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SERIES II — PTERIDOPHYTA
Ferns and Fern Allies

Volume 2, part 1: Tectaria Group

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THE NEW YORK BOTANICAL GARDEN

BRONX, NEW YORK 10454

FLORA MALESIANA

SERIES Il — PTERIDOPHYTA

Volume 2

Published for Foundation Flora Malesiana
by Riksherbarium/ Hortus Botanicus « Leiden University « The Netherlands

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FLORA MALESIANA

SERIES II — PTERIDOPHYTA

Volume 2 — part 1 — 1991

Tectaria Group
by
R.E. Holttum +

CIP-GEGEVENS KONINKLIJKE BIBLIOTHEEK, DEN HAAG
Flora

Flora Malesiana / founded by C.G.G.J. van Steenis

(1901-1986). — Leiden : Rijksherbarium / Hortus Botanicus

Series II: Pteridophyta : Ferns and fern allies

Vol. 2, pt. 1: Tectaria group / by R.E. Holttum. — Ill.

Comp. and publ. under the auspices of Foundation Flora Malesiana
Met index.

ISBN 90-71236-11-0

Trefw.: varens.

All rights reserved
© 1991 Foundation Flora Malesiana

No part of the material protected by this copyright notice may be reproduced or utilized
in any form or by any means, electronic or mechanical, including photocopying,
recording, or by any information storage and retrieval system, without written permis-
sion from the copyright owner.

Flora Malesiana ser. II, Vol. 2! (1991) 1-132

TECTARIA GROUP (R.E. Holttum +, Kew)*

Polypodiaceae subfam. Dryopteridoideae section A, auct.: C. Chr. in Verdoorn, Man.
Pteridol. (1938) 543, p.p.

Aspidiaceae tribe Aspidieae auct.: Ching, Sunyatsenia 5 (1940) 250, excl. Lomariopsis
and related genera. — Aspidiaceae, group of Ctenitis Copel., Gen. Fil. (1947) 153.

Aspidiaceae auct.: Pichi Sermolli, Webbia 31 (1977) 460—468, p. p.

Dennstaedtiaceae subfam. Tectarioideae Holttum, J. Linn. Soc. Bot. 53 (1947) 152;
Revis. Fl. Malaya 2 (1955) 494; Studies fern genera I, Fern Gaz. 12 (1984) 313-319;
Ibid. VI, Gard. Bull. Sing. 39 (1986) 153 —167.

Caudex in almost all cases erect or suberect, always with a radially organized dictyo-
stele; scales narrow and rather firm, marginal teeth (if present, except in Cyclopeltis) al-
most always formed at the junction of two cells; i.e., the wall separating the two adjacent
cells is elongated at right angles to the margin. Vascular structure of stipe consisting of
small subequal vascular strands arranged in a U-formation as seen in transverse section
(except in Pleocnemia where the arrangement is more complex), the two on the adaxial side
variously enlarged. Fronds amply divided with free veins, or with broader divisions in
which veins anastomose variously, in a few species simple, in most (except Heterogoni-
um) the basal pinnae or lobes longest with elongate basal basiscopic lobes or pinnules;
costae or costules of ultimate lamina-elements + prominent on the upper surface, branch-
ing directly from the prominent upper surface of the rachis bearing them, the margins of
the lamina decurrent on the sides of the rachis. Hairs consisting of several cells, not inter-
grading with scales, variously present on most parts of the frond, the prominent upper
surface of costae of lamina-segments in a majority of cases (at least near the base) covered
with short erect hairs of several cells, the outer cell-walls thin and collapsing on drying (in
other cases hairs in this position may remain firm on drying); unicellular glands usually
+ cylindric, colourless or red to yellow, variously present on surfaces of leaflets, on
indusia, and on stalks of sporangia in some genera. Sori usually orbicular, covered with
reniform or peltate indusia, or exindusiate, in the latter case sometimes spreading along
veins. Fertile pinnae or leaflets in some species much contracted, with sporangia borne all
along the veins to produce an acrostichoid appearance. Spores in most cases with a peri-
spore folded into wings or crests, the wings variously anastomosing, the perispore in other
cases spinulose or with obtuse projections. Base chromosome number 40 or 41.

Distribution — Pantropic with greatest diversity in the Old World; about 20 genera.

Taxonomy — In the 19th century the form of sori and condition of venation (free or
anastomosing) were by most authors regarded as the most important bases for classifica-
tion. Thus the free-veined members of the present group were associated with Dryopteris
and free-veined Thelypteridaceae, and exindusiate species were separated generically.
This led to confusions of various kinds.

*) Dr. Holttum submitted the manuscript of his revision of the Tectaria Group for publication in Flora
Malesiana not long before he passed away in September 1990. Some distributional data were added by
the Editor. The illustrations are by P.J. Edwards, Kew.

2 Flora Malesiana ser. II, Vol. 2! (1991)

The first major work in the 20th century was Christensen’s subdivision of the unnatu-
ral aggregate of species under Dryopteris in his Index Filicum (1905) which included all
free-veined ferns with reniform indusiate sori and their obvious exindusiate relatives. For
the first time he distinguished Crenitis from Dryopteris, pointing out the relationship of the
former to Tectaria and also the distinctive characters of thelypteroid ferns (some of which
have anastomosing veins). In his last major survey of the classification of all ferns (1938)
he retained a major family Polypodiaceae with Dryopteridoideae as one subfamily; this
included, in section A, Dryopteris and Tectaria with other genera related to them, with
Thelypteridaceae in section B.

In 1940 Ching divided Christensen’s Polypodiaceae into several families, one being
Aspidiaceae, similar to Christensen’s Dryopteridoideae section A of 1938. Ching divided
it into two tribes, Aspidieae and Dryopterideae. In 1947 Holttum accepted Ching’s two
tribes, with minor changes, but placed them as subfamilies in a major family Dennstaed-
tiaceae, with the idea that Polypodium and its immediate allies form a very different group
of ferns. Copeland, writing independently in 1947, recognized a large family Aspidiaceae,
including Thelypteris and its allies, also Athyrium/Diplazium and the Lomariopsis group
of genera. In his conspectus (1947: 153) the genera here dealt with appear as allies of
-Ctenitis; to them should be added Cyclopeltis which he associated with Polystichum, and
Dryopolystichum excluded. Pichi Sermolli (1977) included both dryopteroid and tectar-
ioid genera in his family Aspidiaceae and did not accept a division of it into two groups.

Holttum (1984) discussed the above history in more detail, still retaining separate
groups associated with Dryopteris and Tectaria and objecting to the Tryons’ arrangement
in their book of 1982 in which the two groups are confused. In 1986 Holttum presented a
conspectus of all Old World genera of the Tectaria alliance, with comments on those of the
New World, pointing out the peculiar characters of Dryopsis Holttum & Edwards [Kew
Bull. 41 (1986) 171—204] which perhaps connects the two groups. Most species of Dry-
opsis are in Mainland Asia but two are Malesian.

The structure of the junction of the bases of leaflets and the rachis that bears them is
here presented as the most clear distinction between the genera allied to Dryopteris and
those allied to Tectaria. This is seen best in Lastreopsis, Ctenitis and Pteridrys (see fig.
311 in Holttum 1955: 524). In Lastreopsis the relationship is exactly as in Davallia;
Ctenitis only differs in the margins of the lamina which are not thickened where they are
decurrent on the rachis-wing. In Cyclopeltis the relationship of pinna-base to rachis is
only evident near the apex of the frond. An arrangement close to Davallia is also shown
by Rumohra, a genus included by several authors in Aspidiaceae. But Rumohra differs
from all the genera here included in its strongly dorsiventral rhizome-structure, compar-
able to that of Davallia though not identical with it, and also in its scales. [Pichi Sermolli
(1977: 465) cites Chandra’s study of the vascular structure in the caudex of Maxonia as
evidence of dorsiventrality in a ally of Dryopteris; but the dorsiventrality in Maxonia is
only a slight modification of the normal dictyostele of Dryopteris, very different from that
in Rumohra.] Rumohra differs from Davallia in its distribution (widely in South America,
also in southern Africa and Australasia) and in its spores. In Malesia, it only occurs in
New Guinea. It is perhaps a connecting link between Davallia and the Tectaria group; it
needs a comprehensive new morphological study.

Holttum — Tectaria Group 3

Pichi Sermolli (1.c. 1977: 238) has stated that there are four different types of rachis-
structure in Aspidiaceae but he deals only with the junctions of main rachis with pinna-
rachises, not with the relationship between lamina-elements and the smaller distal rachises
which I believe to be significant. I have expressed disagreement with him in 1984 (l.c.:
314).

KEY TO THE GENERA

la. Teeth present at the bases of sinuses between pinna- or pinnule-lobes, the teeth pro-

jecunecout of the plane of the lamina: “iy spe paracniye eae ae ee eee enn eae 2

balecthros this kind lacking’. 2... Ss..cs0drw ss) som Gendt eax ete legious: eee nena akon obs eas 5)
2a. Fronds simply pinnate with free veins; no unicellular glands on lamina or in sori

Pteridrys (p. 4)

b. Fronds mostly bipinnate, their veins forming at least costal areoles; cylindric glands

[DIRECTS ee a En Men cr eee NSE oe RR ce tne ohn Eee Pleocnemia (p. 8)

3a. All axes of the frond bearing copious scales, the smaller ones at least partly clathrate
tS OGAIME II CuCOll Se fai. cus: secre chereee taueeceaer eee nce roe Ctenitis (p. 21)

b. Smaller axes of the frond bearing few scales which are not thus clathrate...... 4
4a. Veins in sterile fronds anastomosing copiously; free veinlets in areoles (including
those along costae) variously directed and in most species forked........... 5

b. Veins anastomosing to form costal areoles which lack free veinlets, or all veins
ERED, AS Sh Rae ae eae eae aia ee amen Oo ot BCA RRP MACE NEU NAC hi oy A pettic a a 7o% c i!

5a. Fertile fronds greatly contracted and bearing indusia................... 6

b. Fertile fronds not greatly contracted, or if so lacking indusia
Tectaria sect. Tectaria (p. 59)
6a. Sterile fronds simple; fertile ones irregularly deeply lobed; indusia reniform
Tectaridium (p. 36)
b. Sterile fronds pinnate; pinnae of fertile fronds linear, with continuous indusia along

Cachipside OfstherCOStac: aan a. cue cee eee Chlamydogramme (p. 37)

7a. Some thick multicellular hairs present between veins on the upper surface, at least
Ne ATECINUSESIDEEWEEN LOWES ae. a sis eer «alee ce ae ee eee 8

b. No thick hairs present between veins on upper surface.................. it
8a. Basal basiscopic lobe or pinnule of basal pinnae longer than the other lobes or
PLUME TLS Si oats e.Fy a: 4 <n) 5 no) ks Vent tar ck as SRP cot Ree A heated Pe no eee 9

b. Basal basiscopic lobe of basal pinnae much reduced, these pinnae widest at about
TMCS LEME thy eu, ns hes yet ka eee Heterogonium (p. 105)
Yamekronds not sreatly longer than. wide. 22.5. ...22.5.¢500.45.055050 06806 10
b. Fronds greatly longer than wide, with many pinnae gradually increasing in size
GOWINWATAS 3.152 Shs. be peleoretier teers aie Renee cde eer a re ate Aenigmopteris (p. 102)
10a. Veins anastomosing in some species; fertile fronds, if greatly contracted, not bear-
INSeVvery small pinmuUles: 2s 4 cis sey eee Tectaria sect. Sagenia (p. 42)

b. Veins all free; fertile fronds bipinnate with very small pinnules, exindusiate
Psomiocarpa (p. 100)
lla. Pinnae articulate to the rachis, entire or crenate ......... Cyclopeltis (p. 116)
b. Pinnae not articulate, deeply lobed or pinnate .......... Lastreopsis (p. 120)

4 Flora Malesiana ser. II, Vol. 21 (1991)

PTERIDRYS

Pteridrys C.Chr. & Ching, Bull. Fan Mem. Inst. Biol. 5 (1934) 129; Copel., Gen. Fil.
(1947) 126; Holttum,.J. Linn. Soc. Bot. 53 (1947) 153; Revis. Fl. Malaya 2 (1955)
529; Copel., Fern Fl. Philipp. (1960) 298; Pichi Sermolli, Webbia 31 (1977) 465. —
Type species: Pteridrys syrmatica (Willd.) C. Chr. & Ching.

Aspidium p.p., Nephrodium p.p., Lastrea p.p., quoad plur. auct.

Caudex short, erect or suberect; scales firm, narrow, entire with cordate base, multi-
cellular hairs present with the scales and variously on the fronds; fronds simple-pinnate;
external form of rachis and its relationship to pinnae as in Davallia; pinnae deeply lobed,
the lobes + crenate or dentate; veins all free, once or twice forked, the basal basiscopic
vein in each lobe springing from the base of the costule (rarely from the costa), the basi-
scopic branch of the basal acroscopic vein (or sometimes both branches) passing to a
tooth in the sinus between that lobe and the next, the tooth projecting out of the plane of
the lamina; unicellular glands absent on the lamina and from sporangium-stalks; sori on
the acroscopic branch of each vein terminal or not, indusiate, indusia reniform, sometimes
asymmetric. — Figs. 1, 2.

Distribution — A genus of 8 known species in SE Asia (including India and Sn
Lanka) and Malesia (6 species).

Cytology —n = 41 (Manton in Holttum 1955).

Note — This genus appears to be most nearly related to Pleocnemia, differing from the
latter in free veins and in the absence of unicellular glands. But in Pteridrys cnemidaria
(Christ) C. Chr. (from S China to NE India) there is occasional anastomosis of the basal
veins of adjacent pinna-lobes and the relationship between these and the teeth in sinuses
between lobes agrees with that in Pleocnemia. Apart from sinus-teeth, Pteridrys differs
from free-veined species of Tectaria in the position of the basal basiscopic vein in each
lobe which in Tectaria springs always from the costa.

KEY TO THE SPECIES

la. Basal basiscopic lobes of basal pinnae shorter than those next to them........ 2
b. Basal basiscopic lobes of basal pinnae elongate, longer than those next tothem.. 4
Paestalkstorbasalspinnae sc. *Oimmblong yy ayer ast lee 1. P. syrmatica
b. Stalks of basal pinnae much shorter, other pinnae mostly sessile............ 3

3a. Pinna-lobes broad and obtuse; costae bearing stiff short hairs at least near their bases
2. P. australis

by Binna-lobes) acute;;costae quitevslabrouse 93.4 eee 3. P. acutissima
4a. Pinnae mostly short-stalked, their basal acroscopic lobes not or little overlapping the
rachis; sinuses between pinna-lobes 2-3 mm wide ................00-- 5)

b. Pinnae (except the basal ones) quite sessile, their basal acroscopic lobes overlapping
the rachis; sinuses between pinna-lobes very narrow ....... 6. P. confertiloba

5a. Basal basiscopic lobes of basal pinnae distinctly longer than the next lobes; pinna-
lobes slightly toothed distally; basal acroscopic veins in larger pinnae twice forked

4. P. microthecia

Holttum — Tectaria Group 5

b. Basal basiscopic lobes of basal pinnae little longer than the next lobes; the pinna-
lobes strongly crenate-serrate almost throughout; basal acroscopic veins in larger
GHESHONCO MOPKCU Nl faust In eters ete tans Grates a ee Oe er roles 5. P. olivacea

PJ. EpwaARDS

Fig. 1. Pteridrys syrmatica (Willd) C. Chr. & Ching. a & b. Upper surface of rachis and bases of a) two
lower pinnae, and b) two upper pinnae, showing structure of the junction, x 8; c. basal basiscopic lobes (one
with sori removed), x 2; d. detail of c showing sinus tooth, x 9 (a—c: van Beusekom & Santisuk 2809).

1. Pteridrys syrmatica (Willd.) C.Chr. & (1960) 298. — Aspidium syrmaticum Willd. in
Ching, Bull. Fan Mem. Inst. Biol. 5 (1934) Linn., Sp. Pl. ed. 4, 5 (1810) 237; Racib.,
131, pl. 11, 17; Holttum, Revis. Fl. Malaya 2. _— Pteridop. Buitenzorg (1898) 171. — Lastrea syr-
(1955) 530, f. 312; Copel., Fern Fl. Philipp. matica (Willd.) T. Moore, Index Fil. (1858) 105;

Flora Malesiana ser. II, Vol. 2! (1991)

OT. Epwarns

Fig. 2. Pteridrys acutissima Ching. a. Base of basal pinna, showing the almost free pinnule, x 1. —
P. microthecia (Fée) C. Chr. & Ching. b. Base of a basal pinna, x 0.6. — P. olivacea (Rosenst.) Copel.
c. Base of a basal pinna, x 0.85. — P. confertiloba Holttum. d. Base of a middle pinna showing basal
lobes overlapping rachis, x 0.85 (a: Ridley 14742; b: Parris 6851; c: Bamler 103; d: Ledermann 7826).

Bedd., Ferns S. India (1863) t. 108; Handb.
Ferns Brit. India (1883) 243, f. 124. — Dryo-
pteris syrmatica (Willd.) Kuntze, Revis. Gen.
Pl. 2 (1891) 813; Christ, Philipp. J. Sci. 2
(1907) Bot. 210 (as var. petiolosa); Alderw.,
Malayan Ferns (1908) 193; Backer & Posth.,
Varenfl. Java (1939) 40. — Type: ‘Habitat in
Peru, Chile’ (B, herb. Willd. 19765).

Aspidium spectabile Blume, Enum. Pl. Javae (1828)
158. — Nephrodium spectabile (Blume) Hook.,
Sp. Fil. 4 (1862) 115, p.p. — Type: Blume
s.n., Java (L; iso K, P).

Polypodium oxyodon Baker, J. Bot. 17 (1879) 66. —
Type: Burbidge s.n., Sulu Archip. (K; iso BM).

Stipe to 40 cm long, pallid when dried, glabres-
cent, basal scales to 10 mm long; lamina to 40 cm
long; pinnae commonly to 10 pairs, largest c. 15
x 3 cm, abruptly narrowed at their bases to stalks
c. 10 mm long, their apices abruptly caudate-acu-
minate, the narrow part crenate, 3—4 cm long, rest
of pinna lobed to 3-4 mm from the costa, lobes 5
mm wide, oblique, slightly falcate, crenate at least
distally; veins usually once forked; very slender
hairs present on the lower surface of costae when
young, also appressed between veins; sori on acro-
scopic branches of veins, usually terminal, indusi-
ate; indusia thin, entire, pale olivaceous when dry,
often asymmetric at their bases. — Fig. 1.

Holttum — Tectaria Group

Distribution — § Thailand, Vietnam; in W Ma-
lesia: Celebes, Philippines.

Habitat — Often but not always on limestone,
otherwise on stream banks or sloping ground, in
forest; mainly in + seasonal areas.

Note — Hooker (1862) did not distinguish be-
tween this and allied species, citing specimens of
two others, under Nephrodium spectabile.

2. Pteridrys australis Ching, Bull. Fan Mem.
Inst. Biol. 5 (1934) 142, pl. 15, 16, 19, f. 12, 13;
Tard. & C.Chr., Fl. Indo-Chine 7, 2 (1941)
306; Holttum, Revis. Fl. Malaya 2 (1955) 532,
f. 313. — Type: S.P. Ko 50556, Kwangtung
(PE, not seen).

Stipe to 50 cm long; basal scales to 15 x 1.5 mm;
lamina dark olivaceous when dried, to 60 cm long;
pinnae to 20 pairs, basal ones a little shorter than
those next above them, narrowed at the base on the
basiscopic side, middle basiscopic lobes + elon-
gate, stalks to 2 mm long; suprabasal pinnae ses-
sile or nearly so, to 20 x 3.5 cm, lobed to 5 mm
from costae, costules to 8 mm apart, lobes obtuse,
slightly crenate distally, apices of pinnae short- to
long-caudate; veins mostly once forked (twice fork-
ed on largest pinnae); a dense tuft of short hairs
present on the lower surface of the base of each
costa (Sometimes also on the upper surface) and
a few at bases of costules, slender hairs to 1 mm
long + abundant on lower surface of rachis, costae
and rarely on the veins; sori terminal on acroscopic
branches of veins; indusia persistent, reniform, en-
tire, on Malesian specimens usually hairless (hairs
present on those of the type).

Distribution — S China and Burma southwards;
in Malesia: Peninsular Malaysia (Selangor).

Habitat — On stream banks in forest, sometimes
near waterfalls, at altitudes to 900 m.

3. Pteridrys acutissima Ching, Bull. Fan
Mem. Inst. Biol. 5 (1934) 138, pl. 13, 19, f. 15;
Holttum, Revis. Fl. Malaya 2 (1955) 531; Gard.
Bull. Sing. 38 (1986) 147. — Type: Haniff
14142, ‘G. Kerbau 6000 ft’ (SING; iso K).

Differing from P. australis as follows: supra-
basal pinnae to 4.5 cm wide, lobed to 3 mm from
costae, lobes + falcate, narrowed to an acute tip,
rather strongly crenate throughout; basal pinnae of

large fronds bearing a pair of pinnules, their largest
basiscopic pinna-lobes 5 cm long; hairs lacking or
very short and sparse at bases of costae. — Fig. 2a.

Distribution — Malesia: Malay Peninsula (Perak),
Bomeo (Sarawak) (one collection from each).

Note — The type was collected by Haniff on a
journey to Gunung Korbu in 1909. The collector’s
label was not preserved. Ridley assigned numbers
in his own series to all Haniff’s specimens and
wrote the locality cited for 1/4/42, indicating a
source near the summit of the mountain. The evi-
dence that the type was not from a high altitude
is as follows. The second collection, reported by
Holttum in 1986, was from ‘slope of bat guano in
cave mouth’ in a limestone area in Gunung Mulu
National Park, Sarawak. There is also a specimen
from a low altitude in Perak, consisting of basal
pinnae and the pair next above them; these are
about intermediate between P. australis and the
type of P. acutissima. The lobes of basal pinnae
are little different from those of P. acutissima; one
of the pair has one almost free basal lobe; hairs at
the bases of costae are like those of P. australis;
the upper pinnae of the frond have not been pre-
served. More information is needed.

4, Pteridrys microthecia (Fée) C.Chr. &
Ching, Bull. Fan Mem. Inst. Biol. 5 (1934)
139, pl. 14, 18, f. 10, 11; Tard. & C.Chr., Fl.
Indo-Chine 7, 2 (1941) 305; Copel., Fern Fl.
Philipp. (1960) 299. — Aspidium microthecium
Fée, Mém. Foug. (1865) 37, t. 41, f. 2. —
Dryopteris microthecia (Fée) C. Chr., Index
Filic. Suppl. 2 (1917) 15; C. Chr., Gard. Bull.
Straits Settlem. 7 (1934) 248. — Type: Cuming
13, Luzon (RB; iso BM, K, L, SING).

Dryopteris mettenianum Christ, Philipp. J. Sci. 2
(1907) Bot. 210; Alderw., Malayan Ferns (1908)
815. — Type: Cuming 13, Luzon (P).

Dryopteris vangenderenstortii Alderw., Bull. Jard.
Bot. Buitenzorg II, 23 (1916) 11; Malayan Ferns
Suppl. (1917) 502. — [Dryopteris subsagenioides
Alderw., Bull. Jard. Bot. Buitenzorg I, 11 (1913)
9; Malayan Ferns Suppl. (1917) 160; non
Christ 1910.] — Type: ’Amdjah 321, N Borneo
Boundary (BO).

Stipe to 60 cm long, lightly rufous and gla-
brous apart from sparse very short hairs near the
base on the abaxial side, basal scales rather thin

Flora Malesiana ser. II, Vol. 2! (1991)

and to more than 2 mm wide; /amina to more than
100 cm long; basal pinnae longest, their bases
very asymmetric, the stalk 2-5 mm long below
the basal acroscopic lobe (which is up to 3.5 cm
long) and more than 20 mm below the basal basi-
scopic lobe which is commonly 8-12 cm long (to
16 cm) and more than 2 cm wide, deeply lobed;
second basiscopic lobe much shorter than the basal
one; suprabasal pinnae to c. 30 cm long and 5-6
cm wide, lobed to 5 mm from costae, sinuses be-
tween lobes 2—3 mm wide, the lobes slightly fal-
cate, distinctly but shallowly crenate-serrate near
their acute tips only; veins in most pinnae once
forked (twice forked only in the largest lobes),
basal basiscopic veins often arising from the costa
below the bases of their costules; all surfaces gla-
brous apart from sparse very short hairs near the
bases of costae (seen only on young fronds); sori at
the apices of acroscopic vein-branches; indusia
little over 0.5 mm in diameter. — Fig. 2b.

Distribution — S$ Vietnam; in Malesia: Philippines
(widely), Borneo, Celebes, New Guinea.

Habitat — In forest, to 1200 m, reported several
times from steep rocky slopes above streams.

5. Pteridrys olivacea (Rosenst.) Copel., Gen.
Fil. (1947) 126. — Dryopteris olivacea Rosenst.,
Hedwigia 56 (1915) 352. — Type: Bamler 103,
E New Guinea, Sattelberg (iso K).

Allied to P. microthecia, differing as follows:
basal basiscopic lobes of basal pinnae to c. 6 cm

long, little longer than the next lobes; suprabasal
pinnae lobed to 2—3 mm from their costae, their
lobes strongly crenate-serrate throughout. — Fig. 2c.

Distribution — Malesia: New Guinea (NE New
Guinea, New Ireland).

Habitat — On steep rocky river banks at alti-
tudes to 900 m.

Note — Several collections, from widely sepa-
rated localities, agree in the specified differences
from P. microthecia. The two species are certainly
closely allied, agreeing in almost wholly glabrous
fronds and the position of basal basiscopic veins
on some pinnae.

6. Pteridrys confertiloba Holttum, nom. nov.
et stat. nov. — Dryopteris mettenianum Christ
var. novoguineensis Brause, Bot. Jahrb. Syst.
56 (1920) 93. — Type: Ledermann 7826, ENew
Guinea, Sepik River Exp. (B; iso K).

Allied to P. microthecia, differing as follows:
all pinnae sessile; sinuses between pinna-lobes very
narrow; basal acroscopic lobes of basal pinnae al-
most free; basal acroscopic lobes of suprabasal
pinnae overlapping the rachis on its abaxial side,
all lobes distinctly though shallowly crenate through-
out, their apices mostly obtuse. — Fig. 2d.

Distribution — Malesia: New Guinea (known
only from the type and one collection from the
Markham valley, Wakefield 1417)).

Habitat — The type from open rocky forest in a
gorge.

PLEOCNEMIA

Pleocnemia Presl, Tent. Pterid. (1836) 183, t. 7, f. 12; Hook., Gen. Fil. (1842) t. 70A,
f. 3 only, & t. 97; Presl, Epim. Bot. (1851) 50; Fée, Mém. Foug. 5. Gen. Filic.
(1852) 311; J.Sm., Ferns Brit. For. (1866) 146, f. 71; Bedd., Handb. Ferns Brit.
India (1883) 223, for P. leuzeana only; Alderw., Malayan Ferns (1908) 170, 810, for
P. leuzeana only; Holttum, Reinwardtia 1 (1951) 171; Kew Bull. 29 (1974) 341. —
Nephrodium § Pleocnemia Hook., Sp. Fil. 4 (1862) 61, for N. leuzeanum only. —
Type species: Pleocnemia leuzeana (Gaud.) Presl.

Dictyopteris Presl, Tent. Pterid. (1836) 194, t. 8, f. 7, excl. D. attenuata Presl; Fée,
Mém. Foug. 5. Gen. Filic. (1852) 267, non Lamour. 1809. — Arcypteris Underw.,
Bull. Torrey Bot. Club 30 (1903) 678; Holttum, Reinwardtia 1 (1951) 191. — Type
species: Dictyopteris irregularis (Presl) Presl.

Dictyopteris auct.: Bedd., Handb. Ferns Brit. India (1883) 298, for D. difformis only.

Aspidium p.p. & Tectaria p.p. quoad auct. plur.

Holttum — Tectaria Group . 9

Caudex usually erect, sometimes to 30 cm or more tall but with little sclerotic tissue;
fronds commonly 150—300 cm long; bases of stipes covered with thin narrow dull brown
scales, their edges entire or with short teeth formed by the projecting wall between adja-
cent cells; vascular strands in stipe more numerous than in Tectaria, with additional small
accessory strands outside the ring which is present in Tectaria, and additional large strands
on the adaxial side also; /amina bipinnatifid to bipinnate-tripinnatifid (or at base tripinnate),
basal pinnae with greatly enlarged basiscopic lobes or pinnules; rachises with very short
septate hairs on their raised upper surfaces; pinnae and pinnules + deeply lobed, with a
tooth projecting out of the plane of the lamina in each sinus between two lobes; veins
always forming a narrow costal areole, lacking free veinlets within it, between the bases
of costules (whether of pinna- or pinnule-lobes) and often similar areoles along costules
(the outer veins of costal areoles never joining the costa at their distal ends as normally in
Tectaria), with additional anastomosis of veins forming + isodiametric areoles below the
sinuses and external to costular areoles where space allows, rarely with short free veinlets
in them; sori variously placed on free or anastomosing veins, round, indusiate or not
(indusia reniform where present); thick glandular hairs, spherical, clavate or cylindrical,
yellow or red, present at the ends of hairs on the stalks of sporangia, often also on the
lower surface of costules and veins. — Figs. 3, 4.

Distribution — Mainly Malesian, extending to NE India and SE China, eastwards to
Samoa; about 20 species.

Habitat — Forest ferns, but generally not in the deepest shade; several are reported
from rocky places, at least one from limestone.

Cytology — n = 41 [Manton in Holttum (1955); M.G. Price in Holttum (1974)].

Taxonomy — There are two groups of species, for the types of which Pres] in 1836
established the genera Pleocnemia and Dictyopteris (later renamed Arcypteris Underw.).
I retained both genera in a monographic treatment in 1951 but remarked on their close
alliance, and united them in 1974. A probable hybrid having one parent from each group
is known (P. x intermedia). The only differences between the two groups are a more com-
plex venation and less distinct sinus-teeth in Arcypteris.

Confusion as regards generic assignment of these ferns in the 19th century resulted
from emphasis on two kinds of characters: pattern of venation, which these ferns share
with Tectaria sect. Sagenia (and less clearly with some thelypteroid ferns), and the pres-
ence or absence of indusia. Until Diels (1899) it was not fully understood that species
lacking indusia could be closely allied to others which have them and thus be congeneric.

Furthermore, the distinctive combination in Pleocnemia of some other characters was
ignored. To anyone seeing a living plant, as did Gaudichaud when he collected specimens
of P. leuzeana, the presence of sinus-teeth which project out of the plane of a pinnule, de-
scribed and figured by him, is striking. The sinus-teeth were noticed by Gaudichaud,
John Smith (who had living plants in his care) and for one species by Van Alderwerelt;
Presl, Fée and Hooker ignored them and omitted them from their illustrations. Unicellular
yellow or orange glands were noticed by various authors but not thought to be of generic
significance. The peculiar vascular anatomy of stipes was observed only by Fée [Gen.
Fil. (1852) t. 21A] for P. macrodonta and shown in a distorted form from a dried speci-
men.

10 Flora Malesiana ser. II, Vol. 2! (1991)

The species here placed as nos 1—4 are closely allied; two of them have been ranked as
varieties of P. irregularis. They agree in exindusiate son, for which reason earlier authors
placed them in Polypodium, or in Dictyopteris which Pres] placed with polypodioid ferns.
The original species of Pleocnemia is also exindusiate and was earlier named Polypodium
leuzeanum by Gaudichaud; in 1836 Pres] placed it with polypodioid ferns. But among
Cuming’s Philippine collections were four numbers, all resembling P. /euzeana in frond-
form and venation, two of them clearly indusiate. John Smith, who made the first survey
of Cuming’s collections (1841) regarded all four as forms of P. leuzeana, which he placed
with aspidioid ferns, remarking that indusia in these ferns were often small and soon shed.
Hooker [Gen. Fil. (1842) t. 97] accepted John Smith’s judgement and illustrated a fertile
pinnule of P. cumingiana and a sterile one of P. conjugata under the name P. leuzeana.
Later Beddome (1866) published as P. /euzeana a drawing of the sterile apex of a frond
(probably P. conjugata) from Moulmein in Burma along with an enlarged drawing of a
fertile pinnule-lobe of P. cumingiana, not stating that the latter was from a non-Burmese
specimen (he cited P. conjugata and P. cumingiana as synonyms).

In 1857 Thomas Moore accepted the genus Pleocnemia, defining it by frond-form,
venation and indusiate sori, and added one species of Heterogonium and one thelypteroid
species. In 1864 Hooker, ranking Pleocnemia as a section of Nephrodium, added two
more thelypteroid species, but John Smith (1875) rejected these additions. Beddome
(1883) added species here included in Tectaria sect. Sagenia and he placed all exindusi-
ate species of Tectaria in Dictyopteris. In general, Van Alderwerelt followed Beddome.
Copeland, in all his later works, included the type species of both Pleocnemia and Dic-
tyopteris Pres] in Tectaria.

Though Copeland lived for several years at Los Banos, at the foot of Mt Makiling in
Luzon, he never discovered distinctions among the plants of Pleocnemia s.s. growing in
the forest on that mountain and in 1960 included all in P. leuzeana. He did recognize as
distinct an exindusiate species in Sumatra (P. olivacea) in 1914 and Van Alderwerelt
another in Java (P. hemiteliiformis). From my field and herbarium studies based on
Singapore I recognized that Pleocnemia is a genus distinct from Tectaria and that (apart
from P. irregularis) three distinct species exist in the Malay Peninsula. After a further
study of specimens in the Bogor herbarium I identified the peninsular species as P. oliva-
cea, P. hemiteliiformis and P. conjugata (Blume) Presl, the last-named based on Aspi-
dium conjugatum Blume (type from Java), a name ignored by Backer and Posthumus in
their fern-flora of Java. I also saw duplicates of all the Cuming Philippine collections and
realized that they do not all belong to one species. I attempted to account for all this new
information in a paper published in 1951. In more recent years M.G. Price has made de-
tailed field studies of Philippine species and has confirmed the distinctions between spe-
cies which I had based on Cuming’s specimens; this further information is included in a
second paper, published in the Kew Bulletin in 1974.

The species of Pleocnemia s.s. (nos 6—18) are all similar in frond-form and venation.
The fronds of mature plants of all are very large and cannot be mounted whole on herbar-
ium sheets. The basal pinnae are of distinctive form on the basiscopic side near the base,
but distally are much like those next above them; more distal pinnae decrease gradually.
Some herbarium specimens are from immature plants; few are annotated to indicate which

Holttum — Tectaria Group a _ 1]

part of a frond they represent. Thus it is sometimes difficult to make comparisons. between
one specimen and another. For purposes of identification one must rely mainly on the
form of pinnules of larger suprabasal pinnae and on the sori they bear. The scales at the
bases of stipes are also sometimes important but are lacking from most specimens. The
abundance of appressed glands on the lower surface of veins appears to be variable in
fronds of the same species, also their thickness; their shape and colour may be distinctive.
More evidence from living plants is needed.

In 1912, dealing with specimens from New Guinea, Rosenstock remarked on the differ-
ences in spore-form from specimens which he regarded as different varieties of P. leuze-
ana. I examined spores of all species and confirmed that they are of three kinds, basing
my key of 1951 on them. They offer important evidence for the distinctions between
P. cumingiana and P. presliana which otherwise are not easy to distinguish except that the
former is indusiate, the latter not. Field study has shown that plants of P. presliana are
smaller and have different scales. Cytological examination might provide further evidence
and experimental hybridization might give further understanding.

Key to the species — The distinctness of some species (e.g. P. cumingiana and P.
conjugata) is very evident, but for others more difficult to state clearly, and more evidence
from field studies is needed. The present key doubtless needs improvement.

KEY TO THE SPECIES

la. Fronds usually bipinnatifid apart from basal pinnae; pinnules, if present, mostly ad-
nate to pinna-rachis; many additional areoles always present below sinuses between

lobes-isinus-teeth only evident in larser'sinuses:.)) 65 8.48 oo ee 2

b. Fronds amply bipinnate on mature plants, most pinnules not adnate; areoles few be-
lowasinuses::sinus-teeth always evident:..2.. 505 24 See Se kes eee a 5

2a. Fronds normally bipinnatifid, the largest sometimes bipinnate near the base; sori scat-
tencduime ularly. very abundant®:......52.....5..6¢.5.+.5. 00 1. P. irregularis

b. Fronds, at least in basal parts, bipinnate with lobed pinnules; sori all, or almost all,
arranged in one row on each side of costules of pinnule-lobes ............. 3)

3a. Sori midway between costules and margins of pinnule-lobes and also some on basal
MINS MR ee Seas cod cat eve A gac see aeeatik sua en Rete eas Et ae 2. P. macrodonta

b. Sori close to margins of pinnule-lobes, lacking on basal veins ............. -
4a. Pinnules of suprabasal pinnae acuminate, to 12 cm long and 2 cm wide at their broad-
ly cuneate bases, not adnate to pinna-rachis ............. 3. P. brongniartii

b. Pinnules of suprabasal pinnae to 4 x 0.8 cm with blunt apices, adnate to pinna-rachis
4. P. andaiensis
5a. Sori present on all vein-branches, in more than | row on each side of costules, sev-

eral sori irregularly arranged on veins below sinuses....... ‘5. P. x intermedia

b. Sori almost always in a single row on each side of the costules (sometimes on both
branches of a vein), additional ones below sinusesfew ................. 6

GDR S OTL ATU SHALE 5 30 sae elec MONE aie Bel heen AYE oe udeie cl cbtauie taped nace ee retire es) ea a 7

12

Flora Malesiana ser. II, Vol. 2! (1991)

7a. Pinnules of middle pinnae lobed c. 2/5 towards costules, lobes distinctly falcate; sinus-

teethinarnrOWA eee ea eee

be Pinnulesimoreideeply, lobed:ysinus-teeth\ triangular] = 455 5. 4. 9

8a. Pinnules short-acuminate, lobes obtuse
b. Pinnules long-acuminate, lobes acute

eh Test sagas Rid ees A 6. P. conjugata
CLE Ca Ree ee eee 7. P. acuminata

9a. Pinnule-lobes subentire (small teeth distally)............. 8. P. pleiotricha

b. Pinnule-lobes crenate to lobulate .....

ieloeelio Mee he ee 10

10a. Stipe dark, glossy; pinnules to 25 x 5 cm, pinnate for half their length
9. P. porphyrocaulos
b. Stipe not dark and glossy; largest pinnules not thus pinnate .............. Le
lla. Pinnules to 15 x 3 cm, their lobes lobulate, drying brown-olivaceous

10. P. cumingiana

b. Pinnules commonly 8 x 1.8 cm, their lobes crenate, drying greenish

dentate tances. Gira Ae montis es:

11. P. seranensis

. Pinnules lobed about 2/3 towards costa; lobes entire or at most minutely crenate-
SURO oa oe 13

b. Pinnules lobed more deeply or not; lobes distinctly crenate .............. 15

lons.43)mm) widesat base. 4.2.22...

. Costules 4—4.5 mm apart; several costular areoles present; basal scales to 5 cm
st ane ae 12. P. hemiteliiformis

b. Costules to 6 mm apart; few costular areoles present; basal scales shorter and nar-

wh AMAA AS oe hen eee 14
+) Sinus-teeth narrow. shorten: baSe:Of Sinus) 2 ..4/45/ 50. oe 13. P. olivacea

b. Sinus-teeth broadly triangular, on basiscopic side of base of sinus

14. P. leuzeana

. Pinnules of suprabasal pinnae to 4.5 cm wide, their lobes acute

15. P. megaphylla

b. Pinnules of suprabasal pinnae to 2.5 cm wide, their lobes obtuse .......... 16

. Pinnules all lobed to c. 1 mm from costa, to 2.5 cm wide, spores not spinulose

16. P. presliana

b. Pinnules lobed less deeply, narrower; spores spinulose ........ 17. P. dahlii

1. Pleocnemia irregularis (Presl) Holttum,
Kew Bull. 29 (1974) 347, f. 1E. — Polypo-
dium irregulare Pres}, Relig. Haenk. (1825) 25,
t. 4, f. 3; Blume, FI. Javae Filic. (1829) 164,
t. 72; Hook., Sp. Fil. 5 (1864) 101; Holttum,
Novit. Bot. Delect. Seminum Horti Bot. Univ.
Carol. Prag. 1968 (1969) 21. — Dictyopteris
irregularis (Presl) Presl, Tent. Pterid. (1836)
194, t. 8, f. 7. — Phegopteris irregularis (Pres)
Mett., Ann. Mus. Bot. Lugd. Bat. 1 (1864) 225.
— Aspidium irregulare (Presl) C. Chr., Index
Filic. (1905) 78. — Tectaria irregularis (Presl)
Copel., Philipp. J. Sci. 2 (1908) Bot. 416;
Backer & Posth., Varenfl. Java (1939) 77;

Copel., Fern Fl. Philipp. (1960) 309, excl.
syn. Dictyopteris macrodonta Fée. — Arcyp-
teris irregularis (Presl) Ching, Sunyatsenia 5
(1940) 251; Holttum, Reinwardtia 1 (1951)
193, f. 1-3; Revis. Fl. Malaya 1 (1955) 538,
f. 317, 318. — Type: Haenke s.n., Philippines
(wrongly stated to be from Mexico); not seen
by Holttum 1969.

Aspidium difforme Blume, Enum. PI. Javae (1828)

160; Copel., Polypod. Philipp. (1905) 35. —
Polypodium difforme (Blume) Blume, FI. Javae
Filic. (1829) 131; Racib., Pteridop. Buitenzorg
(1898) 118. — Phegopteris difformis (Blume)
Mett., Fil. Hort. Bot. Lips. (1856) 84, t. 25, f. 8.

Holttum — Tectaria Group

— Dictyopteris difformis (Blume) T. Moore,
Index Fil. (1858) 90; Bedd., Handb. Ferns Brit.
India (1883) 300. —Arcypteris difformis (Blume)
Underw., Bull. Torrey Bot. Club 30 (1903)
678. — Type: Blume s.n., Java (L 908,300-30).

Polypodium confertum Roxb., Calcutta J. Nat.
Hist. 4 (1844) 493; Morton, Contr. U.S. Natl.
Herb. 38 (1974) 337. — Lectotype (Morton):
W. Roxburgh Jr, ‘Chittagong’ (Roxburgh, I.c.),
more likely from Penang (BR).

Polypodium multiflorum Roxb., Calcutta J. Nat.
Hist. 4 (1844) 493, non Roth 1797; Morton,
Contr. U.S. Natl. Herb. 38 (1974) 351.—
Type: C. Smith s.n., Ambon (G).

Polypodium eximium Kunze, Bot. Zeitung (Ber-
lin) 4 (1846) 424; Morton, Contr. U.S. Natl.
Herb. 38 (1974) 338. — Lectotype (Morton):
Zollinger 514A, Java (G; iso LE).

Caudex short, its apex and bases of stipes den-
sely covered with narrow dark brown scales 3—4
cm long; stipe to 80 cm long, green when living,
glabrescent; lamina to at least 100 cm long and 60
cm wide, deeply bipinnatifid or just bipinnate near
the base; pinnae in many pairs, the lowest with a
deeply lobed basiscopic pinnule to 20 cm long and
6 cm wide, much longer than the next pinnule;
suprabasal pinnae to 40 x 12 cm, the largest with
a few pairs of + sessile pinnules, distally deeply
lobed, the lobes falcate, the largest lobes with
crenate margins and acute apex, c. 1 cm wide, a
short broad tooth present at the bases of the larger
sinuses; upper pinnae gradually less deeply lobed;
rachis and bases of costae bearing short hairs on
the upper surface, the rest glabrous, lower surface
of rachis and costae bearing some hairs 1 mm
long, surfaces of costules and veins bearing red
glands like those in the sori; veins forming single
narrow areoles along costae between one costule
and the next and shorter areoles along costules, rest
of the lamina below sinuses filled with 4—6-sided
areoles, rarely with included free veins; sori small,
close, scattered irregularly, exindusiate, round or
often extending along the veins and sometimes
confluent; sporangia often bearing red spherical
glands on the hairs attached to their stalks. — Fig. 3a.

Distribution — Southern Burma, Thailand, Viet-
nam; throughout Malesia; Carolines, Solomons,
Fiji.

Habitat — In lowland forest; in western Malesia
‘tolerating drier conditions than many other terres-

trial forest species’ (Holttum 1955); ‘especially on
steep valley slopes’ (Backer & Posthumus 1939).

Note — In Thailand and Vietnam occur very large
plants which have more fully bipinnate fronds
than those in Malesia. These are named Tectaria
cumingiana by Tard. & C.Chr., Fl. Indo-Chine
7 (1941) 412 [= Pleocnemia macrodonta (Fée)
Holttum] but are much less fully bipinnate than in
that species and have sori arranged as in P. irregu-
laris.

2. Pleocnemia macrodonta (Fée) Holttum,
Kew Bull. 29 (1974) 348. — Dictyopteris ma-
crodonta Reinw. (‘macrodon’ ) ex. [Presl, Tent.
Pterid. (1836) 194, nom. nud.; J. Sm., J. Bot.
3 (1841) 396, for Cuming 9 only, nom. nud.]
Fée,*Gen, Fils1852)) 267, pls 21, ft. AZ. —
Phegopteris macrodonta (Fée) Mett., Farngatt.
IV (1858) 31. — Polypodium macrodon Reinw.
ex Baker, Syn. Fil. (1867) 318. — Tectaria
irregularis var. macrodon Copel., Philipp. J.
Sci. 2 (1907) Bot. 417. — Tectaria macrodus
(Baker) C. Chr., Index Filic. Suppl. 3 (1934)
181. — Arcypteris mecrodonta (Fée) Holttum,
Reinwardtia 1 (1951) 194. — Type: Cuming 9,
Luzon (Fée’s specimen not at RB; iso K, L; the
isotype in K is here chosen as lectotype. It is
the holotype of Polypodium cumingianum and
P. macrodon).

Polypodium cumingianum Hook., Sp. Fil. 5 (1864)
103, excl. pl. ex Samoa, Fiji (not Pleocnemia
cumingiana Presl). — Tectaria cumingiana
(Hook.) C. Chr., Notul. Syst. 7 (1938) 96, but
not sensu Tard. & C.Chr., Fl. Indo-Chine 7
(1941) 412.

Aspidium whitfordii Copel. in Perkins, Fragm. FI.
Philipp. (1905) 176. — Type: Whitford 201,
Luzon, Bataan Prov. (coll. lost).

Tectaria irregularis auct.: Copel., Fern Fl. Philipp.
(1960) 309, p.p.

Differing from P. irregularis as follows: scales to
2 cm long and more than 1 mm wide; pinnae con-
spicuously pinnate, the pinnules sessile and most-
ly + adnate, unequally cuneate at the base (broader
on the acroscopic side), their margins lobed to one
third or more towards the costa; sori in two close
rows on either side of each costule, often some-
what confluent, with additional sori near the sinuses
in the broader pinnules. — Fig. 3b, ec.

14 Flora Malesiana ser. II, Vol. 2! (1991)

PT. Epwaeps

Fig. 3. Pleocnemia irregularis (Presl) Holttum. a. A single lobe, showing position of sori, x 2.2. — P.
macrodonta (Fée) Holttum. b. A basal basiscopic pinnule, x 0.45; c. detail of lobes showing venation and
sinus teeth, x 3.3. — P. andaiensis (Baker) Holttum. d. Two lobes from near the apex of a pinna, x 4; e.
one segment from near the base of the same pinna, showing venation, sori and sinus teeth between lobes
(sori removed from the right side), x 2.2 (a: cult. Singapore, as in Reinwardtia 1 (1951) 192; b, c:
Holttum s.n., Sarawak, cult. Kew 335-58; d, e: Beccari s.n., Andai).

Holttum — Tectaria Group

15

Distribution — Malesia: Borneo (Sarawak), Phi-
lippines (Luzon, Sibuyan, Mindoro), E New Gui-
nea, Admiralty Is., New Britain.

Notes — Fée did not print a description, but his
figures (overlooked by Hooker) adequately distin-
guish this species from P. irregularis and P. brong-
niartii. The specimens named Tectaria cumingiana
by Tardieu & C. Christensen (1941) represent the
large form of P. irregularis referred to above.

Pres! in 1836 based his new name Dictyopteris
macrodonta on Polypodium macrodon Reinw.; in
1867 the latter binomial was preferred by Baker to
Polypodium cumingianum Hook.; but neither
Presl nor Baker cited a particular publication by
Reinwardt, and I find no evidence that Reinwardt
published the name which is not in either part of
his Sylloge Plantarum Novarum (1824).

3. Pleocnemia brongniartii (Bory) Holttum,
Kew Bull. 29 (1974) 350. — Polypodium
brongniartii Bory in Duperry, Voy. Monde 1
(1828) 263, t. 34; Hook., Sp. Fil. 5 (1864)
103. — Phegopteris brongniartii (Bory) Mett.,
Farngatt. TV (1858) 31. — Aspidium brongniartii
(Bory) Diels in E. & P., Nat. Pflanzenfam. I, 4
(1899) 186; Copel., Polypod. Philipp. (1905) 35.
— Tectaria irregularis var. brongniartii Copel.,
Philipp. J. Sci. 2 (1907) Bot. 417. — Arcy-
pteris brongniartii (Bory) Holttum, Reinwardtia
1 (1951) 195. — Type: d’Urville s.n., Waigeo
Is. (P).

Dictyopteris pteroides Presl, Tent. Pterid. (1836)
194; Fée, Gen. Fil. (1852) t. 21, f. Al; Alderw..,
Malayan Ferns (1908) 515. — [Polypodium
pteroides Presl, Reliq. Haenk. (1825) 25, non
Retz. 1791; Holttum, Novit. Bot. Delect. Sem.
Horti Bot. Univ. Carol. Prag. 1968 (1969) 21.]
— Type: Haenke s.n., Luzon (PRC).

Similar to P. macrodonta in the size and branch-
ing of its fronds and in the shape of pinnules,
differing in the arrangement of sori which are all
near the margins of the lobes of pinnules or of
pinnae near the frond-apex.

Distribution — Malesia: New Guinea (Waigeo
Is.), Philippines (southern Luzon, Sibuyan).

4, Pleocnemia andaiensis (Baker) Holttum,
Kew Bull. 29 (1974) 350. — Polypodium (Dic-
tyopteris) andaiense Baker in Beccari, Malesia 3

(1886) 45. — Dictyopteris andaiensis (Baker)
Alderw., Malayan Ferns (1908) 514. — Tectaria
andaiensis (Baker) C. Chr., Index Filic. Suppl.
3 (1934) 177; Dansk Bot. Ark. 9 (1937) 49;
Copel., Philipp. J. Sci. 78 (1951) 414. — Type:
Beccari s.n., Andai, W New Guinea (FI; iso K).
Nephrodium giganteum auct. non (Blume) Baker
1874: Cesati, Rendiconti Reale Accad. Sci. Fis.
(Napoli) 16 (1877) 26. — Arcypteris gigantea
Holttum, Reinwardtia 1 (1951) 195, nom. illeg.

Differs from P. brongniartii as follows: supra-
basal pinnae to 30 cm long, near their bases bear-
ing a few pairs of separately adnate pinnules, the
largest 4.5 x 0.9 cm, lobed to almost halfway to
the costa, lobes broadly rounded, basal acroscopic
ones not enlarged, costules 3.5—4 mm apart con-
nected by 1 row of costal areoles; rather sparse
small scales and short hairs present on upper sur-
face of rachis and costae, very slender hairs on the
lower surface; unicellular + elongate reddish glands
present on lower surface, mostly on veins, some
also on the upper surface; sori in a row near (but
not touching) the margin of each pinnule-lobe, the
distal ones on free veins; glands not seen in sori. —
Fig. 3d, e.

Distribution — Malesia: W New Guinea (known
from two collections, the second being van Royen
3144, Sorong District).

Note — The name Nephrodium giganteum, pub-
lished by Cesati in 1877, was a misidentification
of Beccari’s specimen later named Polypodium an-
daiense by Baker.

5. Pleocnemia x intermedia Holttum, Kew
Bull. 29 (1974) 349. — Type: Wright & Isma-
wi § 32555, Sarawak, Limbang Dist. (K; iso L).

Basal scales c. 2.5 cm long, very narrow, dis-
tally filiform and tangled; basal pinnae to 80 cm
long, their basal basiscopic branches 45 cm long,
both with separate pinnules almost to the apex;
middle pinnules of suprabasal pinnae mostly 8 x 2
cm with winged stalks 2 mm long, lobed 3/4 to-
wards costae, basal lobes deeply crenate, distal ones
shallowly; sinus-teeth broad and very short in
lower sinuses, narrow and short in distal ones; cos-
tules 5 mm apart; veins forming costal areoles
with a few additional ones below sinuses, veins in
pinnule-lobes twice forked to paucipinnate, the
lower veinlets anastomosing irregularly, distal ones

16

(in crenatures) free; lower surface glabrous apart
from thick glandular hairs on costules and veins;
sori small, exindusiate, on almost all vein-branches
both in pinnule-lobes and below sinuses; spores
few and shrivelled.

Distribution — Known only from the type.

Habitat — Lower slopes of a ridge in low Dip-
terocarp forest.

Note — This is evidently a hybrid between P.
irregularis and one of the amply bipinnate species,
perhaps P. olivacea. No other specimens of such
possible hybrids are known.

6. Pleocnemia conjugata (Blume) Presl, Epim.
Bot. (1851) 259; Holttum, Reinwardtia 1 (1951)
177, f. 1, 5, 7, 9; Revis. Fl. Malaya 2 (1955)
534, f. 314; Molesworth Allen, Gard. Bull.
Sing. 17 (1959) 268; Holttum, Kew Bull. 29
(1974) 350. — Aspidium conjugatum Blume,
Enum. Pl. Javae (1828) 169. — Type: ex Herb.
Blume, Moluccas (L; iso K).

Pleocnemia javanica Presl, Epim. Bot. (1851) 259;
Holttum, Novit. Bot. Delect. Seminum Horti
Bot. Univ. Carol. Prag. 1968 (1969) 46. —
Lectotype (selected here): Zollinger 1459 (BM,
not in PRC).

Pleocnemia fimbrillifera Alderw., Bull. Jard. Bot.
Buitenzorg II, 16 (1914) 28. — Tectaria fim-
brillifera (Alderw.) C.Chr., Index Filic. Suppl.
3 (1934) 179. — Type: C.G. Matthew s.n.,
Sumatra, G. Singgalang, 750 m (BO).

Pleocnemia conjugata vat. elatior Holttum, Rein-
wardtia 1 (1951) 179. — Type: C.G. Matthew
512, Sumatra, G. Singgalang, 1500 m (BO).

Pleocnemia leuzeana auct.: Hook., Gen. Fil. (1842) t.
97, f. 1, 2 only; Bedd., Ferns Brit. India (1866)
t. 134 for sterile frond only; Handb. Ferns Brit.
India (1883) 228, p.p.; Alderw., Malayan Ferns
(1908) 172, p.p. — Tectaria leuzeana auct.:
Backer & Posth., Varenfl. Java (1939) 73, p.p.

Basal scales long-attenuate, conspicuously den-
tate distally; lamina to 120 cm long; suprabasal
pinnae to 70 cm long, their pinnules sessile, 8—13
cm long, 1.8—2.3 cm wide, lobed a little more
than halfway to costa, very firm, almost glaucous
beneath when living, brown-olivaceous when dried;
costules 5—6(—7) mm apart; lobes distinctly fal-
cate and minutely dentate distally, sinuses between
them very narrow; sinus-teeth small, narrow and
curved; veins forming costal areoles with a few ad-

Flora Malesiana ser. II, Vol. 2! (1991)

ditional ones below sinuses, costular areoles usu-
ally near bases of lobes only; glands present on
lower surface of costules and veins; sori in a single
row midway between costule and margin of a pinna-
lobe, rather large, indusiate; spores with a narrow
+ continuous wing and a few cross-wings. — Fig.
4a, b.

Distribution — Hong Kong, Burma; throughout
Malesia.

Habitat — Apparently in areas with a short reg-
ular dry season (in the Peninsula only in the north)
and in rather open forest.

7. Pleocnemia acuminata Holttum, Rein-
wardtia 1 (1951) 182; Kew Bull. 29 (1974)
351. — Type: Lérzing 5644, Sumatra, Sibo-
langit (BO).

Differs from P. conjugata as follows: lamina
more rigid; pinnae and pinnules narrowly acumi-
nate; pinna-lobes more widely spaced and acute;
spores as in P. leuzeana.

Distribution — Malesia: Sumatra (Sibolangit) (2
collections).

Note — This is very near P. conjugata and speci-
mens of the latter have also been collected at Sibo-
langit. A local survey is needed.

8. Pleocnemia pleiotricha Holttum, Rein-
wardtia 1 (1951) 182, f. 12, 13; Kew Bull. 29
(1974) 351. — Type: Elmer 21421, Sabah, Ta-
wau (SING; iso BO, K).

Basal scales to 3 cm long and 1.5 mm wide,
rather firm, attenuate but the tips not filiform nor
dentate; pinnules of suprabasal pinnae to 11 x 2.2
cm, with stalks 2 mm long, rather abruptly acumi-
nate, lobed 3/4 towards costae, lobes well spaced,
minutely dentate distally (about as in P. conjuga-
ta); sinus-teeth broadly triangular; veins forming
costal areoles with no additional ones below si-
nuses, costular areoles usually 1 pair only; red
glands on costules and veins abundant; sori indusi-
ate, indusia thin. — Fig. 4d, e.

Distribution — Malesia: S Sumatra, N Borneo.

9. Pleocnemia porphyrocaulos Alderw., Bull.
Jard. Bot. Buitenzorg III, 5 (1922) 215; Holttum,
Reinwardtia 1 (1951) 188. — Type: Béguin
1123, Ternate (BO; iso L).

