecuador,
Peru, Brazil, Bolivia, Brazil.
MO). Morazán: R. Seiler 1016 (F, MHES).
Elaphoglossum muscosum (Sw.) T. Moore, Index. Fil. 12. 1857.
Acrostichum muscosum Sw., Elaphoglossum hookerianum Underw. ex Maxon
Colombia, Venezuela, Ecuador, Peru.
Salvadoran distribution: Sonsonate: R. Seiler 601 (MHES). Santa Ana: B. PfeifferBerendsohn & W. Berendsohn 4, 24 (LAGU, MO); M.L. Reyna de Aguilar 665
(MHES); R. Seiler 90, 354b, 653 (MHES). Chalatenango: R. Seiler 428 (JBL00097)
(LAGU, MHES); R. Seiler 740 (MHES). Morazán: R. Seiler 994 (F, MHES).
Elaphoglossum paleaceum (Hook. & Grev.) Sledge, Bull. Brit. Mus. (Nat. Hist.) Bot.
4: 95. 1967.
Acrostichum paleaceum Hook. & Grev., A. squamosum Sw., Elaphoglossum
squamosum J. Sm., E. hirtum sensu auct., non Acrostichum hirtum Sw.
Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, Hawaii.
Salvadoran distribution: Santa Ana: J. Monterrosa & R. Carballo 857 (B, LAGU); R.
Seiler 291, 645 (F, MHES); S. Winkler 55 (ITIC). San Salvador: R. Seiler 337
(MHES). Chalatenango: R. Seiler 403 (JBL00099) (F, LAGU, MHES); 741 (MHES).
Elaphoglossum peltatum (Sw.) Urb., Symb. Antill. 4: 60. 1903.
Osmunda peltata Sw., Acrostichum peltatum (Sw.) Sw., Rhipidopteris peltata (Sw.)
Schott ex Fée, Peltapteris peltata (Sw.) C.V. Morton .
Local names: sombrillita.
Antilles, Colombia, Venezuela, Guayana, Suriname, French Guyana, Ecuador, Peru,
Bolivia.
et al. 16754, 16850 (F; B). Pfeiffer-Berendsohn & W. Berendsohn 27 (B, F, LAGU,
MO); G. Davidse et al. 37298 (B, BM, ITIC, LAGU, MO); K. Sidwell et al. 846 (B,
BM, ITIC, LAGU, MO); M.C. Carlson 882 (F); O. Rohweder s.n. (JBL01258), 2649
(JBL01243) (LAGU); R. Seiler 97 (F, ITIC, MHES); 868 (MHES). Morazán: R. Seiler
993 (F, MHES).
Elaphoglossum petiolatum (Sw.) Urb., Symb. Antill. 4: 61. 1903.
Acrostichum petiolatum Sw., A. viscosum Sw., A. schiedei Kunze, A. schmitzii Mett. ex
Kuhn, Elaphoglossum viscosum (Sw.) J. Sm.
Antilles, Colombia, Venezuela, Ecuador, Peru, Brazil.
MO). Morazán: R. Seiler 1029 (F, ITIC, NY).
Elaphoglossum piloselloides (C. Presl) T. Moore, Index Fil. 13. 1857.
Acrostichum piloselloides C. Presl
Colombia, Venezuela, Guayana, Suriname, French Guyana, Ecuador, Peru, Brazil,
Bolivia, Chile.
Salvadoran distribution: Santa Ana: J.L. Linares & C.A. Martínez 1519 (JBL01916)
(LAGU).
Elaphoglossum sartorii (Liebm.) Mickel, Mem. New York Bot. Gard. 46: 181. 1988.
Acrostichum sartorii Liebm., A. scapellum Kunze ex Fée, Elaphoglossum scapellum
(Kunze ex Fée) T. Moore, E. subarborescens Rosenst.
Salvadoran distribution: San Vicente: R. Seiler 327 (F, MHES).
Elaphoglossum setosum (Liebm.) T. Moore, Index Fil. 366. 1862.
Acrostichum setosum Liebm., A. recognitum Kunze ex E. Fourn., Elaphoglossum
catharinae Underw. ex Maxon
Global distribution: Mexico, Guatemala, Nicaragua, Costa Rica.
Salvadoran distribution: Chalatenango: J. Monterrosa et al. 1619 (BM, LAGU,
MHES, NY).
Elaphoglossum squamipes (Hook.) T. Moore, Index Fil. 15. 1857.
Acrostichum squamipes Hook., A. ovatum Liebm., Elaphoglossum subaequale T.
Moore
Global distribution: Mexico, Guatemala, Costa Rica, Panama, Antilles, Colombia,
Venezuela, Ecuador, Peru, Brazil, Bolivia.
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FERN GAZ. 18(4): 120-215. 2008
Salvadoran distribution: Chalatenango: R. Seiler 425 (JBL00096) (LAGU); R. Seiler
149 (F, MHES); 745 (MHES).
Elaphoglossum tenuiculum (Fée) T. Moore ex Baker, Ann. Bot. (Oxford) 5: 491. 1891.
Acrostichum tenuiculum Fée, Elaphoglossum stellatum Mickel
Global distribution: Mexico, Guatemala, Honduras, Costa Rica, Colombia, Venezuela,
Ecuador, Peru, Brazil, Bolivia.
Salvadoran distribution: San Salvador: R. Seiler 31, 54, 515 (MHES). Chalatenango:
W. Heed 31 (ITIC).
Elaphoglossum yourkeorum Mickel, Mem. New York Bot. Gard. 88: 314-315. 2004.
Salvadoran distribution: Chalatenango: R. Seiler 578 (F, MHES, NY).
♦Lastreopsis effusa subsp. divergens (Willd.) Tindale, Contr. New South Wales Natl.
Herb. 3: 299, pl. 21. 1965.
Polypodium divergens Willd.
Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay.
Salvadoran distribution: Santa Ana: J.Monterrosa, D.Rodríguez & E.Pérez 1553 (BM,
LAGU).
♦ Megalastrum pulverulentum (Poir.) A. R. Sm. & R. C. Moran, Amer. Fern. J. 77:
129. 1987.
Ctenitis pulverulenta (Poir.) Copel., C. karsteniana (Klotzsch) Vareschi, C.
pulverulenta (Poir.) Copel. var. heydei (C. Chr.) Stolze, Polypodium pulverulentum
Poir. in Lamarck, P. karstenianum Klotzsch, Dryopteris pulverulenta (Poir.) C. Chr. in
Urban, D. karsteniana (Klotzsch) Hieron, D. karsteniana (Klotzsch) Hieron var. heydei
C. Chr., Megalastrum pulverulentum (Poir.) A. R. Sm. & R. C. Moran var. heydei (C.
Chr.) A.R. Sm. & R.C. Moran
Salvadoran distribution: Santa Ana: J. Monterrosa & R. Martínez 944 (LAGU); J.
Monterrosa et al. 961 (LAGU, NY).
♦Olfersia cervina (L.) Kunze, Flora 7: 312. 1824.
Global distribution: Mexico, Guatemala, Belice, Honduras, Nicaragua, Costa Rica,
Salvadoran distribution: Cabañas: G. Cerén 1788 (LAGU, MEXU, MHES); J.
Monterrosa & D. Rodríguez 1633 (BM, LAGU, MHES, MO).
Phanerophlebia juglandifolia (Humb. & Bonpl. ex Willd.) J. Sm., J. Bot. (Hooker) 4:
187. 1841.
Polypodium juglandifolium Humb. & Bonpl. ex Willd., Cyrtomium juglandifolium
(Humb. & Bonpl. ex Willd.) T. Moore
Salvadoran distribution: Tucker 1288, US (Yatskievych in Davidse et al. 1995).
Sonsonate: J.L. Linares & C.A. Martínez 3045 (JBL01790) (LAGU).
Phanerophlebia macrosora (Baker) Underw., Bull. Torrey Bot. Club 26: 213. 1899.
Aspidium juglandifolium (Humb. & Bonpl. ex Willd.) Kunze ex Klotzsch var.
macrosorum Baker, Cyrtomium macrosorum (Baker) C.V. Morton, C. guatemalensis
(Underw.) C.V. Morton, Phanerophlebia guatemalensis Underw.
Global distribution: Mexico, Guatemala, Costa Rica, Panama.
Salvadoran distribution: Chalatenango: R. Seiler 418 (JBL00081) (F, LAGU, MHES).
Polystichum alfaroi (Christ) Barrington, Rhodora 94: 328. 1992.
Polystichum aculeatum (L.) Roth var. alfaroi Christ
Global distribution: Mexico, Guatemala, Costa Rica, Panama.
Salvadoran distribution: Sin loc: P. Standley 21567, NY (Mickel & Smith, 2004: 514).
Ahuachapán: J. Monterrosa & D. Rodríguez 1434 (B, BM, LAGU, MO, NY).
Polystichum distans E. Fourn., Mexic. Pl. I: 91. 1872.
Polystichum pallidum E. Fourn.
Salvadoran distribution: Santa Ana: R. Seiler 12, 18, 504 (MHES). San Salvador: R.
Seiler 207 (MHES). Chalatenango: R. Seiler 381 (MHES). San Vicente: R. Seiler 315
(MHES).
Polystichum fournieri A. R. Sm., Amer. Fern J. 70: 27. 1980.
Polystichum muelleri Mett. ex E. Fourn.
Salvadoran distribution: Sonsonate: R. Seiler 613 (MHES). Santa Ana: M.L. Reyna
de Aguilar 667 (MHES); R. Seiler 181, 461 (MHES). Chalatenango: W. Berendsohn
et al. 1575 (B, LAGU).
Polystichum hartwegii (Klotzsch) Hieron., Hedwigia 46: 355. 1907.
Aspidium hartwegii Klotzsch, A. trejoi Christ, Polystichum grande Fée, P. drepanoides
E. Fourn., P. trejoi (Christ) Christ
Local name: helecho trenza de sansón.
Colombia?, Venezuela, Ecuador, Bolivia.
Salvadoran distribution: Ahuachapán: D. Witsberger 654 (MHES); F. Chinchilla & E.
López s.n. (ISB00311) (LAGU); N. Herrera s.n. (JBL03863) (B, EAP, LAGU, MO).
Santa Ana: D. Rodríguez & G. Cerén s.n. (JBL04613) (LAGU); J. Monterrosa et al.
208 (B, LAGU); J. Monterrosa & R. Martínez 940 (LAGU, NY); 948 (B, LAGU, NY);
K. Sidwell et al. 833 (B, BM, ITIC, LAGU, MO); R. Seiler 293, 345 (MHES); V.M.
Martínez s.n. (CMC00845) (B, LAGU); s.n. (CMC00880) (B, LAGU, MO); s.n.
(CMC00879), s.n. (CMC00897), s.n. (CMC00972) (B, EAP, LAGU, MO); W.
Berendsohn et al. 1541 (B, LAGU, MO). Usulután: A.K. Monro et al. 3007 (B, BM,
ITIC, LAGU, MO). Morazán: R. Seiler 1011 (MHES); R. Seiler 1033 (JBL04820)
(LAGU, MHES).
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FERN GAZ. 18(4): 120-215. 2008
Polystichum mickelii A. R. Sm., Amer. Fern. J. 70: 25. 1980.
Salvadoran distribution: P.Standley 20135, GH (Barrington in Davidse et al., 1995:
222).
Sources: Barrington in Davidse et al. (1995): 222; Mickel & Smith (2004): 518.
♦ Polystichum muenchii (Christ) C. Chr., Index Fil. 585. 1906.
Aspidium muenchii Christ
MO).
Polystichum muricatum (L.) Fée, Mém. Foug. 5: 278. 1852.
Polypodium muricatum L.
Local names: helecho pata de león, helecho rosquilla de altura, helecho trenza de
sansón.
Antilles, Venezuela.
Salvadoran distribution: Ahuachapán: E. Sandoval 1134 (B, LAGU); F. Chinchilla et
al. s.n. (ISB00091) (B, LAGU, MO); M. Sandoval & E. Sandoval 92 (B, F, LAGU); R.
Villacorta et al. 906 (B, LAGU, MO).Usulután: D. Williams 58 (LAGU, MO).
Polystichum turrialbae Christ, Bull. Herb. Boissier, sér. 2, 6: 163. 1906.
Polystichum smithii Mickel & Beitel
Local name: helecho trenza de sansón.
Global distribution: Mexico, Costa Rica, Panama, Bolivia.
LAGU, MO); W. Berendsohn et al. 1427 (B, LAGU, MO).
EQUISETACEAE
Equisetum giganteum L., Syst. Nat. ed. 10, 2: 1318. 1759.
Local name: cola de caballo.
Global distribution: Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Antilles,
Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, Paraguay, Uruguay, Argentina,
Chile.
Salvadoran distribution: Santa Ana: M.B. Siu s.n. (ITIC); R. Villacorta & H. Sipman
1260 (B, LAGU, MO); V.M. Martínez s.n. (CMC00048) (B, BM, LAGU, MEXU,
MO); V.M. Rosales 10 (ITIC). Chalatenango: B. López s.n. (ITIC); E.A. Montalvo
5435 (ITIC); R. Seiler 255 (F, ITIC, MHES).
Equisetum hyemale A. Braun ex Engelm. L. var. affine (Engelm.) A. A. Eaton, Fern
Bull. II: III. 1903.
Local name: cola de caballo.
Equisetum robustum A. Braun ex Engelm., E. robustum A. Braun ex Engelm. γ affine
Engelm., E. hyemale L. subsp. affine (Engelm.) Calder & Roy Taylor
Global distribution: Canada, USA, Mexico, Guatemala, Asia.
Salvadoran distribution: La Libertad: M. Renderos & R. Villacorta 291 (B, LAGU, MO).
Equisetum myriochaetum Schltdl. & Cham., Linnea 5: 623. 1830.
Equisetum mexicanum Milde
Salvadoran distribution: Santa Ana: R. Villacorta et al. 1049 (B, LAGU, MO).
Chalatenango: E.A. Montalvo 5279 (ITIC). Morazán: M. Renderos et al. 628 (B, BM,
LAGU, MO).
Equisetum x schaffneri Milde, Verh. Zool.-Bot. Ges. Wien II: 345. 1861.
Global distribution: Mexico, Guatemala a Peru.
Salvadoran distribution: S. Calderón 1864 (F, ITIC, US).
Note: Equisetum x schaffneri is believed to be a hybrid between E. giganteum and E.
myriochaetum (Hauke, 1995: 326).
GLEICHENIACEAE
Dicranopteris flexuosa (Schrad.) Underw., Bull. Torrey Bot. Club 34: 254. 1907.
Mertensia flexuosa Schrad., Gleichenia flexuosa (Schrad.) Mett.
Suriname, French Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay.
Salvadoran distribution: La Libertad: M.C. Carlson 251 (F). Usulután: D. Williams
141 (LAGU, MO); J. Lagos 1777 (ITIC). Morazán: K. Sidwell et al. 736 (B, BM,
ITIC, LAGU, MO); R. Seiler 1049b (F, ITIC, MHES); R. Villacorta et al. (B, LAGU).
Diplopterygium bancroftii (Hook.) A. R. Sm., Amer. Fern. J. 70: 26. 1980.
Gleichenia bancroftii Hook., Mertensia bancroftii (Hook.) Kunze, M. bancroftii
(Hook.) Kunze var. vitellina Kunze, Dicranopteris bancroftii (Hook.) Underw.,
Hicriopteris bancroftii (Hook.) Ching
Global distribution: Mexico, Guatemala, Honduras, Costa Rica, Antilles, Colombia,
Salvadoran distribution: Santa Ana: J. Tucker 1184, F, K; Puller s.n. (12-548)
(MHES); R. Seiler 238 (F, ITIC, MHES); S. Winkler 68 (ITIC).
Gleichenella pectinata (Willd.) Ching, Sunyatsenia 5: 276. 1940.
Mertensia pectinata Willd., M. glaucescens Humb. & Bonpl. ex Willd. var. mexicana
Fée, Gleichenia pectinata (Willd.) C. Presl, G. nitida C. Presl, Dicranopteris pectinata
(Willd.) Underw.
Salvadoran distribution: Santa Ana: A. Molina et al. 12577 (F); R. Seiler 460 (F,
MHES); 1199 (F). Chalatenango: E.A. Montalvo s.n. (ITIC); M.C. Carlson 590 (F); R.
Villacorta & A. Araniva 968 (B, LAGU, MO). Morazán: R. Seiler 983 (F, MHES).
Sticherus bifidus (Willd.) Ching, Sunyatsenia 5: 282. 1940.
Mertensia bifida Willd., M. fulva Desv., M. gleichenioides Liebm., Gleichenia bifida
(Willd.) Spreng., G. liebmannii T. Moore, Dicranopteris bifida (Willd.) Maxon, D. fulva
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FERN GAZ. 18(4): 120-215. 2008
(Desv.) Underw.
Local name: helecho cien pié.
Guyana, Ecuador, Peru, Brazil, Bolivia.
J.M. Rosales 1294 (B, LAGU, MO). Santa Ana: H. Weber 293 (ITIC); M.C. Carlson
997 (F). La Libertad: M.A. Hernández s.n. (JCG00590) (B, LAGU, MO). Morazán:
J. Tucker 792 (F).
Sticherus brevipubis (Christ) A. R. Sm., Amer. Fern J. 70: 27. 1980.
Gleichenia brevipubis Christ
Antilles, Colombia, Ecuador.
Salvadoran distribution: Ahuachapán: J.L. Linares 482 (JBL01980) (LAGU). Santa
Ana: A. Molina et al. 12520 (F); G. Cerén & D. Rodríguez s.n. (JBL04661) (LAGU);
G. Davidse et al. 37309 (B, BM, ITIC, LAGU, MO); M.L. Reyna de Aguilar 682
(MHES); R. Villacorta et al. 1065 (B, ITIC, LAGU, MO). Chalatenango: J. Tucker
1183 (F); R. Seiler 725 (F, MHES). Morazán: R. Seiler 989 (F, MHES).
Sticherus palmatus (W. Schaffn. ex E. Fourn.) Copel., Gen. Fil. 28. 1947.
Mertensia palmata W. Schaffn. ex Fée, M. palmata W. Schaffn. ex E. Fourn.,
Gleichenia palmata T. Moore, G. palmata (W. Schaffn. ex E. Fourn.) C. Chr.,
Dicranopteris palmata J. H. Schaffn. ex Underw.
Panama, Antilles.
Salvadoran distribution: Santa Ana: J. Monterrosa & R. Carballo 273 (B, BM, LAGU,
MO); R. Seiler 233 (F, ITIC, MHES); 241 (ITIC).
Sticherus underwoodianus (Maxon) Nakai, Bull. Natl. Sci. Mus. Tokyo 29: 31. 1950.
Dicranopteris underwoodiana Maxon, Gleichenia underwoodiana (Maxon) C. Chr.
Local names: helecho de mano, helecho pié de gallina.
Salvadoran distribution: Santa Ana: A. Molina et al. 16747 (F; B). PfeifferBerendsohn 38 (B, LAGU, MO); B. López s.n. (ITIC); G. Davidse et al. 37269 (B, BM,
ITIC, LAGU, MO); R. Seiler 195 (MHES); 230 (ITIC, MHES); 251 (F, ITIC, MHES);
W. Berendsohn et al. 1547 (B, LAGU). Chalatenango: R. Seiler 726 (F, MHES).
HYMENOPHYLLACEAE
Hymenophyllum crassipetiolatum Stolze, Amer. Fern J. 66: 15. f. I. 1976.
Salvadoran distribution: Santa Ana: A. Molina et al. 12638, 16767, 16811 F; 12488,
12504 (F, ITIC); M.C. Carlson 894 (F); R. Seiler 102 (F, MHES); 687, 876 (MHES);
Chalatenango: J. Tucker 1062a. Morazán: J.L. Linares & A. Pool 2296 (JBL01929)
(LAGU).
♦Hymenophyllum crispum Kunth, Nov. Gen. Sp. I: 26. 1815.
Sphaerocionium crispum (Kunth) Klotzcsh, S. schiedeanum C. Presl, Hymenophyllum
schiedeanum (C. Presl) Bosch
Global distribution: Mexico, Guatemala, Honduras, Costa Rica, Antilles, Colombia,
Venezuela, Ecuador, Peru, Brazil, Bolivia.
Salvadoran distribution: Chalatenango: J.Monterrosa et al. 1618 (BM, LAGU,
MHES).
Hymenophyllum fucoides (Sw.) Sw., J. Bot. (Schrader) 1800(2): 99. 1801.
Trichomanes fucoides Sw., Meringium fucoides (Sw.) Copel.
Salvadoran distribution: Santa Ana: B. Pfeiffer-Berendsohn & W. Berendsohn 22
(LAGU, MO); 31 (B, LAGU, MO); M.L. Reyna de Aguilar 877 (MHES); R. Seiler
284a (ITIC, MHES); 295 (F, MHES); 297 (F, ITIC, MHES). Chalatenango: R. Seiler
433 (F). Morazán: R. Seiler 996 (F, ITIC, MHES).
Hymenophyllum microcarpum Desv., Mém. Soc. Linn. Paris 6: 333. 1827.
Sphaerocionium microcarpum (Desv.) Copel., Hymenophyllum angustifrons Christ, H.
microcarpum Desv. var. lanceolatum C. V. Morton
Antilles, Dominica, Colombia, Venezuela, Guayana, Ecuador, Peru, Brazil, Bolivia.
Source: Stolze (1976): 66; Berendsohn (1989) 1: 13; Pacheco in Davidse et al. (1995):
68; Mickel & Smith (2004): 344.
Hymenophyllum myriocarpum Hook., Sp. Fil. I: 106, t. 37D. 1844.
Mecodium myriocarpum (Hook.) Copel., M. nigricans (C. Presl ex Klotzsch) Copel.,
Hymenophyllum nigrescens Liebm., Sphaerocionium nigricans C. Presl ex Klotzsch
Colombia, Venezuela, Guayana, Suriname, French Guyana, Ecuador, Peru, Bolivia.
Salvadoran distribution: Santa Ana: R. Seiler 285 (F, ITIC, MHES); R. Villacorta 1119
(B, LAGU, MO); S. Winkler 92 (ITIC). Chalatenango: R. Seiler 434 (JBL00105) (F,
ITIC, LAGU, MHES). San Vicente: R. Seiler 334 (F, ITIC, MHES). Morazán: R.
Seiler 995 (F, ITIC, MHES).
Hymenophyllum polyanthos (Sw.) Sw., J. Bot. (Schrader) 1800(2): 102. 1801.
Trichomanes polyanthos Sw., Mecodium polyanthos (Sw.) Copel., Hymenophyllum
jalapense Schltdl. & Cham., H. millefolium Schltdl. & Cham., H. botryoides Bosch
French Guyana, Ecuador, Peru, Brazil, Bolivia, Africa, Asia.
Salvadoran distribution: Sonsonate: A. Molina et al. 21767 (F, ITIC); R. Seiler 599 (F,
MHES). Santa Ana: A. Molina et al. 12629, 12661 (F, ITIC); 12685 F; G. Davidse et
al. 37215 (B, BM, ITIC, LAGU, MO); R. Seiler 282 (F, MHES). Morazán: R. Seiler
988 (F).
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FERN GAZ. 18(4): 120-215. 2008
Hymenophyllum pulchellum Schltdl. & Cham., Linnea 5: 618. 1830.
Sphaerocionium pulchellum (Schltdl. & Cham.) C. Presl, Hymenophyllum pannosum
Christ
Brazil.
Source: Seiler (1982): 384; Berendsohn (1989) 1: 13; Mickel & Smith (2004): 345.
Hymenophyllum tegularis (Desv.) Proctor & Lourteig, Bradea 5: 385. 1990.
Davallia tegularis Desv., Hymenophyllum elegantulum Bosch, Sphaerocionium
elegantulum (Bosch) Copel.
Salvadoran distribution: Santa Ana: A. Molina et al. 12505 (F, ITIC); 16813 (F); R.
Seiler 231 (F, ITIC, MHES); 683 (MHES); S. Winkler 91 (ITIC). Chalatenango: R.
Seiler 435a (JBL00106) (F, ITIC, LAGU, MHES).
Hymenophyllum tunbrigense (L.) Sm., Mém. Acad. Roy. Sci. (Turín) 5: 418. 1793.
Trichomanes tunbrigense L., Hymenophyllum fucoides (Sw.) Sw. var. frigidum Liebm.
Antilles, Venezuela, Ecuador, Bolivia, Chile, Argentina; Europe, Africa, Australia.
Salvadoran distribution: Santa Ana: R. Seiler 125 (F, ITIC, MHES); 284 (F, MHES).
Chalatenango: R. Seiler 432 (F, MHES).
Trichomanes capillaceum L., Sp. Pl. 2: 1099. 1753.
Vandenboschia capillacea (L.) Copel., Trichomanes tenellum Hedw., T. trichoideum
Sw., T. trichoides Sw., T. schiedeanum Müll., T. hypnoides Christ
Global distribution: Mexico, Guatemala, Belize, Honduras, Costa Rica, Panama,
Antilles, Colombia, Venezuela, Ecuador, Peru.
Salvadoran distribution: Santa Ana: A. Molina et al. 16870 (F); E.A. Montalvo 4152
(ITIC); G. Davidse et al. 37305 (B, BM, ITIC, LAGU, MO); R. Seiler 188, 286 (F,
ITIC, MHES); R. Villacorta et al. 1277 (B, ITIC, LAGU, MO); S. Winkler 90 (ITIC).
Trichomanes diaphanum Kunth, Nov. Gen. Sp. 1: 25. 1816.
Trichomanes bradei Christ
Global distribution: Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Panama,
Antilles, Colombia, Venezuela, Guayana, Ecuador, Peru, Brazil.
Salvadoran distribution: Ahuachapán: E. Sandoval & F. Chinchilla 731 (B, LAGU).
Santa Ana: A. Molina et al. 16804 (F); R. Seiler 250 (F, MHES); 303 (F, ITIC, MHES).
Usulután: D. Williams 247 (LAGU).
Trichomanes hymenoides Hedw., Fil. Gen. Sp., t. 3. 1799.
Trichomanes muscoides Sw., T. apodum Hook., Didymoglossum hymenoides (Hedwig)
Copel., Microgonium schaffneri Bosch ex Fée
Global distribution: Mexico, Guatemala, Belize, Nicaragua, Costa Rica, Panama,
Antilles, Colombia, Venezuela, Trinidad and Tobago, Ecuador, Peru, Brazil, Bolivia,
Argentina, Uruguay.
Salvadoran distribution: Ahuachapán: R. Seiler 964 (F, MHES). Santa Ana: R. Seiler
302 (F, MHES); 357 (F, ITIC, MHES); S. Winkler 64 (ITIC).
♦Trichomanes hymenophylloides Bosch, Ned. Kruidk. Arch. 5: 209. 1863.
Trichomanes leptophyllum Bosch, Vandenboschia hymenophylloides (Bosch) Copel.
Antilles, Colombia, Venezuela, Trinidad and Tobago, Guyana, Suriname, French
Guyana, Ecuador, Brazil.
Salvadoran distribution: Santa Ana: J.Monterrosa & D.Rodríguez 1586 (BM, LAGU,
MO).
Trichomanes krausii Hook. & Grev., Icon. Filic. t. 149. 1831.
Didymoglossum krausii (Hook. & Grev.) C. Presl, Hemiphlebium krausii (Hook. &
Grev.) Prantl
Panama, Antilles, Venezuela, Trinidad and Tobago, Guayana, Suriname, French
Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina.
Local names: helecho musgo de pimiento
Santa Ana: S. Winkler 94 (ITIC). Morazán: R. Seiler 1020 (F, MHES).
Trichomanes petersii A. Gray, Amer. J. Sci. Arts 15: 326. 1853.
Microgonium petersii (A. Gray) Bosch, Hemiphlebium petersii (A. Gray) Prantl,
Didymoglossum petersii (A. Gray) Copel., Trichomanes schaffneri Schltdl.
Global distribution: USA, Mexico, Guatemala, Honduras, Nicaragua, Costa Rica.
Salvadoran distribution: Chalatenango: R. Seiler 390 (JBL00107) (F, ITIC, LAGU,
MHES).
Trichomanes radicans Sw., J. Bot. (Schrader) 1800(2): 97. 1801.
Trichomanes scandens Hedw., T. kunzeanum Hook., T. Mexicanum Bosch,
Vandenboschia radicans (Sw.) Copel.
Local names: helecho plumajillo, helecho pluma de quetzal.
Brazil, Bolivia, Paraguay, Europe, Asia, Africa.
LAGU, MO); E. Sandoval & F. Chinchilla 44, 518 (B, LAGU); J.G. Sandoval et al. s.n.
(ISB00026) (B, LAGU, MO); M. Sandoval & E. Sandoval s.n. (ISB00799) (AAU, F,
LAGU); R. Seiler 812 (F, ITIC, MHES); 958 (F, MHES); W. Berendsohn et al. 1363
(B, BG, LAGU, MO). Sonsonate: A. Molina et al. 21771 (F); J.L. Linares & C.A.
Martínez 3035 (JBL01905) (LAGU); R. Seiler 618 (F, ITIC, MHES). Santa Ana: A.
Molina et al. 12648, 16905 (F); J. Monterrosa et al. 172 (B, BM, LAGU, MO); N.
López et al. s.n. (JBL04627) (LAGU); R. Seiler 116, 364 (F, ITIC, MHES); R.
Villacorta & M. Calderón 368 (B, LAGU, MO); S. Winkler 93 (ITIC); W. Berendsohn
et al. 1542 (B, LAGU, MO). San Salvador: R. Seiler 341 (F, MHES). Chalatenango:
J. Tucker 1217 (F).
Trichomanes reptans Sw., Prodr. 136. 1788.
Didymoglossum reptans (Sw.) C. Presl, Hemiphlebium reptans (Sw.) Prantl
171
FERN GAZ. 18(4): 120-215. 2008
Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina.
G. Davidse et al. 37423 (B, BM, ITIC, LAGU, MO). Sonsonate: R. Seiler 598 (F,
MHES). Santa Ana: S. Winkler 95 (ITIC). Chalatenango: R. Seiler 391 (F, MHES).
Usulután: A.K. Monro et al. 3005 (B, BM, ITIC, LAGU, MO).
Trichomanes robinsonii Baker, J. Linn. Soc. Bot. 9: 339. 1866.
Global distribution: Honduras.
Source: Löetschert (1954): 25; Berendsohn (1989) 1: 13.
LINDSAEACEAE
Lindsaea stricta (Sw.) Dryand. var. stricta, Trans. Linn. Soc. London 3: 42. 1797.
Adiantum strictum Sw., Lindsaea portoricensis Desv., L. microphylla C. Presl, L.
multifrondosa Fée, L. moritziana Klotzsch
Salvadoran distribution: Morazán: A.K. Monro et al. 3856 (B, BM, ITIC, LAGU,
MO); K. Sidwell et al. 733, 734 (B, BM, ITIC, LAGU, MO); M. Renderos et al. 605
(AAU, B, LAGU, MO); R. Seiler 1073 (F, MHES).
Note: Seiler (1982) and Berendsohn (1989) treat L. stricta and L. portoricensis as
distinct species.
Lonchitis hirsutus L., Sp. Pl. 2: 1078. 1753.
Anisosorus hirsutus (L.) Maxon, Pteris lacinata Willd.
Antilles, Colombia, Venezuela, Trinidad and Tobago, Suriname, Ecuador, Peru, SE
Brazil, Bolivia.
Source: Seiler (1982): 386; Berendsohn (1989) 1: 8.
LOMARIOPSIDACEAE
Cyclopeltis semicordata (Sw.) J. Sm., Bot. Mag. 72 (Compendium): 36. 1846.
Polypodium semicordatum Sw., Aspidium semicordatum (Sw.) Sw., Dryopteris
semicordata (Sw.) Kuntze, Polypodium semicordatum (Sw.) T. Moore
Salvadoran distribution: Ahuachapán: J.M. Rosales 488 (AAU, B, BM, LAGU, MO);
R. Seiler 925 (F, MHES). Sonsonate: D. Rodríguez & M. Trejo 99 (LAGU). San
Miguel: A. Salgado 70 (F, MHES); J. Tucker 925 (F).
Nephrolepis biserrata (Sw.) Schott, Gen. Fil., t. 3. 1834.
Aspidium biserratum Sw., Nephrolepis mollis Rosenst.
Suriname, French Guyana, Ecuador, Peru, Brazil, Bolivia, Africa, Asia, Polynesia.
Local names: helecho cola de pescado
Salvadoran distribution: Ahuachapán: M. Magaña 17 (MP-00030) (LAGU). La
Libertad: R. Seiler 1046 (F, MHES). Chalatenango: L. Lara 211 (MHES).
Nephrolepis cordifolia (L.) C. Presl, Suppl. Tent. Pterid. 79. 1836.
Polypodium cordifolium L.
Local names: centavito, helecho cola de ardilla.
Global distribution: USA, Mexico, Guatemala, Honduras, Panama, Antilles, Peru,
Brazil, Trópicos del Viejo Mundo.
Salvadoran distribution: La Libertad: K. Prestegard 47 (MP-00028) (LAGU); M.
Magaña 46 (MP-00027) (LAGU). San Salvador: R. Seiler 509 (F, MHES); Vaquerano
s.n. (12-923) (MHES).
Nephrolepis hirsutula (G. Forst.) C. Presl, Suppl. Tent. Pterid. 79. 1836.
Polypodium hirsutulum G. Forst., Davallia multiflora Roxb., Nephrolepis multiflora
(Roxb.) Jarret ex C.V. Morton
Local name: helecho cola de quetzal.
Global distribution: USA, Mexico, Belize, Guatemala, Honduras, Nicaragua, Costa
Rica, Panama, Antilles, Colombia, Venezuela, Trinidad and Tobago, Guayana, Ecuador,
Peru, Brazil, Bolivia, Asia, Polynesia.
E. Sandoval & E. Fuentes 1300 (B, LAGU); E. Sandoval & M. Sandoval 1016 (B,
LAGU); R. Seiler 945 (F); S. Castillo s.n. (ISF00687) (B, F, LAGU, MO); T. Croat
42159 (ITIC). La Libertad: E.A. Montalvo 73a (ITIC); K. Prestegard 27 (MP-00034)
(LAGU). San Salvador: R. Seiler 223, 373 (F, MHES). Chalatenango: L. Lara 141
(MHES). La Paz: R. Villacorta et al. 2855 (AAU, B, BM, LAGU, MO). Usulután: D.
Williams 51 (B, EAP, LAGU, MO). Morazán: R. Seiler 1044 (F, ITIC); 1088a (F, ITIC,
MHES).
Nephrolepis undulata (Afzel. ex Sw.) J. Sm., Bot. Mag. 72 (Compendium): 35 bis.
1846.
Aspidium undulatum Afzel. ex Sw., Nephrolepis occidentalis Kunze, N. intermedia Fée
Local names: helecho cola de iguana, helecho cola de quezallio, helecho quetzal.
Antilles, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, Africa, Asia.
Salvadoran distribution: Ahuachapán: E. Sandoval et al. 773, 1070 (B, LAGU); M.
Sandoval & E. Sandoval 70 (B, F, LAGU). Santa Ana: B. Pfeiffer-Berendsohn 58 (B,
LAGU, MO); R. Seiler 1215 (F, MHES). San Salvador: R. Seiler 44 (F, MHES); 637
(MHES).
LYGODIACEAE
Lygodium venustum Sw., J. Bot. (Schrader) 1801(2): 303. 1803.
Lygodium mexicanum C. Presl
Local names: alambrillo, helecho bejuco de san pablo, helecho crespillo.
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FERN GAZ. 18(4): 120-215. 2008
French Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay.
Salvadoran distribution: Ahuachapán: A. Sermeño 42 (JBL00946) (B, F, LAGU,
MO); D. Witsberger s.n. (12-998) (MHES); E. Sandoval & M. Sandoval 1150 (B,
LAGU); F. Chinchilla s.n. (ISB00307), s.n. (ISB00377) (LAGU); J.L. Linares & C.A.
Martínez 3085 (JBL01902) (LAGU); J.M. Rosales 241, 477 (AAU, B, BM, LAGU,
MO); 1128 (B, BM, LAGU, MO); 1858, 2403 (B, BM, LAGU, MEXU, MO); M.
Sandoval & E. Sandoval 106 (B, LAGU); O. Guerrero s.n. (ISF00448) (AAU, B, F,
LAGU); O. Rivera D. & E. Sandoval s.n. (ISB00695) (AAU, B, F, LAGU); S. Castillo
s.n. (ISF00108) (B, LAGU); W. Berendsohn et al. 1412 (B, LAGU, MO). Santa Ana:
J.C. González & R. Villacorta 121 (B, LAGU, MO); J.L. Linares & C.A. Martínez 1169
(JBL04295) (LAGU); K. Prestegard 22 (MP-00045) (LAGU). La Libertad: L.O.
Williams et al. 15218 (F); N. Ventura 5008 (ITIC); O. González et al. s.n. (JBL04501)
(LAGU); R. Villacorta 1187 (B, LAGU, MO). San Salvador: M.C. Carlson s.n., 11 (F).
Chalatenango: M.L. Reyna de Aguilar 269 (MHES). La Paz: R. Villacorta & S.
Villacorta 1147 (B, ITIC, LAGU, MO); W. Berendsohn et al. 1194 (B, F, HBG, LAGU).
San Vicente: P. Standley et al. 3516a, 3669 (F). San Miguel: A. Salgado 68 (MHES);
J. Tucker 816 (F). Cabañas: G. Davidse et al. 37095 (B, BM, ITIC, LAGU, MO); R.A.
Carballo et al. 222 (B, LAGU, MO). Morazán: J. Monterrosa & C. Rivera 670 (B, BM,
LAGU, MO). La Unión: A.K. Monro et al. 2762 (B, BM, ITIC, LAGU, MO); G.W.
Barclay 2632 (F).
MARATTIACEAE
Marattia excavata Underw., N. Amer. Fl. 16: 22. 1909.
Local names: helecho casco de mula, helecho casco de burro.
Salvadoran distribution: Sonsonate: J.L. Linares & C.A. Martínez 3046 (JBL01932)
(LAGU). Santa Ana: A. Molina et al. 21493 (ITIC); G. Cerén & D. Rodríguez s.n.
(JBL04652) (LAGU); J. Monterrosa & R. Carballo 854 (LAGU); J. Monterrosa & R.
Martínez 947 (LAGU); L. González s.n. (ITIC); L. Lara 229 (MHES); R. Seiler 268
(MHES); R. Villacorta et al. 1054 (B, LAGU, MO). Chalatenango: R. Seiler 436
(JBL00112) (LAGU, MHES). Morazán: R. Seiler 1031a (ITIC, MHES).
Note: Used by local people in Montecristo National Park as a contraceptive.
Marattia weinmanniifolia Liebm., Mexic. Bregn. 308 (reprint 156). 1849.
Salvadoran distribution: Santa Ana: R. Seiler 234 (F, MHES).
MARSILEACEAE
Marsilea ancylopoda A. Braun, Monatsber. Königl. Preuss. Akad. Wiss. Berlin 1863:
434. 1864.
Zaluzianskia ancylopoda (A. Braun) Kuntze, Z. mexicana (A. Braun) Kuntze, Marsilea
mexicana A. Braun
Antilles, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina,
Uruguay.
Salvadoran distribution: La Unión: N.C. Fassett 28654 (ITIC); R. Villacorta et al. 2708
(LAGU); J. Monterrosa & R. Villacorta 1636 (BM, LAGU, MHES, MO).
OPHIOGLOSACEAE
Botrychium schaffneri Underw., Bull. Torrey Bot. Club 30: 51. 1903.
Sceptridium schaffneri (Underw.) Lyon, S. pusillum (Underw.) Lyon, Botrychium
pusillum Underw., B. schaffneri Underw. var. pusillum (Underw.) R. T. Clausen
Ecuador, Peru, Bolivia, Argentina.
Source: Seiler (1982): 382; Berendsohn (1989) 1: 15; Mickel & Smith (2004): 163.
Botrychium underwoodianum Maxon, Bull. Torrey Bot. Club 32: 222. 1905.
Botrychium ternatum Hook. & Baker var. daedaleum Christ
Global distribution: Guatemala, Honduras, Costa Rica, Panama, Antilles, Colombia,
Venezuela, Brazil.
Salvadoran distribution: Santa Ana: B. Pfeiffer-Berendsohn 46 (B, LAGU, MO); 49
(LAGU, MO); M.L. Reyna de Aguilar 778 (MHES); S. Winkler 8a (ITIC).
Botrychium virginianum (L.) Sw., J. Bot. (Schrader) 1800(2): III. 1801.
Osmunda virginiana L., O. Cicutaria Savigny in Lamark, Botrychium cicutarium
(Savigny) Sw., B. virginianum (L.) Sw. var. mexicanum Hook. & Grev., B.
brachystachys Kunze, B. virginianum (L.) Sw. var. meridionale Butters, B. β
virginianum subsp. meridionale (Butters) R. T. Clausen
Global distribution: Canada, USA, Mexico, Guatemala, Honduras, Nicaragua, Costa
Rica, Panama, Antilles, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia; Europe,
Asia.
Salvadoran distribution: Santa Ana: F. Wunderlin 1230 (ITIC); J. Monterrosa et al.
197 (LAGU); 1023 (B, LAGU, NY); M.L. Reyna de Aguilar 679 (MHES); R. Seiler
132 (F, MHES); 252 (F, ITIC); 1146 (F). Chalatenango: R. Seiler 382 (JBL00093)
(LAGU, MHES).
Ophioglossum reticulatum L., Sp. Pl. 2: 1063. 1753.
Brazil, Bolivia, Argentina, Africa, Asia.
OSMUNDACEAE
Osmunda cinnamomea L., Sp. pl. 2: 1066. 1753.
Osmundastrum cinnamomeum (L.) C. Presl
Global distribution: Canada, USA, Mexico, Guatemala, Honduras, Costa Rica,
Antilles, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, Paraguay, Asia.
Salvadoran distribution: Sonsonate: R. Villacorta 768 (B, LAGU, MO). Morazán: R.
Seiler 1009, 1075 (F, MHES); S. Winkler 7b (ITIC).
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FERN GAZ. 18(4): 120-215. 2008
Osmunda regalis L. var. spectabilis (Willd.) A. Gray, Manual, ed. 2, 600. 1856.
Osmunda spectabilis Willd., O. mexicana Fée, O. palmeri A. E. Bobrov
Local names: helecho real.
Global distribution: Canada, USA, Mexico, Guatemala, Honduras, Nicaragua, Costa
Rica, Antilles, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, Paraguay,
Argentina, Uruguay.
Salvadoran distribution: Ahuachapán: G. Davidse et al. 37401 (B, BM, ITIC, LAGU,
MO); N.C. Fassett 28718a (ITIC); N. Herrera s.n. (JBL03779) (B, LAGU, MO).
Sonsonate: R. Villacorta 771 (B, LAGU, MO). Santa Ana: R. Seiler 779 (F).
Morazán: R. Seiler 977 (F, ITIC, MHES).
POLYPODIACEAE
Campyloneurum amphostemon (Kunze ex Klotzsch) Fée, Mém. Foug. 5: 258. 1852.
Polypodium amphostemon Kunze ex Klotzsch, P. angustifolium Sw. var. amphostemon
(Kunze ex Klotzsch) Baker in Martius, Campyloneurum angustifolium (Sw.) Fée var.
amphostemon (Kunze ex Klotzsch) Farw.
Salvadoran distribution: Santa Ana: A. Molina et al. 16764 (F, ITIC); J. Monterrosa et
al. 193 (B, BM, LAGU, MO); R. Seiler 89, 113 (F, MHES); 187 (MHES); 656, 669 (F,
MHES); S. Winkler 49, 58 (ITIC). Chalatenango: R. Seiler 385 (JBL00203) (LAGU,
MHES); 395 (JBL04945) (F, LAGU, MHES); 392, 754, 755 (F, MHES); W.
Berendsohn et al. 1592 (B, LAGU, MO).
Campyloneurum angustifolium (Sw.) Fée, Mém. Foug. 5: 257. 1852.
Polypodium angustifolium Sw.
Rica, Panama, Antilles, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia.
Salvadoran distribution: Ahuachapán: P. Standley et al. 19932 (F). Santa Ana: R.
Seiler 707 (MHES). Chalatenango: R. Seiler 393 (JBL00196) (F, LAGU); 394
(JBL00202) (F, LAGU, MHES). San Vicente: S. Winkler 69 (ITIC).
Campyloneurum costatum (Kunze) C. Presl, Suppl. Tent. Pterid. 190. 1836.
Polypodium costatum Kunze
Suriname, French Guyana, Ecuador, Brazil.
Source: Seiler (1982): 385 (as Polypodium costatum); Berendsohn (1989) 1: 17 (as
Polypodium costatum).
Campyloneurum ensifolium (Willd.) J. Sm., Cat. Cult. Ferns 12. 1857.
Polypodium ensifolium Willd., P. angustifolium Sw. var. ensifolium (Willd.) Hicken,
Campyloneurum angustifolium (Sw.) Fée var. ensifolium (Willd.) Farw.
Local names: ajillo, helecho barba de cacique, helecho barbiquejo.
Salvadoran distribution: Jaurequi 87 (UC) (Mickel & Smith 2004). Ahuachapán: A.K.
Monro et al. 2971, 3425 (B, BM, ITIC, LAGU, MO); E. Sandoval 1258 (B, LAGU);
J.L. Linares 516 (JBL01863) (LAGU); P. Standley et al. 2640 (F); R. Seiler 969 (F,
MHES); R. Villacorta et al. 869 (B, ITIC, LAGU); S. de Moore s.n. (12-926) (MHES);
W. Berendsohn et al. 1370, 1433 (B, LAGU, MO). Sonsonate: R. Seiler 3 (F, MHES).
Santa Ana: A. del Cid 1614 (ITIC); A. Mangandi et al. s.n. (JBL04469) (LAGU); A.K.
Monro et al. 2304 (B, BM, ITIC, LAGU, MO); E.A. Montalvo s.n. (ITIC); J.
Monterrosa & R. Carballo 512 (B, LAGU, MO); M. Magaña 32 (MP-00039) (LAGU);
R. Villacorta 2173 (B, LAGU, MO); S. Winkler s.n., 50 (ITIC); V.M. Martínez s.n.
(CMC00256) (LAGU). La Libertad: E.A. Montalvo 4298 (ITIC). San Salvador: P.
Standley et al. 22672 (F). La Paz: S. Flores 9 (ITIC). San Vicente: E.A. Montalvo s.n.
(ITIC). Usulután: D. Williams & R.W. Herrera 455 (LAGU). San Miguel: A. Salgado
67 (F, MHES).
Note: This species name has been placed in synonomy, or as a variety of C.
angustifolium. Lellinger (1988) and León (1992) clarify the diagnostic characters (scale
cell shape). See discussion by Mickel & Smith (2004) under C. ensifolium.
Campyloneurum phyllitidis (L.) C. Presl, Suppl. Tent. Pterid. 190. 1836.
Polypodium phyllitidis L.
Global distribution: USA, Mexico, Guatemala, Belize, Honduras, Costa Rica, Panama,
Salvadoran distribution: San Salvador: R. Seiler 971 (F, MHES).
Campyloneurum xalapense Feé, Mém. Foug. 5: 258. 1852.
Polypodium xalapense (Fée) Christ, P. weatherbyanum F. Seym., Campyloneurum
caudatum Fée . Local name: cincho.
Salvadoran distribution: Ahuachapán: M.L. Reyna de Aguilar 651 (MHES); R. Seiler
907 (F, ITIC, MHES). Sonsonate: R. Seiler 595 (F, MHES); R. Villacorta et al. 329
(LAGU, MO). Santa Ana: A. Molina et al. 12658; B. Pfeiffer-Berendsohn 48 (B,
LAGU, MO); 76 (LAGU, MO); J. Monterrosa et al. 178, 968 (LAGU); J. Monterrosa
& R. Carballo 278 (B, BM, LAGU, MO); 978 (B, LAGU, NY); M.B. Siu s.n. (ITIC);
R. Seiler 122 (F, ITIC, MHES); 274, 670 (F, MHES); S. Winkler 57 (ITIC). San
Salvador: M.C. Carlson 505 (F); S. Winkler 59 (ITIC). San Vicente: E.A. Montalvo
4655 (ITIC); R. Seiler 324 (F); 325 (F, MHES). San Miguel: M. Renderos et al. 672
(LAGU).
Ceradenia jungermannioides (Klotzsch) L. E. Bishop, Amer. Fern J. 78: 4. 1988.
Polypodium jungermannioides Klotzsch, Grammitis jungermannioides (Klotzsch)
Ching
Colombia, Venezuela, Brazil; Azores.
Salvadoran distribution: R. Seiler 1194 (UC) (A.R. Smith in Davidse et al. 1995).
Santa Ana: R. Seiler 124 (F, ITIC, MHES).
Ceradenia margaritata (A. R. Sm.) L. E. Bishop, Amer. Fern J. 78: 5. 1988.
Grammitis margaritata A. R. Sm.
Salvadoran distribution: Tucker 1075 (UC) (A.R. Smith in Davidse et al. 1995).
177
FERN GAZ. 18(4): 120-215. 2008
Source: Smith in Davidse et al. (1995): 370; Mickel & Smith (2004): 173.
Cochlidium rostratum (Hook.) Maxon ex C. Chr., Dansk Bot. Ark. 6(3): 23. 1929.
Monogramma rostrata Hook., Grammitis rostrata (Hook.) R. M. Tryon & A. F. Tryon
Antilles, Colombia, Venezuela.
et al. 12492, 16796 (F); J. Toledo s.n. (ITIC); R. Seiler 98 (F, MHES); 647 (MHES).
Morazán: R. Seiler 997 (F, MHES); S. Winkler 46 (ITIC).
Cochlidium serrulatum (Sw.) L. E. Bishop, Amer. Fern J. 68: 80. 1978.
Acrostichum serrulatum Sw., Grammitis serrulata (Sw.) Sw., Xiphopteris serrulata
(Sw.) Kaulf., Polypodium serrulatum (Sw.) Mett., P. duale Maxon
Panama, Antilles, Colombia, Venezuela, Trinidad and Tobago, Guayana, Ecuador, Peru,
Brazil, Bolivia, Africa.
MO). Chalatenango: S. Winkler 45 (ITIC). Morazán: R. Seiler 1004 (F, MHES, NY).
Lellingeria apiculata (Kunze ex Klotzsch) A. R. Sm. & R. C. Moran, Amer. Fern J.
81: 83. 1991.
Polypodium apiculatum Kunze ex Klotzsch, Ctenopteris apiculata (Kunze ex Klotzsch)
Copel., Grammitis apiculata (Kunze ex Klotzsch) F. Seym.
Colombia, Venezuela, Guayana, Ecuador, Peru, Bolivia, Brazil.
Source: Seiler (1982): 385 (as Grammitis appiculata); Berendsohn (1989) 1: 16 (as
Grammitis appiculata).
Lellingeria prionodes (Mickel & Beitel) A. R. Sm. & R. C. Moran, Amer. Fern J. 81:
86. 1991.
Grammitis prionodes Mickel & Beitel
Salvadoran distribution: Sonsonate: R. Seiler 602 (F, MHES). Santa Ana: E.A.
Montalvo 4155 (ITIC); J. Monterrosa et al. 226 (B, LAGU); L. Lara 237 (MHES); R.
Villacorta & L. Lara 2664 (B, LAGU). Chalatenango: S. Winkler 81 (ITIC). Morazán:
Loxogramme mexicana (Fée) C. Chr., Index Filic. Suppl. 3: 25. 1934.
Selliguea mexicana Fée, Gymnogramma mexicana (Fée) Baker in Hooker & Baker, G.
salvinii (Hook.) Hook., Grammitis salvinii Hook., Loxogramme salvinii (Hook.) Maxon
Global distribution: Mexico, Guatemala, Nicaragua, Costa Rica, Panama.
Salvadoran distribution: Ahuachapán: E.A. Montalvo 57 (ITIC); G. Davidse et al.
37411 (B, BM, ITIC, LAGU, MO); R. Seiler 822 (F, MHES). Santa Ana: R. Seiler 359,
503 (F, MHES). San Salvador: R. Seiler 342 (F, MHES).
Melpomene deltata (Mickel & Beitel) A. R. Sm. & R. C. Moran, Novon 2: 429.
1992.
Grammitis deltata Mickel & Beitel
Global distribution: Mexico, Honduras.
Source: Mickel & Beitel (1988): 199; Moran & Smith in Davidse et al. (1995): 381;
Mickel & Smith (2004): 390.
Melpomene firma (J. Sm.) A. R. Sm. & R. C. Moran, Novon 2: 430. 1992.
Polypodium firmum Klotzsch, P. aromaticum Maxon, Ctenopteris firma J. Sm., C.
aromatica (Maxon) Copel., Grammitis firma (J. Sm.) C. V. Morton, G. aromatica
(Maxon) Proctor
Colombia, Venezuela, Guayana, Ecuador, Peru, Bolivia.
Salvadoran distribution: Sonsonate: R. Seiler 603 (MHES). Santa Ana: J. Monterrosa
et al. 182 (LAGU); R. Seiler 96 (F, ITIC, MHES). Chalatenango: R. Seiler 737
(MHES). Morazán: R. Seiler 998 (F, MHES).
Melpomene leptostoma (Fée) A. R. Sm. & R. C. Moran, Novon 2: 430. 1992.
Polypodium leptostomum Fée, Polypodium productum Maxon, Ctenopteris leptostoma
(Fée) Copel., Grammitis leptostoma (Fée) F. Seym.
Source: Stolze (1981): 258 (as Grammitis leptostoma); Berendsohn (1989) 1: 16 (as
Grammitis leptostoma).
Melpomene moniliformis (Lag. ex Sw.) A. R. Sm. & R. C. Moran, Novon 2: 430. 1992.
Polypodium moniliforme Lag. ex Sw., P. moniliforme Lag. ex Sw. var. major Liebm.
Ctenopteris moniliformis (Lag. ex Sw.) J. Sm., Grammitis moniliformis (Lag. ex Sw.)
Proctor
Salvadoran distribution: Santa Ana: L. Lara 98 (MHES); M.C. Carlson 900a (F); W.
Berendsohn et al. 1550 (B, LAGU). Chalatenango: R. Seiler 431 (JBL04944) (F,
LAGU, MHES); R. Seiler 748 (F, MHES); 758 (MHES); R. Villacorta et al. 2074 (B,
LAGU, MO); W. Berendsohn et al. 1568 (B, LAGU, MO).
Melpomene pilosissima (M. Martens & Galeotti) A. R. Sm. & R. C. Moran, Novon
2: 431. 1992.
Polypodium pilosissimum M. Martens & Galeotti, Ctenopteris pilosissima (M. Martens
& Galeotti) Copel., Grammitis pilosissima (M. Martens & Galeotti) C. V. Morton
Venezuela, Suriname, Ecuador, Peru, Bolivia, Brazil.
Source: Seiler (1982): 386 (as Grammitis pilossisima); Berendsohn (1989) 1: 16 (as
Grammitis pilossisima).
Melpomene xiphopteroides (Liebm.) A. R. Sm. & R. C. Moran, Novon 2: 431. 1992.
Polypodium xiphopteroides Liebm., P. rigens Maxon, Ctenopteris rigens (Maxon)
Copel., C. megaloura Copel., Grammitis xiphopteroides (Liebm.) A. R. Sm., Grammitis
rigens (Maxon) Proctor
Salvadoran distribution: Santa Ana: R. Seiler 93 (F, MHES); 95 (ITIC, MHES).
179
FERN GAZ. 18(4): 120-215. 2008
Micropolypodium basiattenuatum (Jenman) A. R. Sm., Novon 2: 422. 1992.
Polypodium basiattenuatum Jenman, Xiphopteris basiattenuata (Jenman) Copel.,
Grammitis basiattenuata (Jenman) Proctor, G. basiattenuata (Jenman) Proctor var.
valens Mickel & Beitel
Global distribution: Mexico, Guatemala, Honduras, Antilles, Venezuela.
Salvadoran distribution: R. Seiler 1133, NY (Mickel & Smith 2004).
Source: Seiler (1982): 385 (as Grammitis basiattenuata); Berendsohn (1989) 1: 16 (as
Grammitis rigens); Smith in Davidse et al. (1995): 383; Mickel & Smith (2004): 400.
Niphidium crassifolium (L.) Lellinger, Amer. Fern J. 62: 106. 1972.
Polypodium crassifolium L., Pessopteris crassifolia (L.) Underw. & Maxon
Local name: cincho.
Salvadoran distribution: Seiler 1529, MO (Pacheco in Davidse et al. 1995). Santa Ana:
R. Seiler 491 (F, MHES). La Libertad: O. Pank s.n. (MP-00041) (LAGU).
Pecluma alfredii (Rosenst.) M. G. Price var. cupreolepis (A. M. Evans) A. R. Sm.,
Mem. New York Bot. Gard. 88: 437. 2004.
Polypodium cupreolepis A. M. Evans, Pecluma cupreolepis (A. M. Evans) M. G. Price
R. Seiler 914 (F, ITIC); 968 (F, MHES). Santa Ana: B. Pfeiffer-Berendsohn 81
(LAGU, MO); G. Cerén & D. Rodríguez s.n. (JBL04647) (LAGU); J. Tucker 1249 (F);
R. Seiler 129 (F, MHES); 287 (MHES). Chalatenango: R. Seiler 385 (F, MHES). San
Vicente: E.A. Montalvo 73b (ITIC). San Miguel: W. Eggler 665 (ITIC). Morazán: R.
Pecluma plumula (Humb. & Bonpl. ex Willd.) M.G. Price, Amer. Fern J. 73: 115. 1983.
Polypodium plumula Humb. & Bonpl. ex Willd., P. pulchrum M. Martens & Galeotti,
P. pulchrum M. Martens & Galeotti var. minus E. Fourn., Ctenopteris plumula (Humb.
& Bonpl. ex Willd.) J. Sm.
Suriname, French Guyana, Ecuador, Peru, Brazil, Bolivia.
Salvadoran distribution: Ahuachapán: E.A. Montalvo 62 (ITIC); H. Weber 35 (ITIC);
Sonsonate: R. Seiler 75 (F, MHES). Santa Ana: M.B. Siu s.n. (ITIC); R.A. Carballo &
E.O. Ramos 675 (B, LAGU, MO). La Libertad: M. Renderos 496, B, K (LAGU, MO);
N. Ventura 5016 (ITIC); S. Guadalupe & F. Lessette s.n. (JBL04481) (LAGU). San
Salvador: M.C. Carlson 356 (F). San Vicente: R. Seiler 323 (F, MHES). Usulután: D.
Williams 56 (LAGU, MO); D. Williams & R.W. Herrera 298 (LAGU). San Miguel: A.
Salgado 64 (F, MHES). Usulután: E.A. Montalvo 5035, 5037 (ITIC). Morazán: J.
Monterrosa & C. Rivera 696 (B, LAGU, MO); R.A. Carballo et al. 1044 (LAGU).
Phlebodium pseudoaureum (Cav.) Lellinger, Amer. Fern J. 77: 101. 1987.
Polypodium pseudoaureum Cav., Polypodium areolatum Humb. & Bonpl. ex Willd., P.
areolatum Humb. & Bonpl. ex Willd. var. loreum J. Bommer, P. aureum auct., en parte
L., Goniophlebium areolatum (Humb. & Bonpl. ex Willd.) C. Presl, Phlebodium
aureum auct., en parte (L.) J. Sm.
Local names: helecho calaguala, helecho cola de pavo real.
Antilles, Colombia, Venezuela, Guayana, Ecuador, Peru, Brazil, Bolivia, Argentina,
Paraguay.
Salvadoran distribution: Ahuachapán: D. Witsberger 696 (MHES); E. Sandoval & M.
Sandoval 1299 (B, LAGU); W. Berendsohn et al. 1458 (LAGU). Santa Ana: G.
Davidse et al. 37233 (B, BM, ITIC, LAGU, MO); J. Monterrosa & R. Carballo 284 (B,
BM, EAP, F, LAGU, MO); R. Seiler 185 (MHES). San Salvador: R. Seiler 28, 56, 513
(MHES). San Vicente: R. Seiler 317 (MHES). Usulután: D. Williams & R.W. Herrera
442 (LAGU). Morazán: R. Seiler 1000 (MHES).
