Phytotaxa 61: 17–36 (2012) ISSN 1179-3155 (print edition)

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Copyright © 2012 Magnolia Press
Article PHYTOTAXA
ISSN 1179-3163 (online edition)

A synopsis of the species of Cyathea (Cyatheaceae-Polypodiopsida) with pinnate

to pinnate-pinnatifid fronds
MARCUS LEHNERT

Staatliches Museum für Naturkunde Stuttgart, Abt. Botanik, Am Löwentor, Rosenstein 1, D-70191 Stuttgart, Germany.
E-mail: [email protected]

Abstract

A summary of the species of Cyathea sensu strictu with pinnate to pinnate-pinnatifid fronds is presented. All species are
strictly Neotropical and are found throughout the entire range of the genus in the Western Hemisphere, but most species
occur in the mountainous regions of northern South America and Central America. This artificially delimited group
includes the species formerly recognized as the segregate genus Cnemidaria, which form a monophyletic group within
Cyathea together with some species previously not recognized as Cnemidaria. This group is characterized by a general
lack of hairs, cartilaginous laminar texture, petiole scales that have white margins or that are completely white, and large
sori with diameter ≥ 1 mm and 30–40 sporangia per sorus. Cnemidaria-type spores with at least three large equatorial
pores dominate in this group but are neither omnipresent nor exclusive to it. The remainder of the pinnate to pinnate-
pinnatifid species of Cyathea includes predominantly exindusiate species that have moderately to densely pilose laminae
and relatively small sori with diameter ≤ 1 mm and ± 20 sporangia per sorus.
Twelve new combinations are made and the following three new names are proposed: Cyathea cnemidaria (=
Cnemidaria tryoniana), C. glandulifera (= Cn. glandulosa), C. suprapilosa (= Cn. suprastrigosa). A key to all species of
Cyathea with pinnate to pinnate-pinnatifid fronds is provided. Hybrids are not treated.

Keywords: Andes, hybrids, tree ferns

Introduction

The family Cyatheaceae, or scaly tree ferns, contains ca. 600 species that are distributed in the humid tropics
and the southern temperate regions. The taxonomic history of the family is marked by frequent changes in the
number and definition of its genera, with various attempts to form a classification based on the different
shapes of indusia (e.g. Christ 1897). Holttum (1957), Holttum & Sen (1961) and Sen (1964) discovered that
the morphology of the petiole scales offers a more reliable method of grouping the species of Cyatheaceae,
because it correlates with other morphological characters and ecological preferences. Holttum (1963),
focussing on Old World taxa, used the scale characters to define subgenera and sections in the then universal
genus Cyathea (Domin 1930). Tryon (1970) made a more daring attempt dividing the entire family into six
genera (eight at first, including Lophosoria Presl (1847: 37) and Metaxya Presl (1836: 59)). His work was
based mainly on the Neotropical species, where a different evolutionary history had lead to clearer differences
between the natural groups. However, Tryon made some poor choices in recognizing genera based on weak
morphological differences (Holttum & Edwards 1983, Lellinger 1987). This fuelled a decade-long dispute
over the two generic systems (Holttum & Tryon 1977), which was eventually settled with a truce but was not
resolved (Holttum & Edwards 1983).
Recent phylogenetic studies (Korall et al. 2006, 2007) favour the taxonomic recognition of at least four
clades that correspond with the established genera Sphaeropteris Bernhardi (1801: 122), Alsophila Brown
(1810: 158), Gymnosphaera Blume (1828: 242), and Cyathea Smith (1793: 416) sensu strictu. The latter genus

Accepted by Maarten Christenhusz, 16 Jul. 2012; published online in PDF: 30 Jul. 2012 17
is almost exclusively Neotropical except for eight species that occur in western Pacific Islands, New Guinea,
and northern Australia (Lehnert 2011a), and includes the obsolete genera Cnemidaria (Stolze 1974),
Hymenophyllopsis (Goebel 1929), Trichipteris (Barrington 1978), and Sphaeropteris subg. Sclephropteris
(Windisch 1977, 1978). The inclusion of Cnemidaria and Hymenophyllopsis in Cyathea was unexpected
because of their distinct morphology (Korall et al. 2007, Christenhusz 2009a).
Hymenophyllopsis is characterized by thin, deeply dissected laminae with marginal sori and bivalved
indusia and contains eight species, which are restricted to the table mountains (“tepuis”) of the Guayana
Highlands (Kramer & Green 1990). Its phylogenetic position was long unclear and was placed therefore in a
separate family. They may be maintained as a subgenus of Cyathea with an unaltered morphological
circumscription (Christenhusz 2009a).
The first and only revision of Cnemidaria recognized 23 species with ten varieties (Stolze 1974). Today
Cnemidaria in the traditional sense would contain 27 species and eight varieties, with four species added since
(Gómez & Gómez-L. 1982, Stolze 1984, Moran 1991) and one newly recognized at the species level (Rojas
2005). The species are characterized by a relatively stiff laminar texture, a general lack of pluricellular hairs,
and spores with three large equatorial pores (Tryon 1970, Stolze 1974). This spore type was long believed to
be the most reliable distinguishing character of Cnemidaria and fossil spores of this type (Kuyrilosporites)
were used for hypothesizing historic distributions and estimating divergence times of the genus (Korall et al.
2007, Janssen et al. 2008).
However, the distinguishing characters of the Cnemidaria-clade are ambiguous. Stolze (1974) not only
mentioned Cyathea speciosa as Cnemidaria look-alike, which has been proven by molecular evidence to truly
belong to the Cnemidaria clade (Korall et al. 2007), but also Cyathea williamsii (Maxon 1922: 46) Domin
(1930: 171), and C. maguirei (Tryon 1972: 447) Lehnert, comb. nov. After having revised material of all
Neotropical species of Cyathea, I agree with Stolze’s account and can add C. stolzei A.R.Sm. ex Lellinger
(1987[1988]: 101) from Costa Rica to this list. Vice versa, C. amabilis (C.V.Morton) Lehnert, comb. nov.,
which Stolze (1974) considered to be the “most primitive” of Cnemidaria-species, bears a strong resemblance
to C. bipinnatifida (Baker 1874: 456) Domin (1929b: 262) and allies, and C. suprapilosa Lehnert, nom. nov.
(= Cnemidaria suprastrigosa) and C. (= Cn.) chiricana (Maxon 1912: 33) Domin (1929b: 263) have hairs.
These ambiguous species have small geographical ranges and are often elusive in the field, so no fresh
material was hitherto available to test at the molecular level if they actually belong to the Cnemidaria-clade or
if their coinciding morphology is merely analogous. Furthermore, in a largely neglected publication Gastony
(1979) demonstrated that spores of the Cnemidaria-type are also found in extant members of the paleotropical
Cyathea decurrens (Hooker 1844: 51) Copeland (1929: 356)-group, which are sister to all remaining Cyathea
sensu strictu, including Cnemidaria (Korall et al. 2007). On the other hand, Cyathea speciosa Willdenow
(1810: 490) of the Cnemidaria-clade has only finely porate spores (Gastony 1979).
These observations indicate that some species previously treated as Cnemidaria may actually not belong
to the Cnemidaria-clade. Consequently, a formal recognition of a subgenus Cnemidaria at this point seems
premature and should await a phylogenetic study that covers a larger sampling of Cyatheaceae, to put a
subgeneric division on a sounder basis. In order to facilitate taxonomic studies and there upon depending
phylogenetic studies, I will shortly summarize all simply pinnate to pinnate-pinnatifid species of Cyathea
sensu strictu (Korall et al. 2008, Lehnert 2011a) in the following and provide a key to these 57 species and
eight varieties. The nothogenus ×Cyathidaria (Caluff 2002) is rendered obsolete by the merging of
Cnemidaria into Cyathea.

Materials and methods

Plant material from following herbaria was revised for this study: AAU, B, BM, CAY, COL, CUZ, F, GH,
GOET, K, MO, P, PR, PRC, QCA, QCNE, QPLS, SP, UC, USM and Z. Online resources were used for
checking typifications (plants.jstor.org) and references for the species (www.ipni.org, www.tropicos.org).

18 • Phytotaxa 61 © 2012 Magnolia Press LEHNERT

The terminology generally follows Lellinger (2002), but some peculiarities have to be shortly explained here.
As the species are all simply pinnate and thus the segments or lobes of the pinnae correspond with the
pinnules of species with bipinnate or more complex laminae, there arises an ambiguity in the application of
the terms costules (main veins braching off the costae) and midveins (main veins of ultimate segments). The
latter term is preferred here.
The term costal is replaced here with the term proximal for the soral position, but is kept for describing the
position of areoles formed by anastomosing veins. Veins can fuse to form areoles either only between the
basal veins of the segments next to the costae (costal areoles) or also between distal veins (lateral areoles).
The laminar apices may be gradually reduced or abruptly reduced. In the latter case, the apical section may
still be more complex than the lateral pinnae, i.e. coarsely dissected or lobed (non-conform apex), or just like
a lateral pinna (conform apex) (Stolze 1974).
The key is designed to lay an emphasis on vegetative characters. Characters of sori and indusia are added
in brackets if they seemed dispensable for general identification. The key does not include hybrids because
more studies are deemed necessary before a satisfying account can be given. However, the key includes eight
species with variable frond dissection that are not considered to be hybrids. These species have predominantly
bipinnate or more complex fronds and are thus assigned to other groups that will be treated in more detail in
subsequent publications. Hence the synonymy is omitted for these species in the systematic treatment.

