The Cyatheaceae (Polypodiopsida) of Peru

MARCUS LEHNERT

Staatliches Museum fr Naturkunde Stuttgart, Abt. Botanik, Am Lwentor, Rosenstein 1, 70191

Stuttgart, Germany; e-mail: lehnert.smns@naturkundemuseum-bw.de; marlehnert@yahoo.com

Abstract. A summary is presented of all Peruvian species of scaly tree ferns (Cyatheaceae).
A total of 83 species is native to Peru (Sphaeropteris 1 sp., Alsophila sect. Alsophila 9
spp., Alsophila sect. Gymnosphaera 1 sp., Cyathea 72 spp.), with 13 endemic species in
Cyathea and one in Alsophila. The following new species are described and illustrated:
Alsophila gastonyi, Cyathea chontilla, C. nephele, C. polliculi, and C. praetermissa.
The genera Cnemidaria and Trichipteris are no longer recognized and merged in Cyathea.
Necessary combinations are made for Cyathea alatissima, C. nervosa, and C. uleana.
Cyathea serpens is presented in detail and its relationship is discussed. An articial key to
the Peruvian species of Cyatheaceae is provided.
Key Words: Alsophila gastonyi, Andes, Peru, Cyathea chontilla, C. nephele,
C. polliculi, C. praetermissa, C. serpens, C. subincisa, scaly tree ferns, endemism.

The montane rainforest of Peru harbors one
of the richest fern oras in the world. Tryon
and Stolze (1989) reported 1060 native
species of ferns, horsetails, and lycophytes
from Peru. Since then the number has grown
by 137 new species and records (Smith et al.,
2005), signifying an increase by 13%. The ca.
1200 native Peruvian species represent a
tenth of the global diversity of the seed-free
vascular plants (Smith et al., 2006).
The scaly tree ferns (Cyatheaceae) are one
of the most conspicuous fern groups in
tropical montane forest because of their size
and frequency. The family shows a pantrop-
ical distribution with more than 600 species
(Kramer & Green, 1990). Tryon and Stolze
(1989) listed 45 species of Cyatheaceae from
Peru but assumed that at least 19 more
species that are known from adjoining coun-
tries may occur here as well. The latest
supplementary account of tree ferns to the
ora of Peru (Smith et al., 2005) is rendered
obsolete because recent taxonomic studies in
the genus Cyathea lead to several new
species circumscriptions, nomenclatural
changes, and range extensions (Lehnert,
2008, 2009a, b). The present work provides
an updated list of all known Peruvian scaly
tree ferns.
The basis for this study was a doctoral
research project at the University of California,
Berkeley, aiming on a revision of all Netropical
Cyatheaceae. Besides using the local herbarium
UC, other herbaria provided loans (F, GH, MO)
or were visited (NY, US) during this period.
Further herbaria consulted prior to and after this
time are AAU, B, BM, COL, COLO, CUZ,
GOET, HUT, K, LIL, LOJA, LPB, MBM, P,
QCA, QCNE, QPLS, S, SCZ, SPA, STU, TUR,
UPCB, USM, and Z. The analysis of the
herbarium specimens was supplemented with
rst hand experience on the habit and ecology
of the plants, which were gained on several
eld trips to the Andes between 2000 and 2004.
Material for this study comprises ca. 1000
collections from Peru, including 59 of the
authors collections. The presented ranges are
based exclusively on material that has been
revised personally or whose determinations
were veried recently by colleagues (Blanca
Len, pers. comm.). Thus some departmentos
may be missing from the distribution compared
to the treatment by Tryon and Stolze (1989).
The generic system used here is based on
the works of Conant et al. (1995, 1996) and
Korall et al. (2006, 2007). Three genera are
formally recognized: Alsophila, Cyathea, and
Sphaeropteris. Alsophila falls into two sub-

Brittonia, 63(1), 2011, pp. 1145 ISSUED: 1 March 2011

2010, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.
12 BRITTONIA [VOL 63

clades, forming an unresolved polytomy with
the monophyletic Cyathea in strict consensus
analyses (Korall et al., 2006, 2007; Janssen et
al., 2008). Both subclades roughly corre-
spond with established taxonomic subgroups
in Alsophila (Holttum, 1963; Holttum &
Edwards, 1983) and are recognized here as
sect. Alsophila (incl. Nephelea; Conant, 1983)
and sect. Gymnosphaera. A monophyletic
genus Cyathea must include the genera Cnemi-
daria and Hymenophyllopsis (Korall et al.,
2006, 2007); the genus Trichipteris (or Trichop-
teris) was previously included in Cyathea based
on morphological characters (Lellinger, 1987).
The necessity for recombinations arises here
only for Cnemidaria and Trichipteris, since
Hymenophyllopsis does not occur in Peru.
Except for the Sphaeropteris horrida group
(Tryon, 1971), previously recognized Neotrop-
ical subgenera and sections in Sphaeropteris
(Tryon, 1970; Windisch, 1977, 1978) were
transferred to Cyathea (Kramer, 1978; Lellinger,
1984), which is corroborated by the results of
Korall et al. (2006, 2007). Terminology follows
Tryon (1970) and Gastony (1973).
Results
In total, 83 species of Cyatheaceae are
known from Peru. Five new species, Also-
phila gastonyi, Cyathea chontilla, C. nephele,
C. polliculi, and C. praetermissa, are
described, and all except for C. polliculi are
endemic to Peru. The inclusion of Cnemida-
ria into Cyathea necessitates new combina-
tions for Cyathea alatissima, C. nervosa, and
C. uleana, and leads to the change of
Cnemidaria speciosa C. Presl into C. sub-
incisa Domin. One previously little known
species, C. serpens, is the last species that
had to be transferred from the obsolete
genus Trichipteris to Cyathea (Lehnert,
2009b) and is illustrated here in detail for
the rst time. Trichipteris ava R. M. Tryon,
which was described from a sterile collection
(Tryon & Stolze, 1989), is synonymized with
C. herzogii Rosenst. Currently 14 species of
Cyatheaceae are regarded as being endemic to
Peru.
The presented articial key does not rely
on indusial characters as much as did pre-
vious keys. Instead, the emphasis is laid on
the petiolar characters, like scales and scurf,
which are more readily available and appa-
rently dene natural groups in Cyathea.
However, fertile material is still needed or at
least very helpful for secure identication on
the species level. Where not essential, the
type of indusium is given as an additional
character in brackets.

Key to the neotropical genera of the Cyatheaceae

1. Cells of the petiole scales of (nearly) the same size and orientation (no distinct margin of smaller cells), but dark
marginal teeth, setae or pale, long cilia may be present, petiole scales basix on truncate bases, never with
indurated bases or with transitions to black spines.
2. Petiole scales with small, unicellular teeth that are the end of one vertical cell row; scales of various sizes but not
intergrading with branched and simple hairs; sori subcostal to costal, with deeply cyatheoid to sphaeropteroid
indusia; fertile plants mostly with stout, erect trunks 210 m tall. . . . . . . . . . . . . . . . . . . . . . .Sphaeropteris
2. Petiole scales without small black teeth but long, often pluricellular cilia, scales may be intergrading with
branched and simple hairs; sori either exindusiate or of marginal position; fertile plants mostly without trunks or
with decumbent trunks to 3 m tall. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyathea
1. Cells of the petiole scale margins smaller and with different orientation than those of the scale bodies, margins may
be very narrow and reduced to one row of small teeth in the distal part of the scales, but are recognizable at the
bases, petiole scales pseudopeltately attached on weakly to strongly cordate bases or basally attached, in the latter
case often intergrading with black spines.
3. Scales with at least one dark apical seta protruding from the scale center (supported by more than one cell row
below); stem and petioles, if spiny, with black squaminate spines (i.e., with sharp color contrast to the cortex,
breaking off cleanly at this point in dried specimens); adventitious pinnae at the petiole bases skeletonized if
present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Alsophila
3. Scales without setae, although sometimes small black teeth or long cellular processes of the margin occur
(supported only by one cell row or less, i.e. several marginal teeth supported by the same cell); stem and petioles
only with corticinate spines (i.e., without color contrast towards the cortex, not breaking off cleanly in dried
specimens); adventitious pinnae near the petiole bases not skeletonized if present.. . . . . . . . . . . . . . Cyathea
2011] LEHNERT: CYATHEACEAE OF PERU 13

Key to the Peruvian species of Alsophila

1. Fronds pinnate-pinnatid; fertile plants without trunks but with stout, often ascending rhizomes. . . A. paucifolia
1. Fronds bipinnate-pinnatid to tripinnate; fertile plants with trunks.
2. Main leaf axes shiny dark brown to atropurpureous and without hairs, petioles not spiny, with highly dissected
basal pinnae (aphlebia); sori exindusiate. Sect. Gymnosphaera. . . . . . . . . . . . . . . . . . . . . . . . . . . A. salvinii
2. Main leaf axes with hairs at least adaxially and not shiny brown, or if shiny then petioles spiny or without highly
dissected basal pinnae (aphlebia); sori indusiate, indusia rarely absent at maturity. Sect. Alsophila
3. Laminar apices gradually reduced; squaminate spines few, restricted to the basal parts of the petioles or
replaced by indurated scales; plants without skeletonized pinnae at petiole bases [indusia cyatheoid to
subsphaeropteroid].
4. Petiole scales brown to pale brown, petioles and leaf axes brown to stramineous; petioles basally mainly
with indurated scales (winged spines); trunks not spiny, with adventitious buds.. . . . . . . . . . A. engelii
4. Petiole scales dark brown, petioles and leaf axes orange-brown to purpurascent; petioles basally with few
spines; trunks spiny, without adventitious buds [if costae abaxially hairy and found below 1000 m=A.
gastonyi]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. minervae
3. Laminar apices abruptly reduced, apical sections often pinna-like (conform); squaminate spines numerous on
petioles, often extending to the basal parts of the rachises, if spines weakly developed then plants with
skeletonized pinnae at petiole bases.
5. Petiole bases with skeletonized pinnae; indusia sphaeropteroid; trunks with old petiole bases, appearing
dark and sulcated, forming small groves of well-separated trunks with short subterranean stolons.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. mostellaria
5. Petiole bases without skeletonized pinnae; indusia meniscoid to sphaeropteroid; trunks without old petiole
bases, without short subterranean stolons but sometimes forming multistemmed plants by means of
adventitious buds.
6. Costae and costules abaxially densely pubescent; petiole scales lanceolate and dark brown, squaminate
spines relatively short and broad, of the same length as scales. . . . . . . . . . . . . . . . . . . A. gastonyi
6. Costae and costules abaxially without hairs or only sparsely hairy; petiole scales short-linear to narrowly
lanceolate, sometimes wooly, stramineous to deep brown, in the latter case always long-linear,
squaminate spines reatively long.
7. Scales of abaxial sides of petiole bases with lateral setae; indusia subsphaeropteroid to sphaeropteroid.
8. Costae distally not alate; indusia pale brown, translucent, fragile; scales on petioles and trunks
relatively short, revealing most of the smaller squamimate spines, lenticels visible on the
trunk. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. imrayana
8. Costae distally alate; indusia dark brown, opaque, rm, splitting into three almost equally sized
parts at maturity; scales on petioles and trunks relatively long, hiding most of the smaller
squamimate spines, lenticels not visible on the trunk. . . . . . . . . . . . . . . . . . . . . . A. cuspidata
7. Scales of the petiole without lateral setae (rarely a second apical seta present); indusia meniscoid to
subsphaeropteroid with irregular dehiscence.
9. Costae distally not alate; stellate hairs and squamellae with many black setae present on the lamina
abaxially; indusia subsphaeropteroid to cyatheoid, usually splitting into two unequally sized parts
at maturity; scales with apical seta to 1 mm long, dark central stripe mostly continuous.A. erinacea
9. Costae distally alate; stellate hairs absent, squamellae present on the lamina abaxially rarely with
one black seta; indusia meniscoid, sometimes abraded to irregular discs; scales with apical seta to
2 mm long, dark central stripe discontinuous or lacking. . . . . . . . . . . . . . . . . . . . . . A. incana

Keys to the Peruvian species of Cyathea

1. Laminae simply pinnnate to pinnate-pinnatisect.
2. Pinnae all sessile, varying from deeply pinnatisect to entire, veins lacking hairs on both sides, costae (as usually
all leaf axes) adaxially without hairs. Cnemidaria-group.
3. Rachises green alate, basal veins of segments always arising from the costae, segments obtuse to round.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. alatissima
3. Rachises not green alate, basal veins of segments always arising from the miveins or from the axil between
midvein and costa, if some basal veins arising directly from the costae then segments long acute and petioles
spiny.
4. Pinnae linear-elliptic to oblanceolate, entire or coarsely crenate (incised 1/5 or less towards the costae).
5. Pinna pairs 812; sori more or less equidistant between margins and costae; indusia hemitelioid, reaching
1/2 to 2/3 around the receptacles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. ewanii
5. Pinna pairs 1214; sori in a zig-zag-pattern along the costa (distally subcostal, proximally medial with
respect to the midveins); indusia cyatheoid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. nervosa
4. Pinnae long triangular to linear, largest ones incised at least 1/3 towards the costae.
14 BRITTONIA [VOL 63

6. Apices gradually reduced to pinnatid sections; petioles spiny; segments long acute to attenuate,
separated by relatively wide, round sinuses. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .C. horrida
6. Apices abruptly reduced to conform or subconform sections; petioles inermous or spasely muricate;
segments obtuse to round, separated by narrow, acute sinuses.
7. Segments incised to 1/3 towards the costae, rarely in large plants the basal segments more deeply
separated; basal veins of segments arising from the midvein, regularly forming costal areoles.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. subincisa
7. Segments incised 2/3 to 3/4 towards the costae, rarely in large plants the basal segments free; basal
veins of segments arising from the axils between midveins and costa, rarely directly from the costae,
only irregularly forming costal areoles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .C. uleana
2. Pinnae either short-pubescent on at least one side of the veins or notably stalked, costae (as usually all leaf axes)
at least sparsely hairy adaxially.
8. Pinnae notably stalked, apparently without hairs (few scattered hairs may occur), leaf axes dark brown to
atropurpureous or black, shiny.
9. 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]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. latevagans
9. 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.
10. Petiole scales to 121.5 mm; veins abaxially only with at squamules; sori with hemitelioid indusia
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. windischiana
10. Petiole scales 15252.54.0 mm; veins abaxially with bullate squamules; sori exindusiate. C. werfi
8. Pinnae sessile, or if short-stalked then veins nely and densely pubescent at least on one side; leaf axes
yellowish to brown, rarely darker, never shiny.
11. Petiole scales narrowly triangular with long marginal cilia, intergrading with hairs; laminar indument
with hairs 35 mm long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. aterrima
11. Petiole scales narrowly to broadly lanceolate, dentate to mbriate but without long marginal cilia
and not intergrading with hairs; laminar indument consisting of short pubescence, rarely hairs
longer than 1 mm.
12. Veins irregularly forming areoles between midveins; indusia hemitelioid. . . . . . . . . . . . C. bradei
12. Veins never forming areoles between midveins; indusia cyatheoid to sphaeropteroid or lacking.
13. Largest pinnae basally pinnate, sinuses between segments wide and predominantly acute;
trunks/rhizomes without adventitious buds [indusia absent]. . . . . . . . . . . . . . . . . . C. werfi
13. Largest pinnae basally only pinnatid to pinnatisect, sinuses between segments narrow or
rounded; trunks/rhizomes regularly with adventitious buds, at least in posterior parts [indusia
absent or sphaeropteroid].
14. Petiole scales pale, white to stramineous, concolorous or nearly so; laminae adaxially short-
pubescent and lustrous dark green [indusia cyatheoid]. . . . . . . . . . . . . . . . .C. peladensis
14. Petiole scales orange to dark brown, or strongly bicolorous; laminae adaxially short-
pubescent to glabrescent, if adaxially short-pubescent then either not lustrous or pale green
[indusia absent or sphaeropteroid].
15. Laminar hairs present between veins (may also be present on veins); pinna pairs 2236
per frond; [indusia absent].
16. Petiole scales concolorous or if bicolorous then with only weakly darker center,
restricted to petiole bases; petioles glabrous or glabrescent with short hairs; plants
with erect trunks. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. bipinnatida
16. Most petiole scales with dark brown to blackish center, persistent and extending to
rachises; petioles persistently long hairy; plants without trunks or trunks short and
decumbent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. phegopteroides
15. Laminar hairs absent between veins (may be present on veins); pinna pairs 1018(24)
per frond [indusia absent or sphaeropteroid].
17. Petioles scurfy, only distally hairy near transistion to rachis, segment veins
usually lacking hairs on both sides (or with trichomidia instead) [indusia absent]
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. thelypteroides
17. Petioles hairy, not scurfy, segment veins hairy on both sides.
18. Pinnae 2.513.01.02.5 cm, paraphyses relatively long (0.40.5 mm) [indusia
absent]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. palaciosii
18. Pinnae to 3.21.2 cm, paraphyses relatively short (0.20.3 mm) [indusia
sphaeropteroid]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. concordia
1. Laminae bipinnate to tripinnate-pinnatid.
19. Fronds only bipinnate, ultimate segments (= pinnules) entire to coarsely crenate, basal segments of largest
pinnules incised less than 1/2 towards the costae.
2011] LEHNERT: CYATHEACEAE OF PERU 15

