Alansmia, a new genus of grammitid ferns (Polypodiaceae)
segregated from Terpsichore
MICHAEL KESSLER1, ANA LAURA MOGUEL VELÁZQUEZ2,3, MICHAEL SUNDUE4,
5
AND PAULO H. LABIAK
1
Systematic Botany, University of Zurich, Zollikerstrasse 107, CH-8008, Zurich, Switzerland;
e-mail: michael.kessler@systbot.uzh.ch
2
Department of Systematic Botany, Albrecht-von-Haller-Institute of Plant Sciences, Georg-AugustUniversity, Untere Karspüle 2, 37073, Göttingen, Germany
3
Present Address: Pfefferackerstr. 22, 45894, Gelsenkirchen, Germany; e-mail: moguelve@yahoo.com
4
The New York Botanical Garden, 200th St. and Southern Blvd., Bronx, NY 10458, USA;
e-mail: msundue@nybg.org
5
Departamento de Botânica, Universidade Federal do Paraná, Caixa Postal 19031( 81531-980,
Curitiba, PR, Brazil; e-mail: plabiak@ufpr.br
Abstract. Alansmia, a new genus of grammitid ferns is described and combinations
are made for the 26 species known to belong to it. Alansmia is supported by five
morphological synapomorphies: setae present on the rhizomes, cells of the rhizome
scales turgid, both surfaces of the rhizome scales ciliate, laminae membranaceous, and
sporangial capsules setose. Other diagnostic characters include pendent fronds with
indeterminate growth, concolorous, orange to castaneous rhizome scales with ciliate
or sometimes glandular margins, hydathodes often cretaceous, and setae simple,
paired or stellate. The group also exhibits the uncommon characteristic of producing
both trilete and apparently monolete spores, sometimes on the same plant. New
combinations are made for Alansmia alfaroi, A. bradeana, A. canescens, A. concinna,
A. contacta, A. cultrata, A. dependens, A. diaphana, A. elastica, A. glandulifera, A.
heteromorpha, A. immixta, A. kirkii, A. lanigera, A. laxa, A. longa, A. monosora, A.
reclinata, A. semilunaris, A. senilis, A. smithii, A. spathulata, A. stella var. stella,
A. stella var. flava, A. turrialbae, A. variabilis, A. xanthotrichia. Lectotypifications
are made for Alansmia concina, A. variabilis, Polypodium ciliare, P. flexile, and
P. ovalescens. The genus is named in honor of pteridologist Alan R. Smith.
Key Words: Africa, Grammitidaceae, Lellingeria, Leucotrichum, Neotropics, taxonomy.
Grammitid ferns are a clade of about 750
species (Parris, 2009) characterized by having
green, tetrahedral spores (Mettenius, 1866),
sporangial stalks of one row of cells (Wilson,
1959), and fronds that lack scales but
typically bear uniseriate, pluricelluar setae
(Ching, 1940; Holttum, 1947). They have
been treated as a distinct family (Newman,
1840; Ching, 1940; Parris, 1990a; Ranker
et al., 2004) or as a subfamily of the
Polypodiaceae (e.g., Tryon & Tryon, 1982;
Lellinger, 1989). Phylogenetic studies using
chloroplast DNA sequence data have shown
that the grammitids are a monophyletic
lineage nested within the Polypodiaceae
(Hasebe et al., 1995; Ranker et al., 2004;
Schneider et al., 2004). Generic delimitation
within grammitid ferns is highly controversial, with only one genus recognized by
Tryon and Tryon (1982), four genera by
Parris (1990b), 12 by Copeland (1947), 18
by Parris (2003), and 25 by Parris (2009).
The difficulty in sorting grammitid ferns into
genera is due to homoplasy of obvious
Brittonia, 63(2), 2011, pp. 233–244
© 2010, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.
ISSUED: 1 June 2011
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BRITTONIA
morphological traits such as blade dissection
and the rather obscure nature of taxonomically informative traits such as hydathodes,
rhizome symmetry, root insertion, or even
presence of fungal bodies on the blades
(Smith, 1993; Ranker et al., 2004; Sundue
et al., 2010). In a series of papers revising the
neotropical grammitids based on more technical characters, Bishop (1988, 1989), Smith
et al. (1991), Bishop and Smith (1992), Smith
(1992, 1993), and Smith and Moran (1992),
recognized ten genera. Three other neotropical genera have been described since then
(Murillo & Smith, 2003; Sundue, 2010a;
Labiak et al., 2010).
One of these neotropical genera, Terpsichore, was described by Smith (1993) to
accommodate about 50 species characterized
by hydathodes that often produce a calcareous deposit, reddish to castaneous (less often
hyaline) setae mostly 1–3 mm long on the
leaves, concolorous rhizome scales that are
orange to castaneous or blackish and usually
setulose along the margins, and pinna
segments with free, unbranched, pinnate
venation. Smith (1993) recognized five informal species groups within Terpsichore and
suggested that these may be regarded as
natural sections of the genus. These groups
were distinguished by characters such as leaf
orientation (pendent to erect), blade growth
(determinate or indeterminate), rhizome symmetry, color, and margin, and cell type of the
rhizome scales, presence or absence of setae
on the sporangia, presence or absence of
black fungal fruiting bodies (Acrospermum
maxonii Farlow ex Riddle), length of petioles,
degree of development of the proximal
pinnae, whether the setae of the fronds are
solitary or clustered, and presence or absence
of calcareous secretions on the hydathodes.
