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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 The basis for this study was a doctoral
of the richest fern oras in the world. Tryon research project at the University of California,
and Stolze (1989) reported 1060 native Berkeley, aiming on a revision of all Netropical
species of ferns, horsetails, and lycophytes Cyatheaceae. Besides using the local herbarium
from Peru. Since then the number has grown UC, other herbaria provided loans (F, GH, MO)
by 137 new species and records (Smith et al., or were visited (NY, US) during this period.
2005), signifying an increase by 13%. The ca. Further herbaria consulted prior to and after this
1200 native Peruvian species represent a time are AAU, B, BM, COL, COLO, CUZ,
tenth of the global diversity of the seed-free GOET, HUT, K, LIL, LOJA, LPB, MBM, P,
vascular plants (Smith et al., 2006). QCA, QCNE, QPLS, S, SCZ, SPA, STU, TUR,
The scaly tree ferns (Cyatheaceae) are one UPCB, USM, and Z. The analysis of the
of the most conspicuous fern groups in herbarium specimens was supplemented with
tropical montane forest because of their size rst hand experience on the habit and ecology
and frequency. The family shows a pantrop- of the plants, which were gained on several
ical distribution with more than 600 species eld trips to the Andes between 2000 and 2004.
(Kramer & Green, 1990). Tryon and Stolze Material for this study comprises ca. 1000
(1989) listed 45 species of Cyatheaceae from collections from Peru, including 59 of the
Peru but assumed that at least 19 more authors collections. The presented ranges are
species that are known from adjoining coun- based exclusively on material that has been
tries may occur here as well. The latest revised personally or whose determinations
supplementary account of tree ferns to the were veried recently by colleagues (Blanca
ora of Peru (Smith et al., 2005) is rendered Len, pers. comm.). Thus some departmentos
obsolete because recent taxonomic studies in may be missing from the distribution compared
the genus Cyathea lead to several new to the treatment by Tryon and Stolze (1989).
species circumscriptions, nomenclatural The generic system used here is based on
changes, and range extensions (Lehnert, the works of Conant et al. (1995, 1996) and
2008, 2009a, b). The present work provides Korall et al. (2006, 2007). Three genera are
an updated list of all known Peruvian scaly formally recognized: Alsophila, Cyathea, and
tree ferns. 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 phila gastonyi, Cyathea chontilla, C. nephele,
the monophyletic Cyathea in strict consensus C. polliculi, and C. praetermissa, are
analyses (Korall et al., 2006, 2007; Janssen et described, and all except for C. polliculi are
al., 2008). Both subclades roughly corre- endemic to Peru. The inclusion of Cnemida-
spond with established taxonomic subgroups ria into Cyathea necessitates new combina-
in Alsophila (Holttum, 1963; Holttum & tions for Cyathea alatissima, C. nervosa, and
Edwards, 1983) and are recognized here as C. uleana, and leads to the change of
sect. Alsophila (incl. Nephelea; Conant, 1983) Cnemidaria speciosa C. Presl into C. sub-
and sect. Gymnosphaera. A monophyletic incisa Domin. One previously little known
genus Cyathea must include the genera Cnemi- species, C. serpens, is the last species that
daria and Hymenophyllopsis (Korall et al., had to be transferred from the obsolete
2006, 2007); the genus Trichipteris (or Trichop- genus Trichipteris to Cyathea (Lehnert,
teris) was previously included in Cyathea based 2009b) and is illustrated here in detail for
on morphological characters (Lellinger, 1987). the rst time. Trichipteris ava R. M. Tryon,
The necessity for recombinations arises here which was described from a sterile collection
only for Cnemidaria and Trichipteris, since (Tryon & Stolze, 1989), is synonymized with
Hymenophyllopsis does not occur in Peru. C. herzogii Rosenst. Currently 14 species of
Except for the Sphaeropteris horrida group Cyatheaceae are regarded as being endemic to
(Tryon, 1971), previously recognized Neotrop- Peru.
ical subgenera and sections in Sphaeropteris The presented articial key does not rely
(Tryon, 1970; Windisch, 1977, 1978) were on indusial characters as much as did pre-
transferred to Cyathea (Kramer, 1978; Lellinger, vious keys. Instead, the emphasis is laid on
1984), which is corroborated by the results of the petiolar characters, like scales and scurf,
Korall et al. (2006, 2007). Terminology follows which are more readily available and appa-
Tryon (1970) and Gastony (1973). rently dene natural groups in Cyathea.
However, fertile material is still needed or at
least very helpful for secure identication on
Results
the species level. Where not essential, the
In total, 83 species of Cyatheaceae are type of indusium is given as an additional
known from Peru. Five new species, Also- 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 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

SYSTEMATIC TREATMENT Alsophila erinacea (H. Karst.) D. S. Conant


Alsophila R. Br. sect. Alsophila Depts. Cajamarca, Cuzco, Pasco. 1780
2800 m. (Costa Rica, Colombia, Ecuador,
Alsophila cuspidata (Kunze) D. S. Conant Peru, Bolivia)
Depts. Amazonas, Cuzco, Huanuco, Lor- Voucher: Cajamarca: San Ignacio, San
eto, Madre de Dios, Pasco, Puno, Ucayali. Jos de Lourdes, 050042S, 785415W,
1001100(1570) m. (Costa Rica, Panama, 1800 m, Feb 2006, Bonino 59 (MO, UC).
Colombia, Venezuela, French Guiana, Ecua- Reference: Tryon & Stolze, 1989.
dor, Peru, Bolivia, Paraguay)
Voucher: Madre de Dios: Tambopata, ca.
Alsophila imrayana (Hook.) D. S. Conant
30 air km or 7080 km by river SSW of
Puerto Maldonado at efuence Ro La Torre Depts. San Martin. Ca. 2000 m. (Venezuela,
(Ro Dorbigny)/Ro Tambopata (SE bank), Trinidad, Colombia, Ecuador, Peru)
Tambopata Nature Reserve, 1249S, 6917W, Voucher: San Martin: Rioja, Carretera
260 m, 14 May 1984, Barbour 5297 (F, MO) Moyobamba-Pedro Ruiz, Km 404.1, 5 Aug
Reference: Tryon & Stolze, 1989. 2002, Lehnert 214 (GOET, USM, UC).
Reference: Gastony, 1973; Conant, 1983.

Alsophila engelii R. M. Tryon Alsophila incana (H. Karst.) D. S. Conant


Depts. Cajamarca, Cuzco, San Martin. 1750 Depts. Junin, Pasco, Ucayali. 1130
2150 m. (Colombia, Venezuela, Ecuador, Peru). 1500 m. (Colombia, Ecuador, Peru)
Voucher: Cusco: La Convencin, 1258S, Voucher: Pasco: Oxapampa, Dist. Huan-
07233W, 2150 m, 24 Nov 2003, Bonino et cabmba, sector Pan de Azucar (Huampal),
al. 1155 (CUZ, MO). 101111S, 753424W, 1130 m, 11 Nov
Reference: Tryon & Stolze, 1989. 2004, Mellado 221 (MO, UC).
2011] LEHNERT: CYATHEACEAE OF PERU 21

References: Tryon & Stolze, 1989; Lehnert, Cyathea J. Sm.


2005.
Cyathea alatissima (Stolze) Lehnert,
comb. nov.
Alsophila gastonyi Lehnert, sp. nov. Dept. Hunuco. 1500 m. (Peru)
Dept. Cuzco, Pasco. 520950 m. (Peru) Voucher: Hunuco: Ro Llulla Pichis water-
Voucher: Pasco: Prov. Oxapampa, Dist. shed, 25 Jul 1969, Dudley 13282 (holotype:
Palcaz, Reserva Comunal Yanesha, Comu- GH-photo F; isotype: NA n.v.).
nidad Nativa San Pedro de Pichanaz, Sector References: Stolze, 1974; Tryon & Stolze,
Azulis, 102644S, 750621W, 910 m, 15 1989.
Sep 2005, Monteagudo A. et al. 9810
(holotype: MO).
Cyathea andina (H. Karst.) Domin
Alsophila minervae Lehnert Dept. Madre de Dios. 200 m. (Greater
Dept. Cuzco, Pasco. 22002900 m. (Peru, Antilles, Colombia, Ecuador, Peru, Bolivia)
Bolivia) Voucher: Madre de Dios: Tambopata,
Voucher: Pasco: Oxapampa, trail to sum- 20 km NW from the mouth of Ro Inambari
mit of Cordillera Yanachaga via Rio San N bank of Ro Madre de Dios, at the end of the
Daniel, 1023S, 7527W, 2500 m, 17 Jul airstrip of Lagarto 1238S, 6951W, 200 m
1984, Smith D. N. et al. 7817 (USM). 11 Nov 1998, Tuomisto et al. 13679 (TUR).
References: Lehnert, 2003; Smith et al., 2005. Reference: Tryon & Stolze, 1989.

Alsophila mostellaria Lehnert Cyathea arnecornelii Lehnert


Depts. Amazonas, Cajamarca, Pasco. 1500 Dept. Cuzco. 2980 m. (Peru, Bolivia)
2800 m. (Ecuador, Peru, Bolivia) Voucher: Cuzco: La Convencin, Dist.
Voucher: Amazonas: Bongar, carretera Quellouno, Yavero, 12 3535S, 722321
Pedro Ruiz-Florida, 0551.7S, 7758.4W, W, 2980 m, Valenzuela et al. 7784 (MO, UC).
2200 m, 5 Aug 2002, Lehnert 243 (holotype: Det. A. R. Smith 2008.
USM; isotypes: GOET, UC). References: Lehnert, 2003; Smith et al.,
Reference: Lehnert, 2003, 2005. 2005.