Pleopeltis angusta Humb. & Bonpl. ex Willd. var. angusta, Sp. Pl., ed. 4, 5(1): 211.
1810.
Polypodium angustum (Humb. & Bonpl. ex Willd.) Liebm.
Local names: helecho mano de mapache, helecho pié de mapache.
LAGU, MO); D. Witsberger 688 (MHES); E. Sandoval & M. Sandoval 1298 (B,
LAGU); F. Chinchilla s.n. (ISB00206) (B, LAGU, MO); M. Magaña 16 (MP-00040)
(LAGU); R. Seiler 913, 967 (F, MHES); S. de Moore s.n. (12-927) (MHES); W.
Berendsohn et al. 1456, 1459 (LAGU). Sonsonate: R. Seiler 627 (F, MHES). Santa
Ana: B. Pfeiffer-Berendsohn et al. 10 (B, F, LAGU, MO); G. Cerén & D. Rodríguez
s.n. (JBL04668) (LAGU); J. Monterrosa et al. 175 (B, BM, LAGU, MO); 176 (B,
LAGU, MO); J. Monterrosa et al. 920 (B, LAGU, NY); 973 (LAGU); K. Sidwell et al.
436 (B, BM, ITIC, LAGU, MO); M.B. Siu s.n. (ITIC); R. Seiler 168 (F, MHES). La
Libertad: W. Berendsohn et al. 1445 (LAGU); W. Eggler 681 (ITIC). San Salvador:
A.K. Monro et al. 2240 (B, BM, ITIC, LAGU, MO); M.C. Carlson 429 (F); O.
Rohweder s.n. (ITIC); R. Seiler 26 (F, ITIC, MHES). Usulután: A.K. Monro et al. 2999
(B, BM, ITIC, LAGU, MO); D. Williams 129 (B, LAGU, MO). San Miguel: K.
Sidwell et al. 884 (B, BM, ITIC, LAGU, MO).
Pleopeltis astrolepis (Liebm.) E. Fourn., Mexic. Pl. I: 87. 1872.
Polypodium astrolepis Liebm., P. elongatum (Sw.) Mett., P. lanceolatum L. var.
elongatum (Sw.) Krug, Phlebodium astrolepis (Liebm.) Conzantti, Grammitis
lanceolata Schkuhr, G. revoluta Spreng. ex Willd., Gymnogramma elongata (Sw.)
Hook., Pleopeltis revoluta (Spreng. ex Willd.) A. R. Sm.
Local names: helecho barba amarilla, helecho barba negra, helecho barbiquejo.
Rica, Panama, Antilles, Colombia, Venezuela, Trinidad and Tobago, Suriname, French
Salvadoran distribution: Ahuachapán: A.K. Monro & M.L. Fomtg 2973 (B, BM,
ITIC, LAGU, MO); D. Witsberger 711 (MHES); E. Sandoval et al. 729, 1430 (B,
LAGU); F. Chinchilla s.n. (AAU, B, LAGU); M. Sandoval & E. Sandoval 138 (F,
LAGU); P. Standley et al. 2516 (F); R. Seiler 928, 965 (F, MHES); W. Berendsohn et
al. 1430 (B, LAGU, MO); 1466 (LAGU). Sonsonete: R. Seiler 77 (F). Santa Ana: B.
181
FERN GAZ. 18(4): 120-215. 2008
Pfeiffer-Berendsohn 20 (LAGU, MO); V.M. Martínez s.n. (CMC00103) (LAGU). La
Libertad: J. Monterrosa et al. s.n. (JBL04485) (LAGU); N. Ventura 5020 (ITIC); O.
Rohweder 2634 (JBL01246), 2635 (JBL01245), 2636 (JBL01244) (LAGU); W. Zelaya
et al. s.n. (LAGU). San Salvador: R. Seiler 507 (F, MHES); 1515 (F). San Vicente:
E.A. Montalvo 5098 (ITIC). Usulután: E.A. Montalvo 5047 (ITIC); J. Lagos 1783
(ITIC). Cuscatlán: C. Salazar s.n. (ITIC).
Pleopeltis mexicana (Fée) Mickel & Beitel, Amer. Fern J. 77: 21. 1987.
Drymaria mexicana Fée, Polypodium lanceolatum L. var. trichophorum Weath.,
Pleopeltis macrocarpa (Bory ex Willd.) Kaulf. var. trichophora (Weath.) Pic. Serm.
Salvadoran distribution: Santa Ana: A. Molina et al. 12652 (F; B). PfeifferBerendsohn 26, 78, 79 (B, LAGU, MO); 56 (LAGU, MO); G. Davidse et al. 37164,
37306 (B, BM, ITIC, LAGU, MO); J. Monterrosa et al. 171 (B, BM, LAGU, MO); R.
Seiler 109 (F, MHES); S. Winkler 56 (ITIC). Chalatenango: R. Seiler 386 (JBL00201)
(ITIC, LAGU); R. Seiler 152, 729 (F, MHES). Morazán: R. Seiler 1005 (F, ITIC,
Pleopeltis polylepis (Roemer ex Kunze) T. Moore var. interjecta (Weath) E. A. Hooper,
Amer. Fern J. 85: 79. 1995.
Polypodium peltatum Cav. var. interjectum Weath., Pleopeltis macrocarpa (Bory ex
Willd.) Kaulf. var. interjecta (Weath.) A. R. Sm., P. interjecta (Weath.) Mickel & Beitel
Salvadoran distribution: Santa Ana: W. Berendsohn et al. 1714 (B, LAGU).
Polypodium alansmithii R. C. Moran, Ann. Missouri Bot. Gard. 77: 845. 1990.
Salvadoran distribution: Sonsonate: R. Seiler 633 (MHES). Santa Ana: A. Molina et
al. 12562 (F; B). Pfeiffer-Berendsohn 77 (B, LAGU, MO); E.A. Montalvo s.n., 3641
(ITIC); G. Cerén & D. Rodríguez s.n. (JBL04667) (LAGU); G. Davidse et al. 37301 (B,
BM, ITIC, LAGU, MO); J. Monterrosa et al. 165 (B, BM, LAGU, MO); 951, 964 (B,
LAGU, NY); 942 (B, LAGU, MO, NY); 1007 (LAGU, NY); K. Sidwell et al. 847 (B,
BM, ITIC, LAGU, MO); M.C. Carlson 726, 727, 970 (F); N.C. Fassett 28292 (ITIC);
R. Seiler 87 (F); 91, 190 (F, MHES); R. Villacorta & Claus 2147 (B, LAGU, MO); W.
Berendsohn 1303 (LAGU, MO). San Salvador: R. Seiler 214 (F, MHES).
Chalatenango: R. Seiler 398 (JBL00198) (F, LAGU, MHES); W. Heed 36 (ITIC). San
Vicente: R. Seiler 326 (MHES).
Polypodium castaneum Maxon & Tejero, Act. Bot. Mex. 67: 87. 2004.
Global distribution: Guatemala.
Salvadoran distribution: Sonsonate: Croat 42213, MO. Santa Ana: J. Monterrosa et al.
967 (B, BM, IZTA, LAGU, MO, NY).
Polypodium colpodes Kunze var. colpodes, Linnea 23: 276, 316. 1850.
Polypodium fuscopetiolatum A. R. Sm.
Local names: helecho calaguala rojo, helecho de árbol, helecho pluma de ganzo.
Colombia.
LAGU, MO); E. Sandoval et al. 548, 1442 (B, LAGU); F. Chinchilla s.n. (ISB00658)
(AAU, B, F, LAGU); J.M. Rosales 1061 (B, LAGU); 1301, 1491 (AAU, B, BM,
LAGU, MO); 1420 (B, BM, LAGU, MO); M. Sandoval & E. Sandoval 58 (B, F,
LAGU). Sonsonate: R. Seiler 38 (F, MHES); T. Croat 42237 (ITIC). Santa Ana: J.
Lagos 1616 (ITIC); V.M. Martínez s.n. (CMC00306) (B, LAGU). La Libertad: N.
Ventura 5017, 5021 (ITIC); R. Seiler 719 (F, MHES). San Salvador: R. Seiler 439, 584
(F, MHES); 514 (MHES). Chalatenango: L. Lara 210 (MHES); R. Seiler 473 (F,
MHES).
♦Polypodium diplotrichum Mickel & Beitel, Mem. New York Bot. Gard. 46: 297.
1988.
Goniophlebium lepidotrichum Fée
Global distribution: Mexico.
Salvadoran distribution: Ahuachapán: J.Monterrosa & D.Rodríguez 1446 (BM, IZTA,
LAGU).
Polypodium fissidens Maxon, Contr. U.S. Natl. Herb. 8: 275. 1903.
Local names: helecho calaguala
Salvadoran distribution: Chalatenango: W. Berendsohn et al. 1571 (B, LAGU); R.
Seiler 156, 164, 756 (F, MHES).
Polypodium furfuraceum Schltdl. & Cham., Linnea 5: 607. 1830.
Polypodium nivosum Fée, P. furfuraceum Schltdl. & Cham. var. coronulatum E. Fourn.,
P. furfuraceum Schltdl. & Cham. var. rufum E. Fourn., P. margallii Rovirosa
Local names: cola de ardilla, helecho peine de indígena, helecho peinetero rojo, palmita
de café.
Peru, Bolivia.
ITIC, LAGU, MO); E. Sandoval et al. 1190, 1403 (B, LAGU); F. Chinchilla & E.
Sandoval s.n. (ISB00428) (LAGU); R. Seiler 829 (ITIC); 970 (F, MHES); S. de Moore
s.n. (12-929) (MHES); W. Berendsohn et al. 1429 (LAGU). Sonsonate: R. Seiler 5, 628
(F, MHES). Santa Ana: B. Pfeiffer-Berendsohn 19 (LAGU); E.A. Montalvo 4297
(ITIC); T. Croat 43037 (ITIC); V.M. Martínez s.n. (CMC00251) (LAGU). La
Libertad: E.A. Montalvo 4302 (ITIC); N. Ventura 5005 (ITIC); V. Henríquez & C.
Bermúdez s.n. (JBL04482) (LAGU); W. Berendsohn 1452 (LAGU). San Salvador:
E.A. Montalvo 5350 (ITIC); L.O. Williams et al. 16746 (F); S. Flores 8 (ITIC). San
Vicente: E.A. Montalvo 5091 (ITIC). Cuscatlán: C. Salazar s.n. (ITIC). Usulután: D.
Williams & R.W. Herrera 300 (LAGU). San Miguel: A. Salgado 65b (F, MHES). La
Unión: A.K. Monro et al. 2123 (B, BM, ITIC, LAGU, MO).
Polypodium hartwegianum Hook. in Bentham, Pl. Hartw. 54. 1840.
Polypodium ellipsoideum Fée, P. pubescens Fée
Salvadoran distribution: Santa Ana: A. Molina et al. 12560 (F, ITIC); J. Monterrosa et
al. 1031, B, IZTA (LAGU, NY); 1044, IZTA (LAGU); L. Lara s.n. (12-992) (MHES);
183
FERN GAZ. 18(4): 120-215. 2008
R. Seiler 80, 81, 189, 672 (F, MHES). Chalatenango: E.A. Montalvo 4811 (ITIC); R.
Seiler 430 (JBL00199) (LAGU); R. Seiler 162, 426 (F, MHES). Morazán: R. Seiler
1092 (F, ITIC, MHES).
Polypodium hispidulum Bartlett, Proc. Amer. Acad. Arts 43: 48. 1907.
Global distribution: Mexico, Guatemala, Belize, Honduras, Nicaragua.
Source: Seiler (1982): 385; Berendsohn (1989) 1: 17; Moran in Davidse et al. (1995):
358; Mickel & Smith (2004): 494.
Polypodium lindenianum Kunze, Farrnkräuter 2: 83. 1851.
Polypodium cancellatum Fée, P. verapax Christ
Source: Stolze (1981): 408; Berendsohn (1989) 1: 17; Moran in Davidse et al. (1995):
362; Mickel & Smith (2004): 495.
Polypodium muenchii Christ, Bull. Herb. Boissier, sér. 2, 3: 147. 1903.
Pleopeltis muenchii (Christ) A. R. Sm., Microgramma muenchii (Christ) Copel.,
Microphlebodium muenchii (Christ) L. D. Gómez
Salvadoran distribution: Chalatenango: R. Villacorta et al. 2075 (B, LAGU); R. Seiler
400 (JBL00200) (LAGU); R. Seiler 153 (MHES); Tucker 1070 (EAP, F).
Polypodium plebeium Schltdl. & Cham., Linnea 5: 607. 1830.
Polypodium cheilostictum Fée, P. karwinskyanum (C. Presl) A. Braun ex Kunze,
Marginaria karwinskyana C. Presl
Salvadoran distribution: Sonsonate: R. Seiler 608 (F, MHES). Santa Ana: R. Seiler 16
(MHES). San Salvador: R. Seiler 206 (MHES). Morazán: R. Seiler 1019 (F, MHES).
Polypodium plesiosorum Kunze var. plesiosorum, Linnea 18: 313. 1844.
Polypodium gonatodes Kunze, P. confluens Liebm., P. anisomeron (Fée) E. Fourn. var.
elongatum E. Fourn., P. anisomeron (Fée) E. Fourn. var. genuinum E. Fourn., P.
anisomeron (Fée) E. Fourn. var. pectinatum E. Fourn., P. anisomeron (Fée) E. Fourn.
var. abortivum E. Fourn., P. feei E. Fourn., P. van-heurckii E. Fourn., P. pachyrhizon
Christ, P. rossii Christ ex H. Ross, Goniophlebium molestum Fée, G. anisomeron Fée,
G. calaguala Fée, G. stramineum Underw. ex Conzatti
Local name: calaguala.
Salvadoran distribution: Ahuachapán: A.K. Monro et al.3428 (B, BM, ITIC, LAGU,
MO). Sonsonate: A. Molina et al. 21758 (F, ITIC); R. Seiler 594 (F, MHES). Santa
Ana: G. Davidse et al. 37240, 37307 (B, BM, ITIC, LAGU, MO); J. Monterrosa et al.
932 (B, LAGU, NY); L. Lara 223 (MHES); R. Seiler 167 (F, ITIC, MHES); 344 (F,
MHES). Chalatenango: R. Seiler 388 (JBL00197) (F, LAGU, MHES); R. Seiler 757
(F, MHES).
Polypodium plesiosorum Kunze var. plesiosori Tejero, Anales Inst. Biol. Univ. Nac.
Autón. Mexico, Bot. 75(1): 24-28 (F). 1, 2a, 3g-h. 2004.
Local name: calaguala.
Global distribution: Mexico, Guatemala, Nicaragua.
Salvadoran distribution: Santa Ana: Linares s.n. (JBL01320) (B, LAGU, MO).
Note: var. rubicundum is found in Mexico, Honduras, Costa Rica and Panama.
Polypodium pleurosorum Kunze ex Mett., Abh. Senckenberg. Naturf. Ges. 2: 101, t. 2
(F). 6, 7. 1856.
Phlebodium inaequale T. Moore, Goniophlebium inaequale (T. Moore) J. Sm.,
Polypodium inaequale (T. Moore) Lowe, P. lowei C. Chr., P. guatemalense Hook.
Local name: helecho calaguala.
Salvadoran distribution: Ahuachapán: D. Witsberger 65b (MHES); E.A. Montalvo
3506 (ITIC); M. Sandoval & E. Sandoval 16 (LAGU); T. Croat 42145 (ITIC).
Sonsonate: M.B. Siu s.n. (ITIC); T. Croat 42207 (ITIC). Santa Ana: D. Rodríguez et
al. 5 (B, LAGU, NY); J. Monterrosa & R. Carballo 224, 285 (B, LAGU); J. Monterrosa
et al. 965 (B, NY, LAGU); M.B. Siu s.n. (ITIC); N. López et al. s.n. (JBL04626)
(LAGU); R. Seiler 191, 280 (MHES); R. Villacorta & M. Calderón 375 (LAGU, MO).
San Salvador: R. Seiler 27 (F, MHES); 512 (MHES). Chalatenango: J. Tucker 1031,
1252 (F); R. Seiler 407 (JBL00195) (F, LAGU, MHES); 732 (MHES).
Polypodium polypodioides (L.) Watt. var. polypodioides, Canad. Naturalist & Quart. J.
Sci., ser. 2, 3: 158. 1867.
Acrostichum polypodioides L., Polypodium incanum Sw., Goniophlebium incanum
(Sw.) J. Sm., Marginaria polypodioides (L.) Tidestr., Pleopeltis polypodioides (L.) E.
G. Andrews & Windham
Local name: helecho peinetero amarillo.
Suriname, French Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Uruguay,
Argentina.
Salvadoran distribution: Ahuachapán: F. Chinchilla s.n. (ISB00568) (AAU, B, F,
LAGU); J.M. Rosales 2537 (B, BM, LAGU, MO); 2673 (B, LAGU, MO). La Paz: R.
Villacorta & S. Villacorta 1173 (B, LAGU, MO). Usulután: A. del Cid 1782 (ITIC); D.
Williams 50 (B, LAGU, MO); R. Seiler 588 (MHES). Morazán: J. Monterrosa & C.
Rivera 705 (B, BM, LAGU).
Polypodium polypodioides (L.) Watt. var. aciculare Weath., Contr. Gray Herb. 124: 33.
1939.
Pleopeltis polypodioides (L.) Watt var. acicularis (Weath.) E. G. Andrews & Windham
Local name: helecho peinetero rojo.
Salvadoran distribution: Ahuachapán: E. Sandoval et al. 540, 1291 (B, LAGU); M.
Sandoval & E. Sandoval 94 (LAGU); R. Seiler 828 (ITIC, MHES); 912 (F, ITIC,
MHES); R. Villacorta et al. 880 (B, LAGU, MO). Sonsonate: R. Seiler 78 (MHES).
Santa Ana: B. Pfeiffer-Berendsohn & W. Berendsohn 11 (LAGU, MO); J. Monterrosa
et al. 225 (B, BM, LAGU, MO); R. Seiler 169 (ITIC, MHES). La Libertad: R. Seiler
47 (MHES). San Salvador: R. Seiler 215 (MHES). Usulután: D. Williams 243, 436
(LAGU); E.A. Montalvo 5044 (ITIC); J. Lagos 36 (ITIC); R.A. Carballo et al. 820
(LAGU). San Miguel: A. Salgado 65a (MHES). La Unión: R. Seiler 216 (MHES).
185
FERN GAZ. 18(4): 120-215. 2008
Polypodium puberulum Schltdl. & Cham., Linnea 5: 607. 1830.
Polypodium ehrenbergianum Klotzsch, P. olivaceum Liebm. var. elatum Liebm., P.
pubescens Fée, P. fournieri C. Chr., P. galeottii Mett. ex Kuhn, P. glaberulum Mickel &
Beitel
Salvadoran distribution: Chalatenango: R. Seiler 752 (F, MHES).
Polypodium rhodopleuron Kunze, Linnea 18: 315. 1844.
Polypodium isomeron E. Fourn., P. lesourdianum E. Fourn.
Source: Moran in Davidse et al. (1995): 360; Mickel & Smith (2004): 506.
Polypodium sanctae-rosae (Maxon) C. Chr., Index Filic., Suppl. I: 62. 1913.
Goniophlebium sanctae-rosae Maxon
Salvadoran distribution: Santa Ana: B. Pfeiffer-Berendsohn & W. Berendsohn 9 (F,
LAGU); 12 (F, LAGU, MO); R.A. Carballo et al. 115 (B, LAGU, MO); R. Seiler 166
(F, MHES). San Salvador: R. Seiler 32 (F, MHES); 55 (MHES); S. Winkler 27 (ITIC).
Chalatenango: R. Seiler 257 (F, MHES). San Vicente: E.A. Montalvo 4610, 5087
Polypodium subpetiolatum Hook. in Bentham, Pl. Hartw. 54. 1840.
Polypodium biserratum M. Martens & Galeotti, P. schaffneri Fée, P. guilleminianum E.
Fourn., P. hahnii E. Fourn., P. firmulum Maxon, P. maxonii C. Chr., P. adelphum
Maxon, P. teresae Maxon
Salvadoran distribution: San Vicente: R. Seiler 328 (F, MHES).
♦Polypodium thyssanolepis A. Braun ex Klotzsch, Linnaea 20: 392. 1847.
Goniophlebium thyssanolepis (A. Braun ex Klotzsch) T. Moore, G. rhagadiolepis (Fée)
Fée, Marginaria thyssanolepis (A. Braun ex Klotzsch) Farw., Polypodium incanum Sw.
var. fimbriatum M. Martens & Galeotti, P. rhagadiolepis Fée, P. purpusii Christ
Panama, Antilles, Colombia, Venezuela, Ecuador, Peru, Bolivia.
Salvadoran distribution: Chalatenango: J.Monterrosa et al. 1612 (BM, LAGU, IZTA,
MHES).
Polypodium ursipes Moritz ex C. Chr., Index. Filic. 572. 1906.
Polypodium ambiguum Mett. ex Kuhn
Global distribution: Costa Rica, Panama.
Source: Moran in Davidse et al. (1995): 354.
Serpocaulon falcaria (Kunze) A.R. Sm., Taxon 55(4): 928. 2006.
Goniophlebium calaguala Fée, G. invertens Fée, Polypodium falcaria Kunze.
Salvadoran distribution: Santa Ana: A. Molina et al. 16774 (F; B). PfeifferBerendsohn 23, 37 (LAGU, MO); J. Monterrosa & R. Carballo 852 (B, LAGU, MO);
M.C. Carlson 728 (F); R. Seiler 123, 176 (F, MHES); 651 (MHES); V.M. Martínez s.n.
(CMC01094) (B, BM, EAP, LAGU, MO). San Salvador: R. Seiler 339 (MHES).
Note: The name Polypodium loriceum has been applied by Löetschert (1954), Stolze
(1981), Seiler (1982) and Moran in Davidse et al. (1995) to this species. See discution
as P. falcaria in Mickel & Smith (2004).
Serpocaulon triseriale (Sw.) A.R. Sm., Taxon 55(4): 929. 2006.
Goniophlebium haenkei C. Presl, Polypodium haenkei (C. Presl) Liebm., P. triseriale
Sw.
Local names: helecho calaguala, helecho yema de huevo.
Salvadoran distribution: Ahuachapán: F. Chinchilla & M.J. López s.n. (ISB00635)
(AAU, B, F, LAGU); M. Sandoval & E. Sandoval s.n. (ISB00929) (B, F, LAGU, MO).
San Salvador: E.A. Montalvo 20 (ITIC). Chalatenango: D. Sloot et al. 41
(MAG00470) (LAGU, U). Morazán: J. Monterrosa & C. Rivera 695 (B, LAGU, MO).
Terpsichore anfractuosa (Kunze ex Klotzsch) B. León & A. R. Sm., Amer. Fern J. 93:
86. 2003.
Polypodium anfractuosum Kunze ex Klotzsch, P. saxicola Baker, P. induens Maxon,
Ctenopteris anfractuosa (Kunze ex Klotzsch) Copel., Grammitis anfractuosa (Kunze ex
Klotzsch) Proctor, Melpomene anfractuosa (Kunze ex Klotzsch) A. R. Sm. & R. C.
Moran
Antilles, Guadalupe, Colombia, Venezuela, Guayana, Ecuador, Peru, Bolivia.
Salvadoran distribution: Sonsonate: R. Seiler 604 (MHES). Santa Ana: B. PfeifferBerendsohn et al. 25 (B, LAGU); R. Seiler 648 (MHES). Chalatenango: E.A.
Montalvo s.n. (ITIC). Morazán: R. Seiler 987 (ITIC, MHES); S. Winkler 80 (ITIC).
Terpsichore asplenifolia (L.) A. R. Sm., Novon 3: 485. 1993.
Polypodium asplenifolium L., P. laxifrons Liebm., Ctenopteris asplenifolia (L.) Copel.,
Grammitis asplenifolia (L.) Proctor
Source: Seiler (1982): 385 (as Grammitis asplenifolia); Berendsohn (1989) 1: 16 (as
Grammitis asplenifolia); Moran & Smith in Davidse et al. (1995): 387. Mickel & Smith
2004: 617.
Terpsichore cultrata (Willd.) A. R. Sm., Novon 3: 486. 1993.
Polypodium cultratum Willd., Ctenopteris cultrata (Willd.) Copel., Grammitis cultrata
(Willd.) Proctor
Salvadoran distribution: R. Seiler 1130 (UC) (Moran & Smith in Davidse et al. 1995:
388). Santa Ana: A. Molina et al. 16903 (F); R. Seiler 105 (F, MHES). San Salvador:
S. Winkler 24 (ITIC).
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FERN GAZ. 18(4): 120-215. 2008
♦Terpsichore delicatula (M. Martens & Galeotti) A. R. Sm., Novon 3: 486. 1993.
Polypodium delicatulum M. Martens & Galeotti, Ctenopteris delicatula (M. Martens &
Galeotti) J. Sm., Grammitis delicatula (M. Martens & Galeotti) Proctor
Global distribution: Mexico, Guatemala, Honduras, Jamaica.
Salvadoran distribution: Chalatenango: J.Monterrosa et al. 1624 (BM, LAGU, UC).
Terpsichore lehmanniana (Hieron.) A.R. Sm., Novon 3: 487. 1993.
Polypodium lehmannianum Hieron., P. sublongipes Christ, Ctenopteris lehmanniana
(Hieron.) Copel., Grammitis lehmanniana (Hieron.) C. V. Morton
Global distribution: Belize, Guatemala, Honduras, Nicaragua, Costa Rica, Panama,
Colombia, Ecuador.
Salvadoran distribution: Santa Ana: J. Monterrosa & R. Carballo 858 (B, LAGU); R.
Seiler 171 (F, ITIC, MHES). Morazán: R. Seiler 1062 (F, MHES).
Terpsichore semihirsuta (Klotzsch) A. R. Sm., Novon 3: 488. 1993.
Polypodium semihirsutum Klotzsch, P. semihirsutum Klotzsch var. fuscosetosum
Hieron., Ctenopteris semihirsuta (Klotzsch) Copel., Grammitis semihirsuta (Klotzsch)
C. V. Morton, G. semihirsuta (Klotzsch) C. V. Morton var. fuscosetosa (Hieron.)
Lellinger
Salvadoran distribution: Santa Ana: B. Pfeiffer-Berendsohn 34 (LAGU, MO); R.
Seiler 94, 101, 652 (MHES). Chalatenango: R. Seiler 751 (MHES).
Terpsichore taxifolia (L.) A. R. Sm., Novon 3: 488. 1981.
Polypodium taxifolium L., P. lherminieri Fée var. costaricense Rosenst., Ctenopteris
taxifolia (L.) Copel., Grammitis taxifolia (L.) Proctor
Global distribution: Costa Rica, Panama, Antilles, Colombia, Venezuela, Suriname,
Source: Seiler (1982): 386 (as Grammitis taxifolia); Berendsohn (1989) 1: 16 (as
Grammitis taxifolia); Moran & Smith in Davidse et al. (1995): 392.
PSILOTACEAE
Psilotum complanatum Sw., J. Bot. (Schrader) 1800(2): 110. 1801.
Local name: helecho palma de esquipulas.
Global distribution: Mexico, Guatemala, Honduras, Antilles, Colombia, Venezuela,
Peru, Malasia, Oceanía.
Salvadoran distribution: Ahuachapán: E. Sandoval s.n. (MS-00325) (B, LAGU, MO);
E. Sandoval & F. Chinchilla 54 (B, LAGU); M. Sandoval & E. Sandoval 247 (B,
LAGU, MO). San Vicente: Löetschert 252 (EAP) (Palacios-Río in Davidse et al.
1995).
♦Psilotum nudum (L.) P. Beauv., Prodr. Aethéogam. 106, 112. 1805.
Lycopodium nudum L., Psilotum triquetrum Sw.
Global distribution: USA, Mexico, Guatemala, Belice, Honduras, Nicaragua, Costa
Rica, Panama, Antilles, Colombia, Venezuela, Guyana, Suriname, French Guyana,
Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Old World tropics, Australia, New
Zealand.
Salvadoran distribution: La Unión: R. Ibarra, R. Juárez, F. Franco, I. Vega & O.
Jiménez s.n. (JBL05114) (BM, LAGU, MHES); J. Monterrosa & R. Ibarra 1608 (BM,
LAGU, MHES, MO).
PTERIDACEAE
Acrostichum aureum L., Sp. Pl. 2: 1069. 1753.
Suriname, French Guyana, Ecuador, Brazil, Paraguay, Trópicos del Viejo Mundo.
Salvadoran distribution: Ahuachapán: R. Seiler 137 (MHES); R. Villacorta & E.A.
Montalvo 826 (B, LAGU, MEXU). La Unión: K. Sidwell et al. 674 (B, BM, ITIC,
LAGU, MO).
Acrostichum danaeifolium Langsd. & Fisch., Pl. Voy. Russes Monde I: 5, t. I. 1810.
Chrysodium lomarioides Jenman, C. lomarioides Jenman var. hastatum Christ,
Acrostichum lomarioides (Jenman) Jenman., A. excelsum Maxon
Local names: helecho de manglar, helecho de pantano.
Suriname, French Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay.
Salvadoran distribution: Ahuachapán: J.M. Rosales 2191 (B, LAGU). La Libertad:
M. Magaña 48 (MP-00008) (LAGU). San Miguel: J. Tucker 908 (F); R. Villacorta
2889 (B, BM, LAGU).
Adiantum amplum C. Presl, Reliq. Haenk. I(I): 63. 1825.
Adiantum princeps T. Moore
Local names: helecho de arriero, centavito.
Salvadoran distribution: Ahuachapán: A.K. Monro et al. 1925, 3642, 3700 (B, BM,
ITIC, LAGU, MO); A. Sermeño 106 (JBL01020) (B, LAGU, MEXU, MO); J.M.
Rosales 5 (AAU, B, BM, LAGU, MEXU, MO); 2415 (B, BM, LAGU, MEXU, MO);
M. Sandoval & E. Sandoval 304 (B, F, LAGU); R. Seiler 938 (F); T, Croat 42060,
42061, 42095, 42102 (ITIC); W. Berendsohn 1311 (LAGU, MO); 1390, 1391 (B,
LAGU, MO). Santa Ana: L. Lara 221 (MHES). La Libertad: A.K. Monro et al. 3770
(B, BM, ITIC, LAGU, MO); B. Pfeiffer-Berendsohn 68 (B, LAGU, MO); E.A.
Montalvo s.n. (ITIC); J. Monterrosa et al. s.n. (JBL04488) (LAGU); K. Sidwell et al.
465 (B, BM, ITIC, LAGU, MO); N. Ventura 5012, 5015 (ITIC). San Salvador: K.
Sidwell et al. 582 (B, BM, ITIC, LAGU, MO); R. Seiler s.n. (12-326) (MHES); 72 (F,
MHES); 711 (F). Cuscatlán: S. Dar s.n. (JBL03659) (B, BM, LAGU, MO). Morazán:
J. Monterrosa & C. Rivera 675 (BM, LAGU). La Unión: A.K. Monro et al. 2753 (B,
BM, ITIC, LAGU, MO).
Adiantum andicola Liebm., Mexic. Bregn. 266 (reprint 114). 1849.
Adiantum cuneatum Langsd. & Fisch. var. angustifolium M. Martens & Galeotti, A.
tenerum Sw. var. dissectum M. Martens & Galeotti, A. amabile Liebm., A.
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FERN GAZ. 18(4): 120-215. 2008
glaucophyllum Hook. Local name: helecho centavito.
Colombia.
MO). Sonsonate: T. Croat 42212 (ITIC). Santa Ana: A. Molina et al. 16881 (F); D.
Rodríguez & G. Cerén s.n. (JBL04597) (B, LAGU); G. Davidse et al. 37231 (B, BM,
ITIC, LAGU, MO); J. Monterrosa et al. 917 (LAGU, NY); 1024 (LAGU); L. Lara 226
(MHES); R. Seiler 135, 277, 463 (F, MHES). La Libertad: M. Renderos et al. 413
(LAGU, MO). San Salvador: E.A. Montalvo 3155 (ITIC); M.C. Carlson 383, 472 (F);
R. Seiler 211, 517 (F, MHES); S. Winkler 18 (ITIC). Chalatenango: R. Seiler 404
(JBL00080) (F, LAGU, MHES); 730 (F, MHES); W. Berendsohn & H. Sipman 1585
(B, LAGU). San Vicente: R. Seiler 320, 329 (F, MHES). Usulután: D. Williams 136
(B, LAGU, MO). Morazán: J. Tucker 607 (F).
Adiantum braunii Mett. ex Kuhn, Linnea 36: 75. 1869.
Adiantum braunii Mett. ex Kuhn var. cuneatum Mett. ex Kuhn, A. convolutum E. Fourn.
Local names: helecho nido de arriero, helecho nido de pijuyo.
Global distribution: Mexico, Guatemala, Nicaragua, Costa Rica, Colombia, Venezuela,
Ecuador.
Salvadoran distribution: Ahuachapán: E. Sandoval & M. Sandoval 1401 (B, LAGU).
Santa Ana: W. Berendsohn et al. 1536 (B, LAGU, MO). La Libertad: A.K. Monro et
al. 3083 (B, BM, ITIC, LAGU, MO); J.C. González & M.A. Hernández 364 (B, LAGU,
MO). San Salvador: R. Seiler 1502 (F). Chalatenango: R. Seiler 550 (MHES).
Adiantum concinnum Humb. & Bonpl. ex Willd., Sp. Pl., ed. 4, 5(1): 451. 1810.
Adiantum lutescens Moungeot ex Fée
Local names: helecho cola de arriero, helecho nido de arriero, helecho palmita.
Panama, Antilles, Colombia, Venezuela, Trinidad and Tobago, Ecuador, Peru, Brazil.
MO); E. Sandoval & F. Chinchilla 704 (B, LAGU); E. Sandoval & M. Sandoval 1402
(LAGU); 1521 (B, LAGU); F. Chinchilla s.n. (ISB00534), s.n. (ISB00736), s.n.
(ISB00768) (AAU, B, F, LAGU); M. Magaña 7 (MP-00002) (LAGU); M. Sandoval &
E. Sandoval 101 (B, F, LAGU); R. Villacorta et al. 942 (B, LAGU, MO); T. Croat
42115 (ITIC). Sonsonate: D. Rodríguez & M. Trejo 83 (LAGU). Santa Ana: L. Lara
222 (MHES); M.L. Reyna de Aguilar & R. Villacorta 1438 (B, LAGU, MO). La
Libertad: A.K. Monro et al. 2335, 3084 (B, BM, ITIC, LAGU, MO); J. Reyes et al.
(JBL04510) (LAGU); J. Williams s.n. (ITIC); K. Sidwell et al. 483, 698 (B, BM, ITIC,
LAGU, MO); M.B. Siu s.n. (ITIC); N. Ventura 5007, 5011, 5025 (ITIC); R. Amador et
al. s.n. (JBL04461) (LAGU); R. Menjivar & J. Sermeño s.n. (JBL04484) (LAGU); V.
Salzman s.n. (JBL02066) (LAGU). San Salvador: A.K. Monro et al.3437 (B, BM,
ITIC, LAGU, MO); J. Williams 94 (ITIC); K. Sidwell et al. 577 (B, BM, ITIC, LAGU,
MO); M.C. Carlson 2 (F); R. Seiler 511 (F, MHES); Tobar s.n. (12-10) (MHES); W.
Heed 25 (ITIC). Chalatenango: L. Lara 207, 214, 217 (MHES); M.B. Siu s.n. (ITIC);
R. Seiler 141 (MHES). Usulután: A.K. Monro et al. 3000 (B, BM, ITIC, LAGU, MO).
San Miguel: J. Tucker 976 (F). Cabañas: A.K. Monro et al. 2775 (B, BM, ITIC,
LAGU, MO); G. Davidse et al. 37111 (B, BM, ITIC, LAGU, MO); R. Seiler 527 (F,
MHES). Morazán: J. Tucker 640 (F).
Adiantum deflectens Mart., Icon. Pl. Crypt. 94. 1834.
Adiantum deflectens Mart. var. tremulum Hieron., A. delicatulum Mart., A. dissimile
Schrad., A. dolabriforme Hook., A. filiforme Gardner, A. flagellum Fée, A. lunatum
Keyserl., A. subaristatum Fée
Colombia, Venezuela, French Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay.
Source: Stolze (1981): 22; Berendsohn (1989) 1: 19; Moran in Davidse et al. (1995):
111; Mickel & Smith (2004): 29.
Adiantum latifolium Lam., Encycl. I: 43. 1783.
Adiantum lucidum Sw. var. bipinnatum Mett. ex E. Fourn.
Antilles, Colombia, Venezuela, Africa, Asia.
Local names: helecho rosquilla
MO). Usulután: J. Lagos 1084 (ITIC); R. Seiler 591 (F, MHES).
Adiantum lunulatum Burm. f., Fl. Indica 235. 1768.
Local names: helecho cola de arriero, helecho pie de zanate, helecho salamanquera.
Antilles, Colombia, Venezuela, Africa, Asia.
Salvadoran distribution: Ahuachapán: E. Sandoval et al. 609 (B, LAGU); 1398 (B, K,
LAGU); F. Chinchilla s.n. (ISB00605), s.n. (ISB00769) (AAU, B, F, LAGU); J.M.
Rosales 1133, 1134, 1167, 1316, AAU (B, BM, LAGU). Sonsonate: D. Rodríguez &
M. Trejo 64, B (LAGU, NY). Santa Ana: J.L. Linares 535 (JBL01985) (LAGU); K.
Prestegard 43 (MP-00003) (LAGU); R. Seiler 709 (MHES); W. Berendsohn et al. 1539
(B, LAGU). La Libertad: A.K. Monro et al. 3404 (B, BM, ITIC, LAGU, MO); N.
Ventura 5006 (ITIC). San Salvador: R. Seiler 376, 446 (F, MHES). La Paz: E.A.
Montalvo 6205 (B, BG, F, HBG, ITIC, LAGU, MO). San Miguel: R. Seiler 544 (F,
MHES). Cabañas: R. Seiler 524 (F, MHES). Morazán: J. Tucker 452 (F).
Adiantum macrophyllum Sw., Prod. 135. 1788.
Local name: helecho palma de esquipulas, helecho palma de roca.
Salvadoran distribution: Ahuachapán: E. Sandoval et al. 106 (LAGU); s.n.
(JBL01386), 1293 (B, LAGU); F. Chinchilla s.n. (ISB00660) (AAU, B, F, LAGU); M.
Sandoval et al. s.n. (ISB00842) (AAU, B, F, LAGU); 273 (B, F, LAGU); P. Standley et
al. 2673 (F); R. Seiler 932 (MHES). La Libertad: D. Matamoro s.n. (JBL01317)
(LAGU); M.C. Carlson 143 (F); R. Seiler 717 (MHES). San Salvador: E.A. Montalvo
15a (ITIC); R. Seiler 449, 508, 530 (F, MHES).
Adiantum patens Willd., Sp. Pl., ed. 4, 5(1): 439. 1820.
Adiantum lobatum C. Presl
Local names: helecho nido de arriero, helecho pata de paloma, petatillo rojo.
Ecuador, Peru, Bolivia, Africa.
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FERN GAZ. 18(4): 120-215. 2008
Salvadoran distribution: Ahuachapán: E. Sandoval et al. 589, 1362 (B, LAGU). Santa
Ana: R. Seiler 133 (MHES); 761 (F, MHES); T. Croat 42288 (ITIC). La Libertad: J.
Monterrosa et al. s.n. (JBL04500) (LAGU); N. Ventura 5018 (ITIC). San Salvador: R.
Seiler 448, 636 (F, MHES). Chalatenango: D. Sloot & A. Reina 135 (MAG00564) (B,
LAGU, U). Morazán: J. Tucker 641 (F); S. Winkler 19 (ITIC).
Adiantum petiolatum Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck.
Gesammten Naturk 5: 326. 1811.
Adiantum lucidum (Cav.) Sw. var. pinnatum E. Fourn.
Salvadoran distribution: Chalatenango: R. Seiler 138 (F, ITIC, MHES). San Vicente:
R. Seiler 973 (F, ITIC, MHES).
Adiantum poiretii Wikstr., Kongl. Vetensk. Akad. Handl. 1825: 443. 1826.
Adiantum thalictroides Willd. ex Schltdl., A. pellucidum M. Martens & Galeotti, A.
gratum Fée
Antilles, Colombia, Venezuela, Ecuador, Peru, Bolivia, Chile, Argentina, Viejo Mundo.
Salvadoran distribution: Santa Ana: R. Villacorta & L. Lara 2517 (B, LAGU).
Chalatenango: W. Berendsohn & H. Sipman 1586 (B, LAGU).
Adiantum pulverulentum L., Sp. Pl. 2: 1096. 1753.
MO); D. Witsberger 702 (MHES); R. Seiler 818 (F, ITIC, MHES). Sonsonate: D.
Rodríguez & M. Trejo 205 (LAGU, NY).
Adiantum raddianum C. Presl, Suppl. Tent. Pterid. 158. 1836.
Adiantum cuneatum Langsd. & Fisch., A. mexicanum C. Presl, A. werckleanum Christ
Local name: helecho colchón de guardabarranco.
Global distribution: Mexico, Guatemala, Nicaragua, Costa Rica, Antilles, Colombia,
Venezuela, Trinidad and Tobago, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina,
Uruguay.
Salvadoran distribution: Ahuachapán: M. Magaña 14 (MP-00001) (LAGU); W.
Berendsohn et al. 1392 (B, LAGU). Usulután: A.K. Monro et al. 3024 (B, BM, ITIC,
LAGU, MO).
Note: A.K. Monro & D. Alexander 3084 is cited erroneously as A. raddianum in Monro
et al. 2002, it is A. concinnum Humb. & Bonpl. ex Willd.
Adiantum tenerum Sw., Prodr. 135. 1788.
Adiantum trapezoides Fée
Rica, Antilles, Venezuela, Trinidad and Tobago.
Source: Maxon & Standley (1930): 172 (as Adiantum trapezoides). Calderón &
Standley (1941): 29 (as Adiantum trapezoides). Seiler (1982): 384. Berendsohn (1989)
1: 20. Moran et al. in Davidse et al. (1995): 110.
Adiantum tetraphyllum Humb. & Bonpl. ex Willd., Sp. Pl., ed. 4, 5(1): 441. 1810.
Adiantum fruticosum Poepp. ex Spreng., A. tetraphyllum Humb. & Bonpl. ex Willd. var.
costaricense Christ
Local names: helecho ciempiés, helecho rosquilla pacha.
Panama, Antilles, Colombia, Venezuela, Trinidad and Tobago, Ecuador, Peru, Bolivia,
Paraguay.
Salvadoran distribution: Ahuachapán: E. Sandoval et al. 1119, 1207 (B, LAGU, MO);
F. Chinchilla s.n. (ISB00315) (LAGU); s.n. (ISB00661) (AAU, B, F, LAGU); M.
Sandoval et al. s.n. (ISB00815), AAU, B, F, AAU; 298 (B, F, LAGU); R. Morales et al.
s.n. (JBL01274) (B, LAGU, MO); R. Seiler 924 (F, ITIC, MHES); R. Villacorta 623 (B,
LAGU, MO); W. Berendsohn et al. 1355, 1369, 1398 (B, LAGU, MO); 1428 (B,
LAGU). Santa Ana: R. Villacorta & J. María 2203 (B, LAGU, MO). San Salvador: R.
Seiler 531 (F, MHES). Chalatenango: R. Seiler 265 (F, MHES). Cabañas: A.K. Monro
et al. 3747 (B, BM, ITIC, LAGU, MO); J. Monterrosa & R. Carballo 230 (AAU, B,
BM, LAGU, MO). Morazán: J. Monterrosa & C. Rivera 673 (B, BM, LAGU, MO).
Adiantum trapeziforme L., Sp. Pl. 2: 1097. 1753.
Adiantum trapeziforme L. var. oblongatum T. Moore, A. trapeziforme L. var. plumieri
T. Moore
Local names: cabello de ángel, helecho nido de arriero, lantrillo.
Antilles, Trinidad and Tobago, Bolivia, Paraguay; naturalised in Asia and the Pacific
Islands.
Salvadoran distribution: Ahuachapán: E. Sandoval & F. Chinchilla 177, 266 (LAGU);
F. Chinchilla & R.E. Sandoval s.n. (ISB00589) (AAU, B, F, LAGU); M. Sandoval & E.
Sandoval s.n. (ISB00838) (AAU, B, LAGU); P. Standley et al. 3024 (F); R. Seiler 937
(F, MHES); T. Croat 42103 (ITIC); W. Berendsohn et al. 1434 (B, LAGU, MO). Santa
Ana: A.K. Monro & R. Douglas 3464 (B, BM, ITIC, LAGU, MO). La Libertad: O.
Pank s.n. (MP-00004) (LAGU). San Salvador: K. Sidwell et al. 580 (B, BM, ITIC,
LAGU, MO); M.C. Carlson 20, 327 (F); R. Seiler 67 (F, ITIC, MHES); 785 (F, MHES).
Chalatenango: R. Seiler 139 (F, MHES). Usulután: E.A. Montalvo 5101 (ITIC); S.
Winkler 16 (ITIC). Cuscatlán: S. Dar s.n. (JBL03658) (LAGU, MO). La Unión: A.K.
Monro et al. 3536 (B, BM, ITIC, LAGU, MO).
Adiantum trichochlaenum Mickel & Beitel, Mem. New York Bot. Gard. 46: 29-30.
1988.
Salvadoran distribution: San Salvador: R. Seiler 455 (F, MHES).
Adiantum villosum L., Syst. Nat., ed. 10, 2: 1328. 1759.
Local names: helecho ciempiés, helecho rosquilla pacha.
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FERN GAZ. 18(4): 120-215. 2008
J.M. Rosales 1419 (LAGU); M. Sandoval et al. 29 (B, LAGU); T. Croat 42087 (ITIC);
W. Berendsohn & R. Villacorta 1350 (LAGU, MO). Santa Ana: M.L. Reyna de Aguilar
663 (MHES). La Libertad: R. Seiler 716 (F, MHES). Chalatenango: R. Seiler 140 (F,
MHES). San Vicente: M.C. Carlson 645 (F). Usulután: R. Seiler 592 (F, MHES). San
Miguel: A. Salgado 71 (MHES); J. Tucker 848 (F). Cabañas: R. Seiler 225
(JBL04805) (F, LAGU, MHES).
Ananthacorus angustifolius (Sw.) Underw. & Maxon in Maxon, Contr. U.S. Natl.
Herb. 10: 487. 1908.
Pteris angustifolia Sw., Taenitis angustifolia (Sw.) Spreng., Vittaria angustifolia (Sw.)
Baker in Martius, Vittaria costata Kunze
Local name: helecho barbiquejo.
Global distribution: Mexico, Guatemala, Belize, Honduras, Costa Rica, Panama,
Salvadoran distribution: Ahuachapán: E. Sandoval et al. 439, 1303 (B, LAGU); J.M.
Rosales 1466 (B, BM, LAGU, MO); 1513 (B, LAGU, MO); M.L. Reyna de Aguilar
652 (MHES); M. Sandoval & E. Sandoval s.n. (ISB00930) (LAGU). San Salvador: R.
Seiler 459, 788 (F, MHES).
Anogramma chaerophylla (Desv.) Link, Fil. Spec. 138. 1841.
Gymnogramma chaerophylla Desv.
Global distribution: Mexico, Antilles, Brazil, Paraguay, Argentina, Uruguay.
Salvadoran distribution: San Salvador: H. Weber 24 (ITIC); R. Seiler 20 (MHES).
Anogramma leptophylla (L.) Link, Fil. Spec. 137. 1841.
Polypodium leptophyllum L., Gymnogramma leptophylla (L.) Desv. var. mexicana
Kunze, Pityrogramma guatemalensis Domin, Anogramma guatemalensis (Domin) C.
Chr.
Global distribution: Mexico, Guatemala, Nicaragua, Costa Rica, Panama, Venezuela,
Ecuador, Peru, Brazil, Eurasia, Africa.
Salvadoran distribution: Sonsonate: R. Seiler 610, 616, 629 (F, MHES). Santa Ana:
E.A. Montalvo 4895 (ITIC); J. Williams 31 (ITIC); R. Seiler 110 (F, ITIC, MHES); 658
(F, MHES). San Salvador: Castañeda s.n. (12-897) (MHES). Usulután: D. Williams &
R.W. Herrera 424 (LAGU).
Bommeria pedata (Sw.) E. Fourn., Bull. Soc. Bot. France 27: 327. 1880.
Hemionitis pedata Sw., Gymnogramma pedata (Sw.) Kaulf., Gymnopteris pedata (Sw.)
C. Chr.
Local name: helecho mano de mapache.
Salvadoran distribution: Ahuachapán: M. Sandoval & E. Sandoval 40 (B, F, LAGU).
Santa Ana: R. Seiler 498 (F, MHES). San Salvador: R. Seiler 46 (F, MHES); 640
(MHES); O. Pank s.n. (12-987) (MHES).
Ceratopteris pteridoides (Hook.) Hieron., Bot. Jahrb. Syst. 34: 561. 1905.
Parkeria pteridoides Hook.
Local name: helecho acuático.
Salvadoran distribution: Ahuachapán: J.M. Rosales 2674 (B, LAGU, MO). La Paz:
N.C. Fassett 28557 (EAP, F); 29180 (F). San Miguel: N.C. Fassett 29391 (F).
Ceratopteris thalictroides (L.) Brongn., Bull. Sci. Soc. Philom. Paris, sér. 3, 8: 186.
1821.
Acrostichum thalictroides L.
Local name: helecho de agua.
Global distribution: USA, Mexico, Guatemala, Nicaragua, Costa Rica, Panama,
Venezuela, Suriname, Ecuador, Brazil, Africa, Asia, Australia, Pacific Islands.
Salvadoran distribution: Ahuachapán: J.M. Rosales 1921 (B, LAGU, MO); 2070
(LAGU). San Vicente: N.C. Fassett 29157 (F); 29162 (EAP, F). Usulután: R.A.
Carballo & S. Santamaría 899 (B, LAGU, MO).
Cheilanthes angustifolia Kunth, Nov. Gen. Sp. I: 21. 1815.
Allosorus angustifolius (Kunth) C. Presl, A. angustifolius (Kunth) C. Presl var. minor
Liebm., Onychium angustifolium (Kunth) Kunze, Pellaea angustifolia (Kunth) Baker in
Hooker & Baker, Cheilanthes venusta Fée var. aurata Fée
Local name: helecho tuya morada.
J.M. Rosales 1230 (B, LAGU, MO); R. Villacorta & Martínez 626 (B, LAGU, MO); W.
Berendsohn & R. Villacorta 1384 (B, LAGU, MO). Sonsonate: R. Seiler 37 (F,
MHES). Santa Ana: B. Pfeiffer-Berendsohn & W. Berendsohn 13 (F, LAGU, MO); S.
Winkler 28 (ITIC). La Libertad: E.A. Montalvo s.n. (ITIC). San Salvador: H.
Castañeda s.n. (12-899) (MHES); E.A. Montalvo s.n. (ITIC). Chalatenango: L. Lara
213, 231 (MHES); R. Seiler 559 (F, MHES); S. Calderón 2476 (F). San Vicente: E.A.
Montalvo 5095 (ITIC). Usulután: D. Williams & R.W. Herrera 301 (LAGU, MO).
Morazán: J. Monterrosa & C. Rivera 689 (B, BM, LAGU, MO); M. Renderos et al.
611 (LAGU, MO).
Cheilanthes aurea Baker in Hooker & Baker var. aurea, Syn. Fil. 453. 1868.
Aleuritopteris aurea (Baker) Ching
Salvadoran distribution: Ahuachapán: J.M. Rosales 1229 (B, LAGU); W. Berendsohn
et al. 1644 (BM, LAGU). San Salvador: R. Seiler 23, 586 (F, MHES).
Cheilanthes bolborrhiza Mickel & Beitel, Mem. New York Bot. Gard. 46: 107. 1988.
Salvadoran distribution: San Salvador: R. Seiler 45, 579 (F, LAGU, MHES, NY).
Cheilanthes brachypus Kunze, Linnea 23: 307. 1850.
Cheilanthes squamosa Hook. & Grev. var. brachypus Kunze, Notholaena brachypus
(Kunze) J. Sm. Local names: helecho de bulto, helecho dorado, helecho pluma de búho.
MO); E. Sandoval & F. Chinchilla 468 (B, LAGU, MO); E. Sandoval & M. Sandoval
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FERN GAZ. 18(4): 120-215. 2008
1404 (B, LAGU); J.M. Rosales 1289, 1587 (AAU, B, BM, LAGU, MO); M. Sandoval
et al. 61 (B, F, LAGU); R. Seiler 926 (F, MHES); R. Villacorta & Martínez 404 (F,
LAGU, MO); T. Croat 42155 (ITIC); W. Berendsohn et al. 1408 (B, LAGU, MO).
Sonsonate: R. Seiler 7 (F, MHES). Santa Ana: K. Prestegard 30 (LAGU); T. Croat
42238 (ITIC). La Libertad: A.K. Monro et al. 2919 (B, BM, ITIC, LAGU, MO); E.A.
Montalvo & R. Villacorta 6266 (B, LAGU, MO); J. Flores s.n. (JF-00106) (LAGU); R.
Villacorta s.n. (JBL00543) (B, LAGU, MO). San Salvador: R. Seiler 444 (F, MHES).
Chalatenango: L. Lara 212, 216 (MHES); R. Seiler 367, 551 (F, MHES). San Vicente:
O. Rohweder 2643 (JBL01248), 2644 (JBL01247) (LAGU). San Miguel: A. Salgado
63 (MHES); R. Seiler 540 (F, MHES); R. Villacorta 2391 (LAGU, MO). Morazán: J.
Monterrosa & C. Rivera 709 (B, BM, LAGU, MO). La Unión: A.K. Monro et al. 3540
(B, BM, ITIC, LAGU, MO).
Cheilanthes chaerophylla (M. Martens & galeotti) Kunze, Linnea 23: 243, 307.
1850.
Allosorus chaerophyllus M. Martens & Galeotti
Salvadoran distribution: Chalatenango: J. Monterrosa et al. 1611 (BM, LAGU, NY).
Cheilanthes decurrens Mickel, Mem. New York Bot. Gard. 88: 191-192. 2004.
Salvadoran distribution: Santa Ana: R. Seiler 100 (F, MHES). San Salvador: R. Seiler
1209 (F, NY).
Cheilanthes kaulfussii Kunze, Linnea 13: 145. 1839.
Cheilanthes viscosa Link, C. glandulifera Liebm.
Global distribution: USA, Mexico, Guatemala, Honduras, Costa Rica, Colombia,
Venezuela.
Salvadoran distribution: Sonsonate: R. Seiler 10 (F, MHES); 1466 (F). Santa Ana: R.
Seiler 680 (F, MHES). Usulután: D. Williams & R.W. Herrera 348 (LAGU). Morazán:
Cheilanthes marginata Kunth, Nov. Gen. Sp. I: 22. 1815; 7: t. 669. 1825.
Allosorus ciliata C. Presl var. minor C. Presl, C. hirsuta Link, C. rufescens Link, C.
crenulata Link
Venezuela, Ecuador, Peru, Bolivia, Argentina.
Salvadoran distribution: Santa Ana: B. Pfeiffer-Berendsohn 47 (LAGU, MO); 54 (B,
LAGU, MO); R. Seiler 298, 473 (F, MHES); 1208 (F); R. Villacorta & J.C. González
1160 (LAGU). Chalatenango: R. Seiler 383 (F, MHES); W. Berendsohn & H. Sipman
1582 (B, LAGU). Morazán: R. Seiler 986 (F, ITIC, MHES); 1069 (F, MHES); 1071
(F).
Note: Seiler (1982), Berendsohn (1989) and R.C. Moran in Davidse et al. 1995
recognise Cheilanthes hirsuta as a distinct species.
Cheilanthes myriophylla Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten
Entdeck. Gesammten Naturk. 5: 328. 1811.
Cheilanthes paleacea M. Martens & Galeotti, C. intermedia Baker, C. fournieri C. Chr.,
Myriopteris intermedia E. Fourn.
Global distribution: Mexico, Guatemala, Honduras, Nicaragua, Antilles, Colombia,
Venezuela, Ecuador, Peru, Brazil, Bolivia, Chile, Argentina.
Salvadoran distribution: R. Seiler 1471 (MO) (Yatskievych & Moran in Davidse et al.
1995).
Cheilanthes pyramidalis Fée, Mém. Foug. 7: 38, t. 25 (F). 3. 1857.
Pellaea angustifolia (Kunth) Baker var. elongata Rovirosa
Salvadoran distribution: Santa Ana: B. Pfeiffer-Berendsohn & S.A. de Reyna 39 (K,
LAGU); M.L. Reyna de Aguilar 766 (MHES); R. Seiler 42 (JBL04800) (F, ITIC,
LAGU, MHES); 351 (F, MHES). San Salvador: R. Seiler 24 (F, MHES).
Cheilanthes skinneri (Hook.) R. M. Tryon & A. F. Tryon, Rhodora 83: 133. 1981.
Pellaea skinneri Hook., P. flavescens E. Fourn., Doryopteris fournieri (Baker) C. Chr.
Local names: helecho cola de pavo real, helecho morado.
Global distribution: Mexico, Guatemala, Nicaragua, Costa Rica, Panama, Colombia.
Salvadoran distribution: Ahuachapán: E. Sandoval et al. 521, 1400 (B, LAGU); J.M.
Rosales 934, 951 (LAGU). Santa Ana: R. Seiler 1201 (F, ITIC, MHES). La Libertad:
R. Seiler 1116 (F, ITIC, MHES). San Miguel: R. Seiler 538 (JBL04808) (F, LAGU,
MHES).
Cheiloplecton rigidum (Sw.) Fée var. rigidum, Mém. Foug. 7: 34, t. 20 (F). 3. 1857.
Pteris rigida Sw., P. cartilaginea C. Presl, Pellaea rigida (Sw.) Hook.
Doryopteris concolor (Langsd. & Fisch.) Kuhn in v.d. Decken, Reisen Ost-Afrika 3(3):
19. 1879.
Pteris concolor Langsd. & Fisch., Cheilanthes concolor (Langsd. & Fisch.) R. M.
Tryon & A. F. Tryon . Local name: helecho de roca.
Global distribution: Mexico, Guatemala, Honduras, Antilles, Colombia, Venezuela,
Ecuador, Peru, Brazil Bolivia, Paraguay, Argentina, Uruguay, Africa, Asia, Australia.
Salvadoran distribution: Sonsonate: M.L. Reyna de Aguilar 1452 (LAGU, MO). San
Salvador: R. Seiler 585 (F, MHES). San Miguel: R. Seiler 539 (F, MHES); R.
Villacorta 2481 (B, LAGU, MO).
Eriosorus flexuosus (Desv.) Copel. var. flexuosus, Gen. Fil. 58. 1947.
Gymnogramma flexuosa Desv., Grammitis flexuosa Kunth
Antilles, Colombia, Venezuela, Guayana, Ecuador, Peru, Brazil, Bolivia.
Salvadoran distribution: Santa Ana: J. Lagos 13 (ITIC); R. Seiler 177 (JBL04804) (F,
LAGU, MHES); R. Villacorta et al. 1048 (B, LAGU, MO); W. Berendsohn 1069 (B,
BM, LAGU, MO). Chalatenango: J. Tucker 1120 (F).
Hemionitis levyi E. Fourn., Bull. Soc. Bot. France 17: 237. 1870.
Hemionitis otonis Maxon
Global distribution: Mexico, Nicaragua, Costa Rica.
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FERN GAZ. 18(4): 120-215. 2008
Salvadoran distribution: S. Calderón 1710, MO (Ranker in Davidse et al. 1995).
Ahuachapán: J.M. Rosales 1187 (B, LAGU). San Miguel: R. Villacorta 2392 (B,
LAGU, MO).
Hemionitis palmata L., Sp. Pl. 2: 1077. 1753.
Salvadoran distribution: P.Standley 19894 (GH) (Ranker in Davidse et al. 1995).
Source: Maxon & Standley 1930: 173; Calderón & Standley (1941): 31; Seiler (1982):
383; Berendsohn (1989) 1: 21; Ranker in Davidse et al. (1995): 132; Mickel & Smith
(2004): 326.
Hemionitis pinnatifida Baker in Hooker & Baker, Syn. Fil. 399. 1868.
Local names: helecho ostra.