Key to the Neotropical species of Cyathea with pinnate to pinnate-pinnatifid fronds

For a key to the genera of Neotropical Cyatheaceae see Lehnert (2011b); for a key to the groups of the genus
Cyathea sensu strictu see Lehnert (2011a).

1 Rhachises densely covered with spreading, basally attached scales 5–8(–10) × 0.5–1.0 mm; pinnae 25–50 pairs per
frond, veins all free [exindusiate] ..........................................................................................................Cyathea ursina
– Rhachises not covered with spreading scales or scales few and appressed; pinnae mostly less than 25 pairs per frond,
if rarely up to 35 pairs then veins forming areoles ....................................................................................................... 2
2 Fronds spirally arranged along a thin climbing rhizome (1.5–2.0 cm diameter), scales weakly discordantly bicolor-
ous, yellowish white to stramineous with brown central stripes; laminae abruptly reduced to a pinna-like section .....
........................................................................................................................................................ Cyathea assurgens
– Fronds clustered at rhizome/trunk apices, scales often darker, concolorous or bicolorous with white margins; if
scales pale and trunk thin then laminae gradually reduced........................................................................................... 3
3 Rhachises notably green-alate throughout or in most parts.......................................................................................... 4
– Rhachises not green-alate or only weakly so in distal parts ....................................................................................... 12
4 Pinnae less than 10 cm long and to 2.2 cm (mostly less than 2 cm) broad; rhachises glabrous or hairy with hairs to
1.2 mm long; exindusiate .............................................................................................................................................. 5
– Pinnae 10–33 cm long and to 3.6 cm broad (sometimes only 1.6 cm broad); rhachises glabrous on both sides, or
sparsely hairy with hyaline hairs to 0.5 mm long, indusiate........................................................................................ 6
5 Rhachises completely glabrous; pinnae 9–12(–20) pairs per frond, to 6.5 × 2.2 cm ...........................Cyathea nanna
– Rhachises densely hairy on both sides, hairs adaxially 0.5–1.0 mm long, brown to tan, antrorsely curved, abaxially
1.0–1.2 mm long, white; pinnae 14–25 pairs per frond, to 7.5 × 1.4–1.8 cm............................ Cyathea pseudonanna
6 All veins free, rarely forming costal areoles................................................................................................................. 7
– Veins regularly merging to form costal, sometimes also lateral areoles....................................................................... 9
7 Pinnae 16–22(–26) pairs per frond [pinnae incised to 3/4 towards the costae, medial ones opposite to subopposite,
distal and proximal ones alternate; pinna bases equilateral, truncate]..............................................Cyathea chiricana
– Pinnae 5–12(–20) pairs per frond ................................................................................................................................. 8
8 Pinnae 5–6 pairs per frond, alternate; sori 0.8 mm diameter, medial, parallel to the midrib; indusia hemitelioid,
reaching to 1/4 around the receptacles..................................................................................................Cyathea minuta
– Pinnae 9–12 pairs per frond, mostly opposite; sori 1.2–1.5 mm diam., in a triangular pattern medial to the midveins;
indusia reaching 1/2 to 2/3 or more around the receptacles......................................................... Cyathea glandulifera
9 Pinnae 14–26 pairs per frond [pinnae incised to 1/4 towards the costae, pinna bases inequilateral with the basiscopi-
cal side pronounced as small auricle] ......................................................................................................................... 10
– Pinnae 5–12(–20) pairs per frond .............................................................................................................................. 11

A SYNOPSIS OF THE SPECIES OF CYATHEA Phytotaxa 61 © 2012 Magnolia Press • 19

10 Petioles not green-alate; pinnae to 16 × 1.6–2.6 cm, strictly opposite; laminae gradually reduced.... Cyathea varians
– Petioles usually green-alate throughout; pinnae to 33.0 × 3.0–3.6 cm, alternate to opposite; laminae gradually
reduced to abruptly reduced to conform apical sections.......................................................... Cyathea decurrentiloba
11 Petiole scales concordantly to discordantly bicolorous, dark brown to castaneous with white margins, in distal peti-
ole parts concolorous white or with a discontinuous brown central streak [indusia reaching 1/2 around the recepta-
cles, rarely more, midveins with 1 cm or more between them] .......................................................Cyathea alatissima
– Petiole scales almost concolorous dark brown to castaneous with paler margins [indusia reaching 2/3 to 3/4 around
the receptacles, rarely discoid, midveins strongly protruding, usually 1 cm or less between them] ..............................
....................................................................................................................................................... Cyathea choricarpa
12 Pinnae entire or incised less than 1/3 (mostly 1/5) towards the costa ........................................................................ 13
– Pinnae pinnatifid to pinnatisect, incised at least 1/3 towards the costa, the largest pinnae sometimes basally fully
pinnate......................................................................................................................................................................... 27
13 Laminar apices gradually reduced, not pinna-like (somewhat transient if lateral pinnae small and entire), pinnae
shorter than 10 cm [veins free, only occasionally with single areoles] ...................................................................... 14
– Laminar apices abruptly reduced, pinna-like or with basal lobes; largest pinnae more than 10 cm long. [Veins free or
forming areoles] .......................................................................................................................................................... 16
14 Pinna bases truncate to cordate, apices ± acute; petiole scales concolorous castaneous or with indistinct paler mar-
gins ........................................................................................................................................................ Cyathea falcata
– Pinna bases round to cuneate, apices obtuse to round; petiole scales almost concolorous white to stramineous ...... 15
15 Sori with small appresssed hemitelioid indusia................................................................................. Cyathea haughtii
– Sori exindusiate...................................................................................................................................... Cyathea brucei
16 Vein anastomosing, regularly forming areoles ........................................................................................................... 17
– Veins free, only occasionally with single areoles, anastomosing veins thinner than free veins................................. 22
17 Largest pinnae elliptic, 25–45 × 6–8 cm, ± entire with subentire to serrulate margins; indusia cyatheoid or hemite-
lioid and reaching 3/4 or more around the receptacles ............................................................................................... 18
– Largest pinnae 2.2–6.5 cm broad, linear to long triangular and incised to 1/3 towards the costae, if pinnae elliptic to
oblanceolate then only to 25 cm long ......................................................................................................................... 19
18 Indusia hemitelioid; midveins less than 1 cm apart; largest pinnae 6–7 cm wide, with truncate bases, margins serru-
late at least at tips .......................................................................................................................... Cyathea karsteniana
– Indusia cyatheoid; midveins more than 1 cm apart; largest pinnae (6.5–)7.5–8.0 cm wide, with cuneate bases, mar-
gins crenate at least at tips....................................................................................................................Cyathea nervosa
19 Indusia discoid; rhachises abaxially hairy ........................................................................................ Cyathea stolzeana
– Indusia hemitelioid, reaching to 1/2 around the receptacles; rhachises glabrous abaxially ...................................... 20
20 Pinnae 15–24 pairs per frond, 2.2–4.5 cm wide, incised commonly 1/3–1/5 towards the costae (in distal parts and in
small plants almost entire); indusia hemitelioid, reaching 1/4 to 1/3 around the receptacles; basal veins always
arising from the midvein ................................................................................................................... Cyathea subincisa
– Pinnae 8–12 pairs per frond, 3.0–6.5 cm wide, margins subentire to incised less than 1/5 towards the costae; indusia
reaching 1/2 or more around the receptacles; basal veins sometimes arising from the costae ................................... 21
21 Basal veins arising from the midveins; indusia reaching 1/2 to 2/3 around the receptacles; largest pinnae 20.0–25.0 ×
3.0–6.5 cm, linear-elliptic to oblanceolate.............................................................................................Cyathea ewanii
– Basal veins arising from the costae or from the axils between costae and midribs; indusia reaching about 1/2 around
the receptacles; largest pinnae 29.0–34.0 × 3.4–4.9 cm broad, lanceolate to long triangular ...... Cyathea roraimensis
22 Sori marginal [with large hemitelioid indusia] ...................................................................................Cyathea speciosa
– Sori in zigzag pattern or in several lines parallel to the costae [mainly exindusiate, rarely hemitelioid indusia]...... 23
23 Pinnae 4–9 pairs per frond, petiole scales 5.0–7.0 × 1.5–2.0 mm ......................................................... Cyathea stolzei
– Pinnae 10–16 pairs per frond; petiole scales to 15.0 × 5.0 mm.................................................................................. 24
24 Indusia hemitelioid, reaching 1/2 to 2/3 or more around the recpetacles; veins glabrous except for abundant reddish
brown trichomidia abaxially, evenly distributed on and between them....................................... Cyathea glandulifera
– Indusia absent or developed as ephemeral mucose layer; veins glabrous or with ephemeral white to tan unicellular
trichomidia on and between them, adaxially glabrous or with few scattered, tortuous, hyaline multicellular hairs to 2
mm long ...................................................................................................................................................................... 25
25 Pinnae basally cuneate to round, apically obtuse to round, margins entire .....................................Cyathea cyclodium
– Pinnae basally cuneate, apically acuminate to attenuate, margins crenulate to dentate ............................................. 26
26 Plants with trunk to 1(–4) m tall; pinnae opposite, strongly dimorphic (fertile ones to 2.5 cm wide, sterile ones
4.0–4.5 cm wide).............................................................................................................................. Cyathea williamsii
– Plants without trunk; pinnae mostly alternate, weakly dimorphic (fertile ones weakly narrower than sterile ones,
2.0–2.5 cm wide)................................................................................................................................Cyathea maguirei
27 (12) Pinnae notably stalked, apparently glabrous (few scattered hairs may occur), sori proximal to subproximal... 28