Holttum — Tectaria Group

17

Stipe very dark, glossy; largest suprabasal pin-
nae 135 cm long; pinnules to 25 cm long, 5 cm
wide at the base, pinnate for half their length;
third-order leaflets to 3 x 0.8 cm, lobed halfway to
their costae, a row of areoles present along each
side of costae, 2 or 3 indusiate sori in each lobe;
largest pinnatifid pinnules with falcate dentate
lobes 3 mm wide separated by more than their own
width; spores as in P. cumingiana.

Distribution — Malesia: Moluccas (known from
the type only).

Note — So far as present information goes, the
smaller pinnules of the type are not distinguish-
able from those of P. cumingiana, but the very
dark glossy stipes are distinctive, and the largest
pinnules are far larger and far more fully pinnate
than those on the largest known specimen of P.
cumingiana which has the basal pinnule-lobes just
free as tertiary leaflets.

10. Pleocnemia cumingiana Presl, Epim. Bot.
(1851) 50; Holttum, Reinwardtia 1 (1951) 188,
f. 20; Novit. Bot. Delect. Seminum Horti Bot.
Univ. Carol. Prag. 1968 (1969) 46; Kew Bull.
29 (1974) 382. — Lectotype (Holttum 1951):
Cuming 107, Luzon (PRC; iso BM, K, SING).

Nephrodium chrysotrichum Baker, Ann. Bot. (Lon-
don) 5 (1891) 328. — Tectaria chrysotricha
(Baker) C. Chr., Index Filic. Suppl. 3 (1934)
178; Bernice P. Bishop Mus. Bull. 177 (1934)
101. — Pleocnemia chrysotricha (Baker) Holt-
tum, Reinwardtia 1 (1951) 187. — Type: Whit-
mee s.n., Samoa (K).

Aspidium leuzeanum var. alsophiloides Christ,
Bot. Jahrb. Syst. 23 (1897) 353; Rechinger,
Akad. Wiss. Wien M.-N. KI., Denkschrift 84
(1908) 48. — Type: Reinecke 90, Samoa (B).

Aspidium angilogense Christ, Bull. Herb. Boissier
II, 6 (1906) 1003; Philipp. J. Sci. 2 (1907)
Bot. 158. — Pleocnemia leuzeana var. angilo-
gense (Christ) Alderw., Malayan Ferns (1908)
810. — Type: Loher s.n., Angilog (P?, n.v.).

Dryopteris rufinervis Hayata, J. Coll. Sci. Imp.
Univ. Tokyo 30 (1911) 420. — Aspidium rufi-
nerve (Hayata) Hayata, Icon. Pl. Formos. 8
(1914) 141, f. 67, 68. — Pleocnemia rufinervis
(Hayata) Nakai, Bot. Mag. (Tokyo) 47 (1933)
163. — Type: Nakahara 274, Taiwan (TAI, n.v.).

Pleocnemia leuzeana auct.: Hook., Gen. Fil. (1842)
t. 97 for f. 3-5 only; Bedd., Ferns Brit. India

(1866) t. 134 for fragm. fert. only; Handb. Ferns
Brit. India (1883) 228. — Nephrodium leuzea-
num auct.: Hook., Sp. Fil. 4 (1862) 61, p.p.
— Tectaria leuzeana auct.: Copel., Fern Fl.
Philipp. (1960) 310, p.p.

Caudex to 50 cm tall, falling and then resuming
upright growth from its apex; stipes not very dark
nor glossy, for some distance above the base den-
sely covered with distally very narrow and conspic-
uously dentate scales to 3 cm long, base of scales
2 mm wide, pinnules of suprabasal pinnae to more
than 15 cm long, to 3 cm wide, very deeply lobed;
costules 5—7 mm apart; lobes usually narrower,
more widely spaced and more deeply crenate-lobu-
late in fertile than in sterile pinnules (but these
characters are very variable); veins forming narrow
costal areoles with no additional ones below si-
nuses, costular areoles variably developed, veins in
crenatures once or twice forked; glands on costules
and veins yellow, usually abundant; sori indusiate;
spores densely spinulose. — Fig. 4h.

Distribution — Taiwan; in Malesia: Moluccas,
Philippines, New Guinea; Solomons, Fiji, Samoa.

Note — Rechinger reported that the trunks of
this species attained a length of 10 m and were
used for house-building in Samoa; this was cer-
tainly an error; trunks so used would be from spe-
cies of Cyatheaceae.

11. Pleocnemia seranensis Holttum, Rein-
wardtia 1 (1951) 187; Kew Bull. 29 (1974) 352. —
Type: Rutten 1850, Ceram (BO; iso L, SING).

Differs from P. cumingiana as follows: Pin-
nules of suprabasal pinnae commonly 8 x 1.8 cm
(largest seen 12 x 2 cm), thinner, drying greenish
(not brown-olivaceous), their lobes at most cre-
nate.

Distribution — Malesia: Moluccas, New Guinea.

Note — The distinction between this species and
P. cumingiana needs to be established through
field study.

12. Pleocnemia hemiteliiformis (Racib.)
Holttum, Reinwardtia 1 (1951) 179, f 11:
Revis. Fl. Malaya 2 (1955) 536, f. 316; Kew
Bull. 29 (1974) 354. — Pleocnemia leuzeana
var. hemiteliiformis Racib., Pteridop. Buitenzorg
(1898) 194; Alderw., Malayan Ferns (1908)

18 Flora Malesiana ser. II, Vol. 2! (1991)

PT. Erwarnos

Fig. 4. Pleocnemia conjugata (Blume) Presl. a. Transverse section of stipe near base, x 2; b. one pinnule
lobe of a middle pinna, x 2, with detail of sinus-tooth. — P. olivacea (Copel.) Holttum. c. A pinnule
lobe of a middle pinna, x 3.3. — P. pleiotricha Holttum. d. A pinnule lobe of a middle pinna, x 3.3;
e. detail of glands, x 23. — P. dahlii (Hieron.) Holttum. f. Pinnule lobes of a middle pinna, x 3.3;
g. pinnule lobes of an adjacent middle pinna for comparison; this shows the condition attributed to P.
dimidiolobata; x 3.3. — P. cumingiana Presl. h. Lobe of a middle pinna, x 3.3 (a, b: from Reinwardtia 1
(1951) 174; c: van Balgooy 4789; d, e: Elmer 21421; f, g: Croft 240; h: Cuming 107).

Holttum — Tectaria Group

173. — Dictyopteris hemiteliiformis (Racib.)
Alderw., Bull. Jard. Bot. Buitenzorg II, 11
(1913) 7; Malayan Ferns Suppl. (1917) 321. —
Tectaria hemiteliiformis (Racib.) C. Chr., Index
Filic. Suppl. 3 (1934) 180. — Type: Ract-
borski s.n., Java, G. Salak (BO; iso L, K).

Dictyopteris compitalis Alderw., Bull. Jard. Bot.
Buitenzorg III, 5 (1922) 194. — Tectaria com-
pitalis (Alderw.) C. Chr., Index Filic. Suppl. 3
(1934) 178. — Type: Lérzing 6378, Sumatra,
Sibolangit (BO).

Lower 30 cm of stipe + persistently covered
with scales, the longest 5 cm long, gradually nar-
rowed from a base 3 mm wide; pinnules of supra-
basal pinnae commonly 8 x 1.7 cm, lobed 2/3
towards their costae, lobes well-spaced, obtuse,
their margins at most sinuous and bearing + abun-
dant short hairs; costules 4-5 mm apart; veins
usually dark on lower surface when dried, forming
one series of areoles between costal areoles and
sinuses, costular areoles many; sinus-teeth short,
broadly triangular and slightly oblique; sori exin-
dusiate, sometimes on both branches of a forked
vein, also on anastomosing veins below sinuses;
spores with a continuous wing and some cross-
wings.

Distribution — Peninsular Thailand; Malesia.

Habitat — Open places in forest at 8300-1400 m.

Note — The type of Dictyopteris compitalis
shows no indication of the reduced auriculiform
pinnae described by the author.

13. Pleocnemia olivacea (Copel.) Holttum,
Reinwardtia 1 (1951) 181, f. 8, 10; Revis. FI.
Malaya 2 (1955) 535, f. 172; Kew Bull. 29
(1974) 355. — Tectaria olivacea Copel., Phi-
lipp. J. Sci. 9 (1914) Bot. 228. — Dictyopteris
olivacea (Copel.) Alderw., Malayan Ferns Suppl.
(1917) 322. — Type: C.J. Brooks 172, Suma-
tra, Lebong Tandai (MICH; iso BM).

Related to P. hemiteliiformis but fronds smaller,
basal scales narrow, to 2.5 cm long, pinnules thin-
ner with costules to 6 mm apart, lobes more close
to each other, usually with minute marginal teeth
distally, lacking hairs; veins forming basal cos-
tular areoles only; sinus-teeth short and narrow,
in bases of sinuses; sori exindusiate, sometimes
on both branches of a vein; spores as those of P.
hemiteliiformis. — Fig. 4c.

Distribution — Western Malesia.
Habitat — In rocky places in lowland forest.

14. Pleocnemia leuzeana (Gaudich.) Presl,
Tent. Pterid. (1836) 183, pl. 7, f. 12; Hook.,
Gen. Fil. (1842) t. 70, f. 3 only; Alderw.,
Malayan Ferns (1908) 172, p.p.; Holttum,
Reinwardtia 1 (1951) 184; Kew Bull. 29 (1974)
356. — Polypodium leuzeanum Gaudich. in
Freycinet, Voy. Uranie (1827) 361, t. 6. — As-
pidium leuzeanum (Gaudich.) Kunze, Bot. Zei-
tung (Berlin) 4 (1846) 474. — Nephrodium
leuzeanum (Gaudich.) Hook., Sp. Fil. 4 (1862)
61, p.p. — Dryopteris leuzeana (Gaudich.)
Kuntze, Revis. Gen. Pl. 2 (1891) 813. — Tec-
taria leuzeana (Gaudich.) Copel., Philipp. J. Sci.
2 (1907) Bot. 417; Fern Fl. Philipp. (1960)
310, p.p. — Type: Gaudichaud s.n., Moluccas,
Pulo Pisang (P; iso K).

Near P. olivacea in form of pinnules; basal
scales not recorded; sinus-teeth broadly triangular,
on basiscopic side of sinus; spores with many
small obtuse protuberances all over them.

Distribution — Eastern Malesia; Fiji, Samoa.

Habitat — Recorded from ‘rocky bank’ (R.F.
Ellen 72, Ceram).

15. Pleocnemia megaphylla Holttum, Kew
Bull. 29 (1974) 357. — Type: Jermy 8149B,
cult. R.B.G. Kew, origin Papua New Guinea,
W Sepik Prov., Bewani Mts (K).

Stipe 100 cm or more long; basal scales 2.5 cm
long, pallid and 5 mm wide at their bases, abruptly
narrowed to 1.5 mm 5 mm above the base, thin,
filiform and dentate distally; /amina 120 cm long;
rachises dark castaneous and minutely hairy on
abaxial surface; basal pinnae 85 cm long with
three diminishing pinnate basiscopic pinnules, the
largest 45 cm long with many deeply lobed tertiary
leaflets to 10 x 2.3 cm, largest acroscopic pinnules
17 cm long; pinnules on first suprabasal pinnae to
16 x 4.5 cm, narrowly acuminate, lobed to less
than 2 mm from the costa, firm and opaque when
dried, costules to 8 mm_.apart, lobes crenate to
lobulate, narrowed to an acute hardly falcate apex;
sinus teeth short and broadly triangular, oblique,
on basiscopic side of sinuses, sometimes present
also between lobules of pinna-lobes; veins form-
ing very narrow costal areoles, also an almost

20

Flora Malesiana ser. II, Vol. 2! (1991)

complete set of costular ones, free veins passing
from the latter to the margin, paucipinnate in
lobules; sori exindusiate, in one row on each side
of costules of lobes, nearer to costules than to
margin, on outer veins of areoles; spores minutely
spinulose; elongate unicellular glands present on
stalks of sporangia.

Distribution — Malesia: New Guinea (known
with certainty from the type only).

Note — Incomplete specimens from Mt Suckling
(Milne Bay Prov.) and Sudest Island may belong
to this species.

16. Pleocnemia presliana Holttum, Rein-
wardtia 1 (1951) 183, f. 14, 15, excl. New
Guinean spec.; Kew Bull. 29 (1974) 357. —
Type: Edano BS 78717, Luzon, Cagayan Prov.,
Mt Bawa (BO; iso SING).

Pleocnemia leuzeana auct.: T. M[asters], Gard.
Chron. II, 2 (1874) 354, f. 74.

Related to P. cumingiana, differing as follows:
fronds of mature plants much smaller; stipe mi-
nutely muricate throughout from bases of fallen
scales; basal scales narrower, filiform and much
tangled; pinnules of suprabasal pinnae to c. 10 x
2.2 cm, abruptly acuminate; cylindric glands on
lower surface of veins red; sori exindusiate; spores
with many small peg-like outgrowths.

Distribution — Malesia: Philippines (Luzon,
Palawan, Leyte, Mindanao).

Habitat — In forest, mainly at lower altitudes
than P. cumingiana.

Notes — Presl cited both Cuming 107 and 33
under P. cumingiana; 33 is P. presliana.

M.G. Price, who made intensive studies of
ferns on Mt Makiling, found both P. cumingiana
and P. presliana growing in the forest there, and
tells me that he regards them as quite distinct.

The illustration in Gard. Chron. (1874) shows
sinus-teeth clearly and also a characteristic spore;
the plant had been imported from the Philippines.

17. Pleocnemia dahlii (Hieron.) Holttum,
comb. nov. — Phegopteris dahlii Hieron., No-
tizbl. Bot. Gard. Berlin-Dahlem 2 (1898) 84. —
Aspidium dahlii (Hieron.) Diels in K. Schum.
& Lauterb, Fl. Schutzgeb. Siidsee (1901) 116.
— Dictyopteris dahlii (Hieron.) Alderw., Bull.
Jard. Bot. Buitenzorg II, 12 (1912) 11; Malayan

Ferns Suppl. (1917) 321. — Tectaria dahlii
(Hieron.) C. Chr., Index Filic. Suppl. 3 (1934)
178. — Type: Dahl s.n., 1896, ‘Neu Lauen-
burg Grup’, N of New Britain (B).

Pleocnemia dimidiolobata Holttum, Reinwardtia 1
(1951) 184, f. 16, 17; Kew Bull. 29 (1974)
356, excl. syn. P. leuzeana var. lobato-crenata
Rosenst. — Type: Brass 2648, Solomon Is.,
San Cristoval (K; iso BO).

Pleocnemia tripinnata Holttum, Reinwardtia 1
(1951) 185. — Type: Lauterbach 560, New
Guinea (SING; iso BO).

Pleocnemia leuzeana var. echinocarpa Rosenst. in
Feddes Repert. Spec. Nov. Regni Veg. 10 (1912)
337. — Type: Bamler LI, E New Guinea, Lo-
gaueng (not seen).

Base of stipe very dark, castaneous distally,
scales (near base only) to 2.5 cm long, 1 mm wide
at base, distally tiliform; pinnules of larger supra-
basal pinnae to 9 x 2 cm with short winged stalks,
lobed 2/3-—3/4 towards costae, lobes distinctly
crenate, obtuse; sinus-teeth broadly triangular,
oblique; veins forming costal areoles and a few
additional ones below sinuses, costular areoles
present only at bases of pinnule-lobes; sori exin-
dusiate, usually about midway between costules
and margin, mostly on acroscopic branches of
forked veins (none seen on both branches); spores
spinulose. — Fig. 4f, g.

Distribution — Malesia: Eastern New Guinea;
Solomons, New Hebrides.

Habitat — In lowland forest.

Note — At Kew is a later collection of P. leuze-
ana var. echinocarpa named by Rosenstock. The
above description is based largely on a plant culti-
vated at Kew. On other specimens the position of
sori is very variable; the condition shown by the
type of P. dimidiolobata represents an extreme
condition.

DOUBTFUL SPECIES

Nephrodium varium Presl, Reliq. Haenk. (1825)
36, excl. syn.; Holttum, Novit. Bot. Delect.
Seminum Horti Bot. Univ. Carol. Prag. 1968
(1969) 18. — Type: Haenke s.n., Luzon (PRC).
The type is a frond from a young plant of a

species allied to Pleocnemia irregularis. It might

represent P. macrodonta, but cannot be identified
specifically with certainty.

Holttum — Tectaria Group 21

CTENITIS

Ctenitis C. Chr. in Verdoorn, Man. Pteridol. (1938) 544; Ching, Bull. Fan Mem. Inst.
Biol. 8 (1938) 375, p.p.; H. Ito in Nakai & Honda, Nov. Fl. Jap. 4 (1939) 205,
p.p.; Copel., Gen. Fil. (1947) 123, p.p.; Holttum, Blumea 31 (1985) 1—38; Gard.
Bull. Sing. 39 (1986) 158. — Dryopteris subg. Ctenitis C.Chr., Kongel. Danske
Vidensk. Selsk. Naturvid. Math. Afh. VII, 10 (1913) 82; ibid. VII, 6 (1920) 31. —
Type species: Ctenitis submarginalis (Langsd. & Fisch.) Ching.

Caudex short, erect or suberect or rarely prostrate, its apex and the bases of stipes den-
sely covered with thin flat scales which are not more than 1 mm wide at their bases, stipes
scaly throughout, with a + abrupt change from the long basal scales to shorter ones of two
different types, distinctive for each species; lamina always bipinnate at the base, the basi-
scopic pinnules of basal pinnae always longer than the acroscopic ones, the basal ones
usually longest and very deeply lobed, in some species pinnate to bipinnate near their
bases; distal pinnae or pinnules + adnate to the rachis and + decurrent at their bases but
their basal basiscopic veins almost always arising from the pinna- or pinnule-midrib; tex-
ture of lamina mostly thin with veins distinct on both sides but in some species opaque
with the smaller veins indistinct; veins all free, those in each pinna-lobe all arising from
the costule of the lobe (not from the costa of the pinna); lower surface of pinna-rachis or
pinna-midrib always bearing scales which are progressively smaller distally, similar scales
also present on costae of pinnules, some ctenitoid hairs also present and often appressed
cylindric unicellular glands, such glands also often present between veins, sometimes
with the addition of short hairs of various kinds; upper surface of pinna-rachis or pinna-
midrib and of costae of pinnules prominent, always covered with ctenitoid hairs which
are thicker than those on the lower surface, similar hairs scattered on veins but not be-
tween veins where appressed cylindric glands or short non-ctenitoid hairs may occur; sori
usually medial on the veins, not terminal; indusia usually present, sometimes very
small and hidden by the mature sporangia, always unpigmented, thin and fragile, usually
bearing marginal cylindric glands when young; spores of two distinct kinds. — Figs.
5-8.

Distribution — Pantropic; c. 100 species (Holttum 1986).

Taxonomy — Christensen’s original study was confined to the species of tropical
America. The first surveys of allied species in Asia were made independently by Ching
(1938) and Ito (1939), both of whom failed to notice the significance of scale-structure
and glands which separate Ctenitis (as here defined) from Tectaria and other genera.
Copeland (1947) added other species which are here included in Tectaria. Holttum (1985)
prepared a list of 47 Old World species included in Ctenitis by previous authors but not
conforming to the present generic concept.

Subdivision of the genus — Christensen (1920) proposed a subdivision of the Ameri-
can species; I have transferred to other genera two of his groups which both extend to
Africa (Holttum 1986). The group which includes the type extends to Africa and Mada-
gascar but not to Asia; its fronds are bipinnatifid. The 43 species of Asia, Malesia and the
Western Pacific (Holttum 1985) are divisible into two groups; as shown in the key, 26

22

Flora Malesiana ser. II, Vol. 2! (1991)

of the 28 Malesian species belong to one group, only 2 (one of them uncertainly) to the
other. No new infrageneric names are here proposed; to establish them effectively a new
conspectus of the whole genus should be made.

la.

KEY TO THE SPECIES

Scales on pinna-rachis narrow with inflexed edges or flat, their bases sometimes
widened but not cordate nor point-attached; isodiametric cells few, nearbase .. 2

. Scales on pinna-rachis flat or nearly so, point-attached at a deeply cordate base;

many isodiametrc/cells in them DaSalipare. - 21) 2 a ee Di

. Basal basiscopic pinnules of basal pinnae not longest, less than twice as long as the

basalfacroscopic pinnules® "2 05 io. os es ete a ele 5 2 3)

. Basal basiscopic pinnules of basal pinnae distinctly longer than the rest, usually

twiceas long asacroscopic OneS*..). 2. Se a es ane) uel suck 9

. Veins thick and prominent on lower surface ............. 1. C. kjellbergii
. Veins not prominent on lower surface, the smaller ones in many cases indistinct 4
. Tertiary leaflets 3 x 3 mm, almost entire, their upper surface densely covered with

SHORGPM AUS Rayer g Rep Ws SPIO tics! ak utes ais 8 ae ese oe ae 2. C. muluensis
. Tertiary leaflets, if present, differently shaped and not thus covered......... 5
. Pinnules or lobes of middle pinnae not lobed nearly to their costae; spreading
bristle-like scales present throughout the main rachis ......... 3. C. aciculata
. Pinnules or lobes of middle pinnae lobed nearly to their costae; such scales not
JOKES) 01 ea ame ey SIMA All ela A ee re arbre eM MPN NACE 5 0 0 6 6 6 6
s ocalesion pinna-rachis flat ornearly SO". 224-2. 22>... > 6 eee Uf
. Scales on pinna-rachis with inflexed edges, at least near their bases......... 8

. Scales on pinna-rachis not widened at their bases; isodiametric cells at base few;

Costalescales very MamOw =e. ss sowie nk locas 4. C. kinabaluensis

. Scales on pinna-rachis widened at their bases, the widened part consisting entirely

of isodiametric cells; costal scales with broad base ........... 5. C. pallens

. Stipe densely scaly throughout, scales at top of stipe 10 mm long

6. C. tabacifera

. Stipe densely scaly near base only; distal scales 3 mm long .. 7. C. subconnexa

. Lamina opaque; smaller veins not distinct on lower surface ... 8. C. propinqua
- Lamina notopaques veins distinct wo... os 6 | ea dS oe 10
. Scales on pinna-rachis very thin, few cells wide, lacking conspicuous isodiametric

EAU ip UR mt te, Oe eat al abhi ath Aly et aan Teta 9. C. boholensis
. Scales on pinna-rachis wider with reflexed margins, widened at their bases with

Some 1sodiametnic'cells oP as oe Se es Shee ee bl
. Indusium very small, hidden by sporangia at maturity, bearing long slender flexu-

OUSTOLANS reer erie ern Renny ata ert oe ote eM tence Copios ake ae Tae ie 12
se indusiumidisinct but trasilesnonlong-frinved 5. a-hr. 4 i eee US
. Basal basiscopic tertiary divisions of basal pinnae lobed ................ 3)

. Basal basiscopic tertiary divisions of basal pinnae entire ................ 14

Holttum — Tectaria Group ; ; _ 23

Pinnae narrowly acuminate; appressed glands present on upper surface
10. C. seramensis

. Pinnae not narrowly acuminate; short erect capitate hairs present between veins on

WP PClASUREACE eaten: 5h RUN wets usea erate ade eRe ere 11. C. bulusanica

. Free pinnae to 8 pairs; basal pinnae of mature plants shorter than suprabasal ones
12. C. vilis

. Free pinnae to c. 4 pairs; basal pinnae longer than suprabasal ones 13. C. silvatica
. Lobes of middle pinnae mostly subentire with rounded apices ............ 16
. Lobes of middle pinnae mostly lobulate, their apices not rounded .......... 9

. Lamina to 13 cm long, bearing many suberect hairs between veins on upper surface

14. C. humilis

. Lamina much longer; hairs on upper surface appressed .... 15. C. subobscura
. Tertiary leaflets of basal pinnae quite free, lobed to their costules, their basal lobes

Leo tanilleate maces ss ee Nee RO te ere gee POP NEC Or an Seay Gara coe MO ee 18

. Tertiary leaflets of basal pinnae adnate and less deeply lobed, their lobes entire . 20
. Lobes of tertiary leaflets of basal pinnae deeply lobulate; thick cylindric glands copi-

OUSIOMBUPPETASUTLACE oa... Gc econ Gee ee Se ee eee ee, ees 16. C. minutiloba
. Lobes of tertiary leaflets of basal pinnae not deeply lobulate; appressed glands on

uppemsurrace neither thickinOMmcoplouS@w ... = oa = oe eee ee 1©
mebasalepinnae toxOOvcm lONS. Sf. 2... ee gee A ts menue hte os 17. C. elata
moasdiepinnac to725 Cm long 2... Sa. 2 Sal hss es omens 18. C. koordersii
mleooules‘or lobes of middle pinnae acutes.... 222.1... 19. C. sumbawensis
meobulcs or lobes of middle pinnae not acutegs sa2 a2 ee dn ee 21
. Basal tertiary leaflets lobed almost to their costules, lobes to 6 pairs . 20. C. croftii
. Basal tertiary leaflets lobed less deeply with fewer lobes ................ DD
wleamina to. 60'cm long; basal pmnae more than;20'cm = 22...4,5.2)5.. 142 - 23
amuna rarely more than 35 cm.long; basal pinna to: lSiem™. 2... 2. 2.8.4. 24

. Costules of basal basiscopic lobes of upper pinnae arising from the main rachis

21. C. decurrentipinnata

. Costules of basal basiscopic lobes of upper pinnae arising from costae of pinnae

22. C. alteroblumei

. Basal basiscopic pinnules of basal pinnae about twice as long as acroscopic ones . 25
. Basal basiscopic pinnules of basal pinnae much less than twice as long as acro-

SEOPIC COMES hw Aen era ale IO Si Oe ae tea en gn on 26. C. angusta

. Tertiary leaflets of basal pinnae deeply lobed at their bases .............. 26
. Tertiary leaflets of basal pinnae lobed less than halfway to their costules

25. C. erythradenia

. Tertiary leaflets to 18 x 10 mm; scales on rachis very narrow, not stiffly spreading

23. C. setosa

. Tertiary leaflets to 8 x 4 mm; scales on rachis stiffly spreading 24. C. atrorubens
. Tertiary leaflets of basal pinnae and lobes of distal pinnae widely spaced, narrow,

al CALS Moat see on eet ee ren ee en Nem Ue Mae ee OO es 27. C. paleolata

. Tertiary leaflets of basal pinnae and lobes of distal pinnae almost contiguous

28. C. subglandulosa

24 Flora Malesiana ser. II, Vol. 2! (1991)

aie
Fi yo
2

sg
Z; 44 Rp

PU. Eowarns

Fig. 5. Ctenitis aciculata (Baker) Ching. a. Base of a basal pinna, x 0.6; b. third acroscopic pinnule of a
basal pinna, x 2.2; c. detail of b, x 23; d. upper surface of pinna rachis showing ctenitoid hairs, x 20. —
C. vilis (Kunze) Ching. e. A basal pinna, x 0.85; f. lower surface of pinna rachis, x 23; g. a middle
pinna, basal acroscopic pinnule, x 4; h. sorus, the indusium bearing long flexuous glands, x 46;
i. subentire distal lobes of a middle pinnule, x 4 (a—d: Parris & Croxall 8941; e-i: Kunstler 3038).

Holttum — Tectaria Group

25

1. Ctenitis kjellbergii (C. Chr.) Ching, Sun-
yatsenia 5 (1940) 250; Holttum, Blumea 31
(1985) 13. — Dryopteris kjellbergii C.Chr.,
Bot. Jahrb. Syst. 66 (1933) 45. — Type: Kjell-
berg 3260, Celebes, Porema (S).

Caudex short, erect, covered with thin castane-
ous scales 8 x 0.5 mm; stipe to 17 cm long,
densely scaly, scales above base 3—4 mm long;
lamina to 26 x 12 cm; pinnae to 8 pairs; basal
pinnae to 7 x 3.5 cm with 2 pairs of free pinnules,
subequal on the two sides of the pinna-rachis,
largest pinnule c. 17 x 7 mm, deeply lobed, the
lobes crenate with rounded apices; distal pinnae
little decurrent at their bases; veins distinctly
prominent on the lower surface; rachis densely
scaly on lower surface, scales to 2.5 x 0.3 mm,
flat, gradually attenuate, with a few isodiametric
cells near the base, rest of cells oblong, all with
rather thick walls, margins of scales minutely den-
ticulate; lower surface of costae of pinnules and
pinna-lobes similarly scaly near their bases, dis-
tally bearing red glands; red cylindric glands also
present on veins, between veins slender pale glands
sometimes present; sori inframedial on the veins;
indusia reniform, thin, bearing superficial red

glands.
Distribution — Malesia: Central Celebes, W
New Guinea (Japen Is.).

Ecology — Type from forest at 1200 m; Japen
Island specimen ‘on the face of limestone rocks’,
at 1100 m.

2. Ctenitis muluensis Holttum, Fern Gaz. 12
(1984) 320; Blumea 31 (1985) 14. — Type:
Jermy 14156, Sarawak, G. Mulu Nat. Park (BM;
iso BO, K).

Stipe to 15 cm long, slender, dark brown, mi-
nutely hairy throughout; basal scales 7 mm long,
0.5 mm wide at their bases, filiform distally, me-
dium brown, thin; scales above base gradually
shorter and darker, very narrow with widened bases;
lamina to 23 cm long, to 10 cm wide at the base,
texture firm and opaque; pinnae to 10 pairs; basal
pinnae to 7 cm long (including stalk of 5 mm)
bearing several pairs of pinnules, basal pinnules
on the two sides about equal, to 16 x 7 mm with
obtuse apex, bearing | or 2 pairs of tertiary leaflets
which are 3 mm long, not quite 3 mm wide, with
truncate base and broadly rounded slightly crenate

apex, other pinnules gradually smaller, the middle
ones deeply lobed near their bases only; lower sur-
face of pinna-rachis and costae of pinnules bearing
very short hairs and copious scales, the largest 3
mm long, 0.2—0.3 mm wide at their bases, flat,
very narrow distally, cells near base isodiametric;
lower surface of leaflets, on and between veins,
bearing many appressed pallid cylindric glands 0.1
mm long; upper surface of pinna-rachis bearing
scales as lower, also abundant thicker short hairs;
upper surface of leaflets densely covered with very
short erect hairs and pale appressed glands; sori
about medial on veins; indusia very small, fugaci-
ous, bearing minute hairs.

Distribution — Malesia: Borneo (G. Mulu Na-
tional Park).

Ecology — In montane limestone forest at 1000—
1700 m.

3. Ctenitis aciculata (Baker) Ching, Bull. Fan
Mem. Inst. Biol. 8 (1938) 292; Holttum, Blu-
mea 31 (1985) 14. — Nephrodium aciculatum
Baker, J. Linn. Soc. Bot. 22 (1886) 226. — Dry-
opteris aciculata (Baker) C.Chr., Index Filic.
(1905) 250; Alderw., Malayan Ferns (1908) 200.
— Type: G.F. Hose 86, Sarawak, G. Matang (K;
iso P).

Caudex erect, to 50 cm tall; stipe to more than
60 cm long, basal scales to 25 mm long and 1 mm
wide, dull brown, thin, decrescent upwards, scales
on upper part and on rachis 5—7 mm long, copi-
ous, very narrow, stiffly spreading with edges in-
rolled; lamina to 75 cm long, firm, opaque; free
pinnae to 5 pairs and 5 pairs adnate; basal pinnae
to 28 cm long, basal pinnules free, then a few
pairs increasingly adnate, basal basiscopic pinnule
to 9.5 x 2 cm, not or little longer than the next
one, deeply lobed, the lobes entire, basal acroscop-
ic pinnule to 4 x 1.2 cm, pinnules and lobes of
middle pinnae not deeply lobed; lower surface of
pinna-rachis bearing very narrow spreading scales
3-4 mm long; some cylindric glands present on
lower surface of pinna-lobes; smaller veins not
evident on either surface; sori mostly on both
branches of a forked vein; indusia very small,
bearing when young elongate glands. — Fig. 5a—d.

Distribution — Malesia: Sumatra, Java, Borneo
(widely), Philippines (Mindanao), SE New Guinea.

Ecology — In forest at altitudes up to 1000 m,
sometimes on rocks or tree-trunks.

26

Flora Malesiana ser. II, Vol. 2! (1991)

4. Ctenitis kinabaluensis Holttum, Blumea
31 (1985) 16. — Type: Holttum SFN 25253,
Sabah, Mt Kinabalu (K).

KEY TO THE VARIETIES

la. Scales on pinna-rachis thin and narrow; upper
surface of lamina not glandular

a. var. kinabaluensis

b. Scales on pinna-rachis very firm; upper surface

densely glandular . b. var. crassisquama

a. var. kinabaluensis

Stipe to 70 cm long, basal scales light brown,
15—25 mm long, hardly 1 mm wide, upper scales
5-6 x 0.3 mm; lamina to 70 cm long, subcoria-
ceous, opaque; basal pinnae to 30 cm long, bear-
ing 6 pairs of pinnules, basal basiscopic pinnule
10 x 3 cm with a single lobed adnate tertiary leaf-
let; pinnules of suprabasal pinnae deeply lobed;
veins hardly distinct on either surface; lower sur-
face of pinna-rachises bearing sparse very narrow
flat scales, upper surface densely covered with
ctenitoid hairs; lower surface of pinnules between
veins bearing slender appressed glands, upper sur-
face glabrous; sori medial on veins; indusia mi-
nute, thin, when young bearing slender glands.

Distribution — Malesia: NE Borneo.

Ecology — In forest at 1000-1500 m, once re-
ported on rocks.

b. var. crassisquama Holttum, Blumea 31
(1985) 16. — Type: K. Iwatsuki et al. B2195,
E Kalimantan (K).

Lamina thinner and veins more distinct; scales
of rachises wider and thicker; upper surface of pin-
nae between veins bearing many appressed glands.

Distribution — Malesia: Borneo (known only
from the type).

5. Ctenitis pallens (Brackenr.) M.G. Price,
Kalikasan 12 (1983) 155; Holttum, Blumea 31
(1985) 15, f. la. — Lastrea pallens Brackenr. in
Wilkes, U.S. Expl. Exped., Filic. 16 (1854)
197. — Type: U.S. Expl. Exp. 1838-1842
s.n., Luzon, forest near Bafios (US).

Dryopteris rizalensis Christ, Philipp. J. Sci. 2
(1907) Bot. 216, not Christ 1906. — Ctenitis

rizalensis (Christ) Copel., Gen. Fil. (1947)
125; Fern Fl. Philipp. (1960) 290. — Type:
Copeland 1659, Mindanao (MICH).

Stipe to more than 50 cm long; basal scales 20
mm long, narrow, thin, scales for 15 cm above
base gradually shorter, those on distal part and on
rachis 4-5 mm long, not stiffly spreading, flat,
very narrow with widened bases; lamina to 75 cm
long, firm and opaque; pinnae to 10 pairs; basal
pinnae to 30 cm long (stalk 2 cm) with 4 pairs of
free and 3 or 4 pairs of adnate pinnules, basal basi-
scopic pinnule to 13 cm long with 1 pair free and
2 or 3 pairs adnate tertiary leaflets, basal tertiary
leaflet 3.5 x 1.4 cm, obtuse, lobed nearly to its
costa at the base, second basiscopic pinnule not
longer than the first, basal acroscopic pinnule to
8.5 cm long with a pair of tertiary leaflets; larger
pinnules on second pair of pinnae to 7.5 x 1.9 cm,
deeply lobed, costules of lobes to 7 mm apart; cos-
tules of pinna-lobes distinct on the lower surface
but not the veins; lower surface of pinna-rachis
bearing copious scales like those of the main ra-
chis but smaller, clathrate at their bases; on both
surfaces between veins some appressed pale glands;
sori about medial on each side of costules of pin-
nule-lobes; indusia very small, fringed with many
pale cylindric glands. — Fig. 6a.

Distribution — Malesia: Philippines (Luzon,
Samar, Negros, Mindanao), New Ireland.

Ecology — In forest at altitudes up to 600 m.

Note — The type of Dryopteris rizalensis Christ
1906 (Loher s.n., Rizal Prov., Luzon) has not been
located; it might represent another species.

6. Ctenitis tabacifera (Alderw.) Ching, Sun-
yatsenia 5 (1940) 250; Holttum, Blumea 31
(1985) 16. — Dryopteris tabacifera Alderw..,
Bull. Jard. Bot. Buitenzorg III, 2 (1920) 147. —
Type: Kornassi 1543, Ceram (BO; iso L).

Stipe to 63 cm long, very densely scaly through-
out; basal scales to 15 x 1 mm, thin, medium
brown, those above base gradually shorter but
without change of colour, largest on upper part of
stipe 8—9 mm long, very narrow, rather weakly
spreading, almost flat, their bases a little widened
with a few isodiametric cells; rachis-scales similar,
on distal part 3—4 mm long; lamina to 70 cm
long, texture very firm, opaque; free pinnae c. 12
pairs; basal pinnae to 28 cm long with 7 pairs of

Holttum — Tectaria Group

Fig. 6. Spores (SEM), all except f, x c. 860. — a. Ctenitis pallens (Brackenr.) M.G. Price; b. C. propin-
qua (Presl) Copel.; c. C. boholensis Holttum; d. C. alteroblumei Holttum; e. C. subglandulosa (Hance)
Ching; f. ibid., x c. 3260 (a: Jermy 7900, New Ireland; b: Cuming 255, Luzon; c: Ramos BS 42983,

Bohol; e, f: Griffith s.n., Assam).

free or adnate pinnules, basal basiscopic pinnule to
10 x 3.5 cm, bearing 1 pair of free deeply lobed
tertiary leaflets to 12 x 7 mm; largest acroscopic
pinnule 7.5 x 2.3.cm, also with a free tertiary
leaflet; middle pinnae bearing c. 6 pairs of deeply
lobed pinnules, their lobes entire or the basal ones
crenate; veins in pinnule-lobes mostly forked, only
the part below the fork visible on the lower sur-
face; scales on lower surface of pinna-rachis flat

except near the base, to 8 cells wide, cells all elon-
gate except a few near the base, many short hairs
also present, no glands on surface between veins;
upper surface of pinna-rachis densely covered with
short thick hairs and some scales; surface of pin-
nules between veins bearing a variable number of
appressed glands; sori at forks of veins in pinnule-
lobes; indusia small and very thin, bearing many
slender glands.

28

Distribution — Malesia: Moluccas (Ceram and
Ambon).
Ecology — In forest at 600 m.

7. Ctenitis subconnexa (Christ) Holttum,
Blumea 31 (1985) 17. — Phegopteris subcon-
nexa Christ in Warb., Monsunia 1 (1900) 83;
Alderw., Malayan Ferns (1908) 496. — Dryo-
pteris subconnexa (Christ) C. Chr., Index Filic.
(1905) 295. — Type: Warburg 17864, Batjan,
Mt Sibella (B).

KEY TO THE VARIETIES

la. Scales on pinna-rachis 3 mm long, very nar-
Tow with few isodiametric cells at base

a. var. subconnexa

b. Scales on pinna-rachis 1—-1.5 mm long, basal
1/3-1/2 formed by isodiametric cells

b. var. alstonii

a. var. subconnexa

Stipe to 52 cm long; basal scales in a tuft, thin
and crinkled, to 20 mm long, 1 mm wide at the
base, apex filiform; scales above base gradually
shorter for 10 cm, on distal part of stipe and on
rachis 2—3 mm long, very narrow, flat with +
widened base; /amina to 70 cm long, texture firm;
pinnae widely spaced, 6 pairs free and 3 or 4 pairs
increasingly adnate; basal pinnae c. 27 cm long
bearing 4 pairs of free pinnules; basal basiscopic
pinnule 8.5 x 3 cm with 1 pair of almost free ter-
tiary leaflets 20 x 7 mm, deeply lobed at their
bases, costules of the lobes 3 mm apart; basal
acroscopic pinnule 7 x 3 cm, also with tertiary
leaflets; lower surface of pinna-rachis bearing
scales 3 mm long, very narrow with widened base,
also copious short hairs; lower surface of pinnae
between veins (which are not distinct) bearing ap-
pressed glands and short erect hairs, glands also
present on upper surface; sori medial on veins; in-
dusia thin and very small, bearing slender flexuous
marginal glands, similar glands on stalks of spo-
rangia.

Distribution — Malesia: Philippines (Batjan, two
collections from Mt Sibella at 760 and 1370 m
altitude).

Flora Malesiana ser. II, Vol. 2! (1991)

b. var. alstonii Holttum, Blumea 31 (1985) 17
— Type: Alston 16937, Batjan, Mt Sibella,
1200 m (BM).

Scales at base of stipe 10 mm long; scales of
rachises 1—-1.5 mm long with dilated bases.

Distribution — Malesia: Philippines (known
from the type only).

8. Ctenitis propinqua (Pres!) Copel., Fern Fl.
Philipp. (1960) 289; Holttum, Novit. Bot.
Delect. Seminum Horti Bot. Univ. Carol. Prag.
1968 (1969) 37; Blumea 31 (1985) 18, f. 1f. —
Lastrea propinqua Presl, Epim. Bot. (1851) 38,
excl. Cuming 80, 151 & 252. — Nephrodium
preslii Baker, Syn. Fil. (1867) 272 (not N. pro-
pinquum R. Br.). — Lectotype (Baker): Cuming
255, Luzon, Ilocos Norte (PRC; BM, K, L).

Stipe incomplete on all specimens seen, basal
scales not known; scales on distal part and on ra-
chis to 3 mm long, narrow, their margins inrolled
above the widened base of isodiametric cells; /a-
mina to 28 cm long, texture firm; pinnae well
spaced, 3 or 4 pairs free and 2 or 3 pairs adnate;
basal pinnae to 8.7 cm long bearing 1 pair of free
and 1 pair of adnate pinnules; basal basiscopic pin-
nule 3.5 x 1.7 cm, distinctly longer than the next
one, very deeply lobed at its base, the lobes entire,
basal acroscopic pinnule 2.2 x 1.4 cm; second pair
of pinnae 8 cm long with 2 pairs of adnate pinnules;
veins in pinnules mostly simple, their distal parts
not distinct on the lower surface; appressed red
cylindric glands present on lower surface of cos-
tules of lobes and on veins; sori medial or infra-
medial on pinnule-lobes; indusia persistent, bear-
ing red glands. — Fig. 6b.

Distribution — Malesia: Philippines (except for
the type, known from one other collection from
Ilocos Norte).

Note — In 1851 Presl, copying John Smith, cited
four Cuming numbers: 80, /57, 252, 255. When
publishing the new name Nephrodium preslii,
Baker cited only Cuming 255. The other Cuming
numbers are now assigned to the species Ctenitis
erythradenia, silvatica and decurrentipinnata.

9. Ctenitis boholensis Holttum, Blumea 31
(1985) 18. — Type: Ramos BS 42983, Bohol
(K; UC).

Holttum — Tectaria Group

Stipe to 53 cm long, light castaneous, near base’
covered with scales 12 x 0.5 mm, scales on distal
part 4 mm long, very narrow, not spreading; la-
mina 36 cm long, thin; free pinnae 3 pairs and 6
pairs adnate; basal pinnae 7 cm below the next
pair, 14 cm long, basal basiscopic pinnules 6.2 x
2.6 cm, lobed almost to the costa, largest lobes 14
x 6 mm, lobed halfway to their costules; basal
acroscopic pinnules 3.7 x 1.4 cm, with crenate
lobes; suprabasal pinnae 12.5 cm long, bearing
adnate deeply lobed pinnules, the pinnule-lobes
mostly entire; veins slender, slightly prominent on
the lower surface; lower surface of pinna-rachises
bearing almost flat narrow scales 1-2 mm long
and slender appressed hairs; sori inframedial in pin-
nule-lobes; indusia thin but persistent, pilose. —
Fig. 6c.

Distribution — Malesia: Philippines (except for
the type, known only from one other collection
from Bohol).

10. Ctenitis seramensis Holttum, Blumea 31
(1985) 19. — Type: De Vriese & Teijsmann
324, Ceram (L 908,294-175).

Basal stipe-scales light brown, narrow, hair-
pointed; upper ones 4—5 mm long with inflexed
margins and basal isodiametric cells; lamina to 45
cm long; free pinnae to 9 pairs, narrowly acumi-
nate; basal pinnae to 16 cm long, bearing 4 pairs
of pinnules, the lowest stipitulate; basal basiscop-
ic pinnule 7 x 2.8 cm, acuminate, deeply lobed,
basal lobe almost free, 19 x 7 mm, deeply lobu-
late, basal acroscopic pinnule 2.5 x 0.9 cm, ob-
tuse; pinnules of suprabasal pinnae obtuse, deeply
lobed, costules of lobes 3.5 mm apart; sori about
medial in lobes of pinnules; indusia very small,
bearing slender glands, glands also present on
stalks of sporangia. — Fig. 7.

Distribution — Malesia: Moluccas (Ceram and
Ambon).

Note — Robinson 1956 from Ambon (BO) was
named Dryopteris intermedia by Alderwerelt.

11. Ctenitis bulusanica Holttum, Blumea 31
(1985) 19. — Type: Elmer 16681, Luzon, Mt
Bulusan (K; iso BM, BO, L, UC).

Stipe to 28 cm long; basal scales 8 x 0.5 mm,
distal scales dark brown, to 5 mm long; lamina to

Fo: Epwarps

Fig. 7. Ctenitis seramensis Holttum. a. Base of a
basal pinna showing difference between basal basi-
scopic and acroscopic pinnules, x 0.6; b. apex of a
middle pinna, x 2; c. upper surface of pinna-rachis
showing ctenitoid hairs and unicellular glandular
hairs between veins, x 20 (a—c: Kato et al. 8031).

23 cm long, firm, brown-olivaceous when dried:

' free pinnae 2 or 3 pairs; basal pinnae to 10 cm
_long including a stalk of 7 mm long, basal basi-

scopic pinnules sessile, to 5 cm long, 2.5 cm wide
at their bases, their basal lobes almost free, 18 x 7
mm, lobed 2/3 towards their costules, lobules 4
pairs; suprabasal pinnae sessile with 1 pair of ad-
nate pinnules; lower surface of pinna-rachises bear-
ing narrow scales 2—3 mm long, their margins

30

Flora Malesiana ser. II, Vol. 2! (1991)

inflexed near the base; upper surface of pinnules
bearing short capitate hairs between veins; sori
inframedial on veins; indusia very small, bearing
slender glands.

Distribution — Malesia: Philippines (known on-
ly from the type collection).

12. Ctenitis vilis (Kunze) Ching, Bull. Fan
Mem. Inst. Biol. 8 (1938) 290; Holttum,
Revis. Fl. Malaya 2 (1955) 406, f. 293; Blumea
31 (1985) 20. — Aspidium vile Kunze, Bot.
Zeitung (Berlin) 4 (1846) 474. — Dryopteris
vilis (Kunze) C. Chr., Index Filic. (1905) 300;
Rosenst., Meded. Rijksherb. 31 (1917) 6;
C. Chr., Gard. Bull. Straits Settlem. 4 (1929)
391; Backer & Posth., Varenfl. Java (1939) 43,
p.p. — Type: Zollinger 1602, Java (B?; iso
BO} KG. P):

Polypodium asplenioides Bory in Bélanger, Voy.
Indes Or. 2 (1833) 33, non Sw. 1801. — Type:
Bory s.n., Java (P).

Lastrea blumei T. Moore, Index Fil. (1858) 94. —
[Aspidium intermedium Blume, Enum. Pl. Javae
(1828) 161, non Willd. 1810.] — Type: Rein-
wardt s.n., Java, G. Burangrang (L 899,314-2).

Nephrodium sarawakense Baker, J. Linn. Soc. Bot.
22 (1886) 225. — Type: Hose 95, Sarawak (K).

Lastrea padangensis Bedd., Handb. Ferns Brit. India
Suppl. (1892) 60. — Type: Kunstler 3038,
Perak (K; iso BM, L, P).

Dryopteris millettii C. Chr., Index Filic. (1905)
278 [not D. setosa (Blume) Kuntze]. — Ne-
phrodium setosum Baker, Syn. Fil. (1867) 274.
— Type: Millett s.n., Java (K).

Stipe to 20 cm long; basal scales 10 mm long,
light brown, scales above base darker, 3-5 mm
long, very narrow with inflexed margins; lamina
commonly 20 cm long (to 35 cm); to 8 pairs of
free pinnae and 5 or 6 pairs adnate; basal pinnae of
mature plants a little shorter than the next pair,
commonly 7 cm long with basal basiscopic pin-
nule 3.5 x 1.4 cm, deeply lobed; middle pinnae
with one pair of free deeply lobed pinnules, distal
part of these pinnae lobed almost to the costa,
lobes oblique, acute, middle ones entire; lower
surface of costae of pinnae bearing very narrow
scales which are widened with some isodiametric
cells at their bases; sori medial on pinna-lobes;
indusia very small, often hidden by sporangia,

bearing many very slender flexuous pale glands. —
Fig. 5e-i.

Distribution — Malesia: Sumatra, Malay Penin-
sula and Peninsular Thailand, Java, Borneo (Sara-
wak), Moluccas (Ambon).

Habitat — Among rocks by streams, often in the
flood-zone, less common in rocky crevices away
from streams, at low to medium altitudes; the type
doubtfully from a mountain summit.

Taxonomy — Under Aspidium intermedium
Blume cited only one collection, from Burangrang.
The only specimen in the Leiden Rijksherbarium
with this name and locality written by Blume was
collected by Reinwardt, and is evidently the type.
It is an unusually large specimen with lamina
39 cm long but only 11 cm wide (the pinnae are
much upcurved). It is closely similar to the type of
Nephrodium setosum Baker. However, in the her-
barium Blume also identified other specimens as
Aspidium intermedium, and sent a pinna of one of
them to Hooker; that specimen is the type of C.
alteroblumei.

The fern described under the name Aspidium vile
by Raciborski in Pteridop. Buitenzorg (1898) is
Macrothelypteris setigera (Blume) Ching; appar-
ently this is also true of Alderw., Malayan Ferns
(1908) 199.

13. Ctenitis silvatica Holttum, Blumea 31
(1985) 20. — Type: M.G. Price 2042, Luzon,
Mt Makiling (K).

Ctenitis vilis auct.: Copel., Fern Fl. Philipp.
(1960) 288.

Ctenitis setosa auct.: Holttum, Novit. Bot. Delect.
Seminum Horti Bot. Univ. Carol. Prag. 1968
(1969) 20, for Cuming 181 only.

Closely related to C. vilis (Kunze) Ching but
with broader and shorter fronds; pinnae to 4 pairs,
basal ones commonly 9 cm long, longer than the
next pair.

Distribution — Malesia: Philippines (many lo-
calities), New Guinea.

Habitat — In lowland forest, not confined to
rocks nor to the flood-zone beside streams.

14. Ctenitis humilis Holttum, Blumea 31
(1985) 22. — Type: Ramos BS 43032, Bohol
(SING; iso MICH, P, UC).

Holttum — Tectaria Group

3]

Allied to C. boholensis Holttum, differing as
follows: fronds much smaller; scales on upper part
of stipe 3 mm long, stiffly spreading, with inrolled
edges; lower surface of pinna-rachises bearing erect
hairs; upper surface of pinnae, between veins, bear-
ing many suberect hairs. Stipe c. 5 cm long; lamina
to 11 cm long; basal pinnae 2.8 cm long with 1
pair of pinnules; basal basiscopic pinnule 13 x 4
mm, deeply lobed near its base; middle pinnae
obtuse, lobed almost to their costae, the lobes fal-
cate, obtuse and mostly entire; sori as in C. bohol-
ensis.

Distribution — Malesia: Philippines (Bohol,
Mindoro, one collection from each).

Habitat — Bohol: ‘at 600 m on rocks in damp
forest’. Mindoro: ‘on a steep cliff’.

Note — This may be a very dwarf habitat-form
of C. boholensis; further collections are needed.

15. Ctenitis subobscura (Christ) Holttum, Fem
Gaz. 12 (1984) 320; Blumea 31 (1985) 23. —
Phegopteris subobscura Christ, Bull. Herb.
Boissier 6 (1898) 836. — Type: G. Schneider
35, Sumatra (P).

Dryopteris squamulifera Alderw., Bull. Jard. Bot.
Buitenzorg II, 16 (1914) 9. — Type: C.G.
Matthew 669, (BO; iso K), Sumatra, Padang
Panjang.

Lastrea intermedia var. blumei auct.: Bedd., Handb.
Ferns Brit. India Suppl. (1892) 61.

Ctenitis manii auct.: Holttum, Revis. Fl. Malaya
2 (1955) 497, for the Malesian plants.

Stipe to 40 cm long, bearing throughout very
narrow dark spreading scales, basal ones 12 mm
long, gradually shorter distally; Jamina to 55 cm
long; pinnae closely placed, to 15 or more pairs,
almost at right angles to the rachis; basal pinnae
to 16 cm long, those on the largest fronds with a
few pairs of free pinnules; basal basiscopic pinnule
to c. 6 x 1.6 cm, lobed almost to its costa, costules
to 5 mm apart, lobes entire; basal acroscopic pin-
nule 2 cm long, lobed near its base only; supra-
basal pinnae sessile, acuminate, very deeply lobed,
lobes mostly subentire and obtuse, of firm texture;
lower surface of rachis and costae bearing scales like
those of the stipe but more sparse, those on costae
with many isodiametric cells in their inflexed basal
part; upper surface of costae densely short-hairy,
a few similar hairs also between veins; veins in

pinna-lobes simple; sori medial on veins; indusia
very small, bearing short slender glands.
Distribution — Malesia: widely in Western Ma-
lesia, also Philippines (Mindanao).
Habitat — In rocky places in forest, often on
stream banks.

16. Ctenitis minutiloba Holttum, Blumea 31
(1985) 23. — Type: W. Meijer SAN 20267,
Sabah, Ranau Dist. (K).

Stipes closely tufted, at the base only bearing
flat brown scales, scales above base darker, seti-
form, the lower ones 10 mm long, shorter upwards;
lamina c. 35 cm long; basal pinnae to 23 cm long
bearing 10 pairs of pinnules, the basal basiscopic
one 12 x 5 cm bearing 7 pairs of deeply lobed ter-
tiary leaflets, 20 x 8 mm, their lobes 4 x 1.5 mm
and lobulate; basal acroscopic pinnule 5 x 1.8 cm,
bearing lobed tertiary leaflets to 10 x 6 mm; mid-
dle pinnae bearing several pairs of well-spaced
lobed pinnules, the lobes to 5 x 3 mm, deeply
lobulate; lower surface of pinna-rachises bearing
scattered narrow scales 2 mm long with inflexed
margins and isodiametric cells at the base; upper
surface of all leaflets copiously glandular, the
glands pale, thick, appressed, c. 0.1 mm long; sori
at apices of veins; indusia firm, reniform, some-

times bearing glands.
Distribution — Malesia: Borneo (Sabah, 2 col-
lections).

Habitat — In forest at 600-1000 m.

17. Ctenitis elata Holttum, Blumea 31 (1985)
24, new name for Dryopteris subarborea var.
glabrior Alderw., Bull. Jard. Bot. Buitenzorg I,
16 (1914) 57. — Type: C.G. Matthew 653,
Sumatra, Korinchi Peak at 1800—2100 m (K;
iso BO).

Stipe to more than 60 cm long, its basal scales
flat, 12 x 1 mm, scales above base 5 mm long,
the basal part clathrate with inflexed margins; /a-
mina to c. 100 cm long; suprabasal pinnae to 60
cm long bearing 12 pairs of well-spaced pinnules,
mostly stalked, the largest, 15 x 5.5 cm bearing 5
pairs of deeply lobed tertiary leaflets, the largest
3.2 x 1 cm, its basal lobes crenate, their costules 4
mm apart; lower surface of rachises bearing scales
2 mm long, dilated and clathrate at their bases,

32

Flora Malesiana ser. II, Vol. 2! (1991)

margins above base inflexed, also many short
hairs; sori near the costules of lobes of pinnules;
indusia small, glandular.

Distribution — Malesia: Sumatra (Korinchi Peak,
2 collections).

Note — The above description is made from
Matthew’s own specimen, which he gave to Kew;
it is much larger than the one he sent to Alderwerelt
but does not include a basal pinna. The second col-
lection (by Holttum, at SING) is about the same
size as Matthew’s at BO.

18. Ctenitis koordersii Holttum, Blumea 31
(1985) 25. — Type: Koorders 17027, Celebes,
Minahassa (BO).

Aspidium obtusilobum auct.: Christ, Ann. Jard.
Bot. Buitenzorg 15 (1898) 130, not Nephro-
dium obtusilobum Baker.

Dryopteris zeylanica Alderw., Malayan Ferns (1908)
203, nomen nov. superfl. for D. obtusiloba
(Baker) Kuntze, for plants of Celebes only.

Base of stipe not preserved; scales on distal part
3 mm long, very narrow, convex and clathrate at
their bases; /amina to at least 50 cm long, thin;
basal pinnae 18—20 cm long with 2 or 3 pairs of
free and 2 or 3 pairs of adnate pinnules, the pin-
nules contiguous or imbricate; basal basiscopic
pinnule 9 x 3.5 cm, bearing several pairs of most-
ly adnate tertiary leaflets, the largest ones 20 x 7
mm, deeply lobed, the lobes 6 pairs, lowest ones
crenate, their costules 3 mm apart; basal acroscopic
pinnule 5.4 x 2 cm bearing 4 pairs of adnate terti-
ary leaflets; pinnules of upper pinnae all deeply
lobed; pinna-rachises covered on the lower surface
by many short scales; upper surface of pinnules
between veins bearing variable slender appressed
glands; sori medial in pinnule-lobes; indusia small,
thin, glanduliferous.

Distribution — Malesia: Celebes (Minahassa, 2
collections).

Note — Nephrodium obtusilobum Baker occurs
only in Ceylon; it belongs to the genus Dryopsis
Holttum & Edwards, the scales of which are very
different from those of Ctenitis.

19. Ctenitis sumbawensis Holttum, Blumea
31 (1985) 26. — Type: Kostermans 18126, W
Sumbawa, Mt Batu Lanteh, 800 m (K;; iso L).