Most of these characters were later found to
be homoplastic, plesiomorphic, or inconsistently distributed among species. When subject to cladistic analysis, based either on
DNA sequence data (Ranker et al., 2004;
Sundue et al., 2010) or entirely upon
morphological characters (Sundue, 2010b;
Sundue et al., 2010), the genus is resolved
as polyphyletic. New combinations (Labiak
& Matos, 2007) and a new genus, Ascogrammitis (Sundue, 2010a) have been
described in efforts to resolve the polyphyly
[VOL 63
of Terpsichore, but additional problems
remain.
As currently understood, the species of
Terpsichore sensu Smith (1993) belong to six
separate clades within the grammitid ferns
(Ranker et al., 2004; Labiak et al., 2010;
Sundue et al., 2010). These clades correspond
in part to informal groups recognized in
Smith’s (1993) subgeneric classification.
Two of the groups (groups 2 and 4, plus
two species from group 3) form a clade with
Melpomene A.R. Sm. & R.C. Moran and part
of Lellingeria A. R. Sm. & R. C. Moran.
Species corresponding to one of these clades
have been combined in Ascogrammitis
(Sundue, 2010a). Group 5 was found nested
within Micropolypodium Hayata, and two of
its species have been combined there
(Labiak & Matos, 2007). Of the remaining
two species groups of Terpsichore, group 1,
which includes the type species of the genus
(T. asplenifolia (L.) A.R. Sm.), consists of
about 12 species and remains as originally
defined by Smith (1993). The last group
(group 3 pro parte), the T. lanigera group,
was recovered as monophyletic by Ranker
et al. (2004) with strong branch support
values (100% parsimony bootstrap, 1.0
posterior probability, Bremer support of 15).
This group differs from Smith’s (1993) circumscription by the exclusion of two aberrant
species (T. delicatula and T. subscabra) that
form the Terpsichore subscabra clade. Monophyly of the T. lanigera group was further
tested by Labiak et al. (2010) who included
seven species in their molecular phylogenetic study focused on circumscribing its
sister clade, Leucotrichum, a new genus of
five species which they segregated from
Lellingeria. Both the monophyly of the
T. lanigera group and its relation as sister
to Leucotrichum had high branch support
values (parsimony bootstrap 100%, posterior
probably 1.0) in analyses using chloroplast
markers (atpB, rbcL) and 87 morphological
characters. Two morphological synapomorphies further support the sister relationship
of Leucotrichum to the T. lanigera group,
these being radially symmetrical rhizomes
and setae that are hyaline or whitish
(Labiak et al., 2010).
The Terpsichore lanigera group is morphologically distinctive and supported by five
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KESSLER ET AL.: ALANSMIA (POLYPODIACEAE)
235
FIG. 1. Characters of Alansmia. A–C. A. reclinata. A. Rhizome scale with cilia and glandular cells in the margin.
B. Branched setae in the segment margin. C. Simple setae and branched hairs on the rachis. D–E. A. bradeana. D.
Branched hair of the laminar tissue. E. Sporangial capsule with cilia. F. A. reclinata, spore. (A–C from Labiak 964,
UPCB; D–E from Prance 10075, NY; F from Labiak 964, UPCB.).
synapomorphies: setae present on the rhizomes,
rhizome scales with turgid cells and ciliate
adaxial and abaxial surfaces, membranaceous
laminae, and episorangial setae (Labiak et al.,
2010) (Figs. 1E, 2H, 3J). Several other
characters, which occur frequently among
the species, are also diagnostic, including
pendent fronds with indeterminate growth
(Figs. 2B, C, G, 3A, C, K), concolorous
(orange to castaneous) rhizome scales with
ciliate or sometimes glandular margins
(Figs. 1A, 2A) (scales absent in a few species),
hydathodes that are often cretaceous (Fig. 2F),
branched hairs on the rachis and laminar tissue
(Fig. 1D), and setae that are simple (Fig. 1C),
paired, or stellate (Figs. 1B, 2D, 3D, G, H).
In keeping with the recent consensus of
dividing the grammitid ferns into genera of
tractable size defined by unique combinations
of morphological characters (Smith, 1993;
Parris, 2003, 2007; Ranker et al., 2004;
Sundue, 2010a; Labiak et al., 2010), this
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[VOL 63
FIG. 2. Characters of Alansmia. A. Alansmia diaphana, rhizome scale. B. A. Reclinata, habit. C. A. senilis. D. A.
stella var. Stella, rachis seta. E. A. Reclinata, detail of the abaxial side of the lamina. F. A. turrialbae. G. A. reclinata,
detail of apex and lower surface. H. A. reclinata, sorus showing the ciliate sporangial capsules. (A from Moguel 32,
GOET; C from Sundue 1682, NY; D from van der Werff 6448, MO; F from Sundue 1740, NY; B, E, G, and H without
voucher, from Paraná, Brazil).
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KESSLER ET AL.: ALANSMIA (POLYPODIACEAE)
237
FIG. 3. A–B. Alansmia cultrata. A. Habit. B. Detail of abaxial lamina. C–H. A. spathulata. C. Habit. D, E. Detail
of abaxial lamina. F. Frond showing branched rachis. G and H. Segment detail. J–K. A. senilis. J. Detail of abaxial
lamina. K. Habit. (A–B from Mickel 6737, NY; C–E from Pringle 4145b, NY; F–H from Bourgeau s.n., NY; J–K
from Breedlove 68357, UC.).
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BRITTONIA
easily diagnosed and well-supported group
deserves recognition at the generic level.
Therefore, we here describe Alansmia and
make the relevant combinations. The description of the new genus is credited to all four
authors because each made a significant
contribution for this part of the study,
whereas the new combinations are credited
to only Moguel and Kessler because they
conducted the taxonomic revision of the
species in this group.
Taxonomic treatment
Alansmia M. Kessler, Moguel, Sundue &
Labiak, gen. nov. Type: Alansmia lanigera (Desv) Moguel & M. Kessler.