Sonsonate: R. Seiler 493 (MHES). Santa Ana: O. Rohweder 2641 (JBL01249), 2689
(JBL01251), 2690 (JBL01250) (LAGU). La Libertad: W. Berendsohn 1024 (LAGU,
MO). San Salvador: M. Renderos 387 (B, LAGU, MO); O. Pank s.n. (12-959)
(MHES); R. Seiler 442 (F, MHES); 505 (MHES). Chalatenango: R. Villacorta 2394
(AAU, B, LAGU, MO). San Miguel: R. Seiler 59 (F, MHES). Morazán: J. Monterrosa
& R. Carballo 535 (B, LAGU, MO).
Mildella intramarginalis (Kaulf. ex Link) Trevis., Rendiconti Reale Ist. Lombardo Sci.,
Cl. Sci. Mat., ser. 2, 9: 810. 1876.
Pteris intramarginalis Kaulf. ex Link, Cheilanthes intramarginalis (Kaulf. ex Link)
Hook., Pellaea intramarginalis (Kaulf. ex Link) J. Sm.
Galápagos.
Salvadoran distribution: Sonsonate: R. Seiler 9 (F, MHES). Santa Ana: B. PfeifferBerendsohn & W. Berendsohn 14, 16 (LAGU); G. Davidse et al. 37247 (B, BM, ITIC,
LAGU, MO); J. Lagos 1422 (ITIC); J. Monterrosa et al. 223 (B, BM, LAGU, MO); 994
(LAGU); M.B. Siu s.n. (ITIC); N. López et al. s.n. (JBL04635) (LAGU); R. Seiler 352
(F, MHES); 898 (F); R. Villacorta et al. 1159 (LAGU); 1052, 1257 (B, LAGU, MO).
San Salvador: A.K. Monro et al. 2234 (B, BM, ITIC, LAGU, MO); M.C. Carlson 424
(F); R. Seiler 30 (JBL04799) (F, LAGU, MHES); 58 (F). Chalatenango: L. González
1711 (ITIC); L. Lara 215 (MHES); R. Seiler 577 (F, MHES). San Vicente: J. Lagos 66
(ITIC). Usulután: D. Williams 57, 60 (B, LAGU, MO). Morazán: R. Seiler 1088b (F,
ITIC, MHES).
Notholaena candida (M. Martens & Galeotti) Hook., Sp. Fil. 5: 110. 1864.
Cheilanthes candida M. Martens & Galeotti, Aleuritopteris candida (M. Martens &
Galeotti) Fée, Chrysochosma candida (M. Martens & Galeotti) Kümmerle
Salvadoran distribution: Sonsonate: R. Seiler 497 (F, MHES). San Miguel: A. Salgado
62 (MHES).
Notholaena meridionalis Mickel, Mem. New York Bot. Gard. 88: 421-422. 2004.
MO). Santa Ana: R. Seiler 88 (F, MHES, NY).
Pellaea ternifolia (Cav.) Link. subsp. ternifolia, Fil. Spec. 59. 1841.
Pteris ternifolia Cav.
Antilles, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, Chile, Paraguay,
Argentina, Hawaii.
Salvadoran distribution: Santa Ana: R. Seiler 679 (MHES); 467 (F).
Pityrogramma calomelanos (L.) Link var. calomelanos, Handbuch 3: 20. 1833.
Acrostichum calomelanos L., Gymnogramma calomelanos (L.) Kaulf., Ceropteris
serrata Fée
Local name: helecho pinta mano.
Suriname, French Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina.
Salvadoran distribution: Ahuachapán: E. Sandoval et al. 279 (B, LAGU, MO); 1103,
1301 (B, LAGU); F. Chinchilla s.n. (ISB00093) (B, LAGU, MO); J.M. Rosales 117,
1228 (B, LAGU); 788 (B, LAGU, MO); M. Magaña 8 (MP-00006) (LAGU); M.
Sandoval et al. 103, 312 (B, F, LAGU); W. Berendsohn et al. 1335 (B, LAGU, MO).
Sonsonate: M.B. Siu s.n. (ITIC); M.L. Reyna de Aguilar 586 (MHES); R. Seiler 631
(F, MHES). Santa Ana: M.C. Carlson 841 (F); M.L. Reyna de Aguilar & R. Villacorta
1440 (B, LAGU, MO); R. Seiler 228 (F, MHES). La Libertad: K. Sidwell et al. 468
(B, BM, ITIC, LAGU, MO); N. Ventura 5014 (ITIC); R. Guerra et al. s.n. (JBL04503)
(LAGU); R. Menjivar & J. Sermeño s.n. (JBL04494) (LAGU). San Salvador: K.
Sidwell et al. 563 (B, BM, ITIC, LAGU, MO); M.C. Carlson 3 (F); N.C. Fassett 28599
(ITIC); R. Seiler 57 (F, MHES); S. Winkler 30 (ITIC); W. Eggler 680 (ITIC); W. Heed
33 (ITIC). La Paz: J.C. González 102 (B, LAGU, MO); R. Villacorta et al. 2854 (B,
LAGU). San Vicente: P. Standley et al. 3689 (F). Usulután: A. del Cid 1941 (ITIC);
D. Williams 14, 297 (LAGU). San Miguel: A. Salgado 75 (MHES); J. Tucker 977 (F).
Morazán: R. Seiler 1047 (F).
Pityrogramma dealbata (C. Presl) Domin, Rozpr. Ceské Akad. Ved., Tr. 2, Vedy Mat.
Prir. 51(15): 7. 1942.
Gymnogramma dealbata C. Presl, Ceropteris schaffneri Fée, Pityrogramma schaffneri
(Fée) Weath.
Salvadoran distribution: Ahuachapán: N.C. Fassett 28718b (F); S. de Moore s.n. (12916) (MHES). Sonsonate: J. Tucker 1340 (F). Santa Ana: A. Molina et al. 16844 (F);
M.B. Siu s.n. (ITIC). La Libertad: N. Ventura 5019 (ITIC); V. Hellebyuck s.n. (12-92)
(F, MHES). San Salvador: A. Molina et al. 21829 (F, ITIC); M.C. Carlson 464 (F); R.
Seiler 66 (F, MHES). Chalatenango: L. Lara 208, 219 (MHES). San Miguel: A.
Salgado s.n. (JBL04801) (LAGU). Morazán: A. Salgado 62 (MHES); J. Monterrosa et
al. 43 (B, BM, LAGU, MO); 671 (BM, LAGU).
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FERN GAZ. 18(4): 120-215. 2008
Pityrogramma ebenea (L.) Proctor, Brit. Fern Gaz. 9: 219. 1965.
Acrostichum ebeneum L., A. tartareum Cav., Pityrogramma tartarea (Cav.) Maxon, P.
tripinnata Domin
Salvadoran distribution: Sonsonate: R. Seiler 630 (F, MHES). Santa Ana: J.
Monterrosa et al.1029 (LAGU, NY); M.L. Reyna de Aguilar 669 (MHES); R. Seiler 40
(F, MHES); 462 (F, ITIC, MHES). San Vicente: S. Winkler s.n. (ITIC).
Pityrogramma ferruginea (Kunze) Maxon, Contr. U.S. Natl. Herb. 17: 173. 1913.
Gymnogramma ferruginea Kunze, G. bommeri Christ
Global distribution: Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Peru.
Salvadoran distribution: Sonsonate: W. Eggler 606 (ITIC). Santa Ana: R. Seiler 499
(F, ITIC, MHES). San Salvador: A.K. Monro et al. 2222 (B, BM, ITIC, LAGU, MO);
M.C. Carlson 448 (F). Morazán: J. Monterrosa et al. 11 (LAGU, MO).
♦ Pityrogramma ochracea (C. Presl) Domin, Spisy Prír. Fak. Karlovy Univ. 88: 8.
1928.
Gymnogramma ochraceae C. Presl, Acrostichum caudatum Cav., Pityrogramma
calomelanos (L.) Link var. ochracea (C. Presl) R. M. Tryon
Global distribution: Honduras, Colombia, Venezuela, Ecuador, Peru, Bolivia.
Salvadoran distribution: Usulután: D. Williams 139 (B, LAGU, MO); 427 (LAGU).
Pityrogramma trifoliata (L.) R. M. Tryon, Contr. Gray Herb. 189: 68. 1962.
Acrostichum trifoliatum L., Trismeria trifoliata (L.) Diels in Engl. & Prantl
Local names: helecho vertical.
Panama, Antilles, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, Chile,
Paraguay, Argentina, Uruguay.
K. Prestegard 49 (MP-00007) (LAGU). Santa Ana: M.C. Carlson 840 (F); P. Standley
et al. 3254 (F); R. Seiler 1207 (F, MHES). La Libertad: L. Williams & A. Molina
15192 (JBL015499), LAGU (F). San Salvador: F. López s.n. (ITIC); R. Seiler 518 (F,
MHES). Chalatenango: F. Wunderlin 717 (ITIC); R. Seiler 264 (MHES); R. Villacorta
et al. 2086 (B, LAGU, MO).
Polytaenium cajenense (Desv.) Benedict, Bull. Torrey Bot. Club 38: 169. 1911.
Hemionitis cajenense Desv., Antrophyum cajenense (Desv.) Spreng., A. lacantunense
Rovirosa
Panama, Antilles; Colombia, Venezuela, Trinidad and Tobago, Guayana, Suriname,
Source: Moran in Davidse et al. (1995): 146 (as Antrophyum cajenense); Mickel &
Smith (2004): 526.
Pteris altissima Poiret in Lamarck, Encycl. 5: 722. 1804.
Pteris kunzeana J. Agardh, P. protea Liebm., Litobrochia kunzeana (J. Agardh) Fée, L.
grandis Fée
Local names: helecho cola de pavo real, helecho palma de cristo.
MO); E. Sandoval & R.Chinchilla 1167 (B, LAGU); F. Chinchilla s.n. (ISB00330)
(LAGU); R. Seiler 944 (MHES); W. Berendsohn et al. 1425 (B, LAGU, MO). Santa
Ana: S. Winkler 31 (ITIC). San Salvador: R. Seiler 536 (MHES).
Pteris biaurita L., Sp. Pl. 2: 1076. 1753.
Local names: helecho cola de ganzo, helecho palma real.
French Guyana, Ecuador, Peru, Brazil, Bolivia, Old World Tropics.
M. Sandoval & E. Sandoval s.n. (ISB00873) (AAU, B, F, LAGU); R. Seiler 921 (ITIC,
MHES); W. Berendsohn et al. 1399 (B, LAGU, MO); 1423 (LAGU); 1426 (B, LAGU).
Sonsonate: D. Rodríguez 157 (LAGU). La Libertad: A.K. Monro et al. 2314 (B, BM,
ITIC, LAGU, MO); J. Monterrosa et al. s.n. (JBL04489) (LAGU); K. Sidwell et al. 699
(B, BM, ITIC, LAGU, MO); N. Ventura 5009 (ITIC); O. Tejada et al. s.n. (JBL04511)
(LAGU). San Salvador: B. López s.n. (ITIC). Cabañas: A.K. Monro et al. 3751 (B,
BM, ITIC, LAGU, MO); G. Davidse et al. 37119 (B, BM, ITIC, LAGU, MO).
Pteris cretica L., Mant. Pl. 130. 1767.
Pteris triphylla M. Martens & Galeotti, P. trifoliata Fée
Local name: helecho pié de zenzontle.
Panama, Antilles, Peru, Brazil, Argentina, Old World Tropics and Subtropics.
Salvadoran distribution: La Libertad: E.A. Montalvo 6227 (B, LAGU). San
Salvador: R. Seiler 222 (MHES). Chalatenango: R. Seiler 377 (JBL00206) (LAGU,
MHES).
♦ Pteris erosa Mickel & Beitel, Mem. New York Bot. Gard. 46: 322-323. 1988.
Salvadoran distribution: Sonsonate: R. Seiler 622 (MHES). Santa Ana: D. Rodríguez
& G. Cerén s.n. (JBL04599) (LAGU); J. Monterrosa et al. 206 (B, LAGU, MO); R.
Seiler 246 (MHES); V.M. Martínez s.n. (CMC00844) (B, LAGU). San Salvador: R.
Seiler 212 (F, MHES). San Vicente: R. Seiler 316 (MHES).
Note: Moran in Davidse et al. (1995) considered this name a synonym of Pteris
orizabae M. Martens et Galeotti. Mickel & Smith (2004) consider P. orizabae and
Pteris erosa distinct based on differences in indusium margin and other characters.
Pteris grandifolia L., Sp. Pl. 2: 1073. 1753.
Litobrochia grandifolia (L.) J. Sm., Heterophlebium grandifolium (L.) Fée
Salvadoran distribution: P. Standley 21242, MO (Moran in Davidse et al. 1995). San
Vicente: R. Seiler 974 (ITIC, MHES).
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FERN GAZ. 18(4): 120-215. 2008
Pteris muricella Fée, Mém. Foug. 8: 73. 1857.
Pteris mollis Christ
Global distribution: Mexico, Guatemala, Nicaragua, Costa Rica, Panama, Venezuela.
Salvadoran distribution: Ahuachapán: R. Seiler 904 (F, ITIC, MHES).
Pteris orizabae M. Martens et Galeotti, Foug. Mexique 53, pl. 13. 1842.
Pteris apicaulis Liebm., P. hemipteris (Fée) Hook., Litobrochia orizabae (M. Martens
& Galeotti) Fée, L. setifera Fée, L. hemipteris Fée
Salvadoran distribution: T. Croat 42417, MO (Moran in Davidse et al. 1995). Santa
Ana: J. Monterrosa & R. Martínez 945 (B, LAGU, MO); R. Seiler 272, 692 (MHES).
Morazán: R. Seiler 1032 (ITIC, MHES).
Pteris paucinervata Fée, Mém. Foug. 8: 73. 1857.
Salvadoran distribution: Chalatenango: R. Seiler 408 (JBL00205) (F, LAGU, MHES).
Pteris podophylla Sw., J. Bot. (Schrader) 1800(2): 67. 1801.
Lonchitis pedata L.
Global distribution: Mexico, Guatemala, Nicaragua, Costa Rica, Panama, Antilles,
Colombia, Venezuela, Suriname, Ecuador, Peru, Brazil, Bolivia.
Pteris quadriaurita Retz., Observ. Bot. 6: 38. 1791.
Pteris plumula Desv., P. nemoralis Willd. var. major M. Martens & Galeotti, P. galeotti
(Fée) Hook., Litobrochia galeotti Fée
French Guyana, Ecuador, Peru, Bolivia, Argentina, Africa, Asia.
MO); R. Seiler 813, 908, 951 (MHES); W. Berendsohn et al. 1347 (B, LAGU, MO).
Sonsonate: R. Seiler 621 (MHES). Santa Ana: K. Prestegard 2 (MP-00035) (LAGU);
M.B. Siu s.n. (ITIC); R. Seiler 1205 (MHES); S. Winkler 36 (ITIC); T. Croat 42306
(ITIC). La Libertad: W. Berendsohn et al. 1048 (B, LAGU). San Salvador: R. Seiler
217 (MHES). Chalatenango: J.C. González & M.A. Hernández 268 (B, LAGU, MO).
Cabañas: A.K. Monro et al. 3743 (B, BM, ITIC, LAGU, MO); R.A. Carballo et al. 253
(LAGU, MO).
Pteris vittata L., Sp. Pl. 2: 1074. 1753.
Local name: helecho de banda.
Global distribution: Mexico, Honduras, Panama, Antilles, Venezuela, Peru, Brazil,
Argentina; Viejo Mundo.
Salvadoran distribution: Ahuachapán: K. Prestegard 6 (MP-00036) (LAGU); R. Seiler
811 (F, ITIC, MHES). La Libertad: E.A. Montalvo 6340 (B, LAGU, MO).
Scoliosorus ensiformis (Hook.) T. Moore, Index Fil. xxix, 1857.
Antrophyum ensiforme Hook. in Bentham, A. falcatum M. Martens & Galeotti, A.
carnosum Liebm., A. galeotti Fée, Polytaenium ensiforme (Hook.) Benedict
Panama.
Salvadoran distribution: Ahuachapán: M. Renderos et al. 430 (LAGU); R. Seiler 915
(ITIC, MHES); 966 (MHES). Sonsonate: E.A. Montalvo s.n. (ITIC); R. Seiler 614
(MHES); T. Croat 42216 (ITIC). Santa Ana: B. Pfeiffer-Berendsohn & W. Berendsohn
30 (B, LAGU, MO); D. Rodríguez et al. 37 (B, LAGU, NY); J. Lagos 1619 (ITIC); J.
Monterrosa et al. 528 (B, LAGU, MO); 938 (B, LAGU, NY); K. Prestegard 25 (MP00009) (LAGU); R. Seiler 15, 131 (ITIC, MHES); 360 (MHES); S. Winkler 54 (ITIC);
W. Berendsohn et al. 1710 (B, BM, LAGU, MO). Chalatenango: R. Seiler 412
(JBL04946) (LAGU, MHES). Morazán: R. Seiler 1017 (ITIC, MHES).
♦Vittaria bradeorum Rosenst., Repert. Spec. Nov. Regni Veg. 22: 18. 1925.
Global distribution: Mexico, Guatemala, Costa Rica. Nicaragua y Panama.
Local names: helecho barba de cacique, helecho cola de gorrión.
W. Berendsohn et al. 1461 (LAGU).
Vittaria graminifolia Kaulf., Enum. Filic. 192. 1824.
Vittaria filifolia Fée, V. dimorpha Müll.
Panama, Antilles, Colombia, Venezuela, Trinidad and Tobago, Suriname, Ecuador,
Peru, Brazil, Bolivia.
Salvadoran distribution: Ahuachapán: R. Seiler 820, 960 (F, MHES). Sonsonate: A.
Molina et al. 21722 (F, ITIC); R. Seiler 605 (F, MHES). Santa Ana: A. del Cid 1621
(ITIC); E.A. Montalvo 3281 (ITIC); F. Wunderlin 559 (ITIC); G. Davidse et al. 37281
(B, BM, ITIC, LAGU, MO); J.L. Linares s.n. (JBL01322) (LAGU); J. Monterrosa et al.
950, 991 (LAGU, NY); M.B. Siu s.n. (ITIC); R. Seiler 17 (ITIC, MHES); 128 (F,
MHES); 99, 283, 356 (F, ITIC, MHES). San Salvador: M.C. Carlson 426 (F).
Chalatenango: R. Seiler 402 (JBL00208) (F, LAGU, MHES); W. Heed 39 (ITIC). San
Vicente: R. Seiler 335 (F, ITIC, MHES). Morazán: R. Seiler 991 (F, MHES).
SALVINIACEAE
Azolla filiculoides Lam., Encycl. I: 343. 1783.
Azolla caroliniana Willd.
Global distribution: USA; Mexico, Guatemala, Honduras, Nicaragua, Colombia,
Venezuela, Bolivia, Chile, Argentina, Uruguay, E Asia, Australia, Hawaii.
References: Maxon & Standley (1930): 177 (as Azolla caroliniana); Calderón &
Standley (1941): 35 (as A. caroliniana); Seiler (1982): 390; Berendsohn (1989): 23.
Azolla microphylla Kaulf., Enum. Filic. 273. 1824.
Azolla mexicana C. Presl
Global distribution: Canada; Mexico, Honduras, Nicaragua, Antilles, Colombia,
Venezuela, French Guyana, Ecuador, Peru, Brazil, Bolivia, Argentina, Uruguay.
Salvadoran distribution: Ahuachapán: R. Seiler 722 (MHES). La Libertad:
E.A.Montalvo 3472 (ITIC). San Vicente: N.C. Fassett 28344 (JBL01484) (ITIC,
LAGU). San Miguel: J. Monterrosa & R. Villacorta 1638 (BM, LAGU, MO).
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FERN GAZ. 18(4): 120-215. 2008
Salvinia auriculata Aubl., Hist. Pl. Guiane 2: 969, t. 367. 1775.
Salvinia rotundifolia Willd.
French Guyana, Ecuador, Peru, Brazil, Bolivia, Chile, Paraguay, Argentina, Uruguay.
Salvadoran distribution: La Unión: R. Villacorta et al. 2706 (B, CR, LAGU, MEXU);
N.C. Fassett 28631 (ITIC, MO) (Moran in Davidse et al. 1995); 28662 (ITIC); J.
Monterrosa & R. Villacorta 1637 (BM, LAGU, MO).
Salvinia minima Baker, J. Bot. 24: 98. 1886.
Salvinia rotundifolia auct. non Willd.
Rica, Panama, Antilles, Colombia, Venezuela, French Guyana, Ecuador, Peru, Brazil,
Bolivia, Paraguay, Argentina, Uruguay.
MO); J.M. Rosales 2272 (B, LAGU, MO); N. Herrera s.n. (JBL03862) (LAGU, MO);
R. Seiler 900 (MHES). Santa Ana: C. Peñate 4 (ITIC). San Salvador: K. Sidwell et al.
534 (B, BM, ITIC, LAGU, MO). La Paz: N.C. Fassett 28515 (ITIC). Usulután: N.C.
Fassett 28683 (ITIC). San Miguel: N.C. Fassett 28556 (ITIC); R. Seiler 723 (MHES).
La Unión: K. Sidwell et al. 689 (B, BM, ITIC, LAGU, MO).
TECTARIACEAE
Tectaria heracleifolia (Willd.) Underw., Bull. Torrey Bot. Club 33: 200. 1906.
Aspidium heracleifolium Willd., Polypodium cordifolium M. Martens & Galeotti,
Tectaria heracleifolia (Willd.) Underw. var. maxima C.V. Morton
Local names: helecho cisne, helecho cola de perro, helecho corazón, helecho palma de
costa.
732 (B, LAGU, MO); J.M. Rosales 231, 745 (AAU, B, BM, LAGU, MO); 946
(LAGU); R. Seiler 941 (JBL04819) (F, LAGU, MHES); R. Villacorta & Martínez 409
(LAGU, MO); W. Berendsohn & R. Villacorta 1110 (B, LAGU); W. Berendsohn et al.
1319, 1324, 1361, 1605 (B, LAGU, MO). Sonsonate: D. Rodríguez & G. Trejo 200 (B,
LAGU, MO). Santa Ana: K. Prestegard 35 (MP-00024) (LAGU). La Libertad: L.O.
Williams et al. 16779 (F); R.A. Carballo et al. 38 (B, BM, LAGU, MO). San Vicente:
P. Standley et al. 3532 (F). San Miguel: A. Salgado s.n. (JBL04802) (LAGU); A.
Salgado 77 (F, MHES). Cabañas: A.K. Monro et al. 3745 (B, BM, ITIC, LAGU, MO).
Tectaria incisa Cav. f. incisa, Descr. Pl. 249. 1801.
Aspidium macrophyllum Sw. var. pittieri Christ, A. martinicense Spreng., Tectaria
martinicensis (Spreng.) Copel.
Local name: helecho samaritana.
Salvadoran distribution: Ahuachapán: E. Sandoval & F. Chinchilla 212 (LAGU);
1352 (B, LAGU); R. Seiler 930 (F, MHES). Sonsonate: D. Rodríguez 161 (LAGU). La
Libertad: S. Winkler 70 (ITIC). San Salvador: S. Winkler 89 (ITIC). San Vicente: P.
Standley et al. 3771 (F).
Tectaria mexicana (Fée) Morton, Amer. Fern. J. 56. 133. 1966.
Sagenia mexicana Fée, Aspidium latifolium C. Presl, Tectaria latifolia (G. Forst.)
Copel., T. mexicana (Fée) C.V. Morton var. pilosula C.V. Morton., T. dilacerata auct.
non Kunze
Local names: helecho cola de pavo, helecho falsa salamandra, helecho pié de león.
Panama, Colombia, Ecuador.
MO); E. Sandoval & H. Rivera 1313 (B, LAGU); F. Chinchilla & E. Sandoval s.n.
(ISB00276) (B, LAGU, MO); s.n. (ISB00670) (AAU, B, LAGU); J.M. Rosales 1314
(LAGU); M. Sandoval & E. Sandoval s.n. (ISB00870) (AAU, B, F, LAGU); M.
Sandoval & E. Sandoval 93 (B, F, LAGU); P. Standley et al. 2607 (F); R. Seiler 816 (F,
MHES); 934 (MHES). Sonsonate: D. Rodríguez & M. Trejo 82 (LAGU); J. Tucker
1351 (F). La Libertad: A.K. Monro et al. 2306, 3090 (B, BM, ITIC, LAGU, MO); J.
Monterrosa et al. s.n. (JBL04486) (B, BM, ITIC, LAGU, MO); M.C. Carlson 141 (F);
M. Magaña 38 (MP-00038), 39 (MP-00014) (LAGU); N. Ventura 5003 (ITIC);
O.González et al. s.n. (JBL04495) (B, BM, ITIC, LAGU, MO). San Salvador: A.K.
Monro et al. 3438 (B, BM, ITIC, LAGU, MO); A. Molina et al. 21848 (F, ITIC); K.
Sidwell et al. 573 (B, BM, ITIC, LAGU, MO); R. Seiler 197 (F, MHES).
Chalatenango: L. González 1721 (ITIC). San Vicente: P. Standley et al. 3772 (F).
Usulután: D. Williams 132 (LAGU, MO). San Miguel: A.K. Monro et al. 2947 (B,
BM, ITIC, LAGU, MO); A. Salgado 74 (MHES); M. Renderos et al. 685 (AAU, B,
LAGU, MO); R. Villacorta et al. 2824 (B, BM, LAGU, MEXU, MO). Cuscatlán: S.
Dar s.n. (JBL03664) (B, LAGU, MO).
Tectaria rivalis (Mett. ex Kuhn) C. Chr., Index Filic., Suppl. 3, 184. 1934.
Tectaria rheosora auct. non. (Baker) C. Chr.
Global distribution: Belize, Guatemala, Nicaragua, Costa Rica, Panama, Colombia,
Ecuador.
Salvadoran distribution: Ahuachapán: R. Seiler 943 (F, MHES).
Tectaria transiens (C. V. Morton) A. R. Sm., Amer. Fern. J. 70: 27. 1980.
Tectaria incisa Cav. subsp. transiens C.V. Morton, T. ramkissoonii Jermy & T. G.
Walker
Antilles, Colombia, Trinidad and Tobago, Ecuador, Peru.
Source: R.C. Moran in Davidse et al. 1995: 209. Mickel & Smith 2004: 614.
THELYPTERIDACEAE
Macrothelypteris torresiana (Gaudich.) Ching, Acta Phytotax. Sin. 8: 310. 1963.
Polystichum torresianum Gaudich., Thelypteris torresiana (Gaudich.) Alston, T.
uliginosa (Kunze) Ching, Aspidium uliginosum Kunze, Dryopteris uliginosa (Kunze)
C. Chr.
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FERN GAZ. 18(4): 120-215. 2008
Local names: helecho amarillo, helecho palma de pozo, helecho pata de paloma.
Brazil, Bolivia, Paraguay, Argentina, Africa, Asia, Pacific Islands.
MO); E. Sandoval & H. Rivera 1276 (B, LAGU); E. Sandoval & O. Rivera s.n.
(ISB00814) (AAU, LAGU); F. Chinchilla & E. Sandoval s.n. (ISB00270) (LAGU);
J.M. Rosales 1052 (B, LAGU); M. Sandoval & E. Sandoval s.n. (ISB00843) (AAU, B,
LAGU); s.n. (JBL01367), 305 (B, LAGU); 208 (B, F, LAGU); S. Castillo s.n.
(ISF00688) (LAGU); S. de Moore s.n. (12-933) (MHES). Santa Ana: G. Davidse et al.
37227 (B, BM, ITIC, LAGU, MO); K. Prestegard 20 (MP-00049) (LAGU); R. Seiler
470 (F, ITIC, MHES); 839 (MHES). La Libertad: A.K. Monro et al. 2309, 3733, 3775
(B, BM, ITIC, LAGU, MO); W. Berendsohn et al. 1047 (B, LAGU); 1444 (LAGU).
San Salvador: M. Renderos 707 (B, LAGU, MO); R. Seiler 369 (F, MHES); 1504 (F).
Cabañas: A.K. Monro et al. 3753 (B, BM, ITIC, LAGU, MO).
Thelypteris subg. Amauropelta
Thelypteris balbisii (Spreng.) Ching, Bull. Fan Mem. Inst. Biol. 10: 250. 1941.
Polypodium balbisii Spreng., Aspidium sprengelii Kaulf., Dryopteris sprengelii
(Kaulf.) Kuntze, D. balbisii (Spreng.) Urb., Thelypteris sprengelii (Kaulf.) Proctor
Local name: helecho palma de río.
Panama, Antilles, Colombia, Venezuela, Trinidad and Tobago, Guayana, Ecuador, Peru,
Bolivia.
MO); E. Sandoval & F. Chinchilla 918 (B, LAGU); M. Sandoval & E. Sandoval 303
(B, F, LAGU); P. Standley et al. 2499 (F); R. Seiler 918, 935 (F, MHES); S. de Moore
s.n. (12-932) (MHES); W. Berendsohn et al. 1346 (B, LAGU, MO); 1722 (B, LAGU).
Sonsonate: J. Tucker 1331 (F). Santa Ana: G. Cerén & D. Rodríguez s.n. (JBL04658)
(B, LAGU, MO); R. Seiler 1203 (F, MHES). La Libertad: E.A. Montalvo & R.
Villacorta 6402 (B, ITIC, LAGU, MO); L. Williams & A. Molina 15203 (JBL01550)
(LAGU). San Salvador: A. Molina et al. 21841 (F, ITIC); R. Seiler 221, 712 (F,
MHES). Chalatenango: L. Lara 235, 236 (MHES). Cuscatlán: E.A. Montalvo 3313
Thelypteris cheilanthoides (Kunze) Proctor var. cheilanthoides, Bull. Inst. Jamaica,
Sci. Ser. 5: 58. 1953.
Phegopteris amplificata Fée, Dryopteris amplificata (Fée) C. Chr., Aspidium kunzei Fée
Salvadoran distribution: Santa Ana: V.M. Martínez s.n. (CMC00980) (B, EAP, LAGU,
MO).
Thelypteris concinna (Willd.) Ching, Bull. Fan Mem. Inst. Biol. 10: 251. 1941.
Polypodium concinnum Willd., Dryopteris concinna (Willd.) Kuntze, Amauropelta
concinna (Willd.) Pic. Serm., Phegopteris elongata E. Fourn., Thelypteris concinna
(Willd.) Ching var. elongata (E. Fourn.) C. F. Reed
Salvadoran distribution: S. Calderón 1994, LAGU (photo), NY. Chalatenango: R.
Seiler 256 (F, MHES). Morazán: J. Tucker 701 (F).
Thelypteris deflexa (C. Presl) R. M. Tryon, Rhodora 69: 5. 1967.
Nephrodium defelxum C. Presl, Dryopteris lindigii C. Chr., Thelypteris lindigii (C. Chr.)
Alston, Amauropelta deflexa (C. Presl) Å. Löve & D. Löve
Salvadoran distribution: Morazán: R. Seiler 1010 (F, LAGU (photo), MHES, NY,
UC).
♦Thelypteris linkiana (C. Presl) R.M. Tryon, Rhodora 69: 6. 1967.
Gymnogramma diplazioides Desv., G. polypodioides Link, G. cheilosora Fée,
Grammitis linkiana C. Presl, Dryopteris linkiana (C. Presl) Maxon, D. diplazioides
(Desv.) Proctor
Antilles, Colombia, Venezuela, Guayana, Ecuador, Peru, Bolivia.
Salvadoran distribution: Santa Ana: J.Monterrosa, D.Rodríguez & R.González 1528
(BM, LAGU, MHES, UC).
Thelypteris melanochlaena (C. Chr.) C. F. Reed, Phytologia 17: 292. 1968.
Dryopteris melanochlaena C. Chr.
Salvadoran distribution: Ahuachapán: R. Seiler 826 (F, MHES). Santa Ana: G.
Davidse et al. 37234 (B, BM, ITIC, LAGU, MO).
Thelypteris oaxacana A. R. Sm., Amer. Fern J. 63: 125 (F). 7, 8. 1973.
Global distribution: Mexico, Costa Rica, Nicaragua, Panama.
Salvadoran distribution: Chalatenango: Tucker 1043 (F, MICH) (A.R. Smith in
Davidse et al. 1995).
Thelypteris oligocarpa (Humb. & Bonpl. ex Willd.) Ching, Bull. Fan Mem. Inst. Biol.
10: 253. 1941.
Polypodium oligocarpum Humb. & Bonpl. ex Willd., Dryopteris oligocarpa (Humb. &
Bonpl. ex Willd.) Kuntze, D. columbiana C. Chr., Amauropelta oligocarpa (Humb. &
Bonpl. ex Willd.) Pic. Serm., Thelypteris columbiana (C. Chr.) C. V. Morton,
Phegopteris pilosula Fée
Local names: helecho trensillo.
Salvadoran distribution: Ahuachapán: E. Sandoval & F. Pérez 1242 (B, LAGU).
Sonsonate: R. Seiler 600 (MHES). Santa Ana: J. Monterrosa et al. 935, 969 (B,
LAGU, NY); R. Seiler 117, 279, 281, 347 (F, MHES); 671 (MHES); 873 (F, ITIC,
MHES). San Salvador: A.K. Monro et al. 3455 (B, BM, ITIC, LAGU, MO); M.C.
Carlson 4b (F); R. Seiler 33, 362 (F, MHES). Chalatenango: R. Seiler 421 (F, MHES).
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FERN GAZ. 18(4): 120-215. 2008
Thelypteris pilosohispida (Hook.) Alston, J. Wash. Acad. Sci. 48: 233. 1958.
Nephrodium pilosohispidum Hook., Dryopteris pilosohispida (Hook.) C. Chr.,
Alsophila pilosa M. Martens & Galeotti
Global distribution: Mexico, Guatemala, Honduras, Nicaragua, Antilles, Colombia,
Salvadoran distribution: Santa Ana: A. Molina et al. 16745 (F); J. Monterrosa et al.
207 (B, LAGU); R. Seiler 248 (F, MHES); S. Winkler 66 (ITIC). Morazán: R. Seiler
1008 (F, MHES).
Thelypteris pilosula (Klotzsch & H. Karst. ex Mett.) R. M. Tryon, Rhodora 69: 7. 1967.
Aspidium pilosulum Mett., Dryopteris pilosula (Klotzsch & H. Karst. ex Mett.) Hieron.,
Amauropelta pilosula (Mett.) Å. Löve & D. Löve
MO); R. Seiler 1193 (F, ITIC, MHES). Chalatenango: R. Seiler 421 (JBL00207)
(LAGU); R. Seiler 728 (MHES).
Thelypteris resinifera (Desv.) Proctor var. resinifera, Bull. Inst. Jamaica, Sci. Ser. 5:
63. 1953.
Nephrodium panamense C. Presl, Dryopteris panamensis (C. Presl) C. Chr., D.
panamensis (C. Presl) C. Chr. var. gonzalezii Christ ex C. Chr., Thelypteris panamensis
(C. Presl) E. P. St. John, Polypodium litiginosum Liebm., Aspidium exsudans E. Fourn.,
A. exsudans E. Fourn. var. majus E. Fourn., A. exsudans E. Fourn. var. feei E. Fourn.,
A. ghiesbreghtii E. Fourn.
Local name: helecho palma de río.
Rica, Panama, Antilles, Colombia, Ecuador.
ITIC, LAGU, MO); E. Sandoval & E. Fuentes 1302 (B, LAGU); F. Chinchilla & E.
Sandoval s.n. (ISB00152) (B, LAGU); J.M. Rosales 431 (B, BM, LAGU, MO); 943 (B,
LAGU, MO); P. Standley et al. 2969, 2982 (F); S. Castillo s.n. (ISF00593) (B, LAGU,
MO). Santa Ana: B. Pfeiffer-Berendsohn 64 (LAGU, MO); E.A. Montalvo 3367
(ITIC). La Libertad: A.K. Monro et al. 2291 (B, BM, ITIC, LAGU, MO); E.A.
Montalvo 3417 (ITIC). San Salvador: A.K. Monro et al. 2239 (B, BM, ITIC, LAGU,
MO); E.A. Montalvo 3585 (ITIC); R. Seiler 784 (F, ITIC, MHES). Chalatenango: A.
Molina et al. 21556 (F, ITIC); R. Seiler 261 (F, MHES); R. Villacorta et al. 982 (B,
LAGU, MO). San Vicente: P. Standley et al. 3409, 3697 (F). Morazán: A.K. Monro et
al. 3803 (B, BM, ITIC, LAGU, MO); J. Tucker 645, 700 (F); K. Sidwell et al. 750 (B,
BM, ITIC, LAGU, MO); R. Seiler 1049a (F, ITIC, MHES).
Thelypteris rudis (Kunze) Proctor, Bull. Inst. Jamaica, Sci. Ser. 5: 64. 1953.
Polypodium rude Kunze, Dryopteris rudis (Kunze) C. Chr., Amauropelta rudis (Kunze)
Pic. Serm., Phegopteris impressa Fée
Antilles, Colombia, Venezuela, Guayana, Ecuador, Peru, Bolivia.
Salvadoran distribution: Santa Ana: A. Molina et al. 16745 (LAGU (photo), NY); R.
Seiler 194 (F).
Thelypteris scalaris (Christ) Alston, J. Wash. Acad. Sci. 48: 234. 1958.
Aspidium scalare Christ, Dryopteris scalaris (Christ) C. Chr., Amauropelta scalaris
(Christ) Å. Löve & D. Löve
MO). Santa Ana: R. Seiler 193 (F, ITIC, LAGU (photo), MHES, NY). La Libertad:
K. Sidwell et al. 708 (B, BM, ITIC, LAGU, MO). Usulután: A.K. Monro et al. 3021
(B, BM, ITIC, LAGU, MO); D. Williams 59 (LAGU, MO). Morazán: R. Seiler 985
(MHES, UC).
Thelypteris thomsonii (Jenman) Proctor, Bull. Inst. Jamaica, Sci. Ser. 5: 65. 1953.
Polypodium thomsonii Jenman, Dryopteris thomsonii (Jenman) C. Chr., D. stuebelii
Hieron., Amauropelta thomsonii (Jenman) Pic. Serm.
Global distribution: Mexico, Guatemala, Costa Rica, Panama, Antilles, Ecuador, Peru.
Source: Stolze (1981): 487; Berendsohn (1989) 1: 26; Mickel & Smith (2004): 637.
Thelypteris subg. Cyclosorus
Thelypteris cretacea A. R. Sm., Univ. Calif. Publ. Bot. 59: 92. 1971.
Christella cretacea (A. R. Sm.) Å. Löve & D. Löve
ITIC, LAGU, MO); J.M. Rosales 945 (B, LAGU, MO); R. Seiler 919 (F, MHES).
Santa Ana: M.B. Siu s.n. (ITIC). La Libertad: R. Seiler 715 (F, MHES). San
Salvador: R. Seiler 254 (F, MHES). Cabañas: R. Seiler 523 (F, MHES).
Thelypteris dentata (Forssk.) E. P. St. John, Amer. Fern J. 26: 44. 1936.
Polypodium dentatum Forssk., Dryopteris dentata (Forssk.) C. Chr., D. mollis (Sw.)
Hieron., Cyclosorus dentatus (Forssk.) Ching, Christella dentata (Forssk.) Brownsey &
Jermy, Aspidium molle Sw.
Local name: palmarín.
Brazil, Paraguay, Argentina, Africa, Asia, Pacific Islands.
Salvadoran distribution: La Libertad: K. Prestegard 44 (MP-00046) (LAGU). San
Salvador: R. Seiler 219 (MHES); 710 (F, MHES). Chalatenango: R. Seiler 259 (F,
MHES). La Unión: R. Seiler 1048 (F, MHES).
Thelypteris hispidula (Decne.) C. F. Reed, Phytologia 17: 283. 1968.
Aspidium hispidulum Decne., Christella hispidula (Decne.) Holttum, Nephrodium
quadrangulare Fée, Dryopteris quadrangularis (Fée) Alston, Thelypteris
quadrangularis (Fée) Schelpe
Local names: helecho de río, helecho palma de río, helecho trensillo, palma de
barranco.
Suriname, French Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina,
209
FERN GAZ. 18(4): 120-215. 2008
Uruguay, Africa, Asia.
Salvadoran distribution: Ahuachapán: E.A. Montalvo 5061 (ITIC); E. Sandoval 206
(LAGU); 1147 (B, LAGU, MO); F. Chinchilla & E. Sandoval s.n. (ISB00426) (LAGU);
J.M. Rosales 43 (AAU, B, BM, LAGU, MO, MEXU); 987 (B, BM, LAGU, MO); 340,
2667 (B, LAGU, MO); 688, 2177 (AAU, B, BM, LAGU, MO); M. Sandoval & E.
Sandoval 251 (B, F, LAGU); R. Seiler 942 (F, MHES); S. Castillo s.n. (ISF00582) (B,
F, LAGU, MO). Santa Ana: K. Prestegard 4 (MP-00047) (LAGU); M. Magaña 3
(LAGU); O. Pank s.n. (12-1000) (MHES). La Libertad: A.K. Monro et al. 3092 (B,
BM, ITIC, LAGU, MO); N. Ventura 5022, 5023 (ITIC); W. Berendsohn et al. 1443
(LAGU). San Salvador: R. Seiler 50, 220, 453 (F, MHES). Usulután: D. Williams &
R.W. Herrera 237 (LAGU). San Miguel: A.K. Monro et al. 2945 (B, BM, ITIC, LAGU,
MO). Cabañas: G. Davidse et al. 37115 (B, BM, ITIC, LAGU, MO); R. Seiler 227, 229
(MHES).
Thelypteris interrupta (Willd.) K. Iwats., Jap. J. Bot. 38: 314. 1963.
Pteris interrupta Willd., Cyclosorus interruptus (Willd.) H. Itô, C. tottum (Thunb.) Pic.
Ser., C. gongylodes (Schkuhr) Link, Dryopteris gongylodes (Schkuhr) Kuntze,
Polypodium tottum Thunb., Thelypteris totta (Thunb.) Schelpe, T. gongylodes
(Schkuhr) Small, Aspidium gongylodes Schkuhr, Nephrodium paludosum Liebm.
Brazil, Bolivia, Paraguay, Argentina, Uruguay, Africa, Asia.
MO); J.M. Rosales 2447 (B, LAGU, MO); N.C. Fassett 28523 (ITIC); N. Herrera s.n.
(JBL03767) (B, LAGU, MO); R. Seiler 950 (F, MHES).
Thelypteris kunthii (Desv.) C.V. Morton, Contr. U.S. Natl. Herb. 38: 53. 1967.
Nephrodium kunthii Desv., Dryopteris normalis C. Chr., Thelypteris normalis (C. Chr.)
Moxley, Christella normalis (C. Chr.) Holttum.
Rica, Antilles, Colombia, Venezuela, Brazil.
Salvadoran distribution: San Miguel: R. Villacorta et al. 2703 (B, BM, LAGU, MO);
2890 (B, BM, EAP, LAGU, MO).
Thelypteris patens (Sw.) Small var. patens, Ferns S.E. States 243. 1938.
Polypodium patens Sw., Dryopteris patens (Sw.) Kuntze, Christella patens (Sw.)
Holttum
Local name: helecho palma de pozo.
Rica, Panama, Antilles, Colombia, Venezuela, Trinidad and Tobago, Peru, Brazil,
Bolivia, Paraguay.
Salvadoran distribution: Ahuachapán: E. Sandoval & F. Chinchilla 81 (B, LAGU); R.
Seiler 936 (F, ITIC, MHES). Sonsonate: D. Rodríguez 160 (LAGU); J. Tucker 1352
(F); M.B. Siu s.n. (ITIC). Santa Ana: E.A. Montalvo 3246 (ITIC); S. Winkler 87
(ITIC). La Libertad: A.K. Monro et al. 3731, 3734 (B, BM, ITIC, LAGU, MO); R.
Villacorta 1068 (B, LAGU, MO). San Salvador: A. Molina et al. 21846 (F); R. Seiler
340, 519 (F, MHES). San Vicente: P. Standley et al. 3675 (F). Usulután: D. Williams
261 (LAGU). Cabañas: A.K. Monro et al. 3744, 3756 (B, BM, ITIC, LAGU, MO); R.
Thelypteris patens (Sw.) Small var. smithiana Ponce, Darwiniana 28: 373. 1987.
Global distribution: Nicaragua, Costa Rica, Panama, Antilles, Colombia, Venezuela,
Ecuador, Peru, Bolivia, Brazil.
Salvadoran distribution: Santa Ana: R. Seiler 762 (F, MHES). San Miguel: A. Salgado
73 (F, MHES). Cabañas: G. Davidse et al. 37118 (B, BM, ITIC, LAGU, MO).
Morazán: Tucker 690 (F, UC) (A.R. Smith in Davidse et al. 1995).
Thelypteris puberula (Baker) C.V. Morton var. puberula, Amer. Fern J. 48: 138. 1958.
Aspidium puberulum Fée, Nephrodium puberulum Baker in Hooker & Baker,
Dryopteris puberula (Baker) Kuntze, D. feei C. Chr., D. augescens (Link) C. Chr. var.
puberula (Baker) C. Chr., Thelypteris feei (C. Chr.) Moxley, Christella puberula
(Baker) Å. Löve & D. Löve
Local name: helecho peloso.
Global distribution: Mexico, Guatemala, Honduras, Nicaragua, Costa Rica, Antilles.
Salvadoran distribution: Santa Ana: J. Lagos 1622 (ITIC); R. Seiler 676 (F); V.M.
Martínez s.n. (CMC00854) (B, LAGU); R. Seiler 876 (MHES). San Salvador: A.
Molina & E.A. Montalvo 21653 (EAP, LAGU (photo), NY); R. Seiler 51 (F, MHES).
La Libertad: J. Williams s.n. (ITIC); L.O. Williams et al. 15092 (F); R. Seiler 1520
(F).
Thelypteris subg. Goniopteris
Thelypteris hatchii A. R. Sm., Amer. Fern J. 63: 118 (F). 1-3. 1973.
Goniopteris hatchii (A. R. Sm.) Å. Löve & D. Löve
Thelypteris minor (C. Chr.) A. R. Sm., Phytologia 34: 232. 1976.
Dryopteris nicaraguensis (E. Fourn.) C. Chr. var. minor C. Chr.
Local name: helecho nido de guachota.
MO); D. Witsberger 653, 690, 691 (MHES); E. Sandoval 42 (LAGU, MO); 1136 (B,
LAGU); G. Davidse et al. 37373, 37376 (B, BM, ITIC, LAGU, MO); N. Herrera s.n.
(JBL03861) (LAGU, MO); R. Seiler 815 (F, ITIC, MHES). Sonsonate: A. Molina et al.
21624 (ITIC). Santa Ana: M.B. Siu s.n. (ITIC); S. Winkler 88 (ITIC). Morazán: R.
Seiler 905 (F, ITIC, MHES).
♦Thelypteris obliterata (Sw.) Proctor, Bull. Inst. Jamaica, Sci. Ser. 5: 62. 1953.
Polypodium obliteratum Sw., P. faucium Liebm., Goniopteris obliterata (Sw.) C. Presl.,
Dryopteris obliterata (Sw.) C. Chr.
Antilles.
Salvadoran distribution: Morazán: J.Monterrosa & C.Bermudez 1383 (BM, LAGU,
MO, NY, UC).
211
FERN GAZ. 18(4): 120-215. 2008
Thelypteris tetragona (Sw.) Small, Ferns S. E. States 256. 1938.
Polypodium tetragonum Sw., Goniopteris tetragona (Sw.) C. Presl, Dryopteris
tetragona (Sw.) Urb., D. subtetragona (Link) Maxon
Suriname, Ecuador, Peru, Brazil, Bolivia.
Salvadoran distribution: P. Standley 21647, GH (A.R. Smith in Davidse et al. 1995).
Ahuachapán: R. Seiler 953 (MHES). Santa Ana: K. Prestegard 11 (MP-00050)
(LAGU).
Thelypteris subg. Meniscium
♦Thelypteris angustifolia (Willd.) Proctor, Bull. Inst. Jamaica, Sci. Ser. 5: 57. 1953.
Meniscium angustifolium Willd., Dryopteris angustifolia (Willd.) Urb., Polypodium
salicifolium Vahl
Antilles, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, Paraguay.
Salvadoran distribution: Morazán: J.Monterrosa & C.Bermudez 1381 (BM, LAGU,
UC).
♦ Thelypteris serrata (Cav.) Alston, Kew Bull. 1932: 309. 1932.
Meniscium serratum Cav., Dryopteris serrata (Cav.) C. Chr.
Local name: helecho gallardete.
Salvadoran distribution: Ahuachapán: F. Chinchilla & E. Sandoval s.n. (ISB00277)
(LAGU, MO). Sonsonate: S. Winkler 34 (ITIC).
♦ Thelypteris standleyi (Maxon & C. V. Morton) R. M. Tryon, Rhodora 69: 8. 1967.
Dryopteris standleyi Maxon & C. V. Morton
Colombia, Ecuador, Peru, Bolivia.
Salvadoran distribution: Morazán: A.K. Monro et al. 3828 (B, BM, ITIC, LAGU,
MO); J. Monterrosa & R. Carballo 795, 871 (LAGU).
WOODSIACEAE
Athyrium bourgeaui E. Fourn., Mexic. Pl. I: 102.1872.
Athyrium paucifrons C. Chr.
Salvadoran distribution: Morazán: R. Seiler 1056 (ITIC, MHES).
Athyrium palmense (Christ) Lellinger, Proc. Biol. Soc. Wash. 89: 708. 1977.
Cystopteris fragilis (L.) Bernh. var. palmensis Christ
Global distribution: Mexico, Costa Rica.
Salvadoran distribution: Santa Ana: M.L. Reyna de Aguilar 760 (MHES); R. Seiler
620 (MHES); 1202 (ITIC). La Libertad: W. Berendsohn et al. 1442 (LAGU). San
Salvador: R. Seiler 450, 580 (MHES).
Athyrium skinneri (Baker) Diels, Nat. Pflanzenfam. I(4): 224.1899.
Asplenium skinneri Baker
Global distribution: Mexico, Guatemala, Nicaragua, Costa Rica, Panama, Colombia,
Ecuador.
J.M. Rosales 944, 1053 (AAU, B, BM, LAGU, MO). Sonsonate: D. Rodríguez & G.
Trejo 107 (LAGU, NY). Santa Ana: K. Prestegard 9 (MP-00013) (LAGU); R. Seiler
701 (MHES). La Libertad: N. Ventura 5002 (ITIC); O. Rohweder 2625 (JBL01253)
(LAGU); 2626 (JBL01252) (LAGU). San Salvador: R. Seiler 583 (JBL04814) (F,
LAGU, MHES); R. Seiler 68, 375, 510, 635 (F, MHES).
Cystopteris fragilis (L.) Bernh., Neues J. Bot. I(2): 26, t. 2 (F). 9. 1806.
Polypodium fragile L., P. diaphanum Bory, Cystopteris fragilis (L.) Bernh. subsp.
diaphana (Bory) Litard., C. diaphana (Bory) Blasdell, C. arcuata Fée, Aspidium fragile
(L.) Sw. var. fumarioides M. Martens & Galeotti
Panama, Antilles, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, Chile,
Argentina, Europe, Africa, Asia, Hawaii.
MO); J. Monterrosa et al. 982 (B, LAGU, MO, NY); R. Seiler 353 (JBL04809) (F,
LAGU, MHES); R. Seiler 278, 343, 502 (F, MHES). Chalatenango: R. Seiler 155 (F,
MHES); R. Seiler 405 (JBL00090) (F, LAGU, MHES).
Diplazium cristatum (Desr.) Alston, J. Bot. 74: 173.1936.
Meniscium cristatum Desr., Asplenium arboreum Willd., A. denticulosum Desv., A.
shepherdii Spreng., Diplazium arboreum (Willd.) C. Presl, D. denticulosum (Desv.) C.
Chr.
Salvadoran distribution: Ahuachapán: J.M. Rosales 922, 947 (AAU, B, BM, LAGU,
MO); M. Magaña 40 (MP-00015) (LAGU); R. Seiler 916 (F, MHES). Santa Ana: M.
B. Siu s.n. (ITIC). San Salvador: R. Seiler 371 (F, ITIC, MHES); 532 (F, MHES).
Diplazium donnell-smithii Christ, Bull. Herb. Boissier, sér. 2, 7: 270. 1907.
MO); R. Seiler 825 (F, ITIC, MHES); S. de Moore s.n. (12-912) (MHES). Santa Ana:
R. Seiler 358 (JBL04810) (LAGU, MHES); 689 (JBL04816) (F, LAGU, MHES); 273
(MHES); 474 (F, MHES); V.M. Martínez s.n. (CMC00896) (B, EAP, LAGU, MO). San
Salvador: A. Molina & E.A. Montalvo 21852 (JBL01464), EAP (F, LAGU); R. Seiler
336 (F, MHES); S. Winkler 32 (ITIC).
♦ Diplazium drepanolobium A.R. Sm., Amer. Fern. J. 70: 21. 1980.
Salvadoran distribution: Sonsonate: R. Villacorta et al. 323 (B, F, LAGU, MO).
Diplazium franconis Liebm., Mexic. Bregn. 256 (reprint 104). 1849.
213
FERN GAZ. 18(4): 120-215. 2008
Asplenium franconis (Liebm.) Mett., A. franconis (Liebm.) Mett. var. gracile E. Fourn.,
A. aspidiiforme Fée, A. distans Fée, A. shepherdii Spreng. var. bipinnatum Christ,
Diplazium camptocarpon Fée .
Local names: helecho pacún, helecho crisalia.
Global distribution: Mexico, Guatemala, Belize, Honduras, Nicaragua, Panama.
E. Sandoval & F. Chinchilla 43 (LAGU). Santa Ana: A. Molina et al. 16896 (F); G.
Cerén & D. Rodríguez s.n. (JBL04662) (LAGU); G. Davidse et al. 37225 (B, BM, ITIC,
LAGU, MO); R. Seiler 178, F (LAGU). Chalatenango: R. Seiler 380 (F, MHES).
Diplazium lindbergii (Mett.) Christ, Prim. Fl. Costaric. 3(I): 27. 1901.
Asplenium lindbergii Mett., A. induratum Christ, Diplazium induratum Diels
Antilles, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, Uruguay.
Salvadoran distribution: Lötschert 414, US (Mickel & Smith, 2004: 260). Morazán: R.
Seiler 981 (MHES); 1028 (ITIC).
Diplazium lonchophyllum Kunze, Linnea 13: 141. 1839.
Diplazium inaequilaterum Liebm., D. actuale Fée .
Local names: helecho de coqueta, helecho palma de ángel, helecho pluma de tucán,
helecho trencilla de bajío.
Colombia, Ecuador.
Salvadoran distribution: Ahuachapán: E. Sandoval & F. Chinchilla 130 (LAGU); E.
Sandoval & F. Chinchilla 487, 1165 (B, LAGU); E. Sandoval & M. Sandoval 1372 (B,
LAGU); F. Chinchilla & E. Sandoval s.n. (ISB00151) (B, LAGU, MO); J.M. Rosales
451 (AAU, B, BM, LAGU, MO); M. Sandoval & E. Sandoval s.n. (ISB00869), 289,
294 (B, F, LAGU); P. Standley et al. 2567 (F); R. Seiler 917, 931 (F, MHES); W.
Berendsohn 1312 (B, LAGU, MO); W. Berendsohn et al. 1400 (B, LAGU, MO).
Sonsonate: J. Tucker 1350 (F); R. Seiler 596 (F, MHES). Santa Ana: K. Prestegard 5
(MP-00019) (LAGU); R. Seiler 244 (F, ITIC, MHES); V.M. Martínez s.n. (CMC00853)
(B, LAGU, MO); V.M. Martínez s.n. (CMC00873), s.n. (CMC00967), s.n.
(CMC01010), s.n. (CMC01032) (B, EAP, LAGU, MO). La Libertad: J. Mauricio et al.
s.n. (JBL04480), s.n. (JBL04490) (B, BM, ITIC, LAGU, MO); R. Seiler 714 (F,
MHES). San Salvador: R. Seiler 218, 370, 451, 452, 533, 786 (F, MHES).
Chalatenango: L. Lara 232, 236a (MHES). Cabañas: A.K. Monro et al. 3749 (B, BM,
ITIC, LAGU, MO). Cuscatlán: Grupo Ecológico Cantón La Bermuda s.n. (JBL03211)
(AAU, B, BM, LAGU, MO). Morazán: R. Seiler 1064, 1086 (F, MHES).
Diplazium striatum (L.) C. Presl, Suppl. Tent. Pterid. 114. 1836.
Asplenium striatum L., Diplazium crenulatum Liebm., D. elongatum Fée, D. matudae
C. D. Adams. Local names: helecho macho de altura, helecho negro de altura, helecho
palma de calvario.
Bolivia.
Salvadoran distribution: Ahuachapán: E. Sandoval & F. Chinchilla 211 (LAGU); E.
Sandoval & M. Sandoval s.n. (JBL01385) (B, LAGU); F. Chinchilla s.n. (ISB00572)
(AAU, F, LAGU), s.n. (ISB00662) (AAU, F, LAGU); R. Seiler 933 (F, MHES).
Sonsonate: J. Tucker 1353 (F). San Salvador: S. Winkler 44 (ITIC). Morazán: R.
Seiler 981 (F).
Diplazium ternatum Liebm., Mexic. Bregn. 252 (reprint 100). 1849.
Asplenium ternatum (Liebm.) Hook.
Salvadoran distribution: Santa Ana: R. Seiler 772 (ITIC); Morazán: J.L. Linares & A.
Pool 2292 (JBL01931) (LAGU).
Diplazium vera-pax (Donn. Sm.) Hieron., Hedwigia 59: 322. 1917.
Asplenium vera-pax Donn. Sm., A. cordovense Baker
Global distribution: Mexico, Guatemala, Belize, Nicaragua, Costa Rica, Panama.
Diplazium werckleanum Christ, Bull. Herb. Boissier, sér. 2, 4: 969. 1904.
Panama, Colombia.
Salvadoran distribution: Ahuachapán: R. Seiler 902 (F, ITIC, MHES); 952 (F,
MHES). Sonsonate: A. Molina & E.A. Montalvo 21625 (JBL01491) (F, EAP, LAGU),
21774 (JBL01555) (F, EAP, LAGU). Santa Ana: B. Pfeiffer-Berendsohn 50 (B,
LAGU, MO); D. Rodríguez & G. Cerén s.n. (JBL04596) (LAGU); J. Tucker 1214 (F);
J. Monterrosa & R. Martínez 939, 946 (LAGU, NY); K. Prestegard 31 (MP-00020)
(LAGU); K. Sidwell et al. 844 (B, BM, ITIC, LAGU, MO); O.Pank s.n. (12-979)
(MHES); R. Seiler 275, 483 (F, MHES); S. Winkler 1b (ITIC). San Salvador: R. Seiler
355 (F, MHES). Chalatenango: R. Seiler 379 (JBL00089) (F, LAGU, MHES).
Morazán: R. Seiler 1025 (F).
Woodsia mollis (Kaulf.) J. Sm., J. Bot. (Hooker) 4: 191. 1841.
Physematium molle Kaulf., Woodsia guatemalensis Hook., W. fragilis Liebm.
Global distribution: Mexico, Guatemala, Nicaragua.
Salvadoran distribution: Sonsonate: R. Seiler 496 (F, MHES). Santa Ana: M.B. Siu
s.n. (ITIC); R. Seiler 43, 661 (F, MHES); 501 (ITIC, MHES); S. Winkler 83 (ITIC).
215
FERN GAZ. 18(4). 2008
will be refereed
aligned.
aligned.
a legible form.
Supplements.
FERN GAZ. 18(5):216-229. 2009
216
A SUMMARY OF INDIAN CHEILANTHOID FERNS AND THE
DISCOVERY OF NEGRIPTERIS (PTERIDACEAE), AN
AFRO-ARABIAN FERN GENUS NEW TO INDIA
C.R. FRASER-JENKINS1 & C.S. DULAWAT2
1
Student Guest House, Thamel, P.O. Box no. 5555, Kathmandu, Nepal
2
Bryology Laboratory, Dept. of Botany, University College of Science, M.L.S.