Alsophila paucifolia Baker


Depts. Ucayali, Amazonas, ca. 9001300 m. Cyathea atahuallpa (R. M. Tryon) Lellinger
(Colombia, Ecuador, Peru) Depts. Amazonas, Cajamarca, La Libertad.
Voucher: Ucayali (Loreto): Prov. Coronel 27003100 m. (Ecuador, Peru)
Portillo, La Divisoria, 29 Jul 1948, Aguilar Voucher: Amazonas: Chachapoyas, Cerros
850 (USM). Calla Calla, E side, 18 km above Leime-
Reference: Tryon & Stolze, 1989. bamba on the road to Balsas, 30003100 m,
Remarks: First conmed report for Peru; 15 Oct 1968, Hutchinson & Wright 6922
the presence was long suspected (Tryon & (holotype: US; isotypes: NY, UC, US).
Stolze 1989). References: Lellinger, 1987; Tryon &
Stolze, 1989.
Alsophila R. Br. sect. Gymnosphaera Blume
Alsohila salvinii Hook. Cyathea aterrima (Hook.) Domin
Dept. San Martin. 1850 m. (Mexico, Depts. Amazonas, San Martin. 700
Guatemala, El Salvador, Belize, Honduras, 1200 m. (Colombia, Ecuador, Peru)
Costa Rica, Panama, Peru) Voucher: San Martin: In monte Guayr-
Voucher: San Martn: Rioja, 054029S, apurima, prope Tarapoto, Peruvia orientalis,
774625W, 1850 m, 25 Mar 1998, van der Sept 1856, Spruce 4713 (holotype: BM;
Werff et al. 15666 (MO, USM). isotypes: K, US).
References: Stolze, 1976; Conant, 1983; References: Tryon & Stolze, 1989; Moran
Mickel & Smith, 2004. et al. 2008.
22 BRITTONIA [VOL 63

Cyathea austropallescens Lehnert Cyathea concordia B. Len & R. C. Moran


Depts. Cuzco, Pasco. 24103200 m. (Peru, Dept. Amazonas. 18002050 m. (Peru)
Bolivia) Voucher: Amazonas: Condorcanqui, Cor-
Voucher: Pasco: Oxapampa Dist. Oxa- dillera del Cndor, Cerro Machinaza, Puesto
pampa, P. N. Yanachaga-Chemilln, Abra de Vigilancia Alfonso Ugarte, cabeceras del
Yanachaga, 1022S, 7527W, 2900 m, 1 Ro Comainas, 0422S, 7811W, 1800
Aug 2003, Vsquez et al. 28458 (UC, MO). 2050 m, 21 Jul 1994, Bltran & Foster
Reference: Lehnert, 2008. 1183 (holotype: USM; isotype: UC-fragm.).
Reference: Len & Moran, 1996.
Cyathea bipinnatida (Baker) Domin
Depts. Amazonas, Cajamarca, Cuzco, Madre Cyathea conjugata (Hook.) Domin
de Dios, Pasco, San Martin. 4002100 m. Depts. La Paz, Cochabamba, Santa Cruz.
(Venezuela, Colombia, Ecuador, Peru, Bolivia) 11002750 m. (Colombia, Ecuador, Peru,
Voucher: Amazonas: Bagua, Quebrada El Bolivia)
Almendro, 051440S, 782124W, 430 m, Voucher: Cuzco: Urubamba, Dist. Machu
8 Mar 1998, van der Werff et al. 14506 (MO, Picchu, Cerro Aguas Calientes, 1309S 72
UC, USM). 31W, 30003500 m, 22 Nov 2003, Huaman-
Reference: Tryon & Stolze, 1989. tupa & Quispe 3838 (MO, UC)
References: Lellinger, 1987; Tryon & Stolze,
1989.
Cyathea bradei (Windisch) Lellinger
Remarks: Cyathea tryonorum (Riba) Lel-
Dept. Loreto, San Martin. 100200 m. linger is believed to be conspecic.
(Colombia, Ecuador, Peru, Bolivia)
Voucher: Loreto: Ro Tigre, 5 km N of the
Cyathea crenata (Sodiro) H. Christ
village Paicheplaya E of the river, 0259S,
7518W, 100200 m, 31 Jan 2005, Tuomisto Dept. Cajamarca. 1900 m. (Ecuador, Peru)
et al. 14768 (GOET, TUR). Voucher: Cajamarca: San Ignacio, Dist.
References: Lellinger, 1984; Tryon & San Jos de Lourdes, Buenos Aires (Cerro El
Stolze, 1989. Parco), 054204S, 775306W, 1900 m,
Feb 2002, Bonino 307 (MO, UC).
Reference: Lehnert, 2009a.
Cyathea carolihenrici Lehnert
Dept. Cuzco. 2050 m. (Peru, Bolivia) Cyathea cystolepis Sodiro
Voucher: Cuzco: Prov. Urubamba, 2050 m, Depts. Amazonas, San Martin. 2060
Bonino 1127 (CUZ). 2750 m. (Colombia, Ecuador, Peru)
References: Lehnert, 2003, Smith et al., 2005. Reference: Lehnert, 2008.
Remarks: Three varieties can be distin-
Cyathea catacampta Alston guished, two of them in Peru:
Depts. Cuzco, San Martin. 20003050 m.
(Colombia, Ecuador, Peru, Bolivia)
Voucher: Cuzco: La Convencin, Dist. 1. Hairs on the segments more abundant adaxially or
equally dense on both sides, generally erect and
Huayopata, Balconpata, 125201S, 7232 abundant on and between the veins; bullate
46W, 2200 m, 21 Apr 2004, Calatayud et al. squamules frequent abaxially, mainly pure white,
2342 (MO rarely with brown body. . . . . . . . . . . . var. leonis
Reference: Lehnert, 2009a. 1. Hairs on the segments more abundant abaxially
than adaxially on and between the veins, except if
hairs generally few and appressed, then sometimes
Cyathea chontilla Lehnert, sp. nov. few hairs adaxially and almost no hairs abaxially,
hairs mainly on the veins; bullate squamules with
Depts. Cajamarca, Pasco. 23002850 m. (Peru) brown body if abundant.
Voucher: Cajamarca: San Ignacio, Huar- 2. Hairs on the segments more abundant abaxially,
ango, localidad Romerillo, Cordillera entre sparse to abundant, mainly erect adaxially,
Romerillo y Nuevo Mundo, 0514S, 7846 appressed adaxially, bullate squamules few to
many, always brown-bodied and pure white
W, 2300 m, 29 Jul 1992, Campos et al. 4294 ones mixed, the brown-bodied squamules often
(holotype: UC; isotypes: MO, USM). dominating. . . . . . . . . . . . . . . . var. cystolepis
2011] LEHNERT: CYATHEACEAE OF PERU 23

2. Hairs sparse on both laminar surfaces, often Cyathea dintelmannii Lehnert


appressed, bullate squamules abaxially few, only Depts. Cuzco, Pasco. 15352250 m. (Peru,
white, or lacking. . . . . . . . . var. boreopallescens Bolivia)
Voucher: Cusco: La Convencin, Dist.
Cyathea cystolepis var. boreopallescens Vilcabamba, Espiritupampa, 125425S, 73
Lehnert 1216W, 1535 m, 22 Jul 2004, Calatayud et
Voucher: Amazonas: Bagua Cordillera al. 2582 (MO, UC).
Coln SE of La Peca, 25002600 m, 30 Sep References: Lehnert, 2006b, 2008.
1982, Barbour 3611 (AAU, MO, USM).
Cyathea divergens Kunze
Cyathea cystolepis var. leonis Lehnert. Depts. San Martin, Huanuco, Pasco. 1590
Voucher: Amazonas: Chachapoyas, Distr. 1800 m. (Costa Rica, Panama, Colombia,
Leymebamba, alrededor de la laguna de los Venezuela, Ecuador, Peru)
Condores, parte sur, 0651.201S, 7740.958W, Voucher: Hunuco: Pampayaco, Jul 1829,
25002700 m, 17 Aug 2002, Quipuscoa et al. Poeppig s.n. (diar. 1163) (holotype: LZ,
1234 (holotype: F; isotypes: GOET, UC). destroyed). Authentic specimens (de Tryon
1976): Poeppig 219 (diar. 1152) (B-fragm.
Cyathea delgadii Sternb. NY, K-fragm. NY, P-photo GH)
Depts. Amazonas. Cuzco, Huanuco, Madre References: Tryon & Stolze, 1989; Lehnert,
de Dios, Pasco, San Martin. 5652200 m. (Costa 2008.
Rica, Panama, Colombia, Venezuela, Ecuador, Remarks: Of the two currently recognized
Peru, Brazil, Bolivia, Paraguay, Argentina) varieties (Tryon, 1976; Tryon & Stolze, 1989),
Voucher: Pasco: Prov. Oxapampa, Dist. only the var. divergens is native to Peru.
Oxapampa, sector Grapanazu, relleno sani-
tario, 103314S, 752535W, 1800 m, 17 Cyathea dombeyi (Desv.) Lellinger.
Mar 2005, Mellado et al. 2904 (MO). Dept. Pasco. 1290 m. (Peru, Bolivia)
References: Tryon & Stolze, 1989; Lehnert, Voucher: In Peruvia, Dombey s.n. (hol-
2006b. otype: P n.v.; isotype: P n.v.).
Remarks: A variable species whose extremes Reference: Barrington, 1978.
can be mistaken for different species: Remarks: The species was previously includ-
ed in C. pungens (Willd.) Domin (Christensen,
1905; Tryon & Stolze, 1989; Lehnert, 2006a).
1. Segments 34 mm broad, margins conspicously
crenulate to crenate; rachises abaxially sparsely
hairy, not scabrous with hair bases, though some- Cyathea dudleyi R. M. Tryon
times muricate with little spines; petioles blackish Depts. Amazonas, Cajamarca, Cuzco, Pasco,
to purpureous. . . . . . . . .C. delgadiitypical form San Martin. 19002700 m. (Ecuador, Peru)
1. Segments 23 mm wide, with margins entire or
inconspicuously crenulate; rachises abaxially Voucher: Cuzco: Prov. Convencin, 12
densely hairy or, if hairs abraded, notably scabrous 37S, 7333W, 26002700 m, 7 Jul 1968,
with hair bases; petioles dull brown to yellowish. Dudley 10867B (holotype: GH; isotypes:
. . . . . . . . . . . . . . . . C. delgadiipubescent form
MO, NA n.v., US).
References: Tryon, 1976; Tryon & Stolze,
Typical form: In perhumid premontane and 1989.
montane forest of Costa Rica, the Andes and
the mata atlantica of Brazil; mainly above Cyathea ebenina H. Karst.
1000 m. Depts. Amazonas, Cajamarca, Cuzco,
Pubescent form: In premontane forest with Pasco, San Martin. 19002700 m. (Colombia,
weak seasonality and poor soils in the Ecuador, Peru)
Venzuelan gran sabana, the Bolivian Chi- Voucher: Cajamarca: San Ignacio, San
quitania, and the Brazilian campo cerrado; Jos de Lourdes, locality Picorana, 0458
only below 900 m. In Peru, in the Huanca- 00S 785301W, 23502450 m, 4 Dec
bamba depression of Depts. San Martin and 2002, Campos et al. 5898 (UC).
Amazonas. Reference: Tryon & Stolze, 1989.
24 BRITTONIA [VOL 63