20 • Phytotaxa 61 © 2012 Magnolia Press LEHNERT

– Pinnae mostly sessile to subsessile, glabrous to densely hairy, if stalked to 20 mm then fronds almost glabrous and
sori marginal and indusiate ......................................................................................................................................... 30
28 Petiole scales concolorous auburn to deep brown; laminar indument with almost circular, pseudopeltately attached
squamules; segments subentire to entire with round tips, basal segments never free [sori exindusiate]........................
........................................................................................................................................................ Cyathea latevagans
– Petiole scales concolorous white to stramineous or bicolorous with dark brown central streak; laminar indument
without circular, pseudopeltately attached squamules; segments crenulate with round to acute tips, basal segments
often free ..................................................................................................................................................................... 29
29 Petiole scales to 12 × 1.5 mm; veins abaxially only with flat squamules; sori with hemitelioid indusia ......................
................................................................................................................................................... Cyathea windischiana
– Petiole scales 15–25 × 2.5–4.0 mm; veins abaxially with bullate squamules; sori exindusiate............Cyathea werffii
30 Largest pinnae 2(–2.5) cm wide; pluricellular hairs ± 1 mm long at least present adaxially on rhachises and costae,
often whole plant abundantly hairy with hairs to 5 mm long; petiole scales of all colors, sometimes very narrow and
hair-like (Cyathea bipinnatifida and look-alikes) ....................................................................................................... 31
– Largest pinnae more than 2 cm wide (mostly 3–8 cm); hairs absent or only 0.5–0.6 mm long, whole plant apparently
glabrous; petiole scales white to bicolorous dark brown to blackish with white margins, always broad-lanceolate to
ovate, never hair-like................................................................................................................................................... 44
31 Laminar indument with hairs 3–5 mm long, especially along rhachises.................................................................... 32
– Laminar indument consisting of short pubescence, hairs rarely longer than 1 mm ................................................... 34
32 Scales on trunks and basal petioles broadly lanceolate to ovate, never transient with hairs, in most petiole parts con-
colorous stramineous to whitish, at petiole bases and on trunks strongly bicolorous dark castaneous to blackish with
stramineous to whitish margins; rhachises strongly hairy [indusia hemitelioid]............................... Cyathea decorata
– Scales on trunks and basal petioles narrowly triangular with long marginal cilia, intergrading with hairs in distal pet-
iole parts, stramineous to rufescent [indusia absent] .................................................................................................. 33
33 Hairs absent between veins abaxially. (Colombia, Venezuela, Colombia, N Peru) .......................... Cyathea aterrima
– Hairs also present between veins abaxially. (SE Brazil) ............................................................. Cyathea myriotricha
34 Veins forming areoles between midveins; indusia hemitelioid ............................................................. Cyathea bradei
– Veins not forming areoles between midveins; indusia cyatheoid to sphaeropteroid or absent................................... 35
35 Pinnae sometimes basally pinnate (may vary within one plant), sinuses between segments wide and predominantly
acute; trunks/rhizomes without adventitious buds [indusia absent]......................................................Cyathea werffii
– Pinnae basally not pinnate, sinuses between segments narrow or rounded; trunks/rhizomes regularly with adventi-
tious buds, at least in posterior parts [indusia absent, cyatheoid or sphaeropteroid].................................................. 36
36 Scales on petioles pale, white to stramineous, concolorous throughout or transient with strongly bicolorous, white-
margined scales on the trunk; laminae dark green, matte to lustrous [indusia absent, hemitelioid or cyatheoid]...... 37
– Scales concolorous orange to dark brown, or strongly bicolorous with margins not white; laminae adaxially either
pale green or not lustrous [indusia absent or sphaeropteroid] .................................................................................... 40
37 Scales on trunks and basal petioles strongly bicolorous, dark castaneous to blackish with stramineous to whitish
margins, in most petiole parts concolorous stramineous to whitish; costae adaxially densely hairy with tan to brown
multicellular hairs to 1.0 mm long; indusia absent ....................................................................Cyathea phalaenolepis
– Scales on trunks and basal petioles like in most petiole parts concolorous stramineous to whitish or rarely pale
brown; either costae glabrous or hemitelioid indusia present..................................................................................... 38
38 Sori submarginal, indusia lacking, laminae coriaceous. (Guayana Highlands).................................. Cyathea demissa
– Sori medial to subproximal, indusia cyatheoid, laminae firm-herbaceous. (N Andes) .............................................. 39
39 Laminae adaxially glabrous except for few hairs on costae, indusial margins short-dentate.............. Cyathea alstonii
– Laminae adaxially short-pubescent on and between veins, indusial margins ciliate...................... Cyathea peladensis
40 Laminar hairs also present between veins; pinna pairs 22–36 per frond [indusia absent] ......................................... 41
– Laminar hairs absent between veins (may be present on veins); pinna pairs 10–18(–24) per frond [indusia absent or
sphaeropteroid]............................................................................................................................................................ 42
41 Petiole scales concolorous or if bicolorous then with only weakly darker center, restricted to petiole bases; petioles
glabrous or glabescent with short hairs; plants with erect trunks ............................................... Cyathea bipinnatifida
– Most petiole scales with dark brown to blackish center, persistent and extending to rhachises; petioles persistently
long hairy; plants without trunks or trunks short and decumbent............................................Cyathea phegopteroides
42 Petioles scurfy, only distally hairy near transistion to rhachis, veins lacking hairs on both sides, or with inconspciu-
ous unicellular trichomidia instead [indusia absent]..................................................................Cyathea thelypteroides
– Petioles hairy, not scurfy, veins hairy on both sides ................................................................................................... 43
43 Largest pinnae 2.5–13.0 × 1.0–2.5 cm, paraphyses relatively long (0.4–0.5 mm) [indusia absent] ..............................
..........................................................................................................................................................Cyathea palaciosii
– Largest pinnae to 3.2 × 1.2 cm, paraphyses relatively short (0.2–0.3 mm) [indusia sphaeropteroid]............................
......................................................................................................................................................... Cyathea concordia

A SYNOPSIS OF THE SPECIES OF CYATHEA Phytotaxa 61 © 2012 Magnolia Press • 21

44 (30) Veins free, only occasionally with costular areoles ............................................................................................ 45
– Vein anastomosing, regularly forming areoles ........................................................................................................... 51
45 Free pinnae 6 pairs or less, pinnatifid apical section comprising ± half of the lamina.................... Cyathea singularis
– Free pinnae in 10–20 pairs, apical section comprising a small fraction of the lamina............................................... 46
46 Lobes (= segments) acute to attenuate, sinuses acute to round .................................................................................. 47
– Lobes (= segments) obtuse to round or truncate, sinuses acute.................................................................................. 49
47 Petiole scales concolorous white or with a narrow brown central stripe.......................................Cyathea cnemidaria
– Petiole scales bicolorous dark brown to black with white margins, the latter often abraded ..................................... 48
48 Sori 1.2(–1.5) mm diameter, medial to supramedial, in a weakly to strongly undulate line parallel to the margins;
largest pinnae to 50.0(–75.0) × 5.0–13.0 cm; indusia reaching 1/2 around the receptacles ................. Cyathea mutica
– Sori 0.6–0.8 mm diameter, submarginal, parallel to the margins; pinnae to 33.0 × 3.0–5.0 cm; indusia reaching about
1/3 around the receptacles.................................................................................................................... Cyathea aristata
49 Pinnae stalked to 2 cm; sori marginal to submarginal [apices gradually reduced, only known from cultivation].........
................................................................................................................................................................ Cyathea bella
– Pinnae sessile; sori ± medial, forming delta-pattern................................................................................................... 50
50 Basal veins arising from costae or from the axils with the midveins, usually more than 1 cm between midveins
(range 0.9–1.4 cm); laminar apices gradually to abruptly reduced....................................................... Cyathea uleana
–. Basal veins arising from midveins; less than 1 cm between midveins; laminar apices gradually reduced....................
..........................................................................................................................................................Cyathea chiricana
51 Basal veins arising from the costae or from the axils between costae and midveins; apices abruptly reduced ......... 52
– Basal veins arising from the midveins (rarely few also from the axils) ..................................................................... 53
52 Sori submarginal, parallel to the margins, or distally closer to the midveins than to the margins .................................
...................................................................................................................................................... Cyathea roraimensis
– Sori proximally/below sinuses close to costae, distally medial to inframedial, parallel to the midveins, on the back of
veins ............................................................................................................................................... Cyathea chocoensis
53 Lobes (= segments) acute to attenuate, sinuses acute to round; laminar apices gradually reduced ........................... 54
– Lobes (= segments) obtuse to round or truncate, sinuses acute; laminar apices gradually to abruptly reduced ........ 56
54 Pinnae incised 3/4 or more towards the costa, to 75.0 × 7.5–18.0(–21.0) cm, 15–20 pairs per frond ...........................
............................................................................................................................................................. Cyathea horrida
– Pinnae incised ± 1/2 towards the costa, to 50 × 6 cm, 20–35 pairs per frond ............................................................ 55
55 Pinnae ca. 20–30 pairs per frond, 23.0–39.0 × 3.7–6.0 cm, distal and proximal ones alternate, medial ones suboppo-
site to opposite ................................................................................................................................ Cyathea spectabilis
– Pinnae 30–35 pairs per frond, to 40–50 × 4.5 cm, strictly opposite ................................................... Cyathea cruciata
56 Petiole scales concolorous to discordantly bicolorous, mainly white with dark brown to castaneous central streaks...
................................................................................................................................................................................... 57
– Petiole scales bicolorous, shiny dark brown to castaneous with often abraded, wide white margins ........................ 58
57 Pinnae 12–14 pairs per frond, 35.0–40.0 × 5.0–6.0 cm, opposite; veins forming costular and lateral areoles, more
than 1 cm between the midveins (1.2–1.5 cm); laminar apices abruptly reduced ...........................Cyathea consimilis
– Pinnae 16–22 pairs per frond, 16.0–23.0 × 3.5–4.5 cm, distal and proximal ones alternate, medial ones subopposite
to opposite; veins forming costular areoles only, less than 1 cm between the midveins (0.6–0.8 cm); laminar apices
gradually reduced............................................................................................................................... Cyathea amabilis
58 Indusia discoid, reaching all around the receptacles..........................................................................Cyathea cocleana
– Indusia reaching 1/2 or less around the receptacles.................................................................................................... 59
59 Veins adaxially with white uniseriate hairs to 1.0 mm long, abaxially with appressed trichomidia and weak brown
scurf containing small flat, tan to brown squamules to 0.5 mm long; laminar apices gradually reduced to pinnatifid
apical sections ............................................................................................................................... Cyathea suprapilosa
– Veins glabrous on both sides, or abaxially with weak whitish scurf; laminar apices abruptly reduced to subconform,
rarely conform (pinna-like) apical sections ................................................................................................................ 60
60 Pinnae incised 3/4 or more towards the costae .............................................................................. Cyathea grandifolia
– Pinnae incised 1/2 or less towards the costae ............................................................................................................. 61
61 Largest pinnae 25–50 × 4.5–6.5 cm, 24–30 pairs per frond; veins forming lateral areoles, fusing below sinuses to
cartilaginous areas; indusia reaching ± 1/2 around recetacles ........................................................... Cyathea andicola
– Largest pinnae 20–35 × 2.2–4.5 cm, 15–24 pairs per frond; veins rarely forming lateral areoles, connivent to sinuses
but not fusing to cartilaginous areas; indusia reaching 1/3 to 1/4 around recpetacles...................... Cyathea subincisa