University, Udaipur, Rajasthan - 313001, India (Email: [email protected])
Key words: Cheilanthes, Aleuritopteris, Notholaena, Negripteris, fern, Rajasthan,
India
ABSTRACT
A summary of Indian cheilanthoid ferns treated under nine genera includes three
new names, Notholaena dipinnata Fras.-Jenk., Cheilanthes bhutanica Fras.Jenk. & Wangdi and Cheilanthes tibetica Fras.-Jenk. & Wangdi, and five new
combinations, Cheilanthes nitidula Hook. subsp. henryi (Christ) Fras.-Jenk.,
Aleuritopteris bicolor (Roxb.) Fras.-Jenk. & Dulawat, Aleuritopteris
subdimorpha (C.B.Clarke & Baker) Fras.-Jenk. and Notholaena muelleri
(Hook.) Fras.-Jenk. Negripteris scioana (Chiov.) Pic.Serm. (Pteridaceae), a
close relative of both Aleuritopteris and Chrysochosma, was discovered by the
second author in semi-arid conditions in the Kumbhalgarh and Sitamata
Reserves of the Aravalli Hills in central Rajasthan, N.W. India, the first record
for the Indian sub-continent. It was known previously only from N.E. Africa,
Socotra and S. Arabia and is an Afro-Arabian species which, as now found,
extends eastwards into the hills of the semi-arid region of W. India.
INTRODUCTION
The cheilanthoid ferns of India have at various times been placed in the families
Sinopteridaceae, Negripteridaceae, Cheilanthaceae and Hemionitidaceae, but are now
generally accepted as belonging to subfamily Cheilanthoideae, within Pteridaceae,
with the other four families, among others, in its synonymy. The common term
cheilanthoid is a vague and undefined one, though it should be recognisable to most
fern-workers. The most obviously “cheilanthoid” ferns of India belong to nine genera,
though various other genera are also allied or within Subfam. Cheilanthoideae, but are
not dealt with here (see, for example, Tryon, Tryon & Kramer 1990). These nine genera
with their Indian subcontinental species are as follows:
1. Notholaena R.Br., includes Cosentinia Tod., Paraceterach Copel. and
Paragymnopteris K.H.Shing, perhaps also Chrysochosma (J.Sm.) Kümmerle. This
genus has long been accepted internationally as typified by N. marantae (L.) Desv.,
following Christensen (1905) and subsequently the carefully reasoned work of Pichi
Sermolli (1981, 1989). An apparent lectotypification by Smith (1875) merely used the
word “type” in the sense of “typical species”, or even “representative species” and in
several other cases Smith cited species not included by the original author of a genus,
or even cited more than one species after the word “type”. His misuse of the word
217
FERN GAZ. 18(5): 216-229. 2009
“type” is therefore not acceptable as effective lectotypification. Christensen’s proper
lectotypification was also followed world-wide, including in N. America (Tryon 1956,
1964 et al.), but Tryon & Tryon (1980) subsequently reversed it in favour of his own
local regional misapplication of the name Notholaena to the N. and C. American
segregate-genus, Chrysochosma (J.Sm.) Kümmerle, on the basis that Smith’s
“lectotypification” was earlier, though that was then rejected by the International
Committee for Nomenclature (Pichi Sermolli 1981), which Tryon did not mention
again. He thus ignored the bulk of literature across four other continents of the Old
World, which created considerable nomenclatural confusion, and was subsequently
supported locally in the USA by Yatskievych & Smith (2003). The conservation of
Notholaena with a conserved type, N. marantae, is therefore advisable to restore the
status quo ante without any room for doubt and allow current Old World authors to
continue using Notholaena in its usual sense. Neither the Australian genus
Paraceterach, nor Shing’s genus, Paragymnopteris, are accepted as replacing
Notholanea for the eight well marked Old World species. Notholaena, in its traditional
and present sense, and Chrysochosma are very close, critical genera, very difficult to
define separately, and maintained largely on molecular grounds of unknown and
possibly minor taxonomic significance. Their separability at a rank as high as genus is
somewhat doubtful, despite being accepted in the USA, but if maintained, Pichi
Sermolli (1989) has already made the great majority of necessary combinations for the
New World species within Chrysochosma. Notholaena species are frequently
exindusiate (without a pseudoindusium) and usually markedly hairy/scaly on the
surfaces and axes; they often have the sori spreading down the veins from the margin,
as a derivative from the marginal condition in many other cheilanthoid ferns. The five
Indian subcontinental species are:
N. himalaica Fras.-Jenk. (1997) (syn.: Gymnopteris vestita (Hook.) Underw., shown to
have been misplaced in that genus by Mickel (1974)). Simply pinnate, with slightly
elongated-ovate, densely silky-hairy pinnae. Widespread throughout the W. and E.
Indo-Himalayan region, except for the far west Himalaya; Tibet; and S.W. China.
N. borealisinensis (Kitag.) Fras.-Jenk. (1997) (probable syn.: Gymnopteris bipinnata
var. auriculata (Franch.) Ching). China; newly reported from the Indian subcontinent
in Bhutan (Thimphu, Taba, CRFJ 31573, 11 Oct. 2005, TAIF). Simply pinnate;
distinguishable from N. himalaica by its smaller, cordate-based pinnae, narrowing
slightly more to their apices and thus hastate-sagitate in shape.
Notholaena dipinnata Fras.-Jenk., nom. nov. for Gymnopteris bipinnata Christ, Not.
Syst. (Lecomte) 1: 55 (1909), non N. bipinnata Liebm. The lower pinnae are again
pinnate and the upper ones lobed. The relationship between this species and N.
borealisinensis is unclear as some intermediate plants appear to connect them together,
though not in the Indian subcontinent. They were therefore treated as two varieties by
Ching, but it is not considered likely here that the two extremes could merely represent
variation within a single species. The intermediates require further study. This species
has not been found in India, as the record by Fraser-Jenkins (2008), from Arunachal
Pradesh (Lohit. B. Krishna 48966, ASSAM) was in error for a scrumpled specimen of
N. borealisinensis, but it is mentioned here to explain its separation from the previous
species and because it occurs very near to the border in S.W. China.
N. marantae (L.) Desv. Sometimes stated to consist of three subspecies in Europe and
Macaronesia, subsp. marantae, subsp. subcordata (Cav.) Benl & Poelt and subsp.
cupripaleacea (Benl) Rivas Mart. et al., but these merely represent minor variation
FRASER-JENKINS & DULAWAT: INDIAN CHEILANTHOID FERNS
218
within the species and are not considered here to be appropriately ranked as subspecies,
or needing recognition. Cape Verdes; Morocco; Algeria; Ethiopia; Macaronesia; across
S. Europe; the Caucasus and Turkey; Cyprus; Lebanon; Syria; Iran; Yemen; from the
west to east Indo-Himalaya; Tibet; and S.W. China. N. marantae is diploid sexual, with
n = 29, including from Uttarakhand, and a report by Khullar, Sharma & Verma (1988)
of n = 58, from Jamunotri, Uttarakhand, is considered to have been in error (FraserJenkins 1997: 184).
N. lanuginosa (Desf.) Desv. ex Poir. (syn.: N. vellaea (Aiton) Desv., non R.Br.;
Cosentinia vellaea (Aiton) Tod.). Pichi Sermolli (1985) treated this species as a separate
genus, Cosentinia, which he revived from earlier Italian literature, on the basis of its
more trilete spores, but this has seldom been accepted. Morocco; Tunisia; Libya;
Macaronesia; across S. Europe; Turkey; Cyprus; Lebanon; Syria; Israel; Yemen; Sudan;
Iran; Afghanistan; Pakistan; N.W. India (Himachal Pradesh). A typical Mediterranean
European element in the W. Himalaya. There are two subspecies, the tetraploid subsp.
lanuginosa, which occurs throughout the range, excluding Pakistan and India, and the
very closely similar, and cryptic diploid subsp. bivalens Reichst. (Badré & Reichstein
1983), apparently confined to Macaronesia, Spain and the Indian subcontinent.
A further species is the Chinese N. sargentii (Christ) Fras.-Jenk. (1997). Two additional
Australian species, Notholaena muelleri (Hook.) Fras.-Jenk., comb. nov., basionym:
Gymnogramma muelleri Hook., Sp. Fil. 5: 143, t. 295 (1864), and N. reynoldsii
F.Muell., were formerly placed in the genus Paraceterach, which was treated as
endemic to Australia, but N. marantae and four other species were placed within
Paraceterach by Tryon (1986) in order to accommodate his misapplication of
Notholaena.
2. Cheilanthes Sw., which includes Cheilosoria Trevis. and Mildella Trevis., normally
consists of efarinose species in Asia, with generally narrow stipe-scales and small
segments. The 11 species present in the Indian subcontinent are:
C. pteridioides (Reichard) C.Chr. subsp. acrosticha (Balb.) O.Bolòs (syn.: C.
acrosticha (Balb.) Tod.,). W. Himalaya westwards to Mediterranean S.W. Asia and
Europe, N. Africa and the Cape Verdes Islands. Subsp. pteridioides (syn.: C. fragrans
(L.f.) Webb & Berth., non Sw; C. maderensis R.Lowe; Negripteris quezelii Tardieu)
occurs from Mediterranean S.W. Asia, west across Mediterranean Europe to
Macaronesia and across N. Africa, but is not present in the Indian subcontinent or near
to it. The two subspecies are different cytotypes with a close reticulate relationship and
are very similar to each other, differing only in the length of the indusium. They have
thus been much confused in earlier literature, usually under the name C. fragrans,
which is illegitimate. Following the rejection by Committee of Nardi & Reichstein’s
(1986) proposal to reject Polypodium pteridioides Reichard, in order to preserve their
nomenclature, the nomenclature of this critical complex is now stable at either the
specific or subspecific rank.
C. persica (Bory) Mett. ex Kuhn (syn.: C. szovitzii Fisch & C.A.Mey.). Far N.W.
Himalaya, ?Himachal Pradesh, W. Asia, E. Mediterranean Europe and N. Africa. The
lamina bears many long, pale hairs beneath.
C. tibetica Fras.-Jenk. & Wangdi, nom. nov., for Pellaea straminea Ching, Bull. Fan
Mem. Inst. Biol., Bot., 2: 203, t. 17 (1931), non Cheilanthes straminea Brause [=
Woodsia indusiosa Christ] (syn.: Mildella straminea (Ching) C.C.Hall & Lellinger). S.
and S.E. Tibet (type) and Bhutan. Newly reported here from the Indian subcontinent in
219
FERN GAZ. 18(5): 216-229. 2009
W. Bhutan (Chele La Pass, Ha District, T. Wangdi, CRFJ no. 31625, 12 Oct. 2005,
THIM, TAIF, det. CRFJ). Specimens from S. Tibet by the N. Sikkim border and from
Lhasa in Herb. Lloyd Botanic Garden, Darjeeling (!). Lamina pale-green, with long
rather unlobed pinnules, a pale-brown stipe and rachis and hair-tipped, linear red-brown
scales up the stipe; fronds deltate-lanceolate, bipinnate; indusia both short and long,
arising at the curled over laminar edge rather than shortly within it as in other species
formerly placed in Mildella.
C. nitidula Hook. subsp. nitidula (syn.: Pellaea nitidula (Hook.) Baker; Mildella
nitidula (Hook.) C.C.Hall & Lellinger), from the W. Himalaya only, reaching
eastwards to W. Nepal.
C. nitidula subsp. henryi (Christ) Fras.-Jenk., comb. nov., basionym: Pellaea henryi
Christ, Bull. Herb. Boissier 7: 7 (1890) (syn.: Mildella henryi (Christ) C.C. Hall &
Lellinger). N.E. India (Arunachal Pradesh), Bhutan (Thimphu, Taba, CRFJ 31496, 8
Oct. 2005, THIM, TAIF), S.E. Tibet, S. China (Yunnan (type), Szechuan, Kweichow,
Kwangtung, Fukien), Taiwan, Vietnam. Reported from Namdapha, Arunachal Pradesh,
by Singh & Panigrahi (2005) sub Aleuritopteris albomarginata (C.B.Clarke) Ching in
error. Their drawing shows A. albomarginata from some other source, that being a
higher altitude species they could not have found in the lower reaches of Namdapha
they visited. However their photograph is of C. nitidula subsp. henryi, whose presence
at Namdapha has been verified by CRFJ, from specimens previously cited by Chauhan
(1996), but misreported by them as the Afro-Arabian C. farinosa (Forssk.) Kaulf. Very
close to subsp. nitidula and differing only in the rachis hairs being denser and more
prominent and sometimes extending further around the sides of the rachis and the
smaller fronds with slightly less lobed pinna and pinna-lobes. This subspecies is a
vicariant of subsp. nitidula and replaces it in the eastern part of its range. Previous
records and collections of C. nitidula from further east referred to the present
subspecies, which merges into it morphologically. The interesting bicentric range of this
species is a well known distribution-pattern for species that prefer a somewhat drier
climate and thus occur scattered behind the Himalayan line in the otherwise wetter parts
of the central and east Himalaya.
C. opposita Kaulf. (syn.: C. mysurensis Wall. ex Hook., C. fragrans Sw.). Sri Lanka, S.
India (N. to Orissa), S. China (Yunnan, Hainan, Kwangtung, Fukien), Taiwan,
Myanmar and the Philippines. This beautiful species with narrow, upright and finely
dissect, bright green leaves has been nomenclaturally confused until recently (see
Fraser-Jenkins 1997), though its nomenclature was clarified by Alston (1936) and in
detail by Fuchs (1961). Its range has sometimes been given as throughout China and
other areas in error for the closely related, but less dissect C. chusana Hook.
C. tenuifolia (Burm.f.) Sw. Sri Lanka, through most of the Indian subcontinent except
the N.W., China, Taiwan, S.E. Asia, Australia, New Guinea and Oceania. Although
reported as diploid apomict by Verma (1961) it is now known that this species is
tetraploid sexual in India and Sri Lanka, but like apomictic species produces only 32
spores per sporangium, as do several other sexually reproducing Cheilanthes species.
C. hancockii Baker. Newly reported from the Indian subcontinent, from Taba, Thimphu,
Bhutan, CRFJ 31497, 8 Oct. 2005, THIM, TAIF; previous records from the E. IndoHimalaya were in error for Aleuritopteris bicolor. S.W. China. ?N. Thailand (C.
delicatula Tagawa & K.Iwats., which may otherwise be distinct).
C. belangeri (Bory) C.Chr. (syn.: C. varians Hook.). N.E. India, from Nepal eastwards;
Bangladesh; Myanmar; S.E. China; Thailand; and S.E. Asia to the Philippines.
220
Reported from S. India by Beddome (1864) in error for C. thwaitesii (below), which he
included within his concept of this species.
C. trichophylla Baker (syn.: Pellaea trichophylla (Baker) Ching; Cheilanthes undulata
C.Hope & C.H.Wright). S.W. China. Reported tentatively by S.C. Verma in Mehra &
Bir (1964), from the Kyangnosla pass, below Chhangu [Tsomgo] Lake, E. Sikkim
(Verma, pers. comm. to CRFJ, 2004), but the specimen lost and unverified, though
perhaps correctly recorded due to its distinct appearance, as recalled by Verma and
illustrated in a drawing in his unpublished Ph.D. thesis of 1962 at Panjab University,
Chandigarh. Distinctive zig-zag rachides and the axes and deeply tripinnatifid lamina
densely covered in short yellowish-brown hairs.
C. bhutanica Fras.-Jenk. & Wangdi, nom. nov., for Pellaea yunnanensis Ching, Act.
Phytotax. Sinica 20(2): 235 (1982), non Cheilanthes yunnanensis Brause [=
Aleuritopteris subvillosa (Hook.) Ching]. China (Yunnan (type), Szechuan), Bhutan.
Newly discovered from the Indian subcontinent by CRFJ as a common species of rocks
and walls around Thimphu and elsewhere in W. Bhutan (Serbithang, CRFJ 31487, 8
Oct. 2005, with T. Wangdi; Taba, CRFJ 31570, 11 Oct. 2005; Paro, CRFJ 31596, 12
Oct. 2005, with R. Pradhan & T. Wangdi; Chuzzom, CRFJ 31676, 15 Oct. 2005, with
T. Wangdi, THIM, TAIF). A small plant similar to a coarser-segmented and slightly less
dissect C. nitidula with more deltate fronds and no hairs above the axes; sori and
indusia continuous around the edges.
C. thwaitesii Mett. ex Kuhn (syn.: Cheilanthes laxa T.Moore ex Bedd., nom. superfl. for
C. thwaitesii; Cheilanthes keralensis N.C. Nair & S.R. Ghosh, Aleuritopteris thwaitesii
(Mett. ex Kuhn) Saiki). Sri Lanka (type) and South India; a Hindu-Lankan endemic.
Type from Sri Lanka. This often overlooked species is important in illustrating that the
presence or absence of farina is not confined to Aleuritopteris, but also occurs in some
true Cheilanthes species.
3. Aleuritopteris Fée, formerly treated by most authors as Cheilanthes subgen.
Aleuritopteris (Fée) W.C.Shieh, and all the species, except A. stenochlamys Ching ex
S.K.Wu, have names in Cheilanthes. But it was recognised generically by Ching in
Ching & Wu (1981, 1983) and subsequent Chinese workers and Saiki (1984). It appears
likely that it constitutes a distinct, if probably heterogeneous entity, which being easily
recognisable is practicably and usefully recognisable as a genus. The species are usually
well-marked and readily distinguishable by their stipe-scales (bicolorous or
concolorous) and their distribution up the axes, though most were at one time referred
to the African and Arabian species, A. farinosa (Forssk.) Kaulf., which does not occur
in Asia. The genus includes Sinopteris C.Chr. & Ching and Leptolepidium Ching &
S.K.Wu (in the sense of its type species and also in the sense of Ching & Wu, which
was based on a misapplication of the name Cheilanthes dalhousiae Hook.). It also has
two sections, the first being the less dissect, more palmate species related to A. argentea
(below) and the rest being the more pinnate species along the pattern of A. farinosa.
Morphologically Aleuritopteris is slightly difficult to define in a way that distinguishes
it from some species of Doryopteris, which are rather close to the A. argentea group,
but are perhaps more likely to be related to the imparipinnate genus, Pellaea. Most
species have rather wide segments and a strong white or yellow farina, with wide stipescales. Due to considerable lack of clarity and confusion, the cytology and reported
aneuploid base-numbers for various species need to be carefully reinvestigated with
accurate identification and preservation of voucher-specimens. The 18 species present
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FERN GAZ. 18(5): 216-229. 2009
in the Indian subcontinent have mostly been detailed by Fraser-Jenkins (1992, 1993 and
1997) and are:
A. argentea (S.G.Gmel.) Fée (syn.: A. flava (Ching & S.K.Wu) S.R.Ghosh, non Saiki,
nec sensu Ghosh [= A. subargentea]). Far N.E. India, a single collection of J.D. Hooker
& T. Thomson’s from the Khasi Hills, Meghalaya (K!), might be correctly localised, but
it has never been collected since in this generally well known area and is normally a
higher-altitude, and usually higher latitude species. A similar case occurs with their
collection at Kew of Lycopodium annotinum L. subsp. alpestre (Hartm.) Å. Löve & D.
Löve, said to be from Khasia, but almost certainly from N. Sikkim. However A.
argentea (S.G.Gmel.) Fée, which otherwise occurs in W., S.W., C., N. and E. China;
Tibet; Taiwan; Korea; Japan; and Siberia, has now been discovered in Bhutan (Taba,
near Thimphu, CRFJ 31500, 8 Oct. 2005, THIM, TAIF), and was previously collected
from N. Lohit, at the Tibetan border in or on the border of Arunachal Pradesh (F.
Kingdon-Ward, BM!). A further species, close to A. argentea, but without white farina
and with small, narrow, less lobed segments has also turned up in N.E. Arunachal
Pradesh (Changlang, Namdapha, Shirung to Hunung, c. 1100 m., B.K. Shukla 88207, 7
Feb. 1986, ASSAM, det. CRFJ). It appears to belong to the efarinose Chinese species,
A. shensiensis Ching, though further comparative study may be required.
A. subargentea Ching ex S.K.Wu. Similar to A. argentea but slightly more dissect and
the frond slightly more pinnately arranged, less palmate. Newly discovered in the
Indian subcontinent (Marpha to Tukuche, Mustang, N.C. Nepal, CRFJ 30509, 27 June
2004, with G. Tamang, TAIF); Songgong, Sikkim, 1,400ft, Ribu & Rhomoo Lepcha, for
G.H. Cave 7634, 4 Oct. 1923 (Herb. Lloyd Botanic Garden, Darjeeling!, det. CRFJ).
Tibet; S.W. China.
A. tamburii (Hook.) Ching. N.C. Nepal; N. Sikkim; Meghalaya; Tibet and S.W. China.
A very distinctive, large species of the A. argentea group, with wide, coarse lobes and
bright white farina.
A. subvillosa (Hook.) Ching (syn.: A. tenella (Ching & S.K.Wu) Saiki). W. to E. IndoHimalaya; Tibet; S.W. China. Efarinose, but closely related to the farinose A. caesia
(Christ) Ching and A. kuhnii (Milde) Ching, from Tibet and S.W. or C. to N.E. China;
Japan (the latter reported from India in error for A. dalhousiae by Dixit (1984)).
A. duthiei (Baker) Ching. W. Indo-Himalaya (very rare); N.W. Nepal; Bhutan.
Efarinose, related to A. leptolepis (Fras.-Jenk.) Fras.-Jenk., but with a wider-triangular
and more dissect lamina; both have wide, pale stipe-scales.
A. leptolepis (Fras.-Jenk.) Fras.-Jenk. (2008) (misapplied name: A. dalhousieae (Hook.)
Ching, nom. utrique rejic.). The totally efarinose, higher-altitude species long known as
A. dalhousieae [“dalhousiae”] in error due to Hooker’s confusing the efarinose Summer
(monsoon) fronds of A. albomarginata (below) with this species. W. to E. IndoHimalaya; Myanmar; Tibet; S.W. China. Misreported sub “C. dalhousiae” sensu auct.
Ind. by Dixit (1996) from Orissa, in error for A. bicolor with the powder washed off by
alcohol during herbarium-poisoning.
A. rufa (D.Don) Ching. Common from the W. to E. Indo-Himalaya; also in S.W. China;
Myanmar; Thailand. The fronds are lanceolate, narrowing slightly to the base, and the
axes are densely covered with narrow, hair-like, fibrillose scales.
A. dubia (C.Hope) Ching (syn.: A. subrufa (Baker) Ching; C. leveillei Christ; A.
humatoides Saiki; C. wusukungii Miyamoto & Ohba). Intermediate between the last
and next species and sharing the same phytochemistry of its white flavonoid powder
(Fraser-Jenkins & Wollenweber in prep.). W. to E. Indo-Himalaya; N. Western Ghats;
222
Myanmar; Tibet; S.W. and S. China; Taiwan; Thailand; the Philippines. Confused in
China and reported as the next species.
A. albomarginata (C.B.Clarke) Panigrahi (syn.: A. dalhousieae (Hook.) Ching, nom.
utrique rejic., non sensu auct. Ind.). Widespread at somewhat higher altitudes from the
far W. to N.E. Himalaya; Orissa; N. Western Ghats; Tibet; S.W. China; Taiwan;
Thailand; Vietnam. The fronds are deltate and instead of hair-like scales have
bicolorous scales extending up the stipe, rachis and costae. The Summer (monsoon)
fronds are taller and more developed and almost or quite without farina, unlike the
smaller, basal Winter fronds.
A. chrysophylla (Hook.) Ching (syn.: A. humatifolia X.C.Zhang & L.Shi; A.
flavopygmaea S.R.Ghosh). The only species with bright sulphur-yellow farina beneath
in the Indian subcontinent. The W. Indo-Himalaya (Simla); C. Nepal to N.E. India;
Myanmar; Tibet; S.W. China; Thailand.
A. formosana (Hayata) Tagawa (syn.: C. brevifrons (Khullar) Khullar). One of the most
widespread species, frequently reported from China under the name A. anceps in error.
W. Africa (Guinea); far W. to the E. Indo-Himalaya; Rajasthan (C.S. Dulawat, det.
CRFJ); Uttar Pradesh; Bihar (Parasnath); Orissa; N.W. Ghats; Arunachal Pradesh;
Tibet; S.W. and S. China; Taiwan; Myanmar; Thailand; the Philippines. It is fairly close
to A. anceps, but has a narrower and usually smaller frond, which is characteristically
bullulate-wrinkled above, and the slightly narrower, bicolorous stipe-scales extend up
the rachis as well, but not (except rarely an odd one) onto the costae. The rachis usually
bears scattered glands.
A. anceps (Blanf.) Panigrahi (syn.: A. pseudofarinosa Ching & S.K.Wu; A. interrupta
Saiki; ?A. javanensis Saiki, non C. javanensis (Willd.) T.Moore). Widespread at lower
altitudes in India etc., far W. to N.E. Himalaya; Rajasthan; N.W. Ghats; C. India; S.
India; Sri Lanka; Tibet; S.W. China (rare); Taiwan; Myanmar; Thailand; ?Java; ?Timor.
The lamina is deltate-lanceolate, but not as triangular and wide-based as in C. bicolor
(below), and has a brighter white farina; the slightly wide, bicolorous stipe-scales do
not normally extend above the top of the stipe.
A. dealbata Fée (syn.: C. dealbata D.Don, non Pursh; A. doniana S.K.Wu ex Ching,
nom. superfl.; C. doniana Fras.-Jenk. & Khullar; A. sikkimensis S.R.Ghosh). E. part of
the W. Himalaya (Uttarakhand); Nepal to N.E. India; Tibet; S.W. China; Myanmar;
??Thailand. This species is closely related to C. anceps, but has a longer, often very
large frond and wider, less lobed segments, with narrower indusia, and a very bright
white farina. The stipe-scales, though vaguely bicolorous, do not have such an obvious
dark central stripe and tend to have a yellower basal region and slightly darker apical
region. It was illustrated from Nagarjun, Kathmandu, Nepal, on the front dust-jacket by
Fraser-Jenkins (1997).
A. bullosa (Kunze) Ching (syn.: ?A. indica Fée; A. flaccida (Bedd.) B.K.Nayar &
S.Kaur; C. flaccida (Bedd.) Mehra & Bir, non sensu Mehra & Bir [= A. bicolor]). Sri
Lanka and S. India. A very large species with long fronds, a bullulate upper surface,
well lobed pinnules, thick stipes and concolorous red stipe-base scales.
A small-sized, more creamy-yellowish powdered segregate of A. dealbata from S. India
(especially the Shevaroy Hills) is A. wollenweberi Fras.-Jenk. (2008), superficially
similar to the Chinese, Taiwan and Japanese A. krameri (Franch. & Sav.) Ching.
A. stenochlamys Ching ex S.K.Wu. E. part of the W. Himalaya (Uttarakhand); Nepal;
Bhutan; Manipur; Tibet, S.W. China. This rare, high-altitude taxon is closely related to
A. grisea and is perhaps of slightly doubtful status, requiring further study, though
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FERN GAZ. 18(5): 216-229. 2009
possibly a good species. Fraser-Jenkins previously confused its type and used the name
C. stenochlamys for it in error.
A. grisea (Blanf.) Panigrahi, non sensu Panigrahi (syn.: A. platychlamys Ching; C.
platychlamys (Ching) Fras.-Jenk.). A high-altitude Himalayan species with concolorous
red stipe-scales; widely misrecorded from C. or S. India and Orissa by Panigrahi (1965)
and Dixit (1996) in error for A. formosana among other species. Far W. to E. IndoHimalaya; Tibet, S.W. China; Taiwan.
A. bicolor (Roxb.) Fras.-Jenk., comb. nov., basionym: Pteris bicolor Roxb. in Griff.,
Calcutta J. Nat. Hist. 4: 507 (1844) (syn.: Cheilanthes farinosa var. tenera C.B.Clarke
& Baker; C. bicolor (Roxb.) Griff. ex Fras.-Jenk.; A. bicolor (Roxb.) Punetha & Kholia,
comb. inval., sin. basionym; A. kathmanduensis Ching & S.K.Wu; misapplied name:
Cheilanthes farinosa sensu auct. Ind. plur., Blanf. et al., non (Forssk.) Kaulf.). A very
common and widespread, rather low-altitude species in India. W. Africa (Nigeria, Jos);
far W. to N.E. Indo-Himalaya; Rajasthan; Bihar (Parasnath); C. India; Orissa; S. India;
Bangladesh; Myanmar; ?Thailand; Laos. A similar taxon, but more delicate and with
russet, concolorous stipe-base scales (like those of A. subdimorpha) occurs in Sumatra.
There is some apparent transition, or some intermediate taxon, between the otherwise
highly distinct A. bicolor and A. anceps (and also somewhat towards A. subdimorpha)
in S.E. Bangladesh (Chittagong Hills), S. India, Myanmar, S.E. China and ?New
Guinea (Papua). But it is not yet clear why this should appear to be so.
A. subdimorpha (C.B.Clarke & Baker) Fras.-Jenk., comb. nov., basionym.:
Cheilanthes farinosa (Forssk.) Kaulf. var. subdimorpha C.B.Clarke & Baker, J. Linn.
Soc., Lond. 24: 411 (1888) (syn.: Cheilanthes subdimorpha (C.B.Clarke & Baker)
Hieron.; A. longipes Ching & S.K.Wu, nom. inval., sin. typ. (which is not A. bicolor, as
tentatively thought by Fraser-Jenkins 1997, in the absence of the type); C. longipes
(Ching & S.K.Wu) Dixit & Bal Krishna, comb. inval.; A. pentagona Saiki; A.
pseudoargentea S.K.Wu). From C. Nepal eastwards to N.E. India; Arunachal Pradesh;
Manipur; abundant in Meghalaya; ?Bangladesh; Myanmar; Thailand; S.W. to S.E.
China; Vietnam. This rather little known species is generally similar to C. bicolor, but
has a longer, thicker stipe, shorter and more coarsely lobed lamina and concolorous,
russet stipe-scales.
4. Negripteris Pic.Serm. is closely similar to Aleuritopteris but differs in having only
rather few (1-4) sporangia per sporangium (also shown by some Aleuritopteris), which
are rather deeply embedded in the laminar powder (“subsessile”) and an apparently
primitive, large sporangium, without a definite stomium area and a very broad annulus
with all its walls thickened (indurated), including the outer one, appearing like a solid,
dark cap to the sporangium. During dehiscence, the sporangium splits laterally and the
whole annulus or top half of the sporangium falls off to release the spores. This may be
a retained adaptive feature connected with a very dry climate, allowing protection of the
developing sporangia against desiccation. However, on the strength of this
hypothetically being supposed to be a very primitive characteristic Pichi Sermolli not
only raised a new genus, but even a new family for the single specimen he had seen
(which was undoubtedly and obviously mistaken), despite pointing out its evident
similarity to Aleuritopteris. In all its other features it is not separable from
Aleuritopteris in its concolorous, lanceolate, pale-reddish stipe-scales, frond-shape,
white farina beneath and shallow, interrupted pseudo-indusia formed by the leafmargin. Weatherby (1948) pointed out tendencies within what is now Chrysochosma
224
from N. America, towards a similar annulus and embedded sporangium, and pointed out
the likeness of N. scioana to some “Notholaena” species (i.e. Chrysochosma spp.).
While not in any way invalidating the genera Aleuritopteris and Notholaena, it must be
said that a few species in either genus can hardly be distinguished morphologically from
the other genus. The likelihood is therefore that Negripteris is either an Aleuritopteris
or a Chrysochosma, or may perhaps be nearer to an ancient ancestor of both. However
it is maintained here as a somewhat dubious genus, resting on its peculiar sporangia,
pending further study. A. rosulata (C.Chr.) Ching (syn.: A. pygmaea Ching), from Tibet
and S.W. China, has similar scales and lamina (with the lobes rather broadly joined at
their bases and fusing at the apex as in Negripteris) and appears to be very close to it,
but without the characteristic sporangia. A rare S.W. Chinese species, A. sichouensis
Ching & S.K.Wu, is again similar and by definition only would most probably be
placed in Chrysochosma.
N. scioana (Chiov.) Pic.Serm. (syn.: Mohria scioana Chiov.; Negripteris tricholepifera
Pic.Serm.). An Afro-Arabian species, formerly known from N.E. Africa and South
Arabia only; Ethiopia, Sudan, Somalia, Socotra (also seen there by the first author in
1967), N. Kenya, Yemen, Saudi Arabia. To this must now be added a remarkable
discovery of it from W. India in the semi-arid hills of Rajasthan. Last year, Dr. C.S.
Dulawat (the second author) sent the first author material of his Rajasthan
Aleuritopteris for identification and most surprisingly there was some fine, unidentified
material of typical Negripteris scioana (Chiov.) Pic.Serm. among them that he had
collected from the Kumbhalgarh and Sitamata Sanctuaries, Aravalli Hills and
Chittorgarh, Rajasthan. This discovery is of considerable phytogeographical interest as
it was otherwise only known to be in N.E. Africa and S. Arabia and fits in with a small
handful of other Afro-Arabian connections in Western India, mainly in Rajasthan. The
photograph purported to be Cheilanthes albomarginata C.B.Clarke sensu lato
published by Chaudhary & Khichi (2007) is actually of the tip of the frond of
Negripteris from Sitamata. Specimens collected are:
1. Rajasthan, Sitamata Wild Life Sanctuary, Ambaretti, crevices of rocks, 500-800
m. C.S. Dulawat CSD/SM/05-147, 14 Oct. 2005, Herb. Botany Dept., M.L.S.
University, Udaipur.
2. Rajasthan, Sitamata Wild Life Sanctuary, Ambaretti, crevices of rocks and bank
of nalah [stream], c. 500-800m. C.S. Dulawat CSD/SM/06-197, 5 Oct. 2006, Herb.
Botany Dept., M.L.S. University, Udaipur.
3. Rajasthan, Khumbhalgarh Wild Life Sanctuary, Ranakpur Ghat, crevices of
rocks, 550-800 m. C.S. Dulawat & B.L. Chaudhary CSD/SM/05-183, 23 Oct. 2005,
BSD.
5. Doryopteris J.Sm. Characterised by the long, uninterrupted sori and often palmate
fronds, but though widely and usefully accepted, is difficult to define exclusively,
especially in relation to the Aleuritopteris argentea group, which also contains some
efarinose species. Two species occur in the Indian subcontinent:
D. concolor (Langsd. & Fisch.) Kuhn (syn.: Doryopteris geraniifolia (Raddi)
Klotzsch). S. and C. America; Africa; Sri Lanka; S. India (N. to Orissa); S. China;
throughout S.E. Asia; Australia; Oceania. D. kirkii (Hook.) Alston from Africa has
occasionally been reported from India in error for D. concolor, but is anyway a
somewhat doubtful taxon. The taxonomic position of this fern has long been in some
doubt as it is very similar to an efarinose member of the Aleuritopteris argentea group
225
FERN GAZ. 18(5): 216-229. 2009
and has often been placed in Cheilanthes. Pending further study, including molecular
DNA work, it is maintained here in Doryopteris, where it appears most likely to belong
and has more usually been placed.
D. ludens (Wall. ex Hook.) J.Sm. A distinctive species with tall black stipes and black
midribs and the horizontal frond varying from cordate to considerably and deeply
palmate-pinnately lobed into long, narrow segments, the lowest lobe basiscopically
lobed again and the fertile fronds taller and more narrowly lobed than the sterile. It
occurs in Orissa (Dixit 1996) and from N.E. India eastwards; Assam; Manipur;
Nagaland; Tripura; Mizoram; Bangladesh; Myanmar; S.W. and S. China; Thailand; and
S.E. Asia. Hope (1901) reported it from Chitral, N.W. Pakistan, in error on the basis of
a specimen given to him by General Gatacre, ostensibly collected during the Chitral
Relief Military Expedition, at Kaffir Rock on the road S. of Ziarat, along with Lygodium
microphyllum (Link) R.Br. The specimens of both species, along with drawings and a
sketch-map provided by Gatacre are in DD (!) and are correct. But anyone who knows
the Pakistan fern-flora (Fraser-Jenkins 1992, 1993) and who has been there, where
Kaffir Rock still exists beside the main road into Chitral from the Lowarai Pass, would
know that neither species could possibly occur there or anywhere within thousands of
miles. The specimens were almost certainly collected during Gatacre’s duty in
Myanmar shortly previous to the Chitral expedition and seem most likely to have been
more in the way of a “military prank”, than due to confusion. It is surprising that such
an experienced expert as Hope could possibly have swallowed it!
6. Pellaea Link, nom. cons. Although various species, including those subsequently
placed in Mildella by Hall & Lellinger (1967) and now in Cheilanthes, have been put
into Pellaea in the past because of their glabrous, efarinose segments with long sori, the
genus is now confined to those species with imparipinnate fronds, the apical segment
being similar to a lateral one. The species may be glabrous or hirsute. Ghosh (1985)
claimed to have “done” the taxonomy of Indian Pellaea and listed 3 species, but the 5
species occurring in India are as follows:
P. falcata (R.Br.) Fée (syn.: P. seticaulis (Hook.) S.R.Ghosh). This species has simple,
unlobed, elongated, nearly glabrous pinnae with a only few hair-like scales beneath, and
a hairy and scaly stipe and rachis, similar in both Australasia and S. India. It is absent
from Malaysia (given by Ghosh), Hooker’s type-locality of “Penang” being in error for
Lady Dalhousie’s collection from Sri Lanka (from where it was reported by Sledge
1982), which was omitted by Ghosh (1985).
P. longipilosa Bonap. (syn.: P. malabarica B.K.Nayar & Geev.). Similar to P. falcata,
but with narrow, tripartite lower and mid pinnae. Africa; S. India; confined to a few
localities in Kerala. Though redescribed as if a new, endemic species, Fraser-Jenkins
has reidentified it as being the African species, P. longipilosa, which is thus one of a
rather small group of tropical African elements in S. India.
P. boivinii Hook. A rather small, bipinnate species with from 2 to 3 pairs of elongated
ovate, articulated-stalked pinnules on the lower pinnae and all the axes densely covered
in very short, dark, blackish-brown hairs. S. and E. Africa; Madagascar; Mascarenes;
Sri Lanka (Sledge 1982, omitted by Ghosh 1985); and S. India. Another tropical African
element in S. India.
P. viridis (Forssk.) Prantl. Another bipinnate species, with slightly larger, thinner and
non-articulate-stalked, often biauriculate-based segments and glabrous axes. Adventive
in S. India and Sri Lanka (Sledge 1982). Also naturalised in Australia and Oceania. S.
226
and E. Africa; Madagascar; and Yemen (type).
P. calomelanos (Sw.) Link non Pityrogramma calomelanos (L.) Link. (syn.: P. hastata
(Thunb.) Prantl, non (L.) Thunb.). A rare and very seldom-collected species occurring
in a few scattered localities in very dry rocky areas. The stalked leaf-segments are
cordate-hastate and without scales, borne on glossy black costae. Africa; Pakistan
(Swat; Hazara); Himachal Pradesh; Uttarakhand; W. Nepal; S.W. China. An AfroArabian element reaching the W. Himalaya.
7. Pityrogramma Link. The “silver ferns” and “gold ferns” of cultivation where they
often spread by self-sporing. They are exindusiate and the sori spread along the veins
and cover the whole surface beneath the segments. Four New World species are
adventive in the Indian subcontinent:
P. calomelanos (L.) Link. The farina beneath the leaf, which is usually rather thin and
weak, is white. S. and C. America. Adventive throughout the world in warmer climates
and so abundant in even remote places throughout nearly all of the Indian subcontinent
that it is sometimes difficult to remember that it is in fact an alien species not recorded
before the later 19th Century.
P. austroamericana Domin (syn.: Pityrogramma calomelanos var. austroamericana
(Domin) Farw.; P. calomelanos (L.) Kaulf. var. aureoflava (Hook.) Weath. ex
F.M.Bailey; misapplied name: P. chrysophylla (Sw.) Link). The farina beneath the leaf
is sulphur yellow. S. and C. America. A common adventive in Sri Lanka and S. India.
In addition to the bright yellow powder, which is often lost in old specimens by
treatment with alcohol (containing pesticides), it also has a shorter lamina with rather
shorter and often less lobed segments. For some reason it has not reached N. India to
date. It was first recognised as a separate species by Domin (1928, 1929) and detailed
further in his later publications. The genus was monographed by Tryon (1962), and
information was thence extracted by Panigrahi (1975) and from the determinations at
Kew.
P. dealbata (C.Presl) R.Tryon. White farina, a small, thin lamina with well lobed and
toothed segments. C. America. Adventive in Sri Lanka, but probably not well
established.
P. sulphurea (Sw.) Maxon. Pale lemon-yellow, rarely white farina and the frond
narrowed towards the base, with the ultimate segments cuneate-based and fanned out or
flabellate at their strongly toothed apices. C. America. Adventive in Samoa and Sri
Lanka, but probably not well established in the latter.
8. Parahemionitis Panigrahi. Following Mickel’s (1974) establishing that the Indian
species is not a true Hemionitis, and Tryon, Tryon & Kramer’s (1990) clear statement
that “H. arifolia” is not readily included in any genus, Panigrahi (1993) took it upon
himself to utilise that information and make several attempts to interject a new generic
name of his own for it (see Fraser-Jenkins 1997: 187-188), finally succeeding in
validating one. Parahemionitis is therefore accepted here as the name for the genus and
its single species-complex.
P. cordifolia (Roxb.) Fras.-Jenk. (misapplied name: Hemionitis arifolia (Burm.f.)
T.Moore). Distinctive cordate-rounded leaves. N.E. India (Bihar; Assam; Manipur;
Nagaland; Tripura; Mizoram); Sri Lanka; S. India; Bangladesh; Myanmar; S. China;
Vietnam; Malesia; Philippines. The widespread Indian triploid cytotype may often be
proliferous with small plantlets developing in the axil of the leaf during the wet season,
227
FERN GAZ. 18(5): 216-229. 2009
but dropping off during the dry season. A Chinese diploid, which is more delicate, is
never proliferous (Huang, Manickam & Chiou 2007).
9. Hemionitis L. (syn.: Gymnopteris Bernh.; Gymnogramme Desv.). The detailed work
carried out by Mickel and his colleagues (Mickel 1974, 1988, Gianassi & Mickel 1979,
Ranker 1989) on this genus has been usefully summarised in Pichi Sermolli &
Bizzarri’s (2005: 66-68) final magnum opus. One species in the Indian subcontinent:
H. tomentosa (Lam.) Raddi (syn.: Gymnopteris tomentosa (Lam.) Underw.) S. America.
Adventive and very common in Sri Lanka.
CONCLUSION
The discovery of the African Negripteris in India is not only of considerable
phytogeographical interest, but also suggests that further collection and study by Indian
pteridologists with a specialist knowledge, able to recognise all the species they see,
may reveal a number of other species hitherto overlooked in the subcontinent. The
discovery of Notholaena borealisinensis, Cheilanthes hancockii, C. bhutanica, C.
tibetica and Aleuritopteris subargentea, all previously unknown in the Indian
subcontinent are further examples.
ACKNOWLEDGEMENTS
The author thanks Dr. Tandin Wangdi of the National Biodiversity Conservation Centre,
Serbithang, Thimphu, Bhutan, who, along with Dr. Rebecca Pradhan, kindly
accompanied him during collection excursions in W. Bhutan in October 2005 and is a
joint authority of some of the resulting new taxa, above.
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Hemionitis, Gymnopteris and Bommeria (Adiantaceae). Amer. J. Bot. 76(2): 297306.
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FERN GAZ. 18(5):230-263. 2009
230
CHECKLIST OF THE FERNS AND LYCOPHYTES OF ACRE STATE,
BRAZIL
J. PRADO1 & R. C. MORAN2
1Instituto de Botânica, Seção de Curadoria do Herbário, C.P. 3005, CEP 01061-970,
São Paulo, SP, Brazil
E -mail: [email protected]
2The New York Botanical Garden, Bronx, NY 10456-5126, U.S.A.
Key words: checklist, Brazil, Acre State, Amazonian forest, ferns, lycophytes
ABSTRACT
One hundred and seventy eight species and five varieties in 24 families and 60
genera of ferns and lycophytes are recorded for state of Acre, Brazil. Data about
habit, habitat, material examined, and distribution of each taxon are also
presented. Acre contains about one-third of the species of ferns and lycophytes
estimated to occur in the Brazilian Amazon region.
INTRODUCTION
The Project of the Flora of Acre State, Brazil, is a collaborative endeavour between the
Federal University of Acre (UFAC) and The New York Botanical Garden (NY). For
some groups of plants, specialists were invited to contribute, such as for ferns and
lycophytes. This checklist is a result of the collaborative project.
The first expeditions to Acre and the Brazilian Amazon took place in 1900-1901
when Ernest Heinrich Georg Ule collected along the rivers Tejo and Juruá-Mirim.
Locally, this area is called “Região do Alto Rio Juruá.” In 1911-1912, a second
expedition was conducted by Ule in the region of the Acre River. The first Brazilian
botanist to explore the state was João Geraldo Kuhlmann, who travelled with Marechal
Rondon through Amazonia in 1923. Later, in 1933, Boris Alexander Krukoff collected
in the region, especially around the Macauã River, in the Basin of Purus River. During
the 1960s and 1970s, several botantists collected in Acre State, the most prominent
being João Murça Pires, Ghillean T. Prance, Enrique Forero, Paul J. Maas, and Paulo G.
Windisch. Other important collections from Acre were made for the Project
RADAMBRASIL during the 1970s by Luis Coêlho, Cid Ferreira, Bruce Nelson, and
Thomas Croat. They were among the members of three expeditions to Acre (Silveira,
2003).
In 1979 the Herbarium of the Federal Universty of Acre (HPZ) was established. Its
main goal was to have a reference collection for the flora of Acre. In 1990 the herbarium
signed an international agreement of cooperation with The New York Botanical Garden
(NY) to promote studies of the Acre flora. During the last twenty years Douglas Daly
(NY), Marcos Silveira (UFAC), and several other collaborators from many institutions
have greatly increased the number of collections in Acre. According to Daly (personal
communication), at the beginning of the project “Floristics and Economic Botany of
Acre, Brazil” the Herbarium of the Federal University of Acre (HPZ) had 2,000
specimens. It now has more than 30,000 specimens, including non-vascular and
vascular plants.
These collections provided many new records of ferns (e.g., Adiantum decoratum
231
FERN GAZ. 18(5): 230-263. 2009
Maxon & Weath., A. poeppigianum (Kuhn) Hieron. A. scalare R. M. Tryon, Bolbitis
oligarchica (Baker) Hennip., Tectaria draconoptera (D. C. Eaton) Copel., Thelypteris
membranacea (Mett.) R. M. Tryon; Labiak & Prado, 2007) and new species (e.g.,
Adiantum windischii J. Prado and Triplophyllum boliviense J. Prado & R. C. Moran;
Prado, 2005; Prado & Moran, 2008).
Acre State is located in northwestern Brazil in the southwestern part of the Brazilian
Amazon. It covers an area about 153,149 km2 and harbours great variation in
topography, soils, and climates. This variation promotes a high diversity of vegetation
types, floristic affinities, and life forms. Several factors divide the state into two
regions: the southeastern region is drained by the Purus River and by the Basin of the
Rivers Acre/Purus. Here there is a pronounced dry season, and several species are
shared with Central Brazil and with other extra Amazonian vegetation types (Prado &
Gibbs, 1993). The second region is northwestern Acre, drained by the Juruá River. This
area is continuously wet, lacking a dry season. Along its border with Peru are the
mountainous Moa Range and Divisor Range (locally called the Serra do Moa and Serra
do Divisor, respectively). These ranges shelter several Andean species and genera such
as Cyathea bipinnatifida (Baker) Domin (Cyatheaceae), Pteris haenkeana C. Presl,
(Pteridaceae), Solanopteris bifrons (Hook.) Copel. (Polypodiaceae); flowering plant
examples include Croton lechleri Müll Arg. (Euphorbiaceae), Ladenbergia lambertiana
(A. Braun ex Mart.) Klotzsch, L. oblongifolia (Rubiaceae), and Monolena primuliflora
Hook. f. (Melastomataceae)). This region has open vegetation called “Campinas” and
“Caatingas Amazônicas” on whitish sandy soils. The flora of this region has affinities
with those of the Rivers Rio Madeira and Negro (Silveira, 2003).
The
preliminary
checklist
of
the
Acre
Flora
(http://www.nybg.org/bsci/acre/www1/checklist.html), which includes all vascular
plants and fungi, lists about 3,273 species (including infraspecific taxa), of which 156
are either ferns or lycophytes, these belonging to 50 genera.
In general, ferns and lycophytes from Acre are poorly known and papers containing
some information about them are few and sparse (Tryon & Conant, 1975; Windisch,
1979; Prado, 2005, and Labiak & Prado, 2007). The present study revealed 178 species,
60 genera, and five varieties in 24 families (Table 1). According to our estimates,
Amazonian Brazil has ca. 550 species of ferns and lycophytes; thus, Acre contains
about 32% of that total number.
LIST OF SPECIES
The present list is based on material from the following herbaria: CEN, HB, HPZ,
HRCB, INPA, K, MG, MO, NY, R, RB, SP, UC. In 2001 the senior author collected
about 250 specimens in Acre. In total, about 820 specimens have been examined for the
present checklist.
The list is organised in alphabetic order by families, genera, species, and varieties.
The arrangement of familes and genera follows Smith et al. (2006). For each species
and variety the full name of the taxon is presented with the reference of the original
publication, as well as additional information such as: habit, habitat, material examined,
and geographic distribution. Voucher information for each species and variety is
organised in alphabetic order by municipality; material collected by same collector for
the same locality appears in ascending numerical order.
Author abbreviations follow Pichi-Sermolli (1996). In general, synonyms are
omitted, except the basionym of the accepted name.
PRADO & MORAN: CHECKLIST OF THE FERNS OF ACRE STATE, BRAZIL 232
Table 1. Synopsis of taxa in Acre State, Brazil.
Families
Number of genera
Number of spp.
Anemiaceae
1
1
Aspleniaceae
1
11
Blechnaceae
2
3
Cyatheaceae
2
4
Dennstaedtiaceae
2
2
Dryopteridaceae
10
21
Gleicheniaceae
2
2
Hymenophyllaceae
4
17
Lindsaeaceae
1
7
Lomariopsidaceae
3
8
Lycopodiaceae
2
2
Lygodiaceae
1
2
Marattiaceae
1
3
Metaxyaceae
1
1
Polypodiaceae
9
28
Psilotaceae
1
1
Pteridaceae
8
30
Saccolomataceae
1
3
Salviniaceae
1
1
Schizaeaceae
1
1
Selaginellaceae
1
7
Tectariaceae
2
6
Thelypteridaceae
1
12
Woodsiaceae
2
5
Totals: 24
60
178
Number of varieties
1
3
1
5
233
FERN GAZ. 18(5): 230-263. 2009
ANEMIACEAE
Anemia phyllitidis (L.) Sw., Syn. Fil. 155. 1806.
Osmunda phyllitidis L., Sp. Pl. 1064. 1753.
Habit/Habitat: Herb; terrestrial on riverside vegetation and on Terra Firme forest.
Material examined: BRAZIL. ACRE: Assis Brasil, D. C. Daly et al. 9779 (HPZ, NY);
Brasiléia, D. C. Daly et al. 9895 (HPZ, NY).
Distribution: Mexico, Greater Antilles, except Puerto Rico, Mesoamerica, tropical and
subtropical South America.
ASPLENIACEAE
Asplenium angustum Sw., Vet. Ak. Handl. 38: 66, tab. 4, fig. 1. 1817.
Habit/Habitat: Herb; epiphyte in forest on Terra Firme.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, G. T. Prance et al. 12637 (NY);
Mâncio Lima, D. C. Daly et al. 8953 (HPZ, NY).
Distribution: N South America.
Asplenium auritum Sw., J. Bot. (Schrader) 1800(2): 52. 1801.
Material examined: BRAZIL. ACRE: Manoel Urbano, D. C. Daly et al. 11466 (NY);
Mâncio Lima, G. T. Prance et al. 11977 (NY); Marechal Thaumaturgo, D.C. Daly
10531 (NY); Porto Valter, P. J. M. Maas et al. P12930 (MG, NY), P. J. M. Maas et al.
P13180 (NY); Santa Rosa, D. C. Daly et al. 11305 (NY); Sena Madureira, M. Silveira
et al. 617 (HPZ, NY); Tarauacá, G. T. Prance 7247 (MG, NY), G. T. Prance et al. 7382
(NY), M. Silveira et al. 921 (NY).
Distribution: S Mexico, Antilles, Mesoamerica, N South America.
Asplenium cirrhatum Rich. ex Willd., Sp. Pl. 5: 321. 1801.
Habit/Habitat: Herb; epiphyte on flooded vegetation.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, G. T. Prance et al. 12202 (NY);
Mâncio Lima, M. Silveira et al. 1245 (NY).
Distribution: Mexico, Antilles, Mesoamerica and tropical South America.
Asplenium cuneatum Lam., Encycl. 2: 309. 1786.
Habit/Habitat: Herb; epiphyte on Terra Firme.
Material examined: BRAZIL. ACRE: Porto Valter, P. J. M. Maas et al. P13103 (NY).
Distribution: Antilles, Mesoamerica and tropical South America, Polynesia, Africa.
Asplenium delitescens (Maxon) L. D. Gómez, Brenesia 8: 52. 1976.
Diplazium delitescens Maxon, Contr. U.S. Natl. Herb. 10: 497. 1908.
Habit/Habitat: Herb; terrestrial grows riverside vegetation and on Terra Firme forest.
Marechal Thaumaturgo, D. C. Daly et al. 7337 (HPZ, NY).
Asplenium hallii Hook., Sp. Fil. 3: 202. 1860.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1166 (HPZ, SP),
J. Prado et al. 1306 (HPZ, SP), J. Prado et al. 1379 (HPZ, SP), L. R. Marinho 251
(NY); P. J. M. Maas et al.P12794 (NY); Mâncio Lima, G. T. Prance et al. 2850 (MG,
NY); G. T. Prance et al. 12246A (MG, NY); J. Prado et al. 1225 (HPZ, SP).
Asplenium juglandifolium Lam., Encycl. 2: 307. 1786.
Habit/Habitat: Herb; epiphyte on flooded vegetation near river margins.
Material examined: BRAZIL. ACRE: Bujari, D. C. Daly et al. 8464 (HPZ, NY), D. C.
Daly et al. 9330 (NY); Cruzeiro do Sul, G. T. Prance et al. 2786 (MG, NY); Mâncio
Lima, J. Prado et al. 1224 (HPZ, SP), J. Prado et al. 1318 (HPZ, SP); Marechal
Thaumaturgo, D. C. Daly et al. 7764 (NY); Porto Acre, A. R. S. Oliveira et al. 761
(NY); Rodrigues Alves, J. Prado et al. 1262 (HPZ, SP).
Distribution: Mexico, Jamaica, Mesoamerica, tropical South America.
Asplenium ortegae Murakami & R. C. Moran, Ann. Missouri Bot. Gard. 80: 23. 1993.
Habit/Habitat: Herb; terrestrial .
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, D. C. Daly et al. 11707 (NY).
Asplenium pearcei Baker, Syn. Fil. (ed. 2) 483. 1874.
Habit/Habitat: Herb; epiphyte on trees in Terra Firme forests.
Material examined: BRAZIL. ACRE: Brasiléia, D. C. Daly et al. 11906 (NY); Cruzeiro
do Sul, J. Prado et al. 1350 (HPZ); Feijó, P. Delprete et al. 8543 (NY); Mâncio Lima,
G. T. Prance et al.12168 (NY); Manoel Urbano, D. C. Daly et al. 11226 (NY); Marechal
Thaumaturgo, D. C. Daly et al. 7336 (HPZ, NY); Porto Valter, G. T. Prance et al.
P13269 (NY), P. J. M. Maas et al.P13295 (NY), S. R. Lowrie et al. 525 (NY); Santa
Rosa, D. C. Daly et al. 9948 (NY); Tarauacá, C. Herinhaus et al. 397 (NY); Without
locality, J. Jangoux et al. 85-096 (MG, NY).
Asplenium pedicularifolium St. Hilaire, Voy. Distr. Diam. 1: 380. 1833.
Habit/Habitat: Herb; epiphyte on forest on Terra Firme.
Material examined: BRAZIL. ACRE: Mâncio Lima, D. C. Daly et al. 8926 (HPZ, NY),
G. T. Prance et al. 12246 (NY).
Distribution: Guianas and Brazil.
Asplenium serratum L., Sp. Pl. 2: 1079. 1753.
Habit/Habitat: Herb; epiphyte on forest on Terra Firme and on flooded vegetation near
river margins.
Brasiléia, D. C. Daly et al. 6771 (HPZ, NY); Bujari, D. C. Daly et al. 9328 (NY), M.
de Pardo et al. 80 (NY); Cruzeiro do Sul, D. C. Daly et al. 11717 (NY), G. T. Prance
et al. 2785 (NY), P. J. M. Maas et al. P12863 (MG, NY), J. Prado et al. 1187 (HPZ,
SP), J. Prado et al. 1197 (HPZ), J. Prado et al. 1317 (HPZ, SP), J. Prado et al. 1340
(HPZ, SP), J. Prado et al. 1382 (HPZ, SP); Mâncio Lima, J. Prado et al. 1135 (HPZ,
SP); Manoel Urbano, D. C. Daly et al. 9136 (NY), D. C. Daly et al. 11241 (NY);
Plácido de Castro, C. Figueiredo & I. Riveiro 592 (NY), L. G. Lohmann & E. C. de
Oliveira 492 (NY); Rodrigues Alves, J. Prado et al. 1237 (HPZ, SP), J. Prado et al.
1255 (HPZ, SP), J. Prado et al. 1291 (HPZ, SP); Santa Rosa, D. C. Daly et al.
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FERN GAZ. 18(5): 230-263. 2009
10053(HPZ, NY), D. C. Daly et al. 10984 (NY), D. C. Daly et al. 11106 (NY); Sena
Madureira, D. C. Daly et al. 7841 (HPZ, NY).
Distribution: S Florida, Mexico, Antilles, Mesoamerica, tropical and subtropical South
America.
Asplenium stuebelianum Hieron., Hedwigia 47: 222. 1908.
Habit/Habitat: Herb; epiphyte or terrestrial, in open forest with bamboo.
Material examined: BRAZIL. ACRE: Feijó, D. C. Daly et al. 8500 (NY); Mâncio Lima,
D. C. Daly et al. 8952 (HPZ, NY); Marechal Thaumaturgo, D. C. Daly et al. 10524
(NY), D. C. Daly et al. 10552 (NY); Rio Branco, D. C. Daly et al. 6871A (HPZ, NY).
Distribution: Tropical South America.
BLECHNACEAE
Blechnum occidentale L., Sp. Pl. 2: 1077. 1753.
Habit/Habitat: Herb; terrestrial in forest on Terra Firme.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, P. J. M. Maas et al. P13242
(NY).
Distribution: S EUA, Mexico, Antilles, Mesoamerica, tropical and subtropical South
America.
Salpichlaena hookeriana (Kuntze) Alston, Bull. Misc. Inform. Kew 1932: 312. 1932.
Spicanta hookeriana Kuntze, Revis. Gen. Pl. 2: 81. 1891.
Habit/Habitat: Herb; climbing on flooded vegetation near stream margins.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1161 (HPZ, SP);
Mâncio Lima, J. Prado et al. 1229 (HPZ, SP).
Salpichlaena volubilis (Kaulf.) J. Sm., J. Bot. (Hooker) 4: 168. 1841.
Blechnum volubile Kaulf., Enum. Filic. 159. 1824.
Habit/Habitat: Herb; climbing on flooded vegetation near stream margins.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1200 (HPZ, SP).
Distribution: Antilles, Mesoamerica, and tropical South America.
CYATHEACEAE
Alsophila cuspidata (Kunze) D. S. Conant, J. Arnold Arbor. 64: 371. 1983.
Cyathea cuspidata Kunze, Linnaea 9: 101. 1834.
Habit/Habitat: Tree-fern; forest on Terra Firme.
Material examined: BRAZIL. ACRE: Brasiléia, D. C. Daly et al. 9881 (NY), D. C. Daly
et al. 11886 (NY); Marechal Thaumaturgo, D. C. Daly et al. 7427 (HPZ, NY).
Distribution: Mesoamerica and tropical South America.
Cyathea microdonta (Desv.) Domin, Pteridophyta 263. 1929.
Polypodium microdontum Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck.