20. Plants with erect trunk; rachises matte green in fresh material, stramineous to gray-brown when dried,
notably hairy on both sides; laminae rm-herbaceous, persistently short-pubescent; indusia discoid to
hemitelioid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. bipinnatida x delgadii
20. Plants without trunk; rachises gray-brown to shiny black in fresh material, not changing when dried, at rst
glance glabrous although appressed hairs may occur adaxially; laminae chartaceaous to coriaceous, glabrous
or glabrescent with sparse hairs; indusia never discoid.
21. Lower petiole scales bicolorous dark brown with distinct white margins; petiole scurf inconspicuous but
notable as amorphous pale brown to whitish layer; indusia absent. . . . . . . . . . . . . . . . . . . . C. werfi
21. Lower petiole scales concolorous reddish to dark brown or with slightly paler margins; petiole scurf
absent or brown and squamular; indusia hemitelioid to sphaeropteroid.
22. Largest pinnules lanceolate to weakly hastate with almost entire margins, sinuses blunt and incised
less than 1/4 the way towards the costules); petiole scurf absent; indusia hemitelioid, covered
entirely by sporangia in intact sori. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. polliculi
22. Largest pinnules lanceolate to oblanceolate without basal auricles, with acute sinuses incised at least
1/3 towards the costules; petiole scurf brown, with squamulae; indusia sphaeropteroid, large
fragments seen at maturity. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. chontilla
19. Fronds bipinnate-pinnatid to tripinnate, basal segments ofpinnules incised at least 1/2 towards the costae,
sometimes free and remote from subsequent segments.
23. Petiole scales completely white or very pale brown, dark marginal teeth or darker brown basal spots may
occur, but color does not reach scale tips; petiole scurf absent to dense, but always pale (usually agreeing
with scale color).
24. Petiole scale margins in distal parts reduced to a continuous row of dark brown teeth, young fronds with
ephemeral tortuous hairs, paraphyses longer than sporangia, abundant, contorted over the sori [indusia
absent]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. poeppigii
24. Petiole scale margins not reduced, marginal dark brown teeth not as a continuous row if present; tortuous
laminar hairs absent; paraphyses straight, shorter than sporangia or of the same length [indusia hemitelioid
to sphaeropteroid].
25. Petioles hairy, petiole scurf absent or very sparse, inconspicuous; segment margins inciso-serrate to double-
serrate; rachises without larger scales; sori medial to supramedial [indusia hemitelioid]. . C. leucolepismata
25. Petioles not hairy, petiole scurf either dense or consisting of distinct, separate squamules; segment margins
subentire to crenate; rachises often with larger scales; sori subcostal to costal.
26. Largest pinnules sessile or subsessile; scurf consisting of distinct squamules [indusia sphaeropteroid].
27. Scurf squamules intergrading with larger scales, lacking dark marginal teeth, scurf near petiole
bases dense, usually covering the petiole surface completely.. . . . . . . . . . . . . C. atahuallpa
27. Scurf squamules not intergrading with larger scales, often with dark marginal teeth, scurf near
petiole bases sparse, usually giving sight of the petiole surface. . . . . . . . . . . . . C. straminea
26. Largest pinnules long stalked, i.e., stalk longer than proximal segment breadth; scurf matted, no
individual squamules discernable [indusia hemitelioid].
28. Largest pinnules to 2.5 cm wide, their proximal segments subentire to crenate, most lateral
veins furcated. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .C. heliophila
28. Largest pinnules to 5 cm wide, their proximal segments partially pinnatid, most lateral veins
bifurcate.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .C. vilhelmii
23. Scales orange-brown, brown, castaneous or blackish, either concolorous or bicolorous with weakly to
strongly contrasting, lighter colored margins, if scales with parts purely white then with continuous brown
to blackish central streak; petiole scurf absent to dense, whitish to dark castaneous.
29. Petioles and rachises densely pubescent on both sides, with pluricellular spreading hairs 15 mm long,
other leaf axes with matching pubescence.
30. Pinnae opposite; largest petiole scales 46 mm wide.
31. Petiole scales bicolorous with dark brown to blackish centers and whitish margins, petioles
spiny, often glaucous in fresh material [exindusiate]. . . . . . . . . . . . . . . . . . . . C. conjugata
31. Petiole scales concolorous orange-brown; petioles inermous to verrucate, never glaucous
[indusia sphaeropteroid]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. obnoxia
30. Pinnae alternate; largest petiole scales less than 4 mm wide, sometimes hair-like.
32. Scales concolorous reddish brown to pale stramineous with darker central streak, often
intergrading with hairs; paraphyses 0.60.8 mm long, longer than the sporangia but distal parts
hyaline, often bent or broken off. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. aterrima
32. Scales orange to dark castaneous to vinaceous, not intergrading with hairs; paraphyses to
1.0 mm long, whitish to yellowish, usually persistent.
33. Petiole bases with dense reddish scurf of branched hairs, petiole scales orange to auburn, with wide
persistent margins, paraphyses abundant, contorted, forming a dense persistent mass.C. lockwoodiana
33. Petiole bases lacking scurf, or with few ephemeral white akey squamules, petiole scales
dark castaneous to vinaceous, shiny, with narrow white, fragile margins, paraphyses
relatively few, straight, not contorted. . . . . . . . . . . . . . . . . . . . . . . . . C. pilosissima
16 BRITTONIA [VOL 63

29. Petioles and rachises without hairs, or only adaxially pubescent with mostly appressed hairs 0.5
1.0 mm long, other leaf axes may have stronger pubescence.
34. Petiole scales very narrowly lanceolate, to 3.5 mm wide, basally often with two hook-like auricles
(hastulate), sometimes hair-like, color stramineous to reddish brown, concolorous or weakly
bicolorous with diffuse transition between centers and margins; plants often precociously fertile
[exindusiate or indusia hemitelioid or sphaeropteroid].
35. Largest pinnules less than 51.5 cm, sessile; plants trunkless [indusia sphaeropteroid].C. chontilla
35. Largest pinnules to 12.02.5 cm, subsessile to notably stalked; plants with trunk [indusia hemitelioid].
36. Largest pinnules subsessile, basally truncate to weakly cuneate; segments short- to long-
acute. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. rufescens
36. Largest pinnules stalked 26 mm, basally cordate (smaller ones also truncate); segments
round to obtuse or short-acute.
37. Sori medial; indusia with dentate margins, relatively large, reaching to 1/51/4 around
the receptacles, arching but not covering the sori; largest pinnules predominantly 20
35 mm wide; laminae abaxially with dark brown to auburn scales with paler margins,
both at and bullate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. traillii
37. Sori supramedial to inframarginal; indusia with entire margins, relatively small,
reaching to 1/101/5 around the receptacle, mostly hidden by the sporangia; largest
pinnules predominantly 920 mm wide; laminae abaxially with concolorous dark
brown to atropurpureous lanceolate scales, bullate ones lacking. . . . . . C. macrosora
34. Petiole scales ovate to ovate-lanceolate, if only lanceolate then more than 3.5 mm wide, never
hastulate or looking like hairs, color pale brown, auburn to blackish, concolorous to strongly
bicolorous with sharp transition; indusia hemitelioid to sphaeropteroid or exindusiate; plants
usually not precociously fertile.
38. Pinnae opposite throughout the frond, rarely a pair askew.
39. Petioles inermous, never glaucous; costae inermous; petiole scales concolorous orange-
brown; costae pependicular or weakly decurrently inserted [indusia sphaeropteroid]
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. obnoxia
39. Plants petioles weakly to strongly spiny, often glaucous in fresh material; costae often
verrucate to muricate, short spines to 2 mm long; petiole scales blackish brown with white
margins to concolorous pale brown or white in one plant; costae with surcurrent insertions
[exindusiate]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. conjugata
38. Pinnae alternate throughout the frond, rarely one of the fronds with most pinnae opposite.
40. Costae with scattered spines to 5 mm long [exindusiate]. . . . . . . . . . . . . C. microdonta
40. Costae inermous or muricate with short spines to 1 mm long.
41. Petiole scales with distinct white margins (sometimes very narrow and abraded in
outer scale layers).
42. Pinnules blunt-tipped, round to short-obtuse, pinnules often obovate-oblong
[exindusiate].
43. Rachises green-alate, laminae short-pubescent, but without tortuous hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. mucilagina
43. Rachises not green-alate, laminae with tortuous hairs but straight hairs sparse
and scattered. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. tortuosa
42. Pinnules with pointed tips, acuminate to long-attenuate, pinnules mostly
lanceolate to triangular, rarely oblanceolate.
44. Sori medial to inframarginal; petiole scurf ephemeral and usually absent,
weakly developed already in croziers, sometimes substituted by hairs.
45. Segment margins dentate to serrate; hemitelioid indusia hidden under the
sori, appressed to lamina or weakly ascending.
46. Leaf axes and veins with many hairs; petiole scales brown with broad
cretaceous margins, often transient with concolorous white scales,
never of reddish color. . . . . . . . . . . . . . . . . . . . C. leucolepismata
46. Leaf axes and veins with scattered hairs; petiole scales dark brown to
brown with narrow, lighter colored margins, sometimes nearly
concolorous and reddish. . . . . . . . . . . . . . . . . . . . . . . .C. lindigii
45. Segment margins entire to weakly crenulate; indusia absent or
hemitelioid indusia arching over the sori.
47. Costae distally not or only weakly alate; costules weakly raised
abaxially, without dark spot at their bases; laminar indument with
few dark-brown at, ovate-lanceolate squamules; pinnule bases
cordate [indusia hemitelioid]. . . . . . . . . . . . . . . . . . . . C. andina
47. Costae distally weakly to strongly alate; costules often strongly
raised abaxially, always with dark spot at their bases; laminar
indument with whitish to pale brown, weakly bullate to lamentous
squamules; pinnule bases mostly truncate to cuneate [exindusiate].
2011] LEHNERT: CYATHEACEAE OF PERU 17

48. Sori medial; petiole scales with relatively long marginal cilia,
sometimes few dark brown; laminar indument brown to dark
brown. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. dombeyi
48. Sori supramedial; petiole scales with short marginal cilia, always white,
never dark brown; laminar indument whitish to tan. . . . . . . C. pungens
44. Sori inframedial to costal on segments; petiole scurf well developed, a dense cover
in croziers, remaining at least in patches near the petiole bases between spines (if
present) and larger petiole scales.
49. Petiole scurf consisting of distinct squamules, these isodiametric to lanceolate,
multiciliate or dentate-mbriate, sometimes tufted but not matted; laminar
indument without linear-lanceolate squamules that are transitional to branched
hairs (which may be present); indusia sphaeropteroid, visible at maturity as
irregularly shaped fragments, sometimes only as lacerate discs around the
receptacles.
50. Petiole scurf consisting of whitish to tan, appressed to ascending, small
squamules with mbriate to ciliate margins, without dark marginal teeth nor
transitional to larger petiole scales.
51. Petiole scurf consisting of at, round to ovate-lanceolate squamules with
paler marginal cilia, these not spreading.. . . . . . . . C. austropallescens
51. Petiole scurf consisting of isodiametric squamules with twisted marginal
cilia (appearing like snowakes or cottonballs).
52. Laminar indument without bullate squamules; petiole scurf squamules
of more or less the same size; segments without hairs adaxially on the
veins; leaf axes abaxially glabrous or glabrescent with sparse scurf;
indusia very fragile and often ephemeral. . . . . . . . . .C. tungurahuae
52. Laminar indument with bullate squamules; petiole scurf squa-
mules of different sizes; segments with few to many hairs
adaxially on the veins; leaf axes abaxially densely covered with
easily abraded white scurf; indusia fragile but ususally persistent
as fragments.. . . . . . . . . . . . . . . . . . . . . . . C. cystolepis
50. Petiole scurf consisting of erect, stramineous, cream, or white lanceolate squamules,
often transitional to larger petiole scales and/or with dark marginal teeth.
53. Petiole scurf diffuse, petiolar surface visible between scurf and petiole
scales; petiole scales with relatively rm, persistent margins; fronds
sometimes with visible black spots between veins.
54. Scurf squamules without dark marginal teeth but usually with
cropped tips; petiole scales concordantly bicolorous to almost
concolorous dark brown to blackish with white to pale brown
margins; laminae usually with visible black spots between the veins
(fungal infection). . . . . . . . . . . . . . . . . . . . . . . . C. dintelmannii
54. Scurf squamules often with dark marginal teeth, petiole scales only pale
brown to brown, discordantly bicolorous with whit eto tan margins
laminae without black spots between the veins. . . . . . . . . C. straminea
53. Petiole scurf dense, obscuring petiolar surface between petiole scales;
petiole scales with brittle, fragmenting margins; laminae without black
spots between the veins.
55. Largest pinnules notably stalked, stalk usually longer than breadth of
basal segment; laminar indument absent to sparse (rarely strongly
developed on leaf axes), appearing concolorous brown, consisting
mainly of hyaline branched hairs with brown tips and thin
concolorous brown squamellae; indusia brown, rm, persisting
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. divergens
55. Largest pinnules sessile to subsessile, stalk length not reaching width
of basal segment; laminar indument sparse to abundant, especially
on costules, not appearing concolorous brown, either white or
bicolorous; indusia light brown to colorless, persisting to ephemeral
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. ruiziana
49. Petiole scurf of squamules or hairs matted or appressed, a closed cover in young
fronds, wearing off in akes, whitish, gray or pale brown; laminar indument with
linear-lanceolate squamules and small scales, both with mbriate to ciliate margins
and often transitional to branched hairs; indusia sphaeropteroid, hemitelioid or absent.
56. Petioles with pairs of basal adventitious pinnae, or if these absent then
laminae either triangular or basally decurrent into a very short petiole;
petiole scurf scarce and appressed, squamules with short hyaline cilia
[indusia sphaeropteroid] . . . . . . . . . . . . . . . . . . . . . . . . . . C. parvifolia
18 BRITTONIA [VOL 63