22 • Phytotaxa 61 © 2012 Magnolia Press LEHNERT

Systematic treatment

Cyathea amabilis (C.V.Morton) Lehnert, comb. nov.

Basionym: Hemitelia amabilis Morton (1951: 10). Cnemidaria amabilis (C.V.Morton) Tryon (1970: 52).
Type:—VENEZUELA. Monagas: NE of Caripe, 13 April 1945, Steyermark 62042 (holotype US-photo F,-
photo GH, isotypes F, VEN n.v.).
Distribution and habitat:—Venezuela at 400–1400 m. Cloud forests, thriving on ridges and in disturbed
sites.

Cyathea andicola Domin (1930: 91), nom. nov. for Hemitelia quitensis Domin (1929c: 215), not Cyathea
quitensis (Christensen 1905: 47) Domin (1929b: 263). Cnemidaria quitensis (Domin) Tryon (1970: 52).
Type:—ECUADOR. Pichincha: “In Andibus Quitensis,” 1875, Sodiro s.n. (holotype K).
Distribution and habitat:—Colombia and Ecuador at 100–1100(–1450) m. In wet forests on the western
flanks of the Andes.

Cyathea aristata Domin (1930: 93), nom. nov. for Hemitelia apiculata Hooker (Hooker & Baker 1865: 29), not
Cyathea apiculata (Rosenstock 1914: 213) Domin (1929b: 262). Cnemidaria apiculata (Hook.) Stolze (1974:
40). Type:—MEXICO. Oaxaca: Sierra San Pedro Nolasco, Talea, 1843–1844, Jürgensen 873 (holotype K-
fragment NY, -fragment US, isotype P-photo F).
Distribution and habitat:—Mexico at 1100–1700 m.

Cyathea assurgens Tryon (1986: 43). Type:—ECUADOR. Carchi: San Marcos de los Coaiqueres, on trail
Chical-Tobar Donoso, 1000 m, Øllgaard, Illum, Korning & Thomsen 57570 (holotype AAU, isotype GH).
Distribution and habitat:—Colombia and Ecuador, Chocó-Esmeraldas region, at 400–1050 m. Climber
in dense understory, perhumid mountain rainforests.

Cyathea aterrima (Hooker [in Hooker & Baker] 1866: 38) Domin (1929b: 262). bipinnate-pinnatifid
species (Lehnert 2011b)
Distribution and habitat:—Colombia, Ecuador, and Peru, at 320–1500(–2000) m. Locally frequent, on
limestone and sandstone.

Cyathea bella (Mett.) Domin (1929b: 263). Hemitelia bella Mettenius (1856: 110). Cnemidaria bella (Mett.)
Tryon (1970: 52). Type:—Cultivated at the Botanical Garden Leipzig (holotype LZ, destroyed, lectotype K,
here designated, isolectotypes MO, P, US)
Distribution and habitat:—Known only from cultivated plants.

Cyathea bipinnatifida (Baker) Domin (1929b: 262). Alsophila bipinnatifida Baker (1874: 456).
Type:—GUYANA. “British Guiana”, without date, Appun 1032 (holotype K-fragment NY, isotype P).
Alsophila pubescens Baker (1868: 449). Trichipteris pubescens (Baker) Tryon (1970: 46). Cyathea
pubens Domin (1929b: 263), nom. nov. for Alsophila pubescens Baker, not Cyathea pubescens Kuhn (1869:
164). Type:—PERU: San Martin: Mt. Guayrapurima, Tarapoto, Aug 1856, Spruce 4712 (lectotype K, chosen
by Barrington (1978: 52), -fragment NY, isolectotypes BM, GH, NY, P).
Distribution and habitat:—Venezuela, Colombia, Ecuador, Peru, Brazil, and Bolivia at
(430–)700–2200(–3450) m. Common in humid mountain forests.

Cyathea bradei (Windisch 1973: 372) Lellinger (1984: 57) bipinnate-pinnatifid species (Lehnert 2011a)
Distribution and habitat:—Colombia, Ecuador, and Peru at 100–200 m, to be expected from western
Brazil. Widespread in western Amazonia in terra firme forests.

A SYNOPSIS OF THE SPECIES OF CYATHEA Phytotaxa 61 © 2012 Magnolia Press • 23

Cyathea brucei Lehnert (2006: 321). Type:—ECUADOR. Zamora-Chinchipe: Campamento Miazi, in forest
and cliffs along Río Nangaritza, ca. 04°18´S, ca. 78°40´W, 900 m, 19 February 1994, van der Werff et al.
13287 (holotype QCNE, isotypes MO, UC).
Distribution and habitat:—Ecuador at 900–1500 m. On limestone cliffs.

Cyathea chiricana (Maxon 1912: 33) Domin (1929b: 263)

Distribution and habitat:—Costa Rica and Panama at 1100–2200 m.

Key to the varieties of Cyathea chiricana

1 Petioles inermous, rhachises with flaccid hairs on both sides, segment tips obtuse to acute .........................................
................................................................................................................................. Cyathea chiricana var. chiricana
- Petioles mostly spiny, rhachises glabrous or with sparse hairs only abaxially, segment tips acute to acuminate..........
................................................................................................................................... Cyathea chiricana var. contigua

Cyathea chiricana (Maxon) Domin var. chiricana. Hemitelia chiricana Maxon. Cnemidaria chiricana
(Maxon) Tryon (1970: 51). Type:—PANAMA. Chiriquí: Between Alto de Las Palmas, top of Cerro de la
Horqueta, 18 March 1911, Maxon 5519 (holotype US).
Distribution and habitat:—Panama at 1100–1750 m in cloud forests.

Cyathea chiricana (Maxon) Domin var. contigua (Maxon) Lehnert, comb. nov.

Basionym: Hemitelia contigua Maxon (1912: 32). Cyathea contigua (Maxon) Domin (1929b: 262).
Cnemidaria chiricana (Maxon) R.M.Tryon var. contigua (Maxon) Rojas (2005: 191). Cnemidaria mutica
(Christ 1909: 233) Tryon (1970: 52) var. contigua (Maxon) Stolze (1974: 37). Type:—COSTA RICA.
Cartago: 5 mi. S of Cartago, 12 May 1906, Maxon 523 (holotype NY, fragment US).
Distribution and habitat:—Costa Rica and Panama at 1700–2200 m in cloud forests.

Cyathea chocoensis (Stolze) Lehnert, comb. nov.

Basionym: Cnemidaria chocoensis Stolze (1974: 59). Type:—COLOMBIA. Chocó: 2 km E of San José del
Palmar, 1550–1650 m, 26 March 1971, Lellinger & de la Sota 763 (holotype US, isotypes BM, COL, CR n.v.,
F, HUA n.v., LP n.v.).
Distribution and habitat:—Colombia at 750–1650 m.