Cyathea ewanii Alston San Daniel, 1015S, 7515W, 2500 m, 21 Aug


Depts. Amazonas, Loreto. 100660 m. 2005, Monteagudo et al. 9562 (MO, UC).
(Ecuador, Peru) Reference: Tryon, 1986.
Voucher: Amazonas: Bagua, Distrito Imaza,
regin del Maran, comunidad de Yamayakat, Cyathea herzogii Rosenst.
Rio Maran, 0455S, 7819W, 600 m 28 Jan Depts. Cuzco, Hunuco, Junin, Pasco.
1995, Rodriguez 285 (UC). 20003200 m. (Peru, Bolivia)
Reference: Tryon & Stolze, 1989. Voucher: Pasco: Oxapampa, Dist. Huanca-
Remarks: Synonym Cnemidaria ewanii bamba, sector quebrada Yanachaga (P.N.
(Alston) R. M. Tryon. Yanachaga-Chemilln), 102326S, 7528
21W, 2545 m, 22 Sep 2004, Mellado &
Cyathea frigida (H. Karst.) Domin Becerra 1884 (MO).
Depts. Cajamarca, Huanuco, Pasco. 2650 References: Smith et al. 2005, Lehnert
3300 m. (Colombia, Ecuador, Peru) 2006b.
Voucher: Hunuco: Huanuco, Huanuco-
Tingo Maria road, steep slopes near Carpish Cyathea horrida (L.) Sm.
divide above mouth of tunnel, 0942S, 76 Depts. Amazonas, Pasco, San Martin. 320
05W, 26502700 m, 3 Feb 1984, Gentry & 770 m. (Costa Rica, Panama, Greater and
D. N. Smith 44863 (MO, NY). Lesser Antilles, Colombia, Venezuela, Ecua-
Reference: Tryon & Stolze, 1989. dor, Peru)
Remarks: Often confused with C. nephele Voucher: Amazonas: Bagua, Distr. Imaza,
Lehnert; Peruvian specimens differ from the Quebrada Almendra, 051440S, 782134
Colombian type material (and most speci- W, 400 m, 17 Mar 2001, van der Werff et al.
mens from Ecuador) by having longer para- 16134 (MO).
physes, fewer or no ciliate laminar squamules, Reference: Tryon & Stolze, 1989.
and on average relatively paler petioles and Remarks: Synonym Cnemidaria horrida
leaf axes. (L.) C. Presl.

Cyathea frondosa H. Karst. Cyathea kalbreyeri (Baker) Domin


Dept. San Martin. 20602100 m. (Vene- Depts. Junin, San Martin. 11701200 m.
zuela, Colombia, Ecuador, Peru) (Colombia, Venezuela, Ecuador, Peru, Bolivia)
Voucher: San Martin: Huallaga, arriba de Voucher: Junin: Satipo, Gran Pajonal, E of
La Ribera-La Meseta, 0652S, 7728W, Chequitavo on trail to Kotampaz, 1045S,
20602100 m, 17 Aug 1997, Quipuscoa-S. 7423W, 1200 m, 5 Apr 1998, Smith D. N.
et al. 1085 (AAU). 6749 (UC).
Reference: Lehnert, 2009a. Reference: Tryon & Stolze, 1989.

Cyathea gracilis Griseb. Cyathea lasiosora (Kuhn) Domin


Depts. Amazonas, Pasco. 7702550 m. Depts. Amazonas, Loreto, Madre de Dios,
(Costa Rica, Jamaica, Colombia, Ecuador, Peru) Pasco, Puno, San Martin, 100800(1030) m.
Voucher: Pasco: Oxapampa, Dist. Palcaz, (Colombia, Venezuela, French Guiana, Ecua-
San Francisco de Pichanaz, 102916S 75 dor, Peru, Brazil, Bolivia)
0358W, 770 m, 28 Feb 2004, Mellado et al. Voucher: Loreto: Loreto, about 2 km N of
894 (MO). the village San Antonio at lower Ro Mar-
Reference: Tryon & Stolze, 1989. aon, 0432S, 7338W, 100200 m, 14 Jan
1996, Tuomisto et al. 8055 (TUR, UC).
Cyathea heliophila R. M. Tryon Reference: Tryon & Stolze, 1989.
Dept. Pasco. 2500 m. (Colombia, Ecuador,
Peru, Bolivia) Cyathea latevagans (Baker) Domin
Voucher: Pasco: Oxapampa, Dist. Huanca- Dept. Cuzco, San Martin. 18503415 m.
bamba, P. N. Yanachaga-Chemilln, sector Hito- (Colombia, Venezuela, Ecuador, Peru)
2011] LEHNERT: CYATHEACEAE OF PERU 25

Voucher: San Martin: Rioja, along road Cyathea lockwoodiana (Windisch) Lellinger
Rioja-Pedro Ruiz, El Mirador, 054029S, Dept. Loreto. 100200 m. (Panama,
774625W, 1850 m, 26 Mar 2002, van der Colombia, Venezuela, Ecuador, Peru)
Werff et al. 15705 (MO, UC, USM). Voucher: Loreto: Prov. Loreto, about 2 km N
Reference: Tryon & Stolze, 1989. of the village San Antonio at lower Ro
Maraon, 0432S, 7338W, 100200 m, 14
Jan 1996, Tuomisto H. et al. 8188 (LSC, TUR).
Cyathea lechleri Mett. Reference: Windisch, 1977.
Depts. Amazonas, Pasco, Puno, San Mar-
tin, 7802160 m. (Colombia, Venezuela, Cyathea macrosora (Baker) Domin
Peru, Bolivia) Depts. Loreto, Pasco, 100700 m. (Colom-
Voucher: Pasco: Oxapampa, Distrito Pal- bia, Venezuela, the Guyanas, Ecuador, Peru)
cazu, P.N. Yanachaga-Chemilln, one hour Remarks: Two varieties can be distinguished:
from Estacin Biologica Paujil, 1020S, 75
15W, 780 m, 25 Oct 2002, Monteagudo et al.
4272 (MO, UC). 1. Petioles and leaf axes with little scurf, often
Reference: Tryon & Stolze, 1989. 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,
Cyathea leucolepismata Alston often partially absent in one frond. . . . var. vaupensis
Depts. Loreto, Madre de Dios. 90500 m.
(Colombia, Ecuador, Peru, Bolivia) Cyathea macrosora var. macrosora
Voucher: Loreto: Maynas, between the Voucher: Pasco: Oxapampa, along road Cha-
roads Ex Petroleros and Bello Horizonte, tarra-Pto. Bermudz, 1030S, 7503W, 700 m,
Km 3840 of the road Iquitos-Nauta, 0404 9 Jul 2003, van der Werff et al. 18171 (MO, UC).
S, 7328W, 100200 m, 6 Jan 1995, Tuo-
misto et al. 6677 (GOET, TUR). Cyathea macrosora var. vaupensis (Windisch)
Reference: Moran, 1995. A. R. Sm.
Remarks: Synonym C. amazonica R. C.
Voucher: Loreto: Loreto, Ro Tigre, about
Moran.
1.5 km E of the river, 0357S, 7417W, 100
200 m, 10 Jan 2005, Tuomisto et al. 13983
Cyathea lindigii (Baker) Domin (TUR, USM).
References: Lellinger, 1984; Tryon &
Depts. Cuzco, Madre de Dios. 400500 m.
Stolze, 1989.
(Colombia, Ecuador, Peru, Bolivia)
Voucher: Madre de Dios: Manu, Parque
Nacional Manu, Ro Manu, Rio Sotileja, 11 Cyathea meridensis H. Karst.
40S, 7155W, 400500 m, 4 Oct 1986, Depts. Amazonas, Cuzco, Cajamarca. 2295
Foster & DAchille, 11627 (F, LPB) 2830 m. (Colombia, Venezuela, Ecuador, Peru)
Reference: Tryon & Stolze, 1989. Remarks: Three varieties can be distinguished:
Remarks: Distinct from C. multiora Sm.,
1. Fertile fronds to 1 m long; pinnules to 65(80) mm
under which it was previously treated (Tryon, long, all sessile or few subsessile; trunks to 1.5 m tall,
1976). 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
Cyathea lindeniana C. Presl 10 m tall, petiole bases persistent or not.
2. Fertile fronds hanging free, arching to distally
Depts. Cajamarca, San Martin. 2000 drooping; trunks to 10 m tall, without
2200 m. (Colombia, Venezuela, Ecuador, Peru) persistent petiole bases, petioles falling
Voucher: Cajamarca: Lower edge of cleanly at an early age. . . . . var. meridensis
Cutervo National Park, 1015 km N of San 2. Fertile fronds leaning and draped over
adjacent vegetation; trunks relatively small
Andres de Cutervo, 0610S, 7840W, 2200 m, compared to the frond length, 0.32.0 m tall,
13 Feb 1992, Gentry et al. 61560 (UC). petiole bases persistent, rotting off only in
Reference: Lehnert, 2009a. older plants. . . . . . . . . . . . . . . var. obtecta
26 BRITTONIA [VOL 63