56. Petioles without basal adventitious pinnae, clearly set apart from the elliptic
laminae, these never basally decurrent; petiole scurf usually dense (but easily
worn off).
57. Sori indusiate, indusia fragmented at maturity and/or hidden by the
paraphyses.
58. Petioles spiny; paraphyses relatively sparse, of the same length or shorter
than the sporangia; remnants of the sphaeropterpoid indusia visible
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. patens
58. Petioles inermous; paraphyses numerous, often tufted, of the same
length or longer than the sporangia; receptacles often with larger scale
that covers the sori, indusia hemitelioid, small, appressed.
59. Fertile plants with erect trunks, fronds patent to arching; pinnae and
pinnules sessile to subsessile; paraphyses of the same length as the
sporangia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. arnecornelii
59. Fertile plants trunkless or with ascending rhizomes, fronds hanging
from cliffs or scrambling over adjacent vegetation; pinnae and
pinnules stalked; paraphyses numerous, longer than the sporangia,
hiding the indusia. . . . . . . . . . . . . . . . . . . . . . . . . . . .C. serpens
57. Sori exindusiate (although at to bullate squamules may be attached to the
receptacles).
60. Petioles inermous, petiole scales ovate-lanceolate; paraphyses longer
than the sporangia [Peru and southern Ecuador only]; segments gibbose,
margins revolute; fertile plants either with trunks several meter tall or
trunkless, in latter case the usually arching to drooping fronds often
contracted and ascending.. . . . . . . . . . . . . . . . . . . . . . . . . C. frigida
60. Petioles spiny, petiole scales lanceolate, dull brown to auburn;
paraphyses of the same length as or shorter than sporangia (0.3
0.4 mm); segments planar, margins not revolute; fertile plants always
with trunk several meter tall, fronds presumably patent-arching and
only distally drooping. . . . . . . . . . . . . . . . . . . . . . . . . . C. nephele
41. Petioles scales without white margins, either concolorous or margins with another
substantial color (orange to yellow) in it.
61. Rachises green alate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. mucilagina
61. Rachises not alate.
62. Ultimate segments obovate-oblong with cuneate bases and obtuse tips, usually so
narrow that sori are extending into sinuses and visible from the adaxial laminar side.
63. Fronds fully tripinnate to tripinnate-pinnatid (free basal segments of pinnules with
almost free lobes), to 2 m long, with ca. 1215 pinna pairs. . . . C. multisegmenta
63. Fronds only bipinnate-pinnatid to partly tripinnate, 0.31.0 m long, with
fewer than 10 pairs of pinnae.
64. Pinnules obtuse to rounded; leaf axes abaxially with many dark brown
bullate squamules.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. microphylla
64. Pinnules short-acute to acuminate; leaf axes abaxially with few brown at
and bullate squamules. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. dudleyi
62. Ultimate segments linear-oblong to triangular, basally adnate to decurrent, sori not
extending into sinuses, not visible adaxially.
65. Veins irregularly forming areoles [indusia hemitelioid]. . . . . . . . . . C. bradei
65. Veins never forming areoles [indusia sphaeropteroid or absent].
66. Petioles verrucate to aculeate, often strongly so, scurf consisting of distinct
squamules, sparse to dense, but often ephemeral [indusia sphaeropteroid].
67. Petiole scales concolorous or nearly so, rather dull brown to lustrous
orange-brown with somewhat paler margins; scurf either scarce or
ephemeral, missing in fully expanded fronds, sometimes scurf replaced
by dense pubescence of muticellular hairs to 2 mm long on upper
petiole parts and rachises.
68. Hairs often present between the veins abaxially but not adaxially;
petiole scales lustrous orange-brown, contrasting with the blackish
epidermis of the lower petiole parts, rarely scales dull brown but
scurf always relatively ne and soon caducous.. . . . . . C. delgadii
68. Hairs present between the veins adaxially; petiole scales dull brown,
petiole epidermis dull brown to stramineous or yellow, with
relatively large, persisting scurf squamules. . . . . . . . . . C. herzogii
67. Petiole scales bicolorous with dark brown to blackish centers at least in
basal petiole parts, if concolorous then dark brown to atropurpureus and
petiole scurf long persisting.
2011] LEHNERT: CYATHEACEAE OF PERU 19