Stipe 40 cm long, slender, at its base bearing
light brown scales 12 mm long with filiform
apices, above the base dark brown spreading scales
with inflexed margins; lamina 40 cm long, thin;
pinnae many pairs, stalked, close to each other;
basal pinnae 17 cm long, its free pinnules 6 pairs,
basal basiscopic pinnule 8—9 cm long with 5 pairs
of deeply lobed tertiary leaflets, the lobes entire
and slightly falcate; basal acroscopic pinnule 3.5 x
1.2 cm, bearing a single tertiary leaflet; pinnules
or lobes of upper pinnae all deeply lobed, the lobes
acute; veins in pinnule-lobes pinnate, the veinlets
simple; lower surface of pinna-rachises bearing
narrow scales 2-3 mm long, clathrate at their
bases, upper surface bearing many hairs to 0.5 mm
long; upper surface of pinnules between veins
bearing slender appressed hairs; sori inframedial on
veins; indusia small, persistent.

Distribution — Malesia: Lesser Sunda Islands
(Sumbawa, Lombok).

20. Ctenitis croftii Holttum, Blumea 31 (1985)
26. — Type: J.R. Croft 583, Papua New Gui-
nea, Mt Misim (K; iso LAE).

Stipe 50 cm long, basal scales narrow, flat, thin,
red-brown, 20 mm long, scales above base gradual-
ly shorter, upper ones 3—4 mm long, spreading,
with inflexed margins; lamina to 50 cm long, firm;
pinnae 12 pairs; basal pinnae to 23 cm long, bear-
ing 3 or 4 pairs of free and several pairs of adnate
pinnules; basal basiscopic pinnules to 8.7 x 2.4
cm, bearing 1 pair of tertiary leaflets which mea-
sure 14 x 6 mm, deeply lobed, the lobes obtuse;
basal acroscopic pinnules 4 cm long; lower surface
of pinna-rachises bearing narrow scales with dilated
clathrate bases and short hairs, upper surface bear-
ing dense thick hairs more than 1 mm long; sori
near costules of lobes of pinnules; indusia minute,
thin, bearing red cylindric glands.

Distribution — Malesia: Papua New Guinea
(Morobe Prov., several collections).

Habitat — At 1000-1600 m, on steep slopes or
stream banks in forest.

Taxonomy — This species is closely allied to C.
fijiensis (Hook.) Copel. but has less deeply divid-
ed pinnules, much smaller indusia, elongate basal
basiscopic pinnules, and stipe-scales gradually de-
crescent upwards.

Holttum — Tectaria Group

33

21. Ctenitis decurrentipinnata (Ching)
Ching, Bull. Fan Mem. Inst. Biol. 8 (1938)
291; Tard. & C.Chr., Fl. Indo-Chine (1941)
346, f. 40; Holttum, Blumea 31 (1985) 27. —
Dryopteris decurrentipinnata Ching, Bull. Fan
Mem. Inst. Biol. 2 (1931) 195, t. 9. — Type:
McClure 8673, Hainan (LU; iso K, SING).

Lastrea propinqua auct.: Presl, Epim. Bot. (1851)
38, p.p.

Caudex massive, suberect; stipe to 75 cm long,
stramineous except at base; basal scales 15 x 1 mm,
medium to dark brown, firm, flat; scales above
base 5—6 mm long, stiffly spreading with inrolled
edges and widened clathrate base; lamina to 50 cm
or more long, thin; pinnae at least 10 pairs, basal
ones 22—30 cm long bearing 5 pairs of free or ad-
nate pinnules; basal basiscopic pinnules 10 cm
long with 1 pair of free deeply crenate tertiary leaf-
lets; basal acroscopic pinnules 4 cm long; second
pair of pinnae 22 cm long, their largest pinnules 6
x 1.7 cm with basal lobes crenate, costules of the
lobes 4 mm apart, several middle pinnules with
much-decurrent basal lobes; upper pinnae with
decurrent basal lobes the costules of which arise
from the rachis; scales of lower surface of pinna-
rachis 2—4 mm long, their edges inrolled, many
cells near their bases clathrate; hairs on upper sur-
face of pinna-rachis 0.5 mm long; sori on the mid-
dle of veins in pinnule-lobes; indusia small, thin,
gland-bearing.

Distribution — Hainan, Vietnam; in Malesia:
Philippines (Luzon).

22. Ctenitis alteroblumei Holttum, Blumea
31 (1985) 27, f. 1c, 3a, b. — Type: Blume s.n.,
Java (L 908,337-1011).

Nephrodium blumei auct.: Hook., Sp. Fil. 4 (1862)
135, for the fragment ex Herb. Blume only, non
Lastrea blumei T. Moore 1858 nec L. blumei
sensu Bedd., Ferns S. India (1864) t. 249.

Aspidium pulvinuliferum auct.: Racib., Pteridop.
Buitenzorg (1898) 177. — Dryopteris pulvinuli-
fera auct.: Alderw., Malayan Ferns (1908) 200.

Caudex short, erect, its apex covered with red-
brown scales to 28 x 1 mm; stipe to more than 60
cm long, its basal scales like those of the caudex,
those above the base dark, bristle-like, spreading at
right angles, 10-12 mm long decreasing distally

to 6-7 mm; lamina to more than 60 cm long;
basal pinnae to 26 cm long (including stalk 1.5
cm) bearing c. 6 pairs of free and 2 or 3 pairs of
adnate pinnules; basal basiscopic pinnule to 12 x
3.6 cm bearing 1 pair of very deeply lobed tertiary
leaflets to 2.8 x 1.2 cm; basal acroscopic pinnule
to 6 cm long; second pair of pinnae bearing up to
5 pairs of free pinnules with basal lobes lobulate
to crenate, distal lobes entire; lower surface of
pinna-rachis bearing scales 2—3 mm long with
inflexed edges and widened clathrate basal part;
appressed glands variably present on and between
veins of pinnules; sori about medial on veins in
pinnule-lobes (sometimes near bases of both
branches of a forked vein); indusia thin, rather
large when young, crumpled when old, bearing
slender glands. — Figs. 6d; 8a, b.

Distribution — Malesia: S Sumatra, Java, Bali.

Habitat — In mountain forest.

Note — For discussion on Aspidium intermedium
Blume see Ctenitis vilis above. Blume sent to
Hooker one pinna of the type of the present species
with the name Aspidium intermedium; Hooker
(1862) merely copied Blume’s description and cited
specimens of other species also.

23. Ctenitis setosa (Presl) Holttum, Novit. Bot.
Delect. Seminum Horti Bot. Univ. Carol. Prag.
1968 (1969) 20, excl. Cuming 80 & 151; Blu-
mea 31 (1985) 28. — Lastrea setosa Presl, Epim.
Bot. (1851) 40. — [Polypodium hirtum Pres],
Reliq. Haenk. (1825) 27, non Sw. 1806.] —
Type: Haenke s.n., Luzon (PR; iso BM, K).

Basal scales of stipe not seen; /Jamina to 30 cm
long, firm but veins distinct on lower surface; free
pinnae 4 pairs and several pairs adnate, the costules
of basal lobes of distal pinnae arising from junction
of pinna-midrib with rachis; basal pinnae to 15 cm
long, bearing 2 pairs of free pinnules, distal parts
of pinnae shaped like the frond-apex; basal basi-
scopic pinnules of basal pinnae to 6.5 x 3 cm with
acute apex, bearing 1 pair of almost free tertiary
leaflets, the largest 18 x 10 mm lobed to its costa
near its base, the lobes entire (or the basal ones
with sinuous margin) with. broadly rounded apices;
basal acroscopic pinnules of basal pinnae 3.7 x 2
cm; pinnules or lobes of second, third and fourth
pairs of pinnae all deeply lobed and with broadly
obtuse apices; lower surface of main rachis and

34

Flora Malesiana ser. II, Vol. 2! (1991)

Fig. 8. Rachis scales (SEM). — a. Ctenitis alteroblumei Holttum, x c. 40; b. ibid., x c. 80. — c. C. sub-
glandulosa (Hance) Ching, x c. 40; d. ibid., x c. 140 (a, b: Sapiin 2793, Java; c, d: Elmer 22162, Luzon).

pinna-rachises bearing minute hairs and many very
narrow scales 2—3 mm long which are not stiffly
spreading, their margins inflexed and their widened
bases formed by isodiametric cells; lower surface
of costae of pinnules bearing sparse very short
hairs of 2 or 3 cells and apparently short slender
appressed red glands (not clearly seen); hairs on up-
per surface of costae 0.5 mm long, thick; no hairs
nor glands seen between veins on upper surface;
sori about medial on simple veinlets in pinnule-
lobes or near the bases of one or both branches of
forked veinlets; indusia very small, bearing a few
slender glands which may be red.

Distribution — Malesia: Philippines (apart from
the type, one specimen from North Luzon).

24. Ctenitis atrorubens Holttum, Blumea 31
(1985) 29. — Dryopteris intermedia (Blume)
Kuntze var. microloba Christ, Philipp. J. Sci.
2 (1907) Bot. 215. — Type: Copeland 1702,
Mindanao, San Ramon (MICH).

Ctenitis vilis auct.: Copel., Fern Fl. Philipp.
(1960) 288, p.p.

Stipe 20 cm long, dark red, its base covered by
tufted narrow red-brown scales 15 mm long, rest of
stipe bearing dark brown rigidly spreading scales;
rachis-scales like those of the stipe but gradually
shorter; lamina to 34 cm long, thin; pinnae imbri-
cate, c. 12 pairs free and several pairs adnate; basal
pinnae 11.5 cm long, short-stalked, their basal
basiscopic pinnules 5.2 x 1.6 cm bearing 1 pair of
deeply lobed tertiary leaflets, basal acroscopic pin-
nules 20 x 8 mm, their basal lobes lobulate; pin-
nules of middle pinnae all deeply lobed, the lobes
slightly oblique, their costules 3 mm apart; lower
surface of pinna-rachises bearing scales 2-3 mm
long, their margins inflexed and bases clathrate,
upper surface bearing hairs 1 mm long; sori supra-
medial on veins; indusia thin, covering young sori.

Distribution — Malesia: Philippines (known on-
ly from the type).

Habitat — In forest at 820 m.

Holttum — Tectaria Group

35

25. Ctenitis erythradenia Holttum, Blumea
31 (1985) 29. — Type: M.G. Price 1773, Lu-
zon, Mt Makiling (K).

Ctenitis setosa auct.: Holttum, Novit. Bot. Delect.
Seminum Horti Bot. Univ. Carol. Prag. 1968
(1969) 20, for Cuming 80 only.

Stipe to 37 cm long, pale when dry, basal scales
flat, light brown, 10 mm or more long, scales
above base very narrow, dark brown, to 8 mm
long with inflexed margins; lamina to 32 cm long,
thin; basal pinnae 15 cm long bearing 1 pair free
pinnules and 2 or 3 pairs adnate, basal basiscopic
pinnules 6.5 x 2 cm, bearing 1 pair of tertiary leaf-
lets 12 x 5 mm, lobed halfway to their costules;
basal acroscopic pinnule 2.5 x 1 cm, deeply lobed,
lobes subentire; lower surface of pinna-rachises bear-
ing very narrow scales 2—3 mm long with inflexed
margins and clathrate base; appressed red glands pres-
ent on and between veins on lower surface of pin-
nules; hairs on upper surfaces of pinna-rachises short;
sori medial; indusia small, thin, bearing red glands.

Distribution — Malesia: Philippines (Luzon,
Mindanao).

Habitat — In ridge forest at 500 m altitude.

Note — This species is allied to C. eatonii
(Baker) Ching of Taiwan, differing in rachis-scales,
indusia and red glands.

26. Ctenitis angusta Holttum, Blumea 31
(1985) 30. — Type: Brass 23208, Papua New
Guinea, Milne Bay Prov. (BM; iso A).

Stipe to 45 cm long, light castaneous, basal
scales thin, flat, narrow, 15 mm long, scales above
base spreading, dark brown, with inflexed margins,
gradually shorter upwards; lamina to 37 cm long,
20 cm wide, thin, with 9 or 10 pairs of free and 3
or 4 pairs of adnate pinnae; basal pinnae 11.5 cm
long, bearing 2 or 3 pairs of pinnules, the lowest
ones 4 x 1 cm, lobed almost to their costae with 1
almost free slightly crenate tertiary leaflet 7 x 3
. mm; pinnules of middle pinnae all deeply lobed;
lower surface of pinna-rachises bearing narrow
scales 2—3 mm long which have inflexed margins
and clathrate bases, also many short hairs, upper
surface covered with thick short hairs; sori a little
inframedial; indusia thin, small, reniform, bearing
pale red glands.

Distribution — Malesia: Papua New Guinea
(known only from the type).

Habitat — In moist valley in oak forest at 1550 m
altitude.

27. Ctenitis paleolata Copel., Philipp. J. Sci.
81 (1952) 24; Fern Fl. Philipp. (1960) 293;
Holttum, Blumea 31 (1985) 30. — Type: Cope-
land s.n. 9 May 1912, Luzon, Benguet Sub-
prov. (MICH).

Stipe nearly SO cm long, its basal scales 15 mm
long, narrow, in a close tuft, above base an abrupt
transition to narrow flat scales 2—3 mm long,
their cells all somewhat elongate but with dark
walls and clear lamina; lamina to at least 50 cm
long, texture thin, wholly catadromous; basal pin-
nae 20 cm long with stalks 3.5 cm and 6 pairs of
widely spaced pinnules; basal basiscopic pinnule 9
cm long, bearing 1 pair of free and several pairs of
adnate tertiary leaflets, all widely spaced; largest
tertiary leaflet 2 cm long, lobed almost to its cos-
ta, the lobes well spaced, very oblique and falcate,
the largest 4—5 x 1.5 mm; basal acroscopic pin-
nule 6.5 cm long; pinnules of suprabasal pinnae
all widely spaced with widely spaced narrow falcate
lobes; scales on lower surface of pinna-rachis nar-
row, evenly attenuate from base to apex, clathrate
but few basal cells isodiametric; sori small, medial
or supramedial; indusia small, distinct, bearing cy-
lindric glands; spores not seen.

Distribution — Malesia: Philippines (known on-
ly from the type).

Note — The scales do not indicate clearly whether
this species belongs to the same group as C. sub-
glandulosa; the form of spores would decide this
question.

28. Ctenitis subglandulosa (Hance) Ching,
Bull. Fan Mem. Inst. Biol. 8 (1938) 302; Ta-
gawa, Col. Illustr. Jap. Pterid. (1959) t. 223;
Copel., Fern Fl. Philipp. (1960) 292; DeVol
& Kuo, Fl. Taiwan 1 (1975) 327; Holttum,
Blumea 31 (1985) 35, f. 2c, d; 3c, d. — Also-
phila subglandulosa Hance, Ann. Sci. Nat. Bot.
V, 5 (1866) 253. — Dryopteris subglandulosa
(Hance) Hayata, Icon. Pl. Formos. 6 (1917) 101,
non (Fée) Kuntze 1891. — Type: Oldham s.n.,
March 1864, N Taiwan, ex Herb. Hance no
11186 (BM; iso K).

Polypodium oldhamii Baker, Syn. Fil. (1867) 311.
— Type: Oldham s.n., 1864, Taiwan, Tamsuy (K).

36

Flora Malesiana ser. II, Vol. 2! (1991)

Aspidium subtripinnatum Miq., Ann. Mus. Bot.
Lugd.-Bat. 3 (1867) 179. — Ctenitis subtripin-
nata (Miq.) H. Ito in Nakai & Honda, Nov. Fl.
Jap. 4 (1939) 74, excl. syn. — Type: ‘Siebold
et Buerger’, Japan [L 908,337-521 (Siebold),
515, 527, 537 (Buerger)].

Nephrodium rhodolepis Clarke, Trans. Linn. Soc.
London II, Bot. 1 (1880) 526, t. 72. — Lastrea
intermedia var. rhodolepis Bedd., Handb. Ferns
Brit. India Suppl. (1892) 62. — Dryopteris rho-
dolepis (Clarke) C.Chr., Index Filic. (1905)
288, excl. syn; Alderw., Malayan Ferns (1908)
202, excl. syn. — Lectotype (Holttum 1985):
Clarke 26934, Darjeeling, 6000 ft (K).

Ctenitis costulisora Ching, Bull. Fan Mem. Inst.
Biol. 8 (1938) 299, excl. W.T. Tsang 20974.
— Type: Y.K. Wang 31589, Kwangtung (not
seen).

Caudex massive; stipe to at least 120 cm long
(Hance), its base covered with light brown thin
scales 20-40 mm long, the scales gradually

shorter upwards for 5-10 cm, then a subabrupt
change to copious appressed overlapping clathrate
scales; lamina as long as stipe; basal pinnae of
lectotype of Nephrodium rhodolepis 45 cm long,
their largest pinnules bearing 4 or 5 pairs of ter-
tiary leaflets, the largest very deeply lobed; scales
on smaller axes of frond to 1 mm wide, consisting
almost entirely of isodiametric cells or at least of
cells not much longer than wide; indusia thin,
rather irregular in shape, bearing cylindric glands
and sometimes also hairs of several cells. — Figs.
6e, f; 8c, d.

Distribution — NE India, S China, Taiwan, Ryu-
kyu Is., S Japan, Guam; in Malesia: E Java, Bali,
Philippines (Luzon, Panay, Mindoro), E New
Guinea; Fiji.

Habitat — In Malesia on mountains in forest at
400-1400 m, in one case on a stream bank.

Taxonomy — Copeland (1960) distinguished C.
rhodolepis from C. subglandulosa in the Philip-
pines, but all specimens seen by me from the Phi-
lippines appear to belong to one species.

TECTARIDIUM

Tectaridium Copel., Philipp. J. Sci. 30 (1926) 329, pl. 1; Gen. Fil. (1947) 131; Fern Fl.
Philipp. (1960) 316. — Lectotype species (Copel., l.c. 1947): Tectaridium macleanii
Copel.

Caudex short, erect; scales narrow, entire; fronds dimorphous; sterile fronds simple,
elongate, entire, with venation as in Tectaria sect. Tectaria; fertile fronds lobed almost or
quite to the costa, the lobes well separated each including one main vein with a very nar-
row lamina on each side of it, the lamina widenend at short intervals, each widened part
bearing a sorus, terminal on a short vein; sori indusiate; spores not distinct from those of
Tectaria and Heterogonium; fronds intermediate in form between sterile and fully fertile
often present. — Fig. 9.

Distribution — A genus of one species, in Malesia native in the Philippines (southern
Luzon, Leyte, Samar, Panay).

Notes — In 1926 Copeland distinguished two species, which are here united.

Sterile fronds in this genus have a venation like that of Tectaria singaporeana, but the
extreme form of sterile fronds, as seen in the type of Tectaridium macleanii is so peculiar
that a generic distinction appears warranted. Pichi Sermolli [Webbia 31 (1977) 467] con-
sidered Tectaridium and Luerssenia to be closely allied, but the type of the latter (from
Sumatra) has sori not significantly different from those of Tectaria pleiosora and the type
species is here transferred to Tectaria, as T. kehdingiana. In my opinion Tectaridium re-
presents a quite separate offshoot from Tectaria. Fronds intermediate between sterile and
fully developed fertile offer significant evidence.

Holttum — Tectaria Group

37

1. Tectaridium macleanii Copel., Philipp.
J. Sci. 30 (1926) 329; ibid. 38 (1929) 138;
Fern Fl. Philipp. (1960) 316. — Type origin-
ally cited destroyed in 1945; neotype (so desig-
nated by Copeland in MICH): Maclean & Cata-
lan 84, Luzon, Cagayan Prov. (MICH).

Tectaridium primitivum Copel., Philipp. J. Sci. 30
(1926) 329; Fern Fl. Philipp. (1960) 316. —
Type: Wenzel 876, Leyte (isoF, MO, UC,
BKL).

Stipe dark castaneous, of sterile fronds 10—17
cm long, of fertile fronds to 36 cm, minutely hairy
near base; scales to 5 mm long, very narrow, hair-
pointed; /amina of sterile fronds commonly to 20
x 3.5 cm, largest seen 39 x 5.5 cm, opaque, entire,
apex short-pointed, base variously cordate to broad-
ly cuneate; veins not distinct on either surface when
dried, glabrous apart from dense short hairs on at
least the basal part of the upper surface of costa;
lobes of extreme form of fertile fronds to 4 cm
long, 8-10 mm apart on each side of the costa,
the base of each lobe widened to join a + contin-
uous very narrow wing on each side of the costa
(the wing occasionally widened to bear a single
sorus); sori borne on small + semicircular lobules
of the fertile lobes, opposite each other or alter-
nate, on very short simple veins, indusiate; indusia
firm, persistent, circular or a littlke asymmetric,
attached along a line from the base to the middle of
the sorus. — Fig. 9.

Distribution — Malesia: Philippines (Southern
Luzon, Leyte, Samar, Panay).

Habitat - Common forest floor fern in primary
forest at 450-500 m (Panay); primary forest on
limestone and limestone-derived soils (Samar);
lowland forest slope, heavy clay soil (Luzon).

Note — A collection in which fertile fronds have
a quite distinct wing joining the bases of successive
lobes was made type of a separate species T. primi-

PY. Epwarods

Fig. 9. Tectaridium macleanii Copel. a. Sterile and
fertile fronds, x 0.33; b. upper surface of a fertile
middle pinna, x 4 (a, b: Sands 3260).

tivum by Copeland. Deeply lobed sterile fronds,
lobes subentire, 4 mm wide, may occur and bear an
occasional sorus on an outer vein of a single row
of areoles. A plant from Panay cultivated at Kew
produced fronds showing a range of forms between
sterile and fertile. One frond 5 cm wide, shallowly
lobed throughout, produced irregularly arranged
sori near the margin, some terminal on short free
veins and some on veins surrounding areoles.

CHLAMYDOGRAMME

Chlamydogramme Holttum, Gard. Bull. Sing. 39 (1986) 157. — Type species: Chlamy-

dogramme hollrungii (Kuhn) Holttum.

Fronds simply pinnate; pinnae entire, basal ones bilobed or not; venation as in Tectaria
sect. Tectaria; pinnae of sterile fronds 2—3 cm or more wide, pinnae of fertile fronds 2—3
mm wide; sori linear, submarginal, protected when young by the thin pinna-margin and

by a continuous inner indusium. — Fig. 10.

38 Flora Malesiana ser. II, Vol. 2! (1991)

Distribution — Malesia: NE New Guinea, 2 known species.

Taxonomy — The type species was described as exindusiate; indusia were first report-
ed by Holttum in 1986. Copeland placed the species in the genus Hemigramma, in which
the sori run along the veins of variably contracted fronds or pinnae. Hemigramma is here
included in Tectaria, with the comment that, as arranged by Copeland, it was not a natural
group.

KEY, TO THE SPECIES

la. Terminal lamina pinna-like; pinnae 2 or 3 pairs, basal ones forked . 1. C. hollrungii
b. Terminal lamina 5-lobed; pinnae to 7 pairs, basal ones not forked .... 2. C. elata

1. Chlamydogramme hollrungii (Kuhn)
Holttum, Gard. Bull. Sing. 39 (1986) 157. —
Gymnopteris hollrungii Kuhn in Schum. &
Hollrung, Fl. Kais. Wilh. Land (1889) 8. —
Acrostichum hollrungii (Kuhn) Baker, Ann.
Bot. (London) 5 (1891) 495. — Leptochilus
hallrungii (Kuhn) C. Chr., Index Filic. (1905)
385; Alderw., Malayan Ferns (1908) 737. —
Hemigramma hollrungii (Kuhn) Copel., Phi-
lipp. J. Sci. 37 (1928) 406. — Type: Hollrung
640, NE New Guinea, Augustafluss (B; iso
BO, K, L).

Hemigramma grandifolia Copel., Philipp. J. Sci.
6 (1911) Bot. 77. — Type: C. King 328, NE
New Guinea, Lakekamu (MICH).

Caudex short, erect, covered with very narrow
entire scales c. 8 mm long. Sterile frond: stipe to
40 cm or more long, mid-castaneous, distally
short-hairy, scales above base caducous; lamina to
more than 30 cm long consisting of 2 or 3 pairs of
pinnae with conform terminal lamina, the upper
pinnae (if 3 pairs) adnate-decurrent, basal pinnae
bilobed; terminal lamina to 25 x 3.5 cm, acumi-
nate; basal pinnae to 25 x 3.5 cm with stalks 5-7
mm, their basiscopic lobes to 20 x 3 cm; sparse
short hairs present on lower surface of costae and
veins with a few between veins; more dense short
hairs present on upper surface of costae. Fertile
frond: stipe to 60 cm long; pinnae c. 2 mm wide. —
Fig. 10.

Distribution — Malesia: NE New Guinea, 3 col-
lections.

Habitat — On hill slopes in lowland forest.

Note — In one specimen there are rounded addi-
tional lobes 5—6 mm long on the acroscopic side
at the base of basal pinnae.

PJ.Eowanrps

Fig. 10. Chlamydogramme hollrungii (Kuhn) Holt-
tum. a. Sterile and fertile fronds, x 0.22; b. trans-
verse section of a fertile pinna, x 18 (a: Hoogland
& Craven 10516; b: Hoogland & Craven 10187).

Holttum — Tectaria Group

39

2. Chlamydogramme elata Holttum, Blumea
35 (1991) 556. — Type: J.R. Croft 1735, Papua
New Guinea, West Sepik Prov. (K; iso LAE).

Sterile frond: stipe to 60 cm or more long, dark
castaneous, sparsely short-hairy; largest basal scales
20 mm long, 3 mm wide at their bases, firm, dark
with thinner margins; lamina to 70 cm long; free
pinnae to 5 pairs and above them 2 or 3 pairs with
decurrent bases grading into a 3—5-lobed apex;
largest pinnae to 22 x 2.5 cm, their sides almost
parallel for much of their length, bases unequally
cuneate; basal pinnae simple, with stalks 3 mm
long, narrowed to a very narrow wing 10 mm long
on the basiscopic side, cuneate at c. 45° on acro-
scopic side; main veins sinuous, smaller veins dis-

tinct but not prominent; lower surface bearing sparse
short hairs on all parts, upper surface glabrous.
Fertile frond: lamina to more than 60 cm long;
pinnae c. 12 pairs, basal ones c. 15 x 0.3 cm, dis-
tal ones with soriferous decurrent bases; spores
similar to those of C. hollrungii.

Distribution — Malesia: New Guinea (known
from type collection only).

Habitat — In narrow gorge in limestone, 1-2 m
above flood level, 300 m.

Note — One sterile frond has only 2 pairs of
pinnae which are 4 cm wide, below a frond-apex
with lobes of similar width. A frond intermediate
between sterile and fertile has most pinnae 5—8
mm wide towards their bases, distally like pinnae
of a fully fertile frond.

TECTARIA

Tectaria Cav., An. Hist. Nat. 1 (1799) 115; Copel., Philipp. J. Sci. 2 (1907) Bot. 409—
418, excl. T. irregularis (Presl) Copel. & T. leuzeana (Gaudich.) Copel.; Ching,
Sinensia 2 (1931) 9—36, excl. T. artinexa (Clarke) Ching & T. leuzeana; Backer &
Posth., Warenfl. Java (1939) 71-78, excl. T. irregularis & T. leuzeana; Copel., Gen.
Fil. (1947) 128, excl. syn. Pleocnemia Presl, Dictyocline Presl, Proferea Presl &
Arcypteris Underw.; Manton & Sledge, Philos. Trans. Sér. B, 238 (1954) 137, 160;
Holttum, Revis. Fl. Malaya 2 (1955) 501-519, 626; Copel., Fern Fl. Philipp. (1960)
302-317, excl. T. irregularis & T. leuzeana. — Type species: Tectaria trifoliata (L.)

Cav.

Aspidium Sw., J. Bot. (Schrader) 1800, 2 (1801) 29, for the type species and few others;
J.Sm., Hist. Fil. (1875) 200—203; Diels in E. & P., Nat. Pflanzenfam. I, 4 (1899)
183-188; C. Chr., Index Filic. (1905) xxii; Alderw., Malayan Ferns (1908) 233-252.
— Aspidium § Euaspidium Hook., Sp. Fil. 4 (1862) 42. — Nephrodium § Sagenia
Hook. & Baker, Syn. Fil. (1867) 296. — Type species (J. Sm. 1875): Aspidium tri-

foliatum (L.) Sw.

Sagenia & Stenosemia Pres] 1836: see Tectaria sect. Sagenia.

Microbrochis Pres] 1851: see Tectaria sect. Sagenia.

Hemigramma Christ, Philipp. J. Sci. 2 (1907) Bot. 170; Copel., ibid. 37 (1928) 402,
excl. H. hollrungii (Kuhn) Copel.; Copel., Fern Fl. Philipp. (1960) 317; M.G. Price,
Kalikasan 3 (1974) 175; Holttum, Gard. Bull. Sing. 39 (1986) 154. — Type species:
Hemigramma zollingeri Christ = Tectaria hilocarpa (Fée) Price.

Quercifilix Copel., Philipp. J. Sci. 37 (1928) 408; Sledge, Kew Bull. 27 (1972) 422. —
Type species: Quercifilix zeilanica (Houtt.) Copel. = Tectaria zeilanica (Houtt.) Sledge.

Ctenitopsis Ching 1938: see Tectaria sect. Sagenia.

Caudex erect or + prostrate, its vascular structure radially arranged; scales narrow,
formed of narrow elongate cells, usually rigid and opaque, in some species with thinner

40 Flora Malesiana ser. II, Vol. 2! (1991)

or fringed margins, in a few small species wholly thin and translucent but the cells not
clathrate-isodiametric as in Ctenitis; fronds in most species at least pinnately divided, the
basal pinnae with elongate basal basiscopic lobes or pinnules, entire or with unlobed basal
pinnae only in a few species which have elaborate anastomosis of veins; veins free with
basal basiscopic one in each pinna-lobe arising from the costa (except in T. manilensis),
or anastomosing to form narrow areoles along costae or costules, or more elaborately
anastomosing with + isodiametric areoles having included free branched veinlets variously
directed; costae of pinnae and costules of pinnules in most species slightly ridged on the
upper surface and covered with ctenitoid hairs, in some species of sect. Tectaria slightly
grooved and glabrous; upper surface of lamina between veins usually bearing scattered
thick ctenitoid hairs at least near the margin, slender hairs variously present on and be-
tween veins on the lower surface; sori usually round with reniform or peltate indusia, in a
few species + elongate or asymmetric or spreading along all veins in contracted fertile pin-
nae; spores usually cristate, the crests forming a more or less definite reticulum, in some
species bearing slender spinules. — Figs. 11-16.

Distribution — Pantropic; c. 210 species, of which c. 40 are neotropic.

Habitat — Almost all species are terrestrial forest plants; a few grow only on rocks,
some species only on limestone; a few are rheophytes.

Cytology — Chromosome number 40.

Taxonomy — In describing the new genus Tectaria, Cavanilles cited only one species
which is therefore the type, but he later included in the same genus other species which
are now placed in Dryopteris, Polystichum, Nephrolepis, Thelypteris and other genera,
their common character being sori covered with reniform or peltate indusia. In 1801
Swartz established the genus Aspidium with a similar wide-ranging content, including the
original species cited by Cavanilles.

Later authors adopted Swartz’s name and treated it variously. Presl, Fée and others at-
tempted to segregate genera on the basis of characters of indusia (whether peltate or reni-
form) and of venation; they are listed by Christensen (1905). But these 19th century
authors did not observe indusia critically and failed to notice other characters now seen to
be important, especially of hairs, scales and glands, and most of their generic names are
no longer recognized, or if recognized are defined mainly by characters not mentioned by
their original authors. Hooker (1862) and Baker (1867), attempting to establish genera of
wider scope, confused together unrelated species. It was not until the classification of
Diels (1899) that it was clearly recognized that species belonging to the same genus could
have, or lack, indusia; he transferred some exindusiate species to Aspidium. But Diels in-
cluded also some extraneous elements and his subdivision of Aspidium is valueless. Chris-
tensen (1905) cited many generic synonyms but his main subdivision of Aspidium is little
better than that of Diels; he did however recognize that, of the synonyms cited, Tectaria
has priority and printed against it ‘nom. opt.’. Soon afterwards Copeland (1907), actively
studying Philippine species, revived the name Tectaria for them, but Van Alderwerelt
(1908) attempted to revert to the form of indusia to characterize infrageneric groups, mis-
using the name Sagenia Presl. Christensen accepted the generic name Tectaria in the third
Supplement to his Index (1934), Ching having already (1931) made the first good study
of the species of mainland Asia.

Holttum — Tectaria Group . 4]

Ching and Copeland accepted in Tectaria only species with anastomosing veins. Cope-
land (1907: 409) however wrote ‘Dryopteris dissecta (Forst.) Kuntze represents, most
nearly of known species, the probable origin of Tectaria.’ But at that time Christensen had
not yet discovered a natural subdivision of the comprehensive Dryopteris of his Index.
When he did so (in his monograph of 1913 and 1920), working entirely with tropical
American species, he distinguished Ctenitis from Dryopteris s.str. (and from thelypteroid
ferns) by the character of its hairs, and pointed out that in its hairs Tectaria agrees with
Ctenitis. Ching dealt with the species of Dryopteris and Ctenitis of mainland Asia in 1938
and in so doing came across a few species which have free veins but in other respects
resemble Tectaria. He proposed a new genus Ctenitopsis for these species (one was
Forster’s dissecta) and also species with free veins later transferred by Holttum to Hetero-
gonium; Copeland (1947) transferred all these to Crenitis and wrote (p. 124) “Tectaria is
probably of ctenitid origin’ and (p. 130) “Ctenitis is more primitive than Tectaria, there-
fore it is the probable parent genus.’ By the latter comment he meant that anastomosing
veins are a specialized character as compared with free veins and that therefore the primi-
tive species of Tectaria had free veins. But Copeland overlooked the fact that the type spe-
cies of Ctenitis, and most other American species included in that genus by Christensen,
differ from Tectaria in their scales and also in bearing cylindric glands which are lacking
in Tectaria; Manton (1954) also showed that they differ in chromosome number. Forster’s
species which was the subject of Copeland’s comment in 1907 differs only from acknowl-
edged species of Tectaria in its free veins, and the species T. fuscipes (Bedd.) C. Chr.
bridges the gap by having veins anastomosing to form costal areoles in its sterile fronds
but all free in fertile ones. The free-veined species here accepted in Tectaria are most abun-
dant in mainland Asia (NE India and China) and in the Philippines. There are a few others
in Malesia and the Pacific but no related species in tropical America or in Africa [T. braun-
iana (Karst.) C. Chr. of Colombia, which also has free veins, is certainly not closely re-
lated to the species of Asia and needs further study]. Thus SE Asia is here regarded as the
centre of origin of the genus. Tectaria ingens (Atk.) Holttum in NE India has fronds up to
3 m tall (including the stipe) and-is perhaps the nearest existing species to a prototype for
the genus.

In considering a subdivision of Tectaria, vein-pattern appears to be our only guide,
and the clearest division [Holttum, Kew Bull. 38 (1983) 108, f. 1] is between species
which have veins free or anastomosing in the pattern indicated for Sagenia by Presl, and
those which have copious anastomosis with branched veinlets in the areoles as in the type
species of Tectaria. There are, however, in Peninsular Thailand and Peninsular Malaysia,
especially on limestone, intermediates between the two. Apart from these species, a dis-
tinct line can be drawn between species which have narrow costal areoles lacking included
free veinlets and those in which costal areoles do contain free veinlets. The division is
here accepted, with sectional rank for the two divisions. As sect. Sagenia is clearly the
primitive group, it is dealt with first.

Description of species — A common feature of all species except a few which have
entire fronds or unlobed basal pinnae is that the basal pinnae are larger than those next
above them and have elongate basal basiscopic lobes or pinnules. In the development of a
young plant, each successive frond has basal pinnae gradually larger and more complex in

42 Flora Malesiana ser. II, Vol. 2! (1991)

structure, and in many species fronds of less than mature size and form may bear sori.
The shape and size of the basal pinnae of fronds of mature plants provide some of the
most distinctive characters for distinguishing between species. The result is that fronds of
immature plants are sometimes difficult to identify with certainty. The arrangement of sori
is independent of size of fronds, also (in some cases) pubescence, but scales, which in
Ctenitis provide good distinguishing characters, are not greatly varied in Tectaria. The
reader is therefore warned that fronds of immature size may be difficult to identify from
the keys. It is thus important for field botanists and collectors to choose fronds from ma-
ture plants for preservation; it is helpful also to include specimens of immature plants with
notes on their status, which few collectors have done. It is also important to collect com-
plete basal pinnae; the type specimen of T. setulosa (Baker) Holttum consists of a pair of
suprabasal pinnae and these have been misinterpreted as basal ones.

KEY TO THE SECTIONS OF TECTARIA

la. Veins free, or anastomosing to form narrow costal or costular areoles lacking free in-
cluded veinlets, such veinlets in other areoles few, short, unbranched and directed

foward Siathve smal Cain sateen niente cmd ane cre Sect. Sagenia (p. 42)
b. Veins freely anastomosing; branched free veinlets variously directed present in many
ARC OLSS parent sie ewan rena pet Fioees caer enney a espera DU Ga Sect. Tectaria (p. 59)

Section Sagenia

Tectaria sect. Sagenia Holttum, Gard. Bull. Sing. 39 (1986) 157. — Sagenia Presl, Tent.
Pterid. (1836) 86, t. II, f. 23-25 (not 22). — Nephrodium § Sagenia auct.: Hook. &
Baker, Syn. Fil. (1867) 296—300, for the smaller part. — Type species: Sagenia lati-
folia Presl = Tectaria mexicana (Fée) Morton.

Stenosemia Presl, Tent. Pterid. (1836) 237, t. X, f. 24; Fée, Mém. Foug. 5. Gen. Filic.
(1852) 53, pl. 3A; Alderw., Malayan Ferns (1908) 727, excl. S. pinnata Copel. & S.
teysmanniana (Baker) Diels; Copel., Gen Fil. (1947) 12; Fern Fl. Philipp. (1960)
301, excl. S. pinnata; S. Chandra, Kalikasan 12 (1983) 157. — Type species: Steno-
semia aurita (Sw.) Presl = Tectaria aurita (Sw.) S. Chandra.

Microbrochis Presl, Epim. Bot. (1851) 51. — Type species: Microbrochis apiifolia
(Schk.) Presl = Tectaria apiifolia (Schk.) Copeland.

Ctenitopsis Ching, Notul. Syst. (Paris) 7 (1938) 86, p.p. excl. spec. typ.

Ctenitis auct.: Copel., Gen. Fil. (1947) 123, p.p.; Fern Fl. Philipp. (1960) 285, p.p.

Distinctive characters as in the above key.

Distribution — Throughout the distribution of the genus; c. 65 species of which c. 10
are neotropic; in Malesia 26 species.

Taxonomy — The type species of Sagenia and Microbrochis are identical in venation
and both are West Indian. Presl distinguished them by the form of their indusia only, pel-
tate in Sagenia and reniform in Microbrochis; but in fact the indusia of the type species of
Sagenia are not truly peltate nor are they clearly distinct from those of the type of Micro-

Holttum — Tectaria Group 43

brochis. Tectaria coadunata (J. Sm.) C. Chr. in Asia is closely related to them both and
all three were regarded as conspecific by Hooker. There are allied species in Africa. Thus
the Sagenia form of venation is pantropic and must have had an early origin from the free-
veined form which, as above noted, is regarded as primitive for the genus Tectaria.

Presl’s name Sagenia was variously misused by later authors. Baker used it (1867:
296—300) as an infrageneric name in Nephrodium, a genus defined solely by the reni-
form indusia of sori and comprising mainly species now placed in the family Thelypter-
idaceae; thus Baker’s Nephrodium § Sagenia differs from Presl’s definition of Sagenia
both in indusia and venation, and is most unnaturally associated with a very different
group of ferns. In 1908 Van Alderwerelt revived the use of Sagenia as an infrageneric
group in Aspidiwm, again defining it in a sense contrary to that of Presl.

Stenosemia Pres| was established for one widely distributed species which has exactly
Sagenia venation in sterile fronds, but much-contracted fertile fronds with a simpler vena-
tion along which the sporangia are so crowded that they appear to be acrostichoid. There
is an allied species in Madagascar. Stenosemia pinnata Copeland has the frond-form of
Heterogonium and is here placed in that genus.

KEY TO THE SPECIES OF SECTION SAGENIA

tae eveinssince infertile fronds)... os. : aise sk sees ne tea ote ee tane tiaras 2
beet easticostaliarcoles present in fertile frondsici.)2)4 2025, fer in es i ee a 14

2a. Fronds very thin, appearing only in the rainy season; caudex creeping, succulent
1. T. manilensis

b. Fronds not very thin, persistent; caudex not succulent .................. 3
3a. All pinnae very deeply lobed; free tertiary leaflets present on basal pinnae ..... =
bayeinnae muchiless deeply lobed: tertiary leafletsirare) 2) 0. 10. 3 0 a a e. 5
4a. Bases of upper pinnae and pinnules narrowly decurrent, their basal lobes sometimes
Sepatatelygattached to rachis ime: tes 2a nese ea hee av ee ae ec ene re oa 5

b. Bases of upper pinnae not narrowly decurrent, their basal lobes never separately
Attached stomhe TACHIS 008 Ge. gas, 1A Nee tae eee se otegel Ate on at aE Aide eee 6

5a. Basal pinnae to 22 cm long; axes glabrous on lower surface ..... 2. T. mesodon
b. Basal pinnae to 45 cm long; axes short-hairy on lower surface ....... 3. T. laxa
6a. Hairs 1 mm long present between veins on upper surface ................ 7

b. Hairs on upper surface between veins few and short, mostly near sinuses
5. T. trichotoma
(ambasalapinnae to. 12 cralong. 4 s.c.y. 5s satus eee esau ea wee ae eee 8
b. Basal pinnae to 27 cm long; fronds not or little dimorphous ...... 4. T. ramosii
8a. Fronds strongly dimorphous; distinct indusia present
4a. T. ramosii x Psomiocarpa apiifolia

b. Fronds not dimorphous; indusia small or lacking ... 4b. T. ramosii x T. aurita
9a. Basal pinnae to at least 40 cm long; suprabasal pinnae bearing 3 or 4 pairs of sessile
CRAG AGC PUMNM ULE Sie cess ogc sate caylee) ches search ean ern a 6. T. setulosa

b. Basal pinnae not over 25 cm long; suprabasal pinnae bearing at least one pair of free
PUNUTUTL ES Reps Seah) Ac eee Sn eter i, EN lett Rays Ne Yat ls ae 10

44 Flora Malesiana ser. II, Vol. 2! (1991)
ase Sporangiabearinseshorucapitateshairs@i poe eee ee 7. T. moussetii
b. Sporangiavlackingsuehjhairsies © i See e eee eee oe Oe eee 11
lla. Fronds dimorphous; irregular slight anastomosis of veins in sterile fronds; scales on
Stiperblacksh1l Once min ee eatecn. ee eee os Mens asin en ae 8. T. fuscipes
b. Fronds not or little dimorphous; no anastomosis, scales otherwise ......... 12
12a. Pinnae very firm, their longest lobes, if sterile, sharply dentate; no hairs near
SiNnUSESKONSUpPPEMSUTLACe AAA eens Sea eee 9. T. rigida
b. Pinnae thinner, their largest lobes not sharply dentate; some hairs present on upper
Surfacesatdeastinear sinuses = Sacunes. Saas Hee See eee 13
13a. Basal pinnae 7 cm long on fronds with lamina 25 cm long; very narrow black scales
presenteonrachesie tins ee eis STORE PT tae 10. T. andersonii
b. Basal pinnae proportionately longer, with free pinnules even on smaller fronds;
Scaleszon rachissfewreand#noteblackwe wa eae eee 11. T. dissecta
14a. Pinnae of fertile fronds much contracted as compared with those of sterile fronds 15
baPinnaeoffertile:trondsinot onlittle;contracted —. == 2-405). 21 7)
15a. Sporangia covering the lobes of fertile pinnae .............. 12. T. aurita
ba Sporangia forming!separate Sori) Qa 2S eee 16
16a. Veins distinct on lower surface; rachis not dark and glossy ... 13. T. impressa
b. Veins not distinct on lower surface; rachis dark and glossy .. 15. T. bamleriana
iijaaaNviddlepinnae,entire or shallowly jlobediia4... nie. 14. T. barberi
b. Middle pinnae deeply lobed or pmnateh. 27) Ve Se eee 18
18a. Hairs between veins on upper surface present only near sinuses........... 19
b. Hairs scattered generally between veins on upper surface ............... 24
9a. Free pinnulesion basal pinnae/atimost Iipairss: 42 454)..4.- 2 ee eee 20
bs Free pinnules|onibasal pinnae/atileast 3/pairsi; 225 5 4 4. = 9-1 eee 22
20a. Fronds opaque; smaller veins not distinct on lower surface .. 15. T. bamleriana
b. Fronds not opaque, or smaller veins distinct on lower surface ............ Ak
21a. Pinna-rachis very dark; basal pinnae not over 30 cm long; New Guinea
16. T. nabirensis
b. Pinna-rachis not dark; basal pinnae to 45 cm long; W Malesia . 17. T. griffithii
22a. Rachis and pinna-rachis densely short-hairy with very narrow dark scales on lower
SUBEACEt.4 evap MAMIE UMRIR e auy a leart 17. T. griffithii var. amplissima
b. Rachis and pinna-rachis neither densely short-hairy nor scaly on lower surface 23
23a. Suprabasal pinnae almost sessile with sessile basal pinnules .. . 18. T. nesiotica
b. Suprabasal pinnae stalked with several stalked basal pinnules .... 19. T. kingii
2A a. SOMMANAUSIALEs 2. hevnce disks eARRRS GEE We es Bee RO eee 25
baySor/exindusiate: nieceienray Mess, Si SEE: REO) eG) TNE ny 29
25 aNo ree pinnules.onjbasalipinnac mane eee ee are 22. T. christii
bavAtleastal painoftree pinnuleson basalipimnaei 45) 4.455 eee 26
26a. Anastomosis confined (or nearly so) to costal and costular areoles ......... Ds)
b. Anastomosis comprising more than one series of small areoles below sinuses
between pinna-lobes) fen: Pac eek es Rs 28
27a. Free tertiary leaflets rarely present on basal pinnae........... 23. T. devexa
b. Free tertiary leaflets present on fronds of mature plants ........ 19. T. kingii

Holttum — Tectaria Group : ‘45

28a.

. Very short hairs abundant between veins on both surfaces . .

Hairs on lower surface of costae and costules to 0.3 mm long; veins not prominent

25. T. coadunata
Hairs on lower surface of costae and costules to 1 mm long; veins prominent on
botsnmSUrraces OL OMed specimens: <).10c)-7s ite ee iene ee 26. T. pubescens
21. T. adenophora
Hairs between veins on upper surface thick, to 0.5 mm or more Jong, none or few
OMMOW CTISUTACE | coh mea ae eN Ee V0.) eM aacaa lathe, 2s fi SRea ESI MS cape at tap aes tee AP Uae 30

. Basal pinnae of mature plants bearing several pairs of free or adnate pinnules, one

tertiary leaflet usually present

oe ee

SAE Napeeatey cues ee ae te eee 24. T. ferruginea

b. Basal pinnae of mature plants bearing one pair of pinnules; tertiary leaflets lacking

1. Tectaria manilensis (Presl) Holttum, In-
dian Fern J. 1 (1984) 36. — Lastrea manilensis
Presl, Epim. Bot. (1851) 39. — Ctenitis
manilensis (Presl) Holttum, Novit. Bot. Delect.
Seminum Horti Bot. Univ. Carol. Prag. 1968
(1969) 36. — Lastreopsis manilensis ( Presl)
Price, Kalikasan 12 (1983) 155. — Type: Meyen
s.n., Manila (PRC).

Nephrodium parishii Hook., Sp. Fil. 4 (1862) 131,
t. 260. — Lastreopsis parishii (Hook.) Ching,
Bull. Fan Mem. Inst. Biol. 8 (1938) 159;
Holttum, Revis. Fl. Malaya 2 (1955) 498, f.
294. — Type: Parish 91, Moulmein (K).

Athyrium loheri Christ, Bull. Herb. Boissier II, 6
(1906) 1001. — Ctenitis loheri (Christ)
Copel., Fern Fl. Philipp. (1960) 287. — Type:
Loher s.n., May 1905, Luzon, Montalban (P).

a. var. manilensis

Caudex short-creeping, succulent, bearing short
narrow thin scales; stipe to 18 cm long, its distal
part + densely hairy; /amina very thin, shrivelling
in the dry season, to 18 cm long and wide, deltoid-
pentagonal with 3 or 4 pairs of free or adnate pin-
nae; basal pinnae to 13 cm long, lobed to a very
narrow wing along the rachis; basal basiscopic
lobe to 8 cm long and 3 cm wide, lobed to a very
narrow wing, its larger lobes deeply lobulate; basal
acroscopic lobe of basal pinna 2.5 cm long; basal
veins in pinna-lobes rarely arising from the costa;
rachis and costae of pinnae bearing hairs 0.5—1
mm long on the upper surface, shorter rather thick
hairs also present on the lamina between veins,
hairs on lower surface fewer and more slender; sori
on short acroscopic branches of veins in pinnule-

20. T. squamipes

lobes; indusia reniform or asymmetric, sometimes
with short hairs. — Fig. 12a-—c.

Distribution — Southern Burma, Thailand; in
Malesia: Malaysia (Langkawi Is. and Perlis), Phi-
lippines (Luzon, Palawan), SW Celebes.

Habitat — Usually on limestone; in Luzon re-
ported on a clay bank, streamside. Fronds of plants
in cultivation in Singapore withered periodically
though not subjected to drought; new ones devel-
oped after a period of rest.

b. var. chupengensis (Ridley) Holttum, comb.
nov. — Lastrea.chupengensis Ridley, J. Malay.
Branch Roy. Asiat. Soc. 59 (1911) 232. —
Lastreopsis parishii var. chupengensis (Ridley)
Holttum, Revis. Fl. Malaya 2 (1955) 498. —
Type: Ridley 14769, Perlis, Chupeng (K).

Stipe to 7 cm, lamina to 10 x 9 cm with only
the basal pinnae free.

Note — Known only from the type locality
whence plants cultivated in Singapore alongside
plants of the typical variety maintained their dis-
tinctive characters over three years. C.G. Matthew
reported that plants grew in a dense mat at the en-
trance to a limestone cave.

2. Tectaria mesodon (Copel.) Price, Kalikasan
1 (1972) 37. — Dryopteris mesodon Copel.,
Philipp. J. Sci. 7 (1912) Bot. 54. — Ctenitis
mesodon (Copel.) Copel., Gen. Fil. (1947) 124;
Fern Fl. Philipp. (1960) 294. — Type: Weber
1176, Mindanao (MICH; iso K, UC).

Caudex short, suberect, bearing firm dark scales
12 x 1 mm; stipe c. 30 cm long, dark, glossy, short-

46

Flora Malesiana ser. II, Vol. 2! (1991)

hairy in the groove only; /amina 30 cm long, very
firm, drying dark, free pinnae 3 or 4 pairs and 2 or
3 pairs adnate; apical lamina very obliquely lobed
almost to its axis, the lobes with narrowly decur-
rent bases, upper pinnae similarly lobed, their basal
lobes sometimes separately attached to the rachis,
all their lobes lobulate, the lobules falcate; basal
pinnae to 25 cm long, bearing 2 pairs of stalked
pinnules and an apical lamina like that of the frond,
basal basiscopic pinnule to 14 cm long with 1 pair
of free tertiary leaflets; largest tertiary leaflet 4 cm
long, very deeply lobed, the lobes crenate, basal
acroscopic pinnule 8 cm long; short hairs present
on upper surface of rachises and costae, also near
sinuses between lobes of pinnae, lacking elsewhere;
veins concolorous at both sides, more prominent
on upper surface than on lower; sori at the ends of
acroscopic branches of forked veins near margin;
indusia small, dark, firm, glabrous.

Distribution — Malesia: Philippines (Mindanao).

3. Tectaria laxa (Copel.) Price, Kalikasan 1
(1972) 37. — Ctenitis laxa Copel., Philipp. J.
Sci. 83 (1954) 98. — Type: Ramos BS 7757,
Luzon, Ilocos Norte (MICH).

Closely allied to T. mesodon and similar in tex-
ture and frond-form, differing in its much larger
fronds with all parts more widely spaced; stipe to
75 cm long, lamina to 60 cm or more; basal pin-
nae to 45 cm long (including stalks 6 cm long),
their basal basiscopic pinnules 24 cm long and
largest tertiary leaflets 8 cm long, basal acroscopic
pinnules 16 cm long; veins distinct on both sur-
faces; very short hairs scattered on all minor axes
on their lower surface, also some between veins. —
Fig. 12d.

Distribution — Malesia: Philippines (Northern
Luzon, several collections).

Habitat — In dry forest at 500-1000 m.

4, Tectaria ramosii (Copel.) Holttum, comb.
nov. — Ctenitis ramosii Copel., Philipp. J. Sci.
81 (1952) 24; Fern Fl. Philipp. (1960) 295. —
Type: Ramos BS 17511, Samar (MICH; iso P).

Caudex short, suberect; stipe to 65 cm long, dark,
glossy, bearing rather thin dark scales 10 x 1 mm
near the base only; /amina to 55 cm long with 5
pairs of free and 2 pairs of adnate pinnae below a

short apical lamina; basal pinnae to 27 cm long
with stalks 2 cm long, bearing 3 pairs of well-
spaced free and 1 or 2 pairs adnate pinnules; basal
basiscopic pinnule 13 x 5.5 cm bearing 2 pairs
free and 1 or 2 pairs adnate tertiary leaflets, its api-
cal lamina very deeply lobed, the lobes obliquely
lobulate; largest tertiary leaflet 2.5 x 1.6 cm, very
deeply lobed; basal acroscopic pinnule 5 x 2.3 cm;
basal basiscopic lobes of distal pinnules on all
pinnae shortly decurrent, their veins arising from
the pinna-rachis; veins all free; hairs on upper sur-
face of costae dense and thick, scattered ones more
than 1 mm long, similar long hairs also present
on and between veins; hairs on lower surface sparse
and shorter; sori supramedial at ends of veins in
lobules of pinna-lobes; indusia thin, persistent,
more than 1 mm diameter when young.

Distribution — Malesia: Philippines (Samar and
Bohol).

Habitat — On limestone-derived soils.

Probable hybrids:

4a. Tectaria ramosii x Psomiocarpa apii-
folia Presl. — Collection: Western Samar,
Price & Hernaez 398, also a sterile frond of
Price & Hernaez 74 (K); habitat as T. ramosii.

Fronds strongly dimorphous; sterile fronds sim-
ilar to those of Psomiocarpa but with larger seg-
ments, lamina to 16 x 16 cm; fertile fronds more
elongate (lamina 18 cm long, basal pinnae 7 cm),
their pinnae and pinnules much contracted; sori in-
dusiate, indusia firm, reniform.

4b. Tectaria ramosii x Tectaria aurita
(Sw.) Chandra. — Collection: Western Samar,
Price & Hernaez 20 & 74 (K).

Fronds very dimorphous; sterile fronds similar
in form to those of T. aurita but with slight and
irregular anastomosis of veins and lack of buds at
bases of pinnae; fertile fronds less contracted than
those of T. aurita with distinct separate small son;
indusia very small or lacking.

5. Tectaria trichotoma (Fée) Tagawa, Acta
Phytotax. Geobot. 25 (1973) 180. — Aspidium
trichotomum Fée, Mém. Foug. 5. Gen. Filic.
(1852) 295. — Ctenitopsis trichotoma (Fée)
C. Chr., Notul. Syst. (Paris) 7 (1938) 88; Tard.

Holttum — Tectaria Group

& C.Chr., Fl. Indo-Chine 7 (1941) 351. —
Ctenitis trichotoma (Fée) Copel., Gen. Fil.
(1947) 125. — Type: Gaudichaud 24, Vietnam,
Tourane (P; iso K).

Dryopteris balabacensis Christ, Philipp. J. Sci. 2
(1907) Bot. 213. — Ctenitis balabacensis (Christ)
Copel., Gen. Fil. (1947) 124; Fern Fl. Philipp.
(1960) 294. — Tectaria balabacensis (Christ)
Price, Kalikasan 1 (1972) 37; Contr. Univ.
Michigan Herb. 16 (1987) 199. — Type: Man-
gubat BS 392, Balabac (P; iso GH, K, NY).

Dryopteris escritorii Alderw., Bull. Jard. Bot. Bui-
tenzorg II, 23 (1916) 10. — Type: Escritor BS
20732, Luzon, Tayabas Prov. (BO; iso F, L, P).

Caudex short, suberect; stipe 30 cm long, cas-
taneous, hairy in the groove only, basal scales 12
x 1 mm, firm, dark; /amina to 50 cm long, firm;
free and adnate pinnae to 6 pairs; apical lamina 13
cm long, acuminate, lobed almost to its axis, its
basal lobes 6 x 1.3 cm, upcurved, deeply crenate;
upper pinnae similar with crenate lobes; basal pin-
nae to 25 cm long (stalk to 5 cm), oblique-ascend-
ing, bearing several pairs of pinnules; basal basi-
scopic pinnule 12.5 x 5.5 cm with 1 or 2 pairs of
free tertiary leaflets; basal acroscopic pinnule 7 x
3.5 cm; veins in pinna-lobes mostly forked, con-
colorous; lower surface glabrous, upper surface
densely short-hairy on rachises, short hairs also
present near sinuses between pinna-lobes. Sori
about medial between costules of pinna-lobes and
margin, terminal on acroscopic branch veinlets; in-
dusia dark, very firm, to 1.5 mm wide, with irreg-
ular small papillae on the upper surface.

Distribution — Vietnam; Malesia: Philippines
(S Luzon, Panay, Sibuyan, Palawan, Balabac), Bor-
neo (Sabah).

Habitat — One specimen from Tonkin was found
on limestone; one from Sabah on ‘ultrabasic steep
cliffs and banks.’

Note — The largest specimens from Vietnam
are considerably larger than any from Malesia (one
has a basal pinna 55 cm long) but otherwise show
no difference.