Gesammten Naturk. 5: 319. 1811.
Habit/Habitat: Tree-fern; forest near river margins.
Manoel Urbano, D. C. Daly et al. 11558 (NY).
Distribution: S Mexico, Antilles, Mesoamerica and N South America.
Cyathea pilosissima (Baker) Domin, Pteridophyta 262. 1929.
Polypodium pilosissimum Baker, Syn. Fil. ed. 2: 457. 1874.
Habit/Habitat: Tree-fern; in disturbed forest on Terra Firme.
Material examined: BRAZIL. ACRE: Mâncio Lima, D. C. Daly et al. 11603 (NY);
Rodrigues Alves, J. Prado et al. 1231 (HPZ, SP).
Distribution: Mesoamerica and N South America.
Cyathea pungens (Willd.) Domin, Pteridophyta 263. 1929.
Polypodium pungens Willd., Sp. Pl. 5: 206. 1810.
Habit/Habitat: Tree-fern; flooded vegetation near stream margins.
Material examined: BRAZIL. ACRE: Brasiléia, D. C. Daly et al. 6823 (HPZ, NY), L.
Ferreira & L. C. Ming 117 (HPZ, NY); Mâncio Lima, D. C. Daly et al. 1252 (NY);
Manoel Urbano, D. C. Daly et al. 9087 (NY); Marechal Thaumaturgo, D. C. Daly et al.
10239 (NY); Santa Rosa, D. C. Daly et al. 10165 (NY), D. C. Daly et al. 11175 (NY).
Distribution: Antilles and N South America.
DENNSTAEDTIACEAE
Dennstaedtia bipinnata (Cav.) Maxon, Proc. Biol. Soc. Wash. 51: 39. 1938.
Dicksonia bipinnata Cav., Descr. Pl. 154. 1802.
Habit/Habitat: Tree-fern; flooded vegetation near stream margins.
Material examined: BRAZIL. ACRE: Manoel Urbano, D. C. Daly et al. 11418 (NY).
Pteridium caudatum (L.) Maxon, Proc. U.S. Natl. Mus. 23: 631. 1901.
Pteris caudata L., Sp. Pl. 2: 1075. 1753.
Habit/Habitat: Herb; terrestrial in cleared land be-side tracks.
Mâncio Lima, G. T. Prance et al. 2864 (NY).
Distribution: S Florida, Mexico, Antilles, Mesoamerica and N South America.
DRYOPTERIDACEAE
Bolbitis lindigii (Mett.) Ching, Index Filic., Suppl. 3: 48. 1934.
Chrysodium lindigii Mett., Ann. Sci. Nat., Bot., sér. 5, 2: 205. 1864.
Habit/Habitat: Herb; terrestrial, in open forest on undulating terrain, except for baixios
(waterlogged low-lying areas).
Material examined: BRAZIL. ACRE: Mâncio Lima, G. T. Prance et al. 12069A (NY);
Marechal Thaumaturgo, D. C. Daly et al. 10252 (NY).
Bolbitis nicotianifolia (Sw.) Alston, Bull. Misc. Inform. Kew 1932: 310. 1932.
Acrostichum nicotianifolium Sw., Syn. Fil. 13: 199. 1806.
Habit/Habitat: Herb; terrestrial, in open forest on undulating terrain, except for baixios
(waterlogged low-lying areas).
Material examined: BRAZIL. ACRE: Bujari, D. C. Daly et al. 9477 (NY), W. R.
Anderson 12122 (NY); Cruzeiro do Sul, J. Prado et al. 1349 (HPZ); Manoel Urbano,
D. C. Daly et al. 9128 (NY), D. C. Daly et al. 11413 (NY); Porto Valter, P. J. M. Maas
et al. P13292 (NY); Sena Madureira, D. C. Daly et al. 8072 (NY).
Distribution: Antilles, Mesoamerica and N South America.
237
FERN GAZ. 18(5): 230-263. 2009
Bolbitis oligarchica (Baker) Henipp., Amer. Fern J. 65: 30. 1975.
Acrostichum oligarchicum Baker, Syn. Fil. 418. 1868.
Habit/Habitat: Herb; terrestrial, in forest.
Material examined: BRAZIL. ACRE: Mâncio Lima, M. Silveira et al. 1361 (NY).
Ctenitis refulgens (Klotzsch ex Mett.) Vareschi, Fl. Venezuela 1: 404. 1969.
Phegopteris refulgens Klotzsch ex Mett., Ann. Sci. Nat., Bot., sér. 5, 2: 240. 1864
Habit/Habitat: Herb; terrestrial, in forest.
Distribution: Mexico, Mesoamerica and N South America.
Cyclodium guianense (Klotzsch) L. D. Gómez, Phytologia 60: 371. 1986.
Aspidium guianense Klotzsch, Linnaea 20: 364. 1847.
Habit/Habitat: Herb; terrestrial, in Terra Firme forest.
Without locality, J. Jangoux et al. 85-034 (NY).
Cyclodium meniscioides (Willd.) C. Presl, Tent. Pterid. 85. 1836.
Aspidium meniscioides Willd., Sp. Pl. 5: 218. 1810.
Infraspecific: var. meniscioides
Mâncio Lima, J. Prado et al. 1226 (HPZ, SP); J. Prado et al. 1228 (HPZ, SP).
Didymochlaena truncatula (Sw.) J. Sm., J. Bot. (Hooker) 4: 196. 1841-1842 [1841].
Aspidium truncatulum Sw., J. Bot. (Schrader) 1800(2): 36. 1801.
Habit/Habitat: Herb; terrestrial in primary forest, unudulating terrain, in places
dissected by many small streams.
Material examined: BRAZIL. ACRE: Brasiléia, D. C. Daly et al. 9880 (HPZ, NY), D.
C. Daly et al. 11893 (NY); Cruzeiro do Sul, G. T. Prance et al. 12087 (MG, NY), G. T.
Prance et al. 12377 (NY); Mâncio Lima, D. G. Campbell et al. 8913 (NY).
Distribution: S Mexico, Antilles, Mesoamerica and N South America. Tropical Asia and
Africa.
Dryopteris patula (Sw.) Underw., Native Ferns ed. 4 117. 1893.
Aspidium patulum Sw., Kongl. Vetensk. Acad. Handl. 1817(1): 64. 1817.
Habit/Habitat: Herb; terrestrial in moist forest on Terra Firme, canopy discontinuous.
Material examined: BRAZIL. ACRE: Plácido de Castro, C. Figueiredo & I. Riveiro 560
(NY); Sena Madureira, D. C. Daly et al. 8117 (NY).
Distribution: S Mexico, Antilles, Mesoamerica and tropical South America.
Elaphoglossum discolor (Kuhn) C. Chr., Index Filic. 306. 1905.
Acrostichum discolor Kuhn, Linnaea 36: 53. 1869.
Habit/Habitat: Herb; terrestrial or epipetric, in Campina.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, D. C. Daly et al. 10607 (NY),
J. Prado et al. 1159 (HPZ, SP). J. Prado et al. 1170 (HPZ, SP), J. Prado et al. 1188
(HPZ, SP), J. Prado et al. 1284 (HPZ, SP), J. Prado et al. 1322 (HPZ, SP), J. Prado et
al. 1324 (HPZ, SP), J. Prado et al. 1329 (HPZ), J. Prado et al. 1388 (HPZ, SP); Mâncio
Lima, J. Prado et al. 1223 (HPZ, SP).
Elaphoglossum flaccidum (Fée) T. Moore, Index Filic. 356. 1862.
Acrostichum flaccidum Fée, Mem. Foug. 2: 35, tab. 7, fig. 2. 1845.
Habit/Habitat: Herb; epiphyte, in Terra Firme forest.
Material examined: BRAZIL. ACRE: Rodrigues Alves, J. Prado et al. 1266 (HPZ, SP);
Xapuri, Silva et al.186 (CEN, SP).
Elaphoglossum glabellum J. Sm., London J. Bot. 1: 197. 1842.
Distribution: S Florida, Mexico, Antilles, Mesoamerica, and N South America.
Elaphoglossum luridum (Fée) H. Christ, Neue Denkschr. Allg. Schweiz. Ges.
Gesammten Naturwiss. 36(1): 33. 1899.
Acrostichum luridum Fée, Mem. Foug. 2: 35, tab. 19, fig. 1. 1845.
J. Prado et al. 1174 (HPZ, SP), J. Prado et al. 1198 (HPZ, SP), J. Prado et al. 1310
(HPZ, SP), J. Prado et al. 1327 (HPZ, SP); Mâncio Lima, J. Prado et al. 1219 (HPZ,
SP), J. Prado et al. 1220 (HPZ, SP); Rodrigues Alves, J. Prado et al. 1238 (HPZ, SP),
(HPZ, SP).
Elaphoglossum plumosum (Fée) T. Moore, Index Filic. 364. 1862.
Acrostichum plumosum Fée, Mem. Foug. 54, tab. 20, fig. 1. 1845.
J. Prado et al. 1285 (HPZ); Mâncio Lima, J. Prado et al. 1221 (HPZ, SP), J. Prado et
al. 1390 (HPZ, SP).
Elaphoglossum raywaense (Jenm.) Aslton, Bol. Soc. Brot., ser. 2, 32: 24. 1958.
Acrostichum raywaense Jenm., Ferns Brit. W. Ind. 341. 1909.
Mâncio Lima, D. C. Daly et al. 8948 (HPZ, NY).
Elaphoglossum styriacum Mickel, Brittonia 39: 326, fig. 4I-K. 1987.
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FERN GAZ. 18(5): 230-263. 2009
Elaphoglossum tantalinum Mickel, Brittonia 39: 326, fig. 4I-K. 1987.
J. Prado et al. 1321 (HPZ, SP), J. Prado et al. 1384 (HPZ, SP).
Lastreopsis effusa (Sw.) Tindale, Victoria Naturalist 73: 184. 1957.
Polypodium effusum Sw., Prodr. 134. 1788.
Habit/Habitat: Herb; terrestrial, forested slope between Baixio and higher terrace.
Distribution: S Mexico, Antilles, Mesoamerica, and tropical South America.
Lomagramma guianensis (Aubl.) Ching, Amer. Fern J. 22: 17. 1932.
Polypodium guianense Aubl., Hist. Pl. Guiane 962. 1775.
Habit/Habitat: Herb; hemiepiphyte, in Terra Firme forest.
C. Daly et al. 6786 (NY), D. C. Daly et al. 11882 (NY); M. Silveira et al. 1662 (NY);
Cruzeiro do Sul, J. Prado et al. 1292 (HPZ, SP); Mâncio Lima, D. C. Daly et al. 11623
(NY); Marechal Thaumaturgo, D. C. Daly et al. 7430 (HPZ, NY); Rodrigues Alves, J.
Prado et al. 1252 (HPZ, SP), J. Prado et al. 1278 (HPZ, SP); Xapuri, L. G. Lohmann
& E. C. de Oliveira 590 (NY).
Distribution: Greater Antilles (except Jamaica) and tropical South America.
Polybotrya caudata Kunze, Linnaea 9: 23. 1834.
Mâncio Lima, J. Prado et al. 1143 (HPZ, SP), J. Prado et al. 1145 (HPZ, SP) G. T.
Prance et al. 12069 (NY), M. Silveira et al. 1662 (NY).
Distribution: S Mexico, Mesoamerica and N South America.
Polybotrya pubens Martius, Icon. Pl. Crypt. 87, tab. 25. 1834.
Rumohra adiantiformis (G. Forst) Ching, Sinensia 5: 70. 1934.
Polypodium adiantiforme G. Forst, Prodr. 82. 1786.
Distribution: Bermuda, Antilles, N South America. Africa, Madagascar, New Zealand
and Australia.
GLEICHENIACEAE
Gleichenella pectinata (Willd.) Ching, Sunyatsenia 5: 276. 1940.
Mertensia pectinata Willd., Kongl. Vetensk. Acad. Nya Handl. 25: 168, tab. 4. 1804.
Habit/Habitat: Herb; terrestrial along forest margins.
Mâncio Lima, G. T. Prance et al. 12119 (MG, NY), M. Silveira et al. 1348 (HPZ, NY).
Sticherus remotus (Kaul.) Chrysler, Amer. J. Bot. 31: 483. 1944.
Mertensia remota Kaulf., Enum. Filic. 39. 1824
Habit/Habitat: Herb; terrestrial along forest margins.
HYMENOPHYLLACEAE
Didymoglossum krausii (Hook. & Grev.) C. Presl, Hymenophyllaceae 115. 1843.
Trichomanes krausii Hook. & Grev., Icon. Filic. 2: pl. 149. 1830
Habit/Habitat: Herb; epiphyte in forest.
Material examined: BRAZIL. ACRE: Manoel Urbano, D. C. Daly et al. 9115 (HPZ,
NY).
Distribution: Florida, Mexico, Antilles, Mesoamerica, tropical South America.
Didymoglossum ovale E. Fourn., Acta Bot. Neerl. 11: 296. 1962.
Habit/Habitat: Herb; epiphyte along river margins.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1288 (HPZ);
Mâncio Lima, J. Prado et al. 1230 (HPZ, SP).
Didymoglossum punctatum (Poir.) Desv., Mém. Soc. Linn. Paris. 6: 330. 1827.
Trichomanes punctatum Poir., Encycl. 8: 64. 1808.
Habit/Habitat: Herb; epiphyte.
Material examined: BRAZIL. ACRE: Marechal Thaumaturgo, D. C. Daly et al. 10330
(NY).
Hymenophyllum angustum Bosch, Ned. Kruidk. Arch. 5(2): 183. 1861.
Habit/Habitat: Herb; epiphyte in waterlogged low-lying areas.
Distribution: Amazonian Brazil.
Trichomanes polyanthos Sw., Prodr. 137. 1788.
J. Prado et al. 1389 (HPZ, SP), P. J. M. Maas et al. 12722 (NY); Mâncio Lima, J.
Prado et al. 1222 (HPZ, SP).
Distribution: Mexico, Mesoamerica and tropical South America.
Trichomanes ankersii C. Parker ex Hook. & Grev., Icon. Filic. 2(11): tab. 201. 1831.
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FERN GAZ. 18(5): 230-263. 2009
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, G. T. Prance et al. 11854 (MG,
NY), J. Prado et al. 1150 (HPZ, SP), J. Prado et al. 1158 (HPZ, SP), J. Prado et al.
1168 (HPZ, SP), J. Prado et al. 1214 (HPZ, SP), J. Prado et al. 1312 (HPZ, SP), J.
Prado et al. 1387 (HPZ, SP); Rodrigues Alves, J. Prado et al. 1236 (HPZ, SP), J. Prado
et al. 1257 (HPZ, SP).
Trichomanes bicorne Hook., Icon. Plan. 9: tab. 892. 1854.
(HPZ, SP), J. Prado et al. 1323 (HPZ, SP), J. Prado et al. 1271 (HPZ, SP).
Trichomanes crispum L., Sp. Pl. 2: 1097. 1753.
J. Prado et al. 1380 (HPZ, SP).
Distribution: S Mexico, Antilles, Mesoamerica, and N South America
Trichomanes elegans Rich., Actes Soc. Hist. Nat. Paris 1: 114. 1792.
Habit/Habitat: Herb; terrestrial, in forest near stream margins.
NY), J. Prado et al. 1212 (HPZ, SP); Mâncio Lima, M. Silveira et al. 1247 (NY), M.
Silveira et al. 1250 (NY); Rodrigues Alves, J. Prado et al. 1364 (HPZ, SP).
Distribution: Antilles, Mesoamerica, tropical and subtropical South America.
Trichomanes hostmannianum (Klotzsch) Kunze, Bot. Zeitung (Berlin) 5: 352. 1847.
Neurophyllum hostmannianum Klotzsch, Linnaea 18: 532. 1844.
Habit/Habitat: Herb; terrestrial, in forest near stream margins on wet soils.
L. R. Marinho 217 (NY); Rodrigues Alves, J. Prado et al. 1272 (HPZ, SP).
Trichomanes macilentum v. d. Bosch, Ned. Kruidk. Arch. 5(2): 146. 1861.
Habit/Habitat: Herb; terrestrial in waterlogged low-lying areas.
Trichomanes martiusii C. Presl, Hymenophyllaceae 15, 36. 1843.
Habit/Habitat: Herb; terrestrial in forest on Terra Forme.
(HPZ, SP), N. A. Rosa 657 (NY); Mâncio Lima, J. Prado et al. 1217 (HPZ, SP).
Trichomanes pilosum Raddi, Opusc Sci. 3: 296. 1819.
Distribution: Bolivia, Paraguay and Brazil (south of the Amazon Basin).
Trichomanes pinnatum Hedw., s.l., Fil. Gen. Sp. tab. 4, fig. 1. 1799.
Habit/Habitat: Herb; terrestrial in forest on Terra Firme, on clay and sand soils.
Cruzeiro do Sul, J. Prado et al. 1183 (HPZ), J. Prado et al. 1311 (HPZ, SP), D. C. Daly
et al. 11640 (NY); Mâncio Lima, D. C. Daly et al.11589 (HPZ, NY), G. T. Prance et al.
12178 (MG, NY); Porto Valter, D. C. Daly et al. 7576 (HPZ, NY), D. C. Daly et al.
11739 (NY), P. J. M. Maas et al.13010 (MG, NY); Rodrigues Alves, J. Prado et al.
1244 (HPZ); Sena Madureira, G. T. Prance et al. 7730 (NY), G. T. Prance et al. 7948
(NY).
Obs.: the material studied for this taxon cited above probably belong to more than two
or three species, but more studies are necessary to clarify this situation.
Trichomanes tanaicum Hook. ex J. W. Sturm., Fl. Bras. 1(2): 260. 1859.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1209 (HPZ);
Rodrigues Alves, J. Prado et al. 1254 (HPZ, SP), J. Prado et al. 1258 (HPZ).
Trichomanes trollii Bergdolt, Flora 127: 256, 264, fig. 3. 1933.
Habit/Habitat: Herb; terrestrial, in forest on Terra Firme.
Material examined: BRAZIL. ACRE: Rodrigues Alves, J. Prado et al. 1366 (HPZ, SP).
Vandenboschia collariata (Bosch) Ebihara & K. Iwats., Blumea 51: 242. 2006.
Trichomanes collariatum Bosch, Ned. Kruidk. Arch. 4: 368. 1859 [1858].
Material examined: BRAZIL. ACRE: Without locality, J. Jangoux et al. 85-053 (MG,
NY), J. Jangoux et al. 850-73 (MG, NY).
LINDSAEACEAE
Lindsaea divaricata Klotzsch, Linnaea 18: 547. 1844.
Habit/Habitat: Herb; terrestrial in forest on Terra Firme, on sandy soils.
J. Prado et al. 1177 (HPZ, SP), J. Prado et al. 1270 (HPZ, SP), T. B. Croat & A. Rosas
Jr. 62671 (NY); Rodrigues Alves, J. Prado et al. 1359 (HPZ, SP).
Lindsaea hemiglossa Kramer, Acta Bot. Neerl. 6(2): 257. 1957.
Habit/Habitat: Herb; terrestrial in forest on Terra Firme, on slopes.
Material examined: BRAZIL. ACRE: Mâncio Lima, G. T. Prance et al. 12142 (NY).
243
FERN GAZ. 18(5): 230-263. 2009
Lindsaea lancea (L.) Bedd., Suppl. Ferns S. Ind. 6. 1876.
Adiantum lanceum L., Sp. Pl. ed. 2 1557. 1763.
Infraspecific: var. lancea
Infraspecific: var. falcata (Dryand.) Rosenst., Hedwigia 46: 79. 1906.
(HPZ, SP).
Lindsaea schomburgkii Klotzsch, Linnaea 18: 545. 1844.
Lindsaea sp.
Habit/Habitat: Herb; terrestrial in forest on Terra Firme, on clay soils.
Distribution: Peru and Brazil.
Lindsaea stricta (Sw.) Dryand. Trans. Linn. Soc. London 3: 42. 1797.
Adiantum strictum Sw., Prodr.: 135. 1788.
Infraspecific: var. stricta
Habit/Habitat: Herb; terrestrial, in Campina.
Distribution: Widespread in the Neotropics.
Lindsaea ulei Hieron., Hedwigia 44: 365. 1905.
J. Prado et al. 1356 (HPZ, SP); Rodrigues Alves, J. Prado et al. 1232 (HPZ, SP), J.
Prado et al. 1243 (HPZ, SP), J. Prado et al. 1367 (HPZ), J. Prado et al. 1369 (HPZ,
SP).
LOMARIOPSIDACEAE
Cyclopeltis semicordata (Sw.) J. Sm., Bot. Mag. 72: 36. 1846.
Polypodium semicordatum Sw., Prodr. 132. 1788.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, D. C. Daly et al. 11721 (NY);
Manoel Urbano, D. C. Daly et al. 11427 (NY), M. Silveira et al. 1574 (NY); Marechal
Thaumaturgo, D. C. Daly et al. 7342 (HPZ, NY), D. C. Daly et al. 10473 (NY), D. C.
Daly et al. 10810 (NY); Porto Acre, C. A. Cid & B. Nelson 2877 (NY), P. J. M. Maas
et al. 13247 (MG, NY); Quixadá, C. A Cid & A. Rosas 2961 (NY); Reserva Extrativista
Chico Mendes, Colocação Simitumba, A. R. S. Oliveira et al. 131 (HPZ, NY); Santa
Rosa, D. C. Daly et al. 11367 (NY); Santa Rosa, D. C. Daly et al. 10978 (NY); Sena
Madureira, D. C. Daly et al. 7857 (HPZ, NY); Rio Branco, D. C. Daly et al. 6881 (HPZ,
NY); Tarauacá, G. T. Prance et al. 7516 (NY), D. C. Daly et al. 8667 (HPZ, NY), D. C.
Daly et al. 8799 (HPZ, NY); Xapuri, D. C. Daly et al. 8420 (NY); Without locality, J.
Jangoux et al. 85-074 (MG, NY).
Lomariopsis japurensis (Mart.) J. Sm., Hist. Fil. 140. 1875.
Acrostichum japurense Mart., Icon. Pl. Crypt. 86, tab. 24. 1834.
Cruzeiro do Sul, P. J. M. Maas et al. P12886 (NY); Mâncio Lima, G. T. Prance et al.
12055 (NY); Marechal Thaumaturgo, L. G. Lohmann et al. 480 (NY); Santa Rosa, D.
C. Daly et al. 9945 (NY).
Lomariopsis nigropaleata Holttum, Bull. Misc. Inform. Kew 1939: 618. 1940.
Material examined: BRAZIL. ACRE: Brasiléia, S. R. Lowrie et al. 726 (NY); Cruzeiro
do Sul, J. Prado et al. 1202 (HPZ, SP), J. Prado et al. 1211 (HPZ, SP), J. Prado et al.
1352 (HPZ, SP); Mâncio Lima, D. C. Daly et al. 11568 (NY); Marechal Thaumaturgo,
L. C. Ming & L. A. Ferreira 324 (NY).
Lomariopsis prieuriana Fée, Mem. Foug. 2: 66, pl. 25, fig. 1. 1845.
Material examined: BRAZIL. ACRE: Mâncio Lima, J. Prado et al. 1137 (HPZ, SP);
Nephrolepis biserrata (Sw.) Schott, Gen. Fil. pl. 3. 1834.
Aspidium biserratum Sw., J. Bot. (Schrader) 1800(2): 32. 1801.
Habit/Habitat: Herb; epiphyte in forest on Terra Firme near stream margins.
J. Prado et al. 1180 (HPZ, SP), J. Prado et al. 1199 (HPZ, SP); Mâncio Lima, J. Prado
Nephrolepis brownii (Desv.) Hovenkamp & Miyam., Blumea 50: 293. 2005.
Nephrodium brownii Desv., Mem. Soc. Linn. Paris 6: 252. 1827.
Habit/Habitat: Herb; epiphyte in forest on Terra Firme near stream margins.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, P. Delprete et al. 8111 (NY);
Mâncio Lima, D. C. Daly et al. 11595 (NY).
Distribution: Florida, S Mexico, Antilles, Mesoamerica, tropical South America,
Paleotropics.
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FERN GAZ. 18(5): 230-263. 2009
Nephrolepis pendula (Raddi) J. Sm., J. Bot. (Hooker) 4: 197. 1841-1842 [1841].
Aspidium pendulum Raddi Opusc. Sci. 3: 289. 1819.
Material examined: BRAZIL. ACRE: Tarauacá, G. T. Prance et al. 7296 (NY).
Nephrolepis rivularis (Vahl) Mett. ex Krug, Bot. Jahrb. Syst. 24: 122. 1897.
Polypodium rivulare Vahl, Eclog. Amer. 3: 51. 1807.
Habit/Habitat: Herb; epiphyte on Várzea forest.
J. Prado et al. 1325 (HPZ, SP); Mâncio Lima, G. T. Prance et al. 12179 (MG, NY).
LYCOPODIACEAE
Huperzia dichotoma (Jacq.) Trevisan, Atti Soc. Ital. Sci. Nat. 17: 248. 1874.
Lycopodium dichotomum Jacq., Enum. Stirp. Vindob. 314. 1762.
Material examined: BRAZIL. ACRE: Without locality, M. Silveira s.n. (HPZ).
Distribution: Florida, Mexico, Antilles, Mesoamerica and N South America.
Lycopodiella cernua (L.) Pic.Serm., Webbia 23: 166. 1968.
Lycopodium cernuum L., Sp. Pl. 2: 1103. 1753
Habit/Habitat: Herb; terrestrial on sandy soils in open places.
J. Prado et al. 1191 (HPZ, SP), J. Prado et al. 1331 (HPZ), A. Rosas et al. 313 (NY),
D. C. Daly et al. 10599 (NY), D. C. Daly et al. 10683 (NY), G. T. Prance et al. 12525
(MG, NY), L. R. Marinho 64 (NY), L. R. Marinho 212 (NY), P. Delprete et al. 8063
(NY).
Distribution: Pantropical.
LYGODIACEAE
Lygodium venustum Sw., J. Bot. (Schrader) 1801(2): 303. 1803.
Habit/Habitat: Herb; vine-like, in forest on Terra Firme.
Material examined: BRAZIL. ACRE: Manoel Urbano, M. Silveira et al. 1543 (HPZ,
NY); Rio Branco, L. Coêlho & A. Rosas 1925 (NY); Tarauacá, C. Ehringhaus et al. 346
(NY).
Lygodium volubile Sw., J. Bot. (Schrader) 1801(2): 303. 1803.
Habit/Habitat: Herb; vine-like, in forest on Terra Firme.
MARATTIACEAE
Danaea leprieurii Kunze, Mém. Acad. Roy. Sci. (Turin) 5(1790-1791): 429 tab. 9.
1793.
Habit/Habitat: Herb; terrestrial in forest on Terra Firme, near stream margins
Distribution: S Mexico, Antilles, Mesoamerica, and N South America.
Danaea nodosa (L.) Sm., Mem. Acad. Roy. Sci. (Turin) 5(1790-1791): 420, tab. 9, fig.
11. 1793.
Achrostichum nodosum L., Sp. Pl. 2: 1070. 1753.
C. Daly et al. 9884 (HPZ, NY); Feijó, P. Delprete et al. 8301A & B (NY); Sena
Madureira, D. C. Daly et al. 8137 (HPZ, NY); Without locality, J. Jangoux et al. 85071 (NY).
Danaea trifoliata Reichenb. ex Kunze, Analecta Pteridogr. 4, tab. 2. 1837.
Habit/Habitat: Herb; terrestrial in forest on Terra Firme, near stream margins.
METAXYACEAE
Metaxya rostrata (Kunth) C. Presl, Tent. Pterid. 60, tab. 1, fig. 5. 1836.
Aspidium rostratum Kuhn, Nov. Gen. Sp. (quarto ed.) 1: 12. 1815 [1816].
Habit/Habitat: Herb; terrestrial in forest on Terra Firme, near stream margins.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, D. C. Daly et al. 11705 (NY), J.
Prado et al. 1186 (HPZ, SP), J. Prado et al. 1301 (HPZ, SP), J. Prado et al. 1320 (HPZ,
SP); Mâncio Lima, D. C. Daly et al. 11624 (NY), M. Silveira et al. 1365 (NY);
Marechal Thaumaturgo, D. C. Daly et al. 10526 (NY); Porto Valter, D. C. Daly et al.
7577 (NY), P. J. M. Maas et al. P13150 (NY); Without locality, J. Jangoux et al. 85043 (NY), J. Jangoux et al. 85-108 (NY).
Distribution: S Mexico, Lesser Antilles, Mesoamerica and N South America.
POLYPODIACEAE
Campyloneurum abruptum (Lindm.) B. Léon, Fieldiana, Bot., n.s. 32: 172. 1993.
Polypodium repens var. abruptum Lindm., Ark. Bot. 1: 245. 1903.
Habit/Habitat: Herb; epiphyte, Várzea forest.
Material examined: BRAZIL. ACRE: Manoel Urbano, D. C. Daly et al. 11225 (NY),
M. Silveira et al. 1582 (NY).
Campyloneurum angustifolium (Sw.) Fée, Mem. Foug. 5: 257. 1852.
Polypodium angustifolium Sw., Prodr. 130. 1788.
Material examined: BRAZIL. ACRE: Bujari, M. de Pardo et al. 93 (NY); Cruzeiro do
Sul, G. T. Prance et al. 12617 (NY), L. R. Marinho 163 (NY); Feijó, P. Delprete et al.
8311 (NY); Marechal Thaumaturgo, D. C. Daly et al. 10936 (NY); Porto Valter, P. J. M.
Maas et al. 12931 (NY); Tarauacá, D. C. Daly et al. 8628 (HPZ, NY), G. T. Prance et
al. 7384 (NY), G. T. Prance et al. 7249 (NY), M. Silveira et al. 1146 (NY).
Distribution: S Mexico, Lesser Antilles, Mesoamerica and tropical South America.
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FERN GAZ. 18(5): 230-263. 2009
Campyloneurum aphanophlebium (Kunze) T. Moore, Index Filic. 223. 1861.
Polypodium aphanophlebium Kunze, Bot. Zeitung (Berlin) 3(17): 288. 1845.
Habit/Habitat: Herb; epiphyte, forest in broad, meandering stream vally in steeply hilly
terrain.
Material examined: BRAZIL. ACRE: Brasiléia, G. P. da Silva et al. 173 (NY); Mâncio
Lima, D. C. Daly et al. 11601 (NY); Marechal Thaumaturgo, D. C. Daly et al. 7496
(HPZ, NY), D. C. Daly et al. 10255 (NY); Porto Valter, P. J. M. Maas et al. P13181
(NY); Rodrigues Alves, J. Prado et al. 1261 (HPZ, SP); Santa Rosa, D. C. Daly et al.
10173 (NY); Tarauacá, D. C. Daly et al. 8630 (HPZ, NY), G. T. Prance et al. 7267
(NY).
Campyloneurum fuscosquamatum Lellinger, Amer. Fern J. 78: 21, fig. 4, 10. 1988.
Habit/Habitat: Herb; epiphyte on Terra Firme forest and Várzea forest.
C. Daly et al. 11894 (NY); Bujari, W. R. Anderson 12123 (NY); Feijó, P. Delprete et al.
8580 (NY); Mâncio Lima, G. T. Prance et al. 12052 (MG, NY), G. T. Prance et al.
12052A (NY); Quixadá, E. Forero et al. 6388 (MG, NY); Santa Rosa, D. C. Daly et al.
10151 (NY), D. C. Daly et al. 10994 (NY); Sena Madureira, D. C. Daly et al. 7856
(HPZ, NY), G. T. Prance et al. 7694 (NY); Tarauacá, D. C. Daly et al. 8620 (HPZ, NY),
D. C. Daly et al. 8760 (HPZ, NY); Without locality, J. Jangoux et al. 85-039 (MG, NY).
Campyloneurum ophiocaulon (Klotzsch) Fée, Mem. Foug. 258. 1852.
Polypodium ophiocaulon Klotzsch, Linnaea 20: 401. 1847.
Habit/Habitat: unknown.
Material examined: BRAZIL. ACRE: Bujari, M. de Pardo et al. 94 (NY).
Campyloneurum phyllitidis (L.) C. Presl, Tent. Pterid. 190. 1836.
Polypodium phyllitidis L., Sp. Pl. 2: 1083. 1753.
do Sul, D. C. Daly et al. 11716 (NY), J. Prado et al. 1315 (HPZ, SP), J. Prado et al.
1377 (HPZ, SP), G. T. Prance et al. 2783 (NY), G. T. Prance et al. 2818 (NY), G. T.
Prance et al. 12530 (NY), G. T. Prance et al. 12622 (MG, NY); Feijó, G. T. Prance et
al. 7317 (NY); Mâncio Lima, J. Prado et al. 1136 (HPZ, SP), J. Prado et al. 1142 (HPZ,
SP), J. Prado et al. 1215 (HPZ, SP); Manoel Urbano, D. C. Daly et al. 9129 (NY);
Marechal Thaumaturgo, D. C. Daly et al. 10528 (NY); Porto Acre, A. R. S. Oliveira et
al. 763 (NY); Porto Valter, P. J. M. Maas et al. P12932 (MG, NY); Rodrigues Alves, J.
Prado et al. 1264 (HPZ, SP), J. Prado et al. 1280 (HPZ, SP); Sena Madureira, D. C.
Daly et al. 7906 (HPZ, NY); Without locality, J. Jangoux et al. 85-098 (NY).
America.
Campyloneurum repens (Aubl.) C. Presl, Tent. Pterid. 190. 1836.
Polypodium repens Aubl., Hist. Pl. Guiane 2: 962. 1775.
Campyloneurum sphenodes (Klotzsch) Fée, Mem. Foug. 5: 258. 1852.
Polypodium sphenodes Klotzsch, Linnaea 20: 402. 1847.
Mâncio Lima, J. Prado et al. 1141 (HPZ, SP); Marechal Thaumaturgo, D. C. Daly et
al. 10345 (NY); Porto Valter, D. C. Daly et al. 7556 (NY).
Dicranoglossum desvauxii (Klotzsch) Proctor, Rhodora 63: 35. 1961.
Taenitis desvauxii Klotzsch, Linnaea 20: 431. 1847.
Material examined: BRAZIL. ACRE: Bujari, D. C. Daly et al. 9466 (NY); Cruzeiro do
Sul, J. Prado et al. 1213 (HPZ), J. Prado et al. 1393 (HPZ, SP); Mâncio Lima, G. T.
Prance et al. 12247 (NY); Rodrigues Alves, J. Prado et al. 1239 (HPZ, SP), J. Prado
et al. 1251 (HPZ); Without locality, J. Jangoux et al. 85-113 (NY).
Distribution: Lesser Antilles, tropical South America.
Microgramma baldwinii Brade, Arch. Jard. Bot. Rio de Janeiro 18: 30, tab. 1. 1965.
Habit/Habitat: Herb; epiphyte in forest on Terra Firme, near stream margins.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, N. A. Rosa 716 (NY), G. T.
Prance et al. 12515 (MG, NY), J. Prado et al. 1353 (HPZ, SP); Mâncio Lima, M.
Silveira et al. 1366 (NY); Rodrigues Alves, J. Prado et al. 1235 (HPZ), J. Prado et al.
1267 (HPZ, SP), J. Prado et al. 1282 (HPZ), J. Prado et al. 1368 (HPZ).
Microgramma bifrons (Hook.) Lellinger., Amer. Fern J. 67: 59. 1977.
Polypodium bifrons Hook., Fil. Exot. 1859.
Habit/Habitat: Herb; epiphyte, in open forest.
Material examined: BRAZIL. ACRE: Mâncio Lima, D. C. Daly et al. 8998 (HPZ, NY).
Obs: This species has long been called Solanopteris bifrons (Hook.) Copel.
Microgramma dictyophylla (Kunze ex Mett.) de la Sota, Novon 17: 27. 2007.
Polypodium dictyophyllum Kunze ex Mett.., Abh. Senckenberg Naturf. Ges. 2: 96.
1856.
Habit/Habitat: Herb; epiphyte, in forest on Terra Firme.
D. G. Campbell et al. 8932 (NY); Tarauacá, G. T. Prance et al. 7258 (MG, NY); Without
locality, J. Jangoux et al. 85-094 (MG, NY).
Obs: This species has long been called Microgramma fuscopunctata (Hook.) Vareschi.
Microgramma megalophylla (Desv.) de la Sota, Bol. Soc. Argent. Bot. 10: 158. 1963.
Polypodium megalophyllum Desv., Mem. Soc. Linn. Paris 6: 227. 1827.
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FERN GAZ. 18(5): 230-263. 2009
Habit/Habitat: Herb; epiphyte, in forest on Terra Firme.
NY), J. Prado et al. 1196 (HPZ, SP), J. Pruski et al. 3465 (NY), P. J. M. Maas et al.
P12725 (MG, NY); Mâncio Lima, M. Silveira et al. 1307 (NY); Rodrigues Alves, J.
Prado et al. 1265 (HPZ, SP); Without locality, J. Jangoux et al. 85-015 (NY).
Microgramma percussa (Cav.) de la Sota, Physis (Buenos Aires) 44(106C): 28. 1986.
Polypodium percussum Cav., Descr. Pl. 243. 1802.
Habit/Habitat: Herb; epiphyte, Terra Firme forest on gently undulating terrain,
undistributed but canopy open.
Material examined: BRAZIL. ACRE: Bujari, M. de Pardo et al. 92 (HPZ, NY);
Cruzeiro do Sul, D. C. Daly et al. 11714 (NY), D. C. Daly et al. 11657 (NY), J. Prado
et al. 1346 (HPZ), J. Prado et al. 1375 (HPZ), P. J. M. Maas et al. P13133B (NY);
Feijó, P. Delprete et al. 8310 (NY), P. Delprete et al. 8488 (NY); Manoel Urbano, D.
C. Daly et al. 11218 (NY), D. C. Daly et al. 11420 (NY); Marechal Thaumaturgo, D. C.
Daly et al. 10297 (NY), L. G. Lohmann et al. 458 (NY); Sena Madureira, M. Silveira et
al. 616 (HPZ, NY); Tarauacá, C. Ehringhaus et al. 382 (NY), G. T. Prance et al. 7380
(NY).
Distribution: S Mexico, Mesoamerica and tropical South America.
Microgramma persicariifolia (Schrad.) C. Presl, Tent. Pterid. 214. 1836.
Polypodium persicariifolium Schrad., Gott. gel. Anz. 1824: 867. 1824.
Habit/Habitat: Herb; epiphyte in Várzea forest.
Material examined: BRAZIL. ACRE: Bujari, D. C. Daly et al. 8497 (HPZ, NY);
Cruzeiro do Sul, D. C. Daly et al. 7300 (HPZ, NY); Mâncio Lima, G. T. Prance et al.
12061 (NY); Santa Rosa, D. C. Daly et al. 10103 (NY), D. C. Daly et al. 11340 (NY);
Sena Madureira, A. R. S. Oliveira et al. 622 (NY), M. de Pardo et al. 148 (HPZ, NY);
Xapuri, L. G. Lohmann et al. 600 (NY); Without locality, D. G. da Silva et al. 57 (NY).
Distribution: Costa Rica, Panama and tropical South America.
Microgramma reptans (Cav.) A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40(8): 230.
1975.
Acrostichum reptans Cav., Anales Hist. Nat. 1: 104. 1799.
Sul, D. C. Daly et al. 11813 (NY); Manoel Urbano, D. C. Daly et al. 11252 (NY);
Marechal Thaumaturgo, D. C. Daly et al. 10571 (NY); Porto Acre, A. R. S. Oliveira et
al. 744 (NY); Porto Valter, P. J. M. Maas et al. 13057 (NY); Sena Madureira, D. C. Daly
et al. 8081 (HPZ, NY).
Distribution: Mexico, Cuba, Mesoamerica and tropical South America.
Microgramma tecta (Kaulf.) Alston, J. Wash. Acad. Sci. 48: 232. 1958.
Polypodium tectum Kaulf., Enum. Fil. 87. 1824.
Habit/Habitat: Herb; epiphyte in forest near stream margins.
J. Prado et al. 1372 (HPZ, SP); Rodrigues Alves, J. Prado et al. 1274 (HPZ, SP).
Microgramma thurnii (Baker) R. M. Tryon & Stolze, Fieldiana, Bot., n.s. 32: 156.
1993.
Polypodium thurnii Baker, Ann. Bot. (Oxford) 5: 476. 1891.
(HPZ, SP), J. Prado et al. 1383 (HPZ, SP), P. Delprete et al. 8092 (NY); Mâncio Lima,
J. Prado et al. 1218 (HPZ, SP); Porto Valter, D. C. Daly et al. 11789 (NY), P. J. M.
Maas et al. 13133A (NY); Rodrigues Alves, J. Prado et al. 1250 (HPZ); Without
locality, J. Jangoux et al. 85-035 (NY).
Niphidium crassifolium (L.) Lellinger, Amer. Fern J. 62: 106. 1972.
Polypodium crassifolium L., Sp. Pl. 2: 1083. 1753.
Material examined: BRAZIL. ACRE: Brasiléia, D. C. Daly et al. 11889 (NY); Marechal
Thaumaturgo, D. C. Daly et al. 10246 (NY); Rodrigues Alves, J. Prado et al. 1263
(HPZ, SP), J. Prado et al. 1283 (HPZ, SP).
Pecluma hygrometrica (Splitg.) M. G. Price, Amer. Fern J. 73: 115. 1983.
Polypodium hygrometricum Splitg., Tijdschr. Natuurl. Gesch. Physiol. 7: 409. 1840.
Habit/Habitat: Herb; epiphyte in open forest on Terra Firme with palms, undulating
terrain.
Santa Rosa, D. C. Daly et al. 10176 (NY), D. C. Daly et al. 10988 (NY).
Pecluma pectinata (L.) M.G. Price, Amer. Fern J. 73: 115. 1983.
Polypodium pectinatum L., Sp. Pl. 2: 1085-1086. 1753.
Habit/Habitat: Herb; epiphyte in Várzea forest.
Sul, G.T. Prance et al. 2784 (NY).
Distribution: Antilles, Mesoamerica and tropical South America.
Pecluma pilosa (A. M. Evans) M. Kessler & A. R. Sm, Candollea 60: 281. 2005.
Polypodium ptilodon Kunze var. pilosum A. M. Evans, Ann. Missouri Bot. Gard. 55:
259, fig. 20. 1969.
G. T. Prance et al. 12282 (NY).
Pecluma plumula (Willd.) M. G. Price, Amer. Fern J. 73: 115. 1983.
Polypodium plumula Willd., Sp. Pl. 5: 178. 1810.
Habit/Habitat: Herb; epiphyte, in riverside vegetation.
Marechal Thaumaturgo, D. C. Daly et al. 10530 (NY); Quixadá, E. Forero et al. 6347
251
FERN GAZ. 18(5): 230-263. 2009
(NY); Rio Branco, D. C. Daly et al. 6923 (NY); Santa Rosa, D. C. Daly et al. 11057
(NY); Sena Madureira, D. C. Daly et al. 7907 (NY); Tarauacá, G. T. Prance et al. 7248
(NY); G. T. Prance et al. 7385 (MG, NY).
Phlebodium decumanum (Willd.) J. Sm., J. Bot. (Hooker) 4: 59. 1841.
Polypodium decumanum Willd., Sp. Pl. 5: 170. 1810.
Porto Valter, P. J. M. Maas et al. P13299 (NY); Rio Branco, C. A. Cid & A. Souza 2986
(NY), D. C. Daly et al. 6871 (HPZ, NY); Santa Rosa, D. C. Daly et al. 10119 (NY), D.
C. Daly et al. 11272 (NY); Sena Madureira, D. C. Daly et al. 8096 (HPZ, NY);
Tarauacá, P. Delprete et al. 8257 (NY); Xapuri, D. C. Daly et al. 8414 (NY).
Distribution: S Florida, Antilles, Mesoamrica, tropical and subtropical South America.
Pleopeltis bombycina (Maxon) A. R. Sm., Candollea 60: 281. 2005.
Polypodium bombycinum Maxon, Contr. U.S. Natl. Herb. 17(7): 592. 1916.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, G. T. Prance et al. 11894 (NY),
G. T. Prance et al. 11927 (NY), G. T. Prance et al. 12529 (NY), J. Prado et al. 1392
(HPZ, SP), D. C. Daly et al. 11656 (NY); Mâncio Lima, G. T. Prance et al. 12040 (MG,
NY); Rio Branco, D. C. Daly et al. 6868 (NY), E. Forero et al. 6346 (NY); Rodrigues
Alves, J. Prado et al. 1234 (HPZ, SP).
Pleopeltis polypodioides (L.) E. G. Andrews & Windham, Contr. Michigan Herb. 19:
46. 1993.
Acrostichum polypodioides L., Sp. Pl. 2: 1068. 1753.
Manoel Urbano, D. C. Daly et al. 9081 (NY), D. C. Daly et al. 11219 (NY), D. C. Daly
et al. 11562 (NY); Rio Branco, B. Nelson 737 (MG, NY), D. C. Daly et al. 9516 (HPZ,
NY), D. C. Daly et al. 9526 (HPZ, NY), C. A. Cid 2893 (MG, NY); Sena Madureira, D.
C. Daly et al. 7908 (HPZ, NY).
Distribution: Antilles, Mesoamerica, tropical South America.
Serpocaulon caceresii (Sodiro) A. R. Sm., Taxon 55: 928, fig. 3A. 2006.
Polypodium caceresii Sodiro, Crypt. Vasc. Quit. 360. 1893.
Mâncio Lima, G. T. Prance et al. 12141 (NY), M. Silveira et al. 1244 (NY); Rodrigues
Alves, J. Prado et al. 1253 (HPZ, SP); Santa Rosa, D. C. Daly et al. 10169 (NY);
Without locality, J. Jangoux et al. 85-037 (NY).
Serpocaulon triseriale (Sw.) A. R. Sm., Taxon 55: 929, figs. 3F, 4L-O. 2006.
Polypodium triseriale Sw., J. Bot. (Schrader) 2: 26. 1800 [1801].
Material examined: BRAZIL. ACRE: Mâncio Lima, D. C. Daly et al. 8944 (HPZ, NY).
PSILOTACEAE
Psilotum nudum (L.) P. Beauv., Prodr. Aetheogam.: 106, 112. 1805.
Lycopodium nudum L., Sp. Pl. 2: 1100. 1753.
Material examined: BRAZIL. ACRE: Brasiléia, D. C. Daly et al. 9693 (NY); Rio
Branco, D. C. Daly et al. 6669 (HPZ, NY); Sena Madureira, B. Nelson 614 (NY), G.T.
Prance et al. 7956 (NY); Xapuri, C. Figueiredo 211 (HPZ, NY), D. C. Daly et al. 8430
(HPZ, NY); Without locality, J. M. Pires et al. 10058 (NY).
Distribution: Cosmopolitan.
PTERIDACEAE
Adiantum argutum Splitg., Tijdschr. Natuurl. Gesch. Physiol. 7: 427, fig. 1-2. 1840.
Habit/Habitat: Herb; terrestrial in forest on Terra Firme near stream margins.
Material examined: BRAZIL. ACRE: Bôca do Acre, G. T. Prance et al. 2533 (MG, NY);
Cruzeiro do Sul, J. Prado et al. 1344 (HPZ, SP); Porto Acre, D. C. Daly et al. 7990
(HPZ); Rio Branco, D. C. Daly et al. 6675 (NY), S. R. Lowrie et al. 650 (GH, NY, R,
RB, US); Sena Madureira, M. Silveira et al. 668 (NY), T. B. Croat & A. Rosas Jr. 62716
(HPZ); Xapurri, K. A. Kainer et al. 126 (NY), L. C. Ming et al. 356 (HPZ), M. Pinard
809 (NY), D. C. Daly et al. 7261 (HPZ, NY).
Adiantum cajennense Willd. ex Klotzsch, Linnaea 18: 552. 1845.
Habit/Habitat: Herb; terrestrial in forest on Terra Firme on gently undulating terrain,
undistributed but canopy open.
D. C. Daly et al. 11704 (NY), J. Prado et al. 1201 (HPZ, SP), J. Prado et al. 1293
(HPZ, SP); Mâncio Lima, J. Prado et al. 1140 (HPZ, SP); Rodrigues Alves, J. Prado et
al. 1233 (HPZ), J. Prado et al. 1245 (HPZ).
Adiantum decoratum Maxon & Weath., Amer. J. Bot. 19: 165. 1932.
Habit/Habitat: Herb; terrestrial, forest in broad, meandering stream vally in steeply hilly
terrain.
Material examined: BRAZIL. ACRE: Santa Rosa, D. C. Daly et al. 10164 (NY), D. C.
Daly et al. 11365 (NY).
Distribution: Mexico, Mesoamerica, N South America.
Adiantum diogoanum Glaz. ex Baker, J. Bot. 20: 310. 1882.
Material examined: BRAZIL. ACRE: Bujari, D. C. Daly et al. 9385 (NY).
Distribution: South America.
Adiantum dolosum Kunze, Linnaea 21: 219. 1848.
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FERN GAZ. 18(5): 230-263. 2009
Adiantum humile Kunze, Linnaea 9: 80. 1834.
J. Prado et al. 1339 (HPZ, SP); Rodrigues Alves, J. Prado et al. 1246 (HPZ, SP).
Distribution: Belize and tropical South America.
Adiantum latifolium Lam., Encycl. 1: 43. 1783.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, Croat & Rosas 62393A (MO,
UC); J. Prado et al. 1294 (HPZ, SP); J. Prado et al. 1374 (HPZ, SP), T. B. Croat & A.
Rosas Jr. 62393A (HPZ); Manoel Urbano, D. C. Daly et al. 9149 (NY). Tarauacá, 23
set. 1968, Prance et al. 7495 (GH, K, NY, R, UC, US).
Distribution: Mexico, Antilles, Mesoamerica and South America.
Adiantum multisorum Samp., Relat. Commiss. Linhas. Estrateg. Matto Grosso
Amazonas 5(Bot. pt. 7): 11, pl. 1, fig. 1. 1916.
Habit/Habitat: Herb; terrestrial in forest on Terra Firme
Adiantum nudum A. R. Sm., Ann. Missouri Bot. Gard. 77: 260, fig. 6A-B. 1990.
Habit/Habitat: Herb; terrestrial in forest on Terra Firme near stream margins, on clay
soils.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, W. C. Steward et al. P12679
(NY); Mâncio Lima, D. C. Daly et al. 8854 (NY).
Adiantum obliquum Willd., Sp. Pl. 5(1): 429. 1810.
J. Jangoux et al. 85-008 (NY); Rodrigues Alves, J. Prado et al. 1259 (HPZ); J. Prado
Adiantum pectinatum Kunze ex Baker, Syn. Fil. 120. 1867.
Habit/Habitat: Herb; terrestrial, riverside vegetation.
Material examined: BRAZIL. ACRE: Basin of Rio Juruá, Rio Juruá Mirim and Igarapé
Periquito (right-bank tributary), across from Santo Antônio, D. C. Daly et al. 11708
(NY); Mâncio Lima, J. Prado et al. 1139 (HPZ, SP); Manoel Urbano, D. C. Daly et al.
11528 (NY); Rodrigues Alves, J. Prado et al. 1275 (HPZ, SP).
Adiantum poeppigianum (Kuhn) Hieron., Hedwigia 48: 231. 1909.
Adiantum lucidum var. poeppigianum Kuhn, Bot. Jahrb. Syst. 1: 340. 1881.
Habit/Habitat: Herb; terrestrial in forest on steep slopes.
Manoel Urbano, D. C. Daly et al. 11473 (NY); Marechal Thaumaturgo, D. C. Daly et
al. 10950 (NY); Sena Madureira, D. C. Daly et al. 7972 (NY); Santa Rosa, D. C. Daly
et al. 10034 (NY).
Habit/Habitat: Herb; terrestrial in primary forest in low-lying area (Baixio) shallowly
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, D. C. Daly et al. 7365 (HPZ,
NY), G. T. Prance et al. 12065 (K, NY), G. T. Prance et al. 12065a (NY), W. C. Steward
et al. P12894 (K, NY), J. Prado et al. 1347 (HPZ, SP); Manoel Urbano, D. C. Daly et
al. 9173 (NY), D. C. Daly et al. 11392 (NY); Rio Azul, J. Jangoux et al. 85-064 (NY);
Rio Branco, S. R. Lowrie et al. 649 (GH, NY, R, US); Tarauacá, C. Figueiredo et al. 883
(NY); Santa Rosa, D. C. Daly et al. 10050 (NY), D. C. Daly et al. 11004 (NY) Rio Juruá
between Mundurucus & Tatajuba, Maas et al. P12894 (GH, K, NY, R, US).
Adiantum scalare R. M. Tryon, Amer. Fern J. 47: 141, tab. 15. 1957.
Habit/Habitat: Herb; terrestrial in moist forest on Terra Firme, tertiary sediments,
undulating terrain.
Marechal Thaumaturgo, D. C. Daly et al. 10304 (NY); Tarauacá, G. T. Prance et al.
7335 (K, NY).
Adiantum terminatum Kunze ex Miq., Verslagen Meded. Vier Kl. Kon. Ned. Inst.
Wetensch. Letterk. Schoone Kunsten 1842: 187. 1843.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Jangoux et al. 85-009 (NY),
J. Prado et al. 1303 (HPZ, SP), J. Prado et al. 1354 (HPZ, SP), G. T. Prance et al.
12203 (INPA, MO, NY); Porto Valter, D. C. Daly et al. 11786 (NY), D. C. Daly et al.
11788 (NY); Rodrigues Alves, J. Prado et al. 1361 (HPZ, SP).
Adiantum tetraphyllum Willd., Sp. Pl. 5(1): 441. 1810.
Material examined: BRAZIL. ACRE: Brasiléia, D. C. Daly et al. 11878 (NY).
Adiantum tomentosum Klotzsch, Linnaea 18: 553. 1844.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, G. T. Prance 11390 (MG, NY),
255
FERN GAZ. 18(5): 230-263. 2009
J. Prado et al. 1165 (HPZ, SP), J. Prado et al. 1195 (HPZ, SP), P. J. M. Maas et al.
P12791 (NY); P. G. Windisch 2574 (HB, HRCB); Porto Valter, D. C. Daly et al. 7578
(HPZ, NY); Rodrigues Alves, J. Prado et al. 1242 (HPZ).
Adiantum windischii J. Prado, Kew Bull. 60: 119-121, fig. 2. 2005.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, G. T. Prance et al. P13045 (UC);
Ananthacorus angustifolius (Sw.) Underw. & Maxon, Contr. U.S. Natl. Herb. 10: 487.
1908.
Pteris angustifolia Sw., Prodr. 129. 1788.
Sul, D. C. Daly et al. 7495 (HPZ, NY), G. T. Prance et al. 12616 (MG, NY); Rio
Branco, B. Nelson 739 (MG, NY); Santa Rosa, D. C. Daly et al. 10160 (NY); Sena
Madureira, M. Silveira et al. 615 (HPZ, NY); Tarauacá, D. C. Daly et al. 8737 (HPZ,
NY), G.T. Prance et al. 7379 (MG, NY).
Distribution: Mexico, Antilles and N South America.
Anetium citrifolium (L.) Splitg., Tijdschr. Natuurl. Gesch. Physiol. 7: 395. 1840.
Acrostichum citrifolium L., Sp. Pl. 2: 1067. 1753.
Material examined: BRAZIL. ACRE: Tarauacá, D. C. Daly et al. 8269 (NY), G. T.
Prance et al. 7426 (MG, NY).
Hecistopteris pumila (Spreng.) Benedict, London J. Bot. 1: 193. 1842.
Gymnogramma pumila Spreng., Tent. Suppl. 31. 1828.
Habit/Habitat: Herb; epiphyte on the base of trunks in forest on Terra Firme.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1208 (HPZ), J.
Prado et al. 1336 (HPZ, SP), J. Prado et al. 1386 (HPZ, SP).
Pityrogramma calomelanos (L.) Link, Handbuch 3: 20. 1833.
Acrostichum calomelanos L., Sp. Pl. 2: 1072. 1753.
Habit/Habitat: Herb; terrestrial in forest on Terra Firme near river margins on clay soils.
Material examined: BRAZIL. ACRE: Brasiléia, S. R. Lowrie et al. 696 (NY); Cruzeiro
do Sul, J. Prado et al. 1173 (HPZ, SP), J. Prado et al. 1304 (HPZ, SP); Assis Brasil, D.
C. Daly et al. 9634 (HPZ, NY); Mâncio Lima, M. Silveira et al. 1294 (NY); Quixadá,
E. Forero et al. 6381 (NY); Sena Madureira, M. Silveira et al. 670 (HPZ, NY);
Tarauacá, G. T. Prance et al. 7302A (NY), M. Silveira et al. 868 (NY).
Distribution: S Florida, Mexico, Antilles, Mesoamerica and tropical South America.
Polytaenium cajenense (Desv.) Benedict, Bull. Torrey Bot. Club 38: 169. 1911.
Hemionitis cajenensis Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck.
Gesammten Naturk 5: 311. 1811.
Feijó, G. T. Prance et al. 7342 (NY), P. Delprete et al. 8475 (NY); Mâncio Lima, D. C.
Daly et al. 8962 (HPZ, NY); Marechal Thaumaturgo, D. C. Daly et al. 7350 (HPZ,
NY), D. C. Daly et al. 10277 (NY); Porto Valter, M. Silveira et al. 1646 (NY); Santa
Rosa, D. C. Daly et al. 11088 (NY), D. C. Daly et al. 10166 (NY), D. C. Daly et al.
11178 (NY); Tarauacá, D. C. Daly et al. 8619 (HPZ, NY), G. T. Prance 7267A (NY);
Without locality, J. Jangoux 85-095 (NY).
Polytaenium guayanense (Hieron.) Alston, Kew Bull. 1932(7): 134. 1932.
Anthrophyum guaynense Hieron., Hedwigia 57: 212. 1915.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, J. Prado et al. 1316 (HPZ), J.
Prado et al. 1351 (HPZ); Marechal Thaumaturgo, D. C. Daly et al. 10254 (NY), D. C.
Daly et al. 10837 (NY); Porto Valter, W. C. Steward et al. P13013 (MG, NY); Rodrigues
Alves, J. Prado et al. 1240 (HPZ, SP), J. Prado et al. 1248 (HPZ, SP), J. Prado et al.
1277 (HPZ, SP); Without locality, J. Jangoux et al. 85-026 (NY), J. Jangoux et al. 85095A (NY).
Pteris altissima Poir., Encycl. 5: 722. 1804.
Habit/Habitat: Herb; terrestrial in forested slope between Baixio and higher terrace,
canopy relatively closed, understory relatvely rich in herbs.
Material examined: BRAZIL. ACRE: Brasiléia, D. C. Daly et al. 9879 (HPZ, NY);
Manoel Urbano, D. C. Daly et al. 9086 (NY), D. C. Daly et al. 11195 (NY), D. C. Daly
et al. 11260 (NY), D. C. Daly et al. 11422 (NY); Marechal Thaumaturgo, D. C. Daly et
al. 7702 (NY); Santa Rosa, D. C. Daly et al. 10054 (HPZ, NY); Tarauacá, G. T. Prance
et al. 799 (NY), G. T. Prance et al. 7259 (INPA, K, NY, US, R), G. T. Prance et al. 7299
(NY), D. C. Daly et al. 8317 (NY), M. Silveira et al. 997 (NY); Without locality, J.
Jangoux et al. 85-063 (NY).
Pteris grandifolia L., Sp. Pl. 2: 1073. 1753.
Habit/Habitat: Herb; terrestrial on substrate hard and brittle, seeping clear water.
Marechal Thaumaturgo, D. C. Daly et al. 10198 (NY); Porto Valter, P. J. M. Maas et al.
13239 (K, NY).
Distribution: Mexico, Antilles, Mesoamerica, N South America.
Pteris haenkeana C. Presl, Reliq. Haenk. 1(1): 55. 1825.
Habit/Habitat: Herb; terrestrial in moist forest on Terra Firme, tertiary sediments,
undulating terrain.
Manoel Urbano, D. C. Daly et al. 11426 (NY); Marechal Thaumaturgo, D. C. Daly et
al. 7341 (NY); Santa Rosa, D. C. Daly et al. 11314 (NY).
257
FERN GAZ. 18(5): 230-263. 2009
Pteris propinqua J. Ag., Recens. Spec. Pter. 65. 1839.