69. Scales of lower parts of petioles with relatively narrow blackish

center and broad, ferrugineous to yellow-brown margins that are
conuent towards the tips; squamules of petiole scurf to 1.0 mm;
rachises sometimes with long-persistent scales; largest pinnules
sessile or nearly so (stalks 1 mm or less). . . . . . . . C. catacampta
69. Scales of lower parts with relatively broad, dark brown to blackish
centers, the marginal color not conuent towards the tips; squamules of
petiole scurf to 0.5 mm; rachises without scales or with ephemeral
concolorous scales; largest pinnules often notably short-stalked to 3 mm.
70. Veins and costules abaxially glabrous or sparsely hairy, hairs ca.
0.5 mm long; petiole scales narrowly lanceolate to lanceolate,
15333.54.5 mm, the brown to dark brown center not sharply
set against the brown to orange margins, often appearing
concolorous; laminar indument with mainly at (rarely sub-
bullate) squamules. . . . . . . . . . . . . . . . . . . . . . C. lindeniana
70. Veins and costules abaxially sparsely to densely hairy, veins
usually more pubescent than costules, hairs 0.51.0 mm long;
petiole scales relatively broadly lanceolate, to 154 mm,
atropurpureus to blackish on the sides and the bases of the
petioles, dark brown on upper petiole parts and towards the
trunks, always with distinct lighter margins, brown to orange,
rarely in parts yellowish or cream-white; laminar indument with
at and bullate squamules. . . . . . . . . . . . . . . . . C. squamipes
66. Petioles inermous, or sparsely verrucate to muricate near the bases, scurf
absent or inconspicuous, i.e. amorphous or squamellae ne, matted and in
color of the petiolar epidermis [indusia sphaeropteroid or absent].
71. Plants without trunk; leaf axes with whitish-gray scurf of matted
tortuous hairs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. pratermissa
71. Plants with trunk; leaf axes without grayish, matted scurf, hairs if present
arching to spreading, not tortuous.
72. Paraphyses longer than the sporangia, distally sometimes contorted.
73. Fronds with 78 pinna pairs; veins connivent to sinuses; largest
pinnules more than 2 cm wide, segments usually appearing
truncate (patent with short but strongly falcate tips). . . .C. ulei
73. Fronds with 1015 pinna pairs, veins not connivent to sinuses;
largest pinnules less than 2 cm wide, segments usually rounded
to obtuse at tips.
74. Costae abaxially glabrous to glabrescent with white hairs to
1 mm long; paraphyses relatively thin, distal parts fragile
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. lasiosora
74. Costae abaxially with reddish spreading hairs to 2 mm long;
paraphyses forming a dense persistent mass. . C. lockwoodiana
72. Paraphyses shorter than or of the same length as the sporangia.
75. Fronds very long (to 7 m), with 1220 pinna pairs, ascending,
distally drooping, supported on surrounding vegetation; sori
exindusiate.. . . . . . . . . . . . . . . . . . . . . . . . . . . C. kalbreyeri
75. Fronds relatively short (to 34 m long), with 715 pinna pairs,
arching, not supported on surrounding vegetation; sori indusiate,
indusia sphaeropteroid.
76. Petiole scales concolorous brown, not contrasting with the
petiolar epidermis.
77. Plants usually epiphytic, rarely terrestrial on open slopes;
trunks to 1.0 m tall; largest pinnules more than 2 cm
broad, broadly lanceolate to triangular, segment tips
acute, margins crenulate to serrate . . . . . .C. gracilis
77. Plants always terrestrial; trunks to 4(12) m tall; largest
pinnules less than 2 cm broad, linear-lanceolate to
weakly oblanceolate; segment apices round to obtuse,
margins subentire to crenulate. . . . . . . . . . .C. lechleri
76. Petiole scales bicolorous to concolorous orange brown, con-
trasting with the petiolar epidermis.
78. Trunks less than 5 cm diam.; receptacles of the sori
medial; green costules contrasting with the shiny
castaneous to blackish costae . . . . . . . . . C. ebenina
20
SYSTEMATIC TREATMENT
Alsophila R. Br. sect. Alsophila
Alsophila cuspidata (Kunze) D. S. Conant
Depts. Amazonas, Cuzco, Huanuco, Lor-
eto, Madre de Dios, Pasco, Puno, Ucayali.
1001100(1570) m. (Costa Rica, Panama,
Colombia, Venezuela, French Guiana, Ecua-
dor, Peru, Bolivia, Paraguay)
Voucher: Madre de Dios: Tambopata, ca.
30 air km or 7080 km by river SSW of
Puerto Maldonado at efuence Ro La Torre
(Ro Dorbigny)/Ro Tambopata (SE bank),
Tambopata Nature Reserve, 1249S, 6917W,
260 m, 14 May 1984, Barbour 5297 (F, MO)
Reference: Tryon & Stolze, 1989.
Alsophila engelii R. M. Tryon
Depts. Cajamarca, Cuzco, San Martin. 1750
2150 m. (Colombia, Venezuela, Ecuador, Peru).
Voucher: Cusco: La Convencin, 1258S,
07233W, 2150 m, 24 Nov 2003, Bonino et
al. 1155 (CUZ, MO).
Reference: Tryon & Stolze, 1989.
BRITTONIA [VOL 63
78. Trunks more than 5 cm diam.; receptacles of the sori
costal to subcostal; costules not contrasting with the
costae.
79. Scales on lower parts of petioles narrowly lanceolate, to
5 cm long; largest pinnules less than 2 cm broad, all
sessile to subsessile (exceptionally short-stalked to
3 mm), usually held upright with respect to the costae
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .C. plicata
79. Scales on lower parts of petioles broadly lanceo-
late, to 3 cm long; largest pinnules usually more
than 2 cm broad and notably stalked (dwarfed
plants may disgress), all held in one plane with
the costae.
80. Larger squamules of petiole scurf asymmetri-
cally lanceolate, squamellar margins not
appearing paler in backlight; laminar indu-
ment dark brown to castaneous, of same
color as the petiole scurf; trunks shedding
petioles when smaller than 2 m, reaching 15
20 cm diam.; stout, massive plants. . . . . . .
. . . . . . . . . . . . . . . . . . C. carolihenrici
80. Larger squamules of petiole scurf oblong to
oblanceolate, squamellar margins appearing
paler in backlight; laminar indument brown to
yellowish brown, paler than the petiole scurf;
trunks with persistent petiole bases at least when
smaller than 2 m, reaching 512 cm diam.; sizes
and proportions of plants variable but habit
always gracile.. . . . . . . . . . . . . C. meridensis
Alsophila erinacea (H. Karst.) D. S. Conant
Depts. Cajamarca, Cuzco, Pasco. 1780
2800 m. (Costa Rica, Colombia, Ecuador,
Peru, Bolivia)
Voucher: Cajamarca: San Ignacio, San
Jos de Lourdes, 050042S, 785415W,
1800 m, Feb 2006, Bonino 59 (MO, UC).
Reference: Tryon & Stolze, 1989.
Alsophila imrayana (Hook.) D. S. Conant
Depts. San Martin. Ca. 2000 m. (Venezuela,
Trinidad, Colombia, Ecuador, Peru)
Voucher: San Martin: Rioja, Carretera
Moyobamba-Pedro Ruiz, Km 404.1, 5 Aug
2002, Lehnert 214 (GOET, USM, UC).
Reference: Gastony, 1973; Conant, 1983.
Alsophila incana (H. Karst.) D. S. Conant
Depts. Junin, Pasco, Ucayali. 1130
1500 m. (Colombia, Ecuador, Peru)
Voucher: Pasco: Oxapampa, Dist. Huan-
cabmba, sector Pan de Azucar (Huampal),
101111S, 753424W, 1130 m, 11 Nov
2004, Mellado 221 (MO, UC).
2011] LEHNERT: CYATHEACEAE OF PERU
References: Tryon & Stolze, 1989; Lehnert,
2005.
Alsophila gastonyi Lehnert, sp. nov.
Dept. Cuzco, Pasco. 520950 m. (Peru)
Voucher: Pasco: Prov. Oxapampa, Dist.
Palcaz, Reserva Comunal Yanesha, Comu-
nidad Nativa San Pedro de Pichanaz, Sector
Azulis, 102644S, 750621W, 910 m, 15
Sep 2005, Monteagudo A. et al. 9810
(holotype: MO).
Alsophila minervae Lehnert
Dept. Cuzco, Pasco. 22002900 m. (Peru,
Bolivia)
Voucher: Pasco: Oxapampa, trail to sum-
mit of Cordillera Yanachaga via Rio San
Daniel, 1023S, 7527W, 2500 m, 17 Jul
1984, Smith D. N. et al. 7817 (USM).
References: Lehnert, 2003; Smith et al., 2005.
Alsophila mostellaria Lehnert
Depts. Amazonas, Cajamarca, Pasco. 1500
2800 m. (Ecuador, Peru, Bolivia)
Voucher: Amazonas: Bongar, carretera
Pedro Ruiz-Florida, 0551.7S, 7758.4W,
2200 m, 5 Aug 2002, Lehnert 243 (holotype:
USM; isotypes: GOET, UC).
Reference: Lehnert, 2003, 2005.
Alsophila paucifolia Baker
Depts. Ucayali, Amazonas, ca. 9001300 m.
(Colombia, Ecuador, Peru)
Voucher: Ucayali (Loreto): Prov. Coronel
Portillo, La Divisoria, 29 Jul 1948, Aguilar
850 (USM).
Reference: Tryon & Stolze, 1989.
Remarks: First conmed report for Peru;
the presence was long suspected (Tryon &
Stolze 1989).
Alsophila R. Br. sect. Gymnosphaera Blume
Alsohila salvinii Hook.
Dept. San Martin. 1850 m. (Mexico,
Guatemala, El Salvador, Belize, Honduras,
Costa Rica, Panama, Peru)
Voucher: San Martn: Rioja, 054029S,
774625W, 1850 m, 25 Mar 1998, van der
Werff et al. 15666 (MO, USM).
References: Stolze, 1976; Conant, 1983;
Mickel & Smith, 2004.
21
Cyathea J. Sm.
Cyathea alatissima (Stolze) Lehnert,
comb. nov.
Dept. Hunuco. 1500 m. (Peru)
Voucher: Hunuco: Ro Llulla Pichis water-
shed, 25 Jul 1969, Dudley 13282 (holotype:
GH-photo F; isotype: NA n.v.).
References: Stolze, 1974; Tryon & Stolze,
1989.
Cyathea andina (H. Karst.) Domin
Dept. Madre de Dios. 200 m. (Greater
Antilles, Colombia, Ecuador, Peru, Bolivia)
Voucher: Madre de Dios: Tambopata,
20 km NW from the mouth of Ro Inambari
N bank of Ro Madre de Dios, at the end of the
airstrip of Lagarto 1238S, 6951W, 200 m
11 Nov 1998, Tuomisto et al. 13679 (TUR).
Reference: Tryon & Stolze, 1989.
Cyathea arnecornelii Lehnert
Dept. Cuzco. 2980 m. (Peru, Bolivia)
Voucher: Cuzco: La Convencin, Dist.
Quellouno, Yavero, 12 3535S, 722321
W, 2980 m, Valenzuela et al. 7784 (MO, UC).
Det. A. R. Smith 2008.
References: Lehnert, 2003; Smith et al.,
2005.
Cyathea atahuallpa (R. M. Tryon) Lellinger
Depts. Amazonas, Cajamarca, La Libertad.
27003100 m. (Ecuador, Peru)
Voucher: Amazonas: Chachapoyas, Cerros
Calla Calla, E side, 18 km above Leime-
bamba on the road to Balsas, 30003100 m,
15 Oct 1968, Hutchinson & Wright 6922
(holotype: US; isotypes: NY, UC, US).
References: Lellinger, 1987; Tryon &
Stolze, 1989.
Cyathea aterrima (Hook.) Domin
Depts. Amazonas, San Martin. 700
1200 m. (Colombia, Ecuador, Peru)
Voucher: San Martin: In monte Guayr-
apurima, prope Tarapoto, Peruvia orientalis,
Sept 1856, Spruce 4713 (holotype: BM;
isotypes: K, US).
References: Tryon & Stolze, 1989; Moran
et al. 2008.
22 BRITTONIA
Cyathea austropallescens Lehnert
Depts. Cuzco, Pasco. 24103200 m. (Peru,
Bolivia)
Voucher: Pasco: Oxapampa Dist. Oxa-
pampa, P. N. Yanachaga-Chemilln, Abra
Yanachaga, 1022S, 7527W, 2900 m, 1
Aug 2003, Vsquez et al. 28458 (UC, MO).
Reference: Lehnert, 2008.
Cyathea bipinnatida (Baker) Domin
Depts. Amazonas, Cajamarca, Cuzco, Madre
de Dios, Pasco, San Martin. 4002100 m.
(Venezuela, Colombia, Ecuador, Peru, Bolivia)
Voucher: Amazonas: Bagua, Quebrada El
Almendro, 051440S, 782124W, 430 m,
8 Mar 1998, van der Werff et al. 14506 (MO,
UC, USM).
Reference: Tryon & Stolze, 1989.
Cyathea bradei (Windisch) Lellinger
Dept. Loreto, San Martin. 100200 m.
(Colombia, Ecuador, Peru, Bolivia)
Voucher: Loreto: Ro Tigre, 5 km N of the
village Paicheplaya E of the river, 0259S,
7518W, 100200 m, 31 Jan 2005, Tuomisto
et al. 14768 (GOET, TUR).
References: Lellinger, 1984; Tryon &
Stolze, 1989.
Cyathea carolihenrici Lehnert
Dept. Cuzco. 2050 m. (Peru, Bolivia)
Voucher: Cuzco: Prov. Urubamba, 2050 m,
Bonino 1127 (CUZ).
References: Lehnert, 2003, Smith et al., 2005.
Cyathea catacampta Alston
Depts. Cuzco, San Martin. 20003050 m.
(Colombia, Ecuador, Peru, Bolivia)
Voucher: Cuzco: La Convencin, Dist.
Huayopata, Balconpata, 125201S, 7232
46W, 2200 m, 21 Apr 2004, Calatayud et al.
2342 (MO
Reference: Lehnert, 2009a.
Cyathea chontilla Lehnert, sp. nov.
Depts. Cajamarca, Pasco. 23002850 m. (Peru)
Voucher: Cajamarca: San Ignacio, Huar-
ango, localidad Romerillo, Cordillera entre
Romerillo y Nuevo Mundo, 0514S, 7846
W, 2300 m, 29 Jul 1992, Campos et al. 4294
(holotype: UC; isotypes: MO, USM).
[VOL 63
Cyathea concordia B. Len & R. C. Moran
Dept. Amazonas. 18002050 m. (Peru)
Voucher: Amazonas: Condorcanqui, Cor-
dillera del Cndor, Cerro Machinaza, Puesto
de Vigilancia Alfonso Ugarte, cabeceras del
Ro Comainas, 0422S, 7811W, 1800
2050 m, 21 Jul 1994, Bltran & Foster
1183 (holotype: USM; isotype: UC-fragm.).
Reference: Len & Moran, 1996.
Cyathea conjugata (Hook.) Domin
Depts. La Paz, Cochabamba, Santa Cruz.
11002750 m. (Colombia, Ecuador, Peru,
Bolivia)
Voucher: Cuzco: Urubamba, Dist. Machu
Picchu, Cerro Aguas Calientes, 1309S 72
31W, 30003500 m, 22 Nov 2003, Huaman-
tupa & Quispe 3838 (MO, UC)
References: Lellinger, 1987; Tryon & Stolze,
1989.
Remarks: Cyathea tryonorum (Riba) Lel-
linger is believed to be conspecic.
Cyathea crenata (Sodiro) H. Christ
Dept. Cajamarca. 1900 m. (Ecuador, Peru)
Voucher: Cajamarca: San Ignacio, Dist.
San Jos de Lourdes, Buenos Aires (Cerro El
Parco), 054204S, 775306W, 1900 m,
Feb 2002, Bonino 307 (MO, UC).
Reference: Lehnert, 2009a.
Cyathea cystolepis Sodiro
Depts. Amazonas, San Martin. 2060
2750 m. (Colombia, Ecuador, Peru)
Reference: Lehnert, 2008.
Remarks: Three varieties can be distin-
guished, two of them in Peru:
1. Hairs on the segments more abundant adaxially or
equally dense on both sides, generally erect and
abundant on and between the veins; bullate
squamules frequent abaxially, mainly pure white,
rarely with brown body. . . . . . . . . . . . var. leonis
1. Hairs on the segments more abundant abaxially
than adaxially on and between the veins, except if
hairs generally few and appressed, then sometimes
few hairs adaxially and almost no hairs abaxially,
hairs mainly on the veins; bullate squamules with
brown body if abundant.
2. Hairs on the segments more abundant abaxially,
sparse to abundant, mainly erect adaxially,
appressed adaxially, bullate squamules few to
many, always brown-bodied and pure white
ones mixed, the brown-bodied squamules often
dominating. . . . . . . . . . . . . . . . var. cystolepis
2011] LEHNERT: CYATHEACEAE OF PERU
2. Hairs sparse on both laminar surfaces, often
appressed, bullate squamules abaxially few, only
white, or lacking. . . . . . . . . var. boreopallescens
Cyathea cystolepis var. boreopallescens
Lehnert
Voucher: Amazonas: Bagua Cordillera
Coln SE of La Peca, 25002600 m, 30 Sep
1982, Barbour 3611 (AAU, MO, USM).
Cyathea cystolepis var. leonis Lehnert.
Voucher: Amazonas: Chachapoyas, Distr.
Leymebamba, alrededor de la laguna de los
Condores, parte sur, 0651.201S, 7740.958W,
25002700 m, 17 Aug 2002, Quipuscoa et al.
1234 (holotype: F; isotypes: GOET, UC).
Cyathea delgadii Sternb.
Depts. Amazonas. Cuzco, Huanuco, Madre
de Dios, Pasco, San Martin. 5652200 m. (Costa
Rica, Panama, Colombia, Venezuela, Ecuador,
Peru, Brazil, Bolivia, Paraguay, Argentina)
Voucher: Pasco: Prov. Oxapampa, Dist.
Oxapampa, sector Grapanazu, relleno sani-
tario, 103314S, 752535W, 1800 m, 17
Mar 2005, Mellado et al. 2904 (MO).
References: Tryon & Stolze, 1989; Lehnert,
2006b.
Remarks: A variable species whose extremes
can be mistaken for different species:
1. Segments 34 mm broad, margins conspicously
crenulate to crenate; rachises abaxially sparsely
hairy, not scabrous with hair bases, though some-
times muricate with little spines; petioles blackish
to purpureous. . . . . . . . .C. delgadiitypical form
1. Segments 23 mm wide, with margins entire or
inconspicuously crenulate; rachises abaxially
densely hairy or, if hairs abraded, notably scabrous
with hair bases; petioles dull brown to yellowish.
. . . . . . . . . . . . . . . . C. delgadiipubescent form
Typical form: In perhumid premontane and
montane forest of Costa Rica, the Andes and
the mata atlantica of Brazil; mainly above
1000 m.
Pubescent form: In premontane forest with
weak seasonality and poor soils in the
Venzuelan gran sabana, the Bolivian Chi-
quitania, and the Brazilian campo cerrado;
only below 900 m. In Peru, in the Huanca-
bamba depression of Depts. San Martin and
Amazonas.
23
Cyathea dintelmannii Lehnert
Depts. Cuzco, Pasco. 15352250 m. (Peru,
Bolivia)
Voucher: Cusco: La Convencin, Dist.
Vilcabamba, Espiritupampa, 125425S, 73
1216W, 1535 m, 22 Jul 2004, Calatayud et
al. 2582 (MO, UC).
References: Lehnert, 2006b, 2008.
Cyathea divergens Kunze
Depts. San Martin, Huanuco, Pasco. 1590
1800 m. (Costa Rica, Panama, Colombia,
Venezuela, Ecuador, Peru)
Voucher: Hunuco: Pampayaco, Jul 1829,
Poeppig s.n. (diar. 1163) (holotype: LZ,
destroyed). Authentic specimens (de Tryon
1976): Poeppig 219 (diar. 1152) (B-fragm.
NY, K-fragm. NY, P-photo GH)
References: Tryon & Stolze, 1989; Lehnert,
2008.
Remarks: Of the two currently recognized
varieties (Tryon, 1976; Tryon & Stolze, 1989),
only the var. divergens is native to Peru.
Cyathea dombeyi (Desv.) Lellinger.
Dept. Pasco. 1290 m. (Peru, Bolivia)
Voucher: In Peruvia, Dombey s.n. (hol-
otype: P n.v.; isotype: P n.v.).
Reference: Barrington, 1978.
Remarks: The species was previously includ-
ed in C. pungens (Willd.) Domin (Christensen,
1905; Tryon & Stolze, 1989; Lehnert, 2006a).
Cyathea dudleyi R. M. Tryon
Depts. Amazonas, Cajamarca, Cuzco, Pasco,
San Martin. 19002700 m. (Ecuador, Peru)
Voucher: Cuzco: Prov. Convencin, 12
37S, 7333W, 26002700 m, 7 Jul 1968,
Dudley 10867B (holotype: GH; isotypes:
MO, NA n.v., US).
References: Tryon, 1976; Tryon & Stolze,
1989.
Cyathea ebenina H. Karst.
Depts. Amazonas, Cajamarca, Cuzco,
Pasco, San Martin. 19002700 m. (Colombia,
Ecuador, Peru)
Voucher: Cajamarca: San Ignacio, San
Jos de Lourdes, locality Picorana, 0458
00S 785301W, 23502450 m, 4 Dec
2002, Campos et al. 5898 (UC).
Reference: Tryon & Stolze, 1989.
24 BRITTONIA
Cyathea ewanii Alston
Depts. Amazonas, Loreto. 100660 m.
(Ecuador, Peru)
Voucher: Amazonas: Bagua, Distrito Imaza,
regin del Maran, comunidad de Yamayakat,
Rio Maran, 0455S, 7819W, 600 m 28 Jan
1995, Rodriguez 285 (UC).
Reference: Tryon & Stolze, 1989.
Remarks: Synonym Cnemidaria ewanii
(Alston) R. M. Tryon.
Cyathea frigida (H. Karst.) Domin
Depts. Cajamarca, Huanuco, Pasco. 2650
3300 m. (Colombia, Ecuador, Peru)
Voucher: Hunuco: Huanuco, Huanuco-
Tingo Maria road, steep slopes near Carpish
divide above mouth of tunnel, 0942S, 76
05W, 26502700 m, 3 Feb 1984, Gentry &
D. N. Smith 44863 (MO, NY).
Reference: Tryon & Stolze, 1989.
Remarks: Often confused with C. nephele
Lehnert; Peruvian specimens differ from the
Colombian type material (and most speci-
mens from Ecuador) by having longer para-
physes, fewer or no ciliate laminar squamules,