Cyathea choricarpa (Maxon) Domin (1929b: 263). Hemitelia choricarpa Maxon (1912: 40). Cnemidaria
choricarpa (Maxon) Tryon (1970: 51). Type:—COSTA RICA. Buenos Aires, Pittier 4835 (holotype US-
photo F, -photo GH).
Distribution and habitat:—Costa Rica and western Colombia at 30–1700 m.

Cyathea cnemidaria Lehnert, nom. nov.

Replaced synonym: Cnemidaria tryoniana Stolze (1974: 38) not Cyathea tryoniana (Gastony 1973: 118)
Nelson-Sutherland et al. (1996: 69). Type:—COLOMBIA. Antioquia: Between Valdivia and Yarumal, 20
February 1942, Metcalf & Cuatrecasas 30122 (holotype US-fragment, photo F, isotypes GH, MO, UC).
Etymology:—Commemorates the now obsolete genus.

24 • Phytotaxa 61 © 2012 Magnolia Press LEHNERT

 Distribution and habitat:—Colombia at 2200 m.
Remarks:—Only known from the type.

Cyathea cocleana (Stolze) Lehnert, comb. nov.

Basionym: Cnemidaria cocleana Stolze (1974: 65). Type:—PANAMA. Cocle: La Mesa, 5 mi. N of El Valle,
10 November 1965, Tyson et al. 2452 (holotype FSU n.v., isotypes FSU n.v.–photos F, GH).
Distribution and habitat:—Costa Rica and Panama at 550–1000 m.

Cyathea consimilis (Stolze) Lehnert, comb. nov.

Basionym: Cnemidaria consimilis Stolze (1974: 66). Type:—VENEZUELA. Sucre: Cerro de Río Arriba,
Peninsula de Paria, 9 August 1966, Steyermark & Rabe 96260 (holotype GH, isotype F).
Distribution and habitat:—Trinidad & Tobago and Venezuela at 600–1200 m.

Cyathea concordia León & Moran (1996: 511). Type:—PERU. Amazonas: Condorcanqui, Cordillera del
Cóndor, Puesto de Vigilancia Alfonso Ugarte, headwaters of Río Comainas, 03º52’48”S 78º26’00”W,
1950–2050 m, 21 July 1994, Beltrán & Foster 1183 (holotype USM, isotypes F, STU).
Distribution and habitat:—Northern Peru at 1950–2050 m, to be expected from southern Ecuador.

Cyathea cruciata (Desv.) Lehnert, comb. nov.

Basionym: Hemitelia cruciata Desvaux (1827: 320). Cnemidaria cruciata (Desv.) Stolze (1974: 65).
Type:—“Habitat in America,” without date, Desvaux s.n. (holotype P-photos F, GH, UC).
Hemitelia leprieurii Kunze (1844: 26). Cyathea leprieurii (Kunze) Domin (1929b: 263).
Type:—FRENCH GUIANA. Leprieur 200 (holotype P).
Distribution and habitat:—French Guiana.

Cyathea cyclodium (R.M.Tryon) Lellinger (1987: 93). Trichipteris cyclodium Tryon (1972: 446).
Type:—VENEZUELA. Amazonas: Cerro Sipapo (Paráque), 6 December 1948, Maguire & Politi 27541
(holotype NY, isotypes GH, US).
Distribution and habitat:—Venezuela, endemic to Cerro Sipapó, at ca. 1000 m.

Cyathea decorata (Maxon) Tryon (1970: 50). Hemitelia decorata Maxon (1946: 439). Type:—COLOMBIA.
El Valle: along the Rio Yurumangui, 5–50 m, Cuatrecasas 15737 (holotype US, isotype F).
Distribution and habitat:—Western Colombia at 100–250(–2000) m. Perhumid mountain rainforest.

Cyathea decurrentiloba Domin (1930: 110), nom.nov. for Hemitelia decurrens Liebmann (1849: 286), not
Cyathea decurrens (Hooker 1844: 51) Copeland (1929: 356). Hemistegia decurrens (Liebm.) Fournier (1872:
135). Cnemidaria decurrens (Liebm.) Tryon (1970: 52). Type:—MEXICO. Oaxaca: Dist. Chinantlá, near
Lobani, 1842, Liebmann 912 (Pl. Mex. 2089) (holotype C n.v.-photo UC,-fragment US).
Hemitelia mexicana Liebmann (1849: 287). Hemistegia mexicana (Liebm.) Fournier (1872: 135).
Type:—MEXICO. Oaxaca: Dist. Chinantlá, near Lacoba, 1842, Liebmann 909, 910, 911 (Pl. Mex. 2105)
(holotype, C n.v.-fragment US).
Hemistegia lucida Fée (1850–1852: 351). Hemitelia lucida (Fée) Maxon (1912: 39). Cyathea lucida (Fée)
Domin (1929b: 264). Type:—MEXICO. Oaxaca: 1842, Galeotti 6537 (holotype BR, isotypes BR, P, S, US).

A SYNOPSIS OF THE SPECIES OF CYATHEA Phytotaxa 61 © 2012 Magnolia Press • 25

 Hemistegia elegantissima Fée (1857: 110). Cyathea elegantissima (Fée) Domin (1930: 11).
Type:—MEXICO. Without date, Linden s.n. (holotype not located); from description (fide Stolze 1974).
Hemitelia guatemalensis Maxon (1912: 40). Cyathea guatemalensis (Maxon) Domin (1929b: 264).
Type:—GUATEMALA. Alta Verapaz: 1800 m, without date, Salvin s.n. (holotype US, isotypes GH n.v., K, P, US).
Cyathea liebmanii Domin (1929b: 264), nom. nov. for Hemitelia mexicana Liebmann (1849: 287), not
Cyathea mexicana Schlechtendal & Chamisso (1830: 616).
Distribution and habitat:—Mexico, Guatemala and Honduras at 100–1350 m.

Cyathea demissa (C.V.Morton) A.R.Sm. ex Lellinger (1987[1988]: 94). Alsophila demissa Morton (1951: 7).
Trichipteris demissa (C.V.Morton) Tryon (1970: 45). Type:—VENEZUELA. Amazonas: Summit Cerro
Duida, 1650–1800 m, 26 November 1928, Tate 402 (holotype US, isotype NY).
Distribution and habitat:—Venezuela at 1540–1830 m, endemic to table mountains.
Remarks:—Barrington (1978) distinguished two varieties, var. demissa and var. thysanolepis, which are
here treated as separate species. Cyathea thysanolepis (Barrington 1976: 1) Smith (1990: 250), which occurs
on the tablemountains further south, has darker petioles and frond axes, and more strongly dissected triangular
laminae.

Cyathea ewanii Alston (1958: 231). Cnemidaria ewanii (Alston) Tryon (1970: 52). Type:—COLOMBIA.
Putumayo: Between Mocoa and Urcusique, 900 m, 9 January 1945, Ewan 16729 (holotype BM-photo F,
isotypes GH, NO n.v., US).
Distribution and habitat:—Colombia, Ecuador and Peru at 100–1000 m.

Cyathea falcata (Kuhn) Domin (1929b: 262). Alsophila falcata Mett. ex Kuhn (1869: 155). Trichipteris
falcata (Kuhn) Barrington (1976: 3). Type:—COLOMBIA. “Panama”, Gorgona Island, November 1847,
Seeman s.n. (holotype B, isotypes K-fragment NY, US).
Nephrodium kuhnii Hieronymus (1905: 440). Dryopteris kuhnii (Hieron.) Christensen (1905: 273).
Alsophila kuhnii (Hieron.) Christensen (1911: 213). Cyathea kuhnii (Hieron.) Domin (1929b: 262).
Trichipteris kuhnii (Hieron.) Tryon (1970: 45). Type:—COLOMBIA. Nariño: Cordillera de Pasto, 1000 m, 10
July 1880, Lehmann 16 (holotype K, isotypes B, US).
Distribution and habitat:—Western Colombia at 200–1000 m.

Cyathea glandulifera Lehnert, nom. nov.

Replaced synonym: Cnemidaria glandulosa Stolze (1984: 101). Type:—PANAMA. Chiriquí: N of San Felix
at Chiriquí-Bocas del Toro border, on Cerro Colorado copper mine road along continental divide, 5000–5500
ft, Mori & Kallunki 5908 (holotype US, isotype MO).
Etymology:—Conserved meaning: “gland-bearing”.
Distribution and habitat:—Panama at ca. 1500–1700 m.

Cyathea grandifolia Willdenow (1810: 490)

Distribution and habitat:—Lesser Antilles, Trinidad & Tobago, Venezuela at 530–1410 m.