6. Tectaria setulosa (Baker) Holttum, comb.
nov. — Nephrodium setulosum Baker, J. Bot.
28 (1890) 265. — Ctenitopsis setulosa (Baker)
C. Chr., Notul Syst. (Paris) 7 (1938) 87; Tard.
& C.Chr., Fl. Indo-Chine 7 (1941) 352. —
Type: Balansa 1836, Tonkin (P; iso K).

Tectaria ingens auct.: Holttum, Revis. Fl. Malaya
2 (1955) 503.

KEY TO THE VARIETIES

la. Larger pinnule-lobes obtuse, fertile ones cre-
nate; sori mostly in one row on each side of
costules; no anastomosis . a. var. setulosa
b. Larger pinnule-lobes acute, fertile ones deeply
lobulate; sori 2 or 3 pairs in each lobule; some
anastomosis of veins . b. var. raciborskii

a. var. setulosa

Caudex short, erect; stipe to 55 cm long, short-
hairy throughout, basal scales 12 x 1 mm, scatter-
ed narrower ones also above base; lamina to 70 cm
long, rather thin; basal pinnae to 40 cm long, their
basal basiscopic pinnules to 16 x 6 cm deeply lobed
with crenate lobes, the basal lobes forming free
tertiary leaflets; suprabasal pinnae bearing 3 or 4
pairs of free or adnate pinnules; distal adnate pin-
nae lobed to 3—4 mm from the costa, their basal
basiscopic lobes decurrent on the rachis; pinna-
rachises and costae short-hairy throughout on
lower surface with sparse shorter hairs on veins,
densely hairy on upper surface, the hairs more than
0.5 mm long, scattered similar hairs present be-
tween veins; lateral veins in lobes of pinnae slen-
der and prominent on the lower surface, forked or
subpinnate; fertile pinnules as large as sterile ones,
their lobes obtuse, crenate; sori about medial in
the lobes, at the ends of acroscopic vein-branches,
or a few additional ones also on basal lobes; indu-
sia large, firm, glabrous.

Distribution — Tonkin, N Burma; in Malesia
only known from the Malay Peninsula (Cameron
Highlands, Pahang).

Habitat — In wet ground under tall Musa plants,
at 1850 m.

Note — The type consists of a pair of supra-
basal pinnae; these are wrongly described as basal
pinnae in the FI. Indo-Chine.

b. var. raciborskii (Alderw.) Holttum, stat. nov.
— Dryopteris raciborskii Alderw., Malayan
Ferns (1908) 197, 816, new name for Aspidi-
um sagenioides auct.: Racib., Pteridop. Buiten-
zorg (1898) 179, non Mett. — Type: Raciborski
s.n., W Java, G. Salak (BO; K, L).

48

Flora Malesiana ser. II, Vol. 2! (1991)

Dryopteris dissecta p.p. auct.: Backer & Posth.,
Varenfl. Java (1939) 95.

Fronds somewhat larger than those of var. setu-
losa, larger lobes of pinnae and pinnules acute,
sterile pinnules broader and less deeply lobed than
fertile; lobes of fertile pinnules deeply lobed with
3 pairs of sori in each lobule; irregular slight ana-
stomosis of veins in sterile pinnules especially
near bases of lobes.

Distribution — Known only from the original
collection.

7. Tectaria moussetii Holttum, Blumea 35
(1991) 553. — Type: Mousset 130, E Java, La-
wang, near Poespo, 700-800 m (K; iso L, UC).

Caudex not seen; stipe to 50 cm long, dark pur-
plish-brown, glabrescent, rachis similar, both bear-
ing narrow dark glossy scales; basal scales to 10 x
1.5 mm; lamina to at least 50 cm long; pinnae 7
pairs or more, well spaced; basal pinnae of type
specimen 16 cm long (stalk 10 mm) with 1 pair of
separately adnate pinnules, basal basiscopic pinnule
7 cm long, deeply lobed, its lobes 7 mm apart;
middle pinnae deeply lobed at their bases, lobes
crenate, costules 10-12 mm apart; lateral veins
in lobes mostly forked; lower surface of pinna-
rachis short-hairy near its base, short scales present
throughout; upper surface of pinnae bearing thick
hairs between the veins; sori terminal on acroscopic
vein-branches; indusia thin, often with short slen-
der hairs; sporangia bearing short capitate hairs
near the annulus.

Distribution — Malesia: East Java, Bali (3 col-
lections).

Note — Zollinger F146 from Bali is larger than
the type, having lamina 60 cm long and basal pin-
nae 23 cm long with 2 pairs of free pinnules. The
capitate hairs on sporangia are unique in the genus.

8. Tectaria fuscipes (Bedd.) C.Chr., Contr.
U.S. Natl. Herb. 26 (1931) 290; Ching, Sinen-
sia 2 (1931) 14. — Aspidium fuscipes Bedd.,
Ferns Brit. India Suppl. (1876) 15, t. 366. —
Ctenitopsis fuscipes (Bedd.) C. Chr., Notul.
Syst. (Paris) 7 (1938) 87; Ching, Bull. Fan
Mem. Inst. Biol. 8 (1938) 313. — Type: Clarke
7050, Cachar, Assam (K).

Nephrodium membranifolium Hook., Sp. Fil.
(1862) 131, p.p., incl. t. 261.

Caudex short, erect; fronds dimorphous, the fer-
tile ones with contracted pinnae; stipes commonly
30 cm long, light castaneous, bearing many nar-
row black scales; lamina to 30 cm long; pinnae 3
or 4 pairs, basal pinnae to 14 cm long, basal basi-
scopic pinnules to 6 cm long; middle pinnae of
sterile frond lobed about 3/5 towards their costae,
the lobes subentire and sinuses between them nar-
row; middle pinnae of fertile fronds very deeply
lobed, the lobes crenate and well-spaced; veins usu-
ally forked in lobes of sterile pinnae, the basal
ones uregularly anastomosing (rarely in Malesian
specimens), often simple in lobes of fertile pinnae;
thick hairs present between veins on upper surface,
most abundant on sterile pinnae, slender ones less
abundant on lower surface; sori on acroscopic
branches of veins; indusia variably hairy.

Distribution — Widely in Southeast Asia; in Ma-
lesia: E Java and SW Celebes.

9. Tectaria rigida Holttum, Blumea 35 (1991)
555. — Type: W. Meijer 7491, Sumatra, G.
Sago (BO; iso SING).

Caudex short, erect; stipe to 35 cm long, slight-
ly castaneous, its basal scales 10 mm long, barely
1 mm wide, firm with filiform tips; lamina c. 30
cm long; pinnae 3 pairs stalked, 2 pairs sessile or
adnate; basal pinnae 15 cm long with 2 pairs of
stalked pinnules, basal basiscopic ones 7 cm long
with basal lobe almost free; lobes of all pinnae
falcate, acute, the margins of sterile ones acutely
dentate; veins as in T. dissecta; very short hairs
present on upper surface of rachis and near bases of
pinna-rachises, none between veins on the upper
surface; sori terminal on acroscopic branches of
veins in pinna-lobes; indusia firm, dark, glabrous.

Distribution — Malesia: Sumatra (G. Sago;
known only from the type collection).

Habitat — Ravine in forest, 900-1000 m.

10. Tectaria andersonii Holttum, Blumea 35
(1991) 547. — Type: J.A.R. Anderson S 31936,
Sarawak, G. Subis, Niah, Miri Dist. (K).

Caudex short, erect; stipe to 22 cm long, dark-
castaneous, bearing short stiff erect hairs and scales
throughout; basal scales to 12 mm long, 0.5 mm
wide at the base, apex filiform, very dark, scales
on stipe and rachis 2-3 mm long, very narrow;
lamina to 25 cm long, firm, drying dark brown on

Holttum — Tectaria Group

lower surface; pinnae to 5 pairs free and 3 pairs
adnate, horizontal, hardly upcurved distally; basal
pinnae 7 cm long with 1 pair of free pinnules,
basal basiscopic one 2.2 x 0.9 cm, acroscopic 1.5
cm long; other pinnae lobed almost to their costae
near the base, costules to 8 mm apart, lobes obtuse
and slightly crenate; lower surface of costae bear-
ing erect hairs 0.2—0.4 mm long, fewer and short-
er hairs on veins, few between veins; upper surface
of costae bearing dense thick hairs 0.4 mm long,
between veins hairs only present near the margin;
sori on acroscopic branches of veins, nearer to
margin than to costule; indusia reniform, firm,
with a few short hairs.

Distribution — Malesia: Borneo: Sarawak (Miri
Dist.; known only from the type collection).

Habitat — Upper slopes of a limestone hill,
limestone rocks outcropping and deep ‘mor’ soil,
in shaded place.

11. Tectaria dissecta (G. Forster) Lellinger,
Amer. Fern J. 58 (1968) 156; Brownlie, Pterid.
Fl. Fiji (1977) 288. — Polypodium dissectum
G. Forster, Fl. Ins. Austr. (1786) 81. — Lastrea
dissecta (G. Forster) Carr. in Seem., Fl. Vit.
(1873) 360. — Dryopteris dissecta (G. Forster)
Kuntze, Revis. Gen. Pl. (1891) 812; Alderw.,
Malayan Ferns (1908) 196; Backer & Posth.,
Varenfl. Java (1939) 45, excl. syn. D. raciborskii
Alderw.; C. Chr., Bernice P. Bishop Mus. Bull.
177 (1943) 95, p.p. — Ctenitopsis dissecta
(G. Forster) Ching, Bull. Fan Mem. Inst. Biol.
8 (1938) 321; DeVol & Kuo, Fl. Taiwan 1
(1975) 330. — Ctenitis dissecta (G. Forster)
H. Ito in Nakai & Honda, Nov. Fl. Jap. 4 (1939)
91, for the name only; Copel., Fern Fl. Philipp.
(1960) 286. — Type: G. Forster s.n., Pacific
Islands (BM; iso K).

Nephrodium membranifolium Presl, Reliq. Haenk.
(1825) 36, t. V, f. 3; Hook., Sp. Fil. 4 (1862)
131, p.p., excl. t. 261; Holttum, Novit. Bot.
Delect. Seminum Horti Bot. Univ. Carol. Prag.
1968 (1969) 17. — Aspidium membranifolium
(Presl) Kunze, Bot. Zeitung (Berlin) 6 (1848)
361; C.Chr., Index Filic. (1905) 82, p.p. —
Ctenitopsis membranifolia (Presl) Ching, Bull.
Fan Mem. Inst. Biol. 8 (1938) 315. — Type:
Haenke s.n., Luzon (PRC).

Caudex short, erect; stipe to 50 cm long or more,
rather dark, minutely hairy; basal scales 10 x 1.2

49

mm, dark, firm, those above base with pale thin-
ner margins, distal scales and those on rachis thin,
mostly caducous; /amina to 60 cm long, always
much longer than wide; pinnae to 10 or more pairs,
oblique and + upcurved, apical lamina lobed al-
most to the rachis; basal pinnae to 20 cm long
(stalk to 2 cm) bearing up to 3 pairs of pinnules
(at least one pair in small plants); basal basiscopic
pinnule to 12 cm long, very deeply lobed but rare-
ly with a free tertiary leaflet; middle pinnae sessile,
short-acuminate, very deeply lobed, the lowest
lobes at least crenate, lateral veins in the lobes
once or twice forked, the basal basiscopic one al-
ways arising from the costa; lower surface of costa
bearing + abundant short hairs, hairs between veins
usually lacking or few; upper surface of costae bear-
ing dense thicker hairs, between veins some thick
hairs (variable); sori on the acroscopic branches of
veins, sometimes terminal, usually nearer to the
margin than to the costule; indusia well developed,
dark when dried, glabrous or variously short-hairy
when hairs are present on the lower surface. — Fig.
lla,e,g.

Distribution — Tahiti, Fiji, Tonga, New He-
brides; Taiwan; in Malesia: Java, Lesser Sunda
Islands, Christmas Is., Borneo, Philippines, Cele-
bes, New Guinea.

Habitat — This species appears to be adapted to
a climate with a regular dry season. The largest
plants are reported as terrestrial in forest. In Borneo
all plants for which the habitat is recorded were
found on limestone or (one from Mt Kinabalu at
1000 m) pendulous from rocks near a stream. In
central Java one plant was found on an old wall.

Variability — The largest plants seen are from
Luzon, the Lesser Sunda Islands and Christmas Is-
land. Most plants from New Guinea have abundant
erect very slender hairs between veins on the lower
surface and also copious hairs on the upper surface,
the latter thinner than those on plants which have
a few thick hairs; some Philippine plants are
similar but those on limestone have few hairs on
the lower surface. Limestone plants in Sarawak are
rather small but typical in pubescence. A few spec-
imens from limestone in the Philippines are very
small, possibly the result of exposure.

12. Tectaria aurita (Sw.) S. Chandra, Kalika-
san 12 (1983) 157. — Acrostichum auritum Sw.,
J. Bot. (Schrader) 1800, 2 (1801) 12; Syn. Fil.

50 Flora Malesiana ser. II, Vol. 2! (1991)

fi Hdl

Fig. 11. Tectaria, types of venation; scales. — T. dissecta (G. Forster) Lellinger. a. Fertile lobes of a mid-
dle pinna with sori removed from the right, x 2.5. — T. devexa (Kunze ex Mett.) Copel. b. Fertile lobes
of a middle pinna with sori removed from the right, x 2.5. — T. melanocaula (Blume) Copel. c. Base of a
middle pinna with sori removed on the right, x 2.5. — T. singaporeana (Hook. & Grev.) Copel. d. Vein
pattern between two lateral veins, x 2.5. — T. dissecta. e. A rhizome scale with margin and tip details,
x 100. — T. devexa. f. Hairs from laminal veins, x 150. — T. dissecta. g. Hairs from rachis and midrib,
x 150 (a, e, g: Price 1763; b, f: Anderson s.n., Sarawak; c: Mousset s.n., Java; d: Parris 10559).

Holttum — Tectaria Group

51

(1806) 13, 198; Hook., Sp. Fil. 5 (1864) 257:
Racib., Pteridop. Buitenzorg (1898) 52. — Poly-
botrya aurita (Sw.) Blume, Fl. Javae Filic.
(1828) 15, t. 1. — Stenosemia aurita (Sw.)
Presl, Tent. Pterid. (1836) 237, t. X, f. 24; Fée,
Mém. Foug. 5. Gen. Filic. (1852) 55, t. WIA;
Alderw., Malayan Ferns (1908) 726; Backer &
Posth., Varenfl. Java (1939) 79; Copel., Fern
Fl. Philipp. (1960) 301; Walker, J. Linn. Soc.
Bot. Suppl. I (1973) 109. — Type Thunberg
s.n., Java (S).

Polybotrya orientalis Blume, Enum. Pl. Javae
(1828) 99. — Type: Blume s.n., Noesa Kam-
bangan (L).

Polybotrya cicutaria Blume, Enum. PI. Javae (1828)
99; Fl. Javae Filic. (1828) 17, t. 2. — Steno-
semia cicutaria (Blume) Presl, Tent. Pterid.
(1836) 237. — Type: Blume s.n., W Java, on
rocky streambank (L 908,24-874).

Caudex short, erect; stipe of sterile frond to 30
cm long, of fertile to at least 45 cm; basal scales
10 mm long, narrow; sterile frond tripartite, broad-
ly deltoid-pentagonal in outline; terminal lamina
deltoid, deeply multi-lobed, the lobes all entire,
acute to acuminate, the lower ones slightly reduced
or in the largest fronds elongate and deeply lobed;
pinnae opposite or nearly so, to 16 cm long, short-
stalked, deeply lobed, the basal basiscopic lobe to
9 cm long, asymmetric, wider and more deeply
lobed on its basiscopic side, a bud present at the
base of each pinna; veins forming areoles along
costae of lobes of apical lamina and of pinnae,
sometimes with additional areoles below sinuses
between lobes; lower surface of costae variously
hairy, hairs on some Sumatran specimens more
than 1 mm long, usually shorter or lacking; upper
surface always densely short-hairy on costae, be-
tween veins scattered thick hairs variously abun-
dant; fertile frond shaped as sterile but all parts nar-
row; venation of very narrow areoles; sporangia
abundant all along veins; spores with very thin
translucent wing; n = 40 (Walker). — Fig. 12e.

Distribution — Throughout Malesia except for
the Malay Peninsula; Solomon Islands, New Heb-
rides.

Habitat — In most cases reported as growing on
rocky streambanks in forest, in some cases on
limestone, also reported terrestrial in forest.

Variability - Some specimens are very small;
one such was named var. minor by Christ [Verh.

Naturf. Ges. Basel 11 (1895) 252, type from lime-
stone in Celebes] but it is not clear whether the
dwarf form is constant; experiment is needed. Poly-
botrya cicutaria Blume was described from a sterile
frond which has very narrow lobes.

Probable hybrids: see 4. Tectaria ramosit.

13. Tectaria impressa (Fée) Holttum, comb.
nov. — Phlebiogonium impressum Fée, Mém.
Foug. 5. Gen. Filic. (1852) 314, t. 24A, f. 2.
— Type: Griffith s.n., Ind. Or. (RB).

Aspidium variolosum Wall. ex Hook., Sp. Fil. 4
(1862) 51; Bedd., Ferns Brit. India Suppl.
(1876) t. 365; Handb. Ferns Brit. India (1883)
216. — Tectaria variolosa (Hook.) C.Chr.,
Contr. U.S. Natl. Herb. 26 (1931) 289; Ching,
Sinensia 2 (1931) 21, pl. I; Holttum, Revis. Fl.
Malaya 2 (1955) 506, p.p.; Gard. Bull. Sing.
38 (1986) 146, f. 1A. — Lectotype (Ching
1931): Wallich 379, p.p. (K, ex Herb. Hook.).

Aspidium immersum Hook., Sp. Fil. 4 (1862) 58,
non Blume 1828. — Type: Wallich s.n. (K).

Aspidium prominens Alderw., Bull. Jard. Bot. Bui-
tenzorg II, 16 (1914) 56; Malayan Ferns, Suppl.
(1917) 198. — Tectaria prominens (Alderw.)
C. Chr., Index Filic. Suppl. 3 (1934) 183. —
Type: Matthew 706, Sumatra (BO).

Aspidium zollingerianum auct.: Bedd., Ferns Brit.
India (1867) t. 251 (non Kunze 1846).

Caudex short-creeping or suberect, bearing di-
morphous fronds with occasional intermediates;
stipe Castaneous, longest on fertile fronds, its basal
scales to 8 mm long, firm with fragile margins;
sterile lamina to c. 30 cm long with 3 or 4 pairs
of pinnae and a deeply lobed terminal lamina; basal
pinnae to 20 cm long, usually with 1 pair (rarely 2
or 3) of pinnules, the basal basiscopic one to 12 x
3 cm, lobed 1/3 towards its costa; middle pinnae
lobed to at least halfway to their costae, the lobes
oblique, obtuse; veins forming narrow areoles
along costae of pinnae and costules of lobes with
additional oblique areoles a few of which have an
included free veinlet; short hairs present on both
surfaces of costae and costules, also some on upper
surface near margins; pinnae of fertile fronds +
contracted and more deeply lobed, the venation
+ modified with absence of regular costular areoles;
sori mostly at the ends of free veins in areoles,
variably impressed (most deeply where the lamina
is most constricted), indusia large, firm.

52

Flora Malesiana ser. II, Vol. 2! (1991)

Distribution — NE India & S China southwards
to Peninsular Thailand; in Malesia known only
from 2 collections, one from Langkawi Is. and one
from Sumatra (type of Aspidium prominens, prob-
ably Pulau Weh).

Habitat — Usually on limestone in forest.

Note — Wallich cited 3 localities for his no 379:
Amherst, Tavoy and Penang. The Penang specimen
in his herbarium at Kew is T. brachiata (Z. & M.)
Morton. Ching’s lectotype is unlocalized. A speci-
men of Wallich 379 from Tavoy in Herb. Hooker
is a syntype of T. barberi (Hook.) Copel. Matthew
collected other specimens on limestone on Pulau
Weh in January 1912.

14. Tectaria barberi (Hook.) Copel., Philipp.
J. Sci. 2 (1907) Bot. 414; Holttum, Revis. FI.
Malaya 2 (1955) 508, 626, f. 300. — Polypo-
dium barberi Hook., Sp. Fil. 5 (1864) 100. —
Dictyopteris barberi (Hook.) Bedd., Ferns Brit.
India (1869) t. 322; Handb. Ferns Brit. India
(1883) 298; Alderw., Malayan Ferns (1908)
515. — Lectotype (selected here): Barber s.n.,
Borneo (K).

Aspidium kunstleri Bedd., Handb. Ferns Brit. India
Suppl. (1892) 44; Alderw., Malayan Ferns
(1908) 236. — Type: Kunstler 405, Perak, Go-
peng (K).

Phegopteris polycarpa Mett., in Kuhn, Linnaea 36
(1869) 124. — Dictyopteris polycarpa (Mett.)
Bedd., Ferns Brit. India Suppl. (1876) 20;
Handb. Ferns Brit. India (1883) 300. — Aspi-
dium kuhnii C. Chr., Index Filic. (1905) 78
(not A. polycarpum Blume). — Type: Jagor 222,
Malacca (B).

Caudex short, erect, its apex covered with stiff
brown scales 15 x 1.5 mm; fronds of young plants
3—5-lobed, of older ones with a trilobed apical
lamina and a pair of bilobed acuminate pinnae, all
lobes entire; fronds of mature plants with stipe to
70 cm long and lamina to 50 cm long, stipe and
rachis dark brown and bearing short erect hairs;
pinnae to 7 pairs; basal pinnae commonly to 18
cm long, stalked (stalk 2 cm), deeply lobed on the
basiscopic side, with basal lobe to 12 cm long,
shallowly lobed on acroscopic side; middle pinnae
with + sinuous margins or irregularly lobed on
the largest fronds; main veins in pinnae 8-10 mm
apart, smaller veins anastomosing between them
with many excurrent free veinlets in the areoles;

Sori small, mostly at the ends of short veinlets in
areoles, when very young having (at least in some
cases) small reniform indusia but later appearing
exindusiate; tetraploid (Manton in Holttum 1955).
— Fig. 12h, i.

Distribution — Southern Burma and Thailand; in
Malesia: Sumatra, Malay Peninsula, Borneo, Sulu
Archipelago.

Habitat — In lowland forest.

Notes — Hooker cited three syntypes for Poly-
podium barberi: Wallich 379 from Tavoy, Wallich
s.n. from Penang and Barber s.n. from Borneo.
The Tavoy specimen was part of the material
named Aspidium variolosum by Wallich. The types
of A. kunstleri Bedd. and Phegopteris polycarpa
Mett. are unusually large specimens. Mettenius
noted the resemblance of the latter to A. barberi
Hook.; Christensen saw the type and confirmed to
me that it is not distinct.

Manton found a plant from Singapore to be tetra-
ploid. Experiment is desirable to indicate the rela-
tionship of this species to others; if it is allotetra-
ploid, one parent is probably T. impressa.

15. Tectaria bamleriana (Rosenst.) C. Chr.,
Index Filic. Suppl. 3 (1934) 177. — Aspidium
bamlerianum Rosenst., in Feddes Repert. Spec.
Nov. Regni Veg. 10 (1912) 338; Alderw.,
Malayan Ferns, Suppl. (1917) 192. — Type:
G. Bamler W10, Wareo, NE New Guinea (L).

Aspidium beccarianum Baker, Malesia 3 (1886)
43; Alderw., Malayan Ferns (1908) 238. — As-
pidium cesatianum C. Chr., Index Filic. (1905)
68, nom. nov. superfl. — Tectaria cesatiana
(C. Chr.) Copel., Philipp. J. Sci. 6 (1911) Bot.
76; ibid. 78 (1951) 416, nom. illeg. — Type:
Beccari s.n., Andai, W New Guinea (FI).

Aspidium ledermannii Brause, Bot. Jahrb. Syst. 56
(1920) 114. — Tectaria ledermannii (Brause)
C. Chr., Index Filic. Suppl. 3 (1934) 181. —
Type: Ledermann 9409, NE New Guinea (B).

Aspidium pleocnemioides Alderw., Nova Guinea 14
(1924) 6. — Tectaria pleocnemioides (Alderw.)
C Chr., Index Filic. Suppl. 3 (1934) 183. —
Type: H.J. Lam 423, W New Guinea, Mamber-
amo River (BO).

Aspidium evenulosum Alderw., Bull. Jard. Bot. Bui-
tenzorg II, 28 (1915) 7. — Tectaria evenulosa
(Alderw.) C. Chr., Index Filic. Suppl. 3 (1934)
179. — Type: Kornassi 835, SE Ceram (BO).

Holttum — Tectaria Group 53

P.J. Epwarns

Fig. 12. Tectaria manilensis (Presl) Holttum. a. Complete frond, x 0.33; b. venation at base of upper
(adnate) pinna, undersurface, x 2.6; c. thick hairs on upper surface of lamina between veins, x 33. —
T. laxa (Copel.) Price. d. One tertiary leaflet, x 2.2. — T. aurita (Sw.) S. Chandra. f. Whole frond, x 0.33;
e. sterile and fertile fronds, x 0.26. — T. bamleriana (Rosenst.) C. Chr. g. Venation of an acroscopic
lobe, x 1.7. — T. barberi (Hook.) Copel. h. Venation of a fertile pinna, x IE venation of a sterile
pinna, x 1.7 (a—c: Smith 3054; d: Price 3345; e: Croft 1682; f, g: Ratcliffe 188; h, 1: Piggott 2968).

54

Flora Malesiana ser. II, Vol. 2! (1991)

Caudex erect, its apical scales firm, to 10 x 1
mm, similar scales at base of stipe, those above
base very narrow; stipe and rachis dark, glossy on
abaxial surface; lamina to 30 cm long, variably
+ dimorphous in fronds of the same collection;
pinnae to 3 pairs; apical lamina deltoid, very deep-
ly lobed, the larger lobes lobulate; basal pinnae
short-stalked, to 18 cm long, lacking free pinnules,
basal basiscopic lobes to 8 cm long deeply lobed
when fertile; middle pinnae sessile, very deeply
lobed, the lobes lobulate or crenate, widest on
sterile fronds with narrow sinuses between them,
narrower with wider sinuses when fertile, a bud
present on the acroscopic margin near base of a
pinna; veins not distinct on lower surface, forming
narrow costal and costular areoles and other ones
about as in T. impressa, included veinlets usually
few; lower surface of pinnae glabrous or with
sparse minute hairs on veins; upper surface of cos-
tae densely short-hairy, a few thick hairs present
between veins near margin; venation of fertile pin-
nae usually less complex, the sori in one row on
each side of costules in pinna-lobes; indusia to
nearly 1 mm diameter, reniform, dark, firm, some-
times bearing short hairs. — Fig. 12g.

Distribution — Malesia: Moluccas (SE Ceram),
widely in New Guinea, Manus Is.

Habitat — In lowland forest, twice reported
growing among rocks, in one case on limestone
(Manus Is.), once at 1100 m on a Steep ridge.

Note — The Ceram specimen is apparently a
young plant; the fronds have only one pair of pin-
nae but the largest is fertile.

16. Tectaria nabirensis Holttum, Blumea 35
(1991) 553. — Type: W. Schénian 38, W New
Guinea, Geelvink Bay, Nabire, 100—400 m (B).

Caudex and bases of stipes lacking; stipe and
rachis very dark, glossy, bearing throughout many
narrow light brown scales; lamina incomplete,
probably 50 cm long with c. 4 pairs of pinnae;
apical lamina 20 cm long, lobed to a narrow wing,
lobes to 10 pairs, lowest 8 x 1.5 cm, deeply close-
ly obliquely lobulate; basal pinnae 28 cm long
(stalk 3 cm) with 1 pair free pinnules and 1 pair
adnate, basal basiscopic pinnule 17.5 x 7 cm bear-
ing one free tertiary leaflet, the rest lobed to a
narrow wing, the lobes close, oblique and deeply
lobulate; veins distinct on lower surface, forming
costal and costular areoles, other areoles below

sinuses few; lower surface glabrous apart from
sparse minute hairs on costae and costules; dense
short hairs present on upper surface of costae and
a few hairs near sinuses; sori on free veins near
apices of pinna-lobes, otherwise on anastomosing
veins; indusia rather large, thin, glabrous.

Distribution — Malesia: W New Guinea (known
only from the type.

17. Tectaria griffithii (Baker) C. Chr., Index
Filic. Suppl. 3 (1934) 180; Tard. & C. Chr., FI.
Indo-Chine 7 (1941) 411; Holttum, Revis. FI.
Malaya 2, ed. 2 (1968) 636. — Nephrodium
griffithii Baker, Syn. Fil. (1867) 300. — Sage-
nia griffithii (Baker) Bedd., Ferns Brit. India
(1870) t. 337. — Type: Griffith s.n., Burma (K).

Nephrodium multicaudatum Clarke, Trans. Linn.
Soc. London II, Bot. 1 (1880) 540, t. 77. —
Aspidium multicaudatum (Clarke) Bedd., Handb.
Ferns Brit. India (1883) 222; Alderw., Malayan
Ferns (1908) 250. — Tectaria multicaudata
(Clarke) Ching, Sinensia:2 (1931) 20; Holttum,
Revis. Fl. Malaya 2 (1955) 507, f. 299. —
Type: Clarke 18427, Sylhet (K).

Aspidium malayense Christ, Philipp. J. Sci. 2
(1907) Bot. 187. — Tectaria malayense (Christ)
Copel., Philipp. J. Sci. 2 (1907) Bot. 416;
C. Chr., Gard. Bull. Straits Settlem. 7 (1934)
260; Copel., Fern Fl. Philipp. (1960) 305. —
Lectotype (selected here): Loher 858, Luzon (P;
iso K).

Pleocnemia trimenii Bedd. forma dissecta Alderw.,
Bull. Jard. Bot. Buitenzorg III, 2 (1920) 164. —
Type: Lérzing 5403, Sumatra, Sibolangit (BO;
iso L).

KEY TO THE VARIETIES

la. Basal pinnae with 1 pair of pinnules..... 2
b. Basal pinnae with 5 pairs of pinnules
c. var. amplissima
2a. Veins amply anastomosing in fertile fronds
a. var. griffithii
b. Veins all free in lobes of fertile pinnae
b. var. singaporeana

a. var. griffithii

Caudex erect, its apex covered with scales to 20
mm long, 2 mm wide at the base; stipe to 60 cm
long, dull dark brown, bearing many very narrow

Holttum — Tectaria Group

55

scales throughout; /amina deltoid-pentagonal, to
60 cm long and wide; free pinnae 2—4 pairs, wide-
ly spaced; basal pinnae commonly 30 cm long (to
45 cm) with 1 pair of free pinnules, the rest lobed
to 5-10 mm from the costa, the costules 3—4 cm
apart, the lobes acuminate; basal basiscopic pin-
nule to 20 x 6 cm, widest in the middle, deeply
lobed with costules 1.5 cm apart, lobes falcate,
acute, entire; acroscopic pinnule to 10 x 2.8 cm,
shallowly lobed; second pinnae of large fronds
sessile with one free pinnule; veins concolorous
but distinct on lower surface, forming narrow cos-
tal and costular areoles, also other areoles below
sinuses (often with short excurrent included vein-
lets) and an additional outer row in lobes of pin-
nae, costular areoles not formed in distal parts of
lobes of fertile pinnae; Jower surface glabrous or
sparsely hairy on costae and costules, usually with
some narrow scales; upper surface of costae and
costules densely covered with short thick hairs,
groups of similar hairs present near margins or at
sinuses; sori on free veins in areoles near base of
pinna-lobes, on acroscopic branches of quite free
veins distally; indusia large, firm, persistent.

Distribution — Assam (Khasia Hills), Burma,
Vietnam, Thailand; in Malesia: N Sumatra, Malay
Peninsula, W Java, Borneo, Celebes, Philippines
(Luzon to Mindanao).

Habitat — In lowland forest, sometimes near
limestone.

b. var. singaporeana Holttum, Blumea 35
(1991) 551. — Type: C.G. Matthew s.n., Jan.
1908, Singapore, Bukit Timah (K).

Differs from the type of the species because the
veins in the lobes of the fertile pinnae are totally
free. Anastomosis in sterile pinnae is also much
less than in var. griffithii, the outer row of areoles
in lobes being lacking.

Distribution — Malesia: Singapore (known only
from Bukit Timah).

c. var. amplissima Holttum, Blumea 35 (1991)
551. — Type: C.G. Matthew s.n., Jan. 1913,
Sumatra, Padang Panjang, Bukit Tilabung (K).

Pinnae to 12-jugate, the lowermost 50 cm long
provided with 6-jugate pinnules, the lowermost
basiscopic pinnules provided with 3-jugate tertiary
leaflets; raches and costae beneath densely short

hairy.

Distribution — Malesia: Sumatra (known only
from the type collection.

18. Tectaria nesiotica Holttum, Blumea 35
(1991) 553. — Type: J.R. Croft & J. Marsh
LAE 71234, Goodenough Is. (K; iso L, LAE).

Caudex erect; frond with stipe 200 cm long, the
stipe 80 cm (collectors); rachis dark brown, glabres-
cent and glossy on abaxial surface, hairy adaxially;
basal pinnae not seen, suprabasal pinnae subses-
sile, to 42 cm long, bearing 12 pairs of sessile free
pinnules and 3 or 4 pairs adnate; pinnules 2.5 cm
apart, to 8.7 x 1.7 cm, lobed to 1 mm from the
costa, lobes oblique, little more than 2 mm wide
and separated by wider sinuses, their margins cre-
nate, apices obtuse, basal basiscopic and acro-
scopic lobes subequal and not reduced; veins form-
ing narrow areoles along costae of pinnules, those
in the lobes all free, a forked one corresponding to
each crenature; upper surface of costae of pinnules
densely short-hairy near the base only, lower sur-
face bearing a few narrow scales; sori on acroscopic
branches of veins, apparently terminal; indusia dark,
firm, glabrous.

Distribution — Malesia: New Guinea (known
only from the type collection).

Habitat — In forest, at 950 m.

19. Tectaria kingii Copel., Philipp. J. Sci. 9
(1914) Bot. 4; ibid. 78 (1951) 414. — Pleo-
cnemia kingii (Copel.) Alderw., Malayan Ferns,
Suppl. (1917) 147; Holttum, Reinwardtia 1
(1951) 186. — Type: C. King 402, Woodlark
Is. (MICH).

Caudex erect or + prostrate; stipe to 45 cm long,
castaneous, adaxially glabrous apart from scales;
basal scales to 20 mm long and more than 2 mm
wide; lamina to 40 cm long; pinnae to 4 pairs free
and 3 pairs + adnate; apical lamina small; basal
pinnae to 30 cm long, free pinnules to 4 pairs,
basal basiscopic pinnule to 17 cm long with 2
pairs of deeply lobed tertiary leaflets (largest 3.5
x 1.7 cm) and 2 pairs adnate; basal pinnules on
second pair of pinnae bearing 1 pair of tertiary
leaflets; pinnules on middle pinnae very deeply
lobed with + crenate lobes; veins forming costal
and costular areoles, usually no additional areoles
below sinuses between pinna-lobes; lower surface
almost glabrous; upper surface of rachis and pinna-

56

Flora Malesiana ser. II, Vol. 2! (1991)

rachises densely short-hairy, surface between veins
bearing very few hairs except near sinuses; sori
mostly at the ends of acroscopic branches of veins
in pinnule-lobes, marked by prominences on the
upper surface; indusia thin, at first covering sori,
shrivelling later and then not conspicuous.

Distribution — Malesia: New Guinea (Woodlark
Is., Sudest Is., New Britain, Admiralty Is.; Milne
Bay Prov. in Papua New Guinea); Santa Cruz Is.

Ecology — In lowland forest, on steeply sloping
ground, in one case on limestone.

20. Tectaria squamipes Holttum, Blumea 35
(1991) 555. — Type: J.R. Croft 1753, Papua
New Guinea, West Sepik Prov. (K).

Caudex erect; stipe at least 40 cm long, dark,
glabrescent on abaxial surface, bearing many nar-
row mid-brown scales throughout, similar scales
also present on rachis and on costae of pinnae;
lamina to 50 cm long; free pinnae 3 pairs and 2
pairs adnate; basal pinnae to 23 cm long (stalk 7
mm) bearing 1 pair of free or adnate pinnules;
basal basiscopic pinnules 12 x 3 cm, lobed to 3
mm from their costae, costules of lobes to 12 mm
apart, lobes 8 mm wide, deeply crenate, obtuse,
basal acroscopic pinnule 5 cm long, rest of basal
pinnae lobed to 2—3 mm on each side of their cos-
tae; suprabasal pinnae almost sessile, deeply lobed
with oblique obtuse deeply lobulate lobes; veins
forming costal and costular areoles with few addi-
tional areoles below sinuses; lower surfaces gla-
brous or nearly so apart from scales; thick hairs
present between veins on upper surface; sori most-
ly on acroscopic branches of free veins in pinna-
lobes; indusia small and apparently caducous.

Distribution — Malesia: Papua New Guinea (also
known from Milne Bay Prov.).

Habitat — The type from forest on exposed
summit ridge at 1100 m; the Milne Bay specimen
from forest at 430 m.

21. Tectaria adenophora Copel. in Elmer,
Leafl. Philipp. Bot. 4 (1911) 1151; Fern FI.
Philipp. (1960) 306. — Type: Elmer 12419,
Sibuyan Is. (MICH, K).

Caudex apparently short-creeping with tufted
fronds: stipe 20—30 cm long, ferrugineous, short-
hairy, scaly near the base only, scales black, rigid,
7 mm long, less than 1 mm wide; lamina to 20

cm long, consisting of 2 pairs of pinnae and a
deeply lobed apical section 12 cm long, dark rusty
brown when dried; basal pinnae of type 8 cm long
(stalk 2 mm), acuminate, lobed to 4-5 mm from
its costa, basal basiscopic lobe 6 x 1.5 cm and
shallowly lobulate, the other lobes subentire, of
Ramos 4710 to 20 cm long; upper pinnae slight-
ly adnate, lobed halfway to costa, lobes oblique,
rounded; veins forming narrow areoles along costae
and costules, also a second series below sinuses
and in broader lobes; many very short erect hairs
between veins on both surfaces; sori mostly on the
outer veins of costal and costular areoles; indusia
small, bearing short hairs, soon caducous.

Distribution — Malesia: Philippines (known
from the type and 2 collections from Zambales
Province, Luzon).

22. Tectaria christii Copel., Philipp. J. Sci. 2
(1907) Bot. 416; in Elmer, Leafl. Philipp. Bot.
4 (1911) 1151; Fern Fl. Philipp. (1960) 305.
— Aspidium coadunatum auct. (non Wall. ex
Hook. & Grev. 1831): Christ, Philipp. J. Sci.
2 (1907) Bot. 187. — Lectotype (Copeland
1911): Copeland 1899, Luzon, Bontoc, Sagada
(MICH).

Caudex short-creeping; stipe to 20 cm long,
slender, castaneous, glabrescent on abaxial side;
basal scales to 8 x 1 mm, thin, light brown; /ami-
na of type 28 cm long, consisting of apical section
and 2 pairs of pinnae; apical section 21 cm long,
deltoid, lobed to 5S—6 mm from its axis, basal lobes
8 x 2.7 cm, obliquely lobulate halfway to costa, the
lobules broadly rounded; upper pinnae adnate, 9 x
3.5 cm, lobed halfway to costae, lobes slightly
crenate with rounded apices, apex acute, not acumi-
nate; basal pinnae 13 cm long, lobed to 5 mm from
costae, the lobes + overlapping, basal basiscopic
lobe 6.5 x 3.3 cm, deeply lobulate with crenate
lobules and a very obtuse apex; acroscopic lobes
all about equal, oblique, 3 x 1.5 cm; veins forming
costal and costular areoles, also several additional
ones below sinuses and an outer row in pinna-
lobes; lower surface of costae bearing slender hairs
to 1 mm long, short erect hairs present on and be-
tween veins; upper surface bearing copious short
hairs between veins; sori in one row on each side
of main veins in pinna-lobes, on the outer veins of
areoles or on short veins within the second row of
areoles; indusia large, thin, glabrous.

Holttum — Tectaria Group

a,

Distribution — North Thailand; Malesia: Philip-
pines (Luzon), Borneo (Sabah; one collection).

Habitat — In Luzon on limestone at 1500 m
(type) and near a waterfall; in Sabah pendulous on
rocks by stream at 1000 m.

Note — This is very near T. coadunata but appears
to differ constantly in lack of pinnules on basal
pinnae and in closely shallowly lobed non-acumi-
nate upper pinnae. The largest Luzon specimen has
a lamina 38 cm long, basal pinnae 19 cm.

23. Tectaria devexa (Kunze ex Mett.) Copel.,
Philipp. J. Sci. 2 (1907) Bot. 415; Ching, Si-
nensia 2 (1931) 16; Backer & Posth., Varenfl.
Java (1939) 72; Holttum, Revis. Fl. Malaya 2
(1955) 505, 626, f. 297; Copel., Fern Fl. Phi-
lipp. (1960) 304; Holttum & Roy, Blumea 13
(1965) 135; Sledge, Kew Bull. 27 (1972) 416;
Holttum, Gard. Bull. Sing. 34 (1981) 136; In-
dian Fern J. 1 (1985) 36. — Aspidium devexum
Kunze, Bot. Zeitung (Berlin) 6 (1848) 259,
nom. nud. — Aspidium intermedium Mett.,
Farngatt. IV (1858) 119, non Willd. 1810. —
Aspidium devexum Kunze ex Mett., Ann. Mus.
Bot. Lugd.-Bat. 1 (1864) 237, in obs. — Pleo-
cnemia devexa (Kunze ex Mett.) Alderw., Ma-
layan Ferns (1908) 174, incl. var. permutata
Alderw. — Type: Zollinger 2717, Java (LZ,
lost; iso B, G, L).

Aspidium membranaceum Hook., Sp. Fil. 5 (1864)
105 in footnote. — Pleocnemia membranacea
(Hook.) Bedd., Ferns Brit. India Suppl. (1875)
15; Handb. Ferns Brit. India (1883) 225. —
Lectotype (Holttum 1981): Cuming s.n., Phi-
lippines (K).

KEY TO THE VARIETIES

la Lobes of upper pinnae deeply lobulate; veins
forming few areoles beyond those along costae

aNGl COMMENTS Ae Gece on ce eee ae eee 2

b. Lobes of upper pinnae at most crenate; addi-
tional areoles frequent below sinuses between
pinna-lobes .... c. var. novoguineensis

2a. Both surfaces densely short-hairy between the
VEU S Herre iaret cite te Slaw ie a. var. devexa

b. Upper surface bearing scattered hairs 0.5 mm
long; lower surface between veins almost gla-
DEOUSH Ee Rie 2 Us Se) pee b. var. minor

a. var. devexa

Caudex short, suberect; stipe to 30 cm long,
slender, light castaneous, glabrescent; basal scales
to 10 mm long, less than 1 mm wide; lamina to
35 cm long, thin; pinnae 3—5 pairs; basal pinnae
to 18 cm long (stalk 1.5 cm) with 1 pair of free
pinnules, basal basiscopic pinnule to 8 x 3.5 cm,
very deeply lobed, the lobes lobulate up to half-
way to their costules, basal one sometimes almost
free; upper pinnae adnate on the basiscopic side,
lobed as pinnules of basal ones, costae of lobes to
10 mm apart; veins forming costal and costular
areoles, additional ones below sinuses few; lower
surface of costae bearing hairs 0.5 mm long, whole
lower surface on and between veins bearing many
very slender shorter hairs; upper surface similarly
hairy with thicker hairs; sori near the margins of
lobules of pinna-lobes, mostly on the acroscopic
branches of free veins; indusia thin, glabrous or
with short hairs; n = 40 (Manton in Holttum
1955). — Fig. 1b, f.

Distribution — Southern Burma and Thailand;
Malesia: Western Malesia, Central and SW Celebes,
Philippines. Specimens from Taiwan, SW China
and Tonkin have sparse longer hairs on the upper
surface but are otherwise similar.

Habitat — Always on limestone, usually at the
base of rocks.

Taxonomy — The first description of this species
(as Aspidium intermedium) was by Mettenius in
1858, with citation of A. devexum as a synonym;
the latter name was not validated until 1864, al-
most simultaneously with the publication of A.
membranaceum Hook.

b. var. minor (Hook.) Holttum, Blumea 35 (1991)
550 — Aspidium giganteum Blume var. minor
Hook., Sp. Fil. 4 (1862) 50. — Sagenia gigan-
tea var. minor Bedd., Ferns S. India (1864) 81,
t. 243. — Type: Thwaites CP 1358, Sri Lan-
ka (K).

Frond-form and size as var. devexa, differing in
almost glabrous lower surface between veins and
sparse thick hairs 0.5 mm-long between veins on
upper surface.

Distribution — Sri Lanka, Christmas Island (In-
dian Ocean).

58

Flora Malesiana ser. II, Vol. 21 (1991)

c. var. novoguineensis Holttum, Blumea 35
(1991) 550. — Type: Holttum s.n., cult. Kew,
origin limestone near Kundiawa, E New Guinea

(K).

The type plant cultivated at Kew was found by
S.K. Roy to be tetraploid [Blumea 13 (1965) 135].
Brass 32188, from a neighbouring locality, has
larger fronds with lamina to 40 cm long and reg-
ular pluriseriate areoles below sinuses; its highest
pinnae have costules of their crenate lobes 12 mm
apart; costuies of upper pinnae of var. devexa of
comparable size are 6 mm apart.

Distribution — Malesia: Eastern New Guinea,
New Hebrides (Efate).

24. Tectaria ferruginea (Mett.) Copel., Phi-
lipp. J. Sci. 6 (1911) Bot. 76; ibid. 78 (1951)
415. — Phegopteris ferruginea Mett., Ann. Mus.
Bot. Lugd.-Bat. 1 (1864) 224, t. 6, f. 2. — As-
pidium zippelianum C. Chr., Index Filic. (1905)
98 (not A. ferrugineum Sw. 1806). — Dictyo-
pteris ferruginea (Mett.) Alderw., Malayan Ferns
(1908) 516. — Type: Zippelius s.n., Western
New Guinea (L).

Pleocnemia membranacea (Hook.) Bedd. var. novo-
guineensis Rosenst., Feddes Repert. Spec. Nov.
Regni Veg. 10 (1912) 335. — Type: G. Bamler
5, Papua New Guinea, Logaueng, 1909 (no 63,
same locality, 1910, at K, L).

Tectaria gymnocarpa Copel., Philipp. J. Sci. 9
(1914) Bot. 4; ibid. 78 (1951) 415. — Type:
G. King 401, Papua, Loane (MICH).

Frond-form as T. kingii but basal pinnae shorter
(to c. 25 cm), their basal basiscopic pinnules bear-
ing at most one pair of adnate tertiary leaflets; veins
often forming additional areoles below sinuses in
sterile fronds, not in fertile ones; hairs always
present on lower surface of costae, costules and
veins, variable, sometimes to 1 mm long, few or
none between veins; hairs on upper surface be-
tween veins abundant, at least 1 mm long (to 1.5
mm); sori exindusiate, usually rather large and
slightly irregular in shape.

Distribution — Malesia: widely in New Guinea
(only the type from the West); Solomon Islands.

Habitat — In forest from low altitudes to 2000
m, twice recorded on limestone. Plants found at
higher altitudes appear to be more hairy on both

surfaces, with longer hairs, than those from low
altitudes, and to have smaller fronds.

Note — The types of Pleocnemia membranacea
var. novoguineensis and Tectaria gymnocarpa both
have fewer and shorter hairs on the abaxial side of
rachises than most other specimens and fewer thick
hairs between veins on the upper surface; they are
thus nearer to T. kingii.

25. Tectaria coadunata (J.Sm.) C. Chr., Contr.
U.S. Natl. Herb. 26 (1931) 331; Ching, Sinen-
sia 2 (1931) 18, p.p. — Aspidium coadunatum
Wall. ex Hook. & Grev., Icon. Filic. (1831) 202,
non Kaulf. 1824. — Sagenia coadunata J.Sm.,
J. Bot. (Hook.) 4 (1841) 184, nom. nov. —
Sagenia macrodonta Fée, Mém. Foug. 5. Gen.
Filic. (1852) 313, nom. nov. superfl. — Tec-
taria macrodonta (Fée) C.Chr., Index Filic.
Suppl. 3 (1934) 181; Tard. & C. Chr., Fl. Indo-
Chine 7 (1941) 410 (p.p.?); — Type: Wallich
377, Nepal (E).

Aspidium cicutarium auct.: Hook., Sp. Fil. 4 (1862)
61, p.p.; Bedd., Handb. Ferns Brit. India (1883)
220, p.p. — Nephrodium cicutarium auct.:
Baker, Syn. Fil. (1867) 299, p.p.; Clarke, Trans.
Linn. Soc. London II, Bot. 1 (1880) 539, excl.
var. coadunatum.

Note on the species — In the North of India
Clarke distinguished two varieties, one with large
fronds glabrous on the lower surfaces and one
somewhat smaller with fronds hairy beneath. He
wrongly called the latter var. coadunatum; it is
variable and has not been clearly distinguished. In
northern Thailand, on limestone, there is a more
densely hairy form which may represent a distinct
variety.

KEY TO THE VARIETIES

la. Basal pinnae of mature plants to 50 cm long,
pinnate; those of small plants deeply lobed but
not pinnate’... a. var. coadunata

b. Basal pinnae to 15 cm long, pinnate
b. var. minor

a. var. coadunata

Fronds large, with basal pinnae to 50 cm long
bearing a pair of free pinnules; lobes of pinnae and

Holttum — Tectaria Group

59.

pinnules very broad; veins forming costal and cos-

tular areoles lacking free included veinlets and many
other areoles, some with short included veinlets;
lower surfaces glabrous, thick hairs present be-
tween veins on upper surface.

Distribution — NE India, SW China, N Thailand.

b. var. minor Holttum, Blumea 35 (1991) 518.
— Tectaria macrodonta auct.: Holttum, Revis.
Fl. Malaya 1, 2 (1955) 505, quoad plantas penin-
sulares. — Type: Henderson SF 21383, Pulau
Langkawi (K; SING).

Caudex short-creeping; stipe to 50 cm long, its
base swollen and persistent; basal scales thin, to 5
mm long; lamina of type 26 cm long consisting
of 3 pairs of pinnae and a deeply lobed deltoid api-
cal portion 15 cm long (its lowest lobes 5.5 x 1.5
cm); basal pinnae 15 crn long (stalk 10 mm), their
basal basiscopic pinnules 9 x 2 cm, lobed halfway
to the costa, the lobes obtuse, basal acroscopic pin-
nules 3.5 x 1.4 cm; suprabasal pinnae sessile, deep-
ly lobed, the lobes falcate, obtuse, the larger ones
with sinuous margins; hairs 0.2—0.3 mm long
present on lower surface of costae and a few very
slender ones between veins, many thicker hairs
present between veins on upper surface; sori most-
ly on free veins in outer areoles, marked by de-
pressions on the upper surface when dried; indusia
rather large, glabrous.

Distribution — Malesia: W Malaysia (Langkawi
Is., Perlis).

Habitat — On limestone.

26. Tectaria pubescens Copel., Univ. Calif.
Publ. Bot. 18 (1942) 221; Philipp. J. Sci. 78
(1951) 415, pl. 16. — Type: C. King 359, Pa-
pua, Lakekamu (MICH; iso UC).

Caudex short, erect; stipe to 50 cm long; basal
scales to 15 x 1 mm, those above base gradually
shorter, very narrow scales present on abaxial
surface of both stipe and rachis, hairs 0.5 mm long
also on rachis; /amina to 60 cm long consisting
of 3—5 pairs of pinnae and a terminal portion c.
15 cm long; basal pinnae to 28 cm long (stalk
3-5 cm) bearing 3 or 4 pairs of pinnules with a
deeply lobed apex like that of the lamina, basal
basiscopic pinnule to 12 cm or more long, broadly
lobed to 3—5 mm from its costa with 1 or 2
tertiary leaflets to 5 x 1.5 cm lobed halfway to the
costule; first suprabasal pinnae with 1 pair of free
pinnules; lobes of fertile pinnae and pinnules nar-
rower than sterile; veins prominent on both sur-
faces, forming additional areoles below sinuses in
less deeply lobed parts of pinnae: rather sparse
hairs up to 1 mm long present on lower surfaces
of costae and costules, also scattered on veins,
thick hairs more than 0.5 mm long, scattered be-
tween veins on upper surface; sori in a row half-
way between costules of lobes and margin, the
lower ones on short veins in areoles, distal ones
on free veins; indusia rather large, glabrous, shriv-
elling when old.

Distribution — Malesia: NE New Guinea; Solo-
mon Islands (Bougainville).

Habitat — In forest at 500-600 m.

Section Tectaria

Taxonomy — The generic names Hemigramma Christ and Quercifilix Copel., cited
as synonyms above, call for comment. The sterile and fertile fronds of the types of both
genera are strongly dimorphous, the sterile ones having venation as in Tectaria sect. Tec-
taria, the fertile ones having a greatly contracted lamina with simplified venation and spor-
angia borne all along the veins, giving an acrostichoid appearance.

Christ established the genus Hemigramma (1907) for a Java fern originally named
Hemionitis zollingeri Kurz (Tectaria zollingeri here) with the allied Philippine species
(here named Tectaria hilocarpa) as a variety. Copeland (1928) added other species, two of
which (from Hong Kong and Tonkin) have broader fertile pinnae in which the venation,
though contracted, is normal for Tectaria sect. Tectaria; these two are not nearly related to
the type of Hemigramma. All these ferns, including the type of Quercifilix, represent dif-
ferent modifications of fertile fronds which have evidently arisen on different evolutionary

60

Flora Malesiana ser. II, Vol. 2! (1991)

lines. The Philippine Tectaria hilocarpa hybridizes naturally with Tectaria crenata. Copeland
in 1928 also included in Hemigramma the very different species here named Chlamydo-
gramme hollrungii, not noticing its peculiar indusia.

The vein-pattern in sect. Tectaria is closely similar to that in several genera of Polypo-
diaceae; this appears to be a remarkable example of parallel evolution.

KEY TO THE SPECIES OF SECTION TECTARIA

la. Fronds pinnate or bipinnate, or elongate and lobed to a winged rachis ....... D
bs Fronds. simple;notielongateifdeeply lobed. 2.52. 24554. 2 2a ee 74
ZasRachismotawhollyswingeds) 1 06% 5.0. 86 ae Geen 2 Ea 3
b. Rachis wholly winged (also part of stipe) or the wing not quite continuous in mature
PLANES Ses ee ees uu Ee Pe Ok Walaa tease 5 gay ae 61
3a. Suprabasal pinnae deeply lobed or pinnate in fronds of mature plants ....... 4
b. Suprabasal pinnae, if any, entire or nearly so in fronds of mature plants; pinnules, if
any,wnatrow, entireror nearly: SO. .k.. eke es aks dln kee ee 19
davRachisivery darkcand/atlength’glossy . 22435202 9 2 5
beeRachisinot very darknoratlengthslossy (425 002. | ee eee 12
Sas Son smallsinrregularly scattered’) 0. 25: en ne eo ee ee 6
b. Sori larger, in one row on each side of main veins or of costules near apices of
PINMA=LODES 44d crashes aa ees, Wee aeensnesy + "o, Suede bep satel, bl tyconctue ys 0 cece eae ae 9
6a. Indusia small: sori not. confluent 4. s 4. 48 4.6 oe eee 7
b. Indusia lacking; sori sometimes confluent ............... 30. T. simonsii
2. bhickshairs on upper surtace confined to sinuses @ 4 ae 2) ee 8
b. Thick hairs between veins generally on upper surface ....... 29. T. holttumii
8a. Lobes of basal pinnae deeply lobulate ............... 27. T. melanocaula
bigeobes of basalypinnac entire Or nearly sol. «hens cee 28. T. nebulosa
9a. Suprabasal pinnae shallowly lobed apart from the acute basal lobes
31. T. durvillei
b. Suprabasal pinnae deeply lobed with acute to acuminate lobes ............ 10
10a. Basal pinnae of largest fronds 30 cm or more long bearing at most 1 pair of
pinnules;/|suprabasal pinnae lacking pinnules. .......4.2...). «eee 11
b. Basal pinnae of largest fronds less than 30 cm long, bearing 2 pairs of pinnules; 2
pairs of suprabasal pinnae each with 1 pair of pinnules .... 34. T. athyriosora
lla. Basal pinnae to 30 cm long; lower surface of costae and veins bearing hairs more
thany Q'S, mam Ome es eas oan Bil hag we hs ears |e oes 32. T. melanorachis
b. Basal pinnae to 60 cm long bearing a pair of stalked pinnules; all parts of lower
surface bearing minute hairs with spherical tips ......... 33. T. danfuensis
12a. Fronds strongly dimorphous; sterile pinnae broad and shallowly lobed
35. T. brachiata
b. Fronds not strongly dimorphous; pinnae deeply lobed.................. 13
IBa\ Son allinearn marcins, of pinna-lobeSian seen le eee 36. T. calcarea
b. Son. mostly notinear Margin yg... hey Seudck tna cee eee 14

Holttum — Tectaria Group 7 61

14a.