Cyathea meridensis var. meridensis Cyathea multisegmenta R. M. Tryon


Voucher: Cuzco: Paucartambo, Kosipata, Depts. Cuzco, Pasco. 18502295 m. (Peru)
Trocha Unin, Km 10, 19 L01221737, UTM Voucher: Cuzco: Prov. La Convencin, Dud-
8552556, 2295 m, 3 Sep 2003, Garca K. et ley 11326 (holotype: NA; isotypes: GH, US).
al. 207 (MO). References: Tryon, 1976, Tryon & Stolze,
1989.

Cyathea meridensis var. nana Lehnert


Voucher: Cajamarca: San Ignacio San Cyathea nephele Lehnert, sp. nov.
Jos de Lourdes, Cerro Picorana, 0458S, Depts. Amazonas, Pasco, San Martin.
7853W, 2830 m, 17 Aug 1998, Campos et 27203550 m. (Peru)
al. 5550 (MO, UC). Voucher: Pasco: Prov. Oxapampa, Distrito
Huancabamba, Parque Nacional Yanachaga-
Chemilln, 1022S, 7528W, 3110 m, 5 Dec
Cyathea meridensis var. obtecta Lehnert 2003, Monteagudo A. et al. 6430 (holotype:
Voucher: Amazonas: Laguna de Pomaco- UC; isotype: MO).
chas, 2550 m, 28 Mar 2002, van der Werff et
al. 15835 (MO, UC).
Cyathea nervosa (Maxon) Lehnert, comb. nov.
Reference: Lehnert, 2009a.
Dept. Amazonas. 300400 m. (Ecuador, Peru)
Voucher: Amazonas: Prov. Condorcanqui,
above Pongo de Manseriche, gully to right of
Cyathea microdonta (Desv.) Domin
mouth of Ro Santiago, 300 m, 18 Dec 1931,
Depts. Cuzco, Loreto, Pasco, San Martin. Mexia 6291 (holotype: F; isotypes: GH, UC).
1001100 m. (Mexico, Central America, Greater Reference: Tryon & Stolze, 1989.
Antilles, Colombia, Venezuela, Trinidad, the
Guyanas, Ecuador, Peru, Brazil, Bolivia)
Voucher: Pasco: Oxapampa, Distr. Palcaz, Cyathea obnoxia Lehnert
El Paujil, 101924S, 751549W, 500 m 11 Depts. Amazonas, Cajamarca, San Martin.
May 2005, van der Werff et al. 19989 (MO). 22002700 m. (Colombia, Ecuador, Peru,
Reference: Tryon & Stolze, 1989. Bolivia)
Voucher: Amazonas: Road Chachapoyas-
Mendoza, a little past Molinopampa, 0614
Cyathea microphylla Mett. 11S, 773549W, 2400 m, 15 Mar 1998,
Dept. Puno. 2100 m. (Peru) van der Werff et al. 15038 (MO, UC, USM).
Voucher: Puno: Prov. Carabaya, St. Gavan, Reference: Lehnert, 2006a.
Lechler 2160 (holotype: LZ, destroyed);
authentic specimen Lechler 2569 (B-fragm.
F, P-photo GH). Cyathea palaciosii R. C. Moran
Reference: Tryon & Stolze, 1989. Dept. Amazonas. 1950 m. (Ecuador, Peru)
Voucher: Amazonas: San Martin, 0541S,
7748W, 1950 m, 4 Mar 2001, van der Werff
Cyathea mucilagina R. C. Moran et al. 16721 (MO, UC).
Dept. Hunuco, Madre de Dios, Pasco, San Reference: Moran, 1995.
Martin. 490770 m. (Costa Rica, Colombia,
Ecuador, Peru, Bolivia)
Voucher: San Martin: Prov. San Martin, Cyathea parvifolia Sodiro
Progreso, village in the rst valley of the Dept. Pasco. 28002900 m. (Colombia,
Cerro Escalera, from Tarapoto after the Ecuador, Peru, Bolivia)
tunnel, along Tarapoto-Yurimaguas, 0624S, Voucher: Pasco: Oxapampa, Dist. Huanca-
7618W, 600 m, 8 Aug 2006, Christenhusz bamba, Lanturachi, sector Santa Barbara,
et al. 2024 (GOET, TUR, USM). road to Milpo, 1022S, 7536W, 2824 m,
Reference: Moran, 1991. 10 Oct 2003, Perea et al. 721 (MO).
2011] LEHNERT: CYATHEACEAE OF PERU 27

References: Moran, 1995; Lehnert, 2008, 14.26S, 7735.96W, 2700 m, 4 Aug 2002,
2009a. Lehnert 240 (GOET, UC, USM)
Remarks: Synonym Cyathea brevistipes R. References: Lehnert, 2006a.
C. Moran
Cyathea polliculi Lehnert, sp. nov.
Cyathea patens H. Karst. Dept. Pasco. 2450 m. (Ecuador, Peru)
Dept. Amazonas, Cajamarca, Cuzco. Voucher: Pasco: Oxapampa, near antenna
(1900)27003300 m. (Colombia, Venezuela, past Chacos, 1038S, 7515W, 2450 m, 19
Ecuador, Peru) Jul 2003, van der Werff et al. 18733 (MO,
Voucher: Cuzco: La Convencin, Dist. UC).
Vilcabamba, Espiritupampa, 1303S, 7305W,
3300 m, 15 Oct 2003, Suclli et al. 1330 (MO).
Reference: Lehnert, 2009a. Cyathea poeppigii (Hook.) Domin
Remarks: The earlier homonym Cyathea Depts. Amazonas, Cajamarca, Huanuco,
patens hort., Houlst. & Moore is an invalidly Junin, Pasco, San Martin. 3001500 m.
published nomen nudum. The later use of this (Costa Rica, Panama, Colombia, Venezuela,
epithet is not illegitimate (Tryon, 1976; Ecuador, Peru, Brazil, Bolivia)
Lehnert, 2009a). Voucher: Cajamarca: Prov. Bagua, entre
Bagua Grande y Chiriaco, 0522.08S, 78
Cyathea peladensis (Hieron.) Domin 27.15W, 400 m, 20 Jul 2002, Lehnert 197
Depts. Cajamarca. 1900 m. (Colombia, (GOET, UC, USM)
Ecuador, Peru) References: Windisch, 1977; Tryon &
Voucher: Cajamarca: San Ignacio, Dist. Stolze, 1989.
San Jos de Lourdes, Buenos Aires (Cerro El
Parco), 054204S, 775306W, 1900 m,
Cyathea praetermissa Lehnert, sp. nov.
Feb 2006, Bonino 238 (MO, UC).
Dept. Amazonas. 2450 m. (Peru)
References: Tryon, 1976, 1986.
Voucher: Amazonas: A few Km from
Molinopampa, [ca. 1038S, 7515W,
Cyathea phegopteroides (Hook.) Domin 2450 m,] 14 Mar 1998, van der Werff et al.
Depts. Cajamarca, San Martin. 1100 14935 (holotype: UC; isotypes: F, MO,
1800 m. (Peru) USM).
Voucher: San Martin: Peruvia occidenta-
lis, Tarapoto, on the top of Mt. Guayrapur-
ina, Spruce 4020 (holotype: K; isotype: B). Cyathea pungens (Willd.) Domin
References: Tryon, 1976. 1986. Depts. Amazonas, Cuzco, Huanuco, Lor-
eto, Madre de Dios, Pasco, Puno, San Martin,
Cyathea pilosissima (Baker) Domin Ucayali. 1001700 m. (Trinidad, Colombia,
Depts. Amazonas, Loreto, Pasco, San Venezuela, the Guyanas, Ecuador, Peru, Bra-
Martin. 3501500 m. (Colombia, Venezuela, zil, Bolivia)
Ecuador, Peru) Voucher: San Martin: Rioja, along road
Voucher: San Martin: Monte Campana, Rioja-Pedro Ruiz, 054027S, 774035W,
Spruce 4322 (holotype: K-photo GH; isotype: 1170 m, 23 Mar 1998, van der Werff et
BR-photos GH, US). al.15511 (MO, UC).
Reference: Tryon & Stolze, 1989. Reference: Tryon & Stolze, 1989.