Key to the varieties of Cyathea grandifolia

1 Segment tips acute to acuminate, small scales on the abaxial side of costae and midveins white to bicolorous. (St.
Lucia northward to Saba)....................................................................................Cyathea grandifolia var. grandifolia
- Segment tips mostly obtuse, small scales on the abaxial side of costae and midveins brown. (St. Vincent southward
to Venezuela).............................................................................................................. Cyathea grandifolia var. Munita

26 • Phytotaxa 61 © 2012 Magnolia Press LEHNERT

Cyathea grandifolia Willd. var. grandifolia. Hemitelia grandifolia (Willd.) Sprengel (1827: 125).
Cnemidaria grandifolia (Willd.) Proctor (1961: 31). Hemistegia grandifolia (Willd.) Presl (1847: 47).
Microstegnus grandifolius (Willd.) Presl (1847: 46). Type:—“Habitat in America Calidiore,” Herb.
Willdenow No. 20167 (holotype B-photos F, GH, -fragment US).
Hemitelia obtusa Kaulfuss (1824: 252), nom. nud. Cnemidaria obtusa (Kaulf.) Presl (1836: 57).
Cnemidaria grandifolia Willd. var. obtusa (Kaulf.) Stolze (1974: 47). Type:—“Habitat in Antillis“ (not
located).
Cnemidaria kohautiana Presl (1836: 57). Hemitelia kohautiana (C.Presl) Kunze (1844: 298. Hemistegia
kohautiana (C.Presl) Presl (1847: 47). Cyathea kohautiana (C.Presl) Domin (1929b: 264). Hemitelia obtusa
var. kohautiana (C.Presl) Domin (1929a: 71). Cyathea obtusa var. kohautiana (C.Presl) Domin (1930: 141).
Type:—MARTINIQUE. Without date, Sieber 375 [Flora Martin.] (lectotype PRC, designated by
Christenhusz 2009b: 236, isolectotypes B, K, MO, P, US fragment).
Hemitelia serrata Smith (1842: 662), nom. nud.
Hemitelia imrayana Hooker (1844: 669). Hemitelia horrida var. imrayana (Hook.) Hooker (1865: 28).
Type:—DOMINICA. 1839, Imray 14 (holotype not located, isotype P).
Hemistegia willdenowii Fée (1850–1852: 351), for Cyathea grandifolia Willd.
Hemistegia insignis Fée (1866: 99). Hemitelia insignis (Fée) Christensen (1905: 349).
Type:—GUADELOUPE. 1861, l’Herminier s.n. (lectotype RB, designated by Christenhusz 2009b: 236,
isolectotype P).
Cyathea antillana Domin (1930: 91), nom. nov. for Hemitelia imrayana Hook., not Cyathea imrayana
Hooker (1844: 18).
Distribution and habitat:—Saba, St. Kitts & Nevis, Montserrat, Guadeloupe, Dominica, Martinique,
and St. Lucia at 300–1100 m.

Cyathea grandifolia Willd. var. munita (Kuhn) Lehnert, comb. et stat. nov.

Basionym: Cyathea munita Kaulfuss (1824: 260), nom. nud. Cnemidaria munita Presl (1836: 57), nom. nud.
Hemitelia munita Hooker (1844: 32), nom. nud. Hemistegia munita Presl (1847: 47), nom. nud. Hemitelia
munita Kuhn (1869: 162). Type:—„America,“ without date, Palisot de Beauvois s.n., Herb. Willdenow No.
20168 (holotype B-fragment US).
Hemitelia bullata Christ (1897: 81). Hemitelia obtusa var. bullata (H.Christ) Domin (1929a: 71). Cyathea
obtusa var. bullata (H.Christ) Domin (1930: 142). Type:—GRENADA. “In collibus prope Monte Felix,“
1400 ft, 28 November 1889, Eggers 6035 (holotype B-fragment NY [Herbarium Krug & Urban], isotypes F,
P, US)
Distribution and habitat:—St. Vincent, Grenada, Trinidad & Tobago, and Venezuela at 300–1100 m.
Remarks:— This variety represents Cnemidaria grandifolia var. obtusa sensu Stolze (1974). Kaulfuss
(1824) did not cite any specimens for his Hemitelia obtusa. Stolze (1974) referred to Krug (1884–1898), who
cited Ryan s.n. (LZ, destroyed) and Sieber fl. mixta 331 (LZ, destroyed) as specimens annotated with this
name in the Kaulfuss herbarium. Although Stolze did not see any authentic specimens of these collections, he
considered them as possible syntypes. He did not realize that both specimens lie outside the proposed range of
his Cnemidaria grandifolia var obtusa: Ryan s.n. is from Montserrat, and Sieber fl. mixta 331 from Martinique
(M. Christenhusz, pers. comm.). Furthermore, the Sieber collection is a published syntype of Hemitelia
kohautiana, a synonym of Cyathea grandifolia var. grandifolia. Therefore I think it is more justified to treat
Hemitelia obtusa as a nomen nudum in the synonym of the latter variety.
The next available basionym is Hemitelia munita Kuhn. The type in the Willdenow herbarium consists of
three fertile pinna pairs and fits Stolze’s description of this variety well. However, the sheet at B bears the
annotation “Pallisot – Beauvois”, very likely referring to Baron Palisot de Beauvois, a French naturalist
(1752–1820) who made extensive collections of plants and insects in West Africa, Haiti and the United States

A SYNOPSIS OF THE SPECIES OF CYATHEA Phytotaxa 61 © 2012 Magnolia Press • 27

but of which most were lost in fires and ship wrecks before they could arrive in Europe (Merrill 1936). It is
not confirmed that he collected on the Lesser Antilles; he may have passed them on his way from Guinea to
Haiti but he was stricken with yellow fever during this journey. Also, the ship would probably have stopped
for new provisions at one of the French colonies, like Guadeloupe or Martinique, i.e. in the range of var.
grandifolia. The only confirmed collecting activity by Palisot de Beauvois in the Neotropics is on Haiti, but
Cyathea grandifolia does not occur here or the rest of Hispaniola; it is replaced here by C. horrida (Linné
1753: 1092) Smith (1793: 416). The few samples collected by Palisot de Beauvois that are preserved are also
sparsely annotated and notoriously mislabelled, e.g. the provenances “Africa” and “America” are often
confused.
Consequently I choose Hemitelia munita only reluctantly as new varietal basionym, given the possibility
that the type specimen in the Willdenow herbarium has been erroneously attributed to Palisot de Beauvois.

Cyathea haughtii (Maxon) Tryon (1976: 48). Alsophila haughtii Maxon (1944: 46). Type:—COLOMBIA.
Santander: Cerro Armas, 1300–1500 m, 26 July 1936, Haught 1957 (holotype US, isotype US).
Distribution and habitat:—Colombia at 1300–1500 m.

Cyathea horrida (L.) Smith (1793: 416). Polypodium horridum Linné (1753: 1092). Hemitelia horrida (L.)
Brown (1810: 158). Cnemidaria horrida (L.) Presl (1836: 57). Actinophlebia horrida (L.) Presl (1847: 48).
Hemistegia horrida (L.) Fée (1850-1852: 351). Cormophyllum horridum (L.) Newman (1854: 238). Type:—
plate in Plumier (1693) Description des plantes de l'Amérique avec leurs figures 3: t. 4, (lectotype, designated
by Proctor 1985: 157; authentic specimen (epitype) LINN-HL1251-59)
Cyathea commutata Sprengel (1804: 146). Hemitelia commutata (Spreng.) Schlechtendal (1856: 474).
Type:—[DOMINICAN REPUBLIC?]. “Santo Domingo,“ without date, anonymous s.n. (holotype LZ,
destroyed).
Hemitelia hookeri Presl (1848: 350), nom. nud.
Hemistegia repanda Fée (1850–1852: 351). Type:—BRAZIL or CUBA. Without date, Linden s.n.
(holotype RB).
Hemitelia acuminata Schlechtendal (1856: 474), nom. nud.
Hemitelia hookeriana Schlechtendal (1856: 474), nom. nud.
Distribution and habitat:—Cuba, Haiti, Dominican Republic, Puerto Rico, Jamaica, Costa Rica,
Panama, Colombia, Venezuela, Ecuador, and Peru at 100–1350(–1500) m. Lower tropical mountain
rainforests and foothills.
Remarks:—An occurrence in Brazil is unconfirmed except for the ambiguously labelled type of
Hemistegia repanda.

Cyathea karsteniana (Klotzsch) Domin (1929b: 264). Hemitelia karsteniana Klotzsch (1852: 42).
Cnemidaria karsteniana (Klotzsch) Tryon (1970: 52). Type:—VENEZUELA. [Cultivated in the private
garden of Privy Councillor Decker, chief court compositor in Berlin] Karsten s.n. (holotype „Columbien,
Decker, 31 December 1851, Karsten 142“ B, isotype „Culta in horto Deckeriano Berolini,“ 18 September
1850, PRC [Herb. Presl]).
Distribution and habitat:—Venezuela and Ecuador at 950–1550 m. Moist lower mountain forests.

Cyathea latevagans (Baker) Domin (1929b: 262). Alsophila latevagans Baker (1881: 203). Trichipteris
latevagans (Baker) Tryon (1970: 45). Type:—COLOMBIA: Antioquia, 6700 ft, 15 January 1880, Kalbreyer
1327 (holotype K, isotype B).
Distribution and habitat:—Colombia, Ecuador and Peru at 1850–3150 m. Along road cuts, rock
escarpments and in low ridge top vegetation.

28 • Phytotaxa 61 © 2012 Magnolia Press LEHNERT

Cyathea liesneri Smith (1990: 250) (Lehnert in preparation).
Distribution and habitat:—Venezuela at 1550 m on table mountains. Terrestrial in dwarfed forests.
Remarks:—Only known from the type.

Cyathea maguirei (R.M.Tryon) Lehnert, comb. nov.

Basionym: Trichipteris maguirei Tryon (1972: 447). Type:—VENEZUELA. Amazonas: Cerro Sipapo
(Paráque), 6 December 1948, Maguire & Politi 27752 (holotype NY, isotypes GH, US).
Distribution and habitat:—Venezuela at 1050–1580 m on table mountains.