Some sori on free veinlets in areoles, or on branches of free veins near apices of
PUTTS O DSSS Pere si ee smn ren aneeestsoatt nay Cie) meee ete Mama leeleaaeuty eon Alas ker etia aplte Uae a 15

mS Otieallionmeconnectedwveinsxea. Et Ae Bind tied. ieee lsat dane teases 17
. Basal pinnae to more than 30 cm long; lobes of upper pinnae acuminate and deeply

NON CCM eee pce Sete ee ee eee het 37. T. curtisii

. Basal pinnae shorter; lobes of upper pinnae obtuse, at most crenate......... 16
. Basal scales 7 x 1.5 mm, thin, light brown; lamina very firm .. 38. T. cherasica
. Basal scales to 15 x 1 mm, firm, dark; lamina thin, translucent

39. T. translucens

. Sori indusiate; basal acroscopic lobes of pinnae not longer than the next lobes. . 18
. Sori exindusiate; basal acroscopic lobes of pinnae much elongate 42. T. macrota
. Thick hairs lacking between veins on upper surface; indusia glabrous

40. T. keckii

. Thick hairs present between veins on upper surface; indusia fringed with hairs

41. T. villosa

mBasalepinnae beanns narrow pinnules!« J.s50.: ci st ee ee 20
mBasalepinnae lacking free pintules..cc2 o a.g suds « espe Ghee cole oe aes nee 23
. Caudex long-creeping; lamina to 80 cm long ......... 43. T. semibipinnata
wa CauGexeshort, erect; lamunaimuch:Shoren 52. te ea. vo eer ia eee DA

. Lamina to 25 cm or more long; pinnae 2—4 pairs, pinnules entire or minutely den-

Mcilatessrac his Nairy/OM Upper SUMAace: 2:t1)Gel. aie ats. faverlets sncvaesnon coat et ae 22

. Lamina to 10 cm long; pinnae 1 pair, deeply crenate-lobate, each with 1 pinnule; no

MAIR OMePACHTS. insides oa Mea he eee ae eae Saree Se wea 46. T. subdigitata

. Distal parts of lower pinnae entire; almost all sori in 2 rows between main veins

44. T. lobbii

. Distal parts of lower pinnae irregularly obliquely lobed; sori small, numerous, irreg-

WlanlVeSCAltCled , ..h:2' a. see ale | Sees CON eae nents 45. T. jacobsii
. Fertile pinnae much contracted, covered beneath with sporangia........... 24
merle pinnae not very Narrow, MOST SOMsdistinct.. 0 42... Ses ZS
mbinnae oOftertile frond 1 pair;cs lecmioners. as). ee 47. T. zeilanica
. Pinnae of fertile frond several pairs, much longer....... 48. T. lombokensis
mberule pinnae to 1.5 cm wide; sterile to 3:5-cm 2.09525: 49. T. teratocarpa
. Fertile pinnae much wider where fronds are dimorphous................ 26
. Pinnae (including basal ones) all narrowly decurrent .................. pH
mEinnae not allinarowly Gecurrenti\sc neice See ae BU ea as 28

. No buds at bases of pinnae; indusia very small, not distinct on mature sori

50. T. subcaudata

. Buds present at bases of pinnae; indusia distinct ............ 51. T. fauriei
. A large acuminate lobe present on acroscopic base of pinnae, at least on basal ones

42. T. macrota

MeACTOSCOpic ibasallobestlackine ¢.s.ver 2a ete NI cee aes eee 29
. Sori irregularly arranged, some elongate or coalescent.................. 30
. Sori not coalescent, or if so in 2 rows between main veins............... 33

62 Flora Malesiana ser. II, Vol. 2! (1991)
30as)Suprabasalipinnaeinarrowed atithem bases) = 274.5 4s os ee 31
b. Suprabasal pinnae with broadly rounded to cordate bases ............... 32
31a. Sori mostly not confluent, some with small indusa; buds present at bases of upper
pinnachasr an. sce Rs URC ES EE Uke FER DA NEES EE raB 1A | 52. T. suluensis
b. Sori mostly confluent, no indusia, no buds ........... 53. T. semipinnata
32a. Many sori running along veins; no indusa ............. 54. T. sumatrana
b. Sori all on free veinlets in areoles, none linear; small indusia present
69. T. angulata
39ay Sonenear mareinsionly. pinnae pain eis. sa ia a 55. T. craspedocarpa
b. Sori not only near margins; pinnae more than 1 pair................... 34
34a. Sori in 1 row on each side of main veins, or with occasional additional ones es-
peciallysmeatmcostaci ei oe6 ie yy BE Sb)
ba Sori not only im) 1 row on’ each'side of main veins <2. ==. 55... se ee Sil
35a: Somoniireésveinletsin“areoles: aan wewsee ee ee 36
bsSorinotallvonsree veins antareolessan 5 6 4 eee 46
BOA SOKamuchmeElON Eaten seine, cccrusec ce Re ener eae 56. T. dolichosora
bs Sormnotrorlittlerclomgate ssai 220) Gn woe a A oi
37as Areolesin fertile pinnaeeach witha single vemletia 2. 7/45. {hae ee 38
b. Areoles in fertile pinnae mostly containing forked veinlets............... 39
38a. Basal pinnae unlobed: indusiaynot peltate W352. 52 57. T. pleiosora
be Basalapinnaclobeds indusiaypeltate = skeen meee ee 58. T. repanda
39aeStipe and rachisidarkvandslossy®: wes. ee) eek Sh: 58). 40
baStpezandirachis not darkiand glossyixt 4). 2. 2.2. oe As co: Se ee 43
40a. 9Pinnae notiauncledioniacroscopic Sideak ak) «alee es oe 41
baseinnaerauricledionythe acroscopic side aye ns ek 31. T. durvillei
41a. Pinnae commonly 25 cm long; basal scales 1.5—3 mm wide .... 59. T. crenata
b. Pinnae to 17 cm long; basal scales not over 1 mm wide................. 42
42a. Basal scales not contorted; middle pinnae to 3 cm wide ..... 60. T. isomorpha
b. Basal scales much contorted; pinnae to5 cm wide ....... 61. T. exauriculata
43a. Basal pinnae bearing at least 1 large basiscopic lobe ................... 44
bayBasalypinnacwunlobedts Segncl seer Pa) eg ehh Aye tee 57. T. pleiosora
44a. Bases of pinnae broad, cordate on basiscopic side ....... 62. T. brevilobata
b:)Bases/of pinnae subequally cuneate 2 ns Se eee 45
45a. Fertile pinnae 3—5 cm wide; indusia not peltate ............ 59. T. crenata
b. Fertile pinnae 2—3°5;cmiwideindusialpeltateyeeae eee 58. T. repanda
46a. Fronds with a simple or trilobed apex and 1 pair of unlobed pinnae
63. T. ternata
b. Fronds mostly with several pairs of pinnae, basal ones lobed. ............ 47
47aeSonlresgularly spacedtand separates j.)spcew tae cae econ ee ae 48
baySori iresular often: elongate and confluent)-4.4 7... > -1e eee 50
48a. Indusia large; bases of pinnae cordate on basiscopic side. ............... 49
b. Indusia very small; bases of pinnae cuneate to rounded both sides

66. T. decastroi

Holttum — Tectaria Group 63

. Stipes not very dark; basal pinnae with 2 basiscopic lobes ... 64. T. tabonensis
. Stipes nearly black; basal pinnae with one large basiscopic lobe

65. T. subcordata

. Pinnae to 2 cm wide, no buds on rachis; sori often elongate or confluent parallel to

EDEN, SST SS Ae Mace nae em ee ene eR AL ee er LEE it oh 67. T. rufescens
. Pinnae wider; buds present at bases of pinnae; lower sori elongate at right angles to

SSMU, SLIDE ee eis ee reek sce eA Re mean eterna Masta U4 he ae 68. T. siifolia
me Gaudexerect or Sulberect: stipes tufted: 2275. 20. 225 32. eu tenaneest eects ee ete 52
miGaudex lone-creeping, fronds Seriate ss. eis ahs sere specie a Mlle ne 60
Mminnae commonly 5 cmior more Wide. W/V 22. ee oe atest rates, cee ay)
Minnacnotor little over 2 cmi wide... 3 ee es ne eee 58
. Buds present on upper surface of midrib of apical lamina or at bases of pinnae . 54
mBudsilacking ieither poston ys 044 LA ANS Se ee ee See 23)

. Pinnae to 4 pairs; scales not twisted, few persistent above bases of stipes

69. T. angulata

. Pinnae | pair; scales twisted, to 20 mm long, many persistent above bases of stipes

70. T. croftii

Malina Map ade «2s ory SALA je ed det ame eee ot Aa Ee ae, Beetle Lites itanene 56
MelsINMAcHOrAtIeASt 3: DAMS. 8 ee ccs: ote rn ene a eae ce cee anna ay ee ui omen ay)
. Rachis very dark and glossy; apical lamina 5-lobed ......... 28. T. nebulosa
mivachisotherwise; apical lamina 3-lobed’-.. i 24.5. 4.0 2% 71. T. schmutzii
MBasesvorpinnae narrowly cuneate. 22816. Sine PL eee 72. T. fissa
. Bases of pinnae rounded to subcordate on basiscopic side .. 73. T. polymorpha
RPUIMNACHOND Palms: 4s sos. een APs YAaesie Ut era ho Cn PR 74. T. microlepis
MPinmacel or 2 pairsy. aia ek, ee ee a Tee Pee Aga FD 59

. Buds present on upper surface of costa of apical lamina; small indusia present

75. T. trifida

PeBudSMackines MO MNduUSIA 0k he fee RR Aetna 76. T. rheophytica
. Pinnae commonly to 3 cm wide, their bases narrowly cuneate

77. T. menyanthidis

. Pinnae commonly 6 cm wide, their basiscopic bases rounded

78. T. herpetocaulos

. Pinnae or pinna-like lobes of fertile fronds 3 mm wide, wholly covered beneath by

SPO RAIMA a Mee fs ated eth eh Wh uated an uate wale alee lenge eet Gee Ninchatn ny + Seen AES 62

. Pinnae or pinna-like lobes of fertile fronds much wider; sori distinctly separate . 63
. Rachis-wing not quite continuous; one pair of pinnae present . 79. T. taccifolia
. Rachis-wing quite continuous, decurrent below lowest lobes 101. T. hilocarpa
. At least the sori remote from costae in 1 row on each side of main veins; indusia

COMSPLSWMOUSHE MS He tied tes cheats af ee eG entra Sees a est NTRS a hc 64
. Sori otherwise arranged; indusia in most cases small or lacking............ 67
weAllsonin aregular row oneach side of main ves: ~. 2.50.2... ae 65

. Some additional sori, irregularly shaped and arranged, present between the two

LOW Sane atthe sCOStai m4 4 ee eo 58. T. repanda x 101. T. hilocarpa

64 Flora Malesiana ser. II, Vol. 2! (1991)
65a. Unwinged stipe very short, winged part broad; lobes of lamina well-spaced
80. T. decurrens
b. Unwinged stipe longer, upper part of wing not broad; lobes of lamina not widely
Spacediieae Reales MEE Soe ouidons an whes ou Al 66
66aseerleslobesiof lamina to!20)< 2 emis vio a 80. T. decurrens hybrid
barkeraleslobessofelaminastos 0; q6lcmene sa see eee 81. T. sulitii
6astMany som confluent). 42.05 60h. RSS i. a 68
b. Sormot confluent, 6.6 soe wale oo BORIC. Ree ae Se 69
68a. Sori indusiate, confluent ones a minority .............. 50. T. subcaudata
b. Sori exindusiate, confluent ones many, irregular ........ 53. T. semipinnata
69a. Indusia large waith*stiffshairs\onithemy 4). ease ee ne 82. T. tricuspis
b.indusianotthus hairy \smallor lacking’ 35.2 2 ee eee 70
70a. Sori very numerous, each consisting of few sporangia; lower surface between veins
hairy Athnousho tees ay eee eae ie es Oe i a er 83. T. schultzei
bsiSon larger; lowersurfacemnot hairy... .. 2.05... 6s 55543 3 45 eee Tl
71a. Sori indusiate; indusia small but conspicuous ..............2.000008% 2
b. Sori exindusiate or with occasional rudimentary indusia ................ 3,
72a. Wing continuous throughout rachis and also most of the stipe; no buds
84. T. vasta
b. Wing formed by narrowly decurrent bases of pinnae, not continuous on mature
plants; buds present on rachis near pinna-bases ............. 51. T. fauriei
(Bas Lobesrot trondimarowly;acuminate, 3). era ees 85. T. grandidentata
b. Lobes of frond abruptly short-acuminate .............. 86. T. beccariana
74a. Fronds simple with a broad, often cordate base, entire or variously lobed... . . 1d
b. Fronds simple, in most cases unlobed, narrowed towards base ........... 92
75a. Eaminacedeeplyslobed asi. Sai SA eee 28 See 76
beneaminatentireion crenatese9 ante, ey ME © on os 87
762 eSordistinctlyand persistently indusiate } .2)...0. 4 ee Vi
b. Sonmexindusiatexorindusiaiminute ey 7-5 ae. © co See 83
77a. Lamina hairy on all parts of lower surface of fertile fronds............... 78
b. Laminaof fertile fronds glabrous on lower surface’, . 0... 92 2 eee 82
i Saxe ADICESHOMIODESEODLUSE, Huser he. Fl SIME ees WHY 87. T. novoguineensis
bavApicesiofilobesiacuminate mye hee G ees oe. ah Le 79
(Say Songnearsmarecingonily arate areata ee 55 T. craspedocarpa
b. Sori generally distributed on fronds of mature plants .................. 80
80a. Stipe to 5 cm long; fronds not dimorphous; indusia bearing hairs
88. T. johannis-winkleri
b. Stipes to 20 cm long; fertile fronds contracted indusia not hairy
91. T. palmata var. dimorpha
81a. Lamina much longer than wide, near base lobed 1/3 way to costa; lobes contiguous
andidecrescentidistalliyaanp saya eee eee ar eee 89. T. nitens
b. Lamina little longer than wide, near base lobed more deeply, the lobes widely diver-

genty <a: hd seats BORA Sos ean cle ii. Ae eee 82

Holttum — Tectaria Group 65

. Basal lobes narrowly acuminate, upcurved; sori irregular... . . 90. T. stalactica
. Basal lobes spreading, not narrowly acuminate, sori uniform ... 91. T. palmata
. Fronds trilobed, lateral lobes lacking basal lobules.................... 84
. Fronds 5-lobed, or 3-lobed with basiscopic lobules on laterallobes ........ 86
. Lateral lobes acuminate, similar in shape to apical lamina................ 85
ealeateral lobes very short, abruptly acutes......2..2.....0. 94. T. subtriloba
. Lamina to 19 cm long; scales hair-pointed ............ 92. T. filisquamata
. Lamina to 28 cm long; scales not hair-pointed ........ 93. T. microchlamys

. Sori small, nearly all discrete, rarely coalescent, sometimes with small indusia

91. T. palmata var. platanifolia

. Sori often elongate along veins, exindusiate ......... 95. T. christovalensis
WES OISINGUSIATC! Sor arsverh ee ee hd cls neh en cay Sone Mie amet REY. Bag cles 88
MES OLN KAMIGUSTALS ces anata soeascay fare ve anu d eB dertsm ae Mtavag Sy ee Oy ace. a 89
. Both surfaces covered with short erect hairs ........ 87. T. novoguineensis
. Both surfaces glabrous ...... PE er Mirra OA ECU Re oS, PAS 96. T. minuta
. Surface between veins densely hairy on both sides ......... 97. T. labrusca
mSuUrrace between veins glabrous'on both sides 2... .... 2.5 bee et we cess 90
. Free included veinlets present in areoles; sori mostly punctate .. 98. T. brooksii
. Free veinlets rare in costal areoles; sori mostly elongate ..... 99. T. inopinata
. Fronds strongly dimorphous, fertile ones very narrow and at maturity almost cover-

CMe e Meal mMWACIN SOLAN GLA cs Jos oS oN ehe na cack ahs tro ena ee yee oe O2
~ronds,1f dimorphous, having distinctly separate'sor ......5..es54..4.. 93

. Fronds, both sterile and fertile, unlobed (margins at most sinuous), sterile fronds

glabrous beneath, fertile commonly 8 mm wide ......... 100. T. zollingeri

. Fronds, both sterile and fertile, sometimes deeply lobed; sterile fronds hairy on

veins beneath; simple fertile fronds and the lobes of others commonly 3 mm wide
101. T. hilocarpa

. Sterile lamina to 20 x 1.5 cm (immature plant) ........ 43. T. semibipinnata
= sterile lamina proportionately much wider 2.444204. «2 shi. el 94
. Fronds dimorphous; sori elliptic, 3-4 mm long, indusiate 102. T. kehdingiana
. Fronds not or little dimorphous; sori otherwise
maOnwansedsupe tod cnmlong sw. foc. ase 6 oe ee we es ee ee 96

SRA AR UE SA ESNet rie Ere Sea 95

mWnwinged stipe longer, at least'on fertile fronds... 20... ss ee 98
MBS OMMGITCAUISTATE ec eats, state os acre cae eat Se ek ae acne na te RS, Rese En Marae 97
MES OL CXIMCUSTALC!., teayrarans cai esicue cores Ratio atone aks 86. T. beccariana
MmOOL IN| TOW. On each Side-of main VeInsies san 214 ay sete -ee 80. T. decurrens
ESOL ONG MUM|CTOUMSiae a cle nearer hata aiieea tee eee 103. T. pandurifolia
mCaucdexcreeping. Gmmaturesplants) .. 222 505 -e5 en 63. T. ternata
ME @ALIGEKAETEE Ee ety tee areal) Restore AN SEM cy MPP crate oer Bate EIU Co eet 99

. Margins of fronds entire; lamina of fertile fronds commonly 30 x 6 cm

104. T. singaporeana

. Margins of fronds sinuate-lobate; lamina of fertile fronds c. 18 x 3 cm

105. )orara

66

Flora Malesiana ser. II, Vol. 2! (1991)

27. Tectaria melanocaula (Blume) Copel.,
Philipp. J. Sci. 2 (1907) Bot. 416; Fern FI.
Philipp. (1960) 306; Backer & Posth., Varenfl.
Java (1939) 74; Holttum, Revis. Fl. Malaya 2,
ed. 2 (1968) 636, Gard. Bull. Sing. 34 (1981)
143. — Aspidium melanocaulon Blume, Enum.
Pl. Javae (1828) 161; Hook., Sp. Fil. 4 (1862)
53, excl. syn. Polypodium latifolium Forst.;
Alderw., Malayan Ferns (1908) 245. — Sagenia
melanocaulis (Blume) T. Moore, Index. Fil.
(1858) 96; Racib., Pteridop. Buitenzorg (1898)
195. — Type: Blume s.n., Java (L 908,302-125;
K).

Polypodium nigripes Hassk., Cat. Hort. Bot. Bogor
(1844) 4. — Type: Cult. Hort. Bog. (BO? n.v.).

Aspidium microsorum Presl, Epim. Bot. (1861) 61;
Holttum, Novit. Bot. Delect. Seminum Horti
Bot. Univ. Carol. Prag. 1968 (1969) 88. —
Type: Cuming 57, Luzon (PRC; iso K, L).

Caudex short, suberect; stipe to 60 cm long,
nearly black, at first minutely hairy, glabrescent;
basal scales firm, to 12 x 2 mm; lamina of young
plants ovate, cordate, deeply lobed; of mature plants
to 60 cm long, consisting of apical lamina and
1—3 pairs of pinnae; apical lamina to 25 cm long,
deeply lobed, the lobes acuminate, lowest ones
deeply lobulate, base short-decurrent; upper pinnae
sessile, deeply lobed, lobes acuminate on large
fronds; middle pinnae to 25 x 11 cm, their costules
2 cm or more apart, lobes acuminate, crenate; basal
pinnae to 30 cm or more long, sometimes bearing
1 pair of stalked pinnules, basal basiscopic pinnule
or lobe 15 cm long, shaped as upper pinnae; sparse
short hairs present on lower surface of costae and
veins, a few also between veins; hairs on upper
surface of costae dense, short, between veins only
near the margin; sori small, scattered irregularly,
mainly on short free veins in areoles; indusia small,
glabrous, subreniform; spores minutely spinulose.
— Fig. lle.

Distribution — Peninsular Thailand; in Malesia:
Sumatra, Malay Peninsula, Java, Philippines (Lu-
zon, Negros, Sulu Arch.), Sulawesi, New Guinea.

Habitat — On or among rocks (including lime-
stone) or steeply sloping ground, in forest.

28. Tectaria nebulosa (Baker) C. Chr., Index
Filic. Suppl. 3 (1934) 182. — Nephrodium ne-
bulosum Baker, J. Bot. 18 (1880) 213. — As-
pidium nebulosum (Baker) C. Chr., Index Filic.

(1905) 84; Alderw., Malayan Ferns (1908) 246.
— Type: Beccari 575, Ajer Mantjoer, W Suma-
tra, 360 m (FI; iso K).

Fronds of immature plants similar to those of
T. melanocaula, their stipes nearly black, lamina
5-7-lobed with entire lobes and cordate base; next
stage with 5-lobed terminal lamina and 1 pair of
bilobed pinnae, all lobes entire or nearly so; lami-
na of fronds of mature plants 55 cm long, pinnae
2 pairs, basal pinnae 28 cm long, their basal basi-
scopic lobes to 14 x 4 cm very shallowly bluntly
lobulate, upper pinnae lobed not more than half-
way to their costae; lower surface almost glabrous
on all parts; sori small and very numerous, much
as in T. angulata (Willd.) Copel. but not all on
free veinlets, indusia distinct, about as in T. mela-
nocaula.

Distribution — Malesia: Sumatra (known only
from 3 collections from the Padang Highlands).

Note — The Kew isotype consists of 2 fertile
fronds, each with 5-lobed apical lamina and 1 pair
of forked basal pinnae. C.G. Matthew collected
specimens with sterile 5-lobed lamina in the Anai
Kloof and one frond of mature size (above describ-
ed) on Bukit Siboga near Padang Panjang. The lat-
ter is intermediate between 7. melanocaula and T.
angulata in frond form and in sori; it differs from
T. angulata in its 5-lobed apical lamina lacking
buds and in the lobing of its pinnae.

29. Tectaria holttumii C. Chr., Gard. Bull.
Straits Settlem. 7 (1934) 259, pl. 55. — Type:
Holttum 25570, Sabah, Mt Kinabalu, Meneten-
dok Gorge at 1000 m (BM; BO, K, SING).

Stipe and rachis very dark, at first rather copi-
ously minutely hairy, glabrescent; lamina to 32
cm long, shaped as in T. melanocaula but the api-
cal part cordate at its base and basal pinnae not
pinnate; basal pinnae to 20 cm long, their basal
basiscopic lobes to 10 x 3 cm, lobulate less than
halfway to their costae; lower surface of costae and
veins bearing many slender hairs 0.5—1 mm long
with shorter ones between veins; hairs on upper sur-
face of veins short, hairs 1 mm long scattered be-
tween veins; sori similar in arrangement, size and
indusia to those of T. melanocaula; spores not seen.

Distribution — Malesia: Borneo (Sabah), known
only from the type collection and Clemens 27313
(Dallas) and 295/76 (Tenompok).

Holttum — Tectaria Group

67

Habitat — The type was found on rocks (not lime-
stone) near the river in a deep gorge; no habitat
notes with the Clemens specimens.

30. Tectaria simonsii (Baker) Ching, Sinensia
2 (1931) 32; Holttum, Gard. Bull. Sing. 34
(1981) 144. — Nephrodium simonsii Baker,
Syn. Fil. ed. 2 (1874) 504. — Aspidium si-
monsii (Baker) Bedd., Ferns Brit. India Suppl.
(1876) 15, t. 367; Handb. Ferns Brit. India (1883)
218. — Type: Simons 31, Nuka & Naga Hills,
Assam (K).

Sagenia longicruris Christ, Bull. Acad. Int. Géogr.
Bot. (Mans) 16 (1906) 250. — Type: Cavalerie
268, Kweichow (P).

Aspidium kwarenkoense Hayata, Icon. Pl. Formos.
8 (1918) 138. — Tectaria kwarenkoensis (Haya-
ta) C.Chr., Index Filic. Suppl. 3 (1934) 181;
DeVol, Fl. Taiwan 1 (1975) 344. — Type:
Faurie s.n., Kwarenko, Taiwan (TI, not seen).

Caudex short-creeping to suberect; stipe to 5O
cm long, very dark, glossy above the base, basal
scales narrow, firm, very dark; lamina to 5O cm
long, ‘dark bluish green’ when living, consisting
of a deeply trilobed apex and 1-3 pairs of pinnae;
upper pinnae sessile, subentire, with + dilated and
auricled subcordate base; basal pinnae of large
fronds stalked, to 24 cm long, with 1 pair of free
pinnules like the upper pinnae or with falcate acute
basal auricles, the distal part trilobed; main veins
in pinnae upcurved, cross-veins irregular, small
veins slender, prominent; hairs 0.1 mm long pres-
ent on lower surface of costae, sparse and variable
elsewhere; hairs on upper surface of costae short
and slender, fewer on main veins, a few near mar-
gins; sori all on connected veins, small and uneven-
ly arranged, often + confluent, exindusiate; spores
cristate or with many small irregular wings.

Distribution — NE India to SW China, south-
wards to Burma and Thailand; in Malesia: northern
half of Malay Peninsula (3 collections), Borneo (one
collection from Sabah).

Habitat — The Peninsula specimens were found
on steeply rising ground or on ‘ledges of rock-face’
in forest at 600-750 m, the Sabah specimen on
limestone rock in forest.

Note — The Sabah specimen differs in longer
hairs on lower surface of costae and many short
slender hairs between veins, also in the presence of
small fragile indusia on some sori.

31. Tectaria durvillei (Bory) Holttum, comb.
nov. — Aspidium durvillei Bory in Duperrey,
Voy. Monde 1 (1828) 266. — Type: d’Urville
s.n., Waigeou Is. at Offak (P; iso K).

Lastrea barclayi Carr. in Seem., Fl. Vit. (1873) 359.
— Nephrodium barclayi (Carr.) Baker, Syn. Fil.
ed. 2 (1874) 504. — Tectaria barclayi (Carr.)
C. Chr., Index Filic. Suppl. 3 (1934) 177. —
Type: Barclay s.n., New Ireland (BM).

Aspidium weinlandii Christ, Bull. Herb. Boissier
II, 1 (1902) 453; Alderw., Malayan Ferns (1908)
244. — Tectaria weinlandii (Christ) Copel., J.
Arnold Arbor. 10 (1929) 177. — Type: Wein-
land s.n., Bumi, Finschhafen (P).

Tectaria papuana Copel., Philipp. J. Sci. 6 (1911)
Bot. 76. — Aspidium papuanum (Copel.) Al-
derw., Malayan Ferns Suppl. (1917) 191. —
Type: C. King 160, Papua (UC; iso BO).

Caudex suberect or short-creeping with tufted
stipes; stipe very dark, to 40 cm long, minutely
hairy, basal scales medium brown, not rigid, to
15 x 1.5 mm; /amina of fronds of young plants
ovate-acuminate, then 3—5-lobed with cordate
base, next stage with 3-lobed apical lamina and
1 pair of entire bilobed pinnae bearing also + de-
veloped basal acroscopic auricles; lamina of fronds
of mature plants very firm, to 40 cm long, consist-
ing of a trilobed apex and 3 pairs of pinnae; basal
pinnae to 25 cm long, broadly crenate-lobate with
one or more large basiscopic lobes and a narrower
elongate acroscopic one, the basal lobes separately
adnate on largest fronds; middle pinnae + crenate-
lobate, usually bearing an obtuse to acute acro-
scopic auricle; at least the midlobe of the apical
lamina of the frond crenate-lobate; costae of pinnae
dark at the base on the lower surface, paler distally,
minutely hairy; main veins 6—7 mm apart, pale
and prominent on the lower surface, variably short-
hairy, as also surface between veins; upper surface
minutely hairy on costae, hairs between veins few,
very short, near margin only; sori in one row on
each side of main veins, often with some additional
ones near the costa, on free veins in areoles, in
some specimens (including the type) distinctly
impressed, always marked by prominences on the
upper surface; indusia reniform, persistent, some-
times bearing short superficial hairs; spores cris-
tate. — Fig. 13c.

Distribution — Malesia: widely in New Guinea;
Solomon Islands.

68 Flora Malesiana ser. II, Vol. 2! (1991)

a er =e PJ. Eowaans

Fig. 13. Tectaria brachiata (Zoll. & Moritzi) Morton. a. Fertile and sterile fronds, x 0.22. — T. melano-
rachis (Baker) Copel. b. Sterile frond, x 0.22. — T. durvillei (Bory) Holttum. c. Sterile frond, x 0.22. —
T. keckii (Luerssen) C. Chr. d. Base of a fertile lobe (sori removed on right side), x 4. — T. curtisii
Holttum. e. Lobe of a basal pinna (sori removed), x 6.6 (a: Matthew s.n., 3-1-1912; b: Hose 233; c:
Sambui 9; d: 1962 Unesco Limestone Exp. 551; e: Curtis 3376).

Holttum — Tectaria Group

69

Habitat — In lowland forest, especially near
stream banks; small plants also collected in more
exposed places on limestone.

Note — The many specimens indicate great vari-
ability. The type has rather large sori strongly im-
pressed, and almost glabrous lower surfaces; there
appears to be every gradation between this and
smaller not-impressed sori. The specimens most
hairy on the lower surface are from eastern New
Guinea. The type of 7. papuana has almost entire
pinnae and strongly impressed sori. The type of
Aspidium wendlandii consists of trifoliate fronds
from immature plants.

$2. Tectaria melanorachis (Baker) Copel.,
Sarawak Museum J. 2 (1917) 370. — Nephro-
dium melanorachis Baker, J. Bot. 26 (1888)
325. — Aspidium melanorachis (Baker) C. Chr.,
Index Filic. (1905) 82; Alderw., Malayan Ferns
(1908) 250. — Type: Hose 233, Sarawak (K).

Caudex short, erect; stipe very dark, to 50 cm
or more long, copiously scaly at the base, scales
to 20 mm long, little over 1 mm wide, firm, twist-
ed, smaller narrower scales less abundant distally
and on lower surface of rachis and costae of pinnae;
lamina very similar in size and shape to that of T.
melanocaula, differing as follows: lower surface of
all axes of the frond, and veins, bearing hairs up to
more than 0.5 mm long, similar shorter hairs +
abundant between veins; upper surface also bearing
many short hairs between veins, also very small
spherical glands; sori larger, in a row on each side
of main veins in pinna-lobes (some additional ones
present near costae of pinnae); indusia large, bear-
ing slender hairs like those between veins on the
lower surface; spores densely covered with very
short spines. — Fig. 13b.

Distribution — Malesia: Borneo.

Habitat — Always on limestone.

33. Tectaria danfuensis Holttum, Blumea 35
(1991) 549. — Type: A.C. Jermy 7915, New
Ireland, Danfu Valley, 270 m (BM; iso BO).

Caudex erect; stipe dark, minutely hairy through-
out, the basal 20 cm persistently scaly, the scales
spreading and not ngid, to 20 x 1 mm; basal pin-
nae to 64 cm long, bearing one pair of short-stalked
pinnules; basiscopic pinnule to 30 cm long and 6

cm wide, near its base lobed more than halfway to
the costa, costules of lobes to 2 cm apart, lobes
falcate, acute, entire; second pair of pinnae bearing
1 pair of adnate pinnules; apical part of frond and
of basal pinnae more than 30 cm long, near its
apex lobed to 2 cm from its axis, less deeply to-
wards its base, the lobes almost contiguous and
narrowly acuminate, the basal ones 5—6 cm wide,
lobulate 1/3 towards their costae; lower surface of
all parts bearing copious hairs not more than 0.1
mm long, the smaller ones appearing to be capi-
tate, the larger ones obtuse; upper surface of costae
and costules densely covered with hairs 0.2 mm
long, no hairs between veins except near margin;
sori rather large, in 1 row on each side of main
veins in lobes of pinnae and of apical lamina,
some additional ones also present near costae
of pinnae, mostly on free veins in areoles, not
forming prominences of the upper surface; indusia
thin, rather irregularly reniform with minute mar-
ginal papillae; spores minutely and copiously spin-
ulose.

Distribution — Malesia: New Guinea (New Ire-
land), known from the type and Jermy 7869 and
8006 (cult. R.B.G. Kew).

Habitat — The type from alluvial silt in water-
channels of Danfu Valley, Jermy 8006 from a shady
limestone rock-face.

34. Tectaria athyriosora M.G. Price, Kali-
kasan 3 (1974) 113, f. 1. — Type: Cachalian
15210, Samar, Mt Purog (PNH).

Closely related to T. melanorachis, differing in
firmer texture of fronds which are proportionately
longer with more deeply lobed pinnae; lamina of
largest frond 60 cm long, basal pinnae 25 cm long
bearing 1| pair of stalked and 1 pair of adnate pin-
nules, basal basiscopic pinnule 14 cm long, acro-
scopic 8—9 cm long; two pairs of suprabasal pinnae
each bearing one pair of sessile or slightly adnate
pinnules; all veins rather strongly prominent on
the lower surface; lower surface of pinna-midrib
and main veins sparsely minutely hairy; upper
surface of pinna-midrib densely very short-hairy,
between veins + abundant very short more slender
hairs; sori rather large with large thin indusia, vari-
able in form, the receptacle variably elongate (to
more than 2 mm long); indusia often asymmetric
at the base, glabrous with minutely papillose mar-
gin.

70

Flora Malesiana ser. II, Vol. 2! (1991)

Distribution — Malesia: Philippines (Samar,
probably also Bohol).
Habitat — On limestone.

35. Tectaria brachiata (Zoll. & Moritzi) Mor-
ton, Contr. U.S. Natl. Herb. 38 (1973) 217,
excl. syn. Aspidium variolosum Hook.,; Holt-
tum, Gard. Bull. Sing. 38 (1986) 146. — As-
pidium brachiatum Zoll. & Moritzi, Natuur-
Geneesk. Arch. Ned. Indié 1 (1844) 399; Alderw.,
Malayan Ferns (1908) 252. — Aspidium zol-
lingerianum Kunze, Bot. Zeitung (Berlin) 4
(1846) 462 (not Bedd., Ferns S. India t. 251). —
Lectotype (Morton 1973): Zollinger 655, Java
(G; iso L).

Tectaria variolosa auct.: Backer & Posth., Varenfl.
Java (1939) 76, p.p.; Holttum, Revis. Fl. Ma-
laya 2 (1955) 506, p.p.

Caudex short, suberect, scales to 5 mm long,
medium brown, irregularly fringed with slender hairs
when young; fronds strongly dimorphous. Sterile
frond: stipe to 20 cm long, light castaneous, scaly
near base only, short-hairy distally; lamina to 25
cm long; pinnae to 3 pairs; basal pinnae to 15 cm
long, the largest bearing 1 pair of entire pinnules,
rest of pinna deeply lobed near its base only;
suprabasal pinnae shallowly lobed to subentire
with rounded to subcordate base; apical lamina
deltoid with few broad entire lobes; both surfaces
minutely hairy on costae and main veins, marginal
hairs on upper surface very few and short; veins
amply anastomosing, with many free veinlets,
some branched, in areoles. Fertile frond: stipe to
50 cm long; lamina as sterile but all parts much
contracted; basal pinnae to 10 cm long; sori all on
free veinlets in areoles, impressed, with large thin
entire indusia. — Fig. 13a.

Distribution — Peninsular Thailand; in Malesia:
Malay Peninsula (north only), Java, Bawean, SE
Sulawesi, Tanimbar Is.; N Queensland.

Habitat — The Peninsular specimens were found
in rocky places but not on limestone; the type also
from rocks (unspecified).

36. Tectaria calcarea (Pres!) Copel., Philipp.
J. Sci. 2 (1907) Bot. 415; Fern Fl. Philipp.
(1960) 304. — Aspidium calcareum Presl, Epim.
Bot. (1851) 63; Mett., Farngatt. TV (1858) 120,
t. 18, f. 1-3; Hook., Sp. Fil. 4 (1862) 46;

Alderw., Malayan Ferns (1908) 236; Holttum,
Novit. Bot. Delect. Seminum Horti Bot. Univ.
Carol. Prag. 1968 (1969) 27. — Type: Cuming
310, Leyte (PRC; iso K, L).

Caudex short, suberect, its scales to 5 mm long,
entire; stipe to 30 cm long, medium brown, gla-
brescent abaxially, not glossy; lamina to 35 cm
long, very firm; free pinnae to 8 pairs; basal pin-
nae to 17 cm long (stalks 2.5 cm) bearing 2 pairs
of deeply lobed pinnules, the lobes obtuse, basal
basiscopic pinnule 6 cm long, acroscopic 4 cm,
apical part of pinna acuminate and deeply lobed
like the frond apex; on largest fronds 2 or 3 pairs
of suprabasal pinnae bearing a pair of pinnules;
distal pinnae stalked and deeply lobed; veins slen-
der and distinct on lower surface but not prominent;
costae of pinnae minutely hairy on both surfaces
near bases of pinnae only: marginal hairs on upper
surface few and very small; sori large, in depres-
sions near edges of pinna-lobes, mostly not on free
veinlets; indusia large, persistent, glabrous.

Distribution — Malesia: Philippines (Leyte, Sa-
mar, Bohol).

Habitat — In forest on limestone and limestone-
derived soils (Price & Hernaez 10, W Samar).
Young plants with lamina of frond 6 cm long
(pinnae 2 pairs) may be fertile.

37. Tectaria curtisii Holttum, Gard. Bull. Sing.
38 (1986) 145. — Type: C. Curtis 3376, Dec.
1895, near Ipoh, on limestone (K; iso SING).

KEY TO THE VARIETIES

la. Upper pinnae lobed to S mm from their costae
a. var. curtisii

b Upper pinnae lobed to 10 mm from their cos-
LAC ie A acc samen os b. var. hendersonii

a. var. curtisii

Stipe to 70 cm long, castaneous near base, paler
distally, its basal scales to 10 mm long, very nar-
row, dark and firm; /amina to at least 60 cm long;
pinnae 3 pairs stalked, 1 pair sessile and 2 pairs
adnate; basal pinnae more than 30 cm long (stalk 4
cm), bearing 1 pair of free pinnules and 1 pair ad-
nate; basal basiscopic pinnule 20 cm long, 7 cm
wide above its base, lobed to 5—6 mm from its

Holttum — Tectaria Group

costa, the lobes well spaced, to 4 x 1.2 cm, lobu-
late, acuminate, upper pinnae similarly lobed; veins
forming narrow costal and costular areoles in which
frequent free veinlets are present, additional areoles
also present below sinuses, some with included
veinlets; lower surface of pinna-midrib, costae of
lobes, and veins bearing rather sparse slender hairs
0.2—0.3 mm long, abundant slender erect hairs
0.2 mm long between veins; upper surface of pin-
na-midribs and costae of lobes bearing copious
short hairs, no hairs between veins except a few
short ones near margins; sori mostly on short veins
in areoles or on acroscopic branches of free distal
veins; indusia rather large, thin, bearing a few short
hairs. — Fig. 13e.

Distribution — Malesia: Malay Peninsula (known
only from the type collection).

b. var. hendersonii Holttum, Blumea 35 (1991)
549. — Type: Henderson 22325, Pahang, G.
Senyum (K; iso BM, SING).

All parts of the frond larger than in var. curtisii;
fourth pair of pinnae 22 cm long, lobed to 10 mm
from costae, costules 2.5 cm apart, largest lobes
5.5 x 1.8 cm; lower surface glabrous between veins.

Distribution — Malesia: Malay Peninsula (be-
sides the type collection, one young plant from
Sungei Keteh, Kelantan, probably belongs to this
variety; it has hairs between veins on the lower
surface, as in var. curtisil).

Note — These specimens are comparable in size
and shape of pinnae with some specimens of T.
coadunata from NE India, but the latter differ in the
presence of veinlets in costal areoles and in the al-
most glabrous upper surface.

38. Tectaria cherasica Holttum, Gard. Bull.
Sing. 34 (1981) 141. — Type: A.G. Piggott
2027, Bukit Cheras, Panching, Pahang, on
limestone ridge (K).

Caudex short-creeping; stipe 27 cm long, gla-
brous distally, its base thickened, its basal scales
to 7 x 1.5 mm, thin, light castaneous; lamina
firm, 40 cm long, 20 cm wide; pinnae 4 pairs be-
low an apical lamina 15 cm long which is lobed to
10 mm from its axis, the basal lobes 8 x 2 cm,
lobulate; basal pinnae 18 cm long (stalk 1.5 cm)
bearing one pair of pinnules; basiscopic pinnule 8
cm long, deeply lobed with crenulate lobes, acro-

scopic pinnule 5.5 cm long; next pinna 16 cm
long with a pair of pinnules 4.5 cm long; veins
forming costal areoles which have frequent free
(sometimes forked) included veinlets, some free
included veinlets also in costular and other areoles;
lower surface of pinna-midribs and costules of lobes
bearing slender pale hairs to 0.3 mm long, sparse
shorter hairs on veins; upper surface densely hairy
on pinna-midribs, no hairs present between veins
except a few near sinuses between lobes; sori
mostly on short free veins in areoles, in one row
on each side of costules of smaller pinna-lobes,
more irregular on larger lobes; indusia rather large,
thin, with many slender short hairs on the upper
surface.

Distribution — Malesia: Malay Peninsula (known
only from the type collection).

39. Tectaria translucens Holttum, Gard. Bull.
Sing. 38 (1986) 145. — Type: B.S. Parris &
P.J. Edwards 10450, Bukit Batu Luas in Taman
Negara, Pahang, Malaysia (K).

Caudex short, erect or suberect; stipe light cas-
taneous, sparsely short-hairy, its basal scales dark,
firm, to 15 x 1 mm with filiform apex; lamina to
40 cm long, thin; pinnae to 4 pairs (2 pairs stalk-
ed) below a multilobed apex, its lower lobes deep-
ly lobulate; basal pinnae to 19 cm long (stalk 1.5
cm) bearing 1 pair of stalked pinnules and 1 pair
adnate; basal basiscopic pinnule 12 cm long, deep-
ly lobed and widest at mid-length; second pair of
pinnae bearing 1 pair of pinnules; veins forming
costal areoles which have many free included vein-
lets, rarely forked, free veinlets also present in
some costular areoles and (sometimes forked) in
additional areoles below sinuses; lower surface
rather sparsely and variably short-hairy on pinna-
midribs and costae of lobes (hairs 0.2—0.3 mm
long), short erect very slender hairs variably pres-
ent between veins; upper surface densely hairy on
costae, many short rather slender hairs usually
present between veins with longer ones near sinuses
between lobes; sori on outer veins of costal and
costular areoles or on free veins near apex of lobes;
indusia thin, reniform, variably very short-hairy.

Distribution — Malesia: Malay Peninsula (Pa-
hang, at the type locality and Panching, 20 km
west of Kuantan, southern Kelantan).

Habitat — In limestone rock-crevices and in forest
at the foot of limestone cliffs.

72

Flora Malesiana ser. II, Vol. 2! (1991)

Note — The type was collected in forest at the
foot of limestone cliffs. It has very thin fronds and
few hairs between veins on the upper surface, but
a small plant from the same locality is copiously
hairy in that position. The type has also fairly nu-
merous forked veinlets in areoles. Specimens from
Panching (an isolated limestone hill free of forest)
are firmer in texture, very hairy between veins on
the upper surface and have rare forked veinlets in
areoles.

40. Tectaria keckii (Luerssen) C. Chr., Index
Filic. Suppl. 3 (1934) 181; Holttum, Gard. Bull.
Sing. 34 (1981) 143. — Aspidium keckii Luers-
sen, Bot. Centralbl. 11 (1882) 76; Alderw..,
Malayan Ferns (1908) 248. — Type: Kehding
2817, ‘Klang bei den Gua Batu’ (not seen).

Aspidium amplifolium Alderw., Bull. Jard. Bot.
Buitenzorg II, 11 (1913) 2; Malay. Ferns Suppl.
(1917) 197. — Tectaria amplifolia (Alderw.)
C.Chr., Index Filic. Suppl. 3 (1934) 176;
Holttum, Revis. Fl. Malaya 2 (1955) 515. —
Type: Matthew 509, Perak, Gopeng, G. Mensa
(BO; iso K).

Caudex short, suberect; stipe to 50 cm long, light
brown when dry, when young short-hairy, glabres-
cent; basal scales to 10 mm long, less than 1 mm
wide, rigid, almost black; lamina to 60 cm long
and 50 cm wide, firm, drying dark olivaceous both
sides; pinnae 1—3 pairs, apical lamina broadly del-
toid and deeply lobed, not or little decurrent, the
lowest lobes + deeply lobulate; upper pinnae of
large fronds deeply lobed, lobes broad, entire, ob-
tuse to acute; basal pinnae of largest fronds bearing
1 pair of pinnules, basal basiscopic pinnule or
lobe deeply lobed, basal acroscopic one lobed on
largest fronds, remaining lobes entire, the lower
ones acuminate; lower surface between veins gla-
brous on fronds of mature plants but rather densely
hairy on those of young plants; the hairs between
veins on the upper surface near margins only on
mature plants but + abundant on young plants;
sori in one row on each side of costules of pinna-
lobes with some additional ones near the costae on
large fronds, always on connected veins and large,
with large thin persistent glabrous indusia. — Fig.
13d.

Distribution — Peninsular Thailand; Malesia:
Peninsular Malaysia.

Habitat — On limestone rocks.

Note — The type may have been destroyed at
Kaliningrad. The original description agrees with
the common limestone fern above described and
with no other at Batu Caves.

41. Tectaria villosa Holttum, Blumea 35
(1991) 556. — Type: De Voogd 2245, Sumba
(BO).

In frond-form and venation agreeing with T.
keckii, differing in rather densely hairy lower sur-
face of rachis and costae, thick hairs 0.5 mm long
between veins on upper surface and indusia fringed
with short hairs. The type consists of an apparent-
ly creeping short caudex bearing two fronds which
are not quite fully expanded, the lamina probably
20 cm long when mature; apical section 9 cm long,
deltoid and deeply lobed, pinnae 2 pairs, basal ones
11 cm long (stalks 15 mm), deeply lobed with one
pair of adnate pinnules. On the same caudex are
bases of much larger stipes which would have borne
larger fronds

Distribution — Malesia: Lesser Sunda Islands
(known from the type only).

Habitat — In wooded ravine.

42. Tectaria macrota Holttum, Blumea 35
(1991) 552. — Type: Mousset 617, Java, Mt
Tengger, Gerbo (P; iso K, L).

Caudex suberect or short-creeping, bearing tuft-
ed stipes; stipe to 60 cm long, light castaneous,
minutely hairy in the groove only, basal scales to
10 x 1 mm, firm, dark; frond of young plants 3—5-
lobed, successive ones with trilobed apex and
1 pair of pinnae lobed on the basiscopic side; lamina
of mature plants to 40 cm long, consisting of a
(usually) trilobed apex and 2 or 3 pairs of pinnae;
lobes of frond-apex short-acuminate with a short
obtuse lobe corresponding to each main vein;
suprabasal pinnae commonly 20 cm long, 4-5
cm wide above the base, basiscopic base rounded,
acroscopic base bearing an acuminate subentire
lobe 6-10 x 1.5—2 cm, margins above base of
pinna lobed as those of frond-apex; basal pinnae
of largest fronds 25 cm long (stalk 3 cm) bearing
1 pair of subequal pinnules to 16 x 3.5 cm, the
basiscopic ones sometimes longest; main veins in
pinnae to 10 mm or more apart, cross-veins irregu-
lar; lower surface bearing sparse hairs (some to 0.4

Holttum — Tectaria Group

mm long) on costae and main veins, sparse short
hairs present between veins; upper surface closely
short-hairy on costae, fewer hairs on veins, none
between veins; sori numerous, rather small, irregu-
larly arranged, sometimes + confluent to form a
row parallel to the costa, mostly not on free vein-
lets in areoles; no indusia seen; spores with a
strongly defined cristate network.

Distribution — Malesia: Java (E Java, Tengger
Mts; known from 3 collections by Mousset and
Backer 36619).

Habitat — Forest, 700-800 m.

43. Tectaria semibipinnata (Hook.) Copel.,
Sarawak Museum J. 2 (1917) 371; Holttum,
Revis. Fl. Malaya 2 (1955) 515. — Aspidium
semibipinnatum Hook., Sp. Fil. 4 (1862) 59,
t. 231. — Type: Wallich 388, Penang (K).

Nephrodium nudum Baker, J. Bot. 17 (1879) 41.
— Tectaria nuda (Baker) Copel., Sarawak Mu-
seum J. 2 (1917) 370. — Type: Burbidge s.n.,
Sarawak, Kuching (K).

Tectaria modesta C.Chr., Dansk Bot. Ark. 9, 3
(1937) 72, t. 6. — Type: Sarawak Mus. Coll.
218, Kuching (BM).

Caudex long-creeping with well-spaced fronds,
8 mm diameter (dried), its scales 2 mm wide, less
than 10 mm long; stipe to 90 cm long, pale or
lightly reddish, glabrous with a few scales near the
base; lamina to 80 cm long, pinnate with pinna-
like apex, basal 1 or 2 pairs of pinnae with stalks
2-3 cm long bearing 1 or 2 pairs of pinnules
about the same size as apical lamina; suprabasal
pinnae sessile, to 25 x 2.5 cm, narrowed gradually
to both ends, entire or with slightly sinuous mar-
gins; main veins c. 10 mm apart, upcurved, with
several series of areoles between them; surfaces of
pinnae both glabrous; sori small, scattered irregu-
larly, not at the ends of veins in areoles; indusia
small, reniform, glabrous; slender paraphyses pres-
ent.

Distribution — Malesia: Malay Peninsula, Bor-
neo, N Celebes.

Habitat — On muddy river banks within the tidal
zone but in fresh water, the pinnae floating at high
tides.

Note — The type of T. modesta is a young plant
with simple fronds, lamina to 20 x 1.5 cm. No-
one has reported the conditions under which spores
can germinate to produce young plants.

Ie

44. Tectaria lobbii (Hook.) Copel., Philipp. J.
Sci. 10 (1915) Bot. 146; Fern Fl. Philipp.
(1960) 309. — Aspidium lobbii Hook., Sp.
Fil. 4 (1862) 59, t. 232; Alderw., Malayan
Ferns (1908) 244. — Sagenia lobbii (Hook.)
Christ, Ann. Jard. Bot. Buitenzorg 20 (1905)
100. — Type: T. Lobb s.n., Sarawak, 1857 (K).

Nephrodium hosei Baker, Ann. Bot. (London) 5
(1891) 330. — [Nephrodium stenophyllum
Baker, J. Linn. Soc. Bot. 22 (1886) 227, t. 11,
non Sodiro 1883.] — Aspidium stenophyllum
(Baker) Diels in E. & P., Nat. Pflanzenfam. 1,
4 (1899) 185; Alderw., Malayan Ferns (1908)
247. — Tectaria hosei (Baker) Copel., Sarawak
Museum J. 2 (1917) 371. — Type: G.F. Hose
110, Sarawak, river banks (K).

KEY TO THE VARIETIES

la. Sori not elongate nor confluent........ 2
b Sori regularly elongate and sometimes con-
PUGH Eee cree nero c. var. allosora

2a. Margins of pinnae and pinnules entire
a. var. lobbii
b Margins of pinnae and pinnules minutely den-
fiCUIALe arctan ae b. var. denticulata

a. var. lobbii

Caudex erect or suberect, bearing a tuft of fronds
and tough tangled roots; stipe to 25 cm long, light
castaneous, glabrous, its basal scales narrow, to 10
mm long, not rigid; lamina very firm, to 25 cm
long, consisting of a pinna-like apex and 2-4
pairs of pinnae; rachis densely short-hairy on the
adaxial surface; upper pinnae commonly 12-15 x
1—1.5 cm, entire, narrowed gradually to base and
apex, a bud sometimes present at the base of up-
permost ones; basal pinnae bearing 1 or 2 (rarely
3) pinnules which are alternate, with decurrent
bases, shaped as upper pinnae; main veins oblique,
uniting in a series of arcs near the margin with ir-
regular areoles between them containing branched
free veinlets; surface glabrous apart from short hairs
on upper surface of costae; sori in 2 + regular rows
between main veins, additional ones sometimes
present; indusia large, firm, peltate or subreniform.

Distribution — Malesia: Borneo, Philippines (Bo-
hol).

Habitat — On rocky stream banks in the flood
zone.

74 Flora Malesiana ser. II, Vol. 2! (1991)

Si

S'S

Ze
S
S

PJ.Eowarps

Fig. 14. Tectaria jacobsii Holttum. a. Venation (sori removed on the right), x 2.2; b. minute hairs on ad-
axial surface of rachis, x 23; c. a basal pinna, x 0.26. — T. suluensis Holttum. d. Apex of frond,
x 0.17; e. detail of buds, x 1.3. — T. dolichosora Copel. f. Venation of a middle lobe of apical part (sori
removed on the left), x 2.2. — T. fauriei Tagawa. g. Venation of a middle pinna (sori removed on the
right), x 2.2; h. bud on rachis near base of a pinna, x 4. — T. pleiosora (Alderw.) C. Chr. i. Fertile frond,
x 0.1; j. detail of venation (sori removed on the right), x 2.2 (a—c: Jacobs 5590; d, e: Kondo & Edano
PNH 38624; f: Ramos & Edano BS 28960; g, h: Holttum SF 20071; 1, j: Rick 22).

Holttum — Tectaria Group

dS

b. var. denticulata Holttum, Blumea 35 (1991)
552. — Type: P.S. Ashton S§ 18175, Sarawak,
Bintulu, Ulu Tubau, on frequently flooded river
banks (K).

Distribution — Malesia: Borneo (known only
from the type collection).

c. var. allosora Holttum, Blumea 35 (1991)
552. — Type: D.R. Pleyte 250, Halmahera, G.
Sembilan, thinned-out wood near river (BO; iso
K, L).

Distribution — Malesia: Moluccas (Halmahera,
Morotai).

45. Tectaria jacobsii Holttum, Blumea 35
(1991) 552. — Type: M. Jacobs 5590, Brunei,
junction of rivers Temburong and Belalang at
500 m (K; iso L).

Growth-habit of T. Jobbii but whole plant larger;
stipe to 35 cm long, its basal scales 2.5 mm wide
at their bases; lamina to nearly 40 cm long; pinnae
to 5 pairs, the uppermost (adnate) one with a bud
at its base, the lamina-apex obliquely lobed near
its base; upper pinnae to 14 x 2.5 cm, narrowly
acuminate; basal pinnae to 22 cm long (stalk to
2.5 cm) bearing 2 pairs of pinnules, those on the
acroscopic side entire (to 10 x 1.4 cm), the basal
basiscopic one (14 cm long) irregularly lobate on
its basiscopic side, the basal lobe 2.5 x 0.6 cm,
almost free with an obtuse apex, apical part of pin-
na lobed as frond-apex; surfaces glabrous apart from
minute hairs on adaxial surface of rachis; main
veins of pinnae c. 9 mm apart, at 45°, almost
reaching the margin with 2 or 3 rows of areoles
between them; sori small and numerous, irregu-
larly scattered, some on free veinlets in areoles;
indusia firm, reniform to almost peltate. — Fig.
14a, b.

Distribution — Malesia: Borneo (known only
from the type collection).

Habitat — In river bed below the regular high-
water level, on shaly wet rocks in shade.

Note — The irregular lobing of the frond apex
and of distal parts of pinnae may indicate a hybrid
origin, in which case T. lobbii is indicated as one
parent, but there is no clear evidence of the other
possible parent. Jacobs collected specimens of
typical T. lobbii at a neighbouring locality.

46. Tectaria subdigitata (Baker) Copel., Sara-
wak Museum J. 2 (1917) 370. — Nephrodium
subdigitatum Baker, J. Bot. 24 (1877) 259. —
Aspidium psilopodum C.Chr., Index Filic.
(1905) 89 (not A. subdigitatum Blume 1828);
Alderw., Malayan Ferns (1908) 243. — Sagenia
subdigitata (Baker) Christ, Ann. Jard. Bot. Bui-
tenzorg 20 (1905) 100. — Type: C. Hose 196,
Sarawak, Niah (K; iso BM).

Caudex short, erect, bearing many tough roots;
stipe to 10 cm long, dark brown, glabrous, basal
scales narrow, not rigid; lamina firm, to 12 cm
long, consisting of an apical leaflet and 1 or 2
pairs of pinnae, apical leaflet 8 x 1.4 cm, its base
narrowly decurrent, its apex acuminate, its lower
middle part obliquely lobulate; basal pinnae short-
stalked, bearing one free elongate narrow entire
pinnule and rarely a second one, apical part of
pinna shaped as frond-apex; main veins in pinnae
close together and very oblique, the areoles be-
tween them having few included veinlets; surfaces
glabrous apart from a few minute hairs in the ad-
axial groove of the rachis; sori often in only one
row between main veins, additional ones some-
times present; indusia firm, glabrous.

Distribution — Malesia: Borneo (known only
from the type collection).

- Habitat — No information, but evidently a rheo-
phyte, as T. lobbii.

Note — The apical parts of the frond and of basal
pinnae are lobed in about the same way as those of
T. jacobsit.

47. Tectaria zeilanica (Houtt.) Sledge, Kew
Bull. 27 (1972) 422. — Ophioglossum zeilani-
cum Houtt., Nat. Hist. II, 14 (1783) 43, t. 94,
f. 1. — Leptochilus zeilanicus (Houtt.) C. Chr.,
Index Filic. (1906) 388. — Quercifilix zeilanica
(Houtt.) Copel., Philipp. J. Sci. 37 (1928) 408;
Gen. Fil. (1947) 132; Holttum, Revis. FI.
Malaya 2 (1955) 528, f. 310; Manton, Philos.
Trans. Sér. B, 238 (1954) 137, 161; Price,
Contr. Univ. Michigan Herb. 16 (1987) 199.
— Type: Thunberg s.n., Sri Lanka (UPS, not
seen) 7

Acrostichum quercifolium Retz., Observ. Bot. 6
(1791) 39. — Gymnopteris quercifolia (Retz.)
Bernh., Neues J. Bot. (Schrader) 1, pt 2 (1806)
20; Hook., Icon. Pl. 10 (1854) t. 905; Bedd.,
Ferns Brit. India (1863) pl. 47; Handb. Ferns

76

Flora Malesiana ser. II, Vol. 2! (1991)

Brit. India (1883) 432. — Type: Koenig s.n.,

Sri Lanka.