Basionym: Polypodium lanigerum Desv.,
Ges. Naturf. Freunde Berlin Mag. Neueste
Entdeck. Gesamte Naturk. 5: 316. 1811.
Plantae epiphyticae, raro rupestres vel terrestres;
rhizomata breve repentia vel decumbentia, radiatim
symmetrica, squamata vel setosa, vel in ambobus
statibus; squamae rhizomatis luteolae, castaneae vel
nigrescentes, non clathratae, pilis margine vel raro in
pagina instructae; folia pendentia, indeterminate crescentia, plerumque pinnatisecta pinnis proximalibus gradatim reductis, adaxialiter hydathodis instructa, interdum
cum secretionibus calcareis; petioli breves vel brevissimi
(<1 cm longis); petioli laminae rhachidesque cum pilis
simplis vel furcatis, hyalinis vel atropurpureis vestiti;
sporangia vulgo setosa; sporae triletae vel ut videtur
monoletae.
Plants epiphytic, rarely saxicolous or terrestrial; roots non-proliferous; rhizomes short
creeping or very short creeping to suberect,
usually radially symmetric, with ventral or
radial root insertion, branch buds apparently
absent, scales present or sometimes absent,
setae present or absent, the scales basifixed,
concolorous, dull to shiny, yellowish, orangish, castaneous or very rarely blackish, with
variously colored setae (hyaline to atropurpureous) and/or hyaline to orangish hairs on the
margins and sometimes surfaces; blades
pendent, of indeterminate growth, not articulate to rhizome; petioles with a single
vascular bundle, usually short to very short
(<1 cm long), setose and puberulent, the setae
simple, spreading, 0.5–3.5 mm, hyaline to
castaneous, the hairs 0.1–0.2(–0.3) mm,
branched or unbranched; blades proximally
[VOL 63
gradually reduced, lacking mycelia and black
clavate ascomes of the fungus Acrospermum
maxonii, pinnatisect (except in the case of one
very deeply pinnatifid species) to 1-pinnate,
rarely 1-pinnate-pinnatifid, forking in two
species (Fig. 3F), essentially monomorphic,
setose and/or puberulent, the setae stramineous,
hyaline, yellowish, orangish, castaneous, deep
red or atropurpureous, simple, paired, irregularly branched or clustered on pinna surfaces,
pinna margins, and rachises, or rarely on the
abaxial surfaces of the rachises, also growing
out of the transverse walls of catenate axes
made of basal cells, up to 1.5 mm long, the
hairs simple or branched, hyaline to orangish or
rarely whitish-sericeous, tubular to globose,
sometimes eglandular; veins free, pinnate
(Fig. 2E); hydathodes present, sometimes cretaceous (producing whitish lime dots); sori round,
not sunken into the blade tissue, without paraphyses; sporangia setose or rarely glabrous;
spores tetrahedral-globose to ellipsoid or rarely
globose, either monolete or trilete; n = 37
(Evans, 1963).
Etymology.—We dedicate this genus to
Alan R. Smith in recognition of his exceptional
work on neotropical ferns, particularly the
grammitids.
Distribution.—Alansmia comprises 26 species of which 25 occur in the New World tropics
and two in Africa, Madagascar, and the Mascarenes. One of the latter species (A. elastica)
also occurs in the Neotropics.
Both Alansmia and its sister clade,
Leucotrichum, have soft whitish setae,
which are uncommon among grammitid
ferns, which typically have setae reddish
to castaneous and stiff (e.g., Ceradenia,
Micropolypodium, Prosaptia). Leucotrichum, however, differs from Alansmia by
fronds usually less than 10 cm long with
determinate growth, rhizome scales blackish, strongly clathrate, eciliate on both
surfaces, and the hairs 3-celled, 1-furcate,
the longer branch cell being acicular.
Whitish setae are also present in the Asian
genera Calymmodon and Tomophyllum (Parris,
1990b), and some species of the latter resemble
Alansmia. The resemblance is particularly
strong in species of Tomophyllum that have
branched hairs, and pendant, indeterminate,
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KESSLER ET AL.: ALANSMIA (POLYPODIACEAE)
and membranaceous laminae (i.e., Tomophyllum beleense (Copel.) Parris, T. polytrichum
(Copel.) Parris). However, the setae of Tomophyllum are simple, unlike those of Alansmia
that are branched.
The spores in Alansmia vary from tetrahedral-globose to ellipsoid or reniform in shape
and from 25–60 μm long. Bicellular spores
occur frequently in Alansmia, particularly in
species with ellipsoid spores. Sometimes the
usually trilete spores appear monolete or
nearly so (Wagner, 1985). These are not truly
monolete, but have an inconspicuous third
“arm” of the laesura which departs at a low
angle to the other two arms, making a narrow
Y-shape instead of the typical trilete mark
which has roughly equal 120° angles. This
phenomenon can be observed in ellipsoid or
reniform spores, whereas in tetrahedral-globose
spores a triradiate laesura is typical. Similar
variation in spore laesurae was reported to
occur in Dasygrammitis brevivenosa (Alderw.)
Parris by Nayar & Devi (1965) (as Ctenopteris
brevivenosa (Alderw.) Holttum), suggesting
that it may not be unique to Alansmia; however,
Wagner (1985) was unable to confirm this
observation.
NEW COMBINATIONS AND SYNONYMY
Alansmia alfaroi (Donn. Sm.) Moguel & M.
Kessler, comb. nov. Polypodium alfaroi
Donn. Sm., Bot. Gaz. 33: 262. 1902, as
“alfari”. Ctenopteris alfaroi (Donn. Sm.)