Cyathea minuta Murillo & Murillo (2003: 47), nom. nov. for Cyathea parva (Maxon) Tryon (1976: 47), not
Cyathea parva Copeland (1942: 219). Alsophila parva Maxon (1944: 48). Type:—COLOMBIA. El Valle:
Near Córdoba, Dagua Valley, 80–100 m, 6–8 May 1922, Killipp 5254 (holotype US, isotypes GH, NY, PH
n.v.).
Distribution and habitat:—Colombia at 80–100 m.

Cyathea mutica (Christ 1909: 233) Domin (1929b: 264)

Distribution and habitat:—Costa Rica and Panama.

Key to the varieties of Cyathea mutica:

1 Basal segments of medial pinnae free or only connected by a thin strand of green tissue, largest segments of medial
pinnae strongly crenate to weakly lobed with pinnate venation .......................................Cyathea mutica var. grandis
- All segments broadly adnate and contiguous, segments with entire margins and veins forked two to four times ........
.......................................................................................................................................... Cyathea mutica var. mutica

Cyathea mutica (H.Christ) Domin var. mutica. Hemitelia mutica Christ (1909: 233). Cnemidaria mutica
(H.Christ) Tryon (1970: 52). Type:—COSTA RICA. Turrialba, 850 m, Wercklé s.n. (lectotype P, chosen by
Stolze 1974: 34, isolectotype US).
Hemitelia arachnoidea Underw. ex Maxon (1912: 34). Type:—COSTA RICA. Vicinity of La Palma,
1450–1550 m, 6–8 May 1906, Maxon 453 (holotype US, isotype NY).
Hemitelia horrida (L.) R.Br. var. heterosora Rosenstock (1912: 275). Type:—COSTA RICA. San José:
La Palma, “Iuan Viñas prope Río Chis,” 20 March 1910, Brade 451 (holotype not located-fragment US).
Hemitelia pittieri Maxon (1912: 32). Cyathea pittieri (Maxon) Domin (1929b: 264). Type:—COSTA
RICA. Cañas Gordas, Valle de Agua Buena, February 1897, Pittier 10969 pro parte (holotype US-fragment F).
Hemitelia subglabra Maxon (1912b: 36). Cyathea subglabra (Maxon) Domin (1929b: 264). Cnemidaria
subglabra (Maxon) Tryon (1970: 52). Type:—COSTA RICA. Vicinity of La Palma, 6–8 May 1906, Maxon
451 (holotype US).
Cyathea subarachnoidea Domin (1929: 264), nom. nov. for Hemitelia arachnoidea Underw. ex Maxon
(1912b: 34), not Cyathea arachnoidea Hooker (1865: 24).
Distribution and habitat:—Costa Rica and Panama at 1100–1830(–2200) m.

Cyathea mutica (Christ) Domin var. grandis (Maxon) Lehnert, comb. nov.

Basionym: Hemitelia grandis Maxon (1912: 37). Cnemidaria mutica (Christ) R.M.Tryon var. grandis
(Maxon) Stolze (1974: 36). Cyathea grandis (Maxon) Domin (1929b: 264). Type:—COSTA RICA. Vicinity
of Coliblanco, 30 April–02 May 1906, Maxon 307 (holotype US, isotype US).

A SYNOPSIS OF THE SPECIES OF CYATHEA Phytotaxa 61 © 2012 Magnolia Press • 29

 Hemitelia rudis Maxon (1914: 413). Cyathea rudis (Maxon) Domin (1929b: 264). Cnemidaria rudis
(Maxon) Tryon (1970: 51). Type:—PANAMA. Chiriquí: Holcomb’s Trail, above El Boquete, Maxon 5682
(holotype US, isotype GH n.v.).
Distribution and habitat:—Costa Rica and Panama at 1100–1600 m.

Cyathea myriotricha (Baker 1885: 217) R.C.Moran & J.Prado (Moran et al. 2008: 365) bipinnate-
pinnatifid species (Lehnert in preparation).
Distribution and habitat:—Southeastern Brazil at 700–1200 m. On wet rocks.

Cyathea nanna (Barrington) Lellinger (1987[1988]: 94). Trichipteris nanna Barrington (1976: 3).
Type:—GUAYANA: „British Guyana,“ Upper Mazarumi River Basin, Mt. Ayanganna, on shoulder of E
flank, about Thompson Camp, 1418 m, 12 August 1960, Tillet, Tillet & Boyan 45119 (holotype US, isotype
NY).
Distribution and habitat:—Guayana at 1400–1545 m on the flanks of Mt. Ayanganna. In low, dense
forests.

Cyathea neblinae Smith (1990: 250) bipinnate species (Lehnert in preparation).

Distribution and habitat:—Venezuela and Brazil at 140–1300 m in the Neblina range. In mountain
forests.
Remarks:—The new record from Brazil (Caravalho et al. 320, INPA) is a plant with larger, fully
bipinnate-pinnatisect fronds with long-stalked pinnules.

Cyathea nervosa (Maxon) Lehnert (2011b: 43). Hemitelia nervosa Maxon (1944: 309). Cnemidaria nervosa
(Maxon) Tryon (1970: 52). Type:—PERU. Loreto: Mouth of Río Santiago, above Pongo de Manseriche, 300
m, 18 December 1931, Mexia 6291 (holotype US, isotypes F, GH, NY, UC).
Distribution and habitat:—Ecuador and Peru at 300–500 m. Wet lowland forests.

Cyathea palaciosii Moran (1995: 57). Type:—ECUADOR. Zamora-Chinchipe: Nangaritza Cantón, Miazi,
detrás Campamento Militar, 04º16´S, 78º42´W, 900–1000 m, 21 October 1991, Palacios et al. 8578 (holotype
MO, isotypes AAU, QCNE, UC, US).
Distribution and habitat:—Ecuador and Peru at 1000 m. On rocks in seepage areas.

Cyathea phalaenolepis (C.Chr.) Domin (1929b: 263). Alsophila phalaenolepis Christensen (1911: 213).
Trichipteris phalaenolepis (C.Chr.) Tryon (1970: 46). Type:—ECUADOR. Esmeraldas: Without locality,
August 1904, Sodiro 15737 (holotype P-photo, fragment US).
Distribution and habitat:—Colombia and Ecuador at 5–300 m. In understory of perhumid mountain
forests.

Cyathea peladensis (Hieron.) Domin (1929b: 263). Alsophila peladensis Hieronymus (1906: 233).
Type:—COLOMBIA. Huila: Cerro Pelado (upper Magdalena Valley), Stübel 1259 (holotype B-photo GH).
Distribution and habitat:—Colombia, Ecuador and Peru at 1600–2500 m. In open ridge-top vegetation.

Cyathea phegopteroides (Hook.) Domin (1929b: 263). Alsophila phegopteroides Hooker (1865: 32).
Type:—PERU. San Martin: Tarapoto, Spruce 4020 (holotype K, isotypes B, P, US).
Distribution and habitat:—Peru at 1000–1800 m. In open ridge-top vegetation.

Cyathea pseudonanna (L.D.Gómez) Lellinger (1985: 376). Trichipteris pseudonanna Gómez (1981: 69).
Type:—PANAMA. Veraguas: Cerro Tuti [Tute], 08°28'56"N, 81°05'53"W, 700–1000 m, 25 May 1977,
Folsom & Edwards 3370 (holotype MO).

30 • Phytotaxa 61 © 2012 Magnolia Press LEHNERT

 Distribution and habitat:—Panama at 700–1000 m.

Cyathea roraimensis (Domin) Domin (1930: 154). Hemitelia roraimensis Domin (1929c: 216). Cnemidaria
roraimensis (Domin) Tryon (1970: 52). Type:—GUYANA. “British Guyana”, Roraima, 1863–1864, Appun
1127 (holotype K-photo F).
Distribution and habitat:—Guyana at 600–1500 m.

Cyathea singularis (Stolze) Lehnert, comb. nov.

Basionym: Cnemidaria singularis Stolze (1974: 40). Type:—COLOMBIA. Putumayo: Cerro Portachuelo,
2300 m, 27 August 1965, Soejarto 1571 (holotype GH n.v.-photo F, isotypes, F, GH n.v.).
Distribution and habitat:—Colombia at 2300 m.

Cyathea speciosa Willdenow (1810: 490). Hemitelia speciosa (Willd.) Kaulfuss (1824: 252), not Cnemidaria
speciosa Presl (1836: Figures 16, 17). Type:—VENEZUELA. Monagas: Caripe, Humboldt s.n. [Herb.
Willdenow no 20179] (holotype B, isotype P-fragments ex Herb. Kaulfuss, Herb. Luerrsen).
Hemitelia integrifolia Klotzsch (1844: 539). Cnemidaria integrifolia (Klotzsch) Tryon (1970: 52).
Type:—VENEZUELA. “Caracas,“ Otto 671 (lectotype B, designated by Tryon 1976: 47).
Hemitelia lindenii Hooker (1848: 706). Cnemidaria lindenii (Hook.) Tryon (1970: 52).
Type:—VENEZUELA. “Caracas,” Linden 663 (holotype K).
Hemitelia funckiana Klotzsch, nom. ined., annotated on Funck 565 (BR).
Distribution and habitat:—Colombia and Venezuela at 950–1500 m, on mountains with Caribbean
influence.