Caudex slender, creeping; scales thin, narrow,
to 4 mm long; fronds dimorphous. Sterile frond:
stipe 3—8 cm long bearing copious slender hairs;
lamina to 10 cm long, consisting of an apical sec-
tion and 1 pair of pinnae; apical lamina to 7 x 4
cm, + ovate in outline, variably lobed to a depth
of 10 mm, apex rounded, base broad-cuneate; pin-
nae short-stalked, to 3 cm long and 2.5 cm wide at
base which is widened on the basiscopic side with
2 or 3 small lobes; venation normal for sect. Tec-
taria but areoles few; lower surface of costae bear-
ing many slender hairs more than 1 mm long,
sparse minute hairs present on upper surface of
costae and between veins near margin. Fertile
frond: stipe to 18 cm long, almost glabrous; lami-
na to 8 cm long, variously divided, at maximum
with apical section 5.5 cm long, 2.5 mm wide dis-
tally with 2 pairs of narrow lobes to 10 mm long;
pinnae to 3 cm long with a narrow basiscopic lobe
to 8 mm long; sporangia borne densely along the
veins which form a single series of areoles.

Distribution — Sri Lanka, Thailand, N Vietnam;
in Malesia known only from Peninsular Malaysia
(Pulau Tioman) and Borneo (Sabah).

Habitat — On earth banks in shade, or on stream-
sides, sometimes on rocks.

Cytology — n= 80 (Manton 1954).

48. Tectaria lombokensis Holttum, nom. nov.
— Leptochilus siifolius Rosenst., Meded. Rijks-
herb. 14 (1912) 32; Alderw., Malayan Ferns
Suppl. (1917) 534 [not Tectaria siifolia (Willd.)
Copel.]. — Hemigramma siifolia (Rosenst.)
Copel., Philipp. J. Sci. 37 (1928) 407. — Type:
Elbert 2485, SW Lombok, Sepi-Berg N (L
942,64-668).

Caudex not known; fronds dimorphous. Sterile
frond: stipe 12 cm long, minutely hairy, its basal
scales to 5 mm long; /amina 22 cm long, consist-
ing of 3 pairs of pinnae and a pinna-like apex 16 x
3.8 cm; basal pinnae 15 x 3 cm, short-stalked,
with a basal basiscopic lobe 12 x 2.3 cm, other-
wise entire; suprabasal pinnae somewhat shorter,
simple and entire, abruptly narrowed at the base on
the basiscopic side; lower surface of costae minute-
ly hairy near bases of pinnae only, upper surface
similarly hairy throughout. Fertile frond of type:

stipe 43 cm long, stramineous; lamina 28 cm
long, consisting of 4 pairs of pinnae and a pinna-
like apex, all c. 1.5 mm wide; apex 15 cm long,
uppermost pinnae much decurrent on rachis, sec-
ond pair of pinnae 9 cm long and decurrent 5 mm,
third pair slightly decurrent, basal pinnae 18 cm
long (including stalk 5 mm), forked 15 mm above
their bases, basiscopic lobes 13 cm long (basal
pinnae of Elbert 673 13 cm long with 2 lobes 8
cm and 5 cm long on the basiscopic side), all pin-
nae and lobes c. 2 mm wide, covered beneath with
Sporangia on each side of the costa except for a
narrow margin.

Distribution — Malesia: Lesser Sunda Islands
(Lombok, known from the type and Elbert 673,
Rindjani Mts)).

Note — Sections of fertile pinnae of Elbert 673
show a series of vascular strands underlying the nar-
row fertile surface area, indicating a venation as in T.
hilocarpa but closer. Fertile pinnae of the type have
inrolled margins (due to immaturity?) and there-
fore appear narrower, but do not otherwise differ.

49. Tectaria teratocarpa (Alderw.) C.Chr.,
Index Filic. Suppl. 3 (1934) 185. — Aspidium
teratocarpum Alderw., Nova Guinea 14 (1924)
6. — Type: H.J. Lam 966, W New Guinea,
100 m (BO; iso L, SING, UC).

Caudex short, erect; fronds dimorphous. Sterile
frond: stipe to 35 cm long, light castaneous, gla-
brous above base; lamina to 24 cm long, consist-
ing of terminal lamina and 1 or 2 pairs of pinnae;
terminal lamina to 19 x 3.5 cm, entire, apex cau-
date-acuminate, base slightly decurrent; upper pin-
nae sessile, to 12 x 2.8 cm, narrowed at their
bases; basal pinnae short-stalked, with or without
a basiscopic lobe; main veins in apical lamina to
10 mm apart along midrib, cross-veins almost
straight, 2 or 3 series of areoles between them;
sparse short hairs present on lower surface of mid-
rib and veins, upper surface glabrous apart from
very short hairs near bases of pinna-midribs. Fer-
tile frond: stipe a little longer than on sterile fronds;
apical lamina to 16 x 1.9 cm with short lateral
lobes at its base; upper pinnae to 12 x 1.5 cm,
their bases decurrent; basal pinnae to 13.5 cm
long, to 1.8 cm wide above their basal lobes the
largest of which are 5.5 x 0.7 cm (on some speci-
mens the pinnae are narrower); venation much
simplified as compared with sterile fronds; sori

Holttum — Tectaria Group

Wd)

irregularly scattered, some of them confluent along
veins especially near to and parallel with the mar-
gins, and in narrower pinnae; indusia distinct ex-
cept on some confluent sori.

Distribution — Malesia: New Guinea (known
only from the type collection).

50. Tectaria subcaudata (Alderw.) C.Chr.,
Index Filic. Suppl. 3 (1934) 184. — Aspidium
subcaudatum Alderw., Bull. Dép. Agric. Ind.
Néerl. i8 (1908) 9. — Type: Teuscher s.n., W
Borneo (BO).

Caudex short, erect; stipe to 30 cm long, nar-
rowly winged at least distally, red-brown, glabres-
cent; basal scales to 7 x 1 mm with thin margins;
lamina to 30 cm long with 3 pairs of pinnae, all
with narrowly decurrent bases which form an al-
most continuous wing on each side of the rachis,
usually not quite reaching the next lower pinnae;
middle pinnae to 17 x 4 cm, narrowly caudate-
acuminate, entire, with narrowly cuneate base;
basal pinnae sometimes with a narrow basiscopic
lobe to 12 x 2 cm; main veins on pinnae 8-9
mm apart along the costa, upcurved, cross-veins
prominent on the lower surface with 2 rows of
areoles between them; upper surface of midribs
bearing very short hairs, rest of frond glabrous;
sori small, irregularly scattered, many of them
coalescent, mostly not on free veinlets; indusia
very small, thin, persistent but + obscured by de-
veloping sporangia.

Distribution — Malesia: Borneo (W Kalimantan
and Sarawak).

Habitat — In lowland forest.

Note — The type consists of a young plant
bearing small fronds which are trilobed or trifoli-
ate, not fully fertile. The above description is based
mainly on P.W. Richards 2245 from Sarawak, near
Mt Dulit (K).

51. Tectaria fauriei Tagawa, J. Jap. Bot. 14
(1939) 102; DeVol & Kuo, Fl. Taiwan 1 (1975)
342; Holttum, Gard. Bull. Sing. 34 (1981) 146.
— Type: Faurie 67, Taiwan (KYO, not seen).

Tectaria hainanensis Ching & Wang, Acta Phyto-
tax. Sin. 9 (1964) 371. — Type: F.C. How
72939, Hainan (PE; iso UC).

Tectaria gymnosora Holttum, Dansk Bot. Ark. 23
(1966) 308. — Type: Hansen et al. 11225,N
Thailand (C; iso K).

Tectaria gemmifera Ching & Wang, Acta Phytotax.
Sin. 19 (1981) 127, non (Fée) Alston 1939, —
Type: C.H. Wang 80044, Yunnan (PE).

Caudex erect; stipe 40—60 cm long, narrowly
winged in its upper part, basal scales 10 mm or
more long, 2 mm wide at the base, firm, marginal
cells thin-walled; /amina firm, to 50 cm long, con-
sisting of a trilobed apex which is narrowly decur-
rent at its base almost to the attachment of the
uppermost pinnae, and 2 or 3 pairs of pinnae, all
with similarly narrowly decurrent bases; buds pres-
ent on rachis at the base of some pinnae on mature
fronds; middle pinnae 20—25 cm long, entire or
nearly so, gradually narrowed towards their bases,
apex narrowly acuminate, sterile pinnae often
wider than fertile, to 6 cm wide; basal pinnae of
small fronds narrowed towards their base on both
sides, unlobed but the lamina on the basiscopic
side wider than on the acroscopic side, of larger
fronds with a narrowly winged stalk and a rather
long basal basiscopic lobe; pinna-midribs and main
veins reddish on the lower surface when dried and
minutely hairy, upper surface glabrous; smaller
veins concolorous and distinctly prominent both
sides; sori small, round, in about 4 irregular rows
between the main veins, not on free veinlets in
areoles; indusia small, thin glabrous, persistent. —
Fig. 14g, h.

Distribution — Assam, Yunnan, Taiwan, N Thai-
land; in Malesia: Malay Peninsula (Pahang).

Habitat — A rheophyte growing on the rocky
banks of forest streams.

52. Tectaria suluensis Holttum, Blumea 35
(1991) 556. — Type: Y. Kondo & G. Edafo
38624, Sulu Archipelago, Siasi (K; iso L, SING,
US).

Fronds dimorphous. Sterile frond: stipe to 47 cm
long, mid-castaneous, very sparsely hairy, basal
scales to 15 x 2 mm, firm, castaneous with paler
fragile margins, those above base of stipe very
narrow; lamina to 55 cm long, apex pinna-like, to
27 x 8 cm with short decurrent base or a little
united with an uppermost pinna; pinnae to 4 pairs;
uppermost pinnae to 24 x 4.5 cm, narrowed to
cuneate bases with a bud at the base of each, apex
abruptly short-acuminate; basal pinnae unlobed, to
21 x 8 cm, narrowly cuneate at their bases with
stalks 5—7 mm long; main veins 10-15 mm apart

78

Flora Malesiana ser. II, Vol. 2! (1991)

along costae, smaller veins very slender and not
prominent but distinct on lower surface; surfaces
glabrous except for sparse hairs near bases of upper
surface of costae. Fertile frond: lamina to 32 cm
long, its apical section sometimes trilobed; basal
pinnae to 15 x 3.5 cm; sori small, irregularly ar-
ranged, very irregular in shape and sometimes con-
fluent, mostly on free veinlets in areoles, small
indusia sometimes present, hidden by mature spo-
rangia. — Fig. 14d, e.

Distribution — Malesia: Philippines (Sulu Ar-
chipelago and Palawan).

53. Tectaria semipinnata (Roxb.) Morton,
Contr. U.S. Natl. Herb. 38 (1974) 286. —
Acrostichum semipinnatum Roxb., Calcutta J.
Nat. Hist. 4 (1844) 480. — Type: ex herb. Rox-
burgh 2367 (or 23377), ‘Malay Islands’, prob-
ably Penang (BM; iso BR).

Polypodium semipinnatum Roxb., Calcutta J. Nat.
Hist. 4 (1844) 486; Morton, Contr. U.S. Natl.
Herb. 38 (1974) 360. — Type: ex herb. Rox-
burgh, s.n., ‘Malay Islands’, probably Penang
(BR, seen by Morton).

Gymnogramma maingayi Baker, Syn. Fil. ed. 2
(1874) 517. — Selliguea maingayi (Baker) Bedd.,
Ferns Brit. India Suppl. (1876) 24; Handb. Ferns
Brit. India (1883) 392. — Polypodium main-
gayi (Baker) Diels in E. & P., Nat. Pflanzen-
fam. 1, 4 (1899) 318. — Hemionitis maingayi
(Baker) Ridley, J. Malayan Branch Roy. Asiat.
Soc. 4 (1926) 106. — Aspidium maingayi
(Baker) Holttum, Gard. Bull. Straits Settlem. 5
(1934) 207, f. 1-8. — Tectaria maingayi (Baker)
C.Chr., Index Filic. Suppl. 3 (1934) 182;
Holttum, Revis. Fl. Malaya 2 (1955) 509
(f. 302), 513, 626. — Type: Maingay 1809,
Malacca (K).

Polypodium heterosorum Baker, Syn. Fil. ed. 2
(1874) 506. — Dictyopteris heterosora (Baker)
Bedd., Ferns Brit. India Suppl. (1876) 20; Handb.
Ferns Brit. India (1883) 302. — Type: W. Grif-
fith s.n., Malacca, ‘Pulo Bissar’ (K).

Phegopteris subdecurrens Luerssen, Bot. Centralbl.
11 (1892) 30. — Aspidium subdecurrens (Luers-
sen) C.Chr., Index Filic. (1905) 94; Gard. Bull.
Straits Settlem. 4 (1929) 393. — Type: Kehding
2960, Singapore, Pulau Ubin (seen by C.Chr.).

Campylogramma trollii Goebel, Flora 125 (1931)
282, f. 1-4. — Type: W. Troll s.n., Sumatra,
Pulau Berhala, cult. Miinchen (M, photo BM).

Caudex erect; fronds of smaller plants with whol-
ly winged rachis, the stipe also at least partly wing-
ed (width of wing variable), basal pinnae of largest
plants free with slightly decurrent bases or joined
to the rachis-wing (see Holttum 1955: f. 302);
Stipe of largest fronds to 60 cm long, basal scales
c. 10 x 1 mm, the lowest with fragile margins;
lamina to 60 cm long, thin, consisting of a 3—5-
lobed apex and 3 pairs of pinnae or pinna-like lobes
which when nearly free are narrowed towards their
bases; basal pinnae sometimes with a winged stalk-
like base and a large basiscopic lobe; pinnae or
lobes of fertile fronds narrower than those of sterile
ones which may be 7 cm wide; minute hairs pres-
ent on both surfaces of costae only; main veins of
pinnae 8-10 mm apart along the costa, upcurved,
the cross-veins rather regular with many areoles
between them; sori numerous, very irregular in
shape, on free veins in areoles and also along con-
nected veins, often coalescent to linear forms, ex-
indusiate.

Distribution — Peninsular Thailand; Malesia:
Malay Peninsula and N Sumatra.

Habitat — In lowland forest; see Holttum 1931
under Aspidium maingayi.

Cytology — Diploid (Manton in Holttum 1955:
626).

Note — This species appears to be closely allied
to the fern of S China and Tonkin first named Gym-
nopteris decurrens Hook. [Hemigramma decurrens
(Hook.) Copel.]. The latter has smaller, firm,
fronds, the stipe never winged and all sori running
along veins, almost filling the lower surface. The
type of Tectaria simulans Ching [Acta Phytotax.
Sinica 19 (1981) 129] is somewhat intermediate
between T. semipinnata and Gymnopteris decur-
rens, having fertile fronds like the former and a
frond-form like the latter.

54. Tectaria sumatrana (C.Chr.) C.Chr., In-
dex Filic. Suppl. 3 (1934) 185. — [Polypodium
hancockii Baker, Ann. Bot. (London) 8 (1894)
127, non Baker 1885.] — Aspidium sumatra-
num C.Chr., Index Filic. (1905) 94. — Dic-
tyopteris hancockii (Baker) Alderw., Malayan
Ferns Suppl. (1917) 518. — Type: Hancock
89, Sumatra, Barisan Range between Kroe and

Liwa (K).

Caudex and base of stipe unknown; stipe dark
castaneous, glabrescent on abaxial surface; fronds

Holttum — Tectaria Group

q9

dimorphous. Sterile frond of young plant trifoliate,
apical lamina with cuneate base, the pinnae close
to it; lamina of type 38 cm long, consisting of a
rather shallowly trilobed apex with strongly cor-
date base and 1 pair of pinnae 10 cm below it; pin-
nae about 18 cm long (tips broken), shallowly bi-
lobed with a very asymmetric base and stalks 5—7
mm long, strongly cordate on the basiscopic side,
narrower and rounded on the acroscopic side, 8.5
cm wide above the basiscopic lobe; main veins
widely spaced, upcurved, other veins slender, dis-
tinct on the lower surface but not prominent,
areoles many with forked included veinlets; lower
surface glabrous apart from sparse minute hairs on
costae. Fertile frond: only known from 2 detached
stalked pinnae 20 x 6 cm, base broadly rounded or
slightly cordate, apex abruptly short-acuminate;
sori very numerous, mostly on free veinlets in
areoles, also running along other veins, very vari-
able in shape and often confluent; no indusia.
Distribution — Malesia: Sumatra (known only
from the type and a frond of a younger plant sent
later to Kew, probably part of the same collection).

55. Tectaria craspedocarpa Copel., J. Arnold
Arbor. 10 (1929) 178. — Type: Brass 557, Papua
New Guinea, Laloki R., 450 m (A; iso UC).

Stipe to 35 cm long, slender, atropurpureous,
glossy, minutely hairy in the adaxial groove; /a-
mina trifoliate, the apical part tilobed with cuneate
base and acuminate lobes, or 5-lobed (basal lobes
small), to 25 cm long and 22 cm wide; main veins
to 10 mm apart along the costae, at a wide angle, dis-
tally upcurved, cross-veins mostly almost straight
with many areoles between them; short pale erect
hairs present between veins on the lower surface
with somewhat longer ones on costae, upper sur-
face not seen; sori all close to the margin, 1.5 mm
diameter, apparently on connected veins; indusia
almost peltate, persistent.

Distribution — Malesia: Papua New Guinea
(known from the type collection only).

Habitat — In damp soil under rocks near river.

56. Tectaria dolichosora Copel., Philipp. J.
Sci. 38 (1939) 136; Fern Fl. Philipp. (1960)
311; M.G. Price, Brit. Fern Gaz. 10 (1972)
262. — Type: McLean & Catalan 160, Luzon,
Cagayan Prov. (MICH).

Caudex short, suberect; stipe to 60 cm long,
nearly black, glossy, its basal scales to 10 mm
long, less than 1 mm wide, firm, entire; lamina to
50 cm long; apical part to 40 cm long, deeply
multilobed, its basal lobes 25 x 5 cm, acuminate,
obliquely shallowly lobulate; upper lobes entire;
pinnae one pair, to 28 cm long, lobed both sides
to 15 mm from the costa, basal basiscopic lobe to
15 cm long, acuminate, repand to obliquely lobu-
late, other lobes gradually shorter, lobes on the
acroscopic side short and subequal, on largest
fronds acute; main veins prominent on the lower
surface, smaller veins not, areoles with included
veinlets many; lower surface of costae black and
glossy, bearing few minute hairs; upper surface of
costae bearing hairs more than ().5 mm long, scat-
tered hairs also present on and between veins; sori
on free veins in areoles, 3-4 mm long, 1.5—2
mm wide; indusia large, persistent, rounded to cor-
date at their bases, margins minutely papillose. —
Fig. 14f.

Distribution — Malesia: Philippines (Luzon, Ca-
gayan, Tayabas and Quezon Provinces).

Habitat — One specimen from stream-side at
75 m altitude, others from steep mountain slopes
(no altitudes given).

57. Tectaria pleiosora (Alderw.) C.Chr., Gard.
Bull. Sing. 7 (1934) 260. — Aspidium pleio-
sorum Alderw., Malayan Ferns Suppl. (1917)
199; Bull. Jard. Bot. Buitenzorg III, 2 (1920)
131; Nova Guinea 14 (1924) 7. — Aspidium
persoriferum Copel. forma latipinna Alderw.,
Malayan Ferns (1908) 251. — Lectotype
(selected here): Cult. Hort. Bog. IT K, III 7 (BO;
iso L).

Aspidium persoriferum Copel. forma paucipinna
Alderw., Bull. Jard. Bot. Buitenzorg II, 7 (1912)
5. — Type: Gjellerup 788, W New Guinea, Tor
R. (BO).

Caudex erect or suberect; stipe to 50 cm long,
flushed pale to deep red-brown, glabrescent, basal
scales to 15 x 1 mm, their margins fringed when
young; /amina rigid, dimorphous, to c. 40 cm
long with 3 or 4 pairs of pinnae (rarely 7 pairs on
fertile fronds), apex pinna-like, a bud rarely present
at the base of uppermost pinna; basal pinnae usu-
ally unlobed and subsessile; middle sterile pinnae
to 18 x 5 cm, their bases cuneate, wider on basi-
scopic than on acroscopic side, apex acuminate,

80

Flora Malesiana ser. II, Vol. 2! (1991)

margins entire; basal sterile pinnae to 20 x 8 cm,
more asymmetric than upper ones, the basiscopic
margin sometimes sinuous; small veins forming
many areoles between main veins, not prominent
and sometimes not detectable on the lower surface;
costae glabrous on both surfaces; fertile pinnae to
c. 10 x 2.5 cm, sometimes with sinuous margins;
sori in 2 rows between main veins, each at the end
of a single veinlet in an areole (fronds of inter-
mediate type may sometimes occur with more
widely-spaced sori not on single veinlets); indusia
large, broadly elliptic with an elongate receptacle.
— Fig. 14i, j.

Distribution — Malesia: Borneo, N Celebes, New
Guinea, New Ireland.

Habitat — In Borneo found in rocky places, both
sandstone and limestone, also near streams in for-
est; in Sabah the fertile fronds appear to be produced
seasonally but no detailed records exist.

Note — When first describing this species (as A.
persoriferum forma latipinna) Alderwerelt mention-
ed a cultivated plant from Borneo; the specimens
from this are here cited as lectotype. As noted by
Christensen (1934) this species is allied to T. cre-
nata, but I have seen no evidence of hybridization.

58. Tectaria repanda (Willd.) Holttum, comb.
nov. — Aspidium repandum Willd. in Linn.,
Sp. Pl. ed. 4, 5 (1810) 216; Mett., Farngatt. IV
(1858) 126. — Type: Philippines (B, herb.
Willd. 19734).

Polypodium involucratum Roxb., Calcutta J. Nat.
Hist. 4 (1844) 491; Morton, Contr. U.S. Natl.
Herb. 38 (1974) 349. — Type: C. Smith 333,
Ambon (BR).

Aspidium platyphyllum Presl, Epim. Bot. (1851)
65, non Willd. 1810. — Type: Cuming 340,
Cebu (PRC, K).

Aspidium persoriferum Copel. in Perkins, Fragm.
Fl. Philipp. (1905) 177. — Type: Copeland
929, Mindanao (MICH).

Aspidium ternatense Alderw., Bull. Dép. Agric. Ind.
Néerl. 18 (1908) 9. — Type: Teijsmann s.n.,
Ternate (BO).

Tectaria dimorpha St John, Occas. Pap. Bernice P.
Bishop Mus. 21 (1954) 185, f. 4. — Type: St
John 19030, Rotuma Is. (BISH; iso BO, L).

Aspidium nudum auct.: K. Schum. & Lauterb., FI.
Schutzgeb. Siidsee (1900) 16.

Tectaria crenata auct.: Copel., Fern Fl. Philipp.
(1960) 312.

Caudex short, suberect; stipe to 80 cm long,
light castaneous, glabrescent; basal scales 1—1.5
mm wide; /amina to 90 cm long, apex usually
simple or trilobed, pinnae to 8 pairs, their margins
subentire to sinuous, distal pinnae adnate and slight-
ly decurrent; middle pinnae to 30 cm long, narrow-
ly acuminate, sterile ones to at least 4.5 cm wide,
fertile ones commonly 2 cm wide (rarely to 3.5
cm); basal pinnae stalked, with at least one large
basiscopic lobe, in Philippine specimens some-
times several other lobes, decreasing distally; mi-
nute hairs variably present on the lower surface of
costae; sori in one row on each side of main veins,
in narrower fertile pinnae distinctly immersed and
sometimes on single veins in areoles, in broader pin-
nae hardly immersed and not on solitary veinlets; in-
dusia peltate, quite circular on young sori, glabrous
or papillate on upper surface; spores spinulose.

Distribution — Malesia: Java (?), Lesser Sunda
Islands (Flores, Sumbawa), Philippines, NE Cele-
bes, Moluccas, New Guinea; Marianas, Solomons,
New Hebrides, Rotuma Is., probably Fiji.

Cytology — Plants in New Guinea diploid (T.G.
Walker, in litt.).

Taxonomy — Dr. B. Zimmer has kindly exam-
ined Willdenow’s type at Berlin and confirms that
the indusia are peltate.

Hybrids — Especially in the Philippines and New
Guinea there appear to be hybrids between T. re-
panda and T. decurrens; these are variously inter-
mediate between the two parent species.

A specimen from Luzon (Price 2245) is inter-
mediate between 7. repanda and T. hilocarpa. Its
distal sori are like those of T. repanda, but those
on veins nearer the costae are confluent, those
nearest the costae forming continuous lines be-
tween successive main veins as in T. siifolia, dif-
fering from the latter in having conspicuous indusia,
also in the absence of buds. The fronds are trilobed,
the middle lobe being much elongate, the lateral
lobes small (one frond has only one lateral lobe).

A specimen from NE Celebes (de la Saviniére
54) has narrow fertile pinnae with peltate indusia
but broader stipe-scales.

59. Tectaria crenata Cav., Descr. Pl. (1802)
250; C.Chr., Dansk Bot. Ark. 9, 3 (1937) 14;
St John, Occas. Pap. Bernice P. Bishop Mus.
21 (1954) 190, f. 5, 6; Holttum, Revis. FI.
Malaya 2 (1955) 510, 626. — Type: Née s.n.,
Marianas (MA; fragment BM).

Holttum — Tectaria Group

81

Aspidium pachyphyllum Kunze, Bot. Zeitung (Ber-
lin) 6 (1848) 259; Mett., Fil. Hort. Bot. Lips.
(1856) 95, t. 21; Hook., Sp. Fil. 4 (1862) 56,
excl. syn.; Morton, Contr. U.S. Natl. Herb. 38
(1974) 349. — Lectotype (Morton l.c.): Zol-
linger 580, Java (L).

Aspidium grandifolium Presl, Epim. Bot. (1851)
64; Holttum, Novit. Bot. Delect. Seminum
Horti Bot. Univ. Carol. Prag. 1968 (1969) 27.
— Aspidium grande Mett., Farngatt. IV (1858)
121, non Fée 1852. — Tectaria grandifolia
(Presl) Copel., Philipp. J. Sci. 2 (1907) Bot.
413; Fern Fl. Philipp. (1960) 314, name only.
— Type: Cuming s.n., Panay (PRC; probable
isotypes: BM, K).

Cardiochlaena laevis Fée, Mém. Foug. 5. Gen.
Filic. (1852) 316. — Type: Cuming s.n., Phi-
lippines (not seen).

Cardiochlaena sinuosa Fée, Mém. Foug. 5. Gen.
Filic. (1852) 316. — Type: Cuming s.n., Phi-
lippines (not seen).

Aspidium heptaphyllum Baker, J. Bot. 17 (1879)
246. — Tectaria heptaphylla (Baker) C.Chr.,
Index Filic. Suppl. 3 (1934) 180. — Type:
Horne 20, Fiji (K).

Aspidium kawakamii Alderw., Bull. Jard. Bot. Bui-
tenzorg IJ, 7 (1912) 4; Malayan Ferns Suppl.
(1917) 196. — Tectaria kawakamii (Alderw.)
C.Chr., Index Filic. Suppl. 3 (1934) 180. —
Type: Kawakami s.n., S Celebes (BO).

Caudex suberect; stipe to 70 cm long, castane-
ous, its basal scales to 15 mm long, 2 mm or
more wide; /amina to 70 cm long, its apical sec-
tion deeply multilobed, the lower lobes grading to
adnate or sessile pinnae; middle pinnae to 30 cm
long, short-acuminate, of sterile fronds to 8 cm
wide, of fertile fronds 3—5 cm wide (of type 4.5
cm), firm, drying brown-olivaceous, margins cre-
nate to almost entire; basal pinnae distinctly stalk-
ed and bearing a large basiscopic lobe; main veins
in pinnae 7-10 mm apart along the costa, in fertile
fronds with 2 fairly regular rows of areoles between
them; minute hairs variously present on lower sur-
face of costae, upper surface of costae glabrous or
nearly so; sori on free veinlets in areoles, rather
large; indusia firm, slightly reniform, attached to a
linear receptacle 0.5 mm long; spores with irregu-
lar thin fimbriate wings; plants in Malay Peninsula
diploid (Holttum 1955: 626).

Distribution — Malesia: Western Malesia, Philip-

pines, New Guinea; Marianas, Carolines, Solo-
mons, New Hebrides, Fiji.

Habitat — In lowland and mid-mountain forest.

Possible hybrids — This species appears to inter-
grade with T. repanda in the Philippines. One spec-
imen from the Malay Peninsula may represent a
hybrid with T. angulata; it has some irregular
small sori in addition to the two rows normal for
T. crenata, also buds at the bases of some pinnae.

60. Tectaria isomorpha Holttum, Dansk Bot.
Ark. 25, 2 (1967) 49. — Type: Dissing et al.
2369, New Ireland (C; iso K).

Caudex short, suberect; stipe commonly 15—20
cm long (to 30 cm), very dark, hairless, densely
scaly in basal 4 cm, basal scales very dark, less
than 1 mm wide, to c. 12 mm long, upper ones
paler, all with thinner entire margins; lamina to 35
cm long with up to 6 pairs of pinnae, very firm,
apex pinna-like or joined to uppermost pinna; mid-
dle pinnae 10-12 x 2—3 cm, short-stalked, short-
acuminate, margins entire to irregularly repand,
base rounded to broadly cuneate; basal pinnae not
or little longer than the next pair, with a large
acute basiscopic lobe, the lobe on largest fronds
sometimes almost free, the acroscopic base round-
ed or in largest fronds forming an obtuse almost
free lobe 8 x 4 mm; all surfaces almost glabrous,
minute hairs sometimes present on lower surface
of veins and near base of upper surface of costae;
lateral veins of pinnae flexuous, prominent on the
lower surface, smaller veins usually also prominent;
sori on short veinlets in areoles, in one row on
each side of lateral veins, not impressed; indusia
firm, about circular, attached along a line from
centre to base, bearing a few minute hairs.

Distribution — Malesia: New Guinea, Bismarck
Archipelago.

Habitat — On limestone cliffs at altitudes to
1300 m.

Note — The Kew isotype is a rather immature
frond which has a mid-castaneous stipe and rachis
and young sori. All other specimens from both
Western and Eastern New Guinea, have a very dark
stipe and rachis. Young plants have deeply 5-lobed
fronds which may sometimes bear a few sori.

61. Tectaria exauriculata Holttum, Blumea
35 (1991) 550. — Type: M.J.S. Sands et al.
2676, Manus Is. (K).

82

Flora Malesiana ser. II, Vol. 2! (1991)

Caudex short, erect; stipes to 25 cm long (ster-
ile fronds) or 37 cm (fertile), dark, glossy on ab-
axial surface, basal scales very dark, rigid, to 12
mm long, less than 1 mm wide, much contorted;
lamina to 25 cm long, isomorphous, firm, consist-
ing of a trilobed apex and 2 pairs of pinnae; upper
pinnae sessile, to c. 17 x 5 cm, base subcordate on
basiscopic side, cuneate and not auricled on acro-
scopic side, margins subentire, apex not acumi-
nate; lower surface quite glabrous; basal pinnae
with one large basiscopic lobe; main veins c. 15
mm apart along costae, sinuous; sori in one row
on each side of main veins, on terminal veinlets in
areoles, rather large, slightly impressed (with prom-
inences on the upper surface); indusia firm, round-
reniform (not peltate).

Distribution — Malesia: New Guinea (known
only from the type collection).

Habitat — On limestone, found at an altitude of
Sem:

62. Tectaria brevilobata Holttum, Blumea 35
(1991) 548. — Type: C.J. Brooks 16871,
Celebes, Maros Waterfall (BM).

Caudex short, erect; stipes lightly flushed red-
brown, to 26 cm long, glabrous on abaxial sur-
face, scales persistent near base only, to 12 x 1 mm,
rigid, not very dark, distally contorted; lamina to
35 cm long; apical section 19 cm long, deeply
trilobed with a broad non-decurrent base, midlobe
to 4 cm wide, its margin irregularly obliquely
obtusely lobulate, lateral lobes 14 x 2.8 cm with
sinuous margins; pinnae 3 pairs, the bases of all
cordate on the basiscopic side, acroscopic bases not
auricled, rounded on the upper pinnae, subcordate
on basal ones; uppermost pinnae 16 x 3.2 cm,
basal ones 18 x 3.5 cm, acroscopic margins of all
sinuous, basiscopic margins lobed as frond-apex;
basal pinnae bearing a basiscopic lobe to 5.5 x 1.8
cm abruptly short-acuminate; main veins c. 8 mm
apart along costae, cross-veins very irregular; lower
surface bearing sparse minute hairs on costae, up-
per surface of costae densely short-hairy; sori ina
row on each side of main veins, each on a short
vein in an areole, slightly impressed; indusia rather
large, shrivelling, glabrous.

Distribution — Malesia: Celebes (known only
from the type).

Habitat — On limestone.

63. Tectaria ternata (Baker) Copel., Philipp.
J. Sci. 12 (1917) Bot. 58. — Nephrodium ter-
natum Baker, Syn. Fil. (1867) 296. — Dryo-
pteris ternata (Baker) Kuntze, Revis. Gen. Pl. 2
(1891) 813; Alderw., Malayan Ferns (1908) 242.
— Lectotype (selected here): T. Lobb s.n., 1857,
Labuan (K).

Tectaria murudensis Copel., Philipp. J. Sci. 12
(1917) Bot. 58. — Type: Native collector 2905,
Sarawak, foot of Mt Murud (MICH).

Caudex quite prostrate, c. 5 mm diameter; stipe
to 50 cm long, glabrescent, dull reddish, basal
scales to 5 mm long, narrow, not persistent; lami-
na of young plants simple, of mature plants tri-
foliate or with a deeply trilobed apex and one pair
of unlobed pinnae; apical lamina or simple frond
to 28 x 8 cm, margins entire, base narrowly cu-
Neate, apex abruptly caudate-acuminate; pinnae to
20 x 5 cm, sessile, with a bud at the base of each
on old fronds, unlobed, shape as apical lamina but
sometimes wider on the basiscopic side of the
costa than on the acroscopic side; costae sparsely
short-hairy near their bases on lower surface only;
main veins c. 10 mm apart along costae, upcurved,
slender, cross-veins irregular with 3 or 4 series of
areoles between them, smaller veins slender and
distinctly prominent when dried; sori in a rather
uneven row on each side of main veins, often with
many additional ones between the rows, not on
free veinlets; indusia firm, persistent, slightly reni-
form.

Distribution — Malesia: throughout Borneo.

Habitat — In forest to about 300 m, often on
rocks and sometimes beside streams, also on steep-
ly sloping ground.

64. Tectaria tabonensis M..G. Price, Kalika-
san 3 (1974) 118, f. 3. — Type: Raynoso PNH
37858, Palawan, Lipuun Is. (PNH; iso L).

Caudex erect or suberect; stipe to 30 cm long,
base very dark, above base castaneous, minutely
hairy, glabrescent; basal scales to 10 x 1.5 mm,
firm with thinner margins; lamina of young plants
multilobed, of mature plants to 45 cm long, tex-
ture very firm, consisting of a 5-lobed broadly tri-
angular apex and 3 or 4 pairs of subopposite pin-
nae; pinnae gradually larger towards the base of the
frond, short-stalked, with cordate basiscopic and
rounded to cuneate acroscopic base, margins repand-

Holttum — Tectaria Group

sinuate, gradually narrowed to an acuminate apex;
basal pinnae 18-25 cm long (stalks 15 mm), 2
cm wide on acroscopic side of costa, on basiscopic
side a basal lobe 11 cm long and another 9 cm,
both acuminate with repand margins, apical part of
pinnae irregularly obliquely bluntly lobed on the
basiscopic side; main veins to 10 mm apart along
the costae, upcurved, prominent on the lower sur-
face, cross-veins irregular with many areoles be-
tween them, smaller veins distinct but not promi-
nent; lower surface of costae sparsely short-hairy,
upper surface of rachis and costae densely covered
with thicker short hairs, rest of the surface gla-
brous; sori rather large, in a rather uneven row on
each side of main veins with a few additional ones,
not impressed, mostly not on free veinlets; indusia
large, thin and shrivelling, slightly reniform, mi-
nutely hairy, glabrescent.

Distribution — Malesia: Philippines (Palawan,
also eastern Mindanao.

Habitat — In forest on limestone rocks at low
altitudes.

65. Tectaria subcordata Holttum, Blumea 35
(1991) 555. — Type: J.R. Croft LAE 65567,
New Ireiand, N Scheinitz Range, 300 m (LAE;
iso L).

Stipe 65 cm long, nearly black, glossy, rachis
similar; lamina c. 50 cm long, firm but not rigid,
consisting of a trilobed apex and 3 pairs of pinnae;
apical lamina 22 cm long, its midlobe 7.5 cm wide
with obliquely shallowly lobed margins, lateral
lobes c. 17 x 5 cm; middle pinnae 26 cm long
(stalk 5—7 mm), their acroscopic bases cuneate at
45° and then rounded, basiscopic bases truncate to
subcordate, 15 mm wide, then gradually wider,
margins almost entire; basal pinnae 25 cm long
(stalk 12 mm), bilobed, basiscopic lobe 19 cm
long, abruptly short-acuminate, 5.3 cm wide, its
basiscopic margin undulate-subcrenate in its widest
part; main veins to 12 mm apart along costae,

- sinuous, upcurved, cross-veins irregular, smaller
veins concolorous, distinct on both surfaces; lower
surface wholly glabrous, upper surface minutely
hairy on costae only; sori in a row on each side of
main veins (no additional ones present), mostly
not on free veins in areoles, somewhat impressed;
indusia apparently peltate (all old and distorted on
type).

83

Distribution — New Ireland; known only from
the type collection.
Habitat — On limestone in lowland hill forest.

66. Tectaria decastroi (Alderw.) C. Chr., In-
dex Filic. Suppl. 3 (1934) 178. — Aspidium
decastroi Alderw., Bull. Jard. Bot. Buitenzorg
II, 7 (1912) 3; Malayan Ferns Suppl. (1917)
197. — Type: De Castro 64, Timor Dilli (BO;
iso L).

Caudex short-creeping, thick; stipe to 55 cm
long, castaneous, minutely hairy, glabrescent; /a-
mina to 45 cm long, thin, glabrous, consisting of
a deeply trilobed apex and 4 pairs of pinnae; upper-
most pinnae sessile, to 13 x 3.5 cm, base rather
narrowly cuneate on the acroscopic side, more broad-
ly on basiscopic side, margins subentire; second
pair of pinnae similar but larger with + sinuous
margins; third pair much wider on basiscopic than
on acroscopic side of costa, basiscopic margin irreg-
ularly shallowly lobed; basal pinnae 22 cm long
(stalks 10 mm) with 2 large acuminate lobes on
basiscopic side, basiscopic margin obliquely obtuse-
ly lobed to a depth of 3-4 mm; main veins up-
curved, not sinuous, with irregular areoles between
them; sori in a regular row on each side of main
veins with a few additional ones, slightly impres-
sed, mostly not on free veins in areoles; indusia
minute, apparently reniform and soon caducous.

Distribution — Malesia: Lesser Sunda Islands
(besides the type from Timor, a frond of a smaller
plant from Flores probably belongs to this species).

67. Tectaria rufescens Holttum, Blumea 35
(1991) 555. — Type: Soegeng Reksodihardjo
408, SE Irian Jaya (BO; iso L).

Caudex short, erect, bearing a tuft of many
stipes; stipe c. 20 cm long, dull castaneous, minute-
ly hairy on the abaxial surface; basal scales to 8 mm
long, less than 1 mm wide, dark with paler margins;
lamina c. 20 cm long, apex very deeply trilobed or
simple with a pair of pinnae close below it; pinnae
2 or 3 pairs, upper ones 10 x 2 cm, almost sessile,
entire, base rather narrowly cuneate, apex acumi-
nate; basal pinnae to 12 x 2 cm with a basiscopic
lobe 8 x 1.5 cm; main veins in pinnae 5 mm apart
along the costa, at 45°, red-brown and prominent
on the lower surface, cross-veins similar with 2
irregular series of areoles between them; lower

84 Flora Malesiana ser. II, Vol. 2! (1991)

PJ. Eowarps

Fig. 15. Tectaria siifolia (Willd.) Copel. a. Venation of a middle pinna (sori removed on the right, to show
attachment), x 2.2. — T. rheophytica Holttum. b. Fertile frond, x 0.33; c. detail of b, x 4. — T. schultzei
(Brause) C. Chr. d. Middle of a lateral lobe, x 4. — T. tricuspis (Bedd.) Copel. e. Middle of a lateral lobe
(venation details only shown in centre), x 2.2; f. single sorus showing indusium with stiff hairs, x 23
(a: Teijsmann s.n., Sumatra; b, c: Croft 1681; d: Croxall & Parris 5963; e, f: Iwatsuki et al. B-762).

surface of costae and veins bearing sparse minute 68. Tectaria siifolia (Willd.) Copel., Philipp. J.

hairs, upper surface of costae closely covered with Sci. 2 (1907) Bot. 414; Backer & Posth., Varen-
short hairs, no other hairs present; sori rather irreg- fl. Java (1939) 75; Copel., Fern Fl. Philipp.
ularly spaced, often confluent or elongate to form (1960) 314. — Polypodium siifolium Willd. in
lines parallel to the main veins (not to costae); in- Linn., Sp. Pl. ed. 4, 5 (1810) 196. — Aspidium
dusia very small and thin, not covering sori. slifolium (Willd.) Mett., Ann. Mus. Bot. Lugd.-

Distribution — Malesia: Irian Jaya (apart from Bat. 1 (1864) 237; Alderw., Malayan Ferns
the type, one collection from lowland forest in (1908) 251. — Type: Ventenat s.n., Java (B,

NW Irian Jaya, Sorong Dist.). herb. Willd. 19689).

Holttum — Tectaria Group

Dryomenis phymatodes Fée, Mém. Foug. 5. Gen.
Filic. (1852) 225, pl. 18A, f. 1. — Dryomenis
menisciicarpon T. Moore, Index Filic. (1857)
xlvii, nom. nov. — Aspidium menisclicarpon
(T. Moore) Mett., Farngatt. [V (1858) 121. —
Type: Cuming 4, Luzon (isotypes BM, K, L).

Aspidium teysmannianum Hook., Sp. Fil. 4 (1862)
41, t. 236. — Aspidium siifolium var. teysman-
nianum Christ, Ann. Jard. Bot. Buitenzorg 15
(1898) 138. — Type: Teijsmann s.n., Sumatra,
Pulu Pisang (K).

Aspidium biseriatum Christ, Bull. Herb. Boissier
II, 6 (1906) 1002. — Type: Loher s.n., Feb.
1906, Luzon, Zambales (not seen).

Aspidium ternifolium Alderw., Bull. Jard. Bot. Bui-
tenzorg II, 11 (1913) 3; Malayan Ferns Suppl.
(1917) 194. — Tectaria ternifolia (Alderw.)
C.Chr., Index Filic. Suppl. 3 (1934) 185;
Holttum, Revis. Fl. Malaya 2 (1955) 516, f.
303. — Type: Matthew 510, Perak, Gopeng
(BO; iso K).

Aspidium papyraceum Alderw., Bull. Jard. Bot. Bui-
tenzorg III, 2 (1920) 131. — Tectaria papyracea
(Alderw.) C.Chr., Index Filic. Suppl. 3 (1934)
183. — Type: Brooks 377/S, Sumatra, Benkoe-
len (BO; iso BM).

Caudex short-creeping; fronds dimorphous, ster-
ile ones with shorter stipe and broader pinnae than
fertile, pinnae to 4 pairs; fronds of young plants
ovate-acute with strongly cordate base. Sterile
fronds: stipe to 30 cm long, glabrous, basal scales
to 10 mm long, dark with thinner pale margins;
apical lamina often trilobed, the middle lobe some-
times with sinuous margins; buds present at bases
of pinnae on old fronds; basal pinnae to 20 cm
long, to 7 cm wide above the basal lobe, abruptly
short-acuminate; middle pinnae broader on basi-
scopic than on acroscopic side of the costa; main
veins to 10 mm apart along costae; upper surface of
costae glabrous except near their base in Sumatra,
closely short-hairy in Celebes and Philippines, low-
er surfaces glabrous. Fertile fronds: stipe to 50 cm
long; middle pinnae to c. 12 x 4 cm with main veins
to 6 mm apart along costae, areoles between main
veins fewer than in sterile pinnae and in smaller pin-
nae lacking included veinlets; sori on the outer veins
of areoles, in a row on each side of main veins,
usually rather large, those near the costae elongate
and often coalescing to form lines parallel to the
costa; indusia thin, soon caducous. — Fig. 15a.

85

Distribution — Malesia: throughout, but local in
occurrence.

Habitat — In Sumatra and the Malay Peninsula
associated with limestone and seasonal in produc-
ing fertile fronds; no good records elsewhere.

69. Tectaria angulata (Willd.) Copel., Sara-
wak Museum J. 2 (1917) 370; Backer & Posth.,
Varenfl. Java (1939) 74; Holttum, Revis. Fl.
Malaya 2 (1955) 511, f. 301, excl. syn. Nephro-
dium nebulosum Baker. — Polypodium angu-
latum Willd. in Linn., Sp. Pl. ed. 4, 5 (1810)
239. — Aspidium angulatum (Willd.) Mett.,
Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 239;
Alderw., Malayan Ferns (1908) 236. — Type:
Ventenat s.n., Java (B, herb. Willd. 19641).

Aspidium sanctum Blume, Enum. Pl. Javae (1828)
143. — Type: Blume s.n., Java (L 908,300-
118).

Dictyopteris pentaphylla Alderw., Bull. Dép. Agric.
Ind. Néerl. 18 (1908) 16; Malayan Ferns (1908)
519. — Aspidium quinquefoliatum C.Chr., In-
dex Filic. Suppl. 1 (1913) 9 (not A. pentaphyl-
lum Willd.). — Tectaria pentaphylla (Alderw.)
C.Chr., Index Filic. Suppl. 3 (1934) 183. —
Type: Treub 232, New Guinea, Skroe (BO; iso
Ker)

Aspidium trifolium Alderw., Bull. Jard. Bot. Bui-
tenzorg 7 (1912) 4; Malayan Ferns Suppl. (1917)
193, excl. var. compitale. — Tectaria trifolia
(Alderw.) C.Chr., Bot. Jahrb. Syst. 66 (1933)
49; Copel., Fern Fl. Philipp. (1960) 307, p.p.
— Type: Elmer 8329, Luzon (BO; iso K, L).

Aspidium terminale Rosenst., Meded. Rijksherb.
31 (1917) 4; C.Chr., Gard. Bull. Straits Settlem.
4 (1929) 393. — Tectaria terminalis (Rosenst.)
C.Chr., Index Filic. Suppl. 3 (1934) 185. —
Type: Hallier 1821, Borneo (L).

Aspidium falcipinnum Alderw., Bull. Jard. Bot.
Buitenzorg II, 28 (1918) 7. — Tectaria falci-
pinna (Alderw.) C.Chr., Index Filic. Suppl. 3
(1934) 179. — Type: Brooks 374, Sumatra,
Lebong Tandai (BO; iso BM).

Caudex short, erect, sometimes branching near
its base; stipe to 60 cm long, glabrous and glossy
above its base, usually dark, basal scales to 15 x
1.5 mn, stiff, not very dark, fringed when young;
lamina to 60 cm long consisting of a rather shal-
lowly trilobed apex and 1—4 pairs of pinnae, thin,
drying brownish; apical lamina to 25 cm long, its

86

Flora Malesiana ser. II, Vol. 2! (1991)

base not decurrent, 1 or more buds present on the
upper surface of its costa near bases of main veins;
pinnae more broadly rounded at their bases on the
basiscopic than acroscopic side, middle ones com-
monly 15-18 x 5—6 cm, basal pinnae with stalks
to 2 cm long and a broad basiscopic lobe; main
veins 10-15 mm apart along costae of pinnae, at
45°, little upcurved, cross-veins not regular, smaller
veins distinct but not prominent, forming areoles
containing copious branched free veinlets; lower
surface glabrous, upper surface of costae densely
short-hairy; sori small, mostly at ends of veinlets
in areoles, nearly round but on some fronds vari-
ously a little elongate; indusia small, reniform,
sometimes lacking.

Distribution — Peninsular Thailand; throughout
Malesia; Solomon Islands.

Habitat — In lowland forest near streams, usually
on rocks but not in the flood zone of swiftly flow-
ing streams.

Note — There is considerable variation in the de-
velopment of indusia and in the abundance of + elon-
gate sori. Indusia are best developed on specimens
from Perak in the Malay Peninsula. Some speci-
mens from Sarawak and Celebes have buds at the
bases of pinnae in addition to those on the costa of
the apical lamina. Buds may be proliferous on plants
in very humid positions (e.g. spray of waterfalls).
One plant from ‘alluvial forest’ in the G. Mulu
National Park, Sarawak, has pinnae with very broad
bases, subcordate on the basiscopic side. Under T.
trifolia, Copeland (1.c. 1960) appears to have in-
cluded specimens of T. polymorpha.

70. Tectaria croftii Holttum, Blumea 35 (1991)
548. — Type: J.R. Croft 1734, Papua New
Guinea, W Sepik, N slopes of Bewani Mts, 300
m, on vertical limestone face (K; iso LAE, L).

Caudex short, erect; stipe to 40 cm long, dull
castaneous, minutely hairy throughout; basal scales
15-20 mm long, abundant, firm and + twisted,
distal scales and those on rachis and lower surface
of costae all much shorter and very narrow; lamina
to 34 cm long; apical section trilobed, 27 cm long,
base short-decurrent, midlobe 11 cm wide, entire,
short-acuminate, narrowed at base to 8 cm, lateral
lobes upcurved, 15 cm long from bases of their
costae, 5.5 cm wide; pinnae 15 cm long with stalks
3 mm long and abrupt acute apices, each with one
short abruptly acute basiscopic lobe; main veins in

pinnae upcurved, 10-15 mm apart along costae,
cross-veins irregular, small veins forming many
areoles between them with fewer free veinlets than
in T. angulata; lower surface of costae and veins
copiously hairy, slender hairs between veins; upper
surface of pinna-midrib densely covered with thick
hairs 0.3 mm long, fewer similar hairs on main and
smaller veins, a few thick hairs 0.5 mm long be-
tween veins; sori small, scattered irregularly, ex-
indusiate, mostly not on free veinlets.

Distribution — Malesia: Northern Papua New
Guinea (W Sepik), Manus Is.

Habitat — The type and Sands 2629 from Manus
Is. were collected from limestone. A plant cultivated
at Kew, brought from near Lae in 1969, is similar
but did not originate from limestone.

71. Tectaria schmutzii Holttum, Blumea 35
(1991) 555. — Type: E. Schmutz 6977, Flores,
Nengke R. (K).

Aspidium trifolium var. compitale Alderw., Bull.
Jard. Bot. Buitenzorg II, 20 (1915) 6; Malayan
Ferns Duppl. (1917) 193. — Type: Saanam 44,
Obi Is.,; P. Bisa (BO).

Caudex short, erect; stipe to 30 cm long, distally
light-castaneous, glabrous and glossy on abaxial
surface, basal scales 7 mm long, no scales above
base; lamina firm, to 40 cm long, consisting of a
+ deeply trilobed apical section and one or two
pairs of pinnae; upper pinnae short-stalked, to 14 x
5 cm, apex narrowly acuminate, base broadly cor-
date on basiscopic side; basal pinnae to 15 cm long,
each with a large basiscopic lobe, stalks to 5 mm
long; main veins at 45° to costae and 10-12 mm
apart along it, cross-veins and areoles very irregu-
lar, smaller veins slender but distinct; surfaces, in-
cluding costae, quite glabrous; sori many, small,
very irregularly scattered, sometimes confluent,
some on free veinlets in areoles but many on con-
nected veins, no indusia seen.

Distribution — Malesia: Lesser Sunda Islands
(Flores, Obi Is.).

Habitat — On limestone rocks on shaded stream-
banks, just above normal water level, sometimes
flooded, altitude 100—200 m (information from
Dr. Schmutz).

Note — The type of Aspidium trifolium var. com-
pitale is a young plant. The fronds consist of a
simple or slightly trilobed lamina shaped as in im-
mature plants of 7. schmutzii; there are few sori.

Holttum — Tectaria Group

87

72. Tectaria fissa (Kunze) Holttum, comb. nov.
— Aspidium fissum Kunze, Bot. Zeitung (Ber-
lin) 6 (1848) 258. — Lectotype (selected here):
Zollinger 2369, Java (L 908,302-348 & 349;
iso BM).

Aspidium oligophyllum Rosenst., Feddes Repert.
Spec. Nov. Regni Veg. 5 (1908) 13; Alderw.,
Malayan Ferns (1908) 237. — Tectaria oligo-
phylla (Rosenst.) C.Chr., Index Filic. Suppl.
3 (1934) 183; Holttum, Gard. Bull. Sing. 34
(1981) 145. — Type: Burchard s.n., Sumatra,
Indragiri (L).

Aspidium burchardii Rosenst., Meded. Rijksherb.
31 (1917) 3. — Tectaria burchardii (Rosenst.)
C.Chr., Index Filic. Suppl. 3 (1934) 177. —
Type: Burchard s.n., Sumatra (L).

Aspidium divergens Rosenst., Meded. Rijksherb. 31
(1917) 3. — Type: Cult. Hort. Bog. (BO).

Aspidium tricuspe Bedd. var. glabrum Alderw.,
Bull. Jard. Bot. Buitenzorg III, 5 (1922) 184.
— Type: Biinnemeijer 5834, Riouw Arch., P.
Toedjoeh, Mt Rani, 400 m (BO).

Tectaria polymorpha (Wall. ex Hook.) Copel. var.
cuneifolia Bonap., Notes Pterid. 14 (1923) 50;
Holttum, Revis. Fl. Malaya 2 (1955) 518, f. 304.
— Tectaria cuneifolia (Bonap.) Love & Love,
Taxon 26 (1977) 326. — Type: Holttum 9566,
Negri Sembilan, G. Tampin (P; iso SING).

Leptochilus rumicifolius Ridley, J. Malay. Branch
Roy. Asiat. Soc. 4 (1926) 116. — Tectaria ru-
micifolia (Ridley) C.Chr., Index Filic. Suppl.
3 (1934) 184; Holttum, Revis. Fl. Malaya 2
(1955) 519; ibid. ed. 2 (1968) 636; Dansk Bot.
Ark. 23, 2 (1965) 241. — Type: Ridley s.n.,
Selangor, Klang Gates (K).

Caudex short, suberect; stipe to 70 cm long,
mid-castaneous, glabrescent, basal scales to 15 x 2
mm, firm, glossy, twisted distally, fringed when
young; /amina to 100 cm long consisting of 1-5
pairs of pinnae and pinna-like apex which may be
joined to the uppermost pair of pinnae; pinnae
to 25 x 6 cm, entire with narrowly cuneate base
and caudate-acuminate apex; basal pinnae (some-
times 2 pairs) with stalk 2—3 cm long and a basi-
scopic lobe which may be more than 20 cm long;
main veins 6-10 mm apart along costae and
at a broad angle to them, distally upcurved; cross-
veins rather regular, between them 2 irregular rows
of areoles with included veinlets; lower surface of
costa and main veins variably minutely hairy,

upper surface usually quite glabrous; sori mostly
on veins surrounding areoles, in a row on each side
of a cross-vein, usually 4—6 in a row in the mid-
dle of a large pinna; indusia reniform, firm, per-
sistent.

Distribution — Malesia: Sumatra, Malay Penin-
sula, Java, Borneo.

Habitat — In lowland forest, occasionally up to
1200 m.

73. Tectaria polymorpha (Wall. ex Hook.)
Copel., Philipp. J. Sci. 2 (1907) Bot. 413;
Ching, Sinensia 2 (1931) 30; Backer & Posth.,
Varenfl. Java (1939) 78, p.p.; Holttum, Dansk
Bot. Ark. 23, 2 (1965) 240; Sledge, Kew Bull.
27 (1972) 419. — Aspidium polymorphum
Wall. ex Hook., Sp. Fil. 4 (1862) 54, excl. syn.
A. repandum Willd.; Bedd., Ferns S. India (1865)
pl. 116, Handb. Ferns Brit. India (1883) 218,
all synonyms excluded. — Type: Wallich 382,
Nepal (K; iso BM).

Nephrodium subpedatum Harr., J. Linn. Soc. Bot.
16 (1877) 30. — Tectaria subpedata (Harr.)
Ching, Sinensia 2 (1931) 23, excl. syn. Nephro-
dium morsei Baker. — Type: Steere s.n., Tai-
wan (MICH; iso K).

Caudex suberect; stipe to 45 cm long, dull red-
dish, minutely hairy, basal scales to 10 x 1.5 mm,
dark with a thinner + fringed margin; lamina to 50
cm long consisting of a pinna-like apex (sometimes
fused with the uppermost pinnae) and 1—4 pairs of
opposite pinnae; middle pinnae to 25 x 5 cm, short-
stalked, rounded to subcordate on their basiscopic
bases, more narrowly rounded on the acroscopic
side, apex abruptly short-acuminate, margins en-
tire; basal pinnae with stalks 15 mm long, bearing
a large basiscopic lobe; main veins 6-8 mm apart
along costae, at a broad angle and upcurved, cross-
veins rather regular between them; lower surface of
costae and main veins short-hairy, short erect hairs
also present between veins, upper surface of costae
glabrous or sparsely hairy; sori small, at maximum
development 4 or 5 in a row on each side of cross-
veins, mainly on veins which surround areoles;
indusia thin, rather small,. shrivelling; n = 80 in
Sn Lanka and S India.

Distribution — Nepal to Yunnan and to N Thai-
land and Tonkin, Taiwan, S India and Sri Lanka;
in Malesia: E Java, Lesser Sunda Islands (Timor),
Philippines (Luzon), and Celebes.

88

Flora Malesiana ser. II, Vol. 2! (1991)

74. Tectaria microlepis Holttum, Blumea 35
(1991) 553. — Type: J.H. Coert 657, E Java (L).

Caudex short, erect, with long tough roots; stipe
33 cm long, glabrous, basal scales not seen; lamina
33 cm long, firm, its apical section 19 x 1.8 cm
joined at its base with the uppermost pinnae; pin-
nae all with narrowly cuneate bases and acuminate
apices, upper ones 12.5 x 1 cm, third pair 16.2 x
1.4 cm, fourth pair 18 x 1.6 cm, basal pair 18 cm
long (stalk 7 mm), 1.8 cm wide above its base with
a basiscopic lobe 12 x 0.8 cm; veins about as in
T. lobbii; both surfaces glabrous apart from sparse
short hairs on lower surface of main veins; sori
irregular in size and arrangement, indusia very
small, reniform.