Habit/Habitat: Herb; terrestrial in undisturbed forest in low-lying area with moist soils,
canopy open.
Sena Madureira, D. C. Daly et al. 7920 (MO); Marechal Thaumaturgo, D. C. Daly et
al.10465 (NY); Rio Branco, L. Coêlho & A. Rosas 1941 (HPZ, NY).
Pteris pungens Willd., Sp. Pl. 5: 387. 1810.
Material examined: BRAZIL. ACRE: Without locality, J. Jangoux et al. 0 (NY).
Vittaria lineata (L.) Sm., Mem. Acad. Roy. Sci. (Turin) 5(1790-1791): 421, pl. 9, fig.
5. 1793.
Pteris lineata L., Sp. Pl. 2: 1073. 1753.
(NY); Mâncio Lima, J. Prado et al. 1227 (HPZ, SP); Rodrigues Alves, J. Prado et al.
1279 (HPZ, SP).
SACCOLOMATACEAE
Saccoloma elegans Kaulf., Berlin. Jahrb. Pharm. Verbundenen Wiss. 21: 51. 1820.
Sena Madureira, C. A. Cid & B. Nelson 2574 (NY); Without locality, J. Jangoux et al.
85-100 (MG, NY).
Saccoloma inaequale (Kunze) Mett., Ann. Sci. Nat., Bot., ser. 4, 15: 80. 1861.
Davallia inaequalis Kunze, Linnaea 9: 87. 1834.
Habit/Habitat: Herb; terrestrial on Terra Firme forest on high terrace.
Sul, G. T. Prance et al. 12194 (MG, NY), G. T. Prance et al. 12333 (NY), G. T. Prance
et al.12574 (NY), J. Prado et al. 1203 (HPZ, SP); Rodrigues Alves, J. Prado et al. 1256
(HPZ, SP); Santa Lucia, J. Pruski et al. 3497 (NY); Without locality, J. Jangoux et al.
85-011 (NY).
Saccoloma membranaceum Mickel, Amer. Fern J. 74: 119, fig. 2A, B. 1984.
Habit/Habitat: Herb; terrestrial in forest on mountains.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, G. T. Prance et al. 12432 (NY).
SALVINIACEAE
Salvinia minima Baker, J. Bot. 24: 98. 1886.
Habit/Habitat: Herb; aquatic in lakes along road margins.
Distribution: S United States, Mexico, Antilles, Mesoamerica and tropical South
America.
SCHIZAEACEAE
Schizaea elegans (Vahl) Sw., J. Bot. (Schrader) 1800(2): 103. 1801.
Acrostichum elegans Vahl, Symb. Bot. 2: 104, tab. 50. 1791.
Habit/Habitat: Herb; terrestrial in forest on Terra Firme
J. Prado et al. 1175 (HPZ, SP), J. Prado et al. 1307 (HPZ, SP); Mâncio Lima, C. A. Cid
et al. 10095 (NY), G. T. Prance et al. 12101 (NY), M. Silveira et al. 1259 (NY).
Distribution: S Mexico, Greater Antilles, Mesoamerica and tropical South America.
SELAGINELLACEAE
Selaginella anceps (C. Presl) C. Presl, Abh. Konigl. Bohm. Ges. Wiss., ser. 5, 3: 581.
1845.
Lycopodium anceps C. Presl, Reliq. Haenk. 1(1): 80. 1825.
Material examined: BRAZIL. ACRE: Sena Madureira, D. C. Daly et al. 7891 (NY), D.
C. Daly et al. 8139 (NY).
Selaginella asperula Spring, Fl. Bras. 1(2): 127. 1840.
Habit/Habitat: Herb; terrestrial in Campina forest, on sandy soils.
G. T. Prance et al. 12643 (NY), J. Prado et al. 1169 (HPZ, SP), J. Prado et al. 1185
(HPZ, SP), J. Prado et al. 1192 (HPZ, SP), J. Prado et al. 1391 (HPZ, SP), T. B. Croat
& A. Rosas Jr. 62643 (HPZ, NY); Porto Valter, D. C. Daly et al. 7573 (HPZ, NY).
Selaginella exaltata (Kunze) Spring, Bull. Acad. Roy. Sci. Bruxelles 10(1): 234. 1843.
Lycopodium exaltatum Kunze, Linnaea 9: 8. 1834[1835].
Habit/Habitat: Herb; terrestrial in forest on Terra Firme and Várzea forest.
Material examined: BRAZIL. ACRE: Bujari, W. A. Anderson 12124 (NY); Brasiléia, D.
C. Daly et al. 6820 (HPZ, NY); Cruzeiro do Sul, J. Prado et al. 1289 (HPZ, SP), L. R.
Marinho 258 (NY); Mâncio Lima, G. T. Prance et al. 12157 (NY), J. Jangoux et al. 85093 (MG, NY); Plácido de Castro, D. C. Daly et al. 6123 (HPZ, NY); Porto Valter, P.
J. M. Maas et al. P12951 (MG, NY); Rio Branco, D. C. Daly et al. 6676 (HPZ, NY);
Sena Madureira, B. Nelson et al. 527 (NY), G. T. Prance et al. 7626 (NY); Tarauacá, G.
T. Prance et al. 7405 (NY); Xapuri, A. R. S. Oliveira et al. 198 (MG, NY); Without
locality, E. Forero et al. 6305 (NY).
Selaginella flagellata Spring, Bull. Acad. Roy. Sci. Bruxelles 10(1): 228. 1843.
Habit/Habitat: Herb; terrestrial in “Baixio”: low terraces possibly flooded occasionally
by overflowing streams.
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FERN GAZ. 18(5): 230-263. 2009
Selaginella haematodes (Kunze) Spring, Fl. Bras. 1(2): 126. 1840.
Lycopodium haematodes Kunze, Linnaea 9: 9. 1834[1835].
Habit/Habitat: Herb; terrestrial in open forest on steep hilly terrain.
Material examined: BRAZIL. ACRE: Santa Rosa, D. C. Daly et al. 11141 (NY).
Selaginella parkeri (Hook. & Grev.) Spring, Bull. Acad. Roy. Sci. Bruxelles 10: 146.
1843.
Lycopodium parkeri Hook. & Grev., Bot. Misc. 2: 388. 1843.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, G. T. Prance et al. 12305 (NY),
(HPZ, SP), J. Prado et al. 1193 (HPZ, SP), J. Prado et al. 1319 (HPZ, SP); Mâncio
Lima, G. T. Prance et al. 12200 (NY); Rodrigues Alves, J. Prado et al. 1362 (HPZ, SP).
Selaginella sulcata (Desv. ex Poir.) Spring ex Mart., Flora 20(2): 126. 1837.
Lycopodium sulcatum Desv. ex Poir., Encycl. 3: 549. 1814.
Habit/Habitat: Herb; terrestrial in forest on Terra firme, on poorly drained soils,
undulating terrain dissected by numerous streams.
Sena Madureira, D. C. Daly et al. 8075 (NY), L. de Lima et al. 540 (HPZ, NY).
TECTARIACEAE
Tectaria draconoptera (D. C. Eaton) Copel., Philipp. J. Sci. 2C: 410. 1907.
Aspidium draconopterum D. C. Eaton, Mem. Amer. Acad. Arts, n.s., 8: 211. 1860.
Habit/Habitat: Hervb; terrestrial, in primary forest, unudulating terrain, in places
C. Daly et al. 11891(NY).
Tectaria incisa Cav., Descr. Pl. 249. 1802.
Habit/Habitat: Herb; terrestrial in forest on Terra Firme near stream margins, on sandy
and clay soils.
NY); Marechal Thaumaturgo, D. C. Daly et al. 10340 (NY), D. C. Daly et al. 10343
(NY); Quixadá, C. A. Cid & A. Rosas 2952 (MG, NY); Rio Branco, C. A. Cid & A.
Rosas 2899 (NY); Santa Rosa, D. C. Daly et al. 10052 (NY), D. C. Daly et al. 11335
(NY); Sena Madureira, G. T. Prance et al. 7667 (MG, NY); Tarauacá, D. C. Daly et al.
8617 (HPZ, NY) ,G. P. da Silva et al. 98 (NY); Xapuri, C. Figueiredo 207 (HPZ, NY),
Xapuri, D. C. Daly et al. 7273 (HPZ, NY).
America.
Tectaria aff. longipinnata A. Rojas, Revista Biol. Trop. 49: 476, fig. 5. 2001.
Habit/Habitat: Herb; terrestrial on Várzea forest.
Material examined: BRAZIL. ACRE: Mâncio Lima, G. T. Prance et al. 12047 (MG,
NY), M. Silveira et al.1229 (NY); Rio Branco, C. A. Cid & A. Souza 2988 (NY); Sena
Madureira, G. T. Prance et al. 7812A (NY).
Tectaria trifoliata (L.) Cav., Descr. Pl. 249. 1802.
Polypodium trifoliatum L., Sp. Pl. 2: 1087. 1753.
Habit/Habitat: Herb; terretrial in forest of Terra Firme, on undulating terrain, canopy
discontinuous.
Material examined: BRAZIL. ACRE: Brasiléia, D. C. Daly et al. 1229 (HPZ, NY).
Triplophyllum boliviense J. Prado & R. C. Moran, Brittonia 60: 106 figs 5, 6. 2008.
Habit/Habitat: Herb; terrestrial in forest on Terra Firme, undulating terrain.
Brasiléia, D. C. Daly et al. 8463 (HPZ, NY); Bujari, D. C. Daly et al. 9856 (HPZ, NY).
Triplophyllum funestum (Kunze) Holttum, Kew Bull. 41: 255. 1986.
Aspidium funestum Kunze, Linnaea 9: 96. 1834.
do Sul, J. Prado et al. 1334 (HPZ, SP); Rodrigues Alves, J. Prado et al. 1247 (HPZ).
THELYPTERIDACEAE
Thelypteris angustifolia (Willd.) Proctor, Bull. Inst. Jamaica, Sci. Ser. 5: 57. 1953.
Meniscium angustifolius Willd., Sp. Pl. 5: 133. 1810.
Habit/Habitat: Herb; terrestrial in forest on Terra Firme, on sandy soil.
Material examined: BRAZIL. ACRE: Mâncio Lima, M. Silveira et al. 1292 (NY), M.
Silveira et al. 1299 (NY), D. C. Daly et al. 8921 (HPZ, NY); Marechal Thaumaturgo,
D. C. Daly et al. 10196 (NY); Tarauacá, D. C. Daly et al. 8764 (HPZ, NY).
Thelypteris arborescens (Humb. & Bonpl. ex Willd.) C. V. Morton, Contr. U.S. Natl.
Herb. 38: 50. 1967.
Meniscium arborescens Humb. & Bonpl. ex Willd., Sp. Pl. 5: 133. 1810.
Thelypteris chrysodioides (Fée) C. V. Morton, Contr. U.S. Natl. Herb. 38: 51. 1967.
Meniscium chrysodioides Fée, Mem. Foug. 5: 225. 1852.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, T. B. Croat & A. Rosas Jr. 62670
(HPZ, NY).
261
FERN GAZ. 18(5): 230-263. 2009
Thelypteris decussata (L.) Proctor, Bull. Inst. Jamaica, Sci. Ser. 5: 59. 1953.
Polypodium decussatum L., Sp. Pl. 2: 1093. 1753.
Infraspecific: var. decussata
Habit/Habitat: Herb; terrestrial in forest on Terra Firme, on steep slopes.
Thelypteris jamesonii (Hook.) R. M. Tryon, Rhodora 69: 6. 1967.
Nephrodium jamesonii Hook., Sp. Fil. 4: 66. 1862.
Habit/Habitat: Herb; terrestrial in forest on Terra Firme, on steep slopes.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, D. C. Daly et al. 11698 (NY).
Distribution: N South America and southern Brazil.
Thelypteris juruensis (C. Chr.) R. M. Tryon & D. S. Conant, Acta Amazon. 5: 33. 1975.
Dryopteris juruensis C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk.
Math. Afd., ser. 7, 10(2): 256, fig. 43d. 1913.
(NY); Manoel Urbano, D. C. Daly et al. 11391 (NY); Rio Branco, C. A. Cid & A. Rosas
2950 (NY); Sena Madureira, G. T. Prance et al. 7812 (NY), G. T. Prance et al. 7853
(NY).
Thelypteris macrophylla (Kunze) C. V. Morton, Amer. Fern J. 61: 17. 1971.
Meniscium macrophyllum Kunze, Flora 22(1): Beibl. 44. 1838.
Thelypteris membranacea (Mett.) R. M. Tryon, Rhodora 69: 7. 1967.
Phegopteris membranacea Mett., Fil. Lechl. 2: 22. 1859.
Material examined: BRAZIL. ACRE: Mâncio Lima, M. Silveira et al. 1249 (NY).
Thelypteris opposita (Vahl) Ching, Bull. Fan Mem. Inst. Biol., Bot. 10: 253. 1941.
Polypodium oppositum Vahl, Eclog. Amer. 3: 53. 1807.
Material examined: BRAZIL. ACRE: Assis Brasil, D. C. Daly et al. 9644 (HPZ, NY),
D. C. Daly et al. 9653 (HPZ, NY).
Thelypteris opulenta (Kaulf.) Fosberg, Smithsonian Contr. Bot. 8: 3. 1972.
Aspidium opulentum Kaulf., Enum. Fil. 238. 1824.
Habit/Habitat: Herb; terrestrial in open forest with palms and weak presence of Guadua
sarcocarpa, canopy ca. 25 m, deeply dissected terrain. Sandy clay soil.
G. T. Prance et al. 12428 (MG, NY), L. R. Marinho 204 (NY), P. J. M. Maas et al.
P13304 (NY), J. Prado et al. 1333 (HPZ, SP); Mâncio Lima, J. Prado et al. 1144 (HPZ,
SP); Manoel Urbano, D. C. Daly et al. 10458 (NY), D. C. Daly et al. 11498 (NY);
Marechal Thaumaturgo, D. C. Daly et al. 10366 (NY); Porto Acre, C. Figueiredo & I.
Riveiro 751 (NY); Santa Rosa, D. C. Daly et al. 10158 (NY), D. C. Daly et al. 11098
(NY), D. C. Daly et al. 11327 (NY); Sena Madureira, G. T. Prance et al. 7662 (NY), L.
de Lima et al. 553 (HPZ, NY); Xapuri, L. G. Lohmann & E. C. de Oliveira 598 (NY).
Distribution: Antilles, Mesoamerica, N South America. Native to Asia and Africa.
Thelypteris patens (Sw.) Small, Ferns S. E. States 243. 1938.
Polypodium patens Sw., Prodr. 133. 1788.
Habit/Habitat: Herb; terrestrial in forest on steep slopes.
Material examined: BRAZIL. ACRE: Assis Brasil, D. C. Daly et al. 9635 (NY); Porto
Valter, D. C. Daly et al. 11780 (NY).
Thelypteris serrata (Cav.) Alston, Bull. Misc. Inform. Kew 1932: 309. 1932.
Meniscium serratum Cav., Descr. Pl. 548. 1802.
Habit/Habitat: Herb; terrestrial in primary moist forest on Terra Firme.
Material examined: BRAZIL. ACRE: Brasiléia, S. R. Lowrie & B. Nelson 695 (MG,
NY); Rio Branco, D. C. Daly et al. 6882 (HPZ, NY), E. Forero et al. 6377 (MG, NY).
Distribution: S Florida, Antilles, Mesoamerica, tropical and subtropical South America.
Thelypteris tristis (Kunze) R. M. Tryon, Rhodora 69: 8. 1967.
Polypodium triste Kunze, Linnaea 9: 47. 1834.
Habit/Habitat: Herb; terrestrial in forest of Terre Firme, on clay soils.
Mâncio Lima, J. Prado et al. 1138 (HPZ, SP); Porto Valter, D. C. Daly et al. 11767
(NY).
WOODSIACEAE
Diplazium ambiguum Raddi, Opusc Sci. 3: 292. 1819.
Habit/Habitat: Herb; terrestrial on disturbed area of Terra Firme.
(NY).
Distribution: tropical South America.
Diplazium grandifolium (Sw.) Sw., J. Bot. (Schrader) 1800(2): 62. 1801.
Asplenium grandifolium Sw., Prodr. 130. 1788.
NY); Marechal Thaumaturgo, D. C. Daly et al. 10302 (NY); Xapuri, M. Pinard et al.
812 (NY).
Distribution: Mexico, Antilles, Mesoamerica and N South America.
Diplazium lechleri (Mett.) T. Moore, Index Filic. 141. 1859.
Asplenium lechleri Mett., Fil. Lechl. 1: 16, tab. 2. 1856.
263
FERN GAZ. 18(5): 230-263. 2009
Diplazium praestans (Copel.) Maxon & C. V. Morton, Contr. U.S. Natl. Herb. 38: 41.
1967.
Athyrium praestans Copel., Amer. Fern J. 38: 132. 1948.
(HPZ, NY); Tarauacá, G. T. Prance et al. 7400 (MG, NY).
Hemidictyum marginatum (L.) C. Presl, Tent. Pterid. 111, tab. 3, fig. 24. 1836.
Asplenium marginatum L., Sp. Pl. 2: 1082. 1753.
Habit/Habitat: Herb; terrestrial in Várzea forest.
Material examined: BRAZIL. ACRE: Cruzeiro do Sul, G. T. Prance et al. 12479 (NY).
Distribution: S Mexico, Antilles, Mesoamerica, tropical South America.
ACKNOWLEDGEMENTS
We thank Dr. Douglas Daly (NY) for inviting us to participate in this project. The senior
author thanks The New York Botanical Garden Herbarium for the use of facilities
during his three visits to compile these data for publication. This study received finacial
support from the Brazilian Research Council CNPq (Proc. n. 303867/2004-3). We also
thank Dr. Alan Smith (UC) for corrections and constructive comments on the
manuscript.
REFERENCES
LABIAK, P.H. & PRADO, J. 2007. New records of pteridophytes from Bolivia and
Brazil. Amer. Fern J. 97(2): 112-122.
PICHI-SERMOLLI, R.E.G. 1996. Authors of scientific names in Pteridophyta. Royal
Botanic Gardens, Kew.
PRADO, J. 2005. A new species and hybrid in Adiantum (Pteridaceae) from South
America. Kew Bulletin 60(2): 117-121.
PRADO, J. & MORAN, R.C. 2008. Revision of the Neotropical species of
Triplophyllum (Tectariaceae). Brittonia 60(2): 103-130.
PRADO, D.E. & GIBBS, P.E. 1993. Patterns of species distribution in the dry seasonal
forest of South America. Ann. Missouri Bot. Gard. 80(4): 902-927.
SILVEIRA, M. 2003. Estudos botânicos no Acre: história, resultados e impactos nas
políticas públicas. Pp. 209-212. In Desafios da Botânica brasileira no novo milênio:
inventário, sistematização e conservação da diversidade vegetal (M.A.G. JARDIM,
M.N.C. BASTOS & J.U.M. SANTOS, eds.). SBB, UFPA, MPEG, EMBRAPA,
Belém.
SMITH, A.R., PRYER, K.M., SCHUETTPELZ, E., KORALL, P., SCHNEIDER, H. &
WOLF, P.G. 2006. A classification for extant ferns. Taxon 55: 705-731.
TRYON, R.M. & CONANT, D. 1975. The ferns of Brazilian Amazonia. Acta
Amazonica 5(1): 23-34.
WINDISCH, P.G. 1979. Adições ao inventário das pteridófitas do Acre. Bradea 3(5):
29-30.
FERN GAZ. 18(5):264-282. 2009
264
DESICCATION TOLERANCE IN SOME BRITISH FERNS
M.C.F. PROCTOR
School of Biosciences, University of Exeter, Geoffrey Pope Building, Stocker Road,
Exeter EX4 4QD
Key-words: Asplenium, chlorophyll fluorescence, drying rate, light responses,
Polypodium, recovery rate, relative humidity, relative water content.
ABSTRACT
Leaves of ten British fern species were tested for their tolerance of desiccation.
Asplenium ruta-muraria, A. septentrionale, A. trichomanes, A. ceterach,
Polypodium cambricum and P. interjectum withstood drying for periods of a
week or more to a relative water content (RWC) of c. 4–7%. This is far below
the RWC (c. 30%) at which most vascular-plant tissues are irretrievably
damaged. One population of Asplenium adiantum-nigrum was desiccation
tolerant, another was not. Aspelnium obovatum was fairly tolerant, behaviour
differing with intensity of desiccation and in old and young growth. Polypodium
cambricum and P interjectum were both highly tolerant. Polystichum aculeatum
was not tolerant. Recovery rates of RWC and the chlorophyll-fluorescence
parameter Fv/Fm did not vary greatly between species, with half-recovery times
around 2–4 h. The small Asplenium species and A. ceterach dried quickly (halfdrying times a few hours), suggesting little stomatal control over drying. The
much slower drying of the Polypodium species suggests that their stomata close
under water stress. Photosynthetic electron flow in most species saturated at a
quarter to a half of full summer sunlight. Asplenium ruta-muraria, A.
septentrionale and A. trichomanes showed a similar tendency to non-saturating
electron flow at high irradiances as many desiccation-tolerant bryophytes. In A.
ceterach and Polypodium cambricum electron flow was somewhat depressed at
high irradiances. The results are discussed in relation to their ecological and
evolutionary implications.
INTRODUCTION
The tolerance to desiccation of the rusty-back fern Asplenium ceterach L., the
widespread Mediterranean Notholaena marantae (L.) Desv. and the American
‘resurrection’ ferns Polypodium polypodioides (L.) Watt and P. virginiacum L. is well
known (Oppenheimer & Halevy, 1962; Schwab et al., 1989; Iljin, 1931; Potts &
Penfound, 1948; Stuart, 1968; Gildner & Larson, 1992; Muslin & Homann, 1992;
Reynolds & Bewley, 1993). Many other ferns have been reported as desiccation
tolerant, including a number of species of Cheilanthes, Pellaea, Asplenium and
Polypodium; a list with references was given by Proctor & Pence (2002) and further
species were added by Kessler & Siorak (2007). Most of these are essentially ferns of
seasonally arid regions, or saxicolous or epiphytic species of intermittently waterstressed habitats. Kappen (1964) investigated freezing, heat and drought tolerance
round the seasons of the common ferns around Göttingen in central Germany. He found
no sharp line between poikilohydric, desiccation-tolerant ferns and homoiohydric, more
sensitive species, as did Kessler & Siorak in North America. In general, Kappen found
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FERN GAZ. 18(5): 264-282. 2009
the summer-green ferns were sensitive to drying at all times, but most winter-green
species were more tolerant of drying in the cold, dry Central European winter and
spring than from midsummer to autumn. The most desiccation-tolerant species (in
descending order of tolerance) were Polypodium vulgare, Asplenium septentrionale, A.
ruta-muraria and A. trichomanes. Perhaps paradoxically, there is as much physiological
information on desiccation tolerance of the moisture-loving Hymenophyllaceae as on
most other ferns (Shreve, 1911; Holloway, 1923a, b; Härtel, 1940a, b; Richards &
Evans, 1972; Proctor, 2003).
The response of Asplenium ceterach to desiccation and its ability to revive after rain
are very apparent in dry summers. The desiccation tolerance of the two common small
saxicolous Asplenium species is less obvious until they are observed closely. The
present project grew out of work on desiccation tolerance in bryophytes, and developed
over a number of years as opportunity offered or material came to hand. Its aims were
to establish which of our common ferns are tolerant to desiccation, and to collect data
on the desiccation-tolerance of ferns (in particular rates of drying and recovery) for
comparison with bryophytes and angiosperm ‘resurrection plants.’
All the measurements in this paper were made on excised leaves. Material of Asplenium
trichomanes, A. ruta-muraria, A. ceterach and Polypodium cambricum was collected
from Devonian limestone rocks and walls near Chudleigh, Devon (most on 10
December 2000 and 5 January 2001; some A. ceterach in March–April 2008). Some
measurements were made on leaves of A. adiantum-nigrum from a mortared pebble
wall facing the sea at Budleigh Salterton, Devon (4 January 2001); most measurements
on this species were on material from a dry Permian conglomerate roadside bank at
Kennford, Devon (8 February 2006). A. obovatum was from schist outcrops on
Gammon Head, near East Prawle, Devon (30 April 2006). A. septentrionale was from
a slate dry-stone roadside wall south of Trefriw, Gwynedd (11 September 2005).
Polypodium interjectum was from a Permian roadside outcrop near Kennford, Devon (8
February 2006). Polystichum aculeatum came from Malham, Yorkshire (26 July 2006).
Chlorophyll fluorescence measurements using a modulated fluorometer (FMS-1,
Hansatech, King’s Lynn) were used throughout this work. Chlorophyll fluorescence
provides a simple and rapid non-invasive method of monitoring various aspects of the
photosynthetic performance of a plant. Schreiber et al., (1995) and Maxwell & Johnson
(2000) provide an introduction to the technique and explain the calculation of the
commonly used parameters. The parameter Fv/Fm measures the maximum quantum
efficiency of photosystem II; it is generally in the range 0.76–0.84 in dark-adapted
unstressed material, and is widely used as a quick general measure of ‘stress’ or
(conversely) ‘viability’, and is so used here. The non-photochemical quenching
parameter NPQ is largely a measure of photo-protection. The photochemical quenching
parameter qP is a measure of the oxidation state of the first electron acceptor QA of
photosystem II; it is often used (as here) in the inverse form 1–qP.
In the light, the effective quantum yield of photosystem II (FPSII) multiplied by the
irradiance in quantum units (PPFD) provides a relative measure of the rate of electron
transport through PSII, and hence of photosynthetic electron flow (RETR). This figure
includes not only electron transport resulting in carbon fixation, but also photorespiration and flow to any other electron sinks. However, under most conditions the
bulk of the electron flow will be to carbon fixation, so for many purposes RETR
PROCTOR: DESICCATION TOLERANCE IN SOME BRITISH FERNS
266
provides a convenient non-invasive surrogate measure of photosynthetic performance;
for comparisons of chlorophyll-fluorescence and gas-exchange data in bryophytes, see
Marschall & Proctor (2004) and Proctor et al. (2007).
In general, the recovery of both Fv/Fm and RWC were well fitted by logistic curves
on a logarithmic timescale. For the RWC curves, and in those cases where Fv/Fm for
the dry material was substantially >0.0, a 4-parameter logistic curve was appropriate.
This has a lower asymptote corresponding to the value of RWC or Fv/Fm in dry
material and an upper asymptote at the maximum (fully recovered) value of Fv/Fm.
Where Fv/Fm in dry material was ~0 or unknown, a 3-parameter logistic starting at 0
was fitted. In either case, the IC-50 value (point of inflection) of the curve was taken as
a measure of the time for ‘half-recovery’. The relation of RETR to irradiance is
generally well fitted by negative-exponential curve of the form y = (A – e–kx), where y
= RETR and x = PPFD and A and k are constants, at least at low irradiances (<400 mmol
m-2 s-1). Deviations may arise through electron flow to sinks other than photosynthetic
carbon fixation (bryophytes often show non-saturating electron flow of this kind
(Marschall & Proctor, 2004)), or through photoinhibition depressing electron flow at
high irradiances. A useful test of goodness-of-fit of the exponential curve is that the
product Ak should approximate to the dark-adapted value of Fv/Fm (c. 7.5–8.5). NPQ
is generally well fitted by a logistic curve starting at 0. The parameter 1–qP is also often
well fitted by a logistic curve, but the fit is very sensitive to the top few data points,
which can lead to misleading estimates of the upper asymptote and IC-50 value; suspect
values have been omitted from Table 4.
Some comment may be made on measures of water content and water availability.
The water content of desiccation-tolerant plants is often given as a percentage of dry
weight. This is useful for comparisons within a population of a single species, but not
for comparisons between different populations or different species because of
differences in the proportion of cell-wall materials and cell contents. Water content
relative to that at full turgor (RWC) is a much more widely useful measurement, and
has been emphasised here. Relative humidity (RH) of the air is only useful if the
temperature is known (in this paper c. 20°C), and its biological relevance is limited.
Evaporation rates depend on saturation deficit, the difference between the absolute
humidity of the air and the saturated absolute humidity at the same temperature (which
can be looked up in tables); it is usually expressed in terms of partial pressure of water
vapour (kPa). Water potential (Ψ) which defines the tendency for water to move from
one place to another, is usually expressed in pressure units (MPa); for air, it can be
calculated from relative humidity and temperature. For the physical and mathematical
relations between these quantities see, e.g., Slatyer (1967), Nobel (1974), Jones (1992).
RESULTS
Table 1 lists some water-content data for the ferns investigated, expressed as a
percentage of oven-dry weight, and as relative water content. Water content at full
turgor on a dry-weight basis varies widely even within the same species; at full turgor
RWC for all species is 1.0 by definition. A number of measurements were made on ‘air
dry’ material. The figures in the table show clearly that ‘air dry’ can embrace a
substantial range in actual water content measured as RWC; cold dry weather outdoors
will tend to give low absolute (and relative) humidity indoors, and warm or humid
weather the opposite. Fern material equilibrated in a desiccator at c. 74% RH had an
RWC around 0.14–0.16; material equlibrated at 43% RH showed RWC mostly below
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FERN GAZ. 18(5): 264-282. 2009
0.10. Of the air dried material, Asplenium trichomanes (1) and Polypodium cambricum
(1) were probably dried to an equivalent of c.60–70% RH at 20 °C, and the other
samples at c.30–40% RH.
Three small saxicolous Asplenium species
The recovery of relative water content and the fluorescence parameter Fv/Fm on rewetting after a few days’ desiccation in Asplenium ruta-muraria L.and A. trichomanes
L. is shown in Figures 1a and 1b and Table 2. A. trichomanes that had been dried to 14%
RWC recovered rather faster than material dried to a RWC of 7.5%; the half-recovery
times of Fv/Fm in the fitted curves of Figure 1b are respectively 1.7 and 3.9 hours, but
the asymptotes of the two curves are not significantly different. A much rarer species
than either of these in Britain, A. septentrionale L. was reported as highly desiccation
tolerant by Kappen. Leaves collected from a population in North Wales in September
2005, were allowed to air dry. The chlorophyll fluorometer was not available from then
until January 2006, by which time the leaves had been dry for four months. However,
on re-wetting (12 Jan.) they recovered their normal fresh appearance and appeared fully
turgid the following day. Ten leaves re-wetted after 3 days moist followed by 2 days air
dry gave Fv/Fm 0.754 ± 0.035 after 24 h rehydration (Figure 1c). Of these three
Asplenium species, A. ruta-muraria showed the highest light-saturation level, followed
by A. septentrionale with A. trichomanes the lowest; all three showed at least some
indication of non-saturating electron flow at high irradiances (Figure 11a).
Two further species of Asplenium
Asplenium adiantum-nigrum L. sometimes accompanies the other small saxicolous
Asplenium species in crevices of rocks and walls, but usually avoids the most sunexposed and water-stressed situations. Leaves of this species from a mortared pebble
wall facing the sea at Budleigh Salterton were promisingly firm textured, and looked
possible candidates for desiccation tolerance. However, re-wetted after a few days’
desiccation at 43% RH they showed only slight temporary signs of recovery of Fv/Fm
for the first few hours, and by the following day it was clear they would not recover
further (Table 2.)
Plants from another population, on a steep dry roadside bank near Kennford, Devon,
behaved quite differently. Figure 2 shows the rapid and progressive loss of weight of
leaf segments dried at 74% and 43% RH, with fitted exponential curves; the asymptotes
approximated to the figures in Table 1; the leaves dried quickly, losing half their water
within 6.2 h at 74% RH (–41 MPa) and 4.7 h at 43% RH (–114 MPa). The recovery of
Fv/Fm on re-wetting after the two desiccation regimes is shown in Figure 3. The data
and the fitted curves suggest that intensity of drying has a marked effect, but neither the
asymptotes nor the half-recovery times are significantly different, and the slope factor
just fails to reach significance at the 5% level.
Asplenium obovatum Viv. dried rather more slowly than A. adiantum- nigrum; the
half-drying times seen in Figure 4 are around 11 h at 74% RH and 9 h at 43% RH
Recovery of Fv/Fm on re-wetting was both less complete and more variable than in the
Kennford A. adiantum-nigrum. After drying at 74% RH, the dark green leaves of the
previous season showed only a transient rise in Fv/Fm on re-wetting; further recovery
failed completely. However, in material dried at 43% RH, the older leaves recovered
consistently and well. The bright green new-season’s leaves recovered well from drying
at 74% RH, but very erratically from the 43% RH treatment (Figure 5).
409 ± 16
139 ± 4
108 ± 11
344 ± 36
170 ± 22
279 ± 82
156 ± 34
110 ± 16
213 ± 10
306 ± 35
Asplenium ruta-muraria
A. trichomanes (1)
A. trichomanes (2)
A. adiantum-nigrum (1)
A. obovatum
A. ceterach
Polypodium cambricum (1)
P. interjectum
Species
Full
turgor%DW
-
-
18.6 ± 8.3
9.4 ± 0.4
-
-
29.6 ± 2.9
7.9 ± 0.8
19.3 ± 4.6
20.9 ± 4.4
%DW
33.2 ± 5.7
44.3 ± 13.7
-
-
-
-
0.167 ± 0.056
0.063 ± 0.018
33.9 ± 9.2
26.1 ± 4.7
-
-
-
-
-
%DW
0.142 ± 0.031
0.161 ± 0.026
-
-
0.106 ± 0.035
0.155 ± 0.045
15.7 ± 4.6
15.8 ± 1.9
-
-
18.1 ± 5.0
17.4 ± 2.2
-
-
-
-
-
-
-
-
%DW
0.052 ± 0.015
0.072 ± 0.006
-
-
0.055 ± 0.035
0.106 ± 0.022
-
-
-
-
RWC
Water content at 43% RH
(-114 MPa)
RWC
Water content at 74% RH
(-41 MPa)
0.087 ± 0.017
0.075 ± 0.014
0.140 ± 0.037
0.068 ± 0.013
RWC
Water content ‘air dry’
Table 1. Water content of fern leaves at full turgor (as % oven-dry weight at 70°C), and ‘air dry’ and in equlibrium with air at 74% RH (c. –40
MPa at 20°C) and 43% RH (c. –114 MPa at 20°C) as % dry weight and as relative water content (RWC). The two sets of measurements on
Asplenium trichomanes and Polypodium cambricum were both from Chudleigh, in Dec. 2000 and Jan. 2001 respectively. A. adiantum-nigrum
(1) is from Budleigh Salterton; A. adiantum-nigrum (2) is from Kennford.
268
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FERN GAZ. 18(5): 264-282. 2009
The rusty-back fern, Asplenium ceterach
The curling of the dry leaves in Asplenium ceterach initially prevented measurement of
Fv/Fm in the early stages of rehydration, so the data in Figure 6 are from leaves dried
under light pressure to keep them flat. The rise in Fv/Fm was rather slower to get going
in A. ceterach, but climbed more steeply once it had started. Light saturation was
reached at around half noon summer sunlight, but showed some depression of electron
flow at higher irradiances.
An experiment with Polystichum aculeatum (L.) Roth
Pinnae of Polystichum aculeatum dried rather quickly, giving half-drying times of 12.9
h at 74% RH and 8.4 h at 43% RH, reaching practical equilibrium with the atmosphere
in the desiccator in the course of a week. The corresponding equilibrium water contents
(asymptotes of the curves) were 0.109 and 0.069 RWC. Remoistened after 9 days, they
showed small but erratic increases of Fv/Fm for an hour or two, but these were not
maintained and by the following day it was clear that all the pinnae were moribund.
The polypodies, Polypodium cambricum L. and P. interjectum Shivas
These two species were very tolerant of desiccation, and differ in important respects
from the ferns just considered. They dried very much more slowly (Figure 7); the halfdrying times for P. interjectum at 74% RH (–41 MPa) and 43% RH (–114 MPa} were
respectively 2.33 days (56 h) and 1.4 days (34 h). These drying times are almost an
order of magnitude longer than those seen in Asplenium adiantum-nigrum, around five
times longer than in A. obovatum. As a consequence, the asymptotes of the exponential
curves, 0.074 and 0.037 RWC respectively, are substantially below the figures in Table
1 from weighings after a week’s drying. Clearly these latter figures do not reflect
equilibrium with the air.
The recovery curves of Fv/Fm in Figures 8 and 9 thus start from relative water
contents above equilibrium with the ambient air – but still well below levels at which
normal metabolism is possible, or survivable by typical vascular-plant mesophyll
tissues. The curves differ significantly only in the initial level of Fv/Fm. Little sign of
recovery is apparent in the first hour after re-wetting, either in the physical appearance
of the leaves (Figure 10) or in Fv/Fm, but after that recovery is rapid. The half-recovery
times are around 2h, and in the same range as in the other species examined (Table 3).
Photosynthetic electron flow saturated at around 40% of noon summer sunlight, but
was markedly depressed at high irradiances so that the effective saturation irradiance
must be below the PPFD95% figure in Table 4 (Figure 11b).
DISCUSSION
The results confirm that the leaves of some ferns are remarkably tolerant to desiccation,
recovering well from drying for a week or more to a water content of 15–20% of their
dry weight, and a tissue relative water content (RWC) of 4–7%. The mesophyll of
typical vascular plants is generally damaged beyond recovery if RWC falls below about
30% (or a water potential below about –7 MPa), so these ferns are far more tolerant of
desiccation than most drought-tolerant vascular plants (Larcher 1995). Their tolerance
is comparable with such mosses as Tortula (Syntrichia) ruralis or Racomitrium
lanuginosum. However it is also clear there are degrees of desiccation tolerance, and
that tolerance may vary within species. Of our Asplenium species, A. septentrionale is
270
Figure 1. Recovery on remoistening of (a) Fv/Fm and RWC in air-dry leaves of
Asplenium ruta-muraria. Fitted logistic curves. Mean ± s.d., n = 3, (b) Fv/Fm in two
samples of air-dry leaves of A. trichomanes (sample (1) from 0.140 RWC, (2) from
0.075 RWC), and recovery of RWC for sample (1). Means (n = 3), error bars omitted
for clarity (log scale), (c) Fv/Fm in A. septentrionale after four months air dry, with
fitted 4-parameter logistic curve. Mean ± s.d., n = 6.
Species
A. trichomanes (1)
A. trichomanes (2)
A. ceterach
P. interjectum
Fv/Fm after 24 h rehydration
‘Air-dry’
74% RH (-41 MPa)
43% RH (-114 MPa)
-
0.756 ± 0.035
-
-
-
0.796 ± 0.020
-
-
-
0.707 ± 0.059
-
-
-
0.075 ± 0.059
-
-
0.785 ± 0.037
-
0.735 ± 0.053
0.646 ± 0.075
0.800 ± 0.017
-
0.520 ± 0.122*
0.417 ± 0.113
0.816 ± 0.018
-
0.812 ± 0.004
0.789 ± 0.023
-
0.703 ± 0.109
-
-
0.868 ± 0.012
-
0.832 ± 0.033
0.769 ± 0.047
0.821 ± 0.017
-
0.824 ± 0.010
0.813 ± 0.019
FERN GAZ. 18(5): 264-282. 2009
A. obovatum
Fv/Fm at full turgor before
desiccation
271
Table 2. Fv/Fm at full turgor before desiccation and 24 h after rehydration.
272
Figure 2. Drying data for leaves of Asplenium adiantum-nigrum in desiccators at ~74%
and 43% RH (–41 and –114 MPa); the fitted exponential curves give half-drying times
of respectively 6.2 h and 4.7 h. Means ± s.d., n = 6.
Figure 3. Recovery of Fv/Fm on re-wetting leaves of Asplenium adiantum-nigrum
after 6 days drying at 74% RH (–41 MPa; circles and full line) and 43% RH (–114
MPa; triangles and broken line; error bars omitted for clarity), with fitted 4-parameter
logistic curves. Means ± s.d., n = 6.
273
FERN GAZ. 18(5): 264-282. 2009
probably the most tolerant, followed by A. ruta-muraria and A. trichomanes. In A.
adiantum-nigrum it appears that some populations are tolerant, but others are not, or
perhaps that tolerance varies with season, as Kappen found. A. obovatum seems (from
one sample) to be less tolerant than A. adiantum-nigrum, and to show some differences
between young and old leaves. Two leaves of A. marinum L. from near Kynance Cove,
Cornwall in early June 2001 did not recover after drying, but this species should be
tested from more sites. Kappen (1964) found the leaves of summer-green ferns such as
Dryopteris filix-mas are similar to flowering plant leaves in their sensitivity to drying,
with little variation through the season. Species which remained green through the
winter were in general more tolerant in the winter and spring months, but the limiting
leaf saturation-deficit given by Kappen for Polystichum lobatum (aculeatum) is little
different from his corresponding figure for Dryopteris filix-mas. The experiment
reported here gave no indication of desiccation tolerance in P. aculeatum. A high degree
of desiccation tolerance is seen in Polypodium cambricum and P. interjectum; the
‘Polypodium vulgare’ that Kappen found the most desiccation tolerant of all his ferns
was probably P. vulgare L. sensu stricto.
The Asplenium species dried quickly, with a half-drying time of a few hours, and
there is clearly little stomatal control over water loss; six leaves of A. ceterach gave
half-drying times ranging from 2.5 to 9.0 h (depending on size), with a mean of 4.8 ±
2.3 h. Drying of Polystichum aculeatum was only a little slower. The Polypodium
species dried much more slowly, with half-drying times in P. interjectum of a day or
two, suggesting that the stomata close under water stress. This may be related to the
habitats of the plants. The Asplenium species primarily occupy open, brightly lit and
often sunny rock-crevice habitats. In bright sunshine in summer, a small Asplenium is
sitting immovably in the steep temperature gradient close to the ground surface. In this
situation a turgid, actively metabolising plant needs the cooling effect of transpiration
to keep its leaves from reaching lethal temperatures (c.45–50°C), and leaves of these
small ferns are correspondingly dependent on a constant supply of water from the root
system to maintain turgor. Once air dry, desiccation-tolerant plants can withstand much
higher temperatures. Plants tall and open enough for the air to circulate freely around
their leaves do not face this overheating hazard, and Polypodium leaves, held free of the
substratum on flexible petioles would be expected to be more closely coupled to the
temperature of the air. Perhaps Asplenium ceterach, with its dry leaves curved away
from the substratum and protected by their thick scaly covering, comes somewhere
between A. ruta-muraria and Polypodium cambricum in this respect. In their
photosynthetic responses, the small Asplenium species have something in common the
desiccation-tolerant mosses in the same habitats. In this they differ from A. ceterach and
P. cambricum, but these latter two are still not, photosynthetically, unequivocal sun
plants.
The ferns rooted in rock crevices can probably all draw on substantial reserves of
water at depth, which dry up only in hot dry spells in summer. Polypodium is probably
drawing mainly on a more superficial reserve of water, which both dries up and is
replenished more frequently, and this must underlie its success as an epiphyte (Zotz &
Hietz, 2001). Recovery took place at similar rates in all the ferns examined, with halfrecovery times around 2–4 hours, in contrast to the wide range of recovery rates in
bryophytes (Table 3).
These desiccation-tolerant ferns are interesting in an ecological and evolutionary
context. Proctor & Tuba (2002) postulated two strategies for plant life on land,
274
Figure 4. The course of RWC during drying and re-wetting with (a) desiccation at 74%
RH (–41 MPa) and (b) at 43% RH (–114 MPa). Note in (a) the lower final RWC and
less complete recovery of new leaves, and lack of significant difference between old
and new growth in (b); half-drying times in (a) are about 11 h, and in (b) abour 9 h.
275
FERN GAZ. 18(5): 264-282. 2009
Figure 5. The course of Fv/Fm over the duration of the experiment. Note more rapid
decline in Fv/Fm in the new growth, parallelling its faster drying, and the varied
recovery of old and new growth following the two desiccation treatments.
Figure 6. Recovery of Fv/Fm and RWC (from different samples) in Asplenium
ceterach. (data set (2) of Table 3) Mean ± s.d., n = 2 (Fv/Fm), n = 3 (RWC). Fitted 4parameter logistic curves. Note the very wide error bars in the lower part of the curve,
reflecting the variation between replicates in the ‘latent’ time before recovery of Fv/Fm
begins.
276
Figure 7. Drying data for RWC in Polypodium interjectum at 74 and 43% RH (–41 and
–114 MPa), with fitted exponential curves.
Figure 8. Recovery of Fv/Fm following re-wetting of Polypodium cambricum after 8
days’ drying at 74% and 43% RH (–41 and –114 MPa). The only significant difference
in the parameters of the fitted curves is in their lower limit.
277
FERN GAZ. 18(5): 264-282. 2009
Figure 9. Recovery of Fv/Fm following re-wetting of Polypodium interjectum after 8
days’ drying at 74% and 43% RH (–41 and –114 MPa). As in P. cambricum, the only
significant difference between the fitted curves is in their lower limit.
Figure 10. The 74% RH recovery curve of Figure 10 annotated with the appearance
of the leaves at intervals during re-wetting; the comments relate to the successive data
points. The mean value of Fv/Fm in dry leaves is arbitrarily plotted at 0.01 h.
278
Figure 11. Light response of photosynthetic electron flow in (a) Asplenium
trichomanes, showing (very variable) non-saturating electron flow at high irradiance,
and (b) Polypodium cambricum, showing (consistent) depression of electron flow at
high irradiance.
Half-recovery
(IC-50) /h
Rate (slope) parameter
3.05 ± 0.48
A. trichomanes (1)
Species
279
Table 3. Parameters of logistic curves fitted to Fv/Fm recovery data; fitted values ± standard errors. Upper and lower asymptotes are given for
4-parameter curves; logistic curves starting from zero have only an upper asymptote, estimating fully-recovered Fv/Fm. The figures for A.
ceterach are from Spring 2008 data.
Upper and lower asymptotes
Lower limit
-1.34 ± 0.28
0.79 ± 0.10
0.04 ± 0.06
1.73 ± 0.27
-0.86 ± 0.11
0.88 ± 0.04
-
A. trichomanes (2)
3.90 ± 0.22
-0.81 ± 0.03
0.87 ± 0.0.02
-
A. septentrionale
1.64 ± 0.08
-1.11 ± 0.06
0.75 ± 0.02
0.25 ± 0.01
A. adiantum-nigrum (2) [-41 MPa]
2.11 ± 0.20
-1.12 ± 0.11
0.79 ± 0.02
-
A. adiantum-nigrum (2) [-114 MPa]
4.46 ± 3.42
-0.68 ± 0.20
0.86 ± 0.17
-
A. ceterach (1)
3.59 ± 0.27
-1.97 ± 0.27
0.94 ± 0.16
0.04 ± 0.02
A. ceterach (2)
5.80 ± 0.33
-3.07 ± 0.44
0.75 ± 0.05
0.08 ± 0.02
Polypodium cambricum (-41 MPa)
2.92 ± 0.20
-3.68 ± 0.55
0.84 ± 0.02
0.48 ± 0.01
Polypodium cambricum (-114 MPa)
2.45 ± 0.27
-1.09 ± 0.13
0.85 ± 0.05
0.04 ± 0.03
P. interjectum (-41 MPa)
2.03 ± 0.18
-2.43 ± 0.43
0.81 ± 0.05
0.18 ± 0.02
P. interjectum (-114 MPa)
1.87 ± 0.43
-1.51 ± 0.50
0.80 ± 0.11
0.02 ± 0.05
FERN GAZ. 18(5): 264-282. 2009
Upper limit
Species and comments
1 - qP
NPQ
PPFD95%
y-range
IC-50
y-range
IC-50
Asplenium ruta-muraria, Chudleigh, Dec 2000
821 ± 40
10.68
511
0.87
1543
Asplenium ruta-muraria, Chudleigh, Jan 2001
772 ± 192
10.91
1077
-
[>1000]
Asplenium septentrionale, Trefriw, Sep 2004
566 ± 82
7.91
1813
-
-
Asplenium trichomanes, Chudleigh, Dec 2000
405 ± 41
7.91
487
0.74
299
Asplenium adiantum-nigrum, Budleigh Salterton, Jan 2001
524 ± 301
5.56
368
1.21
1201
Asplenium obovatum, Gammon Head, (last season’s growth)
421 ± 81
4.99
295
0.94
573
Asplenium obovatum, Gammon Head (new-season’s leaves)
550 ± 91
6.12
440
-
[>500]
Asplenium ceterach, Chudleigh, Jan 2001 (last season’s growth)
1142 ± 104
7.85
772
-
[>1000]
Asplenium ceterach, Chudleigh, Jan 2001 (new-season’s leaves)
834 ± 84
8.33
644
-
[>1000]
Polypodium cambricum, Chudleigh, Jan 2001
772 ± 215
5.43
400
1.02
839
280
/ì mol m-2 s-1
Table 4. Light-response parameters of some desiccation tolerant ferns. The 95% saturation values (mean ± SD) are calculated from negativeexponential curves fitted to the data-points for irradiances below 400 mmol m-2 s-1; all the species give a good fit to this part of the curve. At
higher irradiances, Asplenium ruta-muraria, A. trichomanes and A. septentrionale all show the non-saturating electron flow commonly
observed in desiccation-tolerant bryophytes (Marschall & Proctor, 2004). Most of the other species show at least some depression of
photosynthetic electron flow at high irradiance, so that Asplenium ceterach and Polypodium cambricum may in practice seldom reach the ‘95%
saturation’ values given above. All species show high NPQ values by typical vascular-plant standards. Parameters have been given for 1-qP
only when the logistic curves gave a sufficiently satisfactory fit to the data; the fit of the curves is very sensitive to error in the highest few data
points.
281
FERN GAZ. 18(5): 264-282. 2009
exemplified by the homoiohydric mainstream vascular plants on the one hand, and the
poikilohydric, desiccation-tolerant bryophytes on the other. Alongside the contrast
between homoiohydry and poikilohydry there is another contrast, between endohydry
and ectohydry. In vascular plants the physiologically important free water is in the
xylem; they are endohydric, and the plant body is isolated from its surroundings by a
waterproof and water-repellent epidermis. In most bryophytes, the physiologically
important water is in capillary spaces outside the plant; they are ectohydric. The
desiccation tolerant fern sporophytes considered in this paper withstand drying (and are
to that extent poikilohydric), but are endohydric. This is also true of most other vascular
‘resurrection plants’. All of these plants function as normal vascular plants when water
is freely available, but they have added to their repertoire of responses the fall-back
option of desiccation tolerance (already inherent in their spores) when the water supply
fails. We are in a region of ecological niche-space where several adaptive strategies
converge, and none is optimal. Desiccation-tolerant ferns share their habitat with fully
poikilohydric (and ectohydric) mosses and liverworts, with small succulents, and
(seasonally) with winter annuals. Fern gametophytes, on the other hand are
unequivocally poikilohydric and ectohydric, and take their place alongside bryophytes
in a diversity of habitats where some of them are as desiccation tolerant as their
bryophyte neighbours (Watkins et al., 2007).
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HÄRTEL, O. 1940a. Physiologische Studien an Hymenophyllaceen. I.
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HÄRTEL, O. 1940b. Physiologische Studien an Hymenophyllaceen. II. Wasserhaushalt
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HOLLOWAY, J.E. 1923a. Studies in the New Zealand Hymenophyllaceae. Part 1. The
distribution of the species in Westland, and their growth-forms. Transactions of the
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HOLLOWAY, J.E. 1923b. Studies in the New Zealand Hymenophyllaceae. Part 2. The
distribution of the species throughout the New Zealand biological region.
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ILJIN, W.S. 1931. Austrocknungsresistenz des Farnes Notochlaena Marantae R.Br.
Protoplasma 13: 322–330.
JONES, H.G. 1992. Plants and microclimate. 2nd edn. Cambridge University Press,
Cambridge.
KAPPEN, L. 1964. Untersuchungen über den Jahreslauf der Frost-, Hitze- und
Austrocknungsresistenz von Sporophyten einheimischer Polypodiaceen (Filicinae).
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KESSLER, M. & SIORAK, Y. 2007. Desiccation and rehyration experiments on the
leaves of 43 pteridophyte species. American Fern Journal 97: 175–185.
LARCHER, W. 1995. Physiological plant ecology, 3rd edn. Springer-Verlag. New York
MARSCHALL, M. & PROCTOR, M.C.F. 2004. Are bryophytes shade plants?
Photosynthetic light responses and proportions of chlorophyll a, chlorophyll b and
total carotenoids. Annals of Botany 95: 593–603.
MAXWELL, K. & JOHNSON, G.N. 2000. Chlorophyll fluorescence – a practical
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MUSLIN, L.E.H. & HOMANN, P.H. 1992. Light as a hazard for the desiccationresistant ‘resurrection’ fern Polypodium polypodioides. Plant, Cell & Environment
15: 81–89.
NOBEL, P.S. 1974. An introduction to biophysical plant physiology. W.H. Freeman,
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polypodioides (L.) A.S, Hitchcock. Ecology 29: 43–53.
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responses, water relations and desiccation tolerance of the filmy ferns
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will be refereed
aligned.
aligned.
a legible form.
Supplements.
FERN GAZ. 18(6):284-285. 2009
284
LECTOTYPIFICATION OF DRYOPTERIS PSEUDOMAS BASED ON
WOLLASTON'S SPECIMEN OF D. AFFINIS
C.R. FRASER-JENKINS
Student Guest House, Thamel, P.O. Box 5555, Kathmandu, Nepal.
e-mail: [email protected]
ABSTRACT
Dryopteris pseudomas (Woll.) Holub & Pouzar is lectotypified from Wollaston's
specimen of D. affinis (Lowe) Fras.-Jenk. at MANCH, which maintains the
stability of the nomenclature of the group.
INTRODUCTION
Dryopteris pseudomas (Woll.) Holub & Pouzar was raised by Holub to provide a name
for the European D. affinis (Lowe) Fras.-Jenk. group, sens. lat., and following the
subdivision of the group as a critical complex, was subsequently applied by him to what
is now called D. borreri Newm. However Wollaston also intended his name to apply to
what is now the whole D. affinis aggregate and had no intention to confine it to D.
borreri. His description emphasised the scaly, truncate-pinnuled, glossy nature of the
plant and would obviously be most aptly applied to what is now D. affinis. But in fact
his original specimen, marked by him as type, contains both species, so could be
lectotypified in either sense. However the most obvious and characteristic morphology,
as emphasisised in Wollaston's description, is that of the larger, left hand and main
specimen on the sheet, which is a specimen of D. affinis, and is therefore selected here
as lectotype. The small right-hand specimen is a compact and scaly D. borreri.
LECTOTYPIFICATION
Lastrea pseudomas Wollaston, Phytologist, n.s. [ser. 2] 1: 171-172 (1855).
Dryopteris pseudomas (Wollaston) Holub & Pouzar, in Holub, Fol. Geobot. Phytotax.
(Praha) 2: 329 332 (1967).
Lectotype of L. pseudomas, designated here: Britain, England: "Lastrea pseudo-mas
(Wol.) Type. Geo. B. Wollaston [G.B. Wollaston scripsit]. Nat. order 93, no 1793 C.
Lastrea filix-mas (Presl) var. paleacea (Moore). Loc.-, Coll. George B. Wollaston, Date
July 1884. Herbarium Walter W. Reeves [printed label of Reeves]. Herbarium
Manchester Museum. [With lectotype labels from H.V. Corley in 1972 [ined.] and C.R.
Fraser-Jenkins in 1979 and 198l]." (MANCH!). Left hand specimen only, which fits
Wollaston's protologue descriptions (Wollaston 1855 and see also 1863 and 1915 and
Wollaston's description in Moore 1855) much better than does D. borreri. The date of
July 1884 is almost certainly, and is presumed here to be the date of acquisition by
Reeves, filled in by him on his herbarium label, printed with the categories, 'Loc.', 'Date'
etc., rather than the date of actual collection by Wollaston, whose own label suggests
that this is original (pre 1855) material and can therefore be chosen as lectotype rather
than neotype. Other Reeves specimens in MANCH collected by him (sometimes with
others) have a full date (including day) and locality given, unlike the present one, and
another specimen from Wollaston of D. oreades, labelled var. abbreviatum, likewise has
the same date of July 1884 and no locality. Manchester is where Wollaston's herbarium
was sent (via Reeves) and there exists no other pre 1855 Wollaston material labelled as
285
FERN GAZ. 18(6): 284-285. 2009
representing L. pseudo-mas at MANCH or at KEW (in Moore's herbarium and the
general herbarium), nor at BM, E, CGE or OXF, but at Paris there are two sheets (Herb.
C.W.W. Hope 29 Jan. 1914) of D. affinis borreri, labelled as cultivated by P. Neill
Fraser from plants sent to him by Wollaston as typical (P!). These specimens (along
with his citing var. paleacea Moore [sensu Moore, pro parte = D. affinis] and var.
borreri Newm. in his protologue) help to indicate that Wollaston's concept of Lastrea
pseudomas was mixed and, as might be expected, he did not distinguish between what
are now the constituent species of the aggregate, but referred to the more typical or
characteristic "D. affinis-like" plants, more-or-less regardless of the modern species in
the group. This further supports the suitability of the above lectotype of D. affinis as
representing the "more characteristic" species.
CONCLUSION
Lectotypification in the present and most obvious sense has the effect of stabilising the
current nomenclature of the group (see Fraser-Jenkins 2007) regarding the widely used
name D. borreri, which might otherwise have been replaced by D. pseudomas had that
been lectotypified from the other specimen on Wollaston's sheet. Although
Fraser-Jenkins (1980) mentioned that L. pseudomas was illegitimate, he was merely
misusing the term at that time to mean a synonym and has not subsequently thought of
it as being illegitimate. In the same paper, when placing D. pseudomas in the synonymy
of D. affinis subsp. borreri (Newm.) Fras.-Jenk., he had not then fully investigated the
identity of many names applied to the group, as he did subsequently in his detailed
unpublished monograph of the D. affinis group (copy at BM). The present
lectotypification helps clarify the views expressed by Ekrt et al. (2009), where it is
suggested that D. pseudomas might apply to D. borreri.
ACKNOWLEDGEMENTS
I am grateful for the helpful discussion and kind acknowledgment from Dr. Libor Ekrt,
University of South Bohemia, České Budějovice, Czech Republic.
REFERENCES
Ekrt, L., Trávníček, P., Jarolímová, V., Vít, P. & Urfus, T. 2009. Genome size and
morphology of the Dryopteris affinis group in Central Europe, Preslia 18(1):
261-280.
Fraser-Jenkins, C.R. 1980. Dryopteris affinis: a new treatment for a complex species in
the European Pteridophyte Flora, Willdenowia 10(1): 107-115.
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FERN GAZ. 18(6):286-306. 2009
286
AN ANNOTATED CHECKLIST OF FERNS AND LYCOPHYTES
FROM THE BIOLOGICAL RESERVE OF UATUMÃ, AN AREA WITH
PATCHES OF RICH-SOILS IN CENTRAL AMAZONIA, BRAZIL
1
G. ZUQUIM1,2, J. PRADO3 & F.R.C. COSTA1
Instituto Nacional de Pesquisas da Amazônia, Coordenação de Pesquisas em
Ecologia, C.P. 478, CEP 69083-000, Manaus, AM, Brazil.
2
Author for correspondence: [email protected]
3
Instituto de Botânica, Seção de Curadoria do Herbário, C.P. 3005, CEP 01061-970,
São Paulo, SP, Brazil.
Key words: Amazonia, Brazil, diversity, edaphic factor, floristic survey, pteridophytes,
rainforest
Palavras-chave: Amazônia, Brasil, diversidade, levantamento florístico, pteridófitas
ABSTRACT
There are few inventories of ferns and lycophytes of the Brazilian Amazon and
most of them were made in areas of poor soils. The general species richness
found in previous studies is low. This study presents an annotated list containing
122 species and three varieties of ferns and lycophytes from Biological Reserve
of Uatumã, a large reserve that includes small patches of rich soils in Central
Amazonia. The study site is located ca. 200 km north of Manaus, AM, Brazil.
Inventories were made more intensively inside a grid system of trails of 5 x 5
km and less intensively in some white-sand forests and waterfalls in the
surroundings of Balbina´s Village. Reference to the original description, habit,
habitat, material examined, geographic distribution and voucher information are
presented for each taxon. This is the first floristic survey of ferns and lycophytes
from areas with relatively rich soils of Central Amazonia. The high number of
species found challenges the concept that Central Amazonia is species-poor for
the two groups studied and the strong relationship with soil conditions reinforces
the proposal to include ferns and lycophytes as indicator groups of the
distribution of biodiversity in the region.