Cyathea plicata Lehnert Cyathea rufescens (Mett. ex. Kuhn) Domin


Depts. Amazonas, Cajamarca. 2100 Dept. San Martin. Ca. 1400 m. (Peru)
2700 m. (Ecuador, Peru) Voucher: San Martin: Tarapoto, Spruce
Voucher: Amazonas: Chachapoyas, Carre- 4212 (holotype: K).
tera Chachapoyas-Mendoza, Km 52 de Cha- References: Windisch, 1978; Tryon &
chapoyas, 10 km detrs de Molinopampa, 06 Stolze, 1989.
28 BRITTONIA [VOL 63

Cyathea ruiziana Klotzsch References: Stolze, 1974; Tryon & Stolze,


Depts. Cuzco, Huanuco, Junin, Pasco. 1989.
20003000 m. (Peru, Bolivia) Remarks: Synonym Cnemidaria speciosa
Voucher: Huanuco [Panhatahuas]: Ruiz C. Presl.
72 (180) (holotype: B).
References: Tryon, 1976; Lehnert, 2008.
Remarks: Synonym Cyathea boliviana R. M. Cyathea thelypteroides A. R. Sm.
Tryon. Dept. Amazonas. 10301200 m. (Peru)
Voucher: Amazonas: Bagua, upper slopes
and summit of Cerro Tayu, 051556S, 78
Cyathea serpens (R. M. Tryon) Lehnert 2207W, 1030 m, 22 Mar 2001, van der
Dept. Cuzco, Pasco. 24502800 m. (Peru) Werff et al. 16323 (holotype: USM; isotypes:
Voucher: Cuzco: La Convencin, Cordil- MO, UC).
lera Vilcabamba, 10 Jul 1968, Dudley 10949 References: Smith et al., 2005, Smith,
(holotype: NA n.v.; isotype: GH). 2006.
References: Tryon & Stolze, 1989; Lehnert,
2009b.
Cyathea tortuosa R. C. Moran
Depts. Amazonas, Junin, Loreto, Madre de
Cyathea squamipes (H. Karst.) Domin Dios, Pasco, Puno, San Martin. 1001000 m.
Depts. Amazonas, Cajamarca, Cuzco, Hua- (Colombia, Ecuador, Peru)
nuco, Pasco, San Martin. 11502900 m. Voucher: Puno: Ro Candamo, la at
(Colombia, Venezuela, Ecuador, Peru, Bolivia) mouth of Ro Guacamayo, 1330S, 6950
Voucher: Amazonas: Laguna de Pomaco- W, 810 m, 25 May 1992, Gentry et al. 77189
chas, 2550 m, 28 Mar 2002, van der Werff et (MO, UC).
al. 15796 (MO, UC, USM). References: Moran, 1991; Smith et al.,
Reference: Tryon & Stolze, 1989; Lehnert, 2006.
2009a.
Remarks: Synonym Cyathea caracasana
var. boliviensis (Rosenst.) R. M. Tryon. Cyathea traillii (Baker) Domin
Depts. Amazonas, Loreto, Pasco. 100
800 m. (Costa Rica, Panama, Colombia,
Cyathea straminea H. Karst. Venezuela, the Guyanas, Ecuador, Peru, Brazil)
Dept. Amazonas, Pasco, San Martin. 3110 Voucher: Amazonas: Bagua, Cerro Tayu,
3400 m. (Colombia, Venezuela, Ecuador, ca. 1 h from Chiriaco, 051556S, 782207
Peru, Bolivia) W, 19 Mar 2001, 800 m, van der Werff et al.
Voucher: Pasco: Oxapampa, Dist. Huanca- 16210 (MO, UC).
bamba, P. N. Yanachaga-Chemilln, 1022 References: Windisch, 1978; Tryon &
32S, 752804W, 3110 m, 6 Apr 2004, Stolze, 1989.
Monteagudo et al. 6715 (MO). Remarks: Synonym Cyathea reginae
References: Tryon & Stolze, 1989; Lehnert, (Windisch) A. R. Sm. ex Lellinger.
2008.

Cyathea tungurahuae Sodiro


Cyathea subincisa (Kunze) Domin Dept. Cuzco. 2200 m. (Colombia, Vene-
Depts. Amazonas, Huanuco, Madre de zuela, Ecuador, Peru)
Dios, Pasco, San Martin, Ucayali. 300 Voucher: Cuzco: La Convencin, Dist.
1515 m. (Ecuador, Peru, Bolivia) Santa Ana, Tunquimayo, 125431S 7248
Voucher: Pasco: Oxapampa, Dist. Palcaz, 45W, 2200 m, 27 Sep 2004, Calatayud et al.
sector Paujil, near the colpa, 101924S, 75 2893 (MO).
1549W, 380 m, 26 Feb 2004, Mellado 2814 Reference: Lehnert, 2008.
(MO). Remarks: First record for Peru.
2011] LEHNERT: CYATHEACEAE OF PERU 29

Cyathea uleana (A. Samp.) Lehnert, Cyathea xenoxyla Lehnert


comb. nov. Depts. Amazonas, Cuzco, Pasco, San Mar-
Dept. Cuzco, Pasco. 11002240 m. (Ecua- tin, Ucayali. 10002770 m. (Colombia, Ecua-
dor, Peru, Brazil, Bolivia) dor, Peru, Bolivia).
Voucher: Pasco: Oxapampa, La Suiza Voucher: Amazonas: Chachapoyas, Carre-
Nueva, 1038S, 7527W, 2240 m, 21 Jun tera Chachapoyas-Mendoza, Km 52 de Cha-
2003, van der Werff et al. 17546 (MO). chapoyas, 10 km detrs de Molinopampa, 06
References: Stolze, 1974; Tryon & Stolze, 14.26S, 7735.96W, 2700 m, 4 Aug 2002,
1989. Lehnert 229 (GOET, UC, USM).
Remarks: Synonym Cnemidaria uleana References: Lehnert, 2003; Smith et al.
(A. Samp.) R. M. Tryon var. uleana. Stolze 2005.
(1974) further distinguished the var. abita-
guensis (Domin) Stolze from Colombia and Sphaeropteris Bernh.
Ecuador.
Sphaeropteris quindiuensis (H. Karst.)
R.M. Tryon
Cyathea ulei (H. Christ) Domin Depts. Amazonas, Cajamarca, Cuzco, Hu-
nuco, Junin, Pasco, San Martn. (1700)
Depts. Cuzco, Hunuco, Pasco, Puno, San
19702400(2700)m. (Colombia, Ecuador,
Martn. 10301300 m. (Colombia, Venezuela,
Peru, Bolivia)
Ecuador, Peru, Bolivia)
Voucher: Pasco: Oxapampa, Dist. Oxa-
Voucher: San Martn: Prov. Picota, Cerro de
pampa, sector quebrada San Alberto, 1032
Ponasa, 1300 m, Mar 1903, Ule 6901 (hol-
46S, 752125W, 2400 m, 21 Aug 2004,
otype: P?; isotypes: B-photo GH, L-photo US).
Mellado & Francis 1543 (MO).
References: Barrington, 1978; Tryon &
References: Tryon, 1971; Tryon & Stolze,
Stolze, 1989.
1989.
NEW SPECIES
Cyathea vilhelmii Domin
Alsophila gastonyi Lehnert, sp. nov. Type:
Dept. Puno. Ca. 2000 m. (Peru)
PERU. Pasco: Prov. Oxapampa, Dist. Pal-
Voucher: Puno: Prov. Carabaya, Tatanara,
caz, Reserva Comunal Yanesha, Comuni-
Ost-Abhang der Cordillera von Peru, Aug
dad Nativa San Pedro de Pichanaz, Sector
1854, Lechler 2654 (lectotype: B).
Azulis, 102644S, 750621W, 910 m,
Reference: Tryon & Stolze, 1989. 15 Sep 2005, A. Monteagudo, F. Pea A. &
Remarks: Probably conspecic with C.
E. Quintuya 9810 (holotype: MO). (Fig.1)
heliophila R. M. Tryon.
Ab Alsophila erinacea (H. Karst.) D. S. Conant et
Alsophila cuspidata (Kunze) D. S. Conant squamis
Cyathea werfi R. C. Moran
petiolorum lanceolatis atrobrunneis costisque abaxia-
Depts. Loreto, Pasco. 120890 m. (Ecua- liter abundante et persistente pilosis (vs. squamis
dor, Peru) linealo-lanceolatis pallide brunneis costisque abaxia-
Voucher: Loreto: Iquitos, 120 m, 10 Apr liter glabrescentibus vel parce pilosis), ab Alsophila
1934, Williams 8236 (F). engelii R. M. Tryon et Alsophila minervae Lehnert in
References: Moran, 1991. presentia pilorum persistentium indusiis (vs. absentia
pilorum persistentium indusiis) absentiaque alarum in
parte distale pinnarum differt.
Cyathea windischiana A. R. Sm.
Dept. Amazonas. 800 m. (Peru) Trunks to 4 m high, 4 cm diam., otherwise
Voucher: Amazonas: Distr. Bagua, Cerro unknown. Fronds of unknown dimensions,
Tayu, ca. 1 hour from Chiriaco, 051556S, presumably 23 m long, patent to arching.
782207W, 800 m, van der Werff et al. Petioles 2756 cm long, basally dark brown
16207 (holotype: UC; isotype: MO) to blackish, distally paler brown to dark
References: Smith, 2006. 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 scurf squamules (Fig. 1B) and marginate
skeletonized basal pinnae absent. Petiole scales squaminate spines to 6 mm long, black, with
to 7 x 2 mm, lanceolate to ovate-lanceolate, broad bases and acuminate, curved tips. Lam-
with differentiated margins of differently ori- inae to ca. 130 cm broad, broadly ovate,
entated cells, dark brown with somewhat paler bipinnate-pinnatid, opaque dark green adax-
margins with only one dark apical setae to ially, opaque pale green abaxially, apices
1 mm, no or few marginal setae, concolorous abruptly reduced, non-conform to conform,
(Fig. 1A), of variable texture, larger scales i.e., pinna-like (Fig. 1C). Rachises and costae
papery, smaller ones rm, intergrading with brown to dark stramineous, inermous, abaxially
2011] LEHNERT: CYATHEACEAE OF PERU 31