Distribution — Malesia: E Java, known only from
the type.

Habitat — River bank; evidently a rheophyte, as
T. lobbii.

75. Tectaria trifida (Fée) M.G. Price, Amer.
Fern J. 64 (1974) 31. — Microsorium trifidum
Fée, Mém. Foug. 5. Gen. Filic. (1852) 269. —
Type: Cuming 31, Luzon (isotypes BM, K).

Aspidium irriguum Presl, Epim. Bot. (1851) 62,
non Smith 1810; Holttum, Novit. Bot. Delect.
Seminum Horti Bot. Univ. Carol. Prag. 1968
(1969) 28; Alderw., Malayan Ferns (1908) 245.
— Tectaria irrigua (Presl) Copel., Philipp. J.
Sci. 2 (1907) Bot. 413; Fern FI. Philipp. (1960)
308. — Type: Cuming 31, Luzon (PRC).

Caudex short, erect, bearing a close tuft of stipes
and many strong roots; stipe to 20 cm long, +
flushed with red, sometimes very dark, its basal
scales flaccid, to 7x 1mm; lamina of young
plants 3—5-lobed, of mature plants with 2 pairs of
pinnae and a simple or trilobed apical lamina, not
or little dimorphous; apical lamina, if simple, com-
monly 12 x 3 cm (to 17 x 6 cm), acuminate, mar-
gins entire or + sinuous, a bud present on the
upper surface near middle of the costa, if trilobed
broadly cuneate at its base; basal pinnae commonly
to 10 x 2 cm (to 17 x 2 cm), acuminate, with a
narrower basiscopic lobe, on largest fronds the
basiscopic lobe almost free and a small acroscopic
lobe present also; main veins 6—7 mm apart along
the costa, at 45°, sinuous, hardly prominent, smaller
veins very slender but distinct; lower surface of
costae sparsely short-hairy, upper surface densely;

sori small, irregularly scattered, often + elongate
and confluent, usually on free branched veinlets in
areoles, indusia very small, apparently lacking in
some sori.

Distribution — Malesia: Philippines (Luzon,
Samar).

Habitat — On rocks in stream-beds, apparently a
true rheophyte.

76. Tectaria rheophytica Holttum, Blumea 35
(1991) 554. — Type: J.R. Croft 1681, Papua
New Guinea, W Sepik, Blackwater Creek log-
ging area (K; iso LAE).

Caudex short, erect; stipe to 28 cm long, sparse-
ly minutely hairy, glabrescent and glossy, dark
purplish brown; basal scales c. 10 x 1 mm, rigid,
dark, persistent, narrower scales above base ca-
ducous; lamina to 25 cm long, rigid, consisting of
a trilobed apex and 1 pair of pinnae; apical lamina
to 20 cm long, its base rather narrowly decurrent,
midlobe 3—4 cm wide, entire or with sinuous
margins and narrowed to 2.5 cm wide at its base,
lateral lobes to 10 x 1.2 cm, often unequal, their
costae at 45° to the midlobe; pinnae 15 cm long
with stalks to 7 mm long, somewhat dimorphous,
sterile ones to 2.5 cm wide, fertile to 1.7 cm; each
with a basiscopic lobe to 10 x 0.8—1.2 cm; main
veins prominent on lower surface, other veins not
prominent, distinct on young fronds only; lower
surface glabrous, upper surface of costae and main
veins sparsely hairy; sori small, irregularly scatter-
ed, exindusiate, many + elongate along veins but
others on free veinlets in areoles. — Fig. 15b.

Distribution — Malesia: New Guinea (known
only from the type and Croft 1623, foot of Mt Bou-
gainville near West Irian border).

Habitat — On rocky or gritty clay river banks in
the flood zone.

Note — The sori are arranged as in T. christovalen-
sis (C.Chr.) Alston but the frond-form is different;
the latter species is not a rheophyte.

77. Tectaria menyanthidis (Presl) Copel.,
Philipp. J. Sci. 2 (1907) Bot. 414; Fern FI.
Philipp. (1960) 313; Brownlie, Pterid. Fl. Fiji
(1977) 290, pl. 31, f. 3, 4. — Aspidium meny-
anthidis Presl, Reliq. Haenk. (1825) 28; Mett.,
Farngatt. [TV (1858) 124, excl. Zollinger 2369;
Hook., Sp. Fil. 4 (1862) 57; Alderw., Malayan

Holttum — Tectaria Group

Ferns (1908) 244; Holttum, Novit. Bot. Delect.
Seminum Horti Bot. Univ. Carol. Prag. 1968
(1969) 10. — Polydictyum menyanthidis (Presl)
Presl, Epim. Bot. (1851) 53. — Cardiochlaena
menyanthidis (Presl) Fée, Mém. Foug. 5. Gen.
Filic. (1852) 315. — Type: Haenke s.n., Luzon
(PRC).

Polydictyum heterophyllum Presl, Epim. Bot.
(1851) 259; Holttum, Nov. Bot. Delect. Sem.
Horti Bot. Univ. Carol. Prag. 1968 (1969) 47.
— Type: Zippel s.n., New Guinea (PRC).

Nephrodium juglandifolium Baker, J. Bot. 17 (1879)
296. — Aspidium juglandifolium (Baker) Christ,
Bot. Jahrb. Syst. 23 (1896) 354, non Kunze
1847. — Tectaria juglandifolia (Baker) C.Chr.,
Index Filic. Suppl. 3 (1934) 180. — Type:
Horne s.n., Fiji.

Caudex long-creeping, commonly 3—5 mm
diameter when dried (to 8 mm in New Guinea);
stipe 20-50 cm long, lightly flushed reddish, gla-
brous, scaly near base only, scales c. 5 mm long
with filiform apex and widened base; /amina not or
little dimorphous, to 50 cm or more long, consist-
ing of a pinna-like apex and 1—4(—6) pairs of pin-
nae, texture firm; apex commonly 20 x 3—4 cm
(to 40 x 4 cm), sometimes united to uppermost
pair of pinnae; basal pinnae usually unlobed (some
New Guinea specimens have a small basiscopic
lobe), commonly 24 x 3 cm (to 35 x 3.5 cm) with
stalk 10 mm long, gradually narrowed to base and
apex, usually entire; main veins 6-8 mm apart
along costae, at 45° and upcurved, slender, cross-
veins irregular, small veins slightly prominent
when dried; both surfaces glabrous; sori small, on
connected veins, mostly in 2 rather irregular rows
between main veins with some additional ones es-
pecially towards the margin; indusia small, thin,
glabrous, reniform.

Distribution — Malesia: Philippines, New Gui-
nea; Solomons, Banks Is., Fiji.

Habitat — In lowland forest, several times re-
ported on or near stream banks but apparently not
a rheophyte; several New Guinea specimens were
found on limestone.

Note — Young plants have simple fronds, the
lamina to 25 x 4.5 cm and fertile. New Guinea
plants often have a thicker caudex and larger fronds
than those from the Philippines, also very dark
basal scales; scales on Philippine plants are dull
brown with paler filiform tips.

89

78. Tectaria herpetocaulos Holttum, Dansk
Bot. Ark. 23, 2 (1965) 241; Gard. Bull. Sing.
34 (1981) 145. — Type: B. Molesworth Allen
4454, Perak, Cameron Highlands rd, 600 m (K).

Caudex long-creeping, 6—10 mm diameter when
dried, bearing stipes c. 1.5 cm apart, scales 5 x ]
mm, base cordate, dark glossy with paler fragile
edges; stipe 30-80 cm long, lightly flushed with
red, glabrescent, scales near base as those of caudex
but 10 mm long; /amina consisting of 1—4 pairs
of pinnae and pinna-like apex; apex to 21 x 11 cm,
entire, base cuneate, apex abruptly short-acuminate;
upper pinnae sessile, to 21 x 6 cm with asymmet-
ric base rounded on the basiscopic side; basal pin-
nae Stalked, to 30 x 10 cm, asymmetric with a
single basiscopic lobe to 20 x 6 cm (largest fronds
may have 2 pairs of pinnae with basal lobes); main
veins pale on lower surface, 8—10 mm apart along
costae, at a wide angle and upcurved, cross-veins
irregular; lower surface of costae and main veins
bearing sparse short hairs, upper surface of costae
densely short-hairy; sori mostly not on free veins
in areoles, rather small, in a single row on each
side of main veins but with additional ones irregu-
larly between the rows; indusia thin, shrivelling
but persistent.

Distribution — Thailand, Burma, Bangladesh; in
Malesia: Malay Peninsula (Perak).

Habitat — Spreading over humus-covered rocks
in steep rocky forest (type); some specimens from
Thailand found on limestone.

79. Tectaria taccifolia (Fée) M.G. Price, Kali-
kasan 3 (1974) 175. — Leptochilus taccifolius
Fée, Mém. Foug. 2 (1845) 89, t. 50. — Gym-
nopteris taccifolia (Fée) Presl, Epim. Bot.
(1851) 151. — Dendroglossa taccifolia (Fée)
Fée, Mém. Foug. 5. Gen. Filic. (1852) 80. —
Acrostichum taccifolium (Fée) Hook., Sp. Fil.
5 (1864) 278, excl. all varieties. — Hemigram-
ma taccifolia (Fée) Copel., Philipp. J. Sci. 37
(1928) 406; Fern Fl. Philipp. (1960) 318. —
Type: Cuming 357, Mindoro (holotype not at
RB; iso K).

Caudex short, suberect; fronds strongly dimor-
phous. Sterile frond: stipe 20—35 cm long (Cope-
land); lamina to 36 cm long; pinnae one pair, to
22 cm long, 4.5 cm wide, narrowed to base and
apex, margins + sinuous, a basiscopic lobe 14 x

90

Flora Malesiana ser. II, Vol. 2! (1991)

3.5 cm arising 3 cm above its base; rest of frond
lobed to a narrow wing on each:side of the rachis,
the lobes 3 pairs, narrowed at their bases, apex of
lamina conform with the upper lobes, the basal
lobes 21 x 4.5 cm with short basiscopic lobules
like the lobes of basal pinnae but shorter. Fertile
frond: stipe to 50 cm long (Copeland); lamina to
30 cm long, divided as sterile frond but its lobes
and pinnae not over 4 mm wide; pinnae of type 19
cm long with lobes 13 cm long; venation consist-
ing of few narrow areoles, the veins bearing dense-
ly packed spongia throughout, so that the lower
surface appears to be covered by them.

Distribution — Malesia: Philippines (Mindoro
only).

80. Tectaria decurrens (Presl) Copel. in El-
mer, Leafl. Philipp. Bot. 1 (1907) 234; Ching,
Sinensia 2 (1931) 22; Backer & Posth., Varenfl.
Java (1939) 73; Copel., Fern Fl. Philipp. (1960)
315; Molesworth Allen, Gard. Bull. Sing. 22
(1967) 177 with photo; Holttum, Revis. FI.
Malaya ed. 2 (1968) 635; Sledge, Kew Bull. 27
(1972) 420; Holttum, Gard. Bull. Sing. 34
(1981) 139. — Aspidium decurrens Pres], Reliq.
Haenk. (1825) 28; Bedd., Handb. Ferns Brit.
India (1883) 219; Alderw., Malayan Ferns (1908)
247; Holttum, Novit. Bot. Delect. Seminum
Horti Bot. Univ. Carol. Prag. 1968 (1969) 9.
— Sagenia decurrens (Presl) T. Moore, Index
Filic. (1857) Ixxxvi; Racib., Pteridop. Buiten-
zorg (1898) 196. — Type: Haenke s.n., Luzon
(PRC):

Aspidium pteropus Kunze, Bot. Zeitung (Berlin) 4
(1846) 462; Mett., Farngatt. [V (1858) 120. —
Sagenia pteropus (Kunze) T. Moore, Index Filic.
(1858) 89; Bedd., Ferns S India (1863) pl. 82.
— Type: Cuming 148, Luzon (holotype lost;
isotypes BM, K, L, SING, Z).

Cardiochlaena alata Fée, Mém. Foug. 5. Gen. Filic.
(1852) 315. — Type: Cuming 148, Luzon (RB).

Sagenia mamillosa Moore, Ill. Hort. IV, 6 (1886)
t. 598. — Aspidium decurrens var. mamillosum
(Moore) Alderw., Malayan Ferns (1908) 247.
— Type: Cult. ex Moluccas (K).

Aspidium heterodon Copel. in Perkins, Fragm. FI.
Philipp. (1905) 177, non Blume 1828. — Type:
Copeland 951, Mindanao (MICH).

Asplenium alatum Ridley, Trans. Linn. Soc. Lond.
9 (1916) 225. — Aspidium ridleyanum Alderw.,

Malayan Ferns (1917) 505. — Tectaria ridley-
ana (Alderw.) C.Chr., Index Filic. Suppl. 3
(1934) 184. — Type: C.B. Kloss s.n., W New
Guinea, Mt Carstensz Exp. (K).

Tectaria peralata Copel., Philipp. J. Sci. 83 (1954)
98, t. 4. — Type: Edano PNH 12297, Luzon
(MICH; iso BM).

Aspidium macrophyllum auct. non Sw. (1806):
Blume, Enum. PI. Javae (1828) 144.

Caudex short, erect; stipe to 50 cm long, winged
except in basal 2-10 cm, the wing 5-10 mm
wide on each side distally, basal scales 10 mm
long, 1.5—2 mm wide at their bases, firm, distal
scales narrow; lamina thin, lobed, to 60 cm long,
the lobes 2—7 pairs joined by a wing, sterile ones
to 7 cm wide, fertile narrower and sometimes with
sinuous or iregularly lobulate margins, basal
lobes sometimes bilobate; main lateral veins in
lobes c. 10 mm apart along the costa, prominent
on the lower surface, smaller veins amply anasto-
mosing in sterile fronds, fertile ones with closer
main veins and smaller areoles; lower surface of
costae and main veins minutely hairy, minute
hairs also sometimes present between veins, nar-
row scales + abundant on lower surface of rachis
and sometimes on costae of lobes (such scales ap-
parently rare in W Malesia); upper surface quite
glabrous; sori in a row on each side of main veins
in lobes of the frond, also on rachis-wings, on free
veinlets in areoles, variably somewhat immersed
with corresponding projections on the upper sur-
face; indusia large (crumpled when old), glabrous,
round or elliptic, sometimes apparently peltate but
more usually attached along a line from the centre
to the base; spores spinulose.

Distribution — Widely in tropical mainland Asia;
Malesia: W Malay Peninsula (in Perak only), Phi-
lippines, New Guinea; in the Pacific eastwards to
Tahiti.

Habitat — In forest at low and medium altitudes.

Note — Young plants collected by M.G. Price
in Luzon have unlobed linear fronds to about 30
cm long; such fronds may be fertile, the smallest
fertile ones seen 9 cm long.

Cytology — T.G. Walker found that three plants
in New Guinea were diploids; Manton found tetra-
ploids in Sri Lanka. Structural differences between
diploid and tetraploid have not been established.

Possible hybrids — In the Philippines and in New
Guinea are intermediates between this species and

Holttum — Tectaria Group

91

T. repanda; they have narrow fertile pinnae decur-
rent to a narrow wing on the rachis and a short stipe-
wing.

81. Tectaria sulitii Copel., Philipp. J. Sci. 81
(1952) 26; Fern Fl. Philipp. (1960) 313. —
Type: Sulit PNH 8727, Mindanao, Bukidnon
Prov., Mt Katanglad (MICH).

Tectaria loheri Copel., Philipp. J. Sci. 81 (1952) 25;
Fern Fl. Philipp. (1960) 315. — Type: Loher
s.n., 1945, Luzon, Rizal Prov., Montalban (UC).

Tectaria grandifolia auct., non Aspidium grandifo-
lium Presl (1851): Copel., Fern Fl. Philipp.
(1960) 314.

Stipe to more than 70 cm long, dark castaneous,
distal part narrowly winged, basal scales to more
than 3 mm wide; /amina to 110 cm long, pinnati-
sect throughout to a wing along the rachis, wing
above lowest lobes 4 mm wide on each side, wider
in distal part of frond; lobes many, closely placed,
entire, to c. 30 x 6 cm (both sterile and fertile),
basal ones with a large lobe on the basiscopic side;
lower surface of rachis and costae minutely hairy,
upper surface glabrous throughout; veins anasto-
mosing as in T. decurrens, the main lateral veins
mostly + sinuous, smaller veins distinct but hard-
ly prominent on dried fronds; sori in one row on
each side of main veins, some on free veins in
areoles; indusia reniform.

Distribution — Malesia: Philippines (Luzon, Pa-
nay, Negros, Mindanao).

Habitat — In forest, to 1200 m altitude.

Note — M.G. Price, who has seen living plants
on Mt Makiling, regards this as a distinct species,
in frond-form between T. decurrens and T. crenata.
It differs from the former in the firmer texture of
the narrow rachis-wing which extends only a short
distance below the lowest lobes of the lamina, in
the more numerous and closer lamina-lobes, darker
stipe and rachis and broad stipe-scales. The type of
T. loheri Copel., though consisting of an almost
complete frond, is imperfectly developed; that of
T. sulitii is much better and is therefore preferred;
they were published simultaneously. There may be
intermediates between T. sulitii and T. crenata.

82. Tectaria tricuspis (Bedd.) Copel., Sarawak
Museum J. 2 (1917) 369. — Aspidium tricuspe
Bedd., Handb. Ferns Brit. India, Suppl. (1892)

44; Alderw., Malayan Ferns (1908) 235. —
Type: Kunstler 978, Gopeng, Perak (K).

Nephrodium everettii Baker, Kew Bull. (1896) 41.
— Aspidium everettii (Baker) C.Chr., Index
Filic. (1905) 73; Alderw., Malayan Ferns (1908)
241. — Tectaria everettii (Baker) C.Chr., Index
Filic. Suppl. 3 (1934) 179. — Type: Everett
s.n., Natuna Is. (K).

Tectaria clemensiae Copel., Brittonia 1 (1931) 73.
— Type: Clemens 22078, Sarawak, Lundu (UC;
iso K, NY).

Caudex short, erect; stipe usually winged in its
distal part (wing 2—8 mm wide on each side in the
type), basal scales to 10 mm long, 1.5 mm wide
at base, firm and dark with paler margins; lamina
on young plants trilobed, on mature plants con-
sisting of a trilobed apical part decurrent by a nar-
row wing along the rachis which may or may not
join with a pair of pinnae, lobes of pinnae usually
wider in sterile than in fertile fronds; pinnae to 20
cm long, to 8 cm wide (sterile) or 4 cm (fertile),
entire, acuminate, lacking basiscopic lobes, a
small scaly bud sometimes present at base of cos-
tae on lower surface; venation similar in sterile and
fertile fronds, forming many areoles containing
branched free veinlets, the cross-veins between main
veins very regular; lower surface of costae and
main veins bearing sparse very short hairs, upper
surface densely covered with thicker hairs; sori in 2
rows between cross-veins (4—5 irregular rows be-
tween main veins) on veins which surround areoles,
some sori present also on rachis-wing; indusia
reniform but when dried sometimes apparently
peltate, bearing stiff hairs 0.3 mm long on their
upper surface. — Fig. 15e, f.

Distribution — Malesia: Malay Peninsula (Perak,
Natuna Is.), Borneo (Sarawak, Kalimantan).

Habitat — In forest at low altitudes.

Note — There is much variation in width of lobes
and pinnae and of the rachis-wing in different spec-
imens. Buds at the bases of lateral lobes (of tri-
lobed fronds) or pinnae occur on the types of A.
tricuspe Bedd. and T. clemensiae Copel..

83. Tectaria schultzei-(Brause) C.Chr., Dansk
Bot. Ark. 9, 3 (1937) 51. — Polypodium schultz-
ei Brause, Bot. Jahrb. Syst. 49 (1912) 53. —
Pleopeltis schultzei (Brause) Alderw., Malayan
Ferns Suppl. (1917) 399. — Type: Schultze 263,
E New Guinea, Sepik Prov., Augusta R. (B).

92

Flora Malesiana ser. II, Vol. 2! (1991)

Caudex short-creeping; stipe to 40 cm long,
winged except near its base, basal scales 10 mm
long, less than 1 mm wide, their apices filiform,
narrow scales + abundant above base of stipe; la-
mina to 75 cm long, consisting of an elliptic apex
and up to 6 pairs of lobes connected by a wing
which is 8-10 mm wide distally and 3-4 mm
wide near base of frond, lobes to 30 x 6 cm, nar-
rowed towards their bases, lower ones more widely
spaced; main veins in lobes connected by rather
regular outwardly curved cross-veins, the areoles
between them forming a very fine reticulum; /ower
surface of costae short-hairy, small scales some-
times present, short hairs also present on veins and
very short slender ones between veins; sori very
numerous, consisting of few sporangia, exindusi-
ate, scattered irregularly on the smaller veins. —
Fig. 15d.

Distribution — Malesia: New Guinea.

Habitat — In forest at low altitudes, to 900 m.

84. Tectaria vasta (Blume) Copel., Philipp.
J. Sci. 2 (1907) Bot. 411; Backer & Posth.,
Varenfl. Java (1939) 74; Holttum, Revis. FI.
Malaya 2 (1955) 512; Gard. Bull. Sing. 34
(1981) 140. — Aspidium vastum Blume, Enum.
Pl. Javae (1828) 142; Bedd., Handb. Ferns Brit.
India (1883) 212; Alderw., Malayan Ferns (1908)
247. — Type: Blume s.n., Java (L 908,303-
5)

Aspidium alatum Hook. & Grev., Icon. Filic.
(1831) t. 184; Hook., Sp. Fil. 4 (1862) 47. —
Sagenia alata (Hook. & Grev.) Bedd., Ferns Brit.
India (1866) pl. 169. — Type: Wallich 378,
Sylhet (K).

Tectaria decurrenti-alata Ching & Wang, Acta Phy-
totax. Sin. 19 (1981) 126. — Type: Yunnan
Complex Exp. 931, Yunnan, Jenping (PE).

Caudex erect or suberect; stipe to 45 cm long,
winged almost to its base, the wing to 10 mm
wide on each side, near base densely scaly on ab-
axial side, largest scales to 20 x 2 mm, very firm,
very narrow scales present on adaxial side; lamina
to 80 cm long, lobed to a wing on each side of the
rachis (wing widest distally), lobes pinna-like, 1-6
pairs, largest 30 x 7 cm, entire, short-acuminate,
the basal ones sometimes with a narrowly acumi-
nate basiscopic lobe; veins amply anastomosing;
lower surface of costae and main veins variably
short-hairy, upper surface of costae glabrous or

with sparse very short hairs; sori small, numerous,
rather irregularly arranged on veins which surround
areoles; indusia distinct, thin, + reniform, their
margins usually short-fringed when young.

Distribution — Assam and W Yunnan southwards
to Burma and Thailand; W Malesia.

Habitat — In lowland forest.

Note — The type of T. decurrenti-alata is an un-
usually large specimen but not much larger than
some from Assam and does not differ appreciably
in other ways.

85. Tectaria grandidentata (Cesati) Holttum,
Revis. Fl. Malaya 2 (1955) 514. — Polypodium
dilatatum var. grandidentatum Cesati, Atti Accad.
Sci. Fis. Napoli 7, no 8 (1876) 27. — Polypo-
dium grandidentatum (Cesati) Baker, Ann. Bot.
(London) 5 (1891) 479; Alderw., Malayan Ferns
(1908) 661. — Pleopeltis grandidentata (Cesati)
Alderw., Malayan Ferns Suppl. (1917) 397. —
Type: Beccari, Sarawak, G. Matang (FI; K).

Dictyopteris carinata Alderw., Bull. Jard. Bot. Bui-
tenzorg II, 28 (1918) 18. — Tectaria carinata
(Alderw.) C. Chr., Index Filic. Suppl. 3 (1934)
178. — Type: Brooks 370, Sumatra, Lebong
Tandai (BO; iso BM).

Fronds of young plants lanceolate with undulate
margins, the lamina c. 20 x 2.5 cm on a slender
stipe 8 cm long, successive fronds longer and more
deeply lobed, the lobes decrescent both towards base
and apex; mature plants with short erect caudex,
stipes to 60 cm long, their upper part winged, a
deeply lobed /amina to c. 85 cm long, lower lobes
a little decrescent and gradually more widely spaced
than the upper ones, the rachis-wing c. 5 mm wide
each side at the base, 10-15 mm distally; largest
lobes c. 20 x 3 cm, hardly narrowed at their base,
very gradually narrowed distally, margins entire
near the base, minutely crenate-dentate distally;
venation prominent on both sides when dried,
forming many areoles between main veins; lower
surface of costae and veins bearing minute hairs,
upper surface quite glabrous; sori small, numerous,
exindusiate, irregularly scattered, mostly on con-
nected veins, rarely terminal on free veinlets in
areoles; branched paraphyses as long as the spo-
rangia present; spores with translucent wing and
few cross-wings.

Distribution — Malesia: Sumatra, Malay Penin-
sula, Borneo (Sarawak).

Holttum — Tectaria Group

93

Habitat — On rocks in forest, not on stream
banks; fronds of mature plants pendulous.

86. Tectaria beccariana (Cesati) C.Chr., Index
Filic. Suppl. 3 (1934) 177; C. Chr., Dansk Bot.
Ark. 9, 3 (1937) 51; Copel., Philipp. J. Sci.
78 (1951) 419; Fern Fl. Philipp. (1960) 308.
— Polypodium beccarianum Cesati, Rendiconti
Reale Accad. Sci. Fis. (Napoli) (1877) 30. —
Aspidium beccarianum (Cesati) Diels in E. & P..,
Nat. Pflanzenfam. I, 4 (1899) 186, non Baker
1886. — Dictyopteris beccariana (Cesati) Al-
derw., Malayan Ferns (1908) 515. — Type:
Beccari s.n., W New Guinea (FI; iso K).

Aspidium bryantii Copel. in Perkins, Fragm. FI.
Philipp. (1905) 175. — Tectaria bryantii
(Copel.) Copel., Philipp. J. Sci. 2 (1907) Bot.
412. — Dictyopteris bryantii (Copel.) Alderw.,
Malayan Ferns (1908) 514. — Type: Copeland
82, Negros Is. (MICH, not seen).

Aspidium bolsteri Copel., Philipp. J. Sci. 1 (1906)
Suppl. 252; ibid. 2 (1907) Bot. 412. — Dic-
tyopteris bolsteri (Copel.) Alderw., Malayan
Ferns (1908) 519. — Type: Bolster 305, Min-
danao (MICH).

Polypodium ingens Brause, Bot. Jahrb. Syst. 56
(1920) 200. — Type: Ledermann 6582, E New
Guinea, Sepik Prov. (B).

Aspidium vastum forma latior Alderw., Nova Gui-
nea 14 (1924) 7. — Type: H.J. Lam 933, W
New Guinea, Mamberamo R. (BO; iso L).

Tectaria celebica C.Chr., Bot. Jahrb. Syst. 66
(1933) 49. — Type: Kjellberg 418, SE Celebes
(S; BO).

Very near Tectaria vasta (Blume) Copel., differ-
ing in firmer texture of fronds with all veins promi-
nent on lower surface, wider wing on stipe (to 2
cm wide on each side) and rachis, wider lamina-
lobes on largest fronds (to 10 cm wide), basal ones
of large fronds always with a large basiscopic lobe,
sori exindusiate or rarely with very small indusia.
— Fig. 16f, g.

Distribution — Malesia: Borneo (Sabah), Philip-
pines (Luzon, Negros, Samar, Mindanao), SE Cele-
bes, widely in New Guinea.

Habitat — In forest at low to medium altitudes.

Notes — The relationship of this species to T.
vasta needs experimental study. The widely distri-
buted indusiate 7. vasta must be the more primi-
tive of the two; it does not occur in Taiwan, so

that the Philippine 7. beccariana may be an eastern
development from T. vasta in Vietnam. In Sabah
are possible hybrids between the two species.

The type of 7. celebica is an immature plant;
the lamina of its largest frond is 50 x 9 cm with
entire or subrepand margins, fertile, with sori as in
T. beccariana.

87. Tectaria novoguineensis (Rosenst.) C.
Chr., Index Filic. Suppl. 3 (1934) 182. — As-
pidium novoguineense Rosenst., Meded. Rijks-
herb. 31 (1917) 4. — Type: Zippel s.n., W
New Guinea (L 908,302-327).

Caudex short, erect or suberect; stipe to 17 cm
long, very dark, minutely hairy throughout, basal
scales 5 mm long, very narrow; lamina simple, of
young plants ovate-acute with deeply cordate base
and irregularly shallowly lobed margin, of mature
plants firm, 3-lobed to sub-5-lobed, to 12 cm long
from top of stipe to apex and 11 cm wide, the
lobes obtuse, irregularly slightly lobulate; both
surfaces bearing many short slender erect hairs
throughout; sori on free veinlets in areoles, rather
large, irregularly arranged; indusia hairy.

Distribution — Malesia: New Guinea; 2 collec-
tions.

Habitat — The second collection is from crevices
in limestone on the north coast, near the border be-
tween W Irian and Papua New Guinea (Croft 1600).
The type was probably also from limestone (some
other Zippel collections certainly are).

88. Tectaria johannis-winkleri (C. Chr.)
C. Chr., Index Filic. Suppl. 3 (1934) 180. — As-
pidium johannis-winkleri C.Chr., Mitt. Inst.
Allg. Bot. Hamburg 7 (1928) 149. — Type:
Hans Winkler 849, Kalimantan, Bidang Mena-
bai (HBG).

Caudex short, suberect, its apex covered with
narrow light brown scales; stipe very dark, gla-
brous, 25 cm long; lamina to 20 cm long and
wide, its base subcordate, near base lobed halfway
to the costa, less deeply distally, lobes subcaudate,
the lowest largest and sometimes with a basiscopic
acuminate lobule; Jower surface rather densely cov-
ered with rufous hairs 0.2 mm long which are more
dense near the margin, upper surface glabrous ex-
cept near the margin; sori small, irregularly scat-
tered, mostly not on free veinlets; indusia firm,

94 Flora Malesiana ser. II, Vol. 2! (1991)

PJ. Epwarps

Fig. 16. Tectaria christovalensis (C. Chr.) Alston. a. Outline of a fertile frond, x 0.22; b. detail of a,
showing pattern of sori mostly on free veins in areoles, x 2.2. — T. inopinata Holttum. c. Outline of a
frond, x 0.4; d. detail of c, showing few simple included veinlets in costal areoles, x 1.7. — T. kehding-
iana (Kuhn) M.G. Price. e. Portion of a fertile frond, with indusia removed on right side, x 2.6. —
T. beccariana (Cesati) C. Chr. f. Outline of frond, x 0.16; g. detail of soral pattern in rachis-ring, x 1.7
(a, b: von Mueller coll. 1879; c, d: Matthew s.n., 29-11-1912; e: Boden Kloss 14766; f, g: Millar &
Holttum NGF 15863).

Holttum — Tectaria Group

95

dark, persistent, subpeltate, bearing a few hairs like
those on the lower surface between veins.
Distribution — Malesia: Borneo (known only
from the type).
Habitat — In forest at 650 m, ‘epiphyte’ (collec-
tor’s label, but no other Tectaria species are thus

reported).

89. Tectaria nitens Copel., Philipp. J. Sci. 56
(1935) 475, t. 6 — Type: Clemens 40652, Sa-
bah, Mt Kinabalu (holotype PNH destroyed; iso
MICH).

Caudex short, suberect, its apex densely covered
with mid-brown narrow scales 5 mm long; stipe
20 cm long, very dark, glossy; Jamina ovate in
outline, to 18 x 12 cm, base deeply cordate with
the basal lobes overlapping, near base lobed 1/3
towards the midrib, distally less deeply, the lobes
contiguous, larger ones falcate and abruptly sub-
caudate; lower surface quite glabrous, upper surface
with some short hairs near the margin; sori irregu-
larly arranged, not on free veins in areoles; indusia
persistent, entire, reniform to peltate.

Distribution — Malesia: Borneo (known only
from the type).

Habitat — Jungle hillside under cliff, 1200 m.

90. Tectaria stalactica M.G. Price, Kalikasan
3 (1974) 116, f. 2 — Type: M.G. Price 2266,
Luzon, Cagayan Prov. (MICH; iso K).

Caudex short with tufted stipes; stipe to 40 cm
long, atropurpureous; basal scales to 12 x 2 mm,
margins conform; /amina deltoid in outline, to 35
cm long and 30 cm wide at its base, deeply 3—S-
lobed, the basal lobes with basal basiscopic lobes
on the largest fronds; apical lobe narrowly acumi-
nate, its margins sinuous on small fronds, lobu-
late with acuminate lobules on large fronds, basal
lobes similar, their apices upcurved; costae beneath
dark at their bases, paler distally, main veins slen-
der, slightly prominent, at c. 45°, almost straight,
smaller veins very slender and not prominent; both
surfaces glabrous except for a few hairs near base
of the midrib on the upper surface; sori rather ir-
regularly arranged, irregular in outline, not on free
veins in the areoles; indusia thin, small, persis-
tent, with a fringe of short slender hairs and some
on the upper surface, the hairs rarely conspicu-
ous.

Distribution — Malesia: Philippines (type local-
ity only).

Habitat — On dripping limestone cliffs, the
fronds pendulous.

91. Tectaria palmata (Mett.) Copel., Sarawak
Museum J. 2 (1917) 369. — Aspidium palma-
tum Mett., Ann. Mus. Bot. Lugd.-Bat. 1 (1864)
238; Alderw., Malayan Ferns (1908) 247. —
Lectotype (selected here): Korthals s.n., Kali-
mantan (L 916,68-77).

KEY TO THE VARIETIES

la. Fertile fronds isomorphous with sterile ones,
or nearly so, their lower surface glabrous be-
EWEENEVEINS es aye eae a Seis ele 2
b. Fertile fronds much contracted, their lower sur-
face bearing many hairs between veins
d. var. dimorpha
2a: Indusiaidistinct oniallisoni.22... 2)53- 3)
b. Indusia very small, hidden by sporangia, or
laCkiN Gia schtec ero: c. var. platanifolia
3a. Costae densely short-hairy on upper surface
a. var. palmata
b. Costae glabrous on upper surface
b. var. sumatrana

a. var. palmata

Caudex suberect with tufted stipes, its scales nar-
rowed distally; stipe very dark, minutely hairy
(glabrescent), to 45 cm long, longer on fertile than
on sterile fronds; Jamina on young plants (some-
times fertile) elongate, unlobed, with cordate base
and acuminate apex, on mature plants deeply S-
lobed, to c. 20 cm long and wide, the lobes entire,
acuminate, a scaly bud present near the base on the
upper surface; costae beneath minutely hairy near
their bases, densely short-hairy on the upper surface;
sori numerous, small, either on veins surrounding
areoles or on free veinlets in areoles; indusia less
than 1mm diameter, reniform, persistent, with
minutely papillose margins.

Distribution — Malesia: Borneo (Central & SE
Kalimantan). ;

b. var. sumatrana Holttum, Blumea 35 (1991)
554. — Type: Lérzing 14739, N Sumatra, G.
Sibajak, 850 m (BO).

96

Flora Malesiana ser. II, Vol. 2! (1991)

Differs from the typical variety in the costae
being entirely glabrous and lightly sulcate above.

Distribution — Malesia: Sumatra (known only
from the type).

c. var. platanifolia (Mett.) Holttum, Blumea 35
(1991) 554. — Aspidium platanifolium Mett.,
Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 239; Al-
derw., Malayan Ferns (1908) 235. — Tectaria
platanifolia (Mett.) Copel., Sarawak Museum
J. 2 (1917) 369. — Type: Wallace s.n., Borneo
(holotype destroyed; iso K).

Aspidium angulatum auct.: Hook., Sp. Fil. 4 (1862)
44, p.p.

Differs from var. palmata: sori much smaller
and more numerous, exindusiate or with very small
indusia hidden by sporangia, those on veins near
costae sometimes a little elongate or confluent.

Distribution — Malesia: Borneo (Sarawak, many
collections; E Kalimantan).

Habitat — In lowland forest, in some cases re-
ported from wet ground.

d. var. dimorpha Holttum, Blumea 35 (1991)
554. — Type: K. Iwatsuki et al. B-980, E Kali-
mantan, Tabang (K; KYO).

Differs from the typical variety in the stipe
being up to 20 cm long and in the contracted
fertile fronds clothed with many 0.5 mm long or
longer hairs between the veins.

Distribution — Malesia: Borneo (known only
from the type locality).

Habitat — In wet ground by stream in shade; alti-
tude 100-150 m.

Notes — The indusia of var. dimorpha are rath-
er smaller than those on specimens of var. palmata
and have papillae on the upper surface as well as
on the margins; the sori are close together on the
contracted lamina.

In both var. platanifolia and var. dimorpha a
scaly bud is sometimes present at the base of the
lamina, on the upper surface of the costa; such buds
have not been seen in var. palmata.

92. Tectaria filisquamata Holttum, Blumea
35 (1991) 550. — Type: T. Lobb 259, Java (K;
iso BM).

Caudex short, erect, its apex covered with nar-
row scales which have filiform apices each consist-

ing of a row of 10 or more cells; stipe to 18 cm
long, slender, mid-castaneous, glabrous; lamina of
fronds of young plants elongate and slightly tri-
lobed, its margins + sinuous distally, of mature
plants subequally trilobed, to 14 cm long, base
shortly cordate, lobes entire, acuminate, midlobe
to 5.5 cm wide, lateral lobes to 3 cm; lower sur-
face bearing very short slender hairs throughout be-
tween veins, upper surface glabrous including cos-
tae of lobes; main veins of midlobe 8—10 mm apart
along the costa, at a wide angle, slender, small
veins forming areoles between them distinct but
not prominent; sori small, irregularly scattered,
mostly not on free veins in areoles, occasionally
+ elongate, exindusiate.

Distribution — Malesia: Java, known from Lobb
258 & 259 only.

Note — Thomas Lobb collected for Veitch in W
Java in 1845-46; if these specimens in fact came
thence, the species is now perhaps extinct.

93. Tectaria microchlamys Holttum, Blumea
35 (1991) 553. — Type: J.R. Croft 1115, Ma-
nus Is. (K; iso LAE).

Caudex short, erect or suberect; stipe dark cas-
taneous, minutely hairy, to at least 22 cm long,
basal scales to 7 mm long, not hair-pointed; lami-
na thin, to 28 cm long, deeply trilobed, its base not
cordate, midlobe to 7 cm wide, narrowed to 1.2 cm at
its base, lateral lobes to 18 x 5 cm, entire, short-acu-
minate; smaller veins concolorous on both surfaces,
+ prominent; no bud present at base of frond; lower
surface of costae and main veins minutely hairy,
very short hairs variably present between veins,
upper surface glabrous; sori small, mostly not
on free veins in areoles, sometimes a little elon-
gate; indusia very small and apparently variable in
shape, bearing many unicellular acicular hairs.

Distribution — Malesia: New Guinea (known on-
ly from the type collection).

Habitat — On vertical clay bank in lowland for-
est; fronds horizontal or pendulous.

Note — Young plants have an almost evenly el-
liptic lamina with rounded apex and broadly cuneate
base (those seen 3.9 x 2.5 cm).

94, Tectaria subtriloba Holttum, Blumea 35
(1991) 556. — Type: Hennipman 5859, SE
Celebes between Soroako and Malili (L; iso K).

Holttum — Tectaria Group

OF)

Caudex short, erect, its apex covered with nar-
row shortly hair-pointed scales 5 mm long; stipe
atrocastaneous, minutely hairy, of sterile fronds to
23 cm long, of fertile to 42 cm; sterile lamina
very firm, rather narrowly ovate, acute, to 14.5 cm
long, its base cordate, with a short acute lobe on
each side near the base, width above the lobes 9.5
cm; fertile lamina similar in shape but somewhat
smaller; lower surface of costae and main veins
sparsely minutely hairy, between veins glabrous;
upper surface of costae densely short-hairy, fewer
hairs on main veins; sori small, abundant, irregu-
larly scattered, exindusiate, often on veinlets in
areoles but also on veins surrounding areoles, of-
ten + elongate or coalescent.

Distribution — Malesia: Celebes (known only
from the type collection).

Habitat — Ultrabasic area, roadside on forest mar-
gin near a small waterfall.

95. Tectaria christovalensis (C. Chr.) Al-
ston, J. Bot. 77 (1939) 290, excl. syn. Aspi-
dium pentaphyllum Alderw. — Polypodium
christovalense C. Chr., Index Filic. (1905) 516,
new name for Gymnogramma palmata Baker,
Syn. Fil. (1868) 389, not Polypodium palma-
tum Blume 1828. — Type: Milne 508, Solo-
mon Is., San Cristobal (K).

Campylogramma pteridiformis Alderw., Bull. Jard.
Bot. Buitenzorg II, 24 (1917) 1; Nova Guinea
14 (1924) 10, t. 1, f. A. — Type: Thomsen 651,
W New Guinea, Mamberamo R. (BO; tso L).

Tectaria diversisora Copel., Philipp. J. Sci. 30
(1926) 328. — Type: C. King 470, Papia New
Guinea, Hydrographers Range (MICH).

Caudex short, erect; stipe of fertile fronds to 35
cm long, of sterile shorter, very dark, glabrescent;
basal scales to 7 mm long, firm, not hair-pointed;
lamina firm, commonly to 15 cm long (largest
seen 25 cm), very deeply trilobed, the lateral lobes,
especially on fertile fronds, usually with small
basiscopic lobules of variable size, base of lamina
not cordate; apical lobe of fertile fronds commonly
3-4 cm wide, of sterile variably wider; a scaly
bud often present near the base of a lateral costa on
the lower surface; small veins concolorous on
lower surface, sometimes hardly distinct though
evident by transmitted light; lower surface glabrous;
upper surface of costae densely short-hairy near
their bases, less so distally; sori mostly on free

veins in areoles, exindusiate, often + elongate or
confluent. — Fig. 16a, b.

Distribution — Malesia: New Guinea; Solomon
Is. (San Cristobal, Malaita).

Habitat — In lowland forest, sometimes on steep
slopes; on limestone in Malaita Is.

Note — In Herb. Kew is a specimen from New
Ireland which has the apical lobe deeply trilobed,
agreeing in other respects.

96. Tectaria minuta Copel., Philipp. J. Sci.
30 (1926) 328. — Type: C. King 493, Papua
New Guinea, Hydrographers Range (MICH).

Caudex short, suberect, scales narrow, to 3 mm
long; stipe to 4.5 cm long, slender, dark, minutely
hairy; lamina firm, ovate, base cordate, apex round-
ed, margins broadly crenate, to 4 cm long and 2.7
cm wide; both surfaces glabrous, including upper
surface of costa; main veins not prominent, not
reaching margin, smaller veins forming areoles
with few included veinlets; sori sparse, some at the
ends of veinlets in areoles; indusia round-reniform,

persistent.
Distribution — Malesia: New Guinea (known
only from the type).

Note — The type consists of one plant bearing a
tuft of about 10 fronds, several of the larger ones
fertile and similar in size.

97. Tectaria labrusca (Hook.) Copel., Philipp.
J. Sci. 2 (1907) Bot. 410. — Polypodium la-
brusca Hook., Sp. Fil. 5 (1863) 73, t. 285B,
p.p.; Baker, Syn. Fil. (1868) 361, p.p. — Lec-
totype (selected here): T. Lobb s.n., 1857, Sa-
rawak (specimen named by Hooker, K).

Nephrodium vitis Racib., Bull. Int. Acad. Sci.
Cracovie (1902) 60. — Sagenia vitis (Racib.)
Christ, Ann. Jard. Bot. Buitenzorg 20 (1905)
108. — Aspidium vitis (Racib.) C. Chr., Index
Filic. (1905) 97. — Dictyopteris vitis (Racib.)
Alderw., Malayan Ferns (1908) 516. — Type:
Cult. Hort. Bog. ex Borneo (KRA; iso BO).

Caudex suberect or partly decumbent; stipe 20-—
40 cm long, dark castaneous, minutely hairy at the
base, basal scales to 8 mm long, narrow, with
long slender tips; Jamina broadly ovate, its base
strongly cordate, apex very shortly apiculate, mar-
gins broadly and shallowly lobed (rarely distinct-
ly trilobed) to sinuous, largest seen 18 x 12 cm,

98

Flora Malesiana ser. II, Vol. 2! (1991)

smallest fertile 9 x 7 cm; main veins prominent
on both surfaces, distinct to the margin, rather
widely spaced, smaller veins forming many areoles
with included veinlets; lower surface densely cov-
ered with slender erect hairs 0.5 mm or more long
between veins, the hairs brownish when dried, up-
per surface more sparsely hairy; sori small, exindu-
siate, sometimes a little elongate, mostly on free
veinlets in areoles.

Distribution — Malesia: Borneo (Sarawak and
Kalimantan).

Habitat — On limestone, in rock crevices.

Notes — Hooker received two collections of speci-
mens from limestone in Sarawak made by Thomas
Lobb in 1857. They are mounted on separate sheets
and evidently Lobb recognized them as distinct. But
Hooker included both in one species; his t. 285B
shows a frond from each sheet. He wrote the name
Polypodium labrusca on one sheet only; which is
now marked as lectotype; the specimens on the
other sheet represent T. brooksii. Hooker omitted
reference to hairs on the fronds, but Baker (1868)
stated that the fronds are glabrous, thus misleading
Raciborski who gave the new name Nephrodium
vitis to a specimen of the true T. labrusca and was
copied by Alderwerelt. See further comment under
T. brooksii.

Probable hybrid: M. Kato et al. B5492 from Ka-
limantan has almost entire fronds, as in T. brooksii
and shorter and more sparse hairs on the lower sur-
face than in T. labrusca; it looks like a hybrid. The
two species grow together, as noted by Brooks.

98. Tectaria brooksii Copel., Philipp. J. Sci. 6
(1911) Bot. 137, pl. 20b. — Aspidium brooksii
(Copel.) C. Chr., Index Filic. Suppl. 1 (1913) 7.
— Dictyopteris brooksii (Copel.) Alderw., Ma-
layan Ferns Suppl. (1917) 322. — Type: C.J.
Brooks 26a, Bidi, Sarawak (MICH; iso BM).

Polypodium labrusca auct.: Baker, Syn. Fil. (1868)
361, p.p. — Aspidium labrusca auct.: Racib.,
Bull. Int. Acad. Sci. Cracovie (1902) 61. -—
Dictyopteris labrusca auct.: Alderw., Bull. Dép.
Agric. Ind. Néerl. 18 (1908) 16, excl. var. ter-
nata; Malayan Ferns (1908) 517, excl. var.

Caudex short, erect or suberect; stipe dark near
its base, less so distally, commonly 15 cm long,
basal scales to 5 mm long and to almost 1 mm
wide at base; /amina firm to rigid, simple, broadly
ovate with strongly cordate base, apex short-acumi-

nate, margins entire or irregularly slightly sinu-
ous, commonly 8 x 5 cm, largest seen 17 x 10.5
cm, rarely + trilobed; main veins sinuous, hardly
prominent, upcurved; lower surface glabrous, up-
per surface of costa variably short-hairy, at least
near the base of the lamina; sori irregularly scatter-
ed, mostly punctate but often + elongate or con-
fluent, exindusiate, on both free veinlets in areoles
and on veins surrounding areoles.

Distribution — Malesia: Borneo, S & Central
Celebes, E New Guinea.

Habitat — On limestone, at 0—1200 m, in rock
crevices, sometimes growing with T. labrusca.

Note — See comment under T. labrusca. Cope-
land (I.c. 1911) was the first person to note that
Aspidium labrusca sensu Hook. (1864) was based
on specimens of two species; he distinguished be-
tween them in the brief diagnosis of his species.

99. Tectaria inopinata Holttum, Blumea 35
(1991) 551. — Type: A.C. Jermy & J.M. Ran-
kin J 15324, Sabah, near Tenom, 2200m (BM).

In shape of lamina very near T. brooksii Copel.,
but thinner; basal scales to 8 mm long, very nar-
rowly attenuate; largest lamina seen 16 cm long
from attachment to stipe to the rather narrrowly
acuminate apex; free veinlets lacking in costal and
costular areoles and very few, unbranched, in other
areoles; sori exindusiate, irregular, nearly all + elon-
gate along veins, those on the outer veins of costal
areoles longest. — Fig. 16c, d.

Distribution — Malesia: Borneo, known only
from the type locality (two collections) and from
G. Matang, Sarawak.

Habitat — In forest, not on limestone, on sloping
ground.

100. Tectaria zollingeri (Kurz) Holttum,
comb. nov. — Hemionitis zollingeri Kurz, Na-
tuurk. Tijdschr. Ned. Ind. 27 (1864) 16; J. Asiat.
Soc. Bengal 39, 2 (1870) 90, pl. 5. — Syn-
gramma zollingeri (Kurz) Diels in E. & P., Nat.
Pflanzenfam. I, 4 (1899) 257. — Hemigramma
zollingeri (Kurz) Christ, Philipp. J. Sci. 2 (1907)
Bot. 170, excl. spec. Philipp.; Alderw., Malayan
Ferns (1908) 831, excl. spec. Philipp. — Type:
Zollinger s.n., Cult. Hort. Bog. (not seen).

Hemigramma latifolia auct.: Backer & Posth., Va-
renfl. Java (1939) 79; Copel., Fern Fl. Philipp.
(1960) 371, p.p.

Holttum — Tectaria Group

Caudex short, erect; fronds very dimorphous.
Sterile frond: stipe commonly less than 5 mm
long (to c. 30 mm); lamina commonly to 15 x 6
cm (largest seen 24 x 7 cm), base + cordate where
stipe is short, cuneate where stipe is longer, mar-
gins entire or + sinuous, not deeply lobed, apex
obtuse; all small veins distinct on both surfaces;
both surfaces quite glabrous. Fertile frond: stipe to
15 cm long, + persistently scaly, scales thin, light
brown, to 7 mm long and 0.5 mm wide; lamina to
12 cm long, commonly 8 mm wide, very gradual-
ly narrowed to base and to apex; margins often +
sinuous; very narrow scales present on lower sur-
face of costa; venation much reduced and lacking
included veinlets in areoles, the main veins very
oblique, connected by simple cross-veins; sori
spreading all along cross-veins.

Distribution — Malesia: Java, SW Celebes, Mo-
luccas (Tanimbar Is., Ambon).

Habitat — On steep earth banks in shaded places,
especially near streams, at low altitudes.

101. Tectaria hilocarpa (Fée) M.G. Price, Ka-
likasan 3 (1974) 175; Holttum, Gard. Bull. Sing.
39 (1986) 154, pl. 1. — Leptochilus hilocarpus
Fée, Mém. Foug. 2 (1845) 87, pl. 48, f. 1. —
Type: Gaudichaud s.n., Voy. Bonite, Manila
(P).

Gymnopteris latifolia Goldm., Nov. Actor. Acad.
Caes. Leop. Nat. Cur. Suppl. 1 (1843) 460;
Presl, Epim. Bot. (1851) 150 [non Tectaria
latifolia (G. Forster) Copel. 1907]. — Lepto-
chilus latifolius (Goldm.) C. Chr., Index Filic.
(1905) 386. — Hemigramma latifolia (Goldm.)
Copel., Philipp. J. Sci. 2 (1907) Bot. 406; ibid.
3 (1908) Bot. 31, pl. 1; Fern Fl. Philipp. (1960)
317, excl. Hemionitis zollingeri et nom. cogn.
— Type: Meyen s.n., Luzon, Rizal Prov. (B).

Leptochilus subquinquefidus Fée, Mém. Foug. 2
(1845) 88, pl. 49. — Dendroglossa subquinque-
fida (Fée) Fée, Mém. Foug. 5. Gen. Filic. (1852)
80, pl. 7, f. Bl. — Type: Cuming 3, Luzon
(RB; iso K).

Gymnopteris trilobata J. Sm. ex Hook., Icon. PI.
10 (1854) t. 7. — Lectotype (selected here): T.
Lobb s.n., Luzon (K).

Hemionitis gymnopteroidea Copel. in Perkins,
Fragm. Fl. Philipp. (1905) 183, incl. forma
major. — Lectotype (selected here): Merrill
3113, Luzon (US).

Hemigramma zollingeri (Kurz) Christ var. major
Christ, Philipp. J. Sci. 2 (1907) Bot. 170. -
Specimens cited not seen.

Closely related to T. zollingeri, differing as fol-
lows: lamina of sterile fronds of larger plants vari-
ably + deeply lobed, sometimes very deeply, with
lamina to 30 cm long having several pairs of lobes
to 14 x 3.5 cm connected by a narrow wing along
the midrib, the wing decurrent below the lowest
lobes; lower surface of midrib of frond, costae of
lobes and main veins always bearing many short
hairs; lamina of simple fertile fronds to 3 mm
wide and up to 15 cm long, larger fertile fronds
irregularly lobed as sterile, the lobes commonly 3
mm wide (intermediates between sterile and fertile
fronds, with wider lobes, may occur).

Distribution — Malesia: N Borneo, Philippines
(Luzon, Leyte, Panay, Mindanao).

Habitat — On damp clay banks in shade, low
country (C.G. Matthew).

Note — Under Hemionitis gymnopteroidea Cope-
land briefly described forma major, based on a
plant with large simple sterile fronds, Copeland
1398, of which no specimens now exist (M.G.
Price). Christ’s new name Hemigramma zollingeri
var. major was based on Copeland’s Hemionitis

‘gymnopteroidea forma major, but Christ described

deeply lobed fronds and cited other specimens.

102. Tectaria kehdingiana (Kuhn) M.G.
Price, Brit. Fern Gaz. 19 (1972) 262. — Luers-
senia kehdingiana Kuhn in Luerssen, Bot. Cen-
tralbl. 11 (1882) 77; Diels in E. & P., Nat.
Pflanzenfam. I, 4 (1899) 180, f. 93A, B; Alderw.,
Malayan Ferns (1908) 233. — Aspidium keh-
dingianum (Kuhn) Christ, Farnkr. Erde (1897)
235. — Type: Kehding 2991, NE Sumatra,
Langkat (B; iso SING).

Caudex short, erect; fronds strongly dimorphous.
Sterile frond: stipe c. 5 cm long, basal scales less
than 1 mm wide; lamina to 35 x 5 cm, widest
above the middle, narrowed very gradually towards
the base, apex hardly acuminate; main veins hardly
prominent on the lower surface, always + sinuous,
free veinlets present in areoles between them; both
surfaces glabrous. Fertile frond: stipe c. 25 cm long,
lamina to 25 x 2 cm; veins forming 3 rows of
+ hexagonal areoles between midrib and margin,
one free vein in each areole; sori on free veins in

100

Flora Malesiana ser. II, Vol. 2! (1991)

areoles; indusia elliptic in outline, c. 3—4 x 2 mm
when young, firm and persistent. — Fig. 16e.
Distribution — Malesia: Sumatra (NE and Men-
tawi Is.). ;
Habitat — Low altitude; no details.

103. Tectaria pandurifolia (C. Chr.) C. Chr.,
Index Filic. Suppl. 3 (1934) 183. — Aspidium
pandurifolium C.Chr., Mitt. Inst. Allg. Bot.
Hamburg 7 (1928) 149. — Type: Hans Winkler
367, W Kalimantan, Lebang Hara (BM).

Fronds simple, differing from those of T. singa-
poreana as follows: caudex perhaps not erect; un-
winged stipes of fertile fronds to 4 cm long; lamina
of fertile fronds c. 30 cm long, 10-13 cm wide,
abruptly narrowed near the base to a wing 1-2 cm
wide on each side of the midrib; upper surface of
midrib densely short-hairy, fewer similar hairs on
main veins.

Distribution — Malesia: Borneo (known from the
type only).

Habitat — In forest at 180 m altitude.

104. Tectaria singaporeana (Hook. & Grev.)
Copel., Sarawak Museum J. 2 (1917) 368;
Holttum, Revis. Fl. Malaya 2 (1955) 512. —
Aspidium singaporeanum Hook. & Grev., Icon.
Filic. (1827) t. 26; Bedd., Ferns Brit. India (1866)
t. 168; Handb. Ferns Bnit. India (1883) 212;
Alderw., Malayan Ferns (1908) 235. — Type:
Wallich s.n., Singapore (E; iso K).

Caudex erect, to 10 cm tall; fronds somewhat
dimorphous; stipe of sterile fronds commonly to
10 cm long, of fertile ones to 25 cm, basal scales

8 mm long, mid-castaneous, narrow, rather closely
fringed when young; /amina simple, widest about
1/3 from the base, sterile to 40 x 12 cm, fertile to
7 cm wide, margins entire, apex short-acuminate,
base + abruptly narrowed (more abruptly in sterile
than in fertile fronds) and narrowly decurrent for a
short distance; main veins c. 10 mm apart near base
of lamina, cross-veins between them rather regular,
2 rows of areoles between cross-veins; both sur-
faces glabrous including midrib; sori to 6 in a row
on each side of cross-veins, not on free veins in
areoles; indusia peltate or nearly so, entire, some-
times with a few short hairs on upper surface.

Distribution — Peninsular Thailand; in Malesia:
Sumatra, Malay Peninsula, Borneo.

Habitat — In lowland forest, rather common in
the Peninsula but little collected elsewhere.

105. Tectaria rara (Alderw.) C. Chr., Index Fi-
lic. Suppl. 3 (1934) 184. — Aspidium rarum
Alderw., Bull. Jard. Bot. Buitenzorg II, 28 (1918)
7. — Type: Biinnemeijer 2258, Bangka (BO).

Similar in habit and frond-form to T. singapore-
ana, differing as follows: fronds smaller, sterile ones
little wider than fertile; largest fertile lamina 18 x
3 cm, its narrowly decurrent base 10-15 mm long;
margins rather regularly sinuate-crenate (one main
vein to each lobe) to a depth of 2-3 mm; main
veins c. 6 mm apart along midrib, cross-veins very
irregular; lower surface of midrib minutely hairy,
upper surface more copiously so; sori fewer, rather
uregular; indusia apparently mostly reniform.