RESUMO
Poucos são os inventários de samambaias e licófitas feitos na Amazônia
brasileira e a maioria destes foram feitos em áreas cujos solos são pobres em
nutrientes. Não obstante, a riqueza de espécies encontrada em estudos prévios
foi baixa. Neste estudo é apresentada uma lista anotada de 122 espécies e três
variedades de samambaias e licófitas da Reserva Biológica do Uatumã, uma
reserva grande em extensão e com pequenas manchas de solo rico na Amazônia
Central. O estudo foi realizado a cerca de 200 km ao norte de Manaus, AM,
Brasil. Os inventários foram mais intensos dentro de uma grade de trilhas de 5 x
5 km e com menor intensidade em áreas de campinas e campinaranas e em
cachoeiras nos arredores da Vila de Balbina. Para cada táxon são apresentadas
informações acerca do local da publicação original, hábito, habitat, material
examinado, distribuição geográfica e voucher. Este é o primeiro estudo sobre
samambaias e licófitas em áreas com solos relativamente ricos na Amazônia
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Central. A alta riqueza de espécies encontrada afeta a idéia geral de que a
Amazônia Central é pobre em espécies destes dois grupos e a forte relação entre
as espécies e as condições do solo reforça a sugestão de se incluir samambaias e
licófitas como grupos indicadores da distribuição da biodiversidade.
INTRODUCTION
Central Amazonian soils are generally deep, highly lixiviated and nutrient-poor
(Chauvel et al., 1987, Laurence, 2001, Mertens, 2004), and this may be the reason for
the low species richness of ferns and lycophytes (39 to 83 species per site) reported in
many inventories in this region (Costa et al., 1999, Costa et al., 2005, Zuquim et al.,
2007). The same pattern is probably true for the Rio Negro Basin, which is mainly
covered by poor sandy soils (Freitas & Prado, 2005). In Western Amazon, Tuomisto et
al. (2002) sampled relatively richer-soil areas and found 140 species in a 20 x 25 km
study site in Ecuador.
To contribute to the knowledge of the Brazilian Amazonian flora, we present here
an annotated checklist of the species of ferns and lycophytes found in the Biological
Reserve of Uatumã. Our evidence suggests that the low species richness in Central
Amazonia sampled until now (as compared with other Neotropical forests) may be due
to low sampling effort from the infrequent patches of rich soils that are difficult to
access.
Study area - the study was conducted at the Biological Reserve of Uatumã (Rebio
Uatumã) and surroundings of Balbina´s Village, about 200 km to the North of the city
of Manaus, AM, Brazil (Fig. 1). It is located in the municipalities of Presidente
Figueiredo, São Sebastião do Uatumã, and Urucará. The Reserve was created in 1990
and it is the largest Biological Reserve in Brazil.
The reserve comprises a mosaic of white-sand shrublands (campinas), white-sand
forests (campinaranas), and dense Terra Firme forests. The gradient of soils is broad
and varies from very poor and coarse sandy soils to rich clayey soils. We have
considered as rich soils those with more than 3 cmol/kg of exchangeable
bases[(Research Program in Biodiversity (PPBio) - unpublished data] since this
concentration represents around three times the amount found in the soils of the most
studied areas in Central Amazonia (see Laurance 2001 - DBFFP Reserves, and Mertens
2004 - Reserva Ducke).
Inventory of ferns and lycophytes
Field work was carried out from July 2006 to February 2008. In 2006, the Research
Program in Biodiversity (hereafter PPBio – the Portuguese acronym) with the support
of the Chico Mendes Institute of Biodiversity Conservation installed a 5 x 5 km grid
system of trails in the southernmost part of Rebio Uatumã (Fig. 1). Sampling effort was
more intense in the trail system, and special attention was given to the streams crossing
the trails. The trail system encompasses only 25 km2 of the 5,600,000 km2 of the limits
of the reserve. The grid comprises non-inundated Tropical Dense Forest (also called
Terra Firme forests) and is crossed by many streams. To cover the variety of
environments found in the region, one waterfall and a surrounding campinarana area
close to the village of Balbina were visited. These two habitats occur in the landscape
but were not included within the grid system.
Manaus
Manaus
160 Km
10 Km
Figure 1. Location of the study site. Black patches represent water and gray scale patches represents vegetation. Solid lines are the limits of the
Biological Reserve of Uatumã, AM, Brazil. The white square shows the area intensively sampled.
ZUQUIM, PRADO & COSTA: A CHECKLIST OF FERNS FROM UATUMÃ
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Presentation of the checklist
The list is arranged in alphabetic order by families, genera, species, and varieties. The
arrangement of families and genera follows Smith et al. (2006).
For each species and variety the full name of the taxon is given, followed by the
reference of original publication, as well as additional information such as: habit,
habitat, material examined, and geographic distribution. Voucher information for each
species and variety is also cited. Author abbreviations follow Pichi-Sermolli (1996).
RESULTS
One hundred and twenty two species and three varieties of pteridophytes distributed in
21 families were found. The richest families were Pteridaceae (with 22 species.),
Polypodiaceae (18 spp.), and Dryopteridaceae (16 spp.). The most diverse genera were
Adiantum (with 13 species) and Trichomanes (12 spp.). Near 35 % of the species are
epiphytes, but these were probably underestimated due to the difficulty to access the
canopy. Four species (Blechnum serrulatum, Gleichenella pectinata, Dicranopteris
flexuosa, and Palhinhaea cernua) were found only in disturbed areas along road
margins. Actinostachys pennula, Adiantum cinnamomeum, Lindsaea tetraptera, L.
schomburgkii, Elaphoglossum plumosum, Thelypteris arborescens, Trichomanes
bicorne, and T. martiusii were collected exclusively in white-sand forests
(campinaranas). A population of around 50 individuals of Hemionitis rufa was found
in a conspicuous rocky patch of ca. 100 m2 covered by open vegetation (mainly grasses
and ground bromeliads). Apparently, H. rufa is strongly related to this kind of habitat
in Amazonia, but more observations are needed. The other species were distributed over
habitats ranging from stream valleys to hillsides, plateaus, rocky soils and tree-fall gaps
in the Terra Firme forests. Images of almost all species can be accessed by downloading
the “Guide to the ferns and lycophytes of REBIO Uatumã – Central Amazonia” at
http://ppbio.inpa.gov.br/Port/guias/. Two species (Schizaea elegans and Gleichenella
pectinata) were accidentally not collected but are common in the region and are easy to
recognise. Both were recorded in photographs.
LIST OF SPECIES
ASPLENIACEAE
Asplenium auritum Sw., J. Bot. (Schrader) 1800(2): 52. 1801.
Habit/Habitat: herb, epiphytic (canopy), Terra Firme forests.
Studied specimen(s): Zuquim & Junqueira 296 (INPA).
Asplenium cirrhatum Rich. ex Willd., Sp. Pl. 5: 321. 1801.
Habit/Habitat: herb, epiphytic (base of trunks and rocks), Terra Firme forest over rich
soils.
Studied specimen(s): Tuomisto & Prado 15699 (INPA, SP, TUR).
Distribution: Mexico, Antilles, Mesoamerica, and tropical South America.
Asplenium cristatum Lam., Encycl. 2(1): 310. 1786.
Habit/Habitat: herb, epiphytic (base of trunks and rocks), Terra Firme forests.
Studied specimen(s): Zuquim 241 (INPA).
290
Asplenium laetum Sw., Syn. Fil.: 79, 271. 1806.
Habit/Habitat: herb, terrestrial or rupiculous, in Terra Firme forests, near streams.
Studied specimen(s): Zuquim & Figueiredo 97 (INPA); 107 (INPA); Zuquim & BragaNeto 138 (INPA); Zuquim & Jakovac 228 (INPA).
Distribution: Mexico, Antilles, Mesoamerica, and tropical South Amarica.
Asplenium pearcei Baker, Syn. Fil. (ed. 2): 483. 1874.
Habit/Habitat: herb, terrestrial, rupiculous or epiphytic, in Terra Firme forests over rich
soils.
Studied specimen(s): Zuquim & Braga-Neto 185 (INPA); Zuquim 232 (INPA).
Asplenium salicifolium L., Sp. Pl. 2: 1080. 1753.
Habit/Habitat: herb, epiphytic (canopy), Terra Firme forests.
Asplenium serratum L., Sp. Pl. 2: 1079. 1753.
Habit/Habitat: herb, epiphytic, in Terra Firme forests.
Studied specimen(s): Zuquim & Figueiredo 105 (INPA); 106 (INPA); Zuquim & Costa
252 (INPA).
America.
Asplenium stuebelianum Hieron., Hedwigia 47: 222. 1908.
Habit/Habitat: herb, terrestrial or epiphytic, in Terra Firme forests over rich soils*.
Studied specimen(s): Tuomisto & Zuquim 15609 (INPA, SP, TUR).
* Prado & Moran (2009) report it in open forest with bamboo in Acre State.
BLECHNACEAE
Blechnum serrulatum Rich., Actes Soc. Hist. Nat. Paris 1: 114. 1792.
Habit/Habitat: herb, terrestrial, in disturbed and loamy open areas.
Studied specimen(s): Prado & Zuquim 1872 (INPA, SP, TUR).
Distribution: S. Florida, Mesoamerica, Antilles, tropical South America. Also occurs in
Malasia and Australia.
Salpichlaena volubilis (Kaulf.) J. Sm., J. Bot. (Hooker) 4: 168. 1841.
Habit/Habitat: herb, terrestrial, climber, in plateaus or slopes of Terra Firme forests.
Studied specimen(s): Zuquim & Figueiredo 103 (INPA); Zuquim & Junqueira 325
(INPA).
CYATHEACEAE
Cyathea sp.
Habit/Habitat: tree-fern, terrestrial, in Terra Firme forests.
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Cyathea lasiosora (Kuhn) Domin, Pteridophyta: 262. 1929.
Habit/Habitat: tree-fern, terrestrial, in Terra Firme forests near streams.
Studied specimen(s): Zuquim & Braga-Neto 386 (INPA).
Cyathea microdonta (Desv.) Domin, Pteridophyta: 263. 1929.
Cyathea pungens Domin, Sp. Pl. 5: 206. 1810.
Studied specimen(s): Zuquim & Jakovac 197 (INPA); Zuquim & Jakovac 216 (INPA).
DENNSTAEDTIACEAE
Dennstaedtia sp.
Habit/Habitat: herb, terrestrial, in Terra Firme forests, near streams.
DRYOPTERIDACEAE
Bolbitis lindigii (Mett.) Ching, Index Filic., Suppl. 3: 48. 1934.
Habit/Habitat: herb, hemi-epiphytic, in Terra Firme forests.
Studied specimen(s): Zuquim & Vieira 444 (INPA).
Bolbitis nicotianifolia (Sw.) Alston, Bull. Misc. Inform. Kew 1932(7): 310. 1932.
Habit/Habitat: herb, terrestrial, rupiculous or hemi-epiphytic, in Terra Firme forests.
Studied specimen(s): Zuquim & Figueiredo 113 (INPA); Zuquim & Braga-Neto 186
(INPA); Zuquim & Costa 286 (INPA)
Distribution: Antilles, Mesoamerica, and N South America.
Bolbitis semipinnatifida (Fée) Alston, Bull. Misc. Inform. Kew 1932(7): 310. 1932.
Habit/Habitat: herbs, terrestrial and rupiculous, in Terra Firme forests over rich soils.
Studied specimen(s): Zuquim & Figueiredo 94 (INPA); Zuquim & Jakovac 217 (INPA);
Zuquim 240 (INPA).
Distribution: Lesser Antilles and N South America.
Bolbitis serratifolia (Mert. ex Kaulf.) Schott, Gen. Fil., tab. 14. 1834.
Habit/Habitat: herb, terrestrial and rupiculous, in Terra Firme forests over rich soils.
Distribution: S Mexico, Mesoamerica, and tropical South America.
Ctenitis refulgens (Klotzsch ex Mett.) Vareschi, Fl. Venezuela 1: 404. 1969.
Habit/Habitat: herb, terrestrial and rupiculous, in Terra Firme forests near streams.
292
Cyclodium guianense (Klotzsch) L. D. Gómez, Phytologia 60(5): 371. 1986.
Habit/Habitat: herb, terrestrial and rupiculous, in Terra Firme forests over rich soils,
near streams.
Studied specimen(s): Zuquim & Jakovac 205 (INPA); 209 (INPA), Zuquim 237 (INPA).
Cyclodium meniscioides (Willd.) C. Presl, Tent. Pterid.: 85. 1836.
Infraspecific: var. meniscioides
Habit/Habitat: herb, terrestrial or hemi-epiphytic, in swamps of Terra Firme forests.
Studied specimen(s): Zuquim & Jakovac 210 (INPA); Zuquim & Fontelles 375 (INPA).
Didymochlaena truncatula (Sw.) J. Sm., J. Bot. (Hooker) 4: 196. 1841-1842 [1841].
Distribution: S Mexico, Antilles, Mesoamerica, and N South America. Tropical Asia
and Africa.
Elaphoglossum flaccidum (Fée) T. Moore, Index Filic.: 356. 1862.
Habit/Habitat: herb, epiphytic (canopy), in Terra Firme forests.
Elaphoglossum luridum (Fée) H. Christ, Neue Denkschr. Allg. Schweiz. Ges.
Gesammten Naturwiss. 36(1): 33. 1899.
Habit/Habitat: herb, epiphytic (canopy), in Terra Firme forests.
Studied specimen(s): Zuquim 337 (INPA); Zuquim & Vieira 400 (INPA).
Distribution: Antilles, Mesoamerica, and N South America.
Elaphoglossum plumosum (Fée) T. Moore, Index Filic.: 364. 1862.
Habit/Habitat: herb, epiphytic, in white-sand forests (campinaranas).
Elaphoglossum raywaense (Jenm.) Aslton, Bol. Soc. Brot., ser. 2, 32: 24. 1958.
Studied specimen(s): Zuquim & Figueiredo 66 (INPA).
Lomagramma guianensis (Aubl.) Ching, Amer. Fern J. 22: 17. 1932.
Habit/Habitat: herb, hemi-epiphytic, in Terra Firme forests over rich soils.
Distribution: Greater Antilles (except Jamaica) and tropical South America.
Olfersia cervina (L.) Kunze, Flora 7: 312. 1824.
Habit/Habitat: herb, terrestrial and hemi-epiphytic.
Studied specimen(s): Sobrinho 1026 (INPA).
Distribution: S. Mexico, Antilles, Mesoamerica, and tropical South America.
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Polybotrya caudata Kunze, Linnaea 9: 23. 1834.
Habit/Habitat: herb, hemi-epiphytic, in Terra Firme forests, near streams.
Distribution: S Mexico, Mesoamerica, and N South America.
Polybotrya sessilisora R. C. Moran, Bull. Illinois Nat. Hist. Surv. 34: 108, fig. 51. 1987.
GLEICHENIACEAE
Dicranopteris flexuosa (Schrad.) Underw., Bull. Torrey Bot. Club 34(5): 254. 1907.
Habit/Habitat: herb, terrestrial, in disturbed and opened areas.
Studied specimen(s): Zuquim & Jakovac 194 (INPA).
Distribution: SE USA, Antilles, Mesoamerica, and tropical South America.
Gleichenella pectinata (Willd.) Ching, Sunyatsenia 5(4): 276. 1940.
Habit/Habitat: herb, terrestrial, in disturbed and opened areas.
HYMENOPHYLLACEAE
Davalliopsis elegans (Rich.) Copel., Philipp. J. Sci. 67(1): 82. 1938.
Habit/Habitat: herb, terrestrial, in Terra Firme forests, near streams
Studied specimen(s): Zuquim & Figueiredo 77 (INPA); Zuquim& Jakovac 215 (INPA).
Distribution: Antilles, Mesoamerica, and tropical and subtropical South America.
Habit/Habitat: herb, epiphytic, in Terra Firme forests
Distribution: Mexico, Mesoamerica, and tropical South America.
Trichomanes ankersii C. Parker ex Hook. & Grev., Icon. Filic. 2(11): tab. 201. 1831.
Habit/Habitat: herb, terrestrial with climbing raquis, in Terra Firme forest.
Studied specimen(s): Zuquim & Costa 245 (INPA); 273 (INPA).
Trichomanes bicorne Hook., Icon. Plan. 9: tab. 892. 1854.
Habit/Habitat: herb, terrestrial or rupiculous, in Terra Firme and White Sand forests, in
both cases, near streams.
Trichomanes cellulosum Klotzsch, Linnae 18: 531. 1844.
Habit/Habitat: herb, terrestrial, in Terra-Firme forests, near streams.
Studied specimen(s): Zuquim & Figueiredo 71 (INPA); Zuquim 121 (INPA).
294
Trichomanes cristatum Kaulf., Enum. Filic. 265. 1824.
Habit/Habitat: herb, terrestrial, in Terra Firme forests.
Studied specimen(s): Zuquim & Figueiredo 78 (INPA); Zuquim 122 (INPA); Zuquim &
Braga-Neto 388 (INPA).
Trichomanes diversifrons (Bory) Mett. ex Sadeb., Nat. Pflanzenfam. I(4): 108. 1899.
Distribution: S Mexico, Antilles, Mesoamerica, Trinidad, and N South America.
Trichomanes ekmanii Wess. Boer, Acta Bot. Neerl. 11: 319, fig. 33. 1962.
Habit/Habitat: herb, rupiculous and epiphytic, in Terra Firme forests.
(INPA); 191 (INPA); Zuquim & Junqueira 292 (INPA).
Distribution: Antilles, Mesoamerica and N South America
Trichomanes martiusii C. Presl, Hymenophyllaceae 15, 36. 1843.
Habit/Habitat: herb, terrestrial, rupiculous and epiphytic, in Terra Firme and white sand
forests, near streams.
Trichomanes pinnatum Hedw., s.l., Fil. Gen. Sp., tab. 4, fig. 1. 1799.
Distribution: Distribution: S Mexico, Antilles, Mesoamerica, and N South America.
NB. The material cited here probably belongs to more than one taxon, but more studies
are necessary to clarify this situation.
Vandenboschia radicans (Sw.) Copel., (Philipp. J. Sci.) 67(1): 54. 1938.
Habit/Habitat: herb, rupiculous and epiphytic, in Terra Firme forests.
Studied specimen(s): Zuquim & Junqueira 291 (INPA); 313 (INPA); Zuquim &
Braga-Neto 382 (INPA).
Distribution: Mexico, Antilles, Mesoamerica, and widespread in South America.
Trichomanes tuerckheimii H. Christ, Hedwigia 44: 361.1905.
Habit/Habitat: herb, terrestrial with climbing rachis, in Terra Firme forests.
Distribution: S Mexico, Mesoamerica, and N South America.
Trichomanes vittaria DC. ex Poir., Encycl. 8: 65. 1808.
Habit/Habitat: herb, terrestrial, in Terra Firme Forest.
Studied specimen(s): Zuquim & Braga-Neto 155 (INPA); Zuquim & Jakovac 196
(INPA); Zuquim 243 (INPA); Zuquim & Junqueira 320 (INPA).
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LINDSAEACEAE
Lindsaea divaricata Klotzsch, Linnaea 18: 547. 1844.
Studied specimen(s): Zuquim & Figueiredo 67 (INPA); Zuquim & Jacovak 218 (INPA);
Zuquim & Vieira 406 (INPA).
Lindsaea guianensis (Aubl.) Dryand., Trans. Linn. Soc. London 3: 42. 1797.
Distribution: Lesser Antilles, Mesoamerica, N South America.
Lindsaea sp.
Studied specimen(s): Zuquim & Costa 248 (INPA); 253 (INPA); Zuquim & Junqueira
295 (INPA).
Lindsaea lancea (L.) Bedd. var. lancea
Studied specimen(s): Zuquim & Figueiredo 74 (INPA); Zuquim & Vieira 402 (INPA).
Lindsaea lancea var. falcata (Dryand.) Rosenst., Hedwigia 46: 79. 1906.
Habit/Habitat: herb, terrestrial, occasionally epiphytic (in decaying wood), in Terra
Firme forests, near streams.
Distribution: Mexico, Mesoamerica, and N South America.
Lindsaea quadrangularis subsp. antillensis, K. U. Kramer, Acta Bot. Neerl. 6(2): 194.
1957.
Habit/Habitat: herb, terrestrial, in white sand forests.
Distribution: Antilles, Guianas, and Brazil.
Lindsaea schomburgkii Klotzsch, Linnaea 18: 545. 1844.
Habit/Habitat: herb, terrestrial and epiphytic, in white sand forests.
Lindsaea tetraptera K. U. Kramer, Acta Bot. Neerl. 6(2): 181, fig. 38. 1957.
Habit/Habitat: herb, terrestrial, in white sand forests.
Studied specimen(s): Zuquim 124 (INPA); Tuomisto & Prado 15718 (INPA SP, TUR).
LOMARIOPSIDACEAE
Cyclopeltis semicordata (Sw.) J. Sm., Bot. Mag. 72: 36. 1846.
Habit/Habitat: herb, terrestrial, in Terra Firme forests over rich soils.
Studied specimen(s): Zuquim & Jakovac 140 (INPA); 221 (INPA).
296
Lomariopsis japurensis (Mart.) J. Sm., Hist. Fil.: 140. 1875.
(INPA); 183 (INPA); 188 (INPA).
Lomariopsis prieuriana Fée, Mem. Foug. 2: 66, Tab. 25, fig. 1. 1845.
Nephrolepis rivularis (Vahl) Ching, Bot. Jahrb. Syst. 24: 122. 1897.
Habit/Habitat: herb, terrestrial, epiphytic, in Terra Firme forests.
Studied specimen(s): Zuquim 126 (INPA); 127 (INPA); Zuquim & Costa 251 (INPA);
Zuquim & Junqueira 289 (INPA).
LYCOPODIACEAE
Palhinhaea camporum (B. Øllg. & P. G.Windisch), Holub, Folia Geobot. Phytotax. 26:
93. 1991.
Habit/Habitat: herb, terrestrial, in disturbed open areas.
LYGODIACEAE
Lygodium volubile Sw., J. Bot. (Schrader) 1801(2): 303. 1803.
Habit/Habitat: herb, terrestrial with climbing rachis, in Terra Firme forests.
MARATTIACEAE
Danaea leprieurii Kunze, Mém. Acad. Roy. Sci. (Turin) 5(1790-1791): 429 tab. 9.
1793.
Danaea nodosa (L.) Sm., Mem. Acad. Roy. Sci. (Turin) 5(1790-1791): 420, tab. 9, fig.
11. 1793.
Danaea trifoliata Reichenb. ex Kunze, Analecta Pteridogr.: 4, tab. 2. 1837.
Studied specimen(s): Zuquim & Braga-Neto 377 (INPA); Zuquim & Junqueira 409
(INPA).
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METAXYACEAE
Metaxya rostrata C. Presl, Tent. Pterid.: 60, tab. 1, fig. 5. 1836.
Studied specimen(s): Zuquim & Costa 249 (INPA); Zuquim & Junqueira 304 (INPA).
Distribution: S Mexico, Lesser Antilles, Mesoamerica, and N South America.
POLYPODIACEAE
Campyloneurum phyllitidis C. Presl, Tent. Pterid.: 190. 1836.
Studied specimen(s): Zuquim & Costa 279 (INPA).
America.
Campyloneurum repens (Aubl.) C. Presl, Tent. Pterid.: 190. 1836.
(INPA); 383 (INPA).
Cochlidium serrulatum (Sw.) L. E. Bishop, Amer. Fern J. 68(3): 80. 1978.
Dicranoglossum desvauxii (Klotzsch) Proctor, Rhodora 63: 35. 1961.
Distribution: Lesser Antilles, tropical South America.
Microgramma dictyophylla (Kunze ex Mett.) de la Sota, Novon 17(1): 27. 2007.
Studied specimen(s): Zuquim & Costa 266 (INPA); Zuquim 336 (INPA).
Microgramma lycopodioides (L.) Copel., Ann. Cryptog. Phytopathol. 5: 185. 1947.
Studied specimen(s): Zuquim & Figueiredo 104 (INPA); Zuquim & Junqueira 312
(INPA); Zuquim & Braga-Neto 384 (INPA).
Distribution: S Mexico, Antilles, Mesoamerica, South America
Microgramma megalophylla (Desv. ) de la Sota, Bol. Soc. Argent. Bot. 10: 158. 1963.
Studied specimen(s): Zuquim & Neres 376 (INPA).
Microgramma percussa (Cav.) de la Sota, Physis (Buenos Aires) 44(106C): 28. 1986.
Distribution: S Mexico, Mesoamerica, and tropical South America.
298
Microgramma reptans (Cav.) A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40(8): 230. 1975.
Studied specimen(s): Zuquim & Jakovac 213 (INPA); Zuquim & Costa 250 (INPA);
Zuquim & Junqueira 290 (INPA).
Distribution: Mexico, Cuba, Mesoamerica, and tropical South America.
Microgramma sp.
Microgramma tecta (Kaulf.) Alston, J. Wash. Acad. Sci. 48: 232. 1958.
Studied specimen(s): Zuquim & Braga-Neto 152 (INPA); 170 (INPA); Zuquim &
Jakovac 229 (INPA).
Microgramma thurnii (Baker) R. M. Tryon & Stolze, Fieldiana, Bot., n.s. 32: 156. 1993.
Studied specimen(s): Zuquim & Mesquita 129 (INPA); Zuquim & Costa 274 (INPA).
Pecluma camptophyllaria (Fée) M. G. Price, Amer. Fern J. 73(4): 113. 1983.
Distribution: Greater Antilles, Mesoamerica, amd N South America.
Pecluma pilosa (A. M. Evans) M. Kessler & A. R. Sm, Candollea 60(1): 281. 2005.
Phlebodium decumanum (Willd.) J. Sm., J. Bot. (Hooker) 4: 59. 1841.
Habit/Habitat: herb, epiphytic, in Terra Firme forests and in open areas.
Distribution: S Florida, Antilles, Mesoamrica, tropical and subtropical South America.
Pleopeltis bombycina (Maxon) A. R. Sm., Candollea 60(1): 281. 2005.
Serpocaulon adnatum (Kunze ex Klotzsch) A. R. Sm., Taxon 55(4): 927. 2006.
Serpocaulon caceresii (Sodiro) A. R. Sm., Taxon 55(4): 928, fig. 3A. 2006.
299
FERN GAZ. 18(6): 286-306. 2009
Studied specimen(s): Zuquim & Jakovac 208 (INPA); Zuquim & Junqueira 301 (INPA);
Zuquim & Braga-Neto 392 (INPA).
PTERIDACEAE
Adiantum adiantoides (J. Sm.) C. Chr., Ind.: 123. 1905.
Studied specimen(s): Zuquim & Braga-Neto 380 (INPA); Zuquim & Vieira 395 (INPA).
Adiantum argutum Splitg., Tijdschr. Natuurl. Gesch. Physiol. 7: 427, fig. 1-2. 1840.
Studied specimen(s): Zuquim & Braga-Neto 142 (INPA); 167 (INPA).
Adiantum cajennense Willd. ex Klotzsch, Linnaea 18: 552. 1845
Studied specimen(s): Zuquim & Braga-Neto 180 (INPA); 379 (INPA); Zuquim 235
(INPA).
Adiantum cinnamomeum Lellinger & J. Prado, Amer. Fern J. 91(1): 4, fig. 1c-f, fig.1.
2001.
Habit/Habitat: herb, terrestrial, in Terra Firme forests, in slightly disturbed areas.
Distribution: Guianas and N. Brazil.
Adiantum dolosum Kunze, Linnaea 21: 219. 1848.
Adiantum glaucescens Klotzsch, Linnaea 18: 552. 1844.
Adiantum humile Kunze, Linnaea 9: 80. 1834.
(INPA).
Distribution: Belize, and tropical South America.
Adiantum lucidum (Cav.) Sw., Syn. fil.: 121. 1806.
Distribution: Meosamerica, Antilles, and South America.
300
Adiantum obliquum Willd., Sp. Pl. 5(1): 429. 1810.
(INPA).
Habit/Habitat: herb, terrestrial, in Terra Firme forests over rich soils.
Studied specimen(s): Zuquim & Braga-Neto 141 (INPA); Zuquim & Jacovak 222
(INPA).
Adiantum terminatum Kunze ex Miq., Verslagen Meded. Vier Kl. Kon. Ned. Inst.
Wetensch. Letterk. Schoone Kunsten 1842: 187. 1843.
Studied specimen(s): Zuquim & Figueiredo 69 (INPA); 99 (INPA); 101 (INPA); Zuquim
& Braga-Neto 148 (INPA); 169 (INPA).
Adiantum tomentosum Klotzsch, Linnaea 18: 553. 1844.
Studied specimen(s): Zuquim & Jakovak 198 (INPA).
Ananthacorus angustifolius (Sw.) Underw. & Maxon, Contr. U.S. Natl. Herb. 10: 487.
1908.
Studied specimen(s): Zuquim & Junqueira 302 (INPA); 309 (INPA).
Distribution: Mexico, Antilles and N South America.
Anetium citrifolium (L.) Splitg., Tijdschr. Natuurl. Gesch. Physiol. 7: 395. 1840.
Hecistopteris pumila (Spreng.) Benedict, London J. Bot. 1: 193. 1842.
Habit/Habitat: herb, epiphyti, in Terra Firme forests.
301
FERN GAZ. 18(6): 286-306. 2009
Hemionitis rufa (L.) Sw., J. Bot. (Schrader) 1800(2): 16. 1801.
Habit/Habitat: herb, terrestrial, in Terra Firme forests, in rocky areas.
Distribution: S Mexico, Greater Antilles, Mesoamerica, and N South America.
Pityrogramma calomelanos (L.) Link, Handbuch 3: 20. 1833.
Distribution: S Florida, Mexico, Antilles, Mesoamerica, and tropical South America.
Polytaenium guayanense (Hieron.) Alston, Kew Bull. 1932(7): 134. 1932.
Pteris pungens Willd., Sp. Pl. 5: 387. 1810.
Studied specimen(s): Zuquim & Junqueira 331 (INPA); Zuquim & Vieira 399 (INPA).
Pteris tripartita Sw., J. Bot. (Schrader) 1800(2): 67. 1801.
(INPA).
Distribution: naturalized and widespread in tropical America, native in Africa, Asia, and
Polynesia.
Vittaria lineata (L.) Sm., Mem. Acad. Roy. Sci. (Turin) 5(1790-1791): 421, pl. 9, fig. 5.
1793.
Studied specimen(s): Zuquim & Junqueira 303 (INPA); 310 (INPA).
SACCOLOMATACEAE
Saccoloma inaequale (Kunze) Mett., Ann. Sci. Nat., Bot., ser. 4, 15: 80. 1861.
Habit/Habitat: herb, terrestrial, in Terra Firme forest
(INPA); Zuquim & Junqueira 311 (INPA).
SALVINIACEAE
Salvinia auriculata Aubl., Hist. Pl. Guiane 2: 969, pl. 367. 1775.
Habit/Habitat: herb, aquatic, in low energy and large water bodies.
Distribution: Mexico, Greater Antilles, Mesoamerica, and tropical South America.
302
SCHIZAEACEAE
Actinostachys pennula (Sw.) Hook., Gen. Fil., tab. 111A. 1842.
Habit/Habitat: herb, terrestrial, in white sand forests and shrub lands.
Studied specimen(s): Prado et al. 1885 (INPA, SP, TUR).
Distribution: Mesoamerica, Trinidad, tropical South America, and Uruguay.
Schizaea elegans (Vahl) Sw., J. Bot. (Schrader) 1800(2): 103. 1801.
Habit/Habitat: herb, terrestrial, in Terra Firme and white sand forests.
Studied specimen(s): not collected
Distribution: S Mexico, Greater Antilles, Mesoamerica, and tropical South America.
SELAGINELLACEAE
Selaginella amazonica Spring, Fl. Bras. 1(2): 124. 1840.
Distribution: Colombia and Brazil.
Selaginella breynii Spring, Fl. Bras. 1(2): 121. 1840.
Distribution: Colombia, French Guiana, and N Brazil.
Selaginella pedata Klotzsch, Linnaea 17: 521. 1844.
Studied specimen(s): Zuquim & Figueiredo 95 (INPA); 96 (INPA); Zuquim &
Braga-Neto 181 (INPA); Zuquim 234 (INPA).
TECTARIACEAE
Tectaria incisa Cav. Descr. Pl.: 249. 1802.
Distribution: S Florida, Mexico, Antilles, Mesoamerica, and tropical and subtropical
South America.
Triplophyllum crassifolium Holttum, Kew Bull. 41(2): 257. 1986.
(INPA); 159 (INPA); 162 (INPA); 193 (INPA); 387 (INPA); Zuquim & Jakovac 200
(INPA); Zuquim 239 (INPA); Zuquim & Costa 246 (INPA).
Triplophyllum dicksonioides (Fée) Holttum, Kew Bull. 41(2): 257. 1986.
Studied specimen(s): Zuquim & Souza 444 (INPA); Zuquim & Neres 445 (INPA).
Distribution: Trinidad and South America.
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FERN GAZ. 18(6): 286-306. 2009
Triplophyllum funestum (Kunze) Holltum, Kew Bull. 41(2): 255. 1986.
Triplophyllum glabrum J. Prado & R. C. Moran, Brittonia 60(1): 121, fig. 13, 14. 2008.
Studied specimen(s): Zuquim & Costa 284 (INPA); Zuquim & Vieira 403 (INPA).
Distribution: Venezuela, Guiana, and N Brazil.
Triplophyllum hirsutum (Holttum) J. Prado & R. C. Moran, Brittonia 60(1): 121, fig. 15,
16, 17. 2008.
Studied specimen(s): Zuquim & Figueiredo 76 (INPA); Zuquim & Costa 268 (INPA);
Zuquim & Vieira 298 (INPA).
Distribution: Mesoamerica, Antilles, and South America.
THELYPTERIDACEAE
Thelypteris abrupta (Desv.) Proctor, Rhodora 61(732): 305-306. 1959[1960].
(INPA); 149 (INPA); 175 (INPA); 176 (INPA); 192 (INPA); 381 (INPA).
Thelypteris arborescens (Humb. & Bonpl. ex Willd.) C. V. Morton, Contr. U.S. Natl.
Herb. 38: 50. 1967.
Habit/Habitat: herb, terrestrial, in Terra Firme and white-sand forests, near streams.
Thelypteris macrophylla (Kunze) C. V. Morton, Amer. Fern J. 61(1): 17. 1971.
Habit/Habitat: herb, terrestrial and rupiculous, in Terra Firme forests, near streams.
(INPA); Zuquim & Jakovac 335 (INPA).
Thelypteris opulenta (Kaulf.) Fosb., Smithsonian Contr. Bot. 8: 3-6. 1972.
Habit/Habitat: herb, terrestrial, in Terra Firme forests, in tree fall gaps or disturbed
areas.
Distribution: Antilles, Mesoamerica, N South America. Native to Asia and Africa.
Thelypteris poiteana (Bory) Proctor, Bull. Inst. Jamaica, Sci. Ser. 5: 63. 1953.
Habit/Habitat: herb, terrestrial or rupiculous, in Terra Firme forests over rich soils.
Distribution: S Mexico, Antilles, and N South America.
304
WOODSIACEAE
Diplazium cristatum (Desr.) Alston, J. Bot. 74: 173. 1936.
Studied specimen(s): Zuquim & Neres 371 (INPA).
Diplazium grandifolium (Sw.) Sw., J. Bot. (Schrader) 1800(2): 62. 1801.
Habit/Habitat: herb, terrestrial or hemi-epiphytic, in Terra Firme forest over rich soils.
Distribution: Mexico, Antilles, Mesoamerica, and N South America.
DISCUSSION
The vegetation is heterogeneous in the Rebio Uatumã region; physiognomies vary from
the open areas of campinas to the dense Terra Firme forests with closed canopy and
emergent trees more than 45 m tall. Within the Terra Firme forests heterogeneity arises
from variations in edaphic factors such as fertility (exchangeable bases content varies,
at least, from 0.16 to 5.68 cmol/kg), texture (clay content varies from 16 to 85 %) and
water availability (from stream margins to high plateaus). The environmental
heterogeneity, including a large edaphic gradient, probably determined the high species
richness (122 spp.) found in the area in comparison with previous inventories made in
the Brazilian Amazon: 83 species in the North of Reserva Ducke (Costa et al., 1999);
48 species in upper Rio Negro (Freitas & Prado, 2005); - 39 species in Madeira-Purus
interfluve (Carvalho et al., 2007); 60 species in DBFF Project areas (Zuquim, 2006); 70 species in Eastern Amazon (Costa & Pietrobom, 2007); 39 species in the
metropolitan region of the city of Belém (Maciel et al., 2007); and 50 species in
northern Amazon (Edwards, 1998).
In fact, ferns and lycophytes have been long recognised as habitat specialists in the
Amazon (Tuomisto & Poulsen, 1996). In the present work, we found some genera that
are not common in Central Amazonia such as Bolbitis and Tectaria. The species of these
genera tend to colonise relatively rich soils (Tuomisto & Poulsen, 1996). On the other
hand, some genera such as Lindsaea, that tends to be more diverse in poor soils
(Tuomisto, 1998), also showed high number of species. High environmental
heterogeneity itself tends to increase species richness for virtually every plant group. To
our knowledge, Prado & Moran (2009) is the only checklist that found a comparative
number of species (178 spp.). Their study comprises almost all the State of Acre and
probably encompasses areas with rich soils since the checklist contains rich soil
indicator species, such as Adiantum pulverulentum, Bolbitis lindgii, B. nicotianifolia,
Didymochlaena truncatula, Tectaria incisa etc (Tuomisto & Poulsen, 1996).
Ferns and lycophytes are sensitive to changes in environmental conditions and land
uses (Paciencia & Prado, 2005). The strong species-environment relationship and the
diversity found in Uatumã region and in previous studies in Amazonia (Costa et al.,
2005, Tuomisto et al., 2002, Tuomisto et al., 2003, and Zuquim et al., 2009) reinforce
the proposal to include ferns and lycophytes as indicator groups of the distribution of
biodiversity in Rapid inventories (RAP). RAP methodology is being applied widely in
Amazonia by NGOs (e.g. World Wild Fund for Nature - WWF, Fundação Vitória
Amazônica - FVA) and governmental institutions (Secretaria de Desenvolvimento
Sustentável do Estado do Amazonas – SDS/AM, Instituto Nacional de Pesquisas da
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FERN GAZ. 18(6): 286-306. 2009
Amazônia - INPA). Fern and lycophyte inventories would enhance the knowledge of
the spatial distribution of diversity and heterogeneity of a given area, providing lowcost information for conservation planning and management.
ACKNOWLEDGEMENTS
We acknowledge the assistance of Joelson Nogueira and Ailton Neres during field
work. The ICMBio staff (especially Caio Pamplona) made it easier to work in REBIO
Uatumã. We thank Carlos H. Franciscon for herbarium facilities at INPA and Hanna
Tuomisto (TUR) for taxonomic and ecological discussions. This work was funded by
CNPq/PPG7.
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will be refereed
aligned.
aligned.
a legible form.
Supplements.
FERN GAZ. 18(7):308-318. 2010
308
THE COLONISATION OF WOODLAND GAPS BY FERNS AND
HORSETAILS
P. BREMER
Wageningen University, Plant Ecology and Nature Conservation, Wageningen,
The Netherlands
(E-mail: [email protected])
Key words: woodland gaps, Voorsterbos, boulder clay, ferns, spore bank,
Equisetum
ABSTRACT
In the Voorsterbos, a planted woodland on a former sea-floor (the Netherlands),
artificial gaps within stands of Fagus sylvatica on boulder clay were monitored
for five or six years after cutting. Ten fern species and three species of horsetail
established in these gaps, with Dryopteris cristata, Thelypteris palustris,
Matteuccia struthiopteris and Equisetum telmateia not previously known from
the Voorsterbos. Athyrium filix-femina was most successful and formed dense
stands in some gaps. Two species (Gymnocarpium dryopteris, Dryopteris
cristata) established but have since disappeared. Thelypteris palustris has built
up a sustainable population. It seems plausible that the germination of ferns and
horsetails took place during the first season following cutting of the gaps and
was derived from a spore bank with an optimal microclimate with constant high
moisture and humidity in the shaded part of the gaps. Competition from
recruiting trees and shrubs was suppressed in some of the gaps for a period of
five year through herbivory by Roe deer.
INTRODUCTION
Woodlands are the climax vegetation over much of the Earth and do not need any
human intervention to be sustained. As humans need timber and fibre, they manage
woodlands and harvest boles. Numerous silvicultural options are available to forest
managers. They include a wide range of harvesting systems, regeneration methods and
rotation lengths. Various studies have been published on the effects of harvest practices
on understory and overstory conditions in managed temperate forest (e.g. Halpern &
Spies 1995, Thysell & Carey 2000, Bergstedt & Milberg 2001), but with the exception
of one study on Pteridium aquilinum (den Ouden 2000), none of them focussed on
ferns.
In 1997 the management objective in one of the planted forest in the province of
Flevoland (Voorsterbos) changed from timber production to ecological forestry.
Homogeneous thinning with a short return interval was banned and replaced by
heterogeneous thinning, artificial gap formation and longer periods of no management,
in order to increase the variation within the stands and to convert single cohort stands
into stands with a mosaic of phases (Koop 1989, Koop & Siebel 1993, van den Burgh
et al. 1995, Hunter 1999). The first series of gaps in the Voorsterbos offered an
opportunity for studying the effect of these gaps on the herb layer. We expected gaps
would be vegetated by recruiting trees while ferns would play a subordinate role as they
had been scarce in the beech stands prior to the gap formation. Natural gaps in beech
309
FERN GAZ. 18(7): 308-318. 2010
stands on sandy soil in the Speulderbos (The Veluwe, the Netherlands) showed hardly
any role for ferns (two species with little cover) while rejuvenation of trees was
suppressed by browsing of deer (personal observations). In the Neuenburger Urwald, an
ancient Hornbeam woodland on loamy soil (Stellario – Carpinetum), Koop (1989)
recorded few fern species colonising the natural gaps.
STUDY SITE
The Noordoostpolder was reclaimed in 1941/1942 and the Voorsterbos (Figure 1)
planted in the period 1944 -1955 (Bremer 2001). The area is below mean sea level: the
altitude varies from -2.5 to -1m. The Voorsterbos has been planted on sand and boulder
clay. Here, Quercus robur dominates (48% of the area), while other tree species are less
important: Pinus spp. (12%), Fagus sylvatica (8%) and Fraxinus excelsior (7%)
(Bremer 2001). Drainage trenches were dug at a density of up to 1 km per ha in the
Voorsterbos (boulder clay); their depth ranges from 0.4 m to more than 1.2m. A large
area on the boulder clay has a water table at a depth of less than 0.5m in the winter.
Currently the trees extract much water, making drainage by trenches less important than
in the years after these woodlands were planted. In winter these trenches have a shallow
water level, while in summer they are dry, only being filled after heavy rainfall. In the
Voorsterbos a management strategy started in 1997, aiming to produce a shifting mosaic
by making artificial gaps in various stands (Koop & Siebel 1993). In the period 2000 2006, 54 gaps were created by cutting down trees. The diameter of these gaps was 1.5
- 2 times tree height, and their size varied from 0.08 to 0.15ha per gap, with the
exception of one gap of 0.25ha (Bremer 2007a). Nearly all the felled trees were
transported with heavy equipment, causing soil disturbance over more than 50% of the
surface of these gaps. All the gaps were intersected with 0.4 – 1m deep drainage
trenches with a density of 1km per ha.
METHODS
This study focuses on 12 gaps in Fagus sylvatica stands on boulder clay in which trees
were felled at the start of the years 2000, 2001 or 2004; monitoring took place in the
Figure 1. The location of the study area (Voorsterbos) in the Netherlands.
BREMER: THE COLONISATION OF WOODLAND GAPS BY FERNS
310
period 2000 – 2008. Each gap was monitored for a period of six years, except for three
plots with a five year period. In these gaps the vegetation and recruitment of tree species
were mapped, and ferns and horsetails recorded. For those ferns and horsetails that are
less common in the Netherlands (listed in Table 1) individuals were mapped on 1:1000
maps, and the length of the longest frond per plant measured and the presence of spore
producing fronds noted. Special attention was paid to trenches, as this habitat can be
rich in fern species (Bremer 2007b). In the second year following tree felling, a 50m
zone surrounding the gaps was surveyed for ferns and horsetails. These `plots` were
used as controls. Shade by the trees surrounding the gaps was mapped on or near 21
June (with largest area of gap floor receiving direct light) and 21 September (the end of
the astronomical summer, approximately the end of the growing season) during more
or less cloudless weather. The shade line of 21 June is defined as the 100% shade line
(the part of the gap south of this line is always in shade), while the shade line of 21
September is defined as the 0% shade (the part of the gap north of this line has no shade
in the period between 21 June and 21 September). The position of each individual fern
and horsetail (n = 25) was recorded with reference to these two lines and the number of
days with shade for each was determined (Figure 2). For the group of common fern
species only Athyrium filix-femina was analysed because it is the species most prone to
desiccation. This species was analysed using a matrix of 1 x 1m plots in one of the gaps
when this gap was five years old and ferns in their subadult or adult phase. Only
sporophytes were included in this study, as gametophytes are difficult to monitor. The
gametophyte phase was frequent in the first year after tree felling and subsequent
sporophytes could be identified when two or three years old. Data were collected in
August. Data on cover of the fern layer, herb layer (< 1m) and young shrub layer (> 1m)
were estimated per gap in late summer.
Statistical analysis
A chi-square test was used to analyse habitat preference. A Kruskall–Wallis test was
used to analyse a gradient in density of Athyrium filix-femina in one of the gaps.
RESULTS
Within three years the gaps were dominated by the herb or initial tree layer (< 1m in
height). Fern cover reached a maximum of 21% in the fifth year (Figure 3) and varied
from 1– 70% per gap. Fraxinus excelsior recruited at a density of more than one sapling
per m2 in 50% of the monitored gaps. In these gaps, after five years, patches of trees
were able to build up a young woodland phase despite heavy browsing by Roe deer
(Capreolus capreolus). In three other gaps shrubs and young trees present before felling
were able to cover these gaps within five years. Ten fern species became established in
the gaps, of which five were present in adjacent woodland zones, and three species of
horsetail (Table 1). The average number of fern species per gap was at its maximum in
the early years and decreased slightly after three years (Figure 4). Three fern species
(Thelypteris palustris, Dryopteris cristata, Matteuccia struthiopteris) and one species
of horsetail (Equisetum telmateia), had not previously been recorded in the woodland.
A single plant of Asplenium scolopendrium was found adjacent to one of the gaps; it
was not recorded within the gap habitat.
Within the gaps, ferns established on trench banks and on the woodland floor.
Athyrium filix-femina, the most successful fern coloniser, had its highest density in the
trenches (χ² = 6.5, p < 0.05). The group of less common fern species (Table 1) showed
no preference for trench or woodland floor habitat in terms of the population size (χ² =
Woodland species
Ferns
Woodland gaps
%
np g
ng
tr
%
wfl
%
f
50 m zone
sh % a
%
np a
311
Table 1. Fern species found within 12 gaps of Fagus sylvatica stands (5 – 6 years after tree-felling) compared with uncut woodland surrounding
these gaps, based on annual monitoring).
n g = number of gaps with species listed, np g = total number of estimated or counted plants/colonies in 12 gaps after six seasons, tr = number
of gaps with species listed growing in drainage trenches, wfl = number of gaps with species listed growing on the woodland floor, f = fertility
of plants after establishing in the gaps; f = with fertile fronds within four years, n.f = not fertile within four years, sh % = average time in shade
during the period 21 June - 21 September based on plants/colonies and presented as percentage (e.g. 100% means plants are shaded during the
whole season within the 21 June shade line), na = not analysed, a = number of 50m zones surrounding gaps with species listed, np a = total
number of plants in 50m zones surrounding the gaps (data of all these gaps summed) after two seasons, VB = species found elsewhere in the
Voorsterbos since 2000.
VB
12
11
8
6
100
91.7
66.7
50.0
> 1000
200-400
200-400
50-100
12
11
8
6
100
91.7
66.7
50.0
12
11
8
5
100
91.7
66.7
41.5
f
f
f
f
na
na
na
na
8
8
8
3
66.7
66.7
66.7
25.0
50 - 100
50 - 100
50 - 100
25 - 50
+
+
+
+
Less common species
Thelypteris palustris
Dryopteris affinis
Matteuccia struthiopteris
Dryopteris cristata
Gymnocarpium dryopteris
Polystichum aculeatum
Asplenium scolopendrium
7
3
2
2
1
1
0
58.3
16.6
16.7
16.7
8.3
8.3
0
13
3
2
2
1
1
0
3
1
1
2
1
0
0
25.0
8.3
8.3
16.7
8.3
0
0
5
2
1
0
0
1
0
41.5
16.7
8.3
0
0
8.3
0
n.f
n.f
n.f
n.f
n.f
n.f
-
84.6
100
70.0
75.0
50.0
100
-
0
1
0
0
0
0
1
0
8.3
0
0
0
0
8.3
0
1
0
0
0
0
1
+
+
+
+
Horsetails
Common species
Equisetum arvense
Equisetum fluviatile
4
2
33.3
16.7
?
3
0
0
0
0
4
3
100
100
f
f
na
100
0
0
0
0
0
0
+
+
Less common species
Equisetum telmateia
1
8.3
2
0
0
2
100
f
100
0
0
0
-
FERN GAZ. 18(7): 308-318. 2010
Common species
Athyrium filix-femina
Dryopteris dilatata
Dryopteris filix-mas
Dryopteris carthusiana
312
Figure 2. Woodland gap within Fagus sylvatica stand showing the shade lines and the distribution of rare ferns and horsetails.
313
based on nine gaps (years 1-5 based on 12 gaps). The x-axis gives the number of growing seasons after gap formation.
FERN GAZ. 18(7): 308-318. 2010
Figure 3. Change in cover layers (average values) during 7 years in woodland gaps in beech stands on boulder clay. The layer of < 1 m
comprises the herb layer including recruiting trees, the layer > 1 m comprises the initial tree layer and shrub layer. Data for the sixth year are
314
Figure 4. The average number of fern species found in the woodland gaps each year from the year before gap formation (year = 0) to six years
after gap formation. Data for the sixth year are based on nine gaps (instead of 12).
315
FERN GAZ. 18(7): 308-318. 2010
0.5 n.s), but when densities were corrected for the area of both habitats there was a
preference for the drainage trench habitat (χ² = 12.7 p < 0.001). Plants of Thelypteris
palustris showed no preference, while Equisetum telmateia was found only on the
woodland floor. Fertility within four years after establishment was observed only in the
group of the most common species (present in more than 49% of the gaps). Some plants
of Athyrium filix-femina reached the adult phase in the third season after cutting and
became co-dominant in five out of the 12 gaps, growing alongside horsetails, grasses
and bryophytes.
The establishment of ferns was most successful in the southern parts of the gaps,
within the 100% shade line: Thelypteris palustris, Dryopteris cristata, Dryopteris
affinis and Polystichum aculeatum were found almost exclusively within this line.
Gymnocarpium dryopteris and Matteuccia struthiopteris were found within the 75%
shade line (Figure 2) and Athyrium filix-femina reached its highest density in this zone
(χ² = 8.3 p < 0.01 n = 12). Within the largest gap (0.25 ha) the average herb layer cover
in the fifth year was 92%, with Athyrium filix-femina contributing 17%. In this gap, a
gradient in density of Athyrium filix-femina could be correlated with the period of shade
in the summer period (χ² = 10.8 p < 0.01). Three species of Equisetum became
established: Equisetum arvense was co-dominant with Athyrium filix-femina in three
gaps, Equisetum telmateia colonised neutral boulder clay in one gap, while Equisetum
fluviatile established within the 100% shade line in two gaps. Forty-two gaps were
created in other stand and soil types. Sandy soils with low pH were poor in fern species.
Dryopteris dilatata was present prior to tree felling and increased afterwards. In a stand
of Carpinus betulus on boulder clay, gaps were created by bark ringing trees. Although
this led to a gradual increase of light intensity, no ferns or horsetails established in these
gaps, but young trees, already present in high density prior to the girdling, formed a
dense stand.
DISCUSSION
Artificial gaps on boulder clay were studied in the Voorsterbos. Recruiting trees were
expected to form a dense juvenile stand, contributing to the horizontal and vertical
structure of the woodland. We had expected that ferns would not expand as they had
been scarce prior to the gap formation, yet they proved to be successful both in cover
and number of species. On boulder clay, Fraxinus excelsior recruits successfully, but
the >1m saplings were intensely browsed by Roe Deer in the first five years, preventing
a dense growth and facilitating vegetation with local dominance of Athyrium filixfemina at the southern parts of gaps. Other ferns were also present from the outset with,
e.g., Athyrium filix-femina outcompeting Gymnocarpium dryopteris. On the boulder
clay, ten ferns and three species of horsetail established; this included species not
previously recorded in the area. From records of young sporophytes it appears that all
these species established from spores in the first year following tree felling and soil
disturbance. Spores of these species may have been derived from dry and wet spore rain
to colonise the disturbed areas. However, there was a high density (establishment of
>50 plants per m2) of Athyrium filix-femina in five out of 12 gaps. As Athyrium filixfemina grew only at a low density prior to tree felling there may have been an extensive
accumulation of spores preserved in the spore bank. Viable spores of Athyrium filixfemina were reported to be the most frequent type of fern spore in the soil spore bank
in a boreal old-growth spruce forest, yet there were no plants in the herb layer (Rydgren
& Hestmark 1996). In contrast in the Kuinderbos fern colonisation by numerous fern
316
species (with no extensive spore bank) always started with one fern individual at a
particular site, followed in subsequent years sometimes by mass colonisation in the
spore shadow of adult plants (Bremer 2007).
In the group of less common fern and horsetail species, the almost simultaneous
colonisation within two years suggests that the spore bank may play a dominant role
rather than spore rain; the spore bank here has had c. 60 years to accumulate. There
appears to be no relationship between the burst of colonisation in the new gaps and the
fern diversity of surrounding populations. This contrasts with the long term study (50
years) of fern colonisation in the Kuinderbos (also on a former sea floor), where Bremer
(2007b) showed that colonisation of 25 fern species was correlated with the population
size within 50 km from the forest.
Gymnocarpium dryopteris, Matteuccia struthiopteris, Dryopteris cristata and
Thelypteris palustris had not previously been recorded from boulder clay in this kind of
habitat within the Netherlands. All three species were able to colonise the bare soils, and
thrive until eliminated by competition. A similar observation was made for a group of
seed plants in the gaps, with establishment of pioneer species (e.g. Puccinellia distans,
Juncus gerardii). These plants established on the boulder clay between the period of
reclamation of the sea floor (1942) and afforestation (c. 1948) and must have survived
the period 1948 – c. 2000 through a seed bank.
Gaps have a characteristic microclimate (Moore & Vankat 1986, Stoutjesdijk &
Barkman 1992). Gaps with a diameter of less than 0.7 times the tree height do not get
any direct sunlight, while in gaps with a diameter equal to the tree height, c. 80% of the
gap area is shaded when the foliage is fully developed. Gaps with a diameter 1 - 2 times
the tree height deviate most from the adjacent woodland, having a microclimate that is
more Atlantic in summer and more continental in winter (Stoutjesdijk & Barkman
1992). As rainwater is not intercepted by the canopy in these gaps and without tree
transpiration, the soil in these gaps becomes constantly wet, with on average a low
saturation deficit during the growing season, while in winter there is more radiation. In
the Voorsterbos, the diameter of most gaps is 1-2 times the tree height which is an
optimal habitat for many terrestrial woodland ferns and horsetails. The dominance of
Athyrium filix-femina is related to soil conditions (boulder clay with reduced
percolation, moisture and neutral substrate). Godefroid et al. (2006) show an effect of
light intensity but also a combined effect of the air temperature and air humidity on the
occurrence of this species in clear cuts. The results of this study show that the less
common species were confined mostly within the 100% shade line; this indicates that
they prefer a constantly wet soil and high air humidity. Comparing gaps formed through
tree-felling with those developed through bark-ringing demonstrates that cutting and
concomitant soil disturbance is important for the germination of fern spores. Equisetum
fluviatile, E. telmateia and to some extent Matteuccia struthiopteris are indicators of
seepage (e.g. Grootjans 1985), indicating a flux of pH neutral,groundwater. That these
species were found growing on impermeable boulder clay shows that seepage is not the
dominant factor, but that a continuously moist, pH neutral habitat is needed, which can
also be achieved by rainwater stagnating on pH neutral boulder clay. Gaps on sandy
soils do not facilitate the growth of ferns, as the water percolates to the water table.
When constant high humidity can be assured, one might expect species such as
Blechnum spicant, Osmunda regalis and Oreopteris limbosperma to establish.
Matteuccia struthiopteris is a common garden plant in the Netherlands, and occurrences
of M. struthiopteris outwith gardens are nearly always related to garden rubbish
317
FERN GAZ. 18(7): 308-318. 2010
deposited at the edge of woodlands by man. The species established for the first time
naturally in one of the planted woodlands of E.-Flevoland (province of Flevoland) in
1988 and still grows there. In the Voorsterbos it was recorded for the first time on
boulder clay; a habitat not known from neighbouring countries (Bremer 2005).
In the woodland experiment, felling trees in Fagus sylvatica stands on boulder clay
in order to make gaps facilitated the establishment and expansion of fern and horsetail
populations with large differences in microclimate between gaps and the adjacent
woodland. It seems likely that the crucial factors are soil composition (weakly acid - pH
neutral boulder clay), moisture, and the light regime. Creating woodland gaps allows
ferns to gain from their anemochoric dispersal capacity and the cumulative effect of
spore dispersal (i.e. spore bank) and from the ability to develop prior to the phase in
which seed plants might become dominant. Ferns, horsetails, bryophytes and liverworts
are the first to establish before seed plants become dominant. Cutting woodland gaps
on soils prone to desiccation would not be beneficial to the fern flora.
ACKNOWLEDGMENTS
Natuurmonumenten (beheerseenheid Voorsterbos/ Zwarte meer) created the woodland
gaps. I thank Jan Akkerman, Klaas Althuis, Dick Buitenhuis and Lykele Zwanenburg
for their cooperation. The analysis of data was financed by Natuurmonumenten. I thank
Prof. Dr. F. Berendse for his comment on and Dr. J. Burrough for improving the English
text. An anonymous reviewer provided helpful suggestions on the manuscript.
REFERENCES
BERGSTEDT, J. & MILBERG, P. 2001. The impact of logging intensity on field-layer
vegetation in Swedish boreal forests. Forest Ecology and Management 154: 105
-115.
BREMER, P. 2001. Flora en vegetatie in het Voorsterbos. Rapport Natuurmonumenten.
BREMER, P. 2005. Is er sprake van inburgering van de Struisvaren (Matteucia
struthiopteris (L.) Tod.) in Nederland? Gorteria 31: 122 – 124.
BREMER, P. 2007. The colonisation of a former sea-floor by ferns. Thesis,
Wageningen University, Wageningen.
BURGH, F. VAN DEN, VAN DER MOLEN, A. & KOOP, H. 1995. Mozaïekmethode.
Omvorming naar meer natuurlijk bos. Nieuwland Advies.
GROOTJANS, A. 1985. Changes of groundwater regime in wet meadows. Thesis,
University of Groningen, Groningen.
GODEFROID, S., RUCQUOIJ, S. & KOEDAM, N. 2006. Spatial variability of
summer microclimates and plant species response along transects within clearcuts
in a beech forest. Plant Ecology 185: 107 - 121.
HALPERN, C. & SPIES, T. 1995. Plant species diversity in natural and managed
forests of the Pacific Northwest. Ecological Applications 5: 913 - 934.
HUNTER, M.L. 1999. Maintaining Biodiversity in Forest ecosystems. Cambridge
University Press
KOOP, H. 1989. Forest Dynamics. Silvi-Star: A Comprehensive Monitoring System.
Springer Verlag. Berlin Heidelberg. Thesis.
KOOP, H. & SIEBEL, H. 1993. Conversion management towards more natural forests:
evaluation and recommendations. In: Broekmeyer, M.E.A., W. Vos & H. Koop
(eds.), 1993. European Forest Reserves. Proceedings of the Forest Reserves
Workshop. PUDOC-DLO. Wageningen; 199 - 204.