and on average relatively paler petioles and
leaf axes.
Cyathea frondosa H. Karst.
Dept. San Martin. 20602100 m. (Vene-
zuela, Colombia, Ecuador, Peru)
Voucher: San Martin: Huallaga, arriba de
La Ribera-La Meseta, 0652S, 7728W,
20602100 m, 17 Aug 1997, Quipuscoa-S.
et al. 1085 (AAU).
Reference: Lehnert, 2009a.
Cyathea gracilis Griseb.
Depts. Amazonas, Pasco. 7702550 m.
(Costa Rica, Jamaica, Colombia, Ecuador, Peru)
Voucher: Pasco: Oxapampa, Dist. Palcaz,
San Francisco de Pichanaz, 102916S 75
0358W, 770 m, 28 Feb 2004, Mellado et al.
894 (MO).
Reference: Tryon & Stolze, 1989.
Cyathea heliophila R. M. Tryon
Dept. Pasco. 2500 m. (Colombia, Ecuador,
Peru, Bolivia)
Voucher: Pasco: Oxapampa, Dist. Huanca-
bamba, P. N. Yanachaga-Chemilln, sector Hito-
[VOL 63
San Daniel, 1015S, 7515W, 2500 m, 21 Aug
2005, Monteagudo et al. 9562 (MO, UC).
Reference: Tryon, 1986.
Cyathea herzogii Rosenst.
Depts. Cuzco, Hunuco, Junin, Pasco.
20003200 m. (Peru, Bolivia)
Voucher: Pasco: Oxapampa, Dist. Huanca-
bamba, sector quebrada Yanachaga (P.N.
Yanachaga-Chemilln), 102326S, 7528
21W, 2545 m, 22 Sep 2004, Mellado &
Becerra 1884 (MO).
References: Smith et al. 2005, Lehnert
2006b.
Cyathea horrida (L.) Sm.
Depts. Amazonas, Pasco, San Martin. 320
770 m. (Costa Rica, Panama, Greater and
Lesser Antilles, Colombia, Venezuela, Ecua-
dor, Peru)
Voucher: Amazonas: Bagua, Distr. Imaza,
Quebrada Almendra, 051440S, 782134
W, 400 m, 17 Mar 2001, van der Werff et al.
16134 (MO).
Reference: Tryon & Stolze, 1989.
Remarks: Synonym Cnemidaria horrida
(L.) C. Presl.
Cyathea kalbreyeri (Baker) Domin
Depts. Junin, San Martin. 11701200 m.
(Colombia, Venezuela, Ecuador, Peru, Bolivia)
Voucher: Junin: Satipo, Gran Pajonal, E of
Chequitavo on trail to Kotampaz, 1045S,
7423W, 1200 m, 5 Apr 1998, Smith D. N.
6749 (UC).
Reference: Tryon & Stolze, 1989.
Cyathea lasiosora (Kuhn) Domin
Depts. Amazonas, Loreto, Madre de Dios,
Pasco, Puno, San Martin, 100800(1030) m.
(Colombia, Venezuela, French Guiana, Ecua-
dor, Peru, Brazil, Bolivia)
Voucher: Loreto: Loreto, about 2 km N of
the village San Antonio at lower Ro Mar-
aon, 0432S, 7338W, 100200 m, 14 Jan
1996, Tuomisto et al. 8055 (TUR, UC).
Reference: Tryon & Stolze, 1989.
Cyathea latevagans (Baker) Domin
Dept. Cuzco, San Martin. 18503415 m.
(Colombia, Venezuela, Ecuador, Peru)
2011] LEHNERT: CYATHEACEAE OF PERU
Voucher: San Martin: Rioja, along road
Rioja-Pedro Ruiz, El Mirador, 054029S,
774625W, 1850 m, 26 Mar 2002, van der
Werff et al. 15705 (MO, UC, USM).
Reference: Tryon & Stolze, 1989.
Cyathea lechleri Mett.
Depts. Amazonas, Pasco, Puno, San Mar-
tin, 7802160 m. (Colombia, Venezuela,
Peru, Bolivia)
Voucher: Pasco: Oxapampa, Distrito Pal-
cazu, P.N. Yanachaga-Chemilln, one hour
from Estacin Biologica Paujil, 1020S, 75
15W, 780 m, 25 Oct 2002, Monteagudo et al.
4272 (MO, UC).
Reference: Tryon & Stolze, 1989.
Cyathea leucolepismata Alston
Depts. Loreto, Madre de Dios. 90500 m.
(Colombia, Ecuador, Peru, Bolivia)
Voucher: Loreto: Maynas, between the
roads Ex Petroleros and Bello Horizonte,
Km 3840 of the road Iquitos-Nauta, 0404
S, 7328W, 100200 m, 6 Jan 1995, Tuo-
misto et al. 6677 (GOET, TUR).
Reference: Moran, 1995.
Remarks: Synonym C. amazonica R. C.
Moran.
Cyathea lindigii (Baker) Domin
Depts. Cuzco, Madre de Dios. 400500 m.
(Colombia, Ecuador, Peru, Bolivia)
Voucher: Madre de Dios: Manu, Parque
Nacional Manu, Ro Manu, Rio Sotileja, 11
40S, 7155W, 400500 m, 4 Oct 1986,
Foster & DAchille, 11627 (F, LPB)
Reference: Tryon & Stolze, 1989.
Remarks: Distinct from C. multiora Sm.,
under which it was previously treated (Tryon,
1976).
Cyathea lindeniana C. Presl
Depts. Cajamarca, San Martin. 2000
2200 m. (Colombia, Venezuela, Ecuador, Peru)
Voucher: Cajamarca: Lower edge of
Cutervo National Park, 1015 km N of San
Andres de Cutervo, 0610S, 7840W, 2200 m,
13 Feb 1992, Gentry et al. 61560 (UC).
Reference: Lehnert, 2009a.
25
Cyathea lockwoodiana (Windisch) Lellinger
Dept. Loreto. 100200 m. (Panama,
Colombia, Venezuela, Ecuador, Peru)
Voucher: Loreto: Prov. Loreto, about 2 km N
of the village San Antonio at lower Ro
Maraon, 0432S, 7338W, 100200 m, 14
Jan 1996, Tuomisto H. et al. 8188 (LSC, TUR).
Reference: Windisch, 1977.
Cyathea macrosora (Baker) Domin
Depts. Loreto, Pasco, 100700 m. (Colom-
bia, Venezuela, the Guyanas, Ecuador, Peru)
Remarks: Two varieties can be distinguished:
1. Petioles and leaf axes with little scurf, often
glabrescent; laminar indument dark brown; indusia
well developed. . . . . . . . . . . . . . . . . var. macrosora
1. Petioles and leaf axes with dense scurf, usually
persistent; laminar indument whitish; indusia small,
often partially absent in one frond. . . . var. vaupensis
Cyathea macrosora var. macrosora
Voucher: Pasco: Oxapampa, along road Cha-
tarra-Pto. Bermudz, 1030S, 7503W, 700 m,
9 Jul 2003, van der Werff et al. 18171 (MO, UC).
Cyathea macrosora var. vaupensis (Windisch)
A. R. Sm.
Voucher: Loreto: Loreto, Ro Tigre, about
1.5 km E of the river, 0357S, 7417W, 100
200 m, 10 Jan 2005, Tuomisto et al. 13983
(TUR, USM).
References: Lellinger, 1984; Tryon &
Stolze, 1989.
Cyathea meridensis H. Karst.
Depts. Amazonas, Cuzco, Cajamarca. 2295
2830 m. (Colombia, Venezuela, Ecuador, Peru)
Remarks: Three varieties can be distinguished:
1. Fertile fronds to 1 m long; pinnules to 65(80) mm
long, all sessile or few subsessile; trunks to 1.5 m tall,
covered with persistent petiole bases.. . . . .var. nana
1. Fertile fronds more than 1 m long: largest pinnules
notably stalked, to 120130 mm long; trunks to 3
10 m tall, petiole bases persistent or not.
2. Fertile fronds hanging free, arching to distally
drooping; trunks to 10 m tall, without
persistent petiole bases, petioles falling
cleanly at an early age. . . . . var. meridensis
2. Fertile fronds leaning and draped over
adjacent vegetation; trunks relatively small
compared to the frond length, 0.32.0 m tall,
petiole bases persistent, rotting off only in
older plants. . . . . . . . . . . . . . . var. obtecta
26 BRITTONIA
Cyathea meridensis var. meridensis
Voucher: Cuzco: Paucartambo, Kosipata,
Trocha Unin, Km 10, 19 L01221737, UTM
8552556, 2295 m, 3 Sep 2003, Garca K. et
al. 207 (MO).
Cyathea meridensis var. nana Lehnert
Voucher: Cajamarca: San Ignacio San
Jos de Lourdes, Cerro Picorana, 0458S,
7853W, 2830 m, 17 Aug 1998, Campos et
al. 5550 (MO, UC).
Cyathea meridensis var. obtecta Lehnert
Voucher: Amazonas: Laguna de Pomaco-
chas, 2550 m, 28 Mar 2002, van der Werff et
al. 15835 (MO, UC).
Reference: Lehnert, 2009a.
Cyathea microdonta (Desv.) Domin
Depts. Cuzco, Loreto, Pasco, San Martin.
1001100 m. (Mexico, Central America, Greater
Antilles, Colombia, Venezuela, Trinidad, the
Guyanas, Ecuador, Peru, Brazil, Bolivia)
Voucher: Pasco: Oxapampa, Distr. Palcaz,
El Paujil, 101924S, 751549W, 500 m 11 Depts. Amazonas, Cajamarca, San Martin.
May 2005, van der Werff et al. 19989 (MO).
Reference: Tryon & Stolze, 1989.
Cyathea microphylla Mett.
Dept. Puno. 2100 m. (Peru)
Voucher: Puno: Prov. Carabaya, St. Gavan,
Lechler 2160 (holotype: LZ, destroyed);
authentic specimen Lechler 2569 (B-fragm.
F, P-photo GH).
Reference: Tryon & Stolze, 1989.
Cyathea mucilagina R. C. Moran
Dept. Hunuco, Madre de Dios, Pasco, San
Martin. 490770 m. (Costa Rica, Colombia,
Ecuador, Peru, Bolivia)
Voucher: San Martin: Prov. San Martin,
Progreso, village in the rst valley of the
Cerro Escalera, from Tarapoto after the
tunnel, along Tarapoto-Yurimaguas, 0624S,
7618W, 600 m, 8 Aug 2006, Christenhusz
et al. 2024 (GOET, TUR, USM).
Reference: Moran, 1991.
[VOL 63
Cyathea multisegmenta R. M. Tryon
Depts. Cuzco, Pasco. 18502295 m. (Peru)
Voucher: Cuzco: Prov. La Convencin, Dud-
ley 11326 (holotype: NA; isotypes: GH, US).
References: Tryon, 1976, Tryon & Stolze,
1989.
Cyathea nephele Lehnert, sp. nov.
Depts. Amazonas, Pasco, San Martin.
27203550 m. (Peru)
Voucher: Pasco: Prov. Oxapampa, Distrito
Huancabamba, Parque Nacional Yanachaga-
Chemilln, 1022S, 7528W, 3110 m, 5 Dec
2003, Monteagudo A. et al. 6430 (holotype:
UC; isotype: MO).
Cyathea nervosa (Maxon) Lehnert, comb. nov.
Dept. Amazonas. 300400 m. (Ecuador, Peru)
Voucher: Amazonas: Prov. Condorcanqui,
above Pongo de Manseriche, gully to right of
mouth of Ro Santiago, 300 m, 18 Dec 1931,
Mexia 6291 (holotype: F; isotypes: GH, UC).
Reference: Tryon & Stolze, 1989.
Cyathea obnoxia Lehnert
22002700 m. (Colombia, Ecuador, Peru,
Bolivia)
Voucher: Amazonas: Road Chachapoyas-
Mendoza, a little past Molinopampa, 0614
11S, 773549W, 2400 m, 15 Mar 1998,
van der Werff et al. 15038 (MO, UC, USM).
Reference: Lehnert, 2006a.
Cyathea palaciosii R. C. Moran
Dept. Amazonas. 1950 m. (Ecuador, Peru)
Voucher: Amazonas: San Martin, 0541S,
7748W, 1950 m, 4 Mar 2001, van der Werff
et al. 16721 (MO, UC).
Reference: Moran, 1995.
Cyathea parvifolia Sodiro
Dept. Pasco. 28002900 m. (Colombia,
Ecuador, Peru, Bolivia)
Voucher: Pasco: Oxapampa, Dist. Huanca-
bamba, Lanturachi, sector Santa Barbara,
road to Milpo, 1022S, 7536W, 2824 m,
10 Oct 2003, Perea et al. 721 (MO).
2011] LEHNERT: CYATHEACEAE OF PERU
References: Moran, 1995; Lehnert, 2008,
2009a.
Remarks: Synonym Cyathea brevistipes R.
C. Moran
Cyathea patens H. Karst.
Dept. Amazonas, Cajamarca, Cuzco.
(1900)27003300 m. (Colombia, Venezuela,
Ecuador, Peru)
Voucher: Cuzco: La Convencin, Dist.
Vilcabamba, Espiritupampa, 1303S, 7305W,
3300 m, 15 Oct 2003, Suclli et al. 1330 (MO).
Reference: Lehnert, 2009a.
Remarks: The earlier homonym Cyathea
patens hort., Houlst. & Moore is an invalidly
published nomen nudum. The later use of this
epithet is not illegitimate (Tryon, 1976;
Lehnert, 2009a).
Cyathea peladensis (Hieron.) Domin
Depts. Cajamarca. 1900 m. (Colombia,
Ecuador, Peru)
Voucher: Cajamarca: San Ignacio, Dist.
San Jos de Lourdes, Buenos Aires (Cerro El
Parco), 054204S, 775306W, 1900 m,
Feb 2006, Bonino 238 (MO, UC).
References: Tryon, 1976, 1986.
Cyathea phegopteroides (Hook.) Domin
Depts. Cajamarca, San Martin. 1100
1800 m. (Peru)
Voucher: San Martin: Peruvia occidenta-
lis, Tarapoto, on the top of Mt. Guayrapur-
ina, Spruce 4020 (holotype: K; isotype: B).
References: Tryon, 1976. 1986.
Cyathea pilosissima (Baker) Domin
Depts. Amazonas, Loreto, Pasco, San
Martin. 3501500 m. (Colombia, Venezuela,
Ecuador, Peru)
Voucher: San Martin: Monte Campana,
Spruce 4322 (holotype: K-photo GH; isotype:
BR-photos GH, US).
Reference: Tryon & Stolze, 1989.
Cyathea plicata Lehnert
Depts. Amazonas, Cajamarca. 2100
2700 m. (Ecuador, Peru)
Voucher: Amazonas: Chachapoyas, Carre-
tera Chachapoyas-Mendoza, Km 52 de Cha-
chapoyas, 10 km detrs de Molinopampa, 06
27
14.26S, 7735.96W, 2700 m, 4 Aug 2002,
Lehnert 240 (GOET, UC, USM)
References: Lehnert, 2006a.
Cyathea polliculi Lehnert, sp. nov.
Dept. Pasco. 2450 m. (Ecuador, Peru)
Voucher: Pasco: Oxapampa, near antenna
past Chacos, 1038S, 7515W, 2450 m, 19
Jul 2003, van der Werff et al. 18733 (MO,
UC).
Cyathea poeppigii (Hook.) Domin
Depts. Amazonas, Cajamarca, Huanuco,
Junin, Pasco, San Martin. 3001500 m.
(Costa Rica, Panama, Colombia, Venezuela,
Ecuador, Peru, Brazil, Bolivia)
Voucher: Cajamarca: Prov. Bagua, entre
Bagua Grande y Chiriaco, 0522.08S, 78
27.15W, 400 m, 20 Jul 2002, Lehnert 197
(GOET, UC, USM)
References: Windisch, 1977; Tryon &
Stolze, 1989.
Cyathea praetermissa Lehnert, sp. nov.
Dept. Amazonas. 2450 m. (Peru)
Voucher: Amazonas: A few Km from
Molinopampa, [ca. 1038S, 7515W,
2450 m,] 14 Mar 1998, van der Werff et al.
14935 (holotype: UC; isotypes: F, MO,
USM).
Cyathea pungens (Willd.) Domin
Depts. Amazonas, Cuzco, Huanuco, Lor-
eto, Madre de Dios, Pasco, Puno, San Martin,
Ucayali. 1001700 m. (Trinidad, Colombia,
Venezuela, the Guyanas, Ecuador, Peru, Bra-
zil, Bolivia)
Voucher: San Martin: Rioja, along road
Rioja-Pedro Ruiz, 054027S, 774035W,
1170 m, 23 Mar 1998, van der Werff et
al.15511 (MO, UC).
Reference: Tryon & Stolze, 1989.
Cyathea rufescens (Mett. ex. Kuhn) Domin
Dept. San Martin. Ca. 1400 m. (Peru)
Voucher: San Martin: Tarapoto, Spruce
4212 (holotype: K).
References: Windisch, 1978; Tryon &
Stolze, 1989.
28 BRITTONIA
Cyathea ruiziana Klotzsch
Depts. Cuzco, Huanuco, Junin, Pasco.
20003000 m. (Peru, Bolivia)
Voucher: Huanuco [Panhatahuas]: Ruiz
72 (180) (holotype: B).
References: Tryon, 1976; Lehnert, 2008.
Remarks: Synonym Cyathea boliviana R. M.
Tryon.
Cyathea serpens (R. M. Tryon) Lehnert
Dept. Cuzco, Pasco. 24502800 m. (Peru)
Voucher: Cuzco: La Convencin, Cordil-
lera Vilcabamba, 10 Jul 1968, Dudley 10949
(holotype: NA n.v.; isotype: GH).
References: Tryon & Stolze, 1989; Lehnert,
2009b.
Cyathea squamipes (H. Karst.) Domin
Depts. Amazonas, Cajamarca, Cuzco, Hua-
nuco, Pasco, San Martin. 11502900 m.
(Colombia, Venezuela, Ecuador, Peru, Bolivia)
Voucher: Amazonas: Laguna de Pomaco-
chas, 2550 m, 28 Mar 2002, van der Werff et
al. 15796 (MO, UC, USM).
Reference: Tryon & Stolze, 1989; Lehnert,
2009a.
Remarks: Synonym Cyathea caracasana
var. boliviensis (Rosenst.) R. M. Tryon.
Cyathea straminea H. Karst.
Dept. Amazonas, Pasco, San Martin. 3110
3400 m. (Colombia, Venezuela, Ecuador,
Peru, Bolivia)
Voucher: Pasco: Oxapampa, Dist. Huanca-
bamba, P. N. Yanachaga-Chemilln, 1022
32S, 752804W, 3110 m, 6 Apr 2004,
Monteagudo et al. 6715 (MO).
References: Tryon & Stolze, 1989; Lehnert,
2008.
Cyathea subincisa (Kunze) Domin
Depts. Amazonas, Huanuco, Madre de
Dios, Pasco, San Martin, Ucayali. 300
1515 m. (Ecuador, Peru, Bolivia)
Voucher: Pasco: Oxapampa, Dist. Palcaz,
sector Paujil, near the colpa, 101924S, 75
1549W, 380 m, 26 Feb 2004, Mellado 2814
(MO).
[VOL 63
References: Stolze, 1974; Tryon & Stolze,
1989.
Remarks: Synonym Cnemidaria speciosa
C. Presl.
Cyathea thelypteroides A. R. Sm.
Dept. Amazonas. 10301200 m. (Peru)
Voucher: Amazonas: Bagua, upper slopes
and summit of Cerro Tayu, 051556S, 78
2207W, 1030 m, 22 Mar 2001, van der
Werff et al. 16323 (holotype: USM; isotypes:
MO, UC).
References: Smith et al., 2005, Smith,
2006.
Cyathea tortuosa R. C. Moran
Depts. Amazonas, Junin, Loreto, Madre de
Dios, Pasco, Puno, San Martin. 1001000 m.
(Colombia, Ecuador, Peru)
Voucher: Puno: Ro Candamo, la at
mouth of Ro Guacamayo, 1330S, 6950
W, 810 m, 25 May 1992, Gentry et al. 77189
(MO, UC).
References: Moran, 1991; Smith et al.,
2006.
Cyathea traillii (Baker) Domin
Depts. Amazonas, Loreto, Pasco. 100
800 m. (Costa Rica, Panama, Colombia,
Venezuela, the Guyanas, Ecuador, Peru, Brazil)
Voucher: Amazonas: Bagua, Cerro Tayu,
ca. 1 h from Chiriaco, 051556S, 782207
W, 19 Mar 2001, 800 m, van der Werff et al.
16210 (MO, UC).
References: Windisch, 1978; Tryon &
Stolze, 1989.
Remarks: Synonym Cyathea reginae
(Windisch) A. R. Sm. ex Lellinger.
Cyathea tungurahuae Sodiro
Dept. Cuzco. 2200 m. (Colombia, Vene-
zuela, Ecuador, Peru)
Voucher: Cuzco: La Convencin, Dist.
Santa Ana, Tunquimayo, 125431S 7248
45W, 2200 m, 27 Sep 2004, Calatayud et al.
2893 (MO).
Reference: Lehnert, 2008.
Remarks: First record for Peru.
2011] LEHNERT: CYATHEACEAE OF PERU
Cyathea uleana (A. Samp.) Lehnert,
comb. nov.
Dept. Cuzco, Pasco. 11002240 m. (Ecua-
dor, Peru, Brazil, Bolivia)
Voucher: Pasco: Oxapampa, La Suiza
Nueva, 1038S, 7527W, 2240 m, 21 Jun
2003, van der Werff et al. 17546 (MO).
References: Stolze, 1974; Tryon & Stolze,
1989.
Remarks: Synonym Cnemidaria uleana
(A. Samp.) R. M. Tryon var. uleana. Stolze
(1974) further distinguished the var. abita-
guensis (Domin) Stolze from Colombia and
Ecuador.
Cyathea ulei (H. Christ) Domin
Depts. Cuzco, Hunuco, Pasco, Puno, San
Martn. 10301300 m. (Colombia, Venezuela,
Ecuador, Peru, Bolivia)
Voucher: San Martn: Prov. Picota, Cerro de
Ponasa, 1300 m, Mar 1903, Ule 6901 (hol-
otype: P?; isotypes: B-photo GH, L-photo US).
References: Barrington, 1978; Tryon &
Stolze, 1989.
Cyathea vilhelmii Domin
Dept. Puno. Ca. 2000 m. (Peru)
Voucher: Puno: Prov. Carabaya, Tatanara,
Ost-Abhang der Cordillera von Peru, Aug
1854, Lechler 2654 (lectotype: B).
Reference: Tryon & Stolze, 1989.
Remarks: Probably conspecic with C.
heliophila R. M. Tryon.
Cyathea werfi R. C. Moran
Depts. Loreto, Pasco. 120890 m. (Ecua-
dor, Peru)
Voucher: Loreto: Iquitos, 120 m, 10 Apr
1934, Williams 8236 (F).
References: Moran, 1991.
Cyathea windischiana A. R. Sm.
Dept. Amazonas. 800 m. (Peru)
Voucher: Amazonas: Distr. Bagua, Cerro
Tayu, ca. 1 hour from Chiriaco, 051556S,
782207W, 800 m, van der Werff et al.
16207 (holotype: UC; isotype: MO)
References: Smith, 2006.
29
Cyathea xenoxyla Lehnert
Depts. Amazonas, Cuzco, Pasco, San Mar-
tin, Ucayali. 10002770 m. (Colombia, Ecua-
dor, Peru, Bolivia).
Voucher: Amazonas: Chachapoyas, Carre-
tera Chachapoyas-Mendoza, Km 52 de Cha-
chapoyas, 10 km detrs de Molinopampa, 06
14.26S, 7735.96W, 2700 m, 4 Aug 2002,
Lehnert 229 (GOET, UC, USM).
References: Lehnert, 2003; Smith et al.
2005.
Sphaeropteris Bernh.
Sphaeropteris quindiuensis (H. Karst.)
R.M. Tryon
Depts. Amazonas, Cajamarca, Cuzco, Hu-
nuco, Junin, Pasco, San Martn. (1700)
19702400(2700)m. (Colombia, Ecuador,
Peru, Bolivia)
Voucher: Pasco: Oxapampa, Dist. Oxa-
pampa, sector quebrada San Alberto, 1032
46S, 752125W, 2400 m, 21 Aug 2004,
Mellado & Francis 1543 (MO).
References: Tryon, 1971; Tryon & Stolze,
1989.
NEW SPECIES
Alsophila gastonyi Lehnert, sp. nov. Type:
PERU. Pasco: Prov. Oxapampa, Dist. Pal-
caz, Reserva Comunal Yanesha, Comuni-
dad Nativa San Pedro de Pichanaz, Sector
Azulis, 102644S, 750621W, 910 m,
15 Sep 2005, A. Monteagudo, F. Pea A. &
E. Quintuya 9810 (holotype: MO). (Fig.1)
Ab Alsophila erinacea (H. Karst.) D. S. Conant et
Alsophila cuspidata (Kunze) D. S. Conant squamis
petiolorum lanceolatis atrobrunneis costisque abaxia-
liter abundante et persistente pilosis (vs. squamis
linealo-lanceolatis pallide brunneis costisque abaxia-
liter glabrescentibus vel parce pilosis), ab Alsophila
engelii R. M. Tryon et Alsophila minervae Lehnert in
presentia pilorum persistentium indusiis (vs. absentia
pilorum persistentium indusiis) absentiaque alarum in
parte distale pinnarum differt.
Trunks to 4 m high, 4 cm diam., otherwise
unknown. Fronds of unknown dimensions,
presumably 23 m long, patent to arching.
Petioles 2756 cm long, basally dark brown
to blackish, distally paler brown to dark
stramineous, with persistent scales and scurf
30 BRITTONIA [VOL 63