Cyathea spectabilis (Kunze) Domin (1929b: 264). Hemitelia spectabilis Kunze (1848: 233). Hemistegia
spectabilis (Kunze) Fée (1850–1852: 351). Cnemidaria spectabilis (Kunze) Tryon (1970: 52).
Type:—SURINAM.”In declivibus montium reg. inter. ad flumen pr. stationem Victoriam,” April 1847,
Kappler 1771 (holotype not located, isotypes K, P-photo F, MPU).
Actinophlebia obtusa Presl (1847: 48), from description, Hooker (1844: pl. 14).
Hemitelia klotzschiana H.Karst. ex Klotzsch (1852: 42). Type:—[Cultivated in the private garden of Privy
Councillor Decker, chief court compositor in Berlin] Authentic material probably “Venezuela, January 1852,
Karsten 143” (B-photo F).
Hemitelia spectabilis Kunze var. trinitensis Domin (1929a: 72). Cyathea spectabilis (Kunze) R.M.Tryon
var. trinitensis (Domin) Domin (1929b: 264). Type:—TRINIDAD & TOBAGO. Trinidad: La Seiva Valley, 16
January 1921, Wakefield s.n. (holotype not located).
Hemitelia spectabilis var. longipinna Domin (1929a: 73). Cyathea spectabilis var. longipinna (Domin)
Domin (1929b: 264). Type:—The same as of Hemitelia spectabilis Kunze.
Cnemidaria spectabilis var. colombiensis Stolze (1974: 63). Type:—COLOMBIA. El Valle [= Valle del
Cauca]: Cordoba, 6–8 May 1922, Killip 5039 (holotype US-photo F, isotypes GH n.v., NY).
Distribution and habitat:—Colombia, Venezuela, Guyana, Suriname and French Guiana at
(75–)130–1300(–2070) m.

Cyathea stolzeana (L.D.Gómez) Lehnert, comb. nov.

Basionym: Cnemidaria stolzeana Gómez (1981: 457). Type:—PANAMA. Coclé: Beyond sawmill in lumber
road above El Cope, 900 m, 20 January 1978, Hammel 1036 (holotype CR, isotype MO).
Distribution and habitat:—Panama at 900 m.

A SYNOPSIS OF THE SPECIES OF CYATHEA Phytotaxa 61 © 2012 Magnolia Press • 31

Cyathea stolzei A.R.Sm. ex Lellinger (1987[1988]: 101), nom. nov. for Trichipteris pinnata Stolze (1984:
103), not Cyathea pinnata Roxburgh (1844: 517). Type:—PANAMA. Colón: Santa Rita Ridge road, 21–26
km from Transisthmian Highway, 500–550 m, Knapp 5881 (holotype MO, isotype F).
Distribution and habitat:—Nicaragua, Costa Rica and Panama at 100–550 m. In upland forests and
seasonally dry, evergreen forest.

Cyathea subincisa (Kunze) Domin (1929b: 264). Hemitelia subincisa Kunze (1844: 296), nom. nov. for
Cnemidaria speciosa C.Presl, not Cyathea speciosa Willd.
Cnemidaria speciosa Presl (1836: Pl. 1, fig. 16, 17). Hemistegia speciosa (C.Presl) Fée (1850–1852: 351).
Type:—PERU. Huánuco: Pampayaco, July 1829, Poeppig 221 [diar. 1144] (holotype PRC-fragment NY, US,
isotypes B, BR n.v., P).
Distribution and habitat:—Peru and Bolivia at 300–1700 m.

Cyathea suprapilosa Lehnert, nom. nov.

Replaced synonym: Cnemidaria suprastrigosa Moran (1990: 246), not Cyathea suprastrigosa (Christ in
Pittier 1901: 44) Maxon (1909: 83). Type:—PANAMA. Veraguas: Cerro Tute, Santa Fé, ridge up from former
Escuela Agricola, 08º35´W, 78º05´W, 1000–1300 m, Hamilton & Dressler 3049 (holotype MO).
Etymology:—Conserved meaning: “hairy above“.
Distribution and habitat:—Panama at 1000–1300 m.

Cyathea thelypteroides Smith (2006: 427). Type:—PERU. Amazonas: Bagua Distr., upper slopes & summit
of Cerro Tayu, 1030 m, 22 March 2001, van der Werff, Vásquez & Gray 16323 (holotype MO, isotype UC).
Distribution and habitat:—Peru at 1030–1200 m. On rocks in open ridge-top vegetation.

Cyathea uleana (Sampaio 1923: 65) Lehnert (2011b: 43)

Distribution and habitat:—Colombia, Ecuador, Peru, Brazil and Bolivia.

Key to the varieties of Cyathea uleana

1 Laminae terminating with a pinnatifid section .........................................................Cyathea uleana var. abitaguensis

– Laminae terminating with a subentire, pinna-like section................................................. Cyathea uleana var. uleana

Cyathea uleana (A.Samp.) Lehnert (2011b: 43) var. uleana. Hemitelia uleana Sampaio (1923: 65).
Cnemidaria uleana (A.Samp.) Tryon (1970: 52). Type:—BRAZIL. Rio de Janeiro: Puerto de Nueva Friburgo,
Alto da Sierra, 1898, Ule s.n. (holotype R n.v.-photo F, GH).
Hemitelia maxonii Rosenstock (1925: 344). Cyathea subarborescens Domin (1930: 162), nom. nov. for
Hemitelia maxonii Rosenst., 1925, not Cyathea maxonii Underw. ex Maxon (1909: 82). Type:—BRAZIL. Sao
Paulo: Serra do Mar, 26 April 1914, Brade 6901 (holotype R n.v., isotype S-PA).
Distribution and habitat:—Ecuador, Peru, Brazil, and Bolivia at (580–)1140–1830(–2000) m. In Mata
Atlântia and wet mountain forests on the eastern Andean flanks.

Cyathea uleana (A.Samp.) Lehnert var. abitaguensis (Domin) Lehnert, comb. nov.

Basionym: Cnemidaria uleana var. abitaguensis (Domin) Stolze (1974: 51). Hemitelia abitaguensis Domin
(1929c: 74). Cyathea abitaguensis (Domin) Domin (1930: 88). Type:—ECUADOR. Pastaza: “Monte
Abitagua, Andibus Ecuadoriensibus,” 1857, Spruce 5364 (holotype K, isotypes GH, NY).

32 • Phytotaxa 61 © 2012 Magnolia Press LEHNERT

 Distribution and habitat:—Colombia and Ecuador at 1450–2100 m. In wet mountain forests on the
eastern Andean flanks.

Cyathea ursina (Maxon) Lellinger (1987[1988]: 101). Alsophila ursina Maxon (1944: 48). Trichipteris
ursina (Maxon) Tryon (1970: 44). Type:—BELIZE: “British Honduras,” Antelope Ridge, Stann Creek Valley,
5 February 1940, Gentle 3197 (holotype US, isotypes MICH n.v., US).
Distribution and habitat:—Belize, Honduras, Nicaragua, Costa Rica and Panama at 4–900 m. In the
understory of forest, preferring swampy sites.

Cyathea varians (R.C.Moran) Lehnert, comb. nov.

Basionym: Cnemidaria varians Moran (1990: 246). Type:—PANAMA. Panamá: Cerro Jefe, 1000 m,
Valdespino & Aranda 139 (holotype MO, isotypes PMA n.v., UC).
Distribution and habitat:—Panama at 1000–1500 m.

Cyathea werffii Moran (1991: 94) bipinnate-pinnatifid species (Lehnert in preparation).

Distribution and habitat:—Ecuador and Peru at 120–1380 m. In ridge-top vegetation, rocky slopes, and
on koalinite soils.

Cyathea williamsii (Maxon) Domin (1930: 171). Alsophila williamsii Maxon (1922: 46). Trichipteris
williamsii (Maxon) Tryon (1970: 44). Type:—PANAMA. Darién: Mountains above Cana, 1800 m, 17
April–8 June 1908, Williams 850 (holotype US, isotype NY).
Distribution and habitat:—Panama at 300–1300(–1500) m.

Cyathea windischiana Smith (2006: 427) bipinnate-pinnatifid species (Lehnert 2011a)

Distribution and habitat:—Peru at 800 m.
Remarks:—Known only from the type.

Acknowledgments

I thank Michael Kessler (Z) and Alan R. Smith (UC) for their scientific guidance. I further thank my
colleagues Dave Conant (LSC), Alexander N. Schmidt-Lebuhn (CANB), Michael Sundue (VT), Florian
Werner (OLD) and Jürgen Homeier (GOET) for their company in the field. I am indebted to the curators of
AAU, B, BM, COL, CUZ, F, GH, GOET, HUT, K, L, LOJA, LPB, MBM, MO, NY, P, Q, QCA, QCNE,
QPLS, S, SCZ, SP, U, UC, USM, UPCB, UPRRP and Z for providing loans or for attending me during my
visits, especially to Asunción Cano (USM), Stefan Beck (LPB) and Alison Paul (BM). Anette Rosenbauer and
Thomas Jossberger (STU) diligently handled loans and mounted new material. I thank Michael Kessler,
Michael Sundue and Maarten Christenhusz for helpful comments on the manuscript.
Field studies that contributed to the presented results were funded by the DAAD (German Academic
Exchange Service) and the DFG (German Research Foundation, grants GR 1588/7, LE 1826/4-1). This
research received support from the SYNTHESYS Project (http://www.synthesys.info/), which is financed by
European Community Research Infrastructure Action under the FP6 “Structuring the European Research
Area” Programme.

A SYNOPSIS OF THE SPECIES OF CYATHEA Phytotaxa 61 © 2012 Magnolia Press • 33

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