318
MOORE, M.R. & VANKAT, J.L. 1986. Responses of the herb layer to the gap
dynamics of a mature Beech-maple forest. The American Midland Naturalist
115(2): 336 - 347.
OUDEN, J. DEN 2000. The role of bracken (Pteridium aquilinum) in forest dynamics.
Thesis, Wageningen University, Wageningen,
RYDGREN, K. & HESTMARK, G. 1996. The soil propagule bank in a boreal
old-growth spruce forest: changes with depth and relationship to aboveground
vegetation. Canadian Journal of Botany 75: 121 - 128.
STOUTJESDIJK, PH. & BARKMAN, J.J. 1992. Microclimate, Vegetation and Fauna.
Opulus Press AB, Uppsala.
THYSELL, D.R. & CAREY, A.B. 2000. Effects of Forest Management on Understory
and Overstory Vegetation: A Retrospective Study. United States Department of
Agriculture. Forest Service.
319
FERN GAZ. 18(7). 2010
BOOK REVIEW
FLORE DES MASCAREIGNES, LA REUNION, MAURICE (MAURITIUS),
RODRIGUES. PTERIDOPHYTES, 1.PSILOTACEES A 26. MARSILEACEES.
Published by Institut de Recherché pour le Developpement, Mauritius Sugar
Industry Research Institute and Royal Botanic Gardens (Kew), Paris 2008.
Pteridophytes by F. Badre. Pp. 452, 69 full page black and white diagnostic plates, 4
pp. col. Plates. 240 x 170 mm. Available from RBG Kew, UK price unspecified but
European price is 40 euro. ISBN: 978-2-7099-1647-9.
Floras of far off tropical islands are always interesting but given that the BPS held a full
excursion to La Reunion only two years ago this flora is one of the most welcome new
publications in recent years. How unfortunate it could not have been published three
years earlier!
Floras of these islands seem to be few and far between. Apart from J. G. Baker,
Flora of Mauritius and Seychelles: Description of flowering plants and ferns of those
islands, 1877, and Jacob de Cordemoy, Flore de l`isle de La Reunion. Fascicule 1:
Cryptogames vasculaires, 1891, no complete fern catalogues seem to have been
published. Although about 25% of the fern flora is endemic to the three islands many
of the remaining taxa occur in Madagascar where floras by Tardieu-Blot and others give
good coverage. The current volume would have been improved by a full history of fern
research on the islands and a full fern bibliography. For a good review of all botanical
research over the three islands see Botanical History of the Mascarene Islands by
Wendy Strahm in Curtis`s Botanical Magazine, Vol.13, Part 4, November 1996, pp.
217-228.
The format here is concise with a very brief introduction on the classification of
ferns, a key to families, then under each family a key to genera with species keys under
each genus. In the body of the book the genera follow a standard systematic order with
the lycophytes included at the front. There is a glossary at the end and finally a very
comprehensive 38 page index. The text is in French throughout.
Each species entry gives full synonymy with authors, first publication, full
description, plate number and page (if illustrated) and full distribution on the three
islands. The authorship of each family varies. F. Badre is the major author but here and
there specialist authors have written chapters. For example, the treatment of the
Cyatheaceae by T. Janssen is very comprehensive, much of it based on his recent
research, as is the treatment of Lomariopsidaceae by D. Lorence and G. Rouhan. Other
major contributors include the late Prof. Holttum, the late M.-L. Tardieu-Blot (proof of
the long gestation period of this volume) and J. P. Roux.
Happily I can find no discrepancies between the full species list given here and that
prepared by Edmond Grangaud for the BPS excursion. I can only conclude that this
volume is an accurate, concise and very welcome addition to the world`s fern floras.
This volume is part of a multi-volume set covering the complete flora of these
islands; so far 25 parts have been published since the first part appeared in 1976.
Currently still in production are volumes on Cyperaceae, Orchidaceae and Gramineae.
Martin Rickard
FERN GAZ. 18(7):320-327. 2010
320
CYATHEA SLEDGEI (CYATHEACEAE): A NEW SPECIES OF
TREE-FERN FROM SRI LANKA
R.H.G. RANIL1, D.K.N.G. PUSHPAKUMARA2, T. JANSSEN3, C.R.
FRASER-JENKINS4 and D.S.A. WIJESUNDARA5
1
Postgraduate Institute of Agriculture, University of Peradeniya, Peradeniya, Sri
Lanka (e-mail: [email protected])
2
Department of Crop Science, Faculty of Agriculture, University of Peradeniya,
Peradeniya, Sri Lanka
3
Department of Botany, Senckenberg Research Institute, Frankfurt, Germany
4
Student Guest House, Thamel, P.O. Box no 5555, Kathmandu, Nepal
(e-mail: [email protected])
5
Department of National Botanical Gardens, Peradeniya, Sri Lanka
Key words: Tree-fern, new species, Cyathea sledgei, hybridisation, Sri Lanka.
ABSTRACT
Cyathea sledgei Ranil, Pushpakumara & Fras.-Jenk. is a new tree-fern species
from Sri Lanka, morphologically intermediate between C. sinuata Hook. &
Grev. and C. hookeri Thwaites. It is easily recognised by its simple but deeply
lobed leaves; each lobe has 8-10 pairs of forked veins and 6-9 pairs of sori. It
was first recorded as a putative hybrid and the origin of this species is postulated
to have been directly through hybridisation between the above two species. It
behaves as a species, readily forming populations and apparently being fertile.
C. sledgei occurs among populations of the two other species, C. hookeri being
similarly restricted in occurrence. All three species are endemic to Sri Lanka,
though Large & Braggins (2004) misreported C. sinuata from South India.
INTRODUCTION
Interspecific and, according to some authors, inter-generic hybridisation is not
uncommon in tree-ferns and has been reported by a number of authors (Conant, 1975;
Tryon, 1976; Conant & Cooper-Driver, 1980; Holttum, 1984; Caluff, 2002; Caluff &
Serrano, 2002; Janssen & Rakotondrainibe, 2007). Tree-fern hybrids have been
reported mainly in the Cyatheaceae, but also in Dicksoniaceae (Large & Braggins,
2004, where a wider sense of the term “tree-fern” is employed, which we do not follow
here). Evidence for hybridisation in the Cyatheaceae was first presented by Holttum and
Sen (1961) from Malaya and from Sumatra and Borneo by Conant and Cooper-Driver
(1980). Tryon (1976) published a list of ten putative hybrids from the West Indies,
Mexico and Central America. Holttum (1981) and Conant et al. (1996) have reported
hybrids from Brazil, Venezuela and Colombia. Caluff (2002) and Caluff and Serrano
(2002) have provided evidence for the occurrence of four natural hybrids from Cuba.
Large and Braggins (2004) tentatively reported hybrids from Australia, Tasmania, New
Zealand and the Philippines and a single case has been reported from mainland Africa
by Edwards (2005). A recent work by Janssen and Rakotondrainibe (2007) reported
three putative hybrids from Madagascar.
However Sri Lanka is the only country in South Asia from which a putative tree-fern
hybrid has been reported (Sledge, 1982). The taxon concerned was first found in
321
FERN GAZ. 18(7): 320-327. 2010
January 1951 by the late Drs W.A. Sledge and F. Ballard as a sterile plant or plants,
labelled by Sledge as being collected in the Kanneliya Forest Reserve near Udugama,
and labelled by Ballard as from near Panangala in Kanneliya Forest Reserve, both
probably referring to the same locality and collection. From the distinctively
intermediate frond-morphology, Sledge postulated that it was not an immature Cyathea
hookeri, but represented a putative hybrid between C. sinuata and C. hookeri. Rather
surprisingly it was omitted by Philcox (2007) from the Revised Handbook of the Flora
of Ceylon, perhaps due to their being sterile collections and uncertainty as to its
taxonomic status. Fifty-three years after its original collection, more than 30 individuals
of the same taxon were rediscovered in Kanneliya forest reserve (Ranil et al., 2004). We
were subsequently able to find four mature plants with ripe spores amid populations of
this taxon at both Kanneliya Forest and a new locality at Sinharaja Forest.
The morphology of this taxon is constant and readily recognisable and though
intermediate between the other two species mentioned there is at present no actual
evidence, other than its morphological intermediacy, that it is a hybrid. This, combined
with its very probable fertility (see below) and apparent formation of populations,
suggests that it should be considered to be a species in its own right, even if its most
likely origin would seem to have been directly from F1 hybrids between the two
species. According to previous studies of Cyathea hybrids (see below) such plants may
be expected to be fertile immediately upon formation.
It is therefore described here as follows and should be added to the list of nonMalaysian species in Asia (Holttum 1965):
Cyathea sledgei Ranil, Pushpakumara & Fras.-Jenk., spec. nov. Filix morphologia
intermedia inter C. hookeri et C. sinuatam. Folia simplicia angusta profunde lobata vel
pinnatifida. Sporae bene evolutae non abortivae. (Plate 1 C, F, I; Plate 2 C, D, E, F).
Etymology: This species is named after the late Dr. W. Arthur Sledge (1904 - 1991), of
the Botany Dept., University of Leeds, UK, to honour his immense contribution to
knowledge of the Sri Lankan Pteridophyte flora.
Type: Sri Lanka, Sabaragamuwa Province, Ratnapura District: Sinharaja forest, 380 m
alt. R.H.G. Ranil 314, 28 July 2008 (3 sheets), (holotype: PDA (314A); isotypes: PDA
(314B, 314C).
Trunk erect, unbranched, 90-135 cm in height, 1.2-1.7cm in diameter, entirely covered
by persistent dead petiole bases, leaf scars not visible, usually with few adventitious
shoots. Crown 112-168 cm in width and 18-28 cm height, with 18-26 leaves crowded
at the apex of the trunk. Petiole (stipe) dark brown, glossy, 4.5-4.7 cm long, 0.3-0.5 cm
in diameter at its base, with blunt, 0.05-0.1 cm long squaminate spines. Petiole-scales
0.4-0.6 cm long, 0.1 cm wide at their base, dark brown, linear to narrowly triangular,
with an apical spine, margin sometimes bearing several conspicuous dark setae, scales
densely covering the young leaves. Lamina simple, pinnatifid about halfway to the
rachis, 60-90 cm long, 3.7-5.1 cm wide, linear, light green adaxially, olive green
abaxially, subcoriaceous to coriaceous. Lobes short, obtuse with a crenate apex, veins
free, once forked, 8-10 pairs in each lobe. Lamina-apex gradually reduced to a serrate
margin. Lamina-base gradually reduced, the lowest lobes deeply cut almost to the
rachis. Rachis grooved and dark brown adaxially, light brown abaxially, glabrous and
RANIL et al.: CYATHEA SLEDGEI
322
glossy. Sori round, about 0.1 cm in diameter, borne at the forks of the veins, 6-9 pairs
in each lobe in two rows on either side of the mid-vein, indusiate. Indusia globose, light
brown, very thin, membranous, only a reduced disc persisting around the base of the
receptacle at maturity. Spores well-formed, trilete, regular and not showing much
variation in size or shape.
Distribution: Endemic to Sri Lanka. This species is known so far only from the
Sinharaja forest in Ratnapura district (Sabaragamuwa Province) and Kanneliya forest in
Galle district (Southern Province), occuring in both places along with its presumed
ancestral species, C. sinuata and C. hookeri.
Habitat: Stream-side or road-side banks in lowland rain forests, 320 - 450 m alt.
Specimens examined: Sri Lanka. Sabaragamuwa Province, Ratnapura district,
Sinharaja forest. R.H.G. Ranil 314, 28 July 2008 (6 sheets) (PDA); idem. R.H.G. Ranil
315, 29 July 2008 (5 sheets) (PDA); idem. R.H.G. Ranil 319, 29 July 2008 (PDA); idem.
R.H.G. Ranil 346, 18 Sep 2008 (PDA); idem. R.H.G. Ranil 347, 18 Sep 2008 (2 sheets)
(PDA). Southern Province, Galle district, Kanneliya forest reserve, near Udugama.
R.H.G. Ranil 117, 118, 119, 120, 05 Dec 2003 (PDA); idem. R.H.G. Ranil 005R, 22
March 2007 (PDA); W.A. Sledge 908, 21 Jan. 1951 (K); Panangala. F. Ballard 1548, 21
Jan. 1951 (K).
Note: Both presumed ancestral species are endemic to Sri Lanka. C. sinuata was listed
by Large & Braggins (2004) as occurring in South India as well, but in error as it is not
known from there. C. sinuata (Plate 1 A, D, G; Plate 2 A) and C. hookeri (Plate 1 B, E,
H; Plate 2 B) are limited to a few isolated pockets in the two main lowland rain forests,
Kanneliya and Sinharaja Forests. C. sinuata is the only Cyathea species known
worldwide with simple leaves (Kramer, 1990) and is fairly common in its places of
occurrence. The intermediate morphology (Table 1) of C. sledgei between a species
with long, narrow, simple, strap-like fronds and another with pinnate fronds makes it
very distinctive and easy to recognise. In contrast to C. sinuata, C. hookeri is very
limited in numbers in both localities. All three species occur in close proximity in the
same microhabitats. Sledge (1982) had mentioned previously that what he had assumed
to be both parents were observed in the immediate vicinity of his collection.
A couple of small sporophytes of C. sledgei transferred from the wild into
cultivation at Peradeniya soon grew normally and were able to produce spores. Spores
of this species were cultured and germinated on a medium of soil and produced prothalli
with both sexes present (Plate 2E and 2F), but did not produce sporophytes in the pots,
perhaps not having the correct stages or conditions to allow fertilisation in culture.
Cytological comments: Chromosome counts of C. sinuata and C. hookeri, have shown
2n = c. 130-140 and n = 69-70, respectively (Manton & Sledge, 1954) and similar
results were obtained by Abraham, Ninan & Mathew (1962) from Sri Lankan material.
C. sledgei has not yet been investigated cytologically and further study is intended to
be carried out at Peradeniya. All species that have been cytologically investigated so far
in Cyathea are sexual diploids with approximately n = 69 (Tryon 1970). Conant et al.
(1996) suggested that the family Cyatheaceae is unusual because it is the only fernfamily known showing such uniformity in chromosome number and is further unusual
323
FERN GAZ. 18(7): 320-327. 2010
A
C
B
D
E
G
a
a
a
b
H
b
F
a
I
a
Plate 1: Habit and frond characters of Cyathea sinuata, C. hookeri and C. sledgei (A)
Habit of Cyathea sinuata; (B) Habit of Cyathea hookeri; (C) Habit of Cyathea sledgei;
(D) Soral arrangement of Cyathea sinuata; (E) Soral arrangement of Cyathea hookeri;
(F) Soral arrangement Cyathea sledgei; (G) Venation of Cyathea sinuata (× 20); (H)
Venation of Cyathea hookeri (× 20); (I) Venation of Cyathea sledgei (×1).
Note: Rachis and secondary rachis represented by a and b, respectively.
324
among ferns due to the occurrence of apparently fertile diploid hybrids (or hybridderived species) with full bivalent formation at meiosis. Thus the cytology of some
hybrids recognised between the sections or genera Alsophila and Nephelea (which we
prefer to treat within Cyathea, following Holttum, in Holttum and Tryon, 1977, due to
their overall strong morphological similarity, in preference to recent molecular-based
classifications) is very unusual in ferns as a full complement of bivalents are formed
during meiosis between the two genomes present and viable spores are produced
(Conant & Cooper-Driver, 1980), which is not due to apomixis. Conant & CooperDriver (1980) reported that the new world hybrids, A. bryophila R.M.Tryon x C.
portoricensis Spring ex Kuhn, A. x dryopteroides (Maxon) R.M.Tryon (A. amintae
Conant x A. bryophila) and A. amintae x C. portoricensis have 69 bivalent
A
B
C
D
E
F
Plate 2: (A) Sori of Cyathea sinuata (x 40); (B) Sori of Cyathea hookeri (x 40); (C)
Sori of Cyathea sledgei (x 40); (D) Cyathea sledgei in ex situ condition; (E) Developed
gametophytes of Cyathea sledgei; (F) Gametophyte of Cyathea sledgei with
archegonia (x 40).
325
FERN GAZ. 18(7): 320-327. 2010
Table 1: Comparison of Cyathea sinuata, C. sledgei and C. hookeri.
Major characters
C. sinuata
C. sledgei
C. hookeri
Trunk height (cm)
70-170 (-200)
90-135
100-360
Trunk diameter (cm) 1.3-1.6
1.2-1.7
2.8-3.5
Crown width (cm)
78-114
112-168
145-195
Crown height (cm)
15-25
18-28
12-28
No. of leaves
20-32
18-26
7-13
Lamina dissection
Simple
Pinnatifid
Simply pinnate
Leaf shape
Narrowly Linear
Linear
Lanceolate
Leaf base
With sinuate or
entire margin
With deeply cut
lobes
With separate pinnae
Petiole length (cm) 3.8-4.2
4.5-4.7
7.5-11.5
Width and length of 0.05-0.1 x 0.3-0.4
petiole-scale (cm)
0.1 x 0.4-0.6
0.1 x 0.4-0.8
Density of
squaminate spines
on the petiole
Length of
squaminate spines
on the petiole (cm)
Length of lamina
(cm)
Absent or slight
Moderate
Dense
≤0.05
0.05-0.1
0.1
45-56
60-90
90-120
Width of lamina
(cm)
2.4-3.5
3.7-5.1
21-37.8
Veinlets
Simple
Forked
Simple
No. of pairs of sori
1-4
6-9
1-3
Soral arrangement
On 1-3 lower pairs
of secondary veins
At fork of tertiary
veins
On 1-3 lower pairs
of quaternary veins
326
chromosomes at meiosis. The usual cytological methods for detecting or confirming
hybridity from failure of chromosome-pairing at meiosis between the different genomes
and thus production of abortive spores have not been useful in this genus. Because
cytological investigation of this sort is apparently not useful to confirm hybridity in
Cyathea, the reproductive biology and gametophyte morphology of the presumed
ancestral species and C. sledgei are being studied further at Peradeniya and it is hoped
that molecular study can also be carried out in the future to help investigate the nature
and origin of C. sledgei. It is suggested that further consideration of the taxonomic
status of what have hitherto been reported as hybrids may be required world-wide if
such apparent hybrids are observed to form populations and behave as species. We
would treat such taxa as species, rather than hybrids, if they demonstrate a high degree
of fertility, form populations which apparently have the potential to spread and are
sufficiently distinct morphologically.
ACKNOWLEDGEMENTS
The authors thank the National Herbarium of Sri Lanka (PDA) for support and the
Forest Department of Sri Lanka for granting permission to carry out study at Sinharaja
and Kenneliya forests. The research was financed by the National Science Foundation
of Sri Lanka (Grant: RG/06/EB/05). We are thankful to Dr. Monika Shaffer-Fehre,
Royal Botanic Garden, Kew, UK (K), for providing photocopies of the first collections
of C. sledgei. CRFJ is grateful to the late Dr. W.A. Sledge for frequent encouragement,
guidance and help during his younger days.
REFERENCES
ABRAHAM, A., NINAN, C.A. & MATHEW, P.M. 1962. Studies on the cytology and
phylogeny of pteridophytes, 7. Observations on one hundred species of south Indian
ferns. J. Indian Bot. Soc. 41: 339-421.
CALUFF, M.G. 2002. A note on the Cuban tree fern hybrid Cyathea × calolepis
(Cyatheaceae) and its parentage. Willdenowia 32: 311-318.
CALUFF, M.G. & SERRANO, G.S. 2002. Cuban novelties in the genus Alsophila
(Cyatheaceae). Willdenowia 32: 303-309.
CONANT, D.S. 1975. Hybrids in American Cyatheaceae. Rhodora 77: 441-455.
CONANT, D.S. & COOPER-DRIVER, G. 1980. Autogamous allohomoploidy in
Alsophila and Nephelea (Cyatheaceae): a new hypothesis for speciation in
homoploid homosporous ferns. Amer. J. Bot. 67: 1269-1288.
CONANT, D.S., RAUBESON, L.A., ATTWOOD, D.K., PERERA, S., ZIMMER, E.A.,
SWEERE, J.A. & STEIN, D.B. 1996. Phylogenetic and evolutionary implications
of combined analysis of DNA and morphology in the Cyatheaceae. In: CAMUS,
J.M., GIBBY, M. and JOHNS, R.J. (eds.) Pteridology in Perspective: 132-248.
Royal Botanic Gardens, Kew.
EDWARDS, P.J. 2005. Cyatheaceae. In: BEENTJE, H.J. & GHAZANFAR, S.A. (eds.)
The Flora of Tropical East Africa: 1-15. Royal Botanic Gardens, Kew.
HOLTTUM, R.E. 1965. Tree-ferns of the genus Cyathea Sm. in Asia (excluding
Malaysia). Kew Bull. 19(3): 463-487.
HOLTTUM, R.E. 1981. The tree-ferns of Africa. Kew Bull. 36: 463-482.
HOLTTUM, R.E. 1984. A remarkable Cyathea hybrid. Amer. Fern J. 74: 19-21.
HOLTTUM, R.E. & SEN, U. 1961. Morphological classification of the tree ferns.
Phytomorphology 11: 406-420.
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HOLTTUM, R.E. & TRYON, R.M. 1977. An exchange of views on the Cyatheaceae.
Flora Malesiana Bull. 30.
JANSSEN, T. & RAKOTONDRAINIBE, F. 2007. An update of the revision of Cyathea
subgen. Alsophila sect. Gymnosphaera (Cyatheaceae) in Madagascar and the
Comoros including a discussion of putative hybridization events. Adansonia 29(2):
195-213.
KRAMER, K.U. 1990. Cyatheaceae. In: KRAMER, K.U. & GREEN, P.S. (eds.) in: K.
KUBITZKI (ed.) The families and genera of vascular plants. 1. Pteridophytes and
Gymnosperms: 69-74. Springer Verlag, Berlin, Heidelberg.
LARGE, M.F. & BRAGGINS, J.E. 2004. Tree Ferns: 1-359. Timber Press, Portland,
USA and Cambridge, UK.
MANTON, I. & SLEDGE, W.A. 1954. Observations on the cytology and taxonomy of
the pteridophyte flora of Ceylon. Philos. Trans. Roy. Soc. Lond., Biol., 238:
127-185.
PHILCOX, D. 2007. Cyatheaceae. In: SHAFFER-FEHRE, M. (ed.) in
DASSANAYAKE, M.D. (ed.) A revised handbook of the flora of Ceylon.
Pteridophyta (Ferns and Fern Allies) 15 Pteridophyta (Ferns and Fern Allies): 48-55.
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RANIL, R.H.G., PUSHPAKUMARA, D.K.N.G., WIJESUNDERA, D.S.A.,
DHANASEKARA, D.M.U.B. & GUNAWARDENA, H.G. 2004. Species diversity
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SLEDGE, W.A. 1982. An annotated check-list of the Pteridophyta of Ceylon. Bot. J.
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FERN GAZ. 18(7):328-332. 2010
328
DRYOPTERIS REMOTA (A. BRAUN EX DÖLL) DRUCE
(DRYOPTERIDACEAE: PTERIDOPHYTA) IN POLAND
Dariusz Tlałka1 & Krzysztof Piątek2
1
Os. nad Sołą 4/19, Kęty, Poland;
Institute of Botany, Jagiellonian University, Kraków, Kopernika 27, 31-501 Kraków,
Poland, e-mail: [email protected]
2
Keywords: Dryopteris remota, Poland, ferns, Pteridophyta, distribution;
ABSTRACT
The presence of Dryopteris remota (A. Braun ex Döll) Druce in Poland was
unclear (Piękoś-Mirkowa 1979). Recently the presence of this species in Poland
has been confirmed and this paper presents a review and description of Polish
localities. Fifteen new localities of D. remota were found in the Beskidy
Mountains (S Poland), and two contemporary and six historical sites confirmed.
Ecological requirements in Poland are described. Determination of the Polish
specimens of D. remota was based on comparison with descriptions and
iconography of this species and in consultation with specialists in Dryopteris. A
map of the distribution of D. remota within Poland is provided.
INTRODUCTION
The presence of Dryopteris remota (A. Braun ex Döll) Druce in Poland was unclear
(Piękoś-Mirkowa,1979). Dryopteris remota had been recorded in Poland from eight
sites, of which six are historical. Fraser-Jenkins (pers. comm.) reported two sites from
Poland, based on a revision of Polish herbarium materials, one of which (Przemyśl)
appeared on a map of the overall distribution of this species (Ekrt et al. 2007). The
species has also been reported by Fiek (1881) and Shube (1903) from four sites. Two
contemporary localities were reported by Kwiatkowski (2006). During 2007-2009 D.
Tlałka found 15 further localities of this species in Poland (Tlałka, 2008; Tlałka &
Rostański, 2008; Tlałka, 2009; Szczęśniak, Tlałka & Rostański, 2009; Wilczek, Tlałka
& Rostański, 2009; Tlałka 2010), and voucher specimens were confirmed by K. Piątek.
This paper presents a review and description of material from all Polish localities. A
map of the distribution of D. remota within Poland is also presented. A detailed
description of the species, nomenclature, general distribution as well as a complete
literature review was published by Ekrt et al. (2007).
The distribution of this species within Poland is based upon examination of specimens
from the KTU herbarium (abbreviations follow Holmgren et al. 1990) and from the
literature (Fiek, 1881; Shube, 1903; Kwiatkowski, 2006). Voucher herbarium
specimens from the Carpathian localities found by Tlałka are preserved at KTU and
duplicates are preserved at KRA. Phytosociological survey was made in accordance
with the methodology proposed by the Zurich-Montpellier school.
ECOLOGICAL REQUIREMENTS IN POLAND
Dryopteris remota is reported from altitudes of 480 to 730 m from ravines, steep rocky
329
FERN GAZ. 18(7): 328-332. 2010
slopes and screes. It grows in forest communities, especially in Carpathian beech forest
Dentario glandulosae-Fagetum. At one of the stations a phytosociological survey
(relevé) was made:
Phytosociological relevé:
Dentario glandulosae-Fagetum. 16.06.2008. Porąbka, Wielka Puszcza valley, steep
(50%) N slope, 550 m a.s.l., Relevé area: 100 m2, cover of tree layer (A): 90%, cover
of shrub layer (B): 1%, cover of herb layer (C): 30%. A: Fagus sylvatica 4. Abies alba
+. Fraxinus excelsior +. Acer pseudoplatanus +. B: Fagus sylvatica +. C: Dryopteris
remota +. D. dilatata 1. D. carthusiana +. D. affinis +. D. filix-mas +. Athyrium filixfemina 2. Oxalis acetosella 1. Rubus sp. +. Acer pseudoplatanus 1. Impatiens parviflora
1. Abies alba 1. Senecio ovatus agg. +. Geranium robertianum +.Allium ursinum +.
Dentaria glandulosa +. Dentaria bulbifera +. Carex sylvatica +. Aruncus dioicus +.
Galeobdolon luteum +. Sorbus aucuparia +.
RECORDS OF DRYOPTERIS REMOTA FROM POLAND
All currently known Polish stations of Dryopteris remota are presented below. The
distribution map of D. remota in Poland is presented in Figure. 1. The map was created
using cartogram method based on an artificial grid of square units 10 x 10 km (Zając A.
& Zając M., 2001).
Historical records
1. Kraków-Bronowice, ATPOL square DF6911, leg. A. Rehman, rev. Fraser-Jenkins C.
R. (pers. inf.);
2. Przemyśl, ATPOL square GF90, rev. Fraser-Jenkins C. R., (pers. inf.);
3. Buchberg (Bukowiec Mount), ATPOL square BE9303 (Fiek 1881);
4. Schierlischskoppe (Kostrzyna Mount), ATPOL square BE9313 (Fiek 1881);
5. Storchberg (Stożek Wielki Mount), ATPOL square BE9302 (Fiek 1881);
6. Freudenkamm (a valley in the vicinity of Sokołowsko (Görbersdorf)), ATPOL
square BE9313 (Shube 1903).
Contemporary localities
7. Góry Ołowiane / Turzec, ATPOL square BE6123 (Kwiatkowski 2006);
8. Ciechanówka, ATPOL square BE7103 (Kwiatkowski 2006);
9. Beskid Mały Mountains, Wielka Puszcza valley, on the SW slope of Wielka
Bukowa Mount and N slope of Beskid Mount, 540-590 a.s.l., 22 individuals in
beech forest. Porąbka, Porąbka gmina, Bielsko-Biała powiat, Silesia Province,
ATPOL square DF9530, KTU;
10. Beskid Mały Mountains, Kocierska Pass, on the N slope, Wielka Puszcza Valley, on
the S slope of Wielka Bukowa Mount and on the N and NE slope of Beskid Mount,
550-610 m a.s.l., 19 individuals in beech forest and fir-spruce forest. Targanice,
Andrychów gmina, Wadowice powiat, Małopolska Province; Porąbka, Porąbka
gmina, Bielsko-Biała powiat, Silesia Province, ATPOL square DF9531;
11. Beskid Mały Mountains, Targaniczanka valley, on the N slope of Kiczora Mount
and on the W slope of Potrójna Mount, Kiczora, on the NW slope, 530-630 m a.s.l.,
several hundred individuals in beech forest and fir-spruce forest, Targanice,
Andrychów gmina, Wadowice powiat, Małopolska Province, ATPOL square
DF9532, KTU;
12. Beskid Mały Mountains, Potrójna Mount between Łamana Skała and Leskowiec,
TLAŁKA & PIĄTEK: DRYOPTERIS REMOTA IN POLAND
330
on the E slope, 630-640 m a.s.l., 2 individuals in beech forest. Targoszów, Stryszawa
gmina, Sucha Beskidzka powiat, Małopolska Province, ATPOL square DF9640,
KTU;
13. Beskid Mały Mountains, Leskowiec Mount, on the SW slope, 700-730 m a.s.l., 26
individuals in beech forest, Targoszów, Stryszawa gmina, Sucha Beskidzka powiat,
Małopolskie Province, ATPOL square DF9630, KTU;
14. Beskid Żywiecki Mountains, Sucha Góra Mount on the SW slope, 600-620 m a.s.l.,
3 individuals in fir-spruce forest, Rajcza, Rajcza gmina, Żywiec powiat, Silesia
Province, ATPOL square DG2334;
15. Beskid Śląski Mountains, Jasionka valley, on the NW slope of Błatna Mount, 600
m a.s.l., 1 plant in beech forest, Brenna, Brenna gmina, Cieszyn powiat, Silesia
Province, ATPOL square DG0202;
16. Beskid Mały Mountains, Łamana Skała Mount, on the SE slope above Komoniecki
Cave, 720-730 m a.s.l., 15 individuals in young fir-spruce forest, Las, Ślemień
gmina, Żywiec powiat, Silesia Province, ATPOL square DF9544, KTU;
17. Beskid Mały Mountains, Beskid Mount near Kocierska Pass on the SE slope, 650-
Figure 1. Map of the distribution of Dryopteris remota in Poland: z – historical
records, – current sites.
331
FERN GAZ. 18(7): 328-332. 2010
660 m a.s.l., 4 individuals in beech forest, Kocierz Rychwałdzki, Łękawica gmina,
Żywiec powiat, Silesia Province, ATPOL square DF9541;
18. Beskid Mały Mountains, Nad Płonem Mount, on the NW slope, 610-620 m a.s.l.,
17 individuals in beech forest, Kocierz Rychwałdzki, Łękawica gmina, Żywiec
powiat, Silesia Province, ATPOL square DG0501;
19. Beskid Mały Mountains, Mały Gibasów Wierch Mount, on the SW slope below
Czarne Działy Caves, 700-710 m a.s.l., 10 individuals in beech forest, Ślemień,
Ślemień gmina, Żywiec powiat, Silesia Province, ATPOL square DG0503;
20. Beskid Mały Mountains, Potrójna (Czarny Groń) Mount, on the W and SW slopes,
620-710 m. a.s.l., 10 individuals in beech forest, Targanice, Andrychów gmina,
Wadowice powiat, Małopolska Province; Kocierz Rychwałdzki, Łękawica gmina,
Żywiec powiat, Silesia Province, ATPOL square DF9542;
21. Beskid Mały Mountains, Kiczera, on the E slope, 530 m a.s.l., 2 individuals in beech
forest, Porąbka, Porąbka gmina, Bielsko-Biała powiat, Silesia Province, ATPOL
square DF9434;
22. Beskid Mały Mountains, Łamana Skała Mount, on the SE slope below Dusica
waterfall, 660-680 m a.s.l., 4 individuals in young fir-spruce forest, Las, Ślemień
gmina, Żywiec powiat, Silesia Province, ATPOL square DG0504;
23. Beskid Mały Mountains, Magurka Ponikiewska Mount, on the NE slope, 480-490
m a.s.l., 3 individuals in beech forest, Koziniec, Mucharz gmina, Wadowice powiat,
Małopolska Province, ATPOL square DF9622.
CONCLUSION
Dryopteris remota should be included in the list of the native species of the flora of
Poland. The species is probably more common, but has been overlooked. It should be
included as a mountain species in the list of Zając (1996). Considering the IUCN
categories and criteria (IUCN 2001), we recommend that D. remota should be
designated as a critically endangered species (CR) in Poland.
ACKNOWLEDGEMENTS
We are very grateful to C. R. Fraser-Jenkins for making available information about
Polish specimens revised by him in herbaria and for confirmation of our identification;
we also thank A. Zając for his help in preparation of the distribution map and translation
of German geographical names.
REFERENCES
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BOOK REVIEW
FLORA OF PENINSULA MALAYSIA. SERIES 1: FERNS AND LYCOPHYTES.
VOLUME 1. Edited by B.S.Parris, R. Kiew, R.C.K. Chung, L.G. Saw & E.
Soepadmo. 2010. (Malaysian Forest Records No. 48). Pp. 250, 12 black & white figs,
18 col. plates. 256 x 800mm. Forest Research Institute Malaysia, 52109 Kepong,
Selangor Darul, Ehsan, Malaysia; Ministry of Natural Resources and Environment,
Malaysia. ISBN:978-967-5221-24-8 (v.1). Available from Forest Research Institute
Malaysia.
The Flora of Peninsula Malaysia is an initiative of the Malaysian Government as part
of its commitment to biodiversity conservation, and it is supporting employment and
training for taxonomists, fieldwork and overseas collaboration. The Flora is founded on
thorough taxonomic review, including examination of type material, and includes
ecological information. A strong focus of the project is new botanical exploration,
coupled with taxonomic training, to gather information from areas where there has
been little collecting in the past and review old records, to acquire up-to-date
information on the status of plants, their diversity and current distributions, in order to
assess conservation needs.
The last published fern flora of Peninsula Malaysia was that of Holttum in 1968
(and covered ferns but not fern allies), but he continued to publish on ferns of this
region during his long retirement at Kew, contributing a number of family accounts for
Flora Malesiana.
The Volume starts with a conspectus by Barbara Parris of Orders, Families and
Genera, to reflect the changes in classification of ferns and fern ‘allies’ that have
emerged from recent phylogenies based on morphological and molecular data. The
editors broadly follow the treatment of Smith et al. (2006) in placing Equisetaceae and
Psilotaceae with the ferns, rather than with lycophytes, but have modified the treatment
for ferns, e.g. Pteridaceae is divided into five monophyletic families, Parkeriaceae,
Adiantaceae, Cryptogrammaceae, Sinopteridaceae and Pteridaceae; Polypodiaceae is
divided into three families. Loxogrammaceae, Polypodiaceae and Grammitidaceae.
Justification for the changes is provided.
A further chapter by Parris deals with the history of botanical collecting in Peninsula
Malaysia, followed by a chapter on species assessment and conservation by L.S.L.
Chua of the Forest Research Institute Malaysia.
Keys to families, genera and species are provided. Each taxonomic account in the
Flora includes information on nomenclature and type material, detailed description,
notes on distribution together with a distribution map, conservation assessment based
on the current IUCN guidelines and notes on ecology. There are useful black and white
figures and coloured plates that illustrate the variation within families, and some fern
habitats.
Volume 1 covers accounts for 9 families, 21 genera and 100 species, and includes
Selaginellaceae, Psilotaceae, Equisetaceae, Osmundaceae, Matoniaceae, Schizaceae,
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FERN GAZ. 18(7). 2010
Cibotiaceae, Loxogrammaceae and Grammitidaceae.
The authors and editors are to be congratulated on a beautiful, scholarly and very
valuable work.
REFERENCE
Holttum, R.E. 1968. Revised Flora of Malaya. Vol. 2 Ferns. 2nd ed. Government
Printing Office, Singapore.
Mary Gibby
335
FERN GAZ. 18(7). 2010
will be refereed
aligned.
aligned.
a legible form.
Supplements.
FERN GAZ. 18(8):336-341. 2010
336
A NEW GENUS FOR THE FERN FLORA OF IRAN: CYRTOMIUM
A. GHOLIPOUR1 & W. GREUTER2
1
Payame Noor University, Sari, Mazandaran. [email protected]
Herbarium Mediterraneum, Orto Botanico, Università degli Studi, Palermo, Italy;
and Botanischer Garten & Botanisches Museum Berlin-Dahlem, Freie Universität
Berlin, Germany. Author for correspondence: [email protected]
2
Keywords: Cyrtomium fortunei, Dryopteridoideae, Mazandaran, Hyrcanian forest,
disjunct area.
ABSTRACT
A stand of Cyrtomium fortunei was found by the first author in a natural habitat
on the northern slope of the Elburs range in N Iran, Mazandaran Province,
growing on sandstone rocks in the Hyrcanian forest. The whole context strongly
suggests it to be indigenous there, but phytogeographically this would result in
a huge, puzzling disjunction between the known native area in SE Asia and the
newly discovered stand.
INTRODUCTION
Cyrtomium C. Presl is a genus of the dryopteridoid ferns, closely related to and
sometimes combined with Polystichum Roth, or alternatively with the New World
Phanerophlebia C. Presl, a name of the same date but which is treated as having priority
over Cyrtomium. Molecular systematics (Yatskievych et al. 1988, Lu et al. 2005) and
karyosystematics (Lu et al. 2006) have helped elucidate the classification of the group,
now thought to consist of four clades equivalent to Polystichum, Phanerophlebia, a
group comprising Cyrtomidictyum Ching, Cyrtogonellum Ching plus some species
formerly misplaced in Polystichum and Cyrtomium, and Cyrtomium sensu stricto. The
latter, thus redefined, comprises ferns with irregularly scattered sori arising from
included veinlets and imparipinnate fronds with a non-pinnatifid terminal segment.
Cyrtomium sensu lato has been monographed twice, by Christensen (1930) who
recognised nine species and Shing (1965) who raised the species number to 59.
According to recent floristic works (Iwatsuki 1995, Kung 2001) a more realistic number
is 35 or less. Except for two species (C. caryotideum (Hook. & Grev.) C. Presl, also
found on Hawai’i, and C. micropterum (Kunze) Ching, which extends to Madagascar
and tropical Africa, south to Natal) the genus is confined to S and SE Asia, with a
marked centre of diversity in China from where it extends to Japan, Korea, Viet-Nâm,
Thailand, India and the Himalaya. The westernmost native occurrences in Eurasia are
in Pakistan, W Himalaya, from where three species have been reported: C.
anomophyllum (Zenker) Fraser-Jenk. (= C. nervosum Ching & Shing), C. caryotideum,
and C. macrophyllum (Makino) Tagawa (Fraser-Jenkins 1992, 2008, Kullar 2000).
Two species of Cyrtomium are widely cultivated as ornamental pot plants: C.
falcatum (Sw.) C. Presl and C. fortunei J. Sm. Both have become established escapes in
various parts of the world with a suitable climate, and either or both are reported as
locally naturalised in, e.g., Britain, Italy, the Macaronesian Islands, the southern United
States, Hawai’i, and Australia. For C. falcatum, diploid, tetraploid and apomictic
triploid karyotypes are known, whereas C. fortunei appears to be consistently triploid
337
FERN GAZ. 18(8): 336-341. 2010
and apomictic, comprising a number of discernible morphotypes that have been treated
as varieties or sometimes species (Iwatsuki 1995).
MATERIAL
Voucher specimens of the Iranian Cyrtomium have been deposited in the herbarium,
Payame Noor University, Sari, Mazandaran, Iran (PNUS; not yet registered in Index
Herbariorum). Duplicates are kept in the Berlin-Dahlem herbarium (B) and the second
author's herbarium (now: PAL-Gr). Identification has been checked by comparison with
herbarium material from all over the generic range present in Berlin-Dahlem (B) as well
as descriptions in the floristic and monographic literature.
RESULTS
Plants belonging to the genus Cyrtomium were discovered by the first author on 10 Sep
2010 in a single locality on the northern slopes of the Elburs range, 25 km S of the city
of Sari and 8.5 km N of the village of Lajim (Fig. 1-3). Cyrtomium had not been
recorded from Iran previously (Khoshravesh et al. 2009). Specimens sent to the second
author for study proved to be typical C. fortunei, characterised e.g. by the shape and
dimensions of its fronds and pinnae, the irregularly and distantly denticulate pinna
margin with minute antrorse to incurved teeth, and the pale, thinly papery, entire indusia
(Fig. 4).
About 30 tufts were observed, scattered over an area of ca. 300 m2, growing in a
small side valley on shaded sandstone rocks, on the trunks of fallen trees and on shallow
soil of the valley bottom (Fig. 5). The dominant vegetation was natural Hyrcanian
deciduous forest, in which Carpinus betulus L. and Fagus sylvatica subsp. orientalis
(Lipsky) Greuter & Burdet predominate, mixed with other characteristic elements such
as Alnus subcordata C. A. Mey., Parrotia persica C. A. Mey., and Acer velutinum Boiss.
Also, the following fern species were noted: Asplenium scolopendrium L., Dryopteris
affinis (Lowe) Fraser-Jenk., Polystichum aculeatum (L.) Roth, and Pteris cretica L. The
area in question, known locally as “Cron forest”, has no protected status.
Label data of the vouchers collected are as follows: Mazandaran, Sari, Aghmashhad,
Cron, 36°20´08.2´´ N, 53°05´25.6´´ E, 520 m, 10 Sep 2010, A. Gholipour 890367
(PNUS, B); same place, 4 Oct 2010, A. Gholipour 890441 (PNUS, PAL-Gr) (Fig. 6).
DISCUSSION
The so far known native range of Cyrtomium fortunei extends from Japan and SE China
to S Korea, Viêt-Nam and E Thailand (Iwatsuki 2000). There is a single record (not
checked by us) from easternmost India (Manipur: Fraser-Jenkins 2008), which if
correct would be the closest known occurrence to our new locality. The distance
between Sari and Manipur is ca. 4200 km as the crow flies, roughly the same as that
which separates Manipur from Tokyo at the eastern edge of the species range
(4500 km). This is a huge disjunction, with the whole Himalayan range and the Iranian
highlands lying in-between.
An easy explanation would be to suspect that the Iranian population, rather than
being truly native, was brought to its new location by the voluntary action or
unintentional assistance of humans. The ease with which cultivated Cyrtomium species,
in various parts of the globe, escape from cultivation and become established in
semi-natural habitats lends credibility to such a hypothesis. For C. fortunei proper,
naturalised occurrence, while much less generalised than for C. falcatum, has been
GHOLIPOUR & GREUTER: A NEW GENUS FOR THE FERN FLORA OF IRAN
338
1
2
3
4
Figures 1-4. Cyrtomium fortunei, photographed in the wild. 1. Habit (with juvenile
Asplenium scolopendrium underneath). 2. Stipe, with large blackish and small fringed
scales. 3. Fertile pinna from below. 4. Detail of same, showing indusia, indumentum,
and crenulate margin. Photography, A. Gholipour.
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FERN GAZ. 18(8): 336-341. 2010
reliably reported from the U.S.A. (4 states: Yatskievych 1993), SE France, where it
enjoys protected status (Salanon 1998), and the southern Alps of Italy (Conti & al.
2005).
However, the entire context in which the plants are growing in Iran makes
naturalisation an unlikely assumption. The place is literally in the middle of nowhere,
and neither at this locality, nor in the surrounding area in general, is there any sign of
degradation or indeed of any interference or activity by humans. Weedy, alien species
are conspicuously absent. The closest village of note, Aghmasshad, is situated at a
distance of ca. 4 km. It does not have any hotel, tourist resort or similar facility in which
alien potted plants might be suspected to be grown for amenity. Even more
significantly, Cyrtomium is not known to be or have been cultivated in Iran, and while
its erratic presence cannot be entirely precluded it is definitely not a popular ornamental
fern in Iran, nor is it available through the local horticultural trade.
We are therefore of the firm opinion that Cyrtomium fortunei is a member of the
native Iranian flora. One may of course theorise that it has settled in it current site as a
result of natural long-distance dispersal, but if so, there is no way to know when that
event may have happened. The Hyrcanian forest of the Elburs range is rich in Tertiary
relic species (and genera), and there is no reason to preclude considering Cyrtomium as
one of their number.
ACKNOWLEDGEMENTS
Ms Sarah Bollendorff, Berlin, kindly produced the scanned specimen image. Brigitte
Zimmer, Berlin, assisted with consultation of the Berlin Herbarium. Christopher FraserJenkins provided valuable information on his Pakistani gatherings of Cyrtomium.
Figure 5. The native habitat of Cyrtomium fortunei, in Mazanderan. Photography, A.
Gholipour.
GHOLIPOUR & GREUTER: A NEW GENUS FOR THE FERN FLORA OF IRAN
Figure 6. One of the Iranian voucher specimens of Cyrtomium fortunei.
340
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FERN GAZ. 18(8): 336-341. 2010
REFERENCES
CHRISTENSEN, C. 1930. The genus Cyrtomium. Amer. Fern J. 20: 41-52.
CONTI, F., ABBATE, G., ALESSANDRINI, A. & BLASI, C. 2005. An annotated
checklist of the Italian vascular flora. Palombi, Roma.
FRASER-JENKINS, C. R. 1992. The ferns and allies of the far west Himalaya.
Pakistan Syst. 5: 85-120.
FRASER-JENKINS, C. R. 2008. Taxonomic revision of three hundred Indian
Subcontinental pteridophytes, with a revised census list. Bishen Singh Mahendra
Pal Singh, Dehra Dun.
IWATSUKI, K. 1995. Dryopteridaceae. In: IWATSUKI, K., YAMAZAKI, T.,
BOUFFORD, D. E. & OHBA, H. (ed.), Flora of Japan. Volume I, Pteridophyta and
Gymnospermae: 120-173. Kodansha Ltd., Tokyo.
KHOSHRAVESH, R., AKHANI, H., ESKANDARI, M. & GREUTER, W. 2009: Ferns
and fern allies of Iran. Rostaniha 10, Suppl. 1.
KHULLAR, S. P. 2000. An illustrated fern flora of the West Himalaya. Volume II.
International Book Distributors, Dehra Dun.
KUNG, H.-S. (ed.) 2001. Flora Reipublicae Popularis Sinicae. Tomus 5(2) Pteriophyta,
Dryopteridaceae (2). Science Press, Beijing [in Chinese].
LU, J.-M., LI, D.-Z., GAO, L.-M., CHENG, X. & WU, D. 2005. Paraphyly of
Cyrtomium (Dryopteridaceae): evidence from rbL and trnL-F sequence data. J. Pl.
Res. 118: 129-135.
LU, J.-M., CHENG, X., WU, D. & LI, D.-Z. 2006. Chromosome study of the fern genus
Cyrtomium (Dryopteridaceae). Bot. J. Linn. Soc. 150: 221-228.
SHING, K.-H. 1965. A taxonomical study of the genus Cyrtomium Presl. Acta Phytotax.
Sin., Add. 1: 1-48 [in Chinese].
SALANON, R. 1998. Mémento de la flore protégée des Alpes-Maritimes. Office
National des forêts, Paris.
YATSKIEVYCH, G. 1993. 13. Cyrtomium. In: Morin, N. R. (ed.), Flora of North
America 2: 299-300. Oxford University Press, Oxford, etc.
YATSKIEVYCH, G., STEIN, D. B. & GASTONY, G. J. 1988. Chloroplast DNA
evolution and systematics of Phanerophlebia (Dryopteridaceae) and related fern
genera. Proc. Natl. Acad. Sci. U.S.A. 85: 2589-2593.
FERN GAZ. 18(8):342-350. 2010
342
TWO NOVEL ASPLENIUM HYBRIDS (ASPLENIACEAE:
PTERIDOPHYTA) FROM TENERIFE, CANARY ISLANDS
F.J. RUMSEY1 & A. LEONARD2
1
Dept. of Botany, Natural History Museum, Cromwell Road, London, SW7 5BD, UK,
2
37 Lower Bere Wood, Waterlooville, Hants., PO7 7NQ, UK
Keywords: Asplenium hemionitis, A. aureum, A. onopteris, A. × tagananaense,
hybridization, Macaronesia
ABSTRACT
A plant closely resembling Asplenium hemionitis L. but with more dissected,
lobed fronds was discovered during a trip to the Anaga mountains, Tenerife,
Canary Islands in 2009. This was found to show almost complete spore abortion,
indicating a hybrid origin. From the associated species and frond form we
suggest the other parent to be A. onopteris L. This represents the first
documented hybrid of the rather taxonomically isolated A. hemionitis. The
hybrid, A. × tagananaense, is described and its distinguishing features given. A
further novel Asplenium hybrid, photographed in 1995 but not subsequently
refound, is identified as that between A. onopteris and A. aureum Cav. In the
absence of a specimen it is not formally described but its distinctive features are
illustrated and its occurrence reported.
INTRODUCTION
In February 2009 a small group of pteridologists led by the second author and
comprising Alison Evans, Michael Hayward, Tim Pyner and Martin Rickard went to
Tenerife. During the excursion, an odd looking fern, which several in the group
considered to be an aberrant form of Asplenium hemionitis was found. The plant had the
palmate frond form unique in the region to this species but closer examination showed
the lobes themselves to be more highly dissected, the lobules not apparent as they were
largely in the same plane as the frond and closely imbricate. The possibility that this
might constitute a hybrid was considered and accordingly photographs were taken (Fig.
1) along with a few voucher fronds (see drawing, Fig. 4a). On the groups’ return to the
British Isles these were examined by FR and found to be producing highly abortive
sporangia, strongly suggesting a hybrid origin. Comparison with recently collected
Azorean material of the very rare hybrid A. × rouyi Viane (A. scolopendrium × A.
onopteris), showed some similarities, suggesting that the other parent might also be
diploid A. onopteris. The group returned to the site in November 2009, this time
accompanied by Patrick Acock. They found the plant still to be present although no new
foliage had developed. It gave the opportunity to make additional observations about
the plant, its surroundings, etc. Apart from A. hemionitis the only other Asplenium
species closely associated (and with juvenile plants in the immediate vicinity) was
found to be A. onopteris (Fig. 2).
The possibility that A. × tagananaense is a hybrid with some other Canarian
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FERN GAZ. 18(8): 342-350. 2010
Asplenium taxon may largely be ruled out on gross morphological grounds; the only
other taxa with an appropriate frond shape are tetraploid and would be expected to show
a greater influence on the hybrid frond form because of gene dosage effects (Bennert et
al., 1991). Asplenium × rouyi (= A. onopteris × A. scolopendrium), another extremely
rare hybrid, only ever found twice (Schäfer & Rasbach, 2000), shares similarities and
the A. onopteris parent. This hybrid with the entire-fronded A. scolopendrium, a very
rare plant in the Canaries, differs in its narrower more distinctly triangular frond form,
the deeper divisions of the central lobe, the pinnules thus created more nearly at right
angles to the rachis. It also appears to have a somewhat fleshier texture common to
many A. scolopendrium hybrids (Rumsey et al., 2004).
Many years earlier AL had found another curious Asplenium on Tenerife which from
its morphology (Figs. 3; 4b) could not be matched with any known species, although
Figure 1. Asplenium × tagananaense – head of Barranco de la Iglesia, Taganana,
Tenerife. Feb. 2009. Photo: Alison Evans.
RUMSEY & LEONARD: ASPLENIUM HYBRIDS FROM TENERIFE
344
from its texture strongly suggested it may be a hybrid of A. aureum Cav., seen growing
nearby. A voucher frond was collected and sent to the Department of Botany at the
Natural History Museum; most unfortunately its present whereabouts have not been
traced. Subsequent visits to the area, by a woodland trail in the hills c. 1km to the west
of Erjos, Los Silos (c. 28° 19’ 49.35’’N 16° 49’ 06.77’’W) at an altitude of just over
720m, revealed the locality to have changed somewhat in character and the plant could
no longer be refound. Photographs taken of the fern when first found, which
unfortunately only reveal the upper surface of the fronds, show the plant to be very
distinctive, clearly differing from A. aureum in its more dissected frond form and in its
somewhat triangular shape, the lowest pinnules just the larger. While other taxa within
the subgenus Ceterach are now known to occur on Tenerife, both morphological (frond
width) and ecological factors indicate that A. aureum is involved. Again, on the basis of
its morphology, only the 2-3 pinnate, deltate fronded A. onopteris, the only other
Asplenium species seen in the vicinity, could be the other parent. Confirmation will
however not be possible until the original specimen or similar wild plants are found.
Few hybrids are known involving members of the morphologically distinctive subgenus
Ceterach, although others are inferred in the origin of allopolyploids within this group
(Trewick et al., 2002; Van den Heede et al., 2004); most examples of F1 hybrids found
in the wild are backcrosses between allopolyploid derivatives and parental taxa (Van
den Heede & Viane, 2002). The Canarian endemic A. aureum has been demonstrated to
Figure 2. Asplenium × tagananaense in situ with juvenile plants of both putative
parents, Nov. 2009. Photo: P. Acock.
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FERN GAZ. 18(8): 342-350. 2010
Figure 3. Asplenium aureum × A. onopteris – by forest trail c. 1km West of Erjos,
Tenerife, Oct. 1995. Photo: A. Leonard.
346
be an allotetraploid (Pintér & Vida, 1993). Although its parentage is still uncertain, it is
now known to be one parent of the Macaronesian endemic (Madeira, Canaries)
hexaploid A. lolegnamense Gibby & Lovis (Pintér et al. 2002) and also involved in the
origin of the Canarian endemic octoploid A. octoploideum Viane & Van den Heede (Van
den Heede et al., 2004). The derived polyploid taxa may be more locally abundant than
the more mesic A. aureum, particularly at altitude but they are more restricted in
distribution across the Canarian archipelago.
Some earlier proposed hybrids involving crosses with taxa of other subgenera have
subsequently been shown to be in error, eg. Asplenoceterach badense; only the hybrid
of tetraploid A. ceterach L. with the allotetraploid A. majoricum (A. × barrancense (W.
Bennert & D.E. Mey.) Pericás & Rosselló) has been cytologically confirmed (Rasbach
et al., 1989). However, another very rare example in which A. aureum has been
suggested to be parental is known (Hansen & Sunding, 1993). Asplenium × newmani
Bolle was described from material collected by C. Bolle in the Barranco del Rio on La
Palma in September 1852. It is a simply pinnate fern with fronds only c. 2cm broad, the
rachis above green. Known only from the type gathering now in Herbarium Berolinense
(B20 0026447), one parent is clearly either the Macaronesian endemic A. anceps or, and
on ecological grounds less likely, its more cosmopolitan relative Asplenium
trichomanes subsp. quadrivalens. Similar plants have never been refound.
DESCRIPTION
Asplenium × tagananaense F.J. Rumsey hybr. nov. (A. hemionitis L. × A. onopteris
L.)
Habitu et textura A. hemionitis similis, praecipue frondis plus divisus, lobatus, plus
minusve pinnatifidus, basi pinnatilobus, lobo terminali multo magis, sporae abortivus
differt.
Holotypus: Canary Islands, Tenerife, Anaga mountains, by track to Taganana near
Casa Forestal, at head of Barranco de la Iglesia, amongst rocks by path on steep slope
in Laurel woodland, with both putative parents, c. 900m, 28° 32’ 43.50” N, 16° 13’
34.50” W, Leonard et al., s.n. 2 February 2009. BM
Perennial from erect-ascending rhizome, leaves flushing annually, wintergreen,
coriaceous, glabrous, to 35 cm; petiole 1.5-2.2 x lamina length, somewhat swollen at
base, dark reddish brown for most of its length; lamina to c. 15 cm long, 12 cm broad,
palmately (3-)5 lobed, the mid lobe distinctly longer, the lobes inconspicuously divided
from ¼ to ¾ of their width, the basal more profoundly, lobes with small erect teeth at
their apices, the apices sub-acute to obtuse. Sori linear, +/- curved, sporangia failing to
mature, spores highly abortive.
Asplenium × tagananaense is most likely to be confused with its A. hemionitis
parent, various frond forms of which have been described from the Canary Islands
(Kunkel, 1966; 1967 and see Benl, 1967). While very distinctive amongst European
Asplenium because of its entire, initially deltate, later palmate frond form, variability
exists in A. hemionitis and the more markedly lobed and deeply divided forms appear
to be more prevalent and perhaps exclusive to the Canary Islands. Rare monstrose
forms such as described by Kunkel, 1966; 1967 may be similar in appearance to the
newly described hybrid, although they would not show the spore abortion it clearly
demonstrates. From the illustration in its original publication f. cristatum Kunkel
(Kunkel, 1966) is clearly not synonymous; we have seen no material of f.
pinnatipartidum Kunkel with which to compare our plants.
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FERN GAZ. 18(8): 342-350. 2010
B
A
Figure 4.
A. (right) Asplenium × tagananaense – underside of frond drawn from Holotype –
Leonard et. al., sn. (BM). Scale bar represents 1cm.
B. (left) Asplenium aureum × A. onopteris – upper surface of frond. Scale bar
represents 1cm.
348
All herbarium material of A. hemionitis at BM has been checked and no further
examples of A. × tagananaense have been found. Given its superficial similarity to A.
hemionitis it may be overlooked but we believe it highly likely that it is genuinely,
extremely rare.
Molecular studies based on the sequencing of small portions of the plastid genome,
by which the maternal parent can be identified (Vogel et al., 1998), have been initiated
but not yet concluded.
DISCUSSION
No hybrids of Asplenium hemionitis have previously been reported (Reichstein, 1981;
Prelli, 2001). A. hemionitis is somewhat isolated within the broad genus Asplenium, the
only European example of the section Hemionitidastrum Fée (Schneider et al., 2004).
Similarly no hybrids of A. aureum have been conclusively found (the likely origin of A.
× newmani, found only once in the mid 19th century, still open to debate).
Hybridization events involving A. aureum are however known to have occurred as
inferred from molecular studies of its allopolyploid progeny (Pintér et al. 2002) (Van
den Heede et al., 2004). Observations in other pteridophyte groups such as Equisetum
(Page, 1972) suggest greater levels of hybridization between phylogenetically closer
taxa, where speciation has proceeded through ecological, hence physical separation and
breeding barriers may not have been so well established. The occurrence of hybrids
may thus give some indication of relationships and where apparent inter-generic
hybrids are found give cause for taxonomic reconsideration (see recent studies in the
Orchidaceae, eg. Bateman, 2006). Generic delimitation within Aspleniaceae has long
been contentious with some apparently morphologically distinctive entities, eg.
Phyllitis, Ceterach, Pleurosorus traditionally recognised by many authors. Molecular
phylogenetic studies (eg. Schneider et al., 2004), are now contributing much to our
understanding of the relationships within this highly speciose and taxonomically
contentious genus. These studies show that these entities are firmly nested within
Asplenium and would by their recognition render it paraphyletic. Essentially, if we
recognise Phyllitis or Ceterach then an equally valid case is made for the recognition
of other segregates; even the limited British flora would see its 13 species split between
8 genera. As evidenced by the finds reported here, even where phylogenetically distant,
rare inter-specific reproductive events may occur, the offspring often instantly
threatened by the very combination of factors which have enabled their production. It
is potentially these unique combinations of more disparate genomes which are the more
likely to promote polyploidy through disrupted cell division and thus fuel the abrupt
speciation of very distinctive entities. As such these plants afford us unique insights into
evolutionary mechanisms and processes but currently have no conservation standing.
ACKNOWLEDGEMENTS
The authors would like to thank the various members of the BPS who were present
during the initial discovery of A. × tagananaense and during the re-visit to Tenerife later
in 2009, particularly to those (Alison Evans, Patrick Acock) who have contributed
photos and thoughts to this publication. Particular mention should go to Martin Rickard
for immediately realising this plant was almost certainly a hybrid.
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PINTÉR, I. & VIDA, G. 1993. Allotetraploid origin of Ceterach aureum (Aspleniaceae:
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