FIG. 1. Alsophila gastonyi. A. Petiole scale. B. Petiolar scurf squamules. C. Laminar apex, abruptly reduced,
conform. D. Medial pinnule of central pinna, adaxially. E. Fertile segment, abaxially, showing hairy costa and
subsphaeropteroid indusia. (AC from Monteagudo A. et al. 9810, MO; DE from Monteagudo A. et al. 9668, MO.)

of appressed lacerate squamules; aphlebiae or
skeletonized basal pinnae absent. Petiole scales
to 7 x 2 mm, lanceolate to ovate-lanceolate,
with differentiated margins of differently ori-
entated cells, dark brown with somewhat paler
margins with only one dark apical setae to
1 mm, no or few marginal setae, concolorous
(Fig. 1A), of variable texture, larger scales
papery, smaller ones rm, intergrading with
scurf squamules (Fig. 1B) and marginate
squaminate spines to 6 mm long, black, with
broad bases and acuminate, curved tips. Lam-
inae to ca. 130 cm broad, broadly ovate,
bipinnate-pinnatid, opaque dark green adax-
ially, opaque pale green abaxially, apices
abruptly reduced, non-conform to conform,
i.e., pinna-like (Fig. 1C). Rachises and costae
brown to dark stramineous, inermous, abaxially
2011] LEHNERT: CYATHEACEAE OF PERU
with fugacious attish brown squamules to
0.5 mm long, white pluricellular hairs to
1.5 mm long at least on distal parts, adaxially
more abundant than abaxially, insertions of
costae into rachises slightly raised abaxially,
bearing a lunular, at to concave blackish
pneumathode to 52.5 mm. Pinnae to 65 cm
long, stalked to 1.5 cm, not green-alate distally;
basal ones reduced, more ore less patent.
Largest pinnules to 10.51.8 cm, sessile,
truncate basally, attenuate apically (Fig. 1D);
costules stramineous to tan abaxially and
adaxially, with at brown, setate scales to
1 mm long, no bullate scales present, with
white to tan pluricellular hairs to 1.5 mm long
abaxially and adaxially, abaxially denser and
less spreading than abaxially (Fig. 1E); seg-
ments to 102.5 mm, adnate, sessile, weakly
ascending, straight (Fig. 1E) to weakly falcate,
the margins subentire to weakly crenate basally
in the proximal segments, at in sterile and
fertile segments; midveins weakly protruding
adaxially and adaxially, with white pluricellular
hairs identical to those on the costules (Fig. 1E),
fertile veins forked once, rarely twice, sterile
ones forked to simple (Fig. 1E), hardly discern-
able adaxially, dark gray abaxially, glabrous on
both sides (Fig. 1E). Sori to 0.8 mm diam., 13
per segment, costal, in vein forks (Fig. 1E);
indusia urceolate to subsphaeropteroid, either
irregularly or not rupturing at maturity, rm,
lustrous dark brown to tan, with persistent
pluricellular hairs to 0.6 mm long (Fig. 1E),
receptacles 0.20.3 mm diam., columellar,
black, paraphyses much shorter than the spor-
angia, 0.10.2 mm long, blackish to tan. Spores
not examined.
Distribution and habitat.Known only
from Peru, growing in the selva central
region in the Dept. Pasco and the Ro Ene
valley in the Dept. Cuzco at 520950 m.
Etymology.The name honors Gerald J.
Gastony in acknowledgment of his contribu-
tions to pteridology.
Additional specimens examined. PERU. Cuzco: La
Convencin, Echarati, Kapiromashi, 750 m, 1630 Apr
2004, Salinas N. et al. 6743 (USM). Pasco: Oxapampa,
Dist. Palcaz, San Pedro de Pichanaz-Azulis, Reserva
Comunal Yanesha, 102840S, 750610W, 670 m, 25 localidad Romerillo, Cordillera entre
Feb 2004, Mellado L. F. 711 (MO), ibid., 25 Feb 2004,
Mellado L. F. 721 (MO); Dist. Palcaz, Reserva
Comunal Yanesha, Comunidad Nativa San Pedro de
Pichanaz, Sector Azulis, 102644S, 750621W, Rodrguez E. & S. Nuez 4294 (holotype:
910 m, 12 Sep 2005, Monteagudo A. et al. 9668 (MO,
31
USM); ibid., 102934S, 750630W, 520640 m, 23
Sep 2005, Monteagudo A. et al. 10127 (MO); Dist.
Palcaz, Reserva Comunal Yanesha, Comunidad Nativa
Lomalinda-Laguna, Sector Nueva Aldea, 102338S,
750558W, 920950 m, 12 Oct 2005, Monteagudo A.
et al. 10400 (MO, USM).
The trunk morphology was not described on
the specimen labels but presumably is like in
the majority of Neotropical Alsophila species,
i.e., densely covered with thin lanceolate scales
and without persistent old petiole bases.
Among its Andean congeners, the species
is similar and supposedly related to Alsophila
engelii R. M. Tryon and A. minervae Lehnert
but differs from these in having long, rela-
tively straight, presistent hairs on both sides
of the midveins and on the indusia. The other
two species have no hairs on the indusia and
on the midveins adaxially, but may have hairs
or trichomidia on the lateral veins abaxially
where A. gastonyi is glabrous; Alsophila
engelii has appressed tortuous hairs and thin
squamules on the leaf axes abaxially but no
spreading straight hairs as they occur in the
same places in A. gastonyi in varying density.
Further, A. gastonyi commonly has abruptly
reduced (but usually not pinna-like) laminar
apices and occurs below 1000 m, whereas A.
engelii and A. minervae have gradually reduced
laminar apices and occur mainly above 1500 m.
Using the general key to the genus (Conant
1983), Alsophila gastonyi will key out to A.
portoricensis (Spreng. ex Kuhn) D. S. Con-
ant, a species restricted to Puerto Rico, and
both populations may well be united under
one species morphologically. Such a biogeog-
rapical pattern would be unequalled among
Neotropical ferns. Other tree ferns that span
such distances do so in a more or less
continuous population (e.g., C. microdonta
and C. delgadii). Either this Alsophila species
has been left undetected in most of its range,
or, a hypothesis favored here, A. gastonyi and
A. portoricensis are an example of convergent
evolution.
Cyathea chontilla Lehnert, sp. nov. Type:
PERU. Cajamarca: San Ignacio, Huarango,
Romerillo y Nuevo Mundo, 0514S, 78
46W, 2300 m, 29 Jul 1992, J. Campos,
UC; isotypes: MO, USM). (Fig. 2)
32 BRITTONIA [VOL 63

FIG. 2. Cyathea chontilla. A. Habit, lateral shoot of many-crowned plant. B. Petiole scale. C. Fertile proximal
pinnule of central pinna adaxially, large, from main shoot. D. Fertile proxial pinnule of central pinna abaxially, small,
from lateral shoot. (From Campos et al. 4294; A from UC; BE from MO.)
2011] LEHNERT: CYATHEACEAE OF PERU
Species pygmaea generis Cyatheae squamis anguste
lanceolatis cupreis, laminis triangularibus gradatim
reductis, rhachidibus sparse vel abundante squamatis,
indusiisque sphaeropteroideis. A Cyathea patente H.
Karst. statura minore, a Cyathea sylvatica Lehnert
laminis subcoriaceis (vs. chartaceis), squamis petiolorum
majoribus (usque 10.02.5 mm vs. 12.020.02.0
3.5 mm) differt.
Trunks absent, rhizomes short-ascending,
branching in posterior parts; apices hidden in
fascicles of petioles (Fig. 2A). Fronds 45
110 cm long, erect (Fig. 2A). Petioles 15
36 cm long, dark-brown to blackish, weakly
shiny, inermous; scurf brown, dense, consisting
of variously sized, erect to appressed, ovate to
lanceolate squamules with mbriate margins
and brown tortuous hairs to 1 mm long; without
pneumathodes; densely scaly proximally, mod-
erately to sparsely so in distal parts. Petiole
scales to 12.020.02.03.5 mm, lanceolate to
narrowly lanceolate (Fg. 2B), shiny, concolo-
rous or weakly bicolorous, deep brown with
margins yellowish to cream white, the tips
elongate, undulate, not twisted, differentiated
margins narrow with cells strongly exerted,
shortly dentate, without setae or cilia. Laminae
to 30852535 cm, bipinnate to bipinnate-
pinnatid, long-triangular to lanceolate
(Fig. 2A), coriaceous, dark green adaxially,
black when dried, gray-green abaxially, apices
gradually reduced, tapering. Rachises inermous,
dark brown to castaneous, densely hairy on both
sides but glabrescent, with brown, twisted,
pluricellular hairs to 1.0 mm long, also with
scattered, lanceolate to linear scales to 4 mm
long. Pinnae to 17 cm long, weakly ascending,
stalked to 5 mm, mostly alternate (Fig. 2A),
fully pinnate only in proximal halves, narrowly
green alate distally; basal pinnae about the size
of the medial pinnae, not reexed, sometimes
opposite. Costae dark-brown to castaneous,
inermous, adaxially densely hairy with whitish
to tan, antrorsely curved, pluricellular, uniseri-
ate hairs to 1.0 mm long, abaxially glabrescent
with sparser twisted hairs and few concolorous
brown, lanceolate scales with whitish, shortly
mbriate margins, most frequently in the axils;
junctures of rachises and costules not swollen,
each abaxially with a circular, prominent
pneumathode to 1 mm diam., dark gray to black
in dried specimens. Largest pinnules/ultimate
segments (10-)20.040.0(4.0)8.014.0 mm,
33
oblong to oblanceolate, subsessile (Fig. 2C) to
sessile (Fig. 2D), ca. 0.7 cm between the
midveins, cuneate to weakly cordate at base,
tips obtuse to rounded, patent, weakly ascend-
ing, straight to distally falcate, margins shal-
lowly to deeply crenate with entire lobes, often
revolute when dried; segments with few whitish
to tan, erect hairs 0.50.8(1.0) mm long on the
veins abaxially, restricted to the midveins/
costules adaxially, none between the veins; with
few to many dark-brown scales on the mid-
veins/costules abaxially, ovate-lanceolate, at
or bullate, to 2.00.8 mm with weakly dentate-
lacerate margins; veins weakly protruding
abaxially, planar or sunken adaxially, sterile
veins forked or simple, fertile veins forked. Sori
to 1.01.2 mm diam., subcostal to inframedial,
on the back of vein (Fig. 2D); indusia sphaer-
opteroid, with prominent umbo, shiny dark
brown, translucent; receptacles globose, 0.3
0.4 mm diam, paraphyses thin, hyaline, tan to
brown, of the same length as the sporangia
(0.4 mm). Spores ca. 0.04 mm diam., exospore
verrucate, perispore lacking (Young 5236,
USM).
Distribution and habitat.Known only
from Peru, Depts. Cajamarca and Pasco, grow-
ing in ridge-top vegetation at 23002850 m.
Etymology.The name is a diminuitive of
chonta, a vernacular name for tree ferns in
northern Peru, referring to the small size of
the plant.
Additional specimens examined. PERU. Pasco:
Prov. Oxapampa, Dist. Oxapampa, Chacos (indumen
telefonica), 1037S, 7517S, 2850 m, 21 Sep 2003,
Rojas R. et al. 1487 (MO, UC); Cerro Pajonal, SE of
Oxapampa, 2800 m, 29 Jul 1996, Young K. R. 5232,
5236 (USM).
Cyathea chontilla probably belongs to the
C. patens group, which is characterized by
mbriate laminar scales and scurf containing
tortuous hairs. This group of sphaeropteroid-
indusiate tree ferns is typical of the upper
mountain rain forests from Costa Rica
through the Andes, and includes other small
species. Cyathea parvifolia Sodiro (Lehnert,
2009a) can be distinguished from C. chontilla
by having longer petiole scales, stronger
dissected pinnules, and the regular presence
of adventitious pinnae at the petiole bases.
Cyathea sylvatica Lehnert can be distin-
34 BRITTONIA
guished from C. chontilla by less indument
abaxially on the laminae and smaller, con-
colorous petiole scales. Cyathea chontilla has
also more hairs abaxially on the veins than
adaxially (vice versa in C. sylvestris) and
fringed margins on the laminar squamules
(vs. margins mostly entire).
Omitting the presence of tortuous hairs,
the similarity of Cyathea chontilla to C.
lechleri becomes more evident. Most speci-
mens of C. lechleri represent plants 4 m or
smaller, which characteristically lack hairs
on the veins and laminar squamules almost
completely. Larger fertile plants (to 12 m)
may have a similar amount of pluricellular
hairs and lanceolate, at to weakly bullate
castanous squamules as found in C. chon-
tilla. Both species are an example for the
often-observed elevational zonation in spe-
cies groups of Cyatheaceae: Cyathea chon-
tilla is clearly a high-elevation counterpart
to C. lechleri, which occurs mainly below
1500 m.
The examined spores (Young 5236, USM;
comm. Blanca Len) lacked a perispore layer,
which may be a species-dening character but
which also could reect just an early devel-
opmental stage. The overall spore morphology
of Cyathea chontilla agrees well with C.
aterrima (Hook.) Domin (Moran et al., 2008).
Cyathea nephele Lehnert, sp. nov. Type: PERU.
Pasco: Prov. Oxapampa, Distrito Huancabamba,
Parque Nacional Yanachaga-Chemilln, 1022S,
7528W, 3110 m, 5 Dec 2003, A. Monteagudo,
J. Perea, C. Mateo, G. Ortiz, & R. Francis 6430
(holotype: UC; isotype: MO). (Fig. 3)
A Cyathea frigida (H. Karst.) Domin
squamis petiolorum longioribus lanceolatis
brunneis pallidioribus bicoloribus sive con-
coloribus (vs. squamis ovato-leanceolatis
bicoloribus atro-brunneis usque castaneis
cum marginibus angustissimis pallide brun-
neis vel albidis), a C. patente H. Karst.
absentia indusiorum (vs. indusiis sphaeropter-
oideis) differt.
Trunks to 8 m tall, 810 cm diam., without
old petiole bases, frond scars round, apices
hidden in fascicles of petioles; adventitious
buds lacking. Fronds 24 m long, presum-
ably arching. Petioles to 40 cm long, prob-
ably longer, dull brown to orange-brown,
[VOL 63
basally blackish, aculeate to muricate, spines
with scales on their tips in young petioles;
with long elliptic brown pneumathodes to
181 mm on the sides basally; scurf persis-
tent (Fig. 3A), white to tan, akey, consisting
of spaced, appressed, round squamules to
0.5 mm diam. with tortuous marginal cilia.
Petiole scales 25.035.0(40.0) 2.54.0(
5.0) mm, long-lanceolate, shiny, plain deep
brown, concolorous or concordantly bicolo-
rous with margins paler brown, sometimes
yellowish or cream white (Fig. 3A); scales
persistent only on the sides of the petioles.
Laminae to 1607080 cm, bipinnate-pinna-
tid, green to dark green adaxially, often
blackish when dried, pale gray-green abax-
ially, apices gradually reduced. Rachises
inermous or weakly muricate basally, dark
yellowish brown to orange-brown, adaxially
with whitish to yellowish, antrorsely curved
hairs to 0.8 mm long, abaxially glabrous or
glabrescent with white to tan squamellae like
in the petiole scurf. Pinnae to 45 cm long
(Fig. 3B), patent to ascending, basal pinnules
only half the size of the medial pinnae,
weakly to strongly reexed. Costae and
costules tan, stramineous or dull orange-
brown, inermous, short-hairy adaxially, hairs
to 1 mm long, white, antrorsely curved;
abaxially glabrescent with white to tan arach-
noid scurf and few bright orange-brown
attish scales with undulated, mbriate mar-
gins, most frequently in the axils. Largest
pinnules 5.06.51.11.7 cm, linear-oblong
to lanceolate, sessile (Fig. 3C) to subsessile
(stalked to 1 mm), 1.01.5 cm between the
stalks, pinnule bases truncate to weakly
cuneate, rarely weakly cordate, tips long-
acute to attenuate, pinnatid to pinnatisect,
basal segments sometimes remote but not
free; segments to 113 mm, weakly ascend-
ing, falcate, the tips obtuse (Fig. 3D) to
round, segment margins crenulate to crenate,
often remaining at when dried; segments
with few to many whitish to tan, erect hairs
0.50.8(1.0) mm long on and between the
veins abaxially, veins adaxially glabrous or
with few short hairs, none between the veins;
with relatively few to many small scales
abaxially, ranging from broadly ovate, at
ones (to 3.02.5 mm) to bullate ones (to
2.51.0 mm) with broadly aring or twisted,
elongated tips, tan to orange-brown, often
2011] LEHNERT: CYATHEACEAE OF PERU 35