Copel., Philipp. J. Sci. 84: 433. 1956.
Grammitis alfaroi (Donn. Sm.) C. V.
Morton, Contr. U.S. Natl. Herb. 38: 103.
1967. Terpsichore alfaroi (Donn. Sm.) A.
R. Sm., Novon 3: 485. 1993. Type: Costa
Rica. Cártago: Sierra Alta de Navarro,
1400 m [as indicated on sheet, protologue
erroneously gives 2000 m], May 1901, A.
Alfaro 8063 (holotype: US).
Polypodium oligosorum Mett. ex Kuhn, Linnaea 36:
132. 1869, nom. illeg., non Klotzsch, 1847.
Alansmia bradeana (Labiak) Moguel & M.
Kessler, comb. nov. Terpsichore bradeana Labiak, Brittonia 52: 251. 2000.
Type: Brazil. Roraima: Serra dos Surucucus, 02º42–47’N, 63º33–36’W, 19 Feb
239
1969, G. T. Prance, J. R. Steward, J. F.
Ramos & L. G. Farias 10075 (holotype:
INPA; isotypes: K, NY).
Alansmia canescens (A. Rojas) Moguel & M.
Kessler, comb. nov. Terpsichore canescens
A. Rojas, Métodos en Ecología y Sistemática 3: 14. 2008. Type. Colombia. Santander:
Municipio de Charalá, Inspección Virolín,
Vereda El Volcán, 1900 m, 30 Jun 1983, J.
Torres 2575 (holotype: COL-n.v).
Alansmia concinna (A. R. Sm.) Moguel &
M. Kessler, comb. nov. Grammitis concinna A. R. Sm., Ann. Missouri Bot. Gard.
76: 338. 1989, nom. nov. for Polypodium
concinnum Mett. ex Kuhn, Linnaea 36:
132. 1869, non Willd. 1810. Terpsichore
concinna (A. R. Sm.) A. R. Sm., Novon 3:
486. 1993. Type: Venezuela. Aragua: High
mountains east of Colonia Tovar, 25 Jun
1854, A. Fendler 212 (lectotype, here
designated: MO; duplicates of lectotype:
K, GH-n.v., YU-n.v.).
Alansmia contacta (Copel.) Moguel & M.
Kessler, comb. nov. Ctenopteris contacta
Copel., Philipp. J. Sci. 84: 447. 1956.
Type: Bolivia. La Paz: Prov. Nor Yungas,
Yungas, 1890, M. Bang 483 (lectotype,
designated by Tryon & Stolze, 1993: US;
duplicates of lectotype: B, F-n.v., K, N).
Alansmia cultrata (Bory ex Willd.) Moguel
& M. Kessler, comb. nov. Polypodium
cultratum Bory ex Willd., Sp. Pl. ed. 4. 5:
187. 1810. Xiphopteris cultrata (Bory ex
Willd.) Schelpe, Bol. Soc. Brot., ser. 2,
41: 217. 1867. Ctenopteris cultrata (Bory
ex Willd.) Copel., Gen. Fil. 219. 1947.
Grammitis cultrata (Bory ex Willd.)
Proctor, Rhodora 63: 35. 1961. Terpsichore cultrata (Bory ex Willd.) A. R. Sm.,
Novon 3: 486. 1993. Type: Jamaica or
Martinique, without precise locality, O.
Swartz s.n. (lectotype, designated by Proctor, 1985: B; duplicate of lectotype: S-n.v).
Alansmia dependens (Baker) Moguel & M.
Kessler, comb. nov. Polypodium dependens Baker, Syn. Fil. 335. 1867. Grammitis
dependens (Baker) C. V. Morton, Contr.
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BRITTONIA
U. S. Natl. Herb. 38: 104. 1967. Xiphopteris heteromorpha var. dependens
(Baker) Crabbe, Brit. Fern Gaz. 9: 318.
1967. Terpsichore dependens (Baker) A.
R. Sm., Novon 3: 486. 1993. Type.
Ecuador. Pichincha: Mt. Pichincha, R.
Spruce 5637 (holotype: K).
Alansmia diaphana (Moguel & M. Kessler)
Moguel & M. Kessler, comb. nov. Terpsichore diaphana Moguel & M. Kessler,
Phytotaxa 2: 38. 2009. Type. Bolivia. La
Paz: Prov. Nor Yungas, hill above
Cotapata on La Paz-Caranavi road, 16°
17’S, 67°50’W, 3200 m, 15 Aug 1990, A.
Fay & L. Fay 3031 (holotype: LPB;
isotypes: LPB, MO, NY, UC).
Alansmia elastica (Bory ex Willd.) Moguel
& M. Kessler, comb. nov. Polypodium
elasticum Bory ex Willd., Sp. Pl., ed. 4. 5:
183. 1810. Polypodium leveilleanum
Desv., Mém. Soc. Linn. Paris 6: 232.
1827, nom superfl. Ctenopteris elastica
(Bory ex Willd.) Copel., Philipp. J. Sci.
84: 426. 1956. Xiphopteris elastica (Bory
ex. Willd.) Alston, Bol. Soc. Brot., ser. 2,
26: 27. 1956. Terpsichore elastica (Bory
ex Willd.) A. R. Sm., Novon 5: 21.
1995. Type: Réunion [“Bourbon”]: forêts
élevée, J. B. G. M. Bory s.n. (holotype:
B-W 19675).
Polypodium mollissimum Fée, Hist. Foug. Antill.
[Mém. Foug. 11]: 47, t. 12, f. 2. 1866. Ctenopteris
mollissima (Fée) Copel., Philipp. J. Sci. 84: 423.