with fugacious attish brown squamules to USM); ibid., 102934S, 750630W, 520640 m, 23
0.5 mm long, white pluricellular hairs to Sep 2005, Monteagudo A. et al. 10127 (MO); Dist.
Palcaz, Reserva Comunal Yanesha, Comunidad Nativa
1.5 mm long at least on distal parts, adaxially Lomalinda-Laguna, Sector Nueva Aldea, 102338S,
more abundant than abaxially, insertions of 750558W, 920950 m, 12 Oct 2005, Monteagudo A.
costae into rachises slightly raised abaxially, et al. 10400 (MO, USM).
bearing a lunular, at to concave blackish The trunk morphology was not described on
pneumathode to 52.5 mm. Pinnae to 65 cm the specimen labels but presumably is like in
long, stalked to 1.5 cm, not green-alate distally; the majority of Neotropical Alsophila species,
basal ones reduced, more ore less patent. i.e., densely covered with thin lanceolate scales
Largest pinnules to 10.51.8 cm, sessile, and without persistent old petiole bases.
truncate basally, attenuate apically (Fig. 1D); Among its Andean congeners, the species
costules stramineous to tan abaxially and is similar and supposedly related to Alsophila
adaxially, with at brown, setate scales to engelii R. M. Tryon and A. minervae Lehnert
1 mm long, no bullate scales present, with but differs from these in having long, rela-
white to tan pluricellular hairs to 1.5 mm long tively straight, presistent hairs on both sides
abaxially and adaxially, abaxially denser and of the midveins and on the indusia. The other
less spreading than abaxially (Fig. 1E); seg- two species have no hairs on the indusia and
ments to 102.5 mm, adnate, sessile, weakly on the midveins adaxially, but may have hairs
ascending, straight (Fig. 1E) to weakly falcate, or trichomidia on the lateral veins abaxially
the margins subentire to weakly crenate basally where A. gastonyi is glabrous; Alsophila
in the proximal segments, at in sterile and engelii has appressed tortuous hairs and thin
fertile segments; midveins weakly protruding squamules on the leaf axes abaxially but no
adaxially and adaxially, with white pluricellular spreading straight hairs as they occur in the
hairs identical to those on the costules (Fig. 1E), same places in A. gastonyi in varying density.
fertile veins forked once, rarely twice, sterile Further, A. gastonyi commonly has abruptly
ones forked to simple (Fig. 1E), hardly discern- reduced (but usually not pinna-like) laminar
able adaxially, dark gray abaxially, glabrous on apices and occurs below 1000 m, whereas A.
both sides (Fig. 1E). Sori to 0.8 mm diam., 13 engelii and A. minervae have gradually reduced
per segment, costal, in vein forks (Fig. 1E); laminar apices and occur mainly above 1500 m.
indusia urceolate to subsphaeropteroid, either Using the general key to the genus (Conant
irregularly or not rupturing at maturity, rm, 1983), Alsophila gastonyi will key out to A.
lustrous dark brown to tan, with persistent portoricensis (Spreng. ex Kuhn) D. S. Con-
pluricellular hairs to 0.6 mm long (Fig. 1E), ant, a species restricted to Puerto Rico, and
receptacles 0.20.3 mm diam., columellar, both populations may well be united under
black, paraphyses much shorter than the spor- one species morphologically. Such a biogeog-
angia, 0.10.2 mm long, blackish to tan. Spores rapical pattern would be unequalled among
not examined. Neotropical ferns. Other tree ferns that span
Distribution and habitat.Known only such distances do so in a more or less
from Peru, growing in the selva central continuous population (e.g., C. microdonta
region in the Dept. Pasco and the Ro Ene and C. delgadii). Either this Alsophila species
valley in the Dept. Cuzco at 520950 m. has been left undetected in most of its range,
Etymology.The name honors Gerald J. or, a hypothesis favored here, A. gastonyi and
Gastony in acknowledgment of his contribu- A. portoricensis are an example of convergent
tions to pteridology. evolution.
Additional specimens examined. PERU. Cuzco: La
Convencin, Echarati, Kapiromashi, 750 m, 1630 Apr
2004, Salinas N. et al. 6743 (USM). Pasco: Oxapampa, Cyathea chontilla Lehnert, sp. nov. Type:
Dist. Palcaz, San Pedro de Pichanaz-Azulis, Reserva PERU. Cajamarca: San Ignacio, Huarango,
Comunal Yanesha, 102840S, 750610W, 670 m, 25 localidad Romerillo, Cordillera entre
Feb 2004, Mellado L. F. 711 (MO), ibid., 25 Feb 2004, Romerillo y Nuevo Mundo, 0514S, 78
Mellado L. F. 721 (MO); Dist. Palcaz, Reserva
Comunal Yanesha, Comunidad Nativa San Pedro de 46W, 2300 m, 29 Jul 1992, J. Campos,
Pichanaz, Sector Azulis, 102644S, 750621W, Rodrguez E. & S. Nuez 4294 (holotype:
910 m, 12 Sep 2005, Monteagudo A. et al. 9668 (MO, 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 33

Species pygmaea generis Cyatheae squamis anguste oblong to oblanceolate, subsessile (Fig. 2C) to
lanceolatis cupreis, laminis triangularibus gradatim sessile (Fig. 2D), ca. 0.7 cm between the
reductis, rhachidibus sparse vel abundante squamatis, midveins, cuneate to weakly cordate at base,
indusiisque sphaeropteroideis. A Cyathea patente H. tips obtuse to rounded, patent, weakly ascend-
Karst. statura minore, a Cyathea sylvatica Lehnert ing, straight to distally falcate, margins shal-
laminis subcoriaceis (vs. chartaceis), squamis petiolorum lowly to deeply crenate with entire lobes, often
majoribus (usque 10.02.5 mm vs. 12.020.02.0 revolute when dried; segments with few whitish
3.5 mm) differt. to tan, erect hairs 0.50.8(1.0) mm long on the
veins abaxially, restricted to the midveins/
Trunks absent, rhizomes short-ascending, costules adaxially, none between the veins; with
branching in posterior parts; apices hidden in few to many dark-brown scales on the mid-
fascicles of petioles (Fig. 2A). Fronds 45 veins/costules abaxially, ovate-lanceolate, at
110 cm long, erect (Fig. 2A). Petioles 15 or bullate, to 2.00.8 mm with weakly dentate-
36 cm long, dark-brown to blackish, weakly lacerate margins; veins weakly protruding
shiny, inermous; scurf brown, dense, consisting abaxially, planar or sunken adaxially, sterile
of variously sized, erect to appressed, ovate to veins forked or simple, fertile veins forked. Sori
lanceolate squamules with mbriate margins to 1.01.2 mm diam., subcostal to inframedial,
and brown tortuous hairs to 1 mm long; without on the back of vein (Fig. 2D); indusia sphaer-
pneumathodes; densely scaly proximally, mod- opteroid, with prominent umbo, shiny dark
erately to sparsely so in distal parts. Petiole brown, translucent; receptacles globose, 0.3
scales to 12.020.02.03.5 mm, lanceolate to 0.4 mm diam, paraphyses thin, hyaline, tan to
narrowly lanceolate (Fg. 2B), shiny, concolo- brown, of the same length as the sporangia
rous or weakly bicolorous, deep brown with (0.4 mm). Spores ca. 0.04 mm diam., exospore
margins yellowish to cream white, the tips verrucate, perispore lacking (Young 5236,
elongate, undulate, not twisted, differentiated USM).
margins narrow with cells strongly exerted, Distribution and habitat.Known only
shortly dentate, without setae or cilia. Laminae from Peru, Depts. Cajamarca and Pasco, grow-
to 30852535 cm, bipinnate to bipinnate- ing in ridge-top vegetation at 23002850 m.
pinnatid, long-triangular to lanceolate Etymology.The name is a diminuitive of
(Fig. 2A), coriaceous, dark green adaxially, chonta, a vernacular name for tree ferns in
black when dried, gray-green abaxially, apices northern Peru, referring to the small size of
gradually reduced, tapering. Rachises inermous, the plant.
dark brown to castaneous, densely hairy on both
sides but glabrescent, with brown, twisted,
Additional specimens examined. PERU. Pasco:
pluricellular hairs to 1.0 mm long, also with Prov. Oxapampa, Dist. Oxapampa, Chacos (indumen
scattered, lanceolate to linear scales to 4 mm telefonica), 1037S, 7517S, 2850 m, 21 Sep 2003,
long. Pinnae to 17 cm long, weakly ascending, Rojas R. et al. 1487 (MO, UC); Cerro Pajonal, SE of
stalked to 5 mm, mostly alternate (Fig. 2A), Oxapampa, 2800 m, 29 Jul 1996, Young K. R. 5232,
5236 (USM).
fully pinnate only in proximal halves, narrowly
green alate distally; basal pinnae about the size
of the medial pinnae, not reexed, sometimes Cyathea chontilla probably belongs to the
opposite. Costae dark-brown to castaneous, C. patens group, which is characterized by
inermous, adaxially densely hairy with whitish mbriate laminar scales and scurf containing
to tan, antrorsely curved, pluricellular, uniseri- tortuous hairs. This group of sphaeropteroid-
ate hairs to 1.0 mm long, abaxially glabrescent indusiate tree ferns is typical of the upper
with sparser twisted hairs and few concolorous mountain rain forests from Costa Rica
brown, lanceolate scales with whitish, shortly through the Andes, and includes other small
mbriate margins, most frequently in the axils; species. Cyathea parvifolia Sodiro (Lehnert,
junctures of rachises and costules not swollen, 2009a) can be distinguished from C. chontilla
each abaxially with a circular, prominent by having longer petiole scales, stronger
pneumathode to 1 mm diam., dark gray to black dissected pinnules, and the regular presence
in dried specimens. Largest pinnules/ultimate of adventitious pinnae at the petiole bases.
segments (10-)20.040.0(4.0)8.014.0 mm, Cyathea sylvatica Lehnert can be distin-
34 BRITTONIA [VOL 63