FIG. 3. Cyathea nephele. A. Detail of petiole base, showing small corticinate spines and granular scurf. B. Pinna.
C. Medial pinnule of medial pinna, abaxially. D. Fertile segment, abaxially. (From Len 2141, UC.)

with the thin margins paler brown to whitish,
mbriate to lacerate with considerably long
processes; sterile veins forked or simple,
fertile veins forked. Sori to 1.0 mm diam.,
subcostal (Fig. 3D), in forks of veins; indusia
absent; receptacles globose to ellipsoid, 0.3
0.4 mm diam, paraphyses thin, hyaline, white
to tan, of the same length as or shorter than
sporangia (0.30.4 mm). Spores not examined.
Distribution and habitat.Frequent in
central Peru, Dept. Pasco, rare in the north,
Prov. Amazonas and San Martin, in upper
montane forest and ridge top dwarf forests at
27203550 m.
Etymology.Named after the Greek god-
dess of the clouds (Greek, nephos=cloud) in
reference to the cloud forests this species
inhabits.
36 BRITTONIA
Additional specimens examined. PERU. Amazonas:
20 km south of Leimatambo [Leymebamba?], 3550 m, 13
Feb 1973, Madison 1144 (GH); Prov. Chachapoyas,
Distrito Leymebamba, road to Celendn, between Ishpingo
and Pomacocha, 28003000 m, 20 Aug 1998, Quipuscoa
et al. 1390 (UC). Pasco: Prov. Oxapampa, Dist. Huanca-
bamba, sector Santa Barbara-Milpo, 102258S, 753723 033726S, 782335W, 2010 m, 16 Dec
W, 2900 m, 1 Feb 2005, Mellado L. F. et al. 2751, 2779
(MO); P.N. Yanachaga-Chemilln, 102232S, 752804
W, 3110 m, 6 Apr 2004, Monteagudo et al. 6687, 6658,
6716 (MO); between Ro Cueva Blanca and Milpo, 1022
33S, 753648W, 2720 m, 18 Sep 2004, Monteagudo A.
et al. 7071 (MO). San Martin: Prov. Mariscal Cceres,
Ro Abiseo National Park, 30003100 m, 13 Jul 1988,
Len B. 2141 (UC, USM).
Most specimens of Cyathea nephele were
previously mistaken for C. frigida (H. Karst.)
Domin, but these species are not considered
here to be closely related. The most striking
difference is the overall appearance of the
specimens, which is summed up from rela-
tively minor differences. Petioles of C. neph-
ele are spiny or muricate whereas they are
generally inermous in C. frigida. The petiole
scales of C. nephele are narrower and paler in
color than in C. frigida (concolorous brown or
bicolorous with paler brown to whitish mar-
gins vs. dark brown to castaneous with white
margins); the paraphyses in C. nephele are not
longer than the sporangia while they may be
much longer than the sporangia in C. frigida;
the segments of C. frigida are gibbose with
revolute margins whereas they are at in C.
nephele, which makes the crenulate margins of
the latter species notable in dried specimens.
Cyathea frigida is very variable in the frond
size, with smaller fronds having stify straight
pinnae while they are characteristically curved
and exuous in larger plants. In C. nephele, the
pinnae have always straight, distally weakly
curved costae. Furthermore, the fronds of C.
frigida often have few long persisting ovate
scales with dentate margins, remaining from
the crozier stage on rachises and costae; if
larger scales are present in C. nephele, they
vary from ovate to long lanceolate or linear
and have mbriate and ciliate margins.
Generally, Cyathea nephele matches C.
patens H. Karst. and allies in the ne indu-
ment of tortuous hairs and ciliate squamules
on axes and laminae; also the petiole scales
clearly align it to C. patens, which is easily
distinguished by its sphaeropteroid indusia
(vs. indusia lacking in C. nephele).
[VOL 63
Cyathea polliculi Lehnert, sp. nov. Type:
ECUADOR. Zamora-Chinchipe: El Pan-
gui, Cordillera del Condor, 2 km S of
Condor Mirador Military post (observa-
torio #1), deep gorge of Ro Tumdayme,
2000, G. Pabn, J. Caranqui & Grupo
Post-Grado MO-QCNE 359 (holotype:
UC; isotype: MO). (Fig. 4)
Species generis Cyatheae squamis anguste lanceo-
latis brunneis conconcoloribus vel bicoloribus cum
margine isabellina vel albida, laminis bipinnatis
lanceolatis, indusiisque hemitelioideis. A Cyathea
bipinnata (R. M. Tryon) R. C. Moran statura minore
squamisque petiolorum atrocioribus (bicoloribus
brunneis cum marginibus albicantibus vs. squamis
usque concoloribus albidis in Cyathea bipinnata),
apicibus laminarum gradatim reductis (vs. abrupte
reductis) differt.
Trunks absent; apices hidden in fascicles of
petioles; adventitious buds lacking. Fronds to
0.7 m long (Fig. 4A), erect. Petioles to 20 cm
long, dull dark brown to blackish, matte,
sparsely muricate; scurf absent; with elliptic,
brown pneumathodes to 51 mm on the sides
basally; scaly only in lower third. Petiole
scales to 10.0 2.53.5 mm, lanceolate
(Fig. 4B), weakly shiny, concolorous auburn
to deep brown to discordantly bicolorous with
margins yellowish to cream-white; scales
persistent only on the sides of the petioles.
Laminae to 5020 cm, pinnate-pinnatid to
bipinnate, long triangular to lanceolate, dark
green adaxially, black when dried, gray-green
abaxially, apices gradually reduced, long-
tapering (Fig. 4A). Rachises inermous or
sparsely verrucate, dark purpureous to casta-
neous, with whitish to tan, antrorsely curved
or appressed, uniseriate hairs, adaxially to
1.5 mm long, abaxially 0.50.8(1.0) mm
long. Pinnae to 12 cm long, weakly ascend-
ing, fully pinnate only in proximal half,
narrowly green alate (Fig. 4A); basal pinnae
about the size of the medial pinnae, not
reexed. Costae dark purpureous to casta-
neous, inermous, adaxially densely pubescent
with whitish to tan, antrorsely curved, uni-
seriate hairs 1.01.5 mm long, abaxially
glabrescent with shorter hairs and few brown,
attish scales with whitish, shortly mbriate
margins (Fig. 4C), most frequently in the
2011] LEHNERT: CYATHEACEAE OF PERU 37

FIG. 4. Cyathea polliculi. A. Habit. B. Petiole scale. C. Fertile proximal pinnules of basal pinna abaxially,
showing hemitelioid indusia. D. Subbullate laminar squamule. (A, D from Pabn et al. 395, UC; BC from van der
Werff et al. 18733, MO.)
38
axils; junctures of rachises and costules not
swollen, each abaxially with a circular,
prominent pneumathode to 1 mm diam., dark
gray to black in dried specimens. Largest
pinnules/ultimate segments to 17.0 3.5
4.0 mm, linear-oblong to hastate, sessile to
subsessile (stalked to 0.5 mm), ca. 0.7 cm
between the midveins, patent, weakly ascend-
ing, straight to distally falcate, bases cuneate
to weakly cordate, tips obtuse to rounded,
segment margins crenulate (Fig. 4C), often
revolute when dried; segments almost gla-
brous or with few to many whitish to tan,
erect hairs 0.50.8(1.0) mm long on the
veins abaxially and adaxially, none between
the veins; with few brown scales on the
midveins abaxially, ovate-lanceolate, at, to
2.00.8 mm with weakly dentate-lacerate
margins (Fig. 4D); sterile veins forked or
simple, fertile veins forked (Fig. 4C). Sori
1.01.2 mm diam., subcostal, in vein forks;
indusia hemitelioid, reaching half way around
the receptacles (Fig. 4C), brown, translucent,
appressed, covered completely by sori; recep-
tacles globose to ellipsoid, 0.30.4 mm diam,
paraphyses thin, hyaline, tan to brown, of the
same length as the sporangia (0.4 mm).
Spores not examined.
Distribution and habitat.Ridge top for-
ests on poor sandstone derived soils and
subparamo at 20102450 m. Known from
southern Ecuador and central Peru; to be
expected at similar sites in northern Peru.
Etymology.The epithet refers to the rela-
tively small stature of the species, which is a
Tom Thumb, in its genus (Latin, polliculus =
small thumb or nger).
Additional specimen examined. PERU. Pasco.
Oxapampa, near antenna past Chacos, 1038S, 7515
W, 2450 m, 19 Jul 2003, van der Werff et al. 18733 (MO,
UC).
Cyathea polliculi may be confused with C.
chontilla, which bears a strong resemblence
in habit, size, and coloration. Despite the
similarity, a close relationship does not have
to exist. Additionally to the characters men-
tioned in the key, the laminar squamules of C.
polliculi resemble those of C. latevagans
more closely than those of C. chontilla (shiny
orange to auburn, ovate to almost round
squamules with strongly cordate to pseudo-
peltate bases vs. dark brown to castaneous,
BRITTONIA [VOL 63
lanceolate, never pseudopeltately attached in
C. chontilla) as does the color of rachises and
petioles (shiny black vs. comparatively matte
castaneous). Cyathea polliculi and C. lateva-
gans are a further example of evidently
closely related species with different indusial
status (hemitelioid vs. exindusiate).
Another species with a similar pinnule
shape is the Ecuadorian Cyathea bipinnata
(R. M. Tryon) R. C. Moran, which has
abruptly reduced frond apices (vs. gradullay
reduced in C. polliculi) and is restricted to the
western slopes of the Andes in the north of
the country (vs. known only from the eastern
Andean escarpment). Further distinguishing
characters of C. polliculi are the narrowly
lanceolate, dark brown, bicolorous petiole
scales with narrow whitish margins and often
with brown marginal teeth (vs. with broad
white margins or almost concolorous white,
without marginal teeth in C. bipinnata), and
few to many short hairs on the veins (vs.
glabrous or single hairs on the midveins).
The Peruvian collection of C. polliculi
differs from the Ecuadorean material in
having petiole scales that are more concolo-
rous and in largely lacking hairs on the veins
of the segments.
Cyathea praetermissa Lehnert, sp. nov.
Type: PERU. Amazonas: A few Km from
Molinopampa, [ca. 0614S, 7736W.
2700 m,] 14 Mar 1998, H. van der Werff,
R. Vasquez, B. Gray, R. Rojas, R. Ortiz &
N. Davila 14935 (holotype: UC; isotypes:
MO, USM n.v.). (Fig. 5)
Species generis Cyatheae acaulescens laminis trian-
gularibus vel anguste lanceolatis gradatim reductis
rhachidibusque etiam costis indumento cinereo obtec-
tis; a Cyathea serpens (R. M. Tryon) Lehnert, Cyathea
heliophila R. M. Tryon et Cyathea frigida (H. Karst.)
Domin indusiis sphaeropteroideis differt.
Trunks absent; rhizome apices hidden in
fascicles of petioles. Fronds to 150 cm long,
presumably arching or pendent from banks
(Fig. 5A). Petioles 40 cm long, probably
more (Fig. 5A), weakly shiny, castaneous to
atropurpureous, basally blackish, inermous,
without pneumathodes, only sparsely scaly in
lower half, with sparse scurf consisting of
appressed, strongly dissected squamules and
2011] LEHNERT: CYATHEACEAE OF PERU 39