1956. Grammitis mollissima (Fée) Proctor,
Rhodora 63: 35. 1961. Terpsichore mollissima
(Fée) A. R. Sm., Novon 3: 487. 1993. Type:
Guadeloupe: Bois élevés de la [illegible] St.
Louis, bras [illegible], 1861, F. L’Herminier 13
(lectotype, designated by Proctor, 1985: P;
duplicates of lectotype: B, K-n.v., US-n.v.).
Polypodium flexile Fée, Synopsis Filicum 54 [Mem.
Foug. 2]: 9, t. 2, f. 3. 1854. Type: Mauritius: F. W.
Sieber 54 (lectotype, here designated: B).
Terpsichore cocosensis A. Rojas, Brenesia 45-46:
47. 1996. Type: Costa Rica. Puntarenas: Isla del
Coco, Cerro Pelón, 5°32’40”N, 87°03’20”W,
200–400 m, 13 Jun 1994, E. Lépiz 337 (holotype:
INB-n.v.; isotypes: CR-n.v., MO-n.v., UC).
Terpsichore esquiveliana A. Rojas, Rev. Biol. Trop.
49: 451. 2001. Type: Costa Rica. Cartago: Paraíso,
Parque Nacional Tapantí, Sendero los Palmitos
(o T6), 09º44’00"N, 83º46’05"W, 1400–1600 m, 3
Sep 1997, A. Rojas 3785 (holotype: INB; isotypes:
CR-n.v., MO-n.v., UC-n.v.).
[VOL 63
Terpsichore lobulata A. Rojas, Métodos en Ecología
y Sistemática 3: 14. 2008. Type. Colombia. Norte
de Santander: Cordillera Oriental, región del
Sararé, Hoya del Río Cubugón, El Indio, 420–
480 m, 13–17 Nov 1941, J. Cuatrecasas 13089
(holotype: COL-n.v.).
Alansmia glandulifera (A. Rojas) Moguel &
M. Kessler, comb. nov. Terpsichore glandulifera A. Rojas, Lankesteriana 6: 96.
2006. Type: Costa Rica. Cartago: Paraíso, Parque Nacional Tapantí, cuenca
del Río Reventazón, Orosi, sendero
Palmitos o T6, 9°43 ’3 5"N, 83°
46’28"W, 1460–1600 m, 3 Sep 1997,
A. Rojas & R. Delgado 3788 (holotype:
CR-n.v.; isotypes: INB-n.v., MO-n.v.).
Alansmia heteromorpha (Hook. & Grev.)
Moguel & M. Kessler, comb. nov. Polypodium heteromorphum Hook. & Grev.,
Icon. Fil. 1: t. 108. 1829. Ctenopteris
heteromorpha (Hook. & Grev.) Copel.,
Philipp. J. Sci. 84: 412. 1956. Grammitis
heteromorpha (Hook. & Grev.) C. V.
Morton, Contr. U. S. Natl. Herb. 38:
102. 1967. Xiphopteris heteromorpha
(Hook. & Grev.) Crabbe, Brit. Fern Gaz.
9: 318. 1967. Terpsichore heteromorpha
(Hook. & Grev.) A. R. Sm., Novon 3:
486. 1993. Type: Ecuador. Pichincha: Mt.
Pichincha, between Mt. Chimborazo and
Pichincha, W. Jameson s.n. (lectotype,
designated by Morton, 1967: K; duplicates of lectotype: FI-n.v., NY).
Alansmia immixta (Stolze) Moguel & M.
Kessler, comb. nov. Grammitis immixta
Stolze, Fieldiana, Bot., n.s. 32: 115. 1993.
Terpsichore immixta (Stolze) A. R. Sm.,
Novon 3: 487. 1993. Type: Peru. Cuzco: La
Convención, Vilcabamba, Ruina Idma-huasi,
4000 m, Jul 1934, G. Bües 2103 (holotype:
US; isotypes: F-n.v., GH-n.v., MO, NY).
Alansmia kirkii (Parris) Moguel & M.
Kessler, comb. nov. Terpsichore kirkii
Parris, Kew Bulletin 57: 431. 2002.
Type: Malawi. Shire, Zambesi Exp., J.
Kirk s.n. (holotype: K).
Alansmia lanigera (Desv.) Moguel & M.
Kessler, comb. nov. Polypodium lanigerum Desv., Ges. Naturf. Freunde Berlin
2011]
KESSLER ET AL.: ALANSMIA (POLYPODIACEAE)
Mag. Neueste Entdeck. Gesamte Naturk.
5: 316. 1811. Ctenopteris lanigera (Desv.)
Copel., Philipp. J. Sci. 84: 420. 1956.
Grammitis lanigera (Desv.) C. V. Morton,
Contr. U.S. Natl. Herb. 38: 105. 1967.
Xiphopteris lanigera (Desv.) Crabbe, Brit.
Fern Gaz. 9: 319. 1967. Terpsichore
lanigera (Desv.) A. R. Sm., Novon 3:
487. 1993. Type: Peru. without precise
locality, J. Dombey s.n. [Herbarium
Jussieu 1099] (lectotype, designated by
Morton, 1967: P; duplicates of lectotype: B, P [P00279983]).
Polypodium sericeolanatum Hooker, Sp. Fil. 4: 221.
1864. Type: Ecuador. Pichincha: ravines of
Pichincha, 3900–4000 m, W. Jameson 235 (lectotype, designated by Morton, 1967: K; duplicate of
lectotype: B).
Alansmia laxa (C. Presl) Moguel & M.
Kessler, comb. nov. Polypodium laxum
C. Presl, Reliq. Haenk. 1: 23, t. 4, f. 1.
1825. Grammitis laxa (C. Presl) C. V.
Morton, Contr. U.S. Natl. Herb. 38: 105.