guished from C. chontilla by less indument basally blackish, aculeate to muricate, spines
abaxially on the laminae and smaller, con- with scales on their tips in young petioles;
colorous petiole scales. Cyathea chontilla has with long elliptic brown pneumathodes to
also more hairs abaxially on the veins than 181 mm on the sides basally; scurf persis-
adaxially (vice versa in C. sylvestris) and tent (Fig. 3A), white to tan, akey, consisting
fringed margins on the laminar squamules of spaced, appressed, round squamules to
(vs. margins mostly entire). 0.5 mm diam. with tortuous marginal cilia.
Omitting the presence of tortuous hairs, Petiole scales 25.035.0(40.0) 2.54.0(
the similarity of Cyathea chontilla to C. 5.0) mm, long-lanceolate, shiny, plain deep
lechleri becomes more evident. Most speci- brown, concolorous or concordantly bicolo-
mens of C. lechleri represent plants 4 m or rous with margins paler brown, sometimes
smaller, which characteristically lack hairs yellowish or cream white (Fig. 3A); scales
on the veins and laminar squamules almost persistent only on the sides of the petioles.
completely. Larger fertile plants (to 12 m) Laminae to 1607080 cm, bipinnate-pinna-
may have a similar amount of pluricellular tid, green to dark green adaxially, often
hairs and lanceolate, at to weakly bullate blackish when dried, pale gray-green abax-
castanous squamules as found in C. chon- ially, apices gradually reduced. Rachises
tilla. Both species are an example for the inermous or weakly muricate basally, dark
often-observed elevational zonation in spe- yellowish brown to orange-brown, adaxially
cies groups of Cyatheaceae: Cyathea chon- with whitish to yellowish, antrorsely curved
tilla is clearly a high-elevation counterpart hairs to 0.8 mm long, abaxially glabrous or
to C. lechleri, which occurs mainly below glabrescent with white to tan squamellae like
1500 m. in the petiole scurf. Pinnae to 45 cm long
The examined spores (Young 5236, USM; (Fig. 3B), patent to ascending, basal pinnules
comm. Blanca Len) lacked a perispore layer, only half the size of the medial pinnae,
which may be a species-dening character but weakly to strongly reexed. Costae and
which also could reect just an early devel- costules tan, stramineous or dull orange-
opmental stage. The overall spore morphology brown, inermous, short-hairy adaxially, hairs
of Cyathea chontilla agrees well with C. to 1 mm long, white, antrorsely curved;
aterrima (Hook.) Domin (Moran et al., 2008). abaxially glabrescent with white to tan arach-
noid scurf and few bright orange-brown
Cyathea nephele Lehnert, sp. nov. Type: PERU. attish scales with undulated, mbriate mar-
Pasco: Prov. Oxapampa, Distrito Huancabamba, gins, most frequently in the axils. Largest
Parque Nacional Yanachaga-Chemilln, 1022S, pinnules 5.06.51.11.7 cm, linear-oblong
7528W, 3110 m, 5 Dec 2003, A. Monteagudo, to lanceolate, sessile (Fig. 3C) to subsessile
J. Perea, C. Mateo, G. Ortiz, & R. Francis 6430 (stalked to 1 mm), 1.01.5 cm between the
(holotype: UC; isotype: MO). (Fig. 3) stalks, pinnule bases truncate to weakly
cuneate, rarely weakly cordate, tips long-
A Cyathea frigida (H. Karst.) Domin acute to attenuate, pinnatid to pinnatisect,
squamis petiolorum longioribus lanceolatis basal segments sometimes remote but not
brunneis pallidioribus bicoloribus sive con- free; segments to 113 mm, weakly ascend-
coloribus (vs. squamis ovato-leanceolatis ing, falcate, the tips obtuse (Fig. 3D) to
bicoloribus atro-brunneis usque castaneis round, segment margins crenulate to crenate,
cum marginibus angustissimis pallide brun- often remaining at when dried; segments
neis vel albidis), a C. patente H. Karst. with few to many whitish to tan, erect hairs
absentia indusiorum (vs. indusiis sphaeropter- 0.50.8(1.0) mm long on and between the
oideis) differt. veins abaxially, veins adaxially glabrous or
Trunks to 8 m tall, 810 cm diam., without with few short hairs, none between the veins;
old petiole bases, frond scars round, apices with relatively few to many small scales
hidden in fascicles of petioles; adventitious abaxially, ranging from broadly ovate, at
buds lacking. Fronds 24 m long, presum- ones (to 3.02.5 mm) to bullate ones (to
ably arching. Petioles to 40 cm long, prob- 2.51.0 mm) with broadly aring or twisted,
ably longer, dull brown to orange-brown, 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, Distribution and habitat.Frequent in
mbriate to lacerate with considerably long central Peru, Dept. Pasco, rare in the north,
processes; sterile veins forked or simple, Prov. Amazonas and San Martin, in upper
fertile veins forked. Sori to 1.0 mm diam., montane forest and ridge top dwarf forests at
subcostal (Fig. 3D), in forks of veins; indusia 27203550 m.
absent; receptacles globose to ellipsoid, 0.3 Etymology.Named after the Greek god-
0.4 mm diam, paraphyses thin, hyaline, white dess of the clouds (Greek, nephos=cloud) in
to tan, of the same length as or shorter than reference to the cloud forests this species
sporangia (0.30.4 mm). Spores not examined. inhabits.
36 BRITTONIA [VOL 63

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

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

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

indusia would have necessitated its trans- bases truncate to weakly cordate, tips long-
ference to Cyathea as dened by Tryon (1970, acute to attenuate, pinnatid to pinnatisect,
1976) despite the fact that both genera had basal segments sometimes remote, connected
already been merged to Cyathea (Lellinger, by thin strands of laminar tissue; segments to
1987). The new combination Cyathea serpens 9.015.02.55.0 mm, weakly ascending,
was already published in the comparison with straight to distally falcate, the sinuses acute
a newly described species (Lehnert, 2009b). to obtuse, to 15 mm wide, the tips obtuse to
An amended description of C. serpens seems rounded, segment margins crenulate to
appropriate in the context of its new taxo- inciso-dentate, the basiscopical margins of
nomic status and the additional collections that the lowest segments often more strongly
brought new information of about its morphol- incised; costules planar to weakly protruding
ogy and distribution. abaxially, ridged adaxially, yellowish to dark
Trunkless; rhizome apices hidden in fas- carnose, with many curved brown hairs to
cicles of petioles; adventitious buds not 1.2 mm long adaxially, glabrous abaxially,
reported and presumably lacking. Fronds to with broadly ovate, at squamules to 3.0
7 m long, pendent from banks or trailing over 2.5 mm, and bullate squamules to 2.5
adjacent vegetation. Petioles 70200 cm 1.0 mm, with mbrate tips, tan to orange-
long, probably longer, shiny, castaneous to brown, often with the thin margins paler
atropurpureous, basally blackish, inermous, brown to whitish, mbriate to lacerate; seg-
with dense, matted, white scurf consisting of ments with few to many deciduous, whitish,
appressed, strongly dissected squamules and contorted, catenate hairs to 2.0 mm long,
tortuous marginal hairs, often abraded in mainly abaxially and along the margins; veins
older material; without pneumathodes, only planar to weakly immersed, yellowish to
scatteredly scaly in lower half. Petiole scales carnose, adaxially with few brown, erect hairs
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 abaxially glabrous or with few trichomidia;
or blackish with white margins (Fig. 6A); sterile veins forked or simple, fertile veins
differentiated margins to 0.6 mm wide, cell forked. Sori to 1.2 mm diam., subcostal to
rows strongly exserted, without cilia or setae. costal, in fork of veins (Fig. 6C), yellowish
Laminae 15050070120 cm, long-triangu- brown to dark orange-brown at maturity;
lar to lanceolate, bipinnate-pinnatid to partly indusia hemitelioid, small, appressed, tan to
tripinnate, green to dark green adaxially, often dull brown, with entire margins (Fig. 6D) or
becoming blackish when dried, pale gray- with one or two marginal cilia, fragile and
green abaxially, apices gradually reduced. evanescent; receptacles globose, 0.40.5 mm
Rachises inermous, dark castaneous to atro- diam. (Fig. 6D), paraphyses thin, hyaline, white
purpureous, densely pubescent with whitish to tan, longer than sporangia (0.40.5 mm).
to tan, cobwebby hairs to 2 mm long, Spores tetrahedral-globose, dark orange-brown.
abaxially often glabrescent. Pinnae to 35 cm Distribution and habitat. Locally com-
long, patent (Fig. 6B), alternate to suboppo- mon in open forests and pajonal-vegetation in
site, stalked to 2.53.0 cm, distally green central and southern Peru at 24502900 m.
alate, the distal segments not decurrent into Etymology. The epithet refers to the
the costae; basal pinnae nearly the size of the trailing fronds.
medial pinnae, weakly reexed. Costae and
costules dark castaneous to atroprurpureous, Additional specimens examined. PERU. Pasco:
inermous, to 2 mm broad, adaxially with tan Prov. Oxapampa, Dist. Oxapampa, sector Chacos, 10
3725S, 751743W, 2750 m, 3 Feb 2004, Mellado et
to brown, antrorsely curved, pluricellular, al. 1664 (MO, UC); Dist. Huancabamba, sector Que-
uniseriate hairs to 1.5 mm long, abaxially brada Yanachaga (P. N. Yanachaga-Chemilln), 1022
glabrescent with white to tan scurf and few 46S, 752743W, 2910 m, 14 Feb 2004, Mellado &
bright orange-brown attish scales with undu- Monteagudo 464 (MO, UC); Prov. Oxapampa, San
lated, mbriate margins, most frequently in Gotardo, 36 km W of Oxapampa, 1037S, 7509W,
27102800 m, 26 Jan 1984, Smith D. N. & Canne 5902
the axils. Pinnules to 5.510.51.53.0 cm, (MO, UC); Prov. Oxapampa, near antenna past Chacos,
long triangular to lanceolate, stalked to 4 mm 1038S, 7515W, 2450 m, 19 Jul 2003, van der Werff et
(Fig. 6B, C), 1.52.0 cm between the stalks, al. 18647 (MO, UC).
2011] LEHNERT: CYATHEACEAE OF PERU 43