FIG. 5. Cyathea praetermissa. A. Habit. B. Petiole scale. C. Fertile segment abaxially, fragmented indusia and
lacerate scales of indument indistinguishable. (From van der Werff et al. 14935, UC.)
40 BRITTONIA
tortuous hairs, apparently ephemeral. Petiole
scales to 18.0 2.53.0 mm, lanceolate,
almost concolorous, dark auburn to brown
with paler brown to yellowish margins, long
attenuate, bases round to cordate, pseudopel-
tately attached (Fig. 5B); differentiated mar-
gins narrow, with cells strongly exerted, with
teeth or short cilia to 0.5 mm long, without
setae. Laminae to 1506070 cm, ovate-
lanceolate (Fig. 5A), bipinnate-pinnatid,
green to dark green adaxially, blackish when
dried, pale gray-green abaxially, apices grad-
ually reduced, tapering. Rachises inermous,
dark castaneous to purpureous, densely hairy
with tan to brown, antrorsely curved, uni-
seriate hairs to 1.5 mm long adaxially, with
white, cobwebby, tortuous hairs to 2.0 mm
long abaxially, glabrescent. Pinnae to 14 cm
long, patent, alternate to subopposite, stalked
to 1.2 cm, distally not or very narrowly green
alate, the distal segments not decurrent; basal
pinnules nearly the size of the medial pinnae,
not reexed. Costae dark brown, inermous,
to 1.0(1.5) mm wide, adaxially with tan to
brown, antrorsely curved, uniseriate hairs to
1.5 mm long, abaxially glabrescent with
white scurf of tortuous hairs (Fig. 5C) and
few tan to brown, attish scales with undu-
late, mbriate to short-ciliate margins, most
frequent in the axils; insertions of costae and
rachises not or only abaxially weakly swol-
len, each with a planar, elliptic pneumathode
to 2.31.5 mm, blackened in dried material.
Largest pinnules to 3.81.0 cm, lanceolate,
sessile (Fig. 5A), 1.01.5 cm between the
costules, bases truncate to weakly rounded,
tips long-acute to attenuate, pinnatid to
pinnatisect, basal segments never remote;
costules adaxially prominent, ridged, with
antrorsely curved, white, pluricellular, uni-
seriate hairs to 1.0 mm, abaxially densely
covered with white, tortuous hairs to 2 mm
long and some tan to brown, subclathrate,
ovate-lanceolate scales to 2.03.0 1.5
2.0 mm with mbriate to ciliate margins;
segments to 9.012.02.55.0 mm, weakly
ascending, straight, the sinuses acute, to 0.5(
1.0) mm wide, segment tips obtuse to
rounded, segment margins crenulate, revo-
lute, the basiscopical margins of the lowest
segments not differently incised (Fig. 5C);
segments with few to many deciduous,
whitish, contorted, catenate hairs to 2.0 mm
[VOL 63
long, mainly abaxially and along the margins;
lateral veins planar on both sides or weakly
raised adaxially, blackish to carnose, with few
persistent, brown, erect hairs to 1.2 mm long,
not between the veins; sterile veins forked or
simple, fertile veins forked. Sori to 1.0 mm
diam., subcostal to costal, in vein forks,
yellowish brown at maturity; indusia sphaer-
opteroid, tan to dull brown, fragile, often
fragmented to a cup or disc; receptacles
globose, 0.30.4 mm diam, paraphyses thin,
hyaline, white to tan, of the same length as
the sporangia (0.4 mm). Spores not examined.
Distribution and habitat. In scrub with
Sphagnum on sandy soil with rock outcrops
in northern Peru at ca. 2700 m. Known only
from the type collection.
Etymology.This species may often be
overlooked and not recognized as a tree fern
due its small stature (Latin, praetermissus=
overlooked, excluded).
Cyathea praetermissa may be confused
with small plants of C. frigida (H. Karst.)
Domin, a species with similar dense indument
of tortuous hairs and a comparable petiole
scale color, but C. frigida differs in lacking
indusia (vs. C. praetermissa with closed
sphaeropteroid indusia) and having larger
and broader scales. Like many other species
of Cyathea from similar localities, they have
both a reduced trunk. Cyathea frigida, how-
ever, may develop a massive trunk to 4 m tall
under more favorable conditions.
Cyathea nephele is sympatric with C. prae-
termissa and matches in most indument char-
acters, but it is much larger and lacks indusia.
REVISED SPECIES
Cyathea serpens (R. M. Tryon) Lehnert,
Phytotaxa 1: 50. 2009. Trichipteris serpens
R. M. Tryon, Fieldiana, Bot. n.s. 20: 126.
1989. Type: PERU. Cuzco: Prov. La Con-
vencin, Cordillera Vilcabamba, 7332S,
1237W, 2900 m, 10 Jul 1968, T. R.
Dudley 10949 (holotype: NA n.v.; isotype:
GH). (Fig. 6)
This species was described as being exin-
dusiate and consequently classied as a
species of the genus Trichipteris by Tryon
(Tryon & Stolze, 1989). The presence of
2011] LEHNERT: CYATHEACEAE OF PERU 41

FIG. 6. Cyathea serpens. A. Petiole scale. B. Pinna. C. Pinnule. D. Fertile segment, abaxially. E. Detail of
receptacle with indusium and paraphyses. (A from Dudley 10949, GH; BD from van der Werff et al. 18647, UC.)
42 BRITTONIA
indusia would have necessitated its trans-
ference to Cyathea as dened by Tryon (1970,
1976) despite the fact that both genera had
already been merged to Cyathea (Lellinger,
1987). The new combination Cyathea serpens
was already published in the comparison with
a newly described species (Lehnert, 2009b).
An amended description of C. serpens seems
appropriate in the context of its new taxo-
nomic status and the additional collections that
brought new information of about its morphol-
ogy and distribution.
Trunkless; rhizome apices hidden in fas-
cicles of petioles; adventitious buds not
reported and presumably lacking. Fronds to
7 m long, pendent from banks or trailing over
adjacent vegetation. Petioles 70200 cm
long, probably longer, shiny, castaneous to
atropurpureous, basally blackish, inermous,
with dense, matted, white scurf consisting of
appressed, strongly dissected squamules and
tortuous marginal hairs, often abraded in
older material; without pneumathodes, only
scatteredly scaly in lower half. Petiole scales
to 11.0(15.0)3.54.0(5.0) mm, lanceolate, to 1.2 mm long, not between the veins;
discordantly bicolorous, brown to dark brown
or blackish with white margins (Fig. 6A);
differentiated margins to 0.6 mm wide, cell
rows strongly exserted, without cilia or setae.
Laminae 15050070120 cm, long-triangu-
lar to lanceolate, bipinnate-pinnatid to partly
tripinnate, green to dark green adaxially, often
becoming blackish when dried, pale gray-
green abaxially, apices gradually reduced.
Rachises inermous, dark castaneous to atro-
purpureous, densely pubescent with whitish
to tan, cobwebby hairs to 2 mm long,
abaxially often glabrescent. Pinnae to 35 cm
long, patent (Fig. 6B), alternate to suboppo-
site, stalked to 2.53.0 cm, distally green
alate, the distal segments not decurrent into
the costae; basal pinnae nearly the size of the
medial pinnae, weakly reexed. Costae and
costules dark castaneous to atroprurpureous,
inermous, to 2 mm broad, adaxially with tan
to brown, antrorsely curved, pluricellular,
uniseriate hairs to 1.5 mm long, abaxially
glabrescent with white to tan scurf and few
bright orange-brown attish scales with undu-
lated, mbriate margins, most frequently in
the axils. Pinnules to 5.510.51.53.0 cm,
long triangular to lanceolate, stalked to 4 mm
(Fig. 6B, C), 1.52.0 cm between the stalks,
[VOL 63
bases truncate to weakly cordate, tips long-
acute to attenuate, pinnatid to pinnatisect,
basal segments sometimes remote, connected
by thin strands of laminar tissue; segments to
9.015.02.55.0 mm, weakly ascending,
straight to distally falcate, the sinuses acute
to obtuse, to 15 mm wide, the tips obtuse to
rounded, segment margins crenulate to
inciso-dentate, the basiscopical margins of
the lowest segments often more strongly
incised; costules planar to weakly protruding
abaxially, ridged adaxially, yellowish to dark
carnose, with many curved brown hairs to
1.2 mm long adaxially, glabrous abaxially,
with broadly ovate, at squamules to 3.0
2.5 mm, and bullate squamules to 2.5
1.0 mm, with mbrate tips, tan to orange-
brown, often with the thin margins paler
brown to whitish, mbriate to lacerate; seg-
ments with few to many deciduous, whitish,
contorted, catenate hairs to 2.0 mm long,
mainly abaxially and along the margins; veins
planar to weakly immersed, yellowish to
carnose, adaxially with few brown, erect hairs
abaxially glabrous or with few trichomidia;
sterile veins forked or simple, fertile veins
forked. Sori to 1.2 mm diam., subcostal to
costal, in fork of veins (Fig. 6C), yellowish
brown to dark orange-brown at maturity;
indusia hemitelioid, small, appressed, tan to
dull brown, with entire margins (Fig. 6D) or
with one or two marginal cilia, fragile and
evanescent; receptacles globose, 0.40.5 mm
diam. (Fig. 6D), paraphyses thin, hyaline, white
to tan, longer than sporangia (0.40.5 mm).
Spores tetrahedral-globose, dark orange-brown.
Distribution and habitat. Locally com-
mon in open forests and pajonal-vegetation in
central and southern Peru at 24502900 m.
Etymology. The epithet refers to the
trailing fronds.
Additional specimens examined. PERU. Pasco:
Prov. Oxapampa, Dist. Oxapampa, sector Chacos, 10
3725S, 751743W, 2750 m, 3 Feb 2004, Mellado et
al. 1664 (MO, UC); Dist. Huancabamba, sector Que-
brada Yanachaga (P. N. Yanachaga-Chemilln), 1022
46S, 752743W, 2910 m, 14 Feb 2004, Mellado &
Monteagudo 464 (MO, UC); Prov. Oxapampa, San
Gotardo, 36 km W of Oxapampa, 1037S, 7509W,
27102800 m, 26 Jan 1984, Smith D. N. & Canne 5902
(MO, UC); Prov. Oxapampa, near antenna past Chacos,
1038S, 7515W, 2450 m, 19 Jul 2003, van der Werff et
al. 18647 (MO, UC).
2011] LEHNERT: CYATHEACEAE OF PERU
Cyathea serpens is very similar to C. helio-
phila R. M. Tryon regarding the overall
appearance of the fronds; it can be distinguished
by the lack of a trunk (vs. C. heliophila with
trunks to 7 m tall), bicolorous brown-white
petiole scales (vs. predominantly concolorous
whitish), the contorted hairs on the laminae
(vs. contorted hairs absent), and the long
paraphyses (vs. paraphyses of the same length
as the sporangia or shorter than those). The
indusia of C. serpens are very thin, appressed
to the lamina and often caduceous, which may
lead to the assumption that the species is
exindusiate. Cyathea heliophila has always
clearly recognizable, persistent indusia. The
evanescence of the indusia of C. serpens may
be an indicator of a hybrid origin of this
species, with the sympatric C. heliophila
(indusia rm) and C. frigida (indusia lacking)
as putative parents. Traits of C. frigida would
be the long paraphyses, the brown color in the
petiole scales, and the tendency not to form real
trunks.
Cyathea serpens has unusually long fronds
that reach 7 m. However, these expanses are
only known from the type collection; the other
collections barely reach half that length. The
only other neotropical tree fern with similarly
long fronds is C. kalbreyeri (Baker) Domin
which grows in the understory of lower
montane forests and supports its fronds on
high branches of adjoining trees. Confusion
with C. serpens is impossible because besides
growing at a completely different elevation,
C. kalbreyeri also lacks indusia and scurf, and
has concolorous orange-brown petiole scales.
NEW COMBINATIONS
Cyathea alatissima (Stolze) Lehnert, comb.
nov. Basionym: Cnemidaria alatissima
Stolze, Fieldiana, Bot. 37: 55. 1974.
Cyathea nervosa (Maxon) Lehnert, comb.
nov. Basionym: Hemitelia nervosa Maxon, J.
Wash. Acad. Sci. 34: 309. 1944. Cnemidaria
nervosa (Maxon) R. M. Tryon, Contr. Gray
Herb. 200: 52. 1970.
Cyathea uleana (A. Samp.) Lehnert, comb.
nov. Basionym: Hemitelia uleana A. Samp.,
Bol. Mus. Nac. Rio de Janeiro 1: 65.1923.
Cnemidaria uleana (A. Samp.) R. M. Tryon,
Contr. Gray Herb. 200: 52. 1970.
43
EXCLUDED AND DUBIOUS NAMES
Alsophila capensis (L. f.) J. Sm. ssp. polypo-
dioides (Sw.) D. S. Conant, J. Arnold Arbor.
64: 369. 1983.
Remarks: Tryon (in Tryon & Stolze 1989)
reports that this species may occur in Peru,
without specifying the reasons for his
assumption. Alsophila capensis occurs in
small, presumably relict populations on
shaded sandstones in the Atlantic Brazilian
rainforest (Paulo Labiak, pers. comm.). It is
characterized by inermous petioles with
highly dissected aphlebiae at their bases.
The only Peruvian tree fern species with
such aphlebiae is A. salvinii, which can be
distinguished by the lack of indusia and the
tripinnate fronds (vs. hemitelioid indusia
present, fronds bipinnate-pinnatifd in A.
capensis).
Cyathea petiolata (Hook.) R. M. Tryon,
Contr. Gray Herb. 206: 42. 1976.
Remarks: Tryon (in Tryon & Stolze 1989)
reports this species from Peru. It is cur-
rently known only from Colombia, Vene-
zuela, and Central America but I have seen
no material from Peru or Ecuador that
could be condently assigned to this spe-
cies. It is easily recognized by its bipinnate
fronds with large, almost entire pinnules and
submarginally positioned sori with hemitelioid
indusia.
Trichipteris ava R. M. Tryon, Fieldiana Bot.
n.s. 20: 126. 1989. Type: PERU. Hunuco:
Above Chinchao, Tingo Mara to Chin-
chao, 1 Aug 1965, Soejarto 1432 (holo-
type: GH). Cyathea herzogii Rosenst.,
Meded. Rijks. Herb. Leiden 19: 7. 1913.
Remarks: This species was described from
a sterile collection. The laminar indument,
which consists of a dense, short pubescence
on and between the veins adaxially and
strongly bullate squamules abaxially, in com-
bination with the dull brown petiole scales is
indicative that the specimen belongs to
Cyathea herzogii, which is quite common in
the central Andes.
44 BRITTONIA
Acknowledgments
I thank Michael Kessler and Alan R.
Smith, who thought studying tree ferns was
a good idea. They were right. I further thank
my colleagues Alexander N. Schmidt-Lebuhn
(Z) and Johana Toivonen. I am indebted to
the curators of AAU, B, BM, COL, CUZ, F,
GH, GOET, HUT, LIL, MBM, MO, P, QCA,
QCNE, S, SCZ, SP, QPLS, UC, USM,
UPCB, and Z for providing loans or for
attending me during my visits, especially to
Asuncin Cano (USM). Hanna Tuomisto
(TUR) and Maarten Christenhusz (BM)
kindly provided additional specimens. I thank
Dave Conant (LSC) and Blanco Len (TEX,
USM) for helpful comments on the manu-
script. Special thanks go to Blanca Len for
additional data that supports this treatment.
Field studies that contributed to the presented
results were funded by the DAAD (German
Academic Exchange Service) and the DFG
(German Research Foundation, grant GR
1588/7). This research received support from
the SYNTHESYS Project (http_//www.syn
thesys.info/), which is nanced by Eurpoean
Community Research Infrastructure Action
under the FP6 Structuring the European
Research Area Programme.
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