1967. Terpsichore laxa (C. Presl) A. R.
Sm., Novon 3: 487. 1993. Type. Peru.
without precise locality, T. Haenke s. n.
(lectotype, designated by Tryon & Stolze,
1993: PRC; duplicate of lectotype: PR).
Alansmia longa (C. Chr.) Moguel & M.
Kessler, comb. nov. Polypodium longum
C. Chr., Index. Fil. 541. 1906, nom. nov. for
P. alternifolium Hook. non Willd., 1810.
Ctenopteris longa (C. Chr.) Copel., Philipp.
J. Sci. 84: 453. 1956. Xiphopteris longa
(C. Chr.) Alston, Bull. Jard. Bot. Etat 27: 35.
1937. Terpsichore longa (C. Chr.) A. R.
Sm., Novon 3: 487. 1993. Type: Ecuador.
Near Esmeraldas and between Cuenca and
Guayaquil, 1000–3300 m, W. Jameson s. n.
(lectotype, designated by Morton, 1967: K,
duplicate of lectotype: US-n.v.).
241
sichore monosora Moguel & M. Kessler,
Phytotaxa 2: 40. 2009. Type. Ecuador.
Pichincha: Andes of Quito, 4000 m, Jun
1877, W. Jameson s.n. [Herbarium Hookerianum 1867] (holotype: K barcode:
K000590671; isotype: K barcode:
K000590672 [Herbarium Hookerianum
1854]).
Alansmia reclinata (Brack.) Moguel & M.
Kessler, comb. nov. Polypodium reclinatum Brack., U. S. Expl. Exped. 16: 11. 1854.
Ctenopteris reclinata (Brack.) Copel.,
Philipp. J. Sci. 84: 424. 1956. Terpsichore
reclinata (Brack.) Labiak, Brittonia 52: 253.
2000. Type: Brazil. Rio de Janeiro. Serra dos
Orgaos, Wilkes Expedition 161. (lectotype,
designated by Copeland, 1956: US; duplicates of lectotype: K, NY).
Polypodium ciliare Fée, Crypt. Vasc. Brésil 1: 94,
t. 27, f. 2. 1869. Ctenopteris ciliare (Fée) Copel.,
Philipp. J. Sci. 84: 425. 1955[1956]. Type: Brazil.
Rio de Janeiro: Habitat in Brasilia Fluminensi, A.
Glaziou 961 (lectotype, here designated: P;
duplicates of lectotype: BR, NY).
Polypodium ovalescens Fée, Crypt. Vasc. Brésil 1:
94, t. 27, f. 3. 1869. Type: Brazil. Rio de Janeiro:
Habitat in Brasilia Fluminensi, A. Glaziou 1722
(lectotype, here designated: P [P00279989];
duplicates of lectotype: BR, P-n.v. [P00632688]).
Alansmia semilunaris (Moguel & M.
Kessler) Moguel & M. Kessler, comb.
nov. Terpsichore semilunaris Moguel &
M. Kessler, Phytotaxa 2: 42. 2009. Type.
Ecuador. Napo: ca. 3 kmW of Oyacachi,
00°12’S, 78°06’W, 3550 m, 27 Mar 1996,
B. Ståhl & H. Navarrete 2295 (holotype:
AAU; isotype: QCA-n.v.).
Terpsichore acrosora A. Rojas, Métodos en Ecología
y Sistemática 3: 13. 2008. Type: Costa Rica.
Limón: Talamanca, Bratsi, Amubri, Alto Lari,
Kivut, quebrada innominada, margen derecha del
Río Dapari, 9°24'20”N, 83°05'35”W, 1000 m, 11
Mar 1992, G. Herrera 5302 (holotype: CR-n.v.;
isotypes: INB-n.v., MO).
Alansmia senilis (Fée) Moguel & M.
Kessler, comb. nov. Polypodium senile
Fée, Mém. Soc. Sci. Hist. Nat. Strasbourg
5 [Mém. Foug. 7]: 60, t. 25, f. 1. 1857.
Ctenopteris senilis (Fée) Copel., Philipp.
J. Sci. 84: 398. 1956. Grammitis senilis
(Fée) C. V. Morton, Contr. U.S. Natl. Herb.
38: 103. 1967. Terpsichore senilis (Fée) A.
R. Sm., Novon 3: 488. 1993. Type: Colombia. Norte de Santander: Ocaña, L. J. Schlim
364 (holotype: RB).
Alansmia monosora (Moguel & M. Kessler)
Moguel & M. Kessler, comb. nov. Terp-
Polypodium subflabelliforme Rosenst. Repert. Spec.
Nov. Regni. Veg. 7: 306. 1909. Ctenopteris
242
BRITTONIA
subflabelliformis (Rosenst.) Copel., Philipp. J.
Sci. 84: 400. 1956. Type: Ecuador. Tungurahua:
Cerro de Abitagua, Andium Quitensium, R.
Spruce 5271 (lectotype, designated by Tryon &
Stolze, 1993: K; duplicate of lectotype: NY).
Polypodium subflabelliforme Rosenst. var. minor
Rosenst., Spec. Nov. Regni. Veg. 7: 307. 1909.
Type: Ecuador: Tungurahua: In Monte Tungurahua,
Andium Quitensium, R. Spruce 5272 (holotype: K).
Polypodium senile Fée var. minor Rosenst.,
Mededeelingen van’s Rijks Herbarium 19: 19.
1913. Type: Bolivia. Cochabamba: Yungas de San
Mateo, 2700 m, Apr 1911, T. Herzog 1985
(lectotype, designated by Tryon & Stolze,
1993: L-n.v.; duplicates of lectotype: B, US).
Alansmia smithii (A. Rojas) Moguel & M.