Cyathea serpens is very similar to C. helio- EXCLUDED AND DUBIOUS NAMES


phila R. M. Tryon regarding the overall
appearance of the fronds; it can be distinguished Alsophila capensis (L. f.) J. Sm. ssp. polypo-
by the lack of a trunk (vs. C. heliophila with
dioides (Sw.) D. S. Conant, J. Arnold Arbor.
trunks to 7 m tall), bicolorous brown-white
64: 369. 1983.
petiole scales (vs. predominantly concolorous
whitish), the contorted hairs on the laminae
Remarks: Tryon (in Tryon & Stolze 1989)
(vs. contorted hairs absent), and the long
reports that this species may occur in Peru,
paraphyses (vs. paraphyses of the same length
without specifying the reasons for his
as the sporangia or shorter than those). The assumption. Alsophila capensis occurs in
indusia of C. serpens are very thin, appressed
small, presumably relict populations on
to the lamina and often caduceous, which may
shaded sandstones in the Atlantic Brazilian
lead to the assumption that the species is rainforest (Paulo Labiak, pers. comm.). It is
exindusiate. Cyathea heliophila has always
characterized by inermous petioles with
clearly recognizable, persistent indusia. The
highly dissected aphlebiae at their bases.
evanescence of the indusia of C. serpens may
The only Peruvian tree fern species with
be an indicator of a hybrid origin of this
such aphlebiae is A. salvinii, which can be
species, with the sympatric C. heliophila
distinguished by the lack of indusia and the
(indusia rm) and C. frigida (indusia lacking)
tripinnate fronds (vs. hemitelioid indusia
as putative parents. Traits of C. frigida would
present, fronds bipinnate-pinnatifd in A.
be the long paraphyses, the brown color in the
capensis).
petiole scales, and the tendency not to form real
trunks.
Cyathea serpens has unusually long fronds Cyathea petiolata (Hook.) R. M. Tryon,
that reach 7 m. However, these expanses are Contr. Gray Herb. 206: 42. 1976.
only known from the type collection; the other
collections barely reach half that length. The Remarks: Tryon (in Tryon & Stolze 1989)
only other neotropical tree fern with similarly reports this species from Peru. It is cur-
long fronds is C. kalbreyeri (Baker) Domin rently known only from Colombia, Vene-
which grows in the understory of lower zuela, and Central America but I have seen
montane forests and supports its fronds on no material from Peru or Ecuador that
high branches of adjoining trees. Confusion could be condently assigned to this spe-
with C. serpens is impossible because besides cies. It is easily recognized by its bipinnate
growing at a completely different elevation, fronds with large, almost entire pinnules and
C. kalbreyeri also lacks indusia and scurf, and submarginally positioned sori with hemitelioid
has concolorous orange-brown petiole scales. indusia.

Trichipteris ava R. M. Tryon, Fieldiana Bot.


n.s. 20: 126. 1989. Type: PERU. Hunuco:
NEW COMBINATIONS
Above Chinchao, Tingo Mara to Chin-
Cyathea alatissima (Stolze) Lehnert, comb. chao, 1 Aug 1965, Soejarto 1432 (holo-
nov. Basionym: Cnemidaria alatissima type: GH). Cyathea herzogii Rosenst.,
Stolze, Fieldiana, Bot. 37: 55. 1974. Meded. Rijks. Herb. Leiden 19: 7. 1913.
Cyathea nervosa (Maxon) Lehnert, comb.
nov. Basionym: Hemitelia nervosa Maxon, J. Remarks: This species was described from
Wash. Acad. Sci. 34: 309. 1944. Cnemidaria a sterile collection. The laminar indument,
nervosa (Maxon) R. M. Tryon, Contr. Gray which consists of a dense, short pubescence
Herb. 200: 52. 1970. on and between the veins adaxially and
Cyathea uleana (A. Samp.) Lehnert, comb. strongly bullate squamules abaxially, in com-
nov. Basionym: Hemitelia uleana A. Samp., bination with the dull brown petiole scales is
Bol. Mus. Nac. Rio de Janeiro 1: 65.1923. indicative that the specimen belongs to
Cnemidaria uleana (A. Samp.) R. M. Tryon, Cyathea herzogii, which is quite common in
Contr. Gray Herb. 200: 52. 1970. the central Andes.
44 BRITTONIA [VOL 63

Acknowledgments Holttum, R. E. 1963. Cyatheaceae. Flora Malesiana, ser.


II, vol 1. 1176.
I thank Michael Kessler and Alan R. & P. J. Edwards. 1983. The tree ferns of Mount
Smith, who thought studying tree ferns was Roraima and neighbouring areas of the Guyana
Highlands with comments on the family Cyatheaceae.
a good idea. They were right. I further thank
Kew Bulletin 38: 155188.
my colleagues Alexander N. Schmidt-Lebuhn Janssen, T., F. Bystriakova, F. Rakotondrainibe, D.
(Z) and Johana Toivonen. I am indebted to Coomes, J.-N. Labat & H. Schneider. 2008. Neo-
the curators of AAU, B, BM, COL, CUZ, F, endemism in Madagascan tree ferns results from
GH, GOET, HUT, LIL, MBM, MO, P, QCA, recent, coincident diversication bursts. Evolution
62: 18761889.
QCNE, S, SCZ, SP, QPLS, UC, USM, Korall, P., K. M. Pryer, J. Metzgar, H. Schneider & D.
UPCB, and Z for providing loans or for S. Conant. 2006. Tree ferns: monophyletic groups
attending me during my visits, especially to and their relationships as revealed by four protein
Asuncin Cano (USM). Hanna Tuomisto coding plastic loci. Molecular Phylogenetics and
(TUR) and Maarten Christenhusz (BM) Evolution 39: 830845.
Korall, P., D. S. Conant, J. Metzgar, H. Schneider &
kindly provided additional specimens. I thank K. M. Pryer. 2007. A molecular phylogeny of scaly
Dave Conant (LSC) and Blanco Len (TEX, tree ferns (Cyatheaceae). American Journal of Botany
USM) for helpful comments on the manu- 94: 873886.
script. Special thanks go to Blanca Len for Kramer, K. U. 1978. The Pteridophytes of Suriname.
Uitgaaven Naturw. Studiekring Suriname Ned. Antil-
additional data that supports this treatment. len 93: 198.
Field studies that contributed to the presented & P. S. Green. (volume editors)1990. Pterido-
results were funded by the DAAD (German phytes and Gymnospermes. In: K. Kubitzki (editor).
Academic Exchange Service) and the DFG The Families and Genera of Vascular Plants. Vol. 1.
(German Research Foundation, grant GR Springer-Verlag, Berlin.
Lehnert, M. 2003. Six new tree ferns from the Andes.
1588/7). This research received support from American Fern Journal 93: 169183.
the SYNTHESYS Project (http_//www.syn . 2005. Reconsideration of Alsophila odonelliana
thesys.info/), which is nanced by Eurpoean (Cyatheaceae), an Andean tree fern. Brittonia 57:
Community Research Infrastructure Action 228236.
. 2006a. Two new tree ferns (Cyatheaceae) from
under the FP6 Structuring the European southern Ecuador. Brittonia 58: 49.
Research Area Programme. . 2006b. The Cyatheaceae and Dicksoniaceae
(Pteridophyta) of Bolivia. Brittonia 58: 229244.
. 2006c. New species and records of tree ferns
(Cyatheaceae, Pteridophyta) in the northern Andes.
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