Kessler, comb. nov. Terpsichore smithii
A. Rojas, Métodos en Ecología y
Sistemática 3: 15. 2008. Type. Costa Rica.
Cartago: El Guarco, Madre Selva, km 64,
Finca Los Lagos, sendero a orillas del río,
9°40'34”N, 83°52'38”W, 2600 m, 21 Apr
1999, A. Rojas & L. Pacheco 5090
(holotype: CR-n.v.; isotypes: INB-n.v.,
MO-n.v.).
Alansmia spathulata (A. R. Sm.) Moguel &
M. Kessler, comb. nov. Terpsichore spathulata A. R. Sm., Novon 3: 483–485.
1993. Type: México. State of México:
Summit of Sierra de las Cruces, 3800 m,
11 Sep 1892, C. G. Pringle 4145b
(holotype: UC; isotypes: K, MO, NY,
US).
Alansmia stella (Copel.) Moguel & M.
Kessler var. stella, comb. nov. Ctenopteris
stella Copel., Philipp. J. Sci. 84: 452. 1956.
Terpsichore stella (Copel.) Moguel & M.
Kessler, Phytotaxa 2: 36. 2009. Type: Peru.
Cuzco: Valley of Río Urubamba, A. Bües 32
(holotype: US).
Ctenopteris fabispora Copel., Philipp. J. Sci. 84: 457.
1956. Terpsichore fabispora (Copel.) A. Rojas.
Type. Panama. Chiriquí: between Alto de Las
Palmas and the top of Cerro Horqueta, 2100–
2268 m, Mar 1911, W. Maxon 5479 (holotype: US).
Alansmia stella (Copel.) Moguel & M.
Kessler var. flava (Moguel & M. Kessler)
Moguel & M. Kessler, comb. nov. Terpsichore stella (Copel.) Moguel & M.
Kessler var. flava Moguel & Kessler,
[VOL 63
Phytotaxa 2: 37. 2009. Type: Peru. Amazonas: Road Chachapoyas-Mendoza, a
little past Molinopampa, 06°14’S, 77°
35’W, 2400 m, 15 Mar 1998, H. van der
Werff, B. Gray, R. Vasquez & R. Rojas
15009 (holotype: MO-n.v.; isotype: UC).
Alansmia turrialbae (H. Christ) Moguel &
M. Kessler, comb. nov. Polypodium
turrialbae H. Christ, Bull. Soc. Roy.
Bot. Belgique 35: Mém 226. 1896. Ctenopteris turrialbae (H. Christ) Copel.,
Philipp. J. Sci. 84: 419. 1956. Grammitis
turrialbae (H. Christ) F. Seym., Phytologia 31: 181. 1975. Terpsichore turrialbae
(H. Christ) A. R. Sm., Novon 3: 488.
1993. Type: Costa Rica. Cartago: Troncs
d’arbres au pied du volcan de Turrialba,
2750 m, 26 Jan 1889, H. F. Pittier 847
(lectotype, designated by Lellinger, 1989:
BR; duplicate of lectotype: US).
Polypodium exsudans H. Christ, Bull. Herb. Boiss.
sér. 2, 4: 1103. 1904. Type: Costa Rica. C.
Wercklé s.n. (holotype: P).
Alansmia variabilis (Mett. ex Kuhn) Moguel
& M. Kessler, comb. nov. Polypodium
variabile Mett. ex Kuhn, Linnaea 36: 133.
1869. Grammitis variabilis (Mett. ex
Kuhn) C. V. Morton, Contr. U.S. Natl.
Herb. 38: 102. 1967. Terpsichore variabilis (Mett. ex Kuhn) A. R. Sm., Novon 3:
488. 1993. Type: Colombia. [Depto.
Caldas or Tolima]: Páramo del Ruiz,
Jun 1826, A. W. Purdie s.n. (lectotype,
here designated: K; duplicate of lectotype: B).
Alansmia xanthotrichia (Klotzsch) Moguel
& M. Kessler, comb. nov. Polypodium
xanthotrichium Klotzsch, Linnaea 20:
376. 1847. Grammitis xanthotrichia
(Klotzsch) Duek & Lellinger, Amer. Fern.
J. 68: 120. 1978. Terpsichore xanthotrichia
(Klotzsch) A. R. Sm., Novon 3: 488. 1993.
Type: Venezuela. Mérida: J. W. K. Moritz
250 (holotype: B).
Polypodium ellipticosorum Fée, Mem. Foug. [Gen.
Filic.] 5: 239. 1850. Type: Venezuela. Mérida:
2000 m, N. Funck & L. J. Schlim 960 (lectotype,
here designated: K; duplicate of lectotype: P).
2011]
KESSLER ET AL.: ALANSMIA (POLYPODIACEAE)
Acknowledgments
We thank Alan R. Smith for his unconditional support of our pteridological studies.
Kessler’s research was supported by the
German Research Association (DFG),
Labiak’s research by a grant from the Brazilian CNPq (PED 201782/2008-1), and Sundue’s research by a grant from the United
States National Science Foundation (NSF) (in
the name of Dr. Robbin C. Moran, DEB
0717056). We thank two anonymous reviewers
for their valuable comments, Patricia Eckel for
revising the Latin diagnosis, and Mr. Haruto
Fukuda for providing the line art illustration as
well as John Mickel for allowing us to
reproduce it here.
Literature Cited
Bishop, L. E. 1988. Ceradenia, a new genus of
Grammitidaceae. American Fern Journal 78: 1–5.
———. 1989. Zygophlebia, a new genus of the
Grammitidaceae. American Fern Journal 79: 103–
118.
——— & A. R. Smith. 1992. Revision of the fern genus
Enterosora (Grammitidaceae) in the New World.
Systematic Botany 17: 345–362.
Ching, R. C. 1940. On natural classification of the
family “Polypodiaceae.” Sunyatsenia 5: 201–268.
Copeland, E. B. 1947. Genera Filicum. Chronica
Botanica, Waltham Mass.
———. 1956 [“1955”]. Ctenopteris in America. Philippine Journal of Science 84: 381–473.
Evans, A. M. 1963. New chromosome observations in
the Polypodiaceae and Grammitidaceae. Caryologia
16: 671–677.
Hasebe, M., P. G. Wolf, K. M. Pryer, K. Ueda, M. Ito,
R. Sano, G. J. Gastony, J. Yokoyama, J. R.
Manhart, N. Murakami, E. H. Crane, C. H.
Haufler & W. D. Hauk. 1995. Fern phylogeny
based on rbcL nucleotide sequences. American Fern
Journal 85: 134–181.
Holttum, R. E. 1947. A revised classification of
leptosporangiate ferns. Botanical Journal of the
Linnean Society 53: 123–158.
Labiak, P. H. & F. B. Matos. 2007. A new hybrid and
two new combinations in neotropical grammitid
ferns. Brittonia 59: 182–185.
———, G. Rouhan & M. Sundue. 2010. Phylogeny and
taxonomy of Leucotrichum (Polypodiaceae), a new
genus of grammitid ferns from the neotropics. Taxon
59: 911–921.
Lellinger, D. B. 1989. The ferns and fern-allies of
Costa Rica, Panama and the Chocó (Part 1:
Psilotaceae through Dicksoniaceae). Pteridologia
2A: 1–364.
243
Mettenius, G. H. 1866. Filices; praesertim Indicae et
Japonicae. Annales Museum Botanicum LugdunoBatavi 2: 149–313.
Morton, C. V. 1967. The genus Grammitis in Ecuador.
Contributions from the United States National Herbarium 38: 85–120.
Murillo, M. T. & A. R. Smith. 2003. Luisma, a new
genus of Grammitidaceae (Pteridophyta) from
Colombia. Novon 13: 313–317.
Nayar, B. K. & S. Devi. 1965. Spore morphology of
Indian ferns. IV. Grammitidaceae. Grana Palynologica 6: 121–127.
Newman, E. 1840. A history of British ferns and allied
plants. J. Van Voorst, London.
Parris, B. S. 1990a. Grammitidaceae. Pp. 153–157. In:
K. Kubitzki (ed.), The Families and Genera of
Vascular Plants, Vol. 1. Springer-Verlag, Berlin,
Heidelberg, New York.
Parris, B. S. 1990b. Noteworthy species of Grammitidaceae from South-East Asia. Hooker’s Icones
Plantarum 40: 1–129.
———. 2003. Distribution of Grammitidaceae inside
and outside Malesia. Telopea 10: 451–466.
———.2007. Five new genera and three new species of
Grammitidaceae (Filicales) and the re-establishment
of Oreogrammitis. Gardens’ Bulletin Singapore 58:
238–274.
———. 2009. New genera of Malesian Grammitidaceae
(Monilophyta). Blumea 54: 217–219.
Proctor, G. R. 1985. Ferns of Jamaica. British Museum
(Natural History), London.
Ranker, T. A., A. R. Smith, B. S. Parris, J. M. O.
Geiger, C. H. Haufler, S. C. K. Straub & H.
Schneider. 2004. Phylogeny and evolution of grammitid ferns (Grammitidaceae): a case of rampant
morphological homoplasy. Taxon 53: 415–428.
Schneider, H., A. R. Smith, R. Cranfill, T. J.
Hildebrand, C. H. Haufler & T. A. Ranker. 2004.
Unravelling the phylogeny of the polygrammoid ferns
(Polypodiaceae and Grammitidaceae): exploring
aspects of the diversification of epiphytic plants.
Molecular Phylogenetics & Evolution 31: 1041–1063.
Smith, A. R. 1992. A review of the fern genus Micropolypodium (Grammitidaceae). Novon 2: 419–425.
———. 1993, Terpsichore, a new genus of Grammitidaceae (Pteridophyta). Novon 3: 478–489.
——— & R. C. Moran. 1992. Melpomene, a new genus
of Grammitidaceae (Pteridophyta). Novon 2: 426–432.
———, ——— & L. E. Bishop. 1991. Lellingeria, a
new genus of Grammitidaceae. American Fern
Journal 81: 76–88.
Sundue, M. A. 2010a. A monograph of Ascogrammitis,
a new genus of grammitid ferns (Polypodiaceae).
Brittonia.
———. 2010b. A morphological cladistic analysis of
Terpsichore (Polypodiaceae). Systematic Botany (in
press).
———, M. Islam & T. A. Ranker. 2010. Systematics
of grammitid ferns (Polypodiaceae): using morphology and plastid sequence data to resolve the
circumscription of Melpomene and the polyphyletic
genera Lellingeria and Terpsichore. Systematic
Botany (in press).
244
BRITTONIA
Tryon, R. M. & A. F. Tryon. 1982. Ferns and allied
plants with special reference to tropical America.
Springer-Verlag, New York.
——— & R. G. Stolze, 1993. Pteridophyta of Peru. Part
V. 18. Aspleniaceae–21. Polypodiaceae. Fieldiana,
Botany, New Series, 27: 1–176.
[VOL 63
Wagner, F. S. 1985. Bilateral spores in new world
grammitid ferns. American Fern Journal 75: 6–11.
Wilson, K. A. 1959. Sporangia of the fern genera
allied to Polypodium and Vittaria. Contributions of
the Gray Herbarium of Harvard University 185:
97–127.
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