Distribution — Malesia: Sumatra (known from
type collection only).

Habitat — In forest at 150 m.

PSOMIOCARPA

Psomiocarpa Presl, Epim. Bot. (1851) 161, for type species only; Copel., Gen. Fil.
(1947) 125; Zamora & Chandra, Kalikasan 6 (1977) 219. — Polybotrya Humb. &
Bonpl. p.p., C. Chr., Index Filic. (1905) 590. — Type species: Psomiocarpa apii-

folia (Kunze) Presl.

Habit, scales and frond-form as Tectaria ramosii but fronds smaller and strongly di-
morphous; leaflets of fertile fronds much contracted, almost circular, their convex lower
surface densely covered with sporangia which obscure the small upper surface. — Fig. 17.

Distribution — Malesia: One species, widely distributed in the Philippines.

Cytology — Chromosome number n = 40 (see the species).

Holttum — Tectaria Group 101

Note — John Smith published the name Polybotrya apiifolia for Cuming 26, but no
description; Kunze published the ‘first description under the same name. Pres] proposed
the new generic name Psomiocarpa, including also three other species now generally re-
tained in Polybotrya, and stated that they differed from Polybotrya in bearing sporangia all
over the upper surface of fertile leaflets or lobes, though Kunze had stated that an upper
surface free from sporangia was visible. The species P. apiifolia was transferred to Acro-
stichum by Hooker and the genus was ignored by other authors until Copeland, who,
however, repeated the erroneous statement that sporangia are borne on both surfaces. In
1947 Copeland regarded Psomiocarpa as ‘in every respect except its dimorphism ... a
typical Crenitis’, but Zamora and Chandra noted resemblances to free-veined Tectaria
species and anatomical differences from Ctenitis (but no species of the latter mentioned).
The close relationship to Tectaria ramosii is confirmed by the existence of hybrids. Tec-
taria ramosii also hybridizes with T. aurita (Sw.) Chandra (type species of Stenosemia).
The retention of Psomiocarpa as a genus distinct from Tectaria is based on the very
specialized structure of its fertile leaflets, a structure which still needs a detailed study.

PJ.Eowarps

Fig. 17. Psomiocarpa apiifolia (Kunze) Presl. a. Outline of a sterile basal pinna, x 0.6; b. enlargement of
upper surface of a pinnule, x 4; c. outline of a fertile frond, x 0.6; h. enlargement of base of basal pinna,
x 20 (a, b: Matthew s.n., 28-11-1906; c, d: Price 1543, cult. Kew).

1. Psomiocarpa apiifolia (Kunze) Presl, Epim. Caudex short, erect; scales 3—4 mm long, nar-
Bot. (1851) 161; Copel., Fern Fl. Philipp. row, dark; fronds strongly dimorphous. Sterile
(1960) 298; Holttum, Blumea 30 (1984) 9, f. le, frond: stipe to 7 cm long, covered with hairs 1 mm
pl. ld. — Polybotrya apiifolia [J.Sm., J. Bot. long and very narrow scales; lamina to 13 cm long,
(Hooker) 3 (1841) 401, nom. nud.] Kunze, bipinnate above base, deltoid-pentagonal in out-
Famkrauter (1844) 142, pl.62.— Acrostichum _line; pinnate, pinnae to 6 pairs, middle ones bear-
aplifolium (Kunze) Hook., Sp. Fil. 5 (1864) ing 3 or 4 pairs of sessile or adnate deeply lobed
248. — Type: Cuming 26, Luzon. pinnules, basiscopic basal veins in larger adnate

102

Flora Malesiana ser. II, Vol. 2! (1991)

ones arising from the major axis; basal pinnae to
6.5 cm long, their basal basiscopic pinnules to 2.8
x 1.3 cm, very deeply lobed above a pair of free
basal leaflets; veins all free; thick hairs present be-
tween veins on upper surface. Fertile frond: stipe to
25 cm long; lamina to 10 cm long, shaped as ster-
ile but all parts much contracted; basal pinnae to
4 cm long, basal basiscopic pinnule to 1.2 cm
long, pinnules on acroscopic side to 1.5 x 1.5
mm, each with a narrow firm upper surface free
from sporangia and a swollen lower surface covered
with them, veins not detectable externally; spores
with finely reticulate crests. — Fig. 17.

Distribution — Malesia: Philippines.

Variability (see Holttum 1984) — M.G. Price,
who saw many plants in the field, reports that
there are two forms of this species, a larger one
with erect sterile fronds and a smaller one with
horizontal sterile fronds, also that spores of the
two differ. Herbarium specimens do not show this
difference clearly. Further field studies are needed.

Cytology — Chromosome number 41 was re-
ported by T.G. Walker in Bot. J. Linn. Soc., Suppl.
1, 67 (1973) 109, but Dr Walker in 1987 has
written to correct this to 40, thus agreeing with
Tectaria.

AENIGMOPTERIS

Aenigmopteris Holttum, Blumea 30 (1984) 3. — Type species: Aenigmopteris dubia
(Copel.) Holttum.

Caudex suberect; scales Tectaria-like; lamina of fronds much longer than wide, bipin-
nate; basal pinnae little longer than the next pair with elongate basal pinnules; middle pin-
nae with many subequal deeply lobed pinnules or pinna-lobes which are decurrent at their
bases to form a very narrow wing along the pinna-rachis; veins free, not reaching the
margin, pinnate in pinnules, the basal basiscopic veins not arising from the pinna-rachis;
scales on smaller axes few, linear or the smallest ovate, not thin nor with isodiametric
cells; scattered thick hairs present on upper surface of lamina between veins; small sub-
spherical unicellular glands present on lower surfaces of veins but not elongate cylindric
ones; sori at the apices of veins (distal ones sometimes apical on short lateral veins); in-
dusia firm, dark, persistent, of sori near the bases of lobes reniform, of distal sori vari-
ously asymmetric; sporangia sometimes bearing a short hair near the annulus; perispore
forming thin ridges or crests with spinulose margins, the ridges forming a few irregular
areoles, the surface within areoles bearing irregular short projections. — Fig. 18.

Distribution — Malesia: Five known species in the Philippines, Western New Guinea
and Borneo.

KEY TO THE SPECIES

la. Upper surface of pinna-rachis, at least distally, prominent and bearing short hairs;

scales of pinna-rachis broad at the base, tapering distally, not dark ... 1. A. dubia

b. Upper surface of pinna-rachis slightly grooved, the groove filled with many short
hairs; scalesion pinna-rachis/dark and narrow 4214-1421) 4 seen eee D

2a. Pinnules of middle pinnae c. 4 mm wide, their lobes c. 1 mm wide . 2. A. pulchra
b. Pinnules of middle pinnae c. 7 mm wide, their lobes c.2 mm wide.......... 3
3a. Thick hairs c. 1 mm long on lower surface of veins ...... 3. A. mindanaensis

bowihickhairs) lacking omlower surface Of VeInS@.6 74s ie oe ee 4

Holttum — Tectaria Group

103

4a. Pinnule-lobes on first suprabasal pinnae deeply lobed; the thick hairs present on up-

per surface rigid, more than 1 mm long, short hairs also present

. 4. A. katoi

b. Pinnule-lobes on first suprabasal pinnae not deeply lobed; thick hairs between veins
on upper surface not rigid, no other hairs present ............. 5. A. elegans

1. Aenigmopteris dubia (Copel.) Holttum,
Blumea 30 (1984) 4, f. a, pl. la. — Dryopteris
dubia Copel. in Elmer, Leafl. Philipp. Bot. 1
(1907) 235. — Ctenitis dubia (Copel.) Copel.,
Gen. Fil. (1947) 124; Fern FI. Philipp. (1960)
293. — Type: Elmer 9016, Luzon (MICH; iso
BORK):

Caudex suberect, bearing few fronds, its apex
covered with dull brown scales 10 x 1 mm; stipes
to 30 cm long, dark castaneous, bearing + persis-
tent scales throughout, also short ctenitoid hairs in
the groove; lamina to 55 cm long and 24 cm wide;
free pinnae to 7 pairs, adnate pinnae to 7 or more
pairs, the distal ones with very narrowly decurrent
bases merging into the apical lamina of the frond
the axis of which is very narrowly winged, as is
the greater part of every pinna-rachis; a bud some-
times present at the base of an upper pinna; basal
pinnae to 14 cm long bearing 4 pairs of free pin-
nules, rest of pinna lobed to a very narrow wing
along the rachis; basal basiscopic pinnule to 7.5
cm long, nearly 3 cm wide, bearing several pairs
of well-spaced tertiary leaflets, all deeply obliquely
lobed with narrowly decurrent bases; basal acro-
scopic pinnule to 3.5 x 1.1 cm, only its basal
lobes lobulate; second pinnae to 11 cm long with
1 pair free pinnules and others adnate; pinnules and
larger pinna-lobes of middle pinnae like tertiary leaf-
lets of basal pinnae; lower surface of pinna-rachis
bearing thick ctenitoid hairs 0.5—1 mm long and a
few ovate-acuminate scales, smallest scales ovate;
lower surface of pinna-lobes bearing a few thick
hairs on costules and veins; upper surface of pinna-
rachis prominent, glabrous, of lamina between veins
and of rachis-wing bearing scattered very thick
hairs 0.5—1 mm long; sori in lobules of pinna-
lobes, apical on veins; indusia firm, reniform, gla-
brous, those of distal sori asymmetric.

Distribution — Malesia: Philippines (S Luzon,
Negros, Leyte, Camiguin de Mindanao).

Habitat — In forest at 850-1200 m.

2. Aenigmopteris pulchra (Copel.) Holttum,
Blumea 30 (1984) 7. — Dryopteris pulchra

Copel., Univ. Calif. Publ. Bot. 18 (1942) 219.
— Ctenitis pulchra (Copel.) Copel., Gen. Fil.
(1947) 124; Philipp. J. Sci. 78 (1951) 410, pl.
14. — Type: Brass 13455, West New Guinea
(MICH; iso BM, BO, L, UC).

Stipe c. 30 cm long, dull castaneous, bearing
long hairs in the groove; basal scales 13 mm long,
less than 1 mm wide, very firm, dark; scales above
base gradually smaller; rachis bearing rather sparse
scales abaxially, adaxially hairs more than 1 mm
long; lamina to 45 cm long and 24 cm wide, firm;
free pinnae c. 15 pairs, contiguous, distal ones
grading into the apical lamina; basal pinnae to
13.5 cm long with 1 pair of free pinnules and sev-
eral with decurrent bases, all acuminate and deeply
lobed with well-spaced, narrow, oblique lobes
which are deeply and obliquely lobulate; basal basi-
scopic pinnule 5.2 x 1.6 cm, acuminate, its lobu-
late lobes 9 x 3.5 mm, their bases 4 mm apart,
their lobules 1 mm wide; basal acroscopic pinnule
3.4 x 1.4 cm; second pair of pinnae with subequal
basal pinnules; fourth pair of pinnae 12 x 3.5 cm;
pinnules of middle pinnae 4 mm wide, their lobes
1 mm wide; lower surface of pinna-rachis bearing
many thick hairs and sparse narrow scales, of cos-
tae of pinnules similarly but more sparse hairy, of
costules of pinna-lobes and on veins a few smaller
hairs and small spherical glands; upper surface of
pinna-rachis bearing many small hairs and scattered
very thick ones more than 1 mm long, similar long
hairs present on pinnule-lobes between the veins;
sori as in A. dubia; indusia sometimes papillate.

Distribution — Malesia: New Guinea (known
only from the Idenburg River; 2 collections).

Habitat — On banks of stream in forest at 850
and 1150 m.

3. Aenigmopteris mindanaensis Holttum,
Blumea 30 (1984) 7. — Type: Ramos & Edano
BS 39139, Mindanao, Bukidnon (BO; iso US).

Stipe 30 cm long; lamina 35 cm long, agreeing
with that of A. dubia but with pinnules or pinna-
lobes more closely placed and the pinna-lobes less

104

Flora Malesiana ser. II, Vol. 2! (1991)

PJ. Epwarps

Fig. 18. Aenigmopteris elegans Holttum. a. Basal part of a basal pinna, lower surface, x 4; b. fertile leaf-
let of a basal pinna, x 6.5 (a: Parris & Croxall 8612; b: Parris & Croxall 9135).

deeply lobulate; upper surface of pinna-rachis slight-
ly grooved and covered with very short hairs; long
thick hairs like those between the veins of leaflets
also occurring on the wing of the pinna-rachis and
sometimes on the margins of the rachis itself with
the short hairs; scales on pinna-rachis few, all
linear; sori as in A. dubia.

Distribution — Malesia: Philippines (known on-
ly from the type collection).

Note — Specimens of the type collection were
distributed from Manila as Dennstaedtia sp.

4, Aenigmopteris katoi Holttum, Blumea 30
(1984) 8, f. 1d. — Type: Kato, Murata & Mogea
B 3820, Kalimantan Selatan, G. Besar (K).

Caudex ¢. 8 mm diameter, bearing stipes in a
lax spiral; shape of pinnae and pinnules as in A.
mindanaensis; hairs on upper surface of leaflets
between veins very thick, rigid (cell-walls not col-
lapsing on drying), tapering to an acute apex, to
1 mm long and consisting of 5 or 6 cells, short
suberect hairs also present; no long hairs on lower
surface of costules and veins.

Distribution — Malesia: Borneo (known only
from the type collection).

Habitat — In forest, in humus-rich soil, 800-
1400 m.

5. Aenigmopteris elegans Holttum, Blumea
30 (1984) 8, f. 1b, c, pl. 1b. — Type: Parris &
Croxall 9135, Sabah, Mt Kinabalu (K).

Stipe 32 cm long, dark castaneous, bearing
throughout narrow dark scales and sparse rather
thin hairs to 1 mm long; basal scales to 12 mm
long, less than 1 mm wide, rachis similarly scaly;
lamina c. 40 cm long; free pinnae 4 pairs, then
c. 10 pairs with adnate decurrent bases, the upper
ones grading into the apical lamina in which the
deeply lobed lobes are connected by a narrow wing
along the rachis; a bud sometimes present at the
base of one or more pinnae; basal pinnae 12-13
cm long including stalk of 5 mm; basal basiscopic
pinnule to 6.5 x 2 cm, lobed to a narrow wing
throughout, the costules of its lobes (or tertiary
leaflets) 5 mm apart, the lobes oblique with obtuse
apex, to 11 x 4 mm, deeply lobulate with 3 or 4

Holttum — Tectaria Group

105

pairs of very oblique lobules 1 mm wide; the suc-
cessive basiscopic lobes of basal pinnae gradually
smaller and with less deeply lobulate lobes; basal
acroscopic pinnule 3.3 x 1.2 cm; middle pinnae
sessile, c. 10 x 3 cm, lobed throughout to a nar-
row wing, the lobes almost all deeply and oblique-
ly lobulate, those of the acroscopic and basiscopic
sides nearly equal, the basal acroscopic lobules
of the largest lobes only slightly crenate; the
lower surface of pinna-rachis bearing many short
hairs and a few narrow dark scales, of costae of
pinna-lobes sparsely short-hairy, of costules of lo-

bules bearing minute spherical glands; the upper
surface of pinna-rachis shallowly grooved and bear-
ing many short hairs with a few long ones, of the
surface of the lobes between veins bearing thick
flaccid hairs consisting of up to 8 cells; sori as in
A. dubia; sporangia sometimes bearing a very
short hair consisting of a few:cells, near the an-
nulus. — Fig. 18.

Distribution — Malesia: Borneo (Mt Kinabalu,
Sabah).

Habitat — In forest at 1400-1600 m, the type
from a ridge slope.

HETEROGONIUM

Heterogonium Presl, Epim. Bot. (1851) 142; Copel., Gen. Fil. (1947) 127; Holttum,
Sarawak Museum J. 5 (1949) 155; Kalikasan 4 (1975) 205. — Type species: Hetero-
gonium aspidioides Pres].

Tectaria auct: C. Chr., Index Filic. Suppl. 3 (1934) 176, p.p.

Ctenitopsis Ching, Bull. Fan Mem. Inst. Biol. 8 (1938) 304, p.p.

Ctenitis auct.: Copel., Gen. Fil. (1947) 123, p.p.

Stenosemia auct.: Copel., Fern Fl. Philipp. (1960) 301, p.p.

Caudex short, erect; stipe and rachis usually blackish; /Jamina simply pinnate with
deeply lobed pinnae; basal pinnae widest in the middle of the basiscopic side and on that
side narrowed to the base, rarely with a pair of free pinnules; veins free or forming a
series of costal and costular areoles, in species with broad pinnae more amply anastomos-
ing, free veins in the areoles none or few, never forked; basal basiscopic vein of each
vein-group, whether free or not, arising from the costa or from the very base of the cos-
tule; upper surface of rachis and costae bearing short multicellular hairs, longer thick mul-
ticellular hairs variously present on the upper surface on and between veins (often fewer
in fertile than in sterile fronds); fertile fronds usually with narrower pinnae than sterile, in
a few cases much contracted and covered beneath with sori; sori round with reniform in-
dusia, or exindusiate and then sometimes spreading along the veins, on free veins where
these are present in the pinna-lobes, or on connected veins. — Fig. 19.

Distribution — Burma to South China; Malesia; Mauritius; 22 known species.

Cytology — Chromosome number n = 41.

Taxonomy — In the 19th century the species here brought together were very various-
ly treated by different authors. Specimens of the type species were assigned to the genera
Stenosemia, Gymnogramma, Digrammaria, Phegopteris and Aspidium, of other species
to Nephrodium, Dryopteris, Lastrea, Polypodium, Acrostichum, Polybotrya and Pleo-
cnemia. As here construed (following Holttum 1949 and 1975) Heterogonium might rank
as a subgenus of Tectaria. As in Tectaria, it comprises species with free and anastomosing
veins, but differs constantly in the shape of basal pinnae and in the venation of species
which have broad pinnae, also in a more frequent tendency to the exindusiate condition

106 Flora Malesiana ser. II, Vol. 2! (1991)

with sori more or less spreading along veins, and to an apparently acrostichoid condition
where the fertile pinnae are greatly contracted.

Presl’s type species has the venation of Sagenia Presl with exindusiate sori spreading a
variable distance along the veins. The genus was ignored by other authors until Copeland
added another species in 1929. Copeland failed to notice the form of basal pinnae. Christen-
sen added two more species in 1934; these are here united as H. stenosemioides which,
having free veins, does not correspond to Copeland’s definition of 1947 and was ignored
by him.

In 1938 Ching made the first attempt to deal with Ctenitis in Asia. In the New World
Christensen had found that Tectaria and Ctenitis agree in having hairs of a peculiar type (a
character ignored in the 19th century) and remarked that they were distinct in the anasto-
mosing veins of Tectaria and the free ones of Ctenitis. But in Asia Ching found that there
were free-veined species which agreed with Tectaria, not Ctenitis, in some details of
venation, and proposed for them the new genus Crenitopsis. In it he included six species
which have basal pinnae as in Tectaria and two with basal pinnae of the Heterogonium
form here recognized; all those published before 1905 had been included in Dryopteris in
Christensen’s Index Filicum. Copeland (1947) included Ctenitopsis in Ctenitus, without
comment on basal pinnae.

Holttum (1949 and 1975) contended that the recognition of the form of basal pinnae as
a distinctive character separated Heterogonium as a natural group from Tectaria when all
relevant species are considered. But there is one known case of apparent hybridization be-
tween a species having basal pinnae as in Tectaria (T. aurita) with one having basal pinnae
as in Heterogonium (H. pinnatum). Experimental study of the relationships between these
and other species of the two genera is needed.

Note on using the key — In nearly all species the pinnae of fertile fronds are + reduced
as compared with pinnae of sterile fronds. In species which normally have much-con-
tracted fertile pinnae intermediates between this condition and sterile pinnae may occur,
and these may mislead users of the key. In some species also, notably H. profereoides,
young plants which do not show the normal frond form and size may be fertile.

KEY TO) THE’ SPECIES

tay Veins free or with ame gular slight/anastomosis: j. <)> «4-1-1 4 aoe 2
bs) Veins|anastomosing in’steriletfronds 23)... 24 s.5 eo seu eee 12
2a. Pinnae c. 3 cm long; veins 3 or 4 pairs, nearly all simple ..... 1. H. ceramense
b. Pinnae much longer; veins 6 or more pairs, mostly forked................ 3
3.aia5 OnlANGUSIate tein eS erro ie aS WR ee 4
bstSorvexindusiate yr. Re ein, GRINS SRR 8
Zan Hairs betweentveins scattered all overupper surbace =) 35.005 47 ee 5)
b. Hairs between veins on upper surface present only near sinuses and margins of pinna-
LODE Sires tesa. eet. RET AE Ptah OU Oe APE Cah. he 6. H. subglabrum

a, Pinna-lobes atmost.crenate sare. Li geese eee ae) Wie a tee 6

b. Pinna-lobes, except those of distal pinnae, distinctly lobulate ... 5. H. lobulatum

Holttum — Tectaria Group 107

6a.

Lamina of largest fronds 30—40 cm long; suprabasal fertile pinnae to 12 x 2.5 cm
2. H. sagenioides

b. Lamina of largest fronds 50—75 cm long; suprabasal fertile pinnae to 15 cm or more
en pare tai CINE WIDE» ice 6: ctieacesneds amsas enunleme- SAsenohss rsitelye. (oy Beistrer = Regewntn tee eet 7
(eStipesmusty Drown, MOt-SlOSSy 2... 06 6 es eee oe oe eo ots 3. H. rufescens
baestipesmeanly black, Slossy se ccejails els eo iis of adele chs) stele. 4. H. wigmanii
8a. Fertile pinnae not greatly contracted; most sori separate................. 10
b. Fertile pinnae with lobes much contracted ......... 10. H. stenosemioides
9a. Basal pinnae of fertile fronds bearing deeply lobed pinnules ... 7. H. wenzelii
be. Basalipinnae of fertile fronds wholly pinnatifid ..........)......5:-......... 10
lOQasdwobes of suprabasal pinnae entire or nearly SO: 3.25 22. hes wee we 11
b. Lobes of suprabasal pinnae distinctly crenate ........ 9. H. novoguineense
11a. Several pairs of pinnae with stalks 3—4 mm long; all pinnae lobed to 3—4 mm from
COSUUIMES MIE rr a mone aetta yeu) amr NNO ne aa oe ae 8. H. calcicola
bekinnae mostly sessile, lobed more deeply ...2.....-.7. 2. H. sagenioides
iZWaw oun fertile fronds with distinctly separate Som<.... 2. 20/2. 2 2h te 13
b. Young fertile fronds with sporangia running along all veins and at maturity covering
lowermsurbace, of pinna=lODES «2. aiey asc cies Ae aie eee vale areas ey Len: 18
13a. Young sori round, indusiate, on pinna-lobes 1.5—2.5 mm wide, completely cover-
ingplobesmwhenwmature 465.22 .k fee ee 11. H. alderwereltii
b. Young sori of varied form; if indusiate on much wider pinna-lobes which they do
OME OVE WIEN WAUUINS Gacy eens ete eateuataice Suet Seamus snd SEN Crcneeoa ee Cates narra 14
14a. Sori all irregularly elongate along veins, exindusiate................... NS
beesoralliround'to: elliptic, not linear, indusiate or not... 2. 2.4.5.4. 52. 16
15a. Suprabasal pinnae of mature plants to 13 x 2.5 cm, lacking costular areoles; no buds
OnwUppPEer sumrace Of Sterile fronds. 2). 9) 09.8 2 ee 12. H. aspidioides
b. Suprabasal fertile pinnae of mature plants to 25 x 5 cm; several pairs of costular
areoles present; buds present on upper surface of some costae of sterile fronds near
LIBRA GIS ars itty ee amt ae aden ayaa. Unto eke eA be eitaaetls 13. H. profereoides
16a. Sori in one row on each side of costules of pinna-lobes................. 17
b. Sori on each side of veins which run to crenatures on margins of pinna-lobes
15. H. pluriseriatum
17a. Basal pinnae of fertile fronds bearing 1 pair of free pinnules and 2 or 3 pairs adnate
7. H. wenzelii
b. Basal pinnae of fertile fronds lacking pinnules .......... 14. H. giganteum
18a. Fronds to 70 cm long; sterile pinnae to 21 x 4cm .... 16. H. teysmannianum
b. Fronds to 30 cm long; sterile pinnae much smaller ......... 17. H. pinnatum
1. Heterogonium ceramense Holttum, Kali- isomorphous; lamina to 17.cm long; free pinnae 9

kasan 4 (1975) 217 — Type: Buwalda 5711, pairs, short-stalked; basal pinnae slightly shorter
Central Ceram, Mt Solagor, 600 m (K; iso L). than those next above them, their middle width 1.1

cm; largest suprabasal pinnae 3 x 1 cm, lobed 2/3

Stipe to 10 cm long, slender, dark brown, near or more towards the costa, lobes slightly crenate
base bearing narrow scales 4—5 mm long; fronds distally; veins 3 or 4 pairs, mostly simple; lower

108

Flora Malesiana ser. II, Vol. 2! (1991)

surface bearing few hairs on costa and costules on-
ly; upper surface of costae densely hairy, a few
thick hairs present between veins especially near
sinuses; sori small, subterminal on veins; indusia
small, caducous.

Distribution — Malesia: Moluccas (Ceram; only
known from two collections).

Habitat — In limestone crevices, in light shade.

2. Heterogonium sagenioides (Mett.) Holt-
tum, Sarawak Museum J. 5 (1949) 161; Revis.
Fl. Malaya 2 (1955) 520, f. 306; Kalikasan 4
(1975) 211. — Aspidium sagenioides Mett.,
Farngatt. IV (1858) 113; Racib., Pteridop. Bui-
tenzorg (1898) 179. — Dryopteris sagenioides
(Mett.) Kuntze, Revis. Gen. Pl. (1891) 813;
Christ, Philipp. J. Sci. 2 (1907) Bot. 211;
Alderw., Malayan Ferns (1908) 191; Backer &
Posth., Varenfl. Java (1939) 41. — Ctenitopsis
sagenioides (Mett.) Ching, Bull. Fan Mem. Inst.
Biol. 8 (1939) 312. — Ctenitis sagenioides
(Mett.) Copel., Gen. Fil. (1947) 124; Fern Fl.
Philipp. (1960) 295. — Type: Zollinger 1803,
Java (no type at B; iso Z).

Nephrodium melanopus Hook., Sp. Fil. 4 (1862)
110, excl. pl. Ambon. — Lastrea melanopus
(Hook.) Bedd., Ferns Brit. India (1865) t. 38. —
Type: Parish 51, Burma, Moulmein (K).

Phegopteris obscura Christ ex Hook., Bull. Herb.
Boissier 6 (1898) 196, name only. — Polypo-
dium obscurum Hook., Sp. Fil. 4 (1862) 237,
non Mett. 1857; Alderw., Malayan Ferns (1908)
492. — Nephrodium obscurum (Hook.) Diels in
E. & P., Nat. Pflanzenfam. I, 4 (1899) 173. —
Dryopteris sagenioides subsp. obscura C. Chr.,
Index Filic. (1905) 290. — Dryopteris obscura
(Hook.) Christ, Philipp. J. Sci. 2 (1907) Bot.
214, not Kuntze. — Ctenitopsis obscura (Hook.)
C. Chr., Notul. Syst. (Paris) 7 (1938) 87. —
Ctenitis obscura (Hook.) Copel., Fern Fl. Phi-
lipp. (1960) 287, for Cuming 302 only. —
Heterogonium obscurum (Hook.) Holttum, Kali-
kasan 4 (1975) 218, f. 1. — Type: Cuming 302,
Leyte (K).

Polypodium viscosum C.H. Wright, Kew Bull.
(1906) 12. — Dryopteris laokaiensis C.Chr.,
Index Filic. Suppl. 1 (1913) 34. — Type: E.H.
Wilson 36, Tonkin (K).

Phegopteris schizoloma Alderw., Bull. Jard. Bot.
Buitenzorg II, 16 (1914) 24; Malayan Ferns,

Suppl. (1917) 306. — Dryopteris schizoloma
(Alderw.) C. Chr., Index Filic. Suppl. 2 (1917)
16. — Type: Amdjah 595, E Kalimantan (BO).

Stenosemia aurita auct.: Hook., Gen. Fil. (1842)
t. 94, for f. 5 & 6 only.

Aspidium membranifolium auct.: Kunze, Bot. Zeit.
(Berlin) 6 (1848) 261.

Phegopteris philippinensis auct.: Mett., Farngatt.
IV (1858) 26, for var. 2 only.

Heterogonium teysmannianum auct.: Holttum,
Reinwardtia 1 (1950) 29, for Cuming 302 only.

Heterogonium gurupahense auct.: Holttum, Rein-
wardtia 3 (1955) 271, for pl. ex Malaya et Thai-
land only.

Stipe to c. 30 cm long, dark, above base min-
utely hairy on abaxial surface, basal scales thin, to
c. 10 x 1.5 mm; fronds slightly dimorphous, those
with broadest pinnae always sterile; lamina com-
monly 30—35 cm long (to 45 cm); pinnae 10-15
pairs, sessile or nearly so; suprabasal pinnae of fer-
tile fronds to 12 x 2.5 cm, lobed to 1-2 mm from
the costa, basal acroscopic lobes sometimes elon-
gate and crenate, other lobes obtuse with + sinuous
margins, of sterile fronds to 15 x 3.5 cm, lobes
often entire or nearly so; basal basiscopic lobes of
basal pinnae always much shorter than acroscopic
ones, middle lobes on basiscopic side longer than
acroscopic ones, the largest 3.5 cm long and cre-
nate-lobate with pinnate veins in the lobes, middle
acroscopic lobes often crenate-lobate on fertile
fronds; veins mostly forked except distal ones;
lower surface variably hairy on costae, veins and
between veins, costal hairs on sterile fronds often
more than 1 mm long, fertile fronds always less
hairy, sometimes almost glabrous; upper surface
of sterile fronds in most cases + copiously short-
hairy between veins with + abundant long thick
hairs also, fertile fronds less hairy than sterile, not
always with some thick hairs between veins; ster-
ile fronds of lowland plants sometimes bearing
thick hairs only between veins; sori terminal on
veins, usually on acroscopic branches of forked
veins; indusia brown, thin, glabrous, soon shrivel-
ling, or lacking. — Fig. 19a, b.

Distribution — Thailand, S Burma, Vietnam,
Hainan; Malesia: West (most abundant in the Pe-
ninsula), Philippines.

Habitat — In forest at c. 0-800 m.

Notes — Philippine specimens are mostly small-
er than those of the Malay Peninsula; some quite

Holttum — Tectaria Group

109

small ones are fertile and very hairy, perhaps from
exposed places.

The only exindusiate specimens known are the
types of Polypodium obscurum Hook. and Phego-
pteris schizoloma Alderw. I see no clear distinc-
tion, other than lack of indusia, between these and
the range of indusiate specimens as above described.

There was much confusion in the synonymies
of various authors, due to various causes discussed
by Holttum in 1975 under H. obscurum. The chief
were Hooker’s mixture of drawings from different
specimens under Stenosemia aurita (1842), the cita-
tion of Phegopteris obscura Fée as a basionym
though a nomen nudum, and the fact that no author
besides Hooker saw specimens of Cuming 302.
Basal pinnae on the specimens of Cuming 302 from
Herb. Hooker at Kew are nearly all defective, and
those of the sterile frond in the photograph repro-
duced as fig. 1 by Holttum in 1975 are lacking.

3. Heterogonium rufescens Holttum, Kalika-
san 4 (1975) 215. — Type: A.C. Jermy 7863,
New Ireland, Danfu Valley (BM).

Stipe and rachis castaneous, minutely hairy on
abaxial surface, basal scales to 15 mm long, hardly
1 mm wide; fronds slightly dimorphous, the lobes
of fertile pinnae narrower and more widely spaced
than those on sterile fronds; lamina to 75 cm long;
pinnae to c. 20 pairs, to 20 cm long, lobed almost
to the costa; basal pinnae to 7 cm wide in the
middle, the longest basiscopic lobes 6 cm long,
very deeply lobulate; suprabasal pinnae to 4 cm
wide, costules 6—8 mm apart (most widely in
sterile fronds), fertile lobes 3—4 mm wide, crenate,
sterile ones to 6 mm wide, crenate-lobate; veins
in lobes almost all once forked (paucipinnate in
largest lobules of basal pinna-lobes); lower surface
of costae of sterile fronds bearing dense short pale
hairs, upper surface between veins bearing many
short slender hairs and some long thick pale ones,
surfaces of fertile pinnae similar but bearing very
few thick hairs between veins on the upper surface;
sori subterminal on the acroscopic branches of
forked veins; indusia thin, light brown.

Distribution — Malesia: New Guinea (Admiralty
Is., New Ireland, Bougainville).

Habitat — In forest at 250-800 m, the type ‘on
well-drained slopes of stiff loam mixed with lime-
stone chips’, a specimen from Los Negros island
“at base of limestone cliff’.

4. Heterogonium wigmanii (Racib.) Holt-
tum, Reinwardtia 3 (1955) 271. — Aspidium
wigmanil Racib., Bull. Int. Acad. Sci. Craco-
vie (1902) 61. — Dryopteris wigmanii (Racib.)
C. Chr., Index Filic. (1905) 301; Alderw., Ma-
layan Ferns (1908) 192. — Type: Cult. Hort.
Bog. 2K.XI1.13, origin Aru Islands (BO; iso L).

Dryopteris sagenioides vat. gurupahensis C. Chr.,
Svensk Bot. Tidskr. 16 (1922) 95, Fig. 2. —
Dryopteris gurupahensis (C. Chr.) C. Chr., Bot.
Jahrb. Syst. 66 (1934) 45. — Heterogonium
gurupahense (C. Chr.) Holttum, Reinwardtia 3
(1955) 272, quoad pl. typ. tantum. — Type:
Kaudern 17, 18, N Celebes, Bolaang Mongon-
dow, Gurupahi (S-PA; iso BM).

Similar to H. rufescens in size and form of
fronds; differing: stipe and rachis very dark and
glossy on the abaxial surface; lobes of both sterile
and fertile pinnae at most crenate; lower surfaces of
costae of sterile pinnae bearing sparse slender ru-
fous hairs 1 mm long, upper surface lacking short
hairs between veins. — Fig. 19c.

Distribution — Malesia: Celebes, Moluccas
(Morotai, Ceram, Aru Is.).

Habitat — In forest, 0Q—-1000 m; on ravine slopes
in limestone area (Ceram).

5. Heterogonium lobulatum Holttum, Blu-
mea 35 (1991) 557. — Type: Price & Hernaez
399, Philippines, W Samar (K; iso MICH).

Closely similar to H. sagenioides in frond-
form, venation and pubescence but possibly less
distinctly dimorphous, differing in the distinctly
lobulate lobes of all pinnae, those on the basiscop-
ic side of basal fertile pinnae most deeply so with
veins pinnate in each lobule; lobes of suprabasal
pinnae lobulate up to halfway towards costules,
the lobules mostly falcate, acute.

Distribution — Malesia: Philippines (known on-
ly from the type collection).

Habitat — In primary forest on limestone and
limestone-derived soils.

6. Heterogonium subglabrum Holttum, Rein-
wardtia 3 (1955) 273; Kalikasan 4 (1975) 216.
— Dryopteris sagenioides forma contracta Al-
derw., Bull. Jard. Bot. Buitenzorg III, 2 (1920)
147. — Type: Lérzing 5520, Sumatra (BO).

110 Flora Malesiana ser. II, Vol. 2! (1991)

P.J.Epwaaps

Fig. 19. Heterogonium sagenioides (Mett.) Holttum. a. Outline of a fertile basal pinna, x 0.6; b. detail of
one median acroscopic lobe of a, x 4. — H. wigmanii (Racib.) Holttum. c. Two median lobes of a middle
pinna, x 4. — H. pinnatum (Copel.) Holttum. d. Outline of a sterile basal pinna, x 0.6; e. fertile lobes
with sori removed from right lobe to show attachment areas, x 4; f. detail of a basiscopic lobe of d, x 4.
— H. profereoides (Christ) Copel. g. Fertile lobes of a middle pinna with sori removed from right lobe, x
4 (a, b: Matier s.n., Penang; c: Kato 7160; d—f: Cuming 237; g: Elmer 10807).

Holttum — Tectaria Group

Closely allied to H. sagenioides, differing con-
stantly in glabrous lower surface apart from rare
short hairs on costules, and the presence of thick
hairs on the upper surface confined to short ones
near sinuses and very near the margins of lobes;
fronds of largest plants differing from those of /7.
sagenioides in broader pinnae which are less deeply
lobed, the lobes broader and acute, those of the
largest fertile suprabasal pinnae distinctly crenate
and subacuminate.

Distribution — Malesia: Sumatra (northern; not
known south of Sibolangit).

Habitat — At low altitudes in forest.

7. Heterogonium wenzelii (Copel.) Holttum,
comb. nov. — Ctenitis wenzelii Copel., Phi-
Lipps We Semis1 (1952). 25; pl. 18; Fern FI.
Philipp. (1960) 296. — Type: Wenzel 165,
Leyte (US; iso MICH).

Stipe to 50 cm long, dark, minutely hairy above
base, near base bearing scales 8 x 1 mm and longer
hairs; sterile lamina not known; fertile lamina to
35 cm long; basal pinnae to 15 cm long with
stalks 10-12 mm, bearing a pair of free pinnules
and 2 or 3 pairs adnate; basal basiscopic pinnule
3.5 x 0.9 cm, deeply lobed, longest basiscopic
pinnule 5 cm long, 4 mm wide, lobes on distal
part of pinnae much narrower, lobulate, widely
spaced and connected by a narrow wing along the
costa; second pair of pinnae 10 cm or more above
basal ones, shaped as distal part of basal pinnae;
veins not visible on the surface, forming narrow
areoles along the costa on distal parts of pinnae,
free and forked in pinna-lobes; lower surface of
costae and costules bearing sparse short hairs; up-
per surface of costae densely hairy, of costules less
sO, a few hairs present near sinuses; sori exindusi-
ate, mostly round, on veins in lobes of pinnae,
those on basal veins sometimes elongate.

Distribution — Malesia: Philippines (known on-
ly from the type collection).

Note — Copeland failed to observe the anasto-
mosis of veins. It is probable that sterile fronds
(not yet known) have wider pinnae than fertile
ones, with wider lobes, and veins anastomosing all
along the costae.

8. Heterogonium calcicola K. Iwatsuki & M.
Kato, Acta Phytotax. Geobot. 34 (1983) 133.

— Type: Kato, Okamoto & Ueda B-11680, Ka-
limantan Timur, 1°30' N, 117°30'E (KYO).

Stipe to 32 cm long, dark but not glossy, scales
on basal 15 cm 10 x 1.5 mm; lamina to 48 cm
long, texture very firm, small veins not visible on
surface, fertile and sterile fronds almost alike; pin-
nae well-spaced, to 12 pairs, all stalked except c. 2
distal adnate pairs, stalks of lower pinnae 5 mm
long; basal pinnae a little reduced, to 3 cm wide,
not greatly widened on the basiscopic side; supra-
basal pinnae to 9 x 2.5 cm, lobed to 3-4 mm
from the costa, base truncate, apex rather abruptly
narrowed, lobes quite entire, c. 6 mm wide at their
bases; veins all free, simple or once forked; lower
surface glabrous apart from some sparse minute
hairs near bases of costae; upper surface of costae
densely short-hairy, a few very short hairs also
present near sinuses; sori exindusiate, usually not
at apices of veins, present on both branches of
forked veins, those near apices of pinna-lobes
round, those on basal acroscopic veins consider-
ably elongate.

Distribution — Malesia: Borneo (known only
from the type locality; with 2 additional collec-
tions).

Habitat — On mountain slopes at 150-1000 m,
in calcareous area.

9. Heterogonium novoguineense Holttum,
Kalikasan 4 (1975) 221. — Type: New Guinea,
without locality or collector’s name (L 908,304-
112, -113, -114).

Stipe to 30 cm long, dark-castaneous, bearing
sparse short hairs abaxially; sterile fronds unknown;
fertile /amina to 35 cm long, bearing 13 pairs of
free pinnae; basal pinnae 10 cm long with stalks
2-3 mm, 2.5 cm wide in the middle, basal basi-
scopic lobe 5 mm long, acroscopic 10 mm, long-
est basiscopic lobe 15 mm long, crenate; supra-
basal pinnae to 10 x 2.2 cm, near their bases lobed
to 1 mm or less from the costa, the lobes well
separated, at most slightly crenate distally (apart
from the basal acroscopic ones); costules 4.5—5
mm apart; veins 6 pairs, forked except the distal
ones; lower surface quite glabrous; upper surface of
costae densely short-hairy, a few thick hairs pres-
ent between veins near the margin, especially at
sinuses; sori exindusiate, terminal on acroscopic
vein-branches.

mi

Flora Malesiana ser. II, Vol. 2! (1991)

Distribution — Malesia: New Guinea, apart from
the type, Biak Island.

Habitat — On limestone (Biak).

Note — Specimens of the type collection were
annotated by Zippel, and were probably collected
by him; he is known to have collected ferns on
limestone in western New Guinea.

10. Heterogonium stenosemioides (Baker)
C.Chr., Index Filic. Suppl. 3 (1934) 111;
Holttum, Reinwardtia 1 (1950) 30; Kalikasan 4
(1975) 223. — Acrostichum (Polybotrya) ste-
nosemioides Baker, J. Linn. Soc. Bot. 22
(1886) 230. — Polybotrya stenosemioides
(Baker) Copel., Polypod. Philipp. (1905) 40;
Alderw., Malayan Ferns (1908) 725. — Type:
G.F. Hose 158, Sarawak, G. Matang (K).

Polybotrya nieuwenhuisii Racib., Bull. Int. Acad.
Sci. Cracovie (1902) 57; Alderw., Malayan
Ferns (1908) 724. — Heterogonium nieuwen-
huisii (Racib.) C. Chr., Index Filic. Suppl. 3
(1934) 111. — Type: Cult. Hort. Bog. ex Bor-
neo leg. Nieuwenhuis, 2K. XIII.2 (BO).

Polybotrya nieuwenhuisii var. brookii Alderw.,
Bull. Jard. Bot. Buitenzorg II, 23 (1916) 19. —
Type: Brooks 195 S, Sumatra (BO, BM).

Stipe castaneous to very dark, of fertile fronds
40 cm or more long, shorter on sterile, minutely
hairy throughout on abaxial surface, basal scales
narrow, to more than 10 mm long. Sterile frond:
lamina to 35 cm long; pinnae to c. 12 pairs; mid-
dle lobes of basal pinnae little longer on basi-
scopic than on acroscopic side, the largest on both
sides often + crenate with some forked veins; supra-
basal pinnae to about 12 cm long and 3 cm wide,
lobed 3/5-—3/4 towards the midrib, the basal
lobes of several pinnae narrowed at their bases,
their apices + acute and margins + crenate; veins
in other lobes mostly simple; lower surface of cos-
tae rather closely hairy, hairs variable in length,
short slender hairs also + abundant on and between
veins (few on the type), upper surface of costae
densely short-hairy, scattered long thick hairs +
abundant between veins. Fertile frond: pinnae to
c. 8 cm long, usually less than 1 cm wide, very
deeply lobed, lobes commonly 2 mm wide; sori all
along veins, at maturity completely covering low-
er surface; fronds of intermediate form may occur.

Distribution — Malesia: Borneo, S Sumatra.

Habitat — In lowland forest.

11. Heterogonium alderwereltii Holttum,
Sarawak Museum J. 5 (1949) 163; Revis. Fl.
Malaya 2 (1955) 522, f. 305; Kalikasan 4 (1975)
229; new name for Pleocnemia stenosemioides
Alderw., Bull. Jard. Bot. Buitenzorg II, 2
(1920) 164, non Christ 1934. — Tectaria ste-
nosemioides (Alderw.) C.Chr., Index Filic.
Suppl. 3 (1934) 184. — Type: Lérzing 5161,
Sumatra, Sibolangit (BO).

Pleocnemia membranifolia auct.: Bedd., Handb.
Ferns Brit. India, Suppl. (1892) 48, p.p.

Differs from H. pinnatum as follows: both ster-
ile and fertile fronds have up to 3 pairs of stalked
or sessile pinnae and to 3 pairs adnate, and in both
there is irregularity in the formation of costal
areoles which in part or whole of a pinna may be
lacking, especially in the narrower fertile pinnae;
lamina of fertile fronds to 30 cm long with a scaly
bud present sometimes at the base of an upper pin-
na (in H. pinnatum such buds only occur on sterile
fronds); suprabasal fertile pinnae 10-15 mm wide,
lobed to 1-2 mm from the costa, lobes to 2.5 mm
wide; sori at first round and clearly separate, termi-
nal on veins, indusiate; indusia thin, soon shrivel-
ling.

Distribution — Peninsular Thailand; Malesia: N
Sumatra, Malay Peninsula.

Habitat — All records from limestone, or lime-
stone areas.

Note — This species always grows in association
with H. pinnatum and may be of hybrid origin,
possibly H. pinnatum x H. sagenioides, in which
case it would be a triploid. Experimental evidence
is desirable. The type is smaller than the Peninsu-
lar specimens from which the above description is
drawn but otherwise agrees with them.

12. Heterogonium aspidioides Presl, Epim.
Bot. (1851) 143; C. Chr., Gard. Bull. Sing. 7
(1934) 258; Holttum, Sarawak Museum J. 5
(1949) 164; Copel., Fern Fl. Philipp. (1960)
300; Holttum, Novit. Bot. Delect. Sem. Horti
Bot. Univ. Carol. Prag. 1968 (1969) 35; Kali-
kasan 4 (1975) 224, excl. syn. Aspidium pro-
fereoides Christ. — Tectaria aspidioides (Presl)
Copel., Philipp. J. Sci. 38 (1929) 137. —
Type: Cuming 295, Leyte (PRC; iso BM, K).

Gymnogramma ambigua Hook., Sp. Fil. 5 (1864)
150. — Digrammaria ambigua auct.: Presl, Tent.
Pterids (1836) t21Ves lS Wi ap spsexcl:

Holttum — Tectaria Group

Asplenium ambiguum Sw. — Tectaria ambigua
(Hook.) Copel., Philipp. J. Sci. 2 (1908) Bot.
415. — Dictyopteris ambigua (Hook.) Alderw.,
Malayan Ferns (1908) 521. — Type: Cuming
154, Luzon (K; iso BM).

Gymnogramma philippinensis Fée, Mém. Foug.
5. Gen. Filic. (1852) 181. — Phegopteris phi-
lippinensis (Fée) Mett., Farngatt. IV (1858) 26,
excl. 2nd var. — Type: Cuming 321, Samar
(iso BM, K).

Stenosemia aurita auct.: J. Sm. in Hook., Gen. Fil.
(1842) t. 94, for Cuming 295, 321 and figs.
1-4 only.

Lastrea spectablis auct.: J. Sm., J. Bot. 3 (1841)
412, for Cuming 154 only.

Fronds dimorphous. Sterile frond: stipe to at
least 35 cm long, not very dark, minutely hairy
throughout, basal scales to c. 12 x 1.5 mm; la-
mina to at least 50 cm long; free pinnae to 10
pairs and 2 or 3 pairs adnate, lower pinnae dis-
tinctly stalked; largest suprabasal pinnae 17 x 3.5
cm, near their bases lobed 3/4 towards costa, less
deeply distally; costules to 8 mm apart; lobes en-
tire, obtuse; veins anastomosing in | row of costal
areoles with few additional areoles below sinuses,
costular areoles rare; basal pinnae to 5 cm wide,
lobed almost to the costa at their bases, lobes all
entire, costular areoles lacking or 1 or 2 in longest
lobes; lower surface of costae bearing sparse slender
hairs to 1 mm long; upper surface of costae bear-
ing dense thicker hairs of variable length, to more
than 1 mn, thick hairs scattered all over upper sur-
face between veins; buds lacking on. upper surface
of costae. Fertile frond: stipe to 60 cm long; lamina
to 50 cm long; suprabasal pinnae to 14 x 2.5 cm,
lobed to c. 2 mm from costa or more deeply near
their bases, basal lobes sometimes + elongate and
rarely crenate; costal areoles present in widest pin-
nae, usually lacking distally and sometimes almost
wholly lacking; costular areoles rare; veins in
lobes often forked; basal pinnae wider, their middle
lobes rarely crenate; hairs on both surfaces of cos-
tae short, no hairs present between veins on up-
per surface; sori exindusiate, spreading along basal
half or more of veins, on both branches of forked
veins.

Distribution — Malesia: Philippines (Luzon,
Mindoro, Leyte, Samar), Central Celebes, doubt-
fully in Borneo.

Habitat — In forest at altitudes to 700 m.

113

13. Heterogonium profereoides (Christ)
Copel., Univ. Calif. Publ. Bot. 16 (1929) 61;
C. Chr., Gard. Bull. Straits Settlem. 7 (1934)
259; Holttum, Sarawak Museum J. 5 (1949)
164; Copel., Fern Fl. Philipp. (1960) 300.
— Aspidium profereoides Christ, Philipp. J.
Sci. 2 (1907) Bot. 158. — Pleocnemia_profer-
eoides (Christ) Alderw., Malayan Ferns (1908)
812. — Type: Copeland 1467A, Mindanao
(MICH).

Aspidium subaequale Rosenst., Feddes Rep. Spec.
Nov. Regni Veg. 12 (1913) 176. — Tectaria
subaequalis (Rosenst.) Copel., Philipp. J. Sci.
9 (1914) Bot. 5. — Dictyopteris subaequalis
(Rosenst.) Alderw., Malayan Ferns (1908) 323.
— Heterogonium subaequale (Rosenst.) Holt-
tum, Reinwardtia 3 (1955) 274. — Type: Keys-
ser 107, NE New Guinea, Sattelberg (B).

Heterogonium aspidioides auct.: Holttum, Kalika-
san 4 (1975) 224, p.p.

Caudex erect, to at least 15 cm tall; fronds vari-
ably a little dimorphous; stipe to at least 50 cm
long, not very dark, minutely hairy; basal scales
firm, to almost 2 mm wide, scales above base
early caducous; lamina to at least 80 cm long; pin-
nae to at least 12 pairs, lobed apical lamina rela-
tively short; suprabasal pinnae to 23 x 5 cm, those
fully fertile more deeply lobed and with more wide-
ly separated lobes than those with few sori (wholly
sterile pinnae rare); costules on largest fronds 10—
12 mm apart; lobes entire, obtuse, or the more
fully fertile ones + attenuate with sinuous margins;
basal pinnae very deeply lobed near their bases, the
basal pair of lobes sometimes almost free; veins
forming costal areoles and many additional ones
below sinuses between lobes, in the lobes many
pairs of costular areoles and usually some between
these and the margin; lower surface of costae gla-
brescent, upper surface densely covered with short
hairs, a few hairs present near sinuses between
lobes on upper surface; scaly buds often present on
upper surface of costae near their bases; sori exin-
dusiate, in one row on each side of costules, also
very irregularly between costae and sinuses be-
tween lobes, variously elongate along veins, al-
most all on connected veins, a few on free veins
near apices of lobes. — Fig. 19g.

Distribution — Malesia: Borneo (Sabah, Sara-
wak), Philippines (Mindanao, Mindoro), New Gui-
nea.

114

Flora Malesiana ser. II, Vol. 2! (1991)

Habitat — In lowland forest and to 1000 m; in
the G. Mulu National Park, Sarawak, often in
limestone areas.

Note — The above description is based on the
largest plants from G. Mulu, but much smaller
plants with pinnae 10 cm long may be fertile, es-
pecially on limestone. One collection from Mt
Kinabalu has narrower fertile pinnae, little different
from those of H. aspidioides but with sori almost
all on connected veins. The larger specimens are
intermediate between H. aspidioides and H. gigan-
teum. They need experimental treatment to arrive
at a better understanding of this relationship. The
presence of scaly buds on the upper surface of cos-
tae is notable.

14. Heterogonium giganteum (Blume) Holt-
tum, Sarawak Museum J. 5 (1949) 166; Kali-
kasan 4 (1975) 226. — Aspidium giganteum
Blume, Enum. Pl. Javae (1828) 159; Hook.,
Sp. Fil. 4 (1862) 50, p.p.; C. Chr., Index Filic.
(1905) 75, p.p. — Pleocnemia gigantea (Blume)
Presl, Epim. Bot. (1851) 259; Racib., Pteridop.
Buitenzorg (1898) 193; Alderw., Malayan Ferns
(1908) 173, excl. syn. — Sagenia gigantea
(Blume) T. Moore, Index Filic. (1858) 92. —
Nephrodium giganteum (Blume) Baker, Syn.
Fil. (1867) 300, p.p. — Dryopteris gigantea
(Blume) Kuntze, Revis. Gen. Pl. 2 (1891) 812.
— Tectaria gigantea (Blume) Copel., Philipp.
J. Sci. 2 (1908) Bot. 410; Backer & Posth., Va-
renfl. Java (1929) 72 [not sensu Ching, Sinen-
sia 2 (1931) 17]. — Type: Blume s.n., Java (L).

Aspidium saxicola Blume, Enum. PI. Javae (1828)
160. — Dictyopteris saxicola (Blume) Alderw.,
Malayan Ferns (1908) 515. — Heterogonium
saxicola (Blume) Holttum, Sarawak Museum J.
5 (1949) 165, f. 2; Revis. Fl. Malaya 2 (1955)
525, f. 308. — Type: Blume s.n., Java (L).

Polypodium petrophyum Blume, FI. Javae Filic.
(1829) 163, t. 71. — Dictyopteris petrophya
(Blume) T. Moore, Index Filic. (1861) 318. —
Phegopteris petrophya (Blume) Mett., Ann.
Mus. Bot. Lugd.-Bat. 1 (1864) 225. — Type: as
for Aspidium saxicola Blume.

Polypodium megalocarpum Hook., Sp. Fil. 5
(1864) 102.—Dictyopteris megalocarpa (Hook.)
J. Sm., Hist. Fil. (1875) 195. — Phegopteris
megalocarpa (Hook.) Salomon, Nomencl. Ge-
fasskrypt. (1883) 276. — Pleocnemia megalo-

carpa (Hook.) Bedd., Handb. Ferns Brit. India,
Suppl. (1902) 48; Alderw., Malayan Ferns
(1908) 172. — Tectaria megalocarpa (Hook.)
C. Chr., Index Filic. Suppl. 3 (1934) 182. —
Type: T. Lobb s.n., Java (K).

Nephrodium oligodictyon Baker, Ann. Bot. (Lon-
don) 5 (1891) 328. — Type: Hutton, Malay Is.
ex Herb. Veitch (K); probably from Java.

Tectaria elliptica Copel., Philipp. J. Sci. 9 (1914)
Bot. 228. — Dictyopteris elliptica (Copel.) Al-
derw., Malayan Ferns Suppl. (1917) 322. —
Type: C.J. Brooks 81S, Sumatra, Lebong Tan-
dai, Benkoelen (MICH; iso BM).

Dictyopteris distincta Alderw., Bull. Jard. Bot.
Buitenzorg II, 5 (1922) 193. — Type: Cult.
Hort. Bot. Bog. ex Sibolangit, Sumatra (BO).

Tectaria calcicola Copel., Brittonia 1 (1931) 72,
pl. 1. — Type: Clemens 20745, Sarawak, Bidi
cave (UC; iso K, NY).

Fronds of largest plants differing from largest
fronds of H. profereoides as follows: apical lamina
commonly 20 cm long, deeply many-lobed, grad-
ing to adnate pinnae; pinnae rarely more than six
pairs, sometimes all slightly adnate except the
basal ones, lacking buds on upper surface of cos-
tae, largest to 28 x 6 cm, lobed near their bases to
3 mm from costa (sterile ones less deeply), lobes
of the largest sometimes slightly crenate; basal
pinnae short-stalked, distinctly shorter than those
next above them, middle basiscopic lobes of larg-
est sometimes much elongate and lobulate; sori
sometimes almost circular, usually + elliptical and
attached by a short linear receptacle, indusiate or
not, the indusia (if present) persistent but not cov-
ering the whole sorus.

Distribution — Malesia: Western, Moluccas.

Habitat — In forest, to 1000 m or more, on
stream banks and on steep slopes in ravines, ap-
parently sometimes on limestone.

Note — I see no distinction in other characters
between fronds with indusiate and exindusiate sori.
I have seen distinctly crenate lobes of suprabasal
pinnae only in Blume’s type collection. One col-
lection from limestone in Sarawak (Paul & Erwin
S 27482) has somewhat elongate indusiate sori,
several pairs of stalked pinnae and a bud on the
upper surface of the costa of one of them; it is per-
haps a hybrid. Of six known collections from the
Malay Peninsula only one is indusiate. A widely
distributed species but nowhere abundant.

Holttum — Tectaria Group

115

15. Heterogonium pluriseriatum Holttum,
Kalikasan 4 (1975) 228. — Type: C.G. Mat-
thew s.n., 1913, Sumatra, G. Singgalang (K).

Suprabasal pinnae to 9 cm apart, to 32 cm long
and 9 cm wide, lobed to 5—6 mm from costae;
costules c. 1.7 cm apart along costae; lobes crenate,
narrowed in distal 2 cm to an almost acute tip,
c. 10 mm wide, the sinuses between them broadly
rounded; veins forming narrow areoles along costae
with one series of small additional areoles below
sinuses, also a regular series of costular areoles
connecting the veins which run from costules to
the marginal crenatures, the veinlets from such
veins also sometimes anastomosing; upper surface
of costae and costules closely short-hairy, surfaces
otherwise glabrous; sori small, round or nearly so,
exindusiate, on veinlets on each side of the small
veins which run from costules to crenatures.

Distribution — Malesia: Sumatra (West, 2 coll.).

Habitat — In forest, the type at 1200 m.

Note — The type consists of 3 suprabasal pinnae
only. The second collection, Alston 14315, above
Indrapura, is a smaller specimen.

16. Heterogonium teysmannianum (Baker)
Holttum, Reinwardtia 1 (1950) 29, excl. syn.
Polypodium obscurum Hook.; Kalikasan 4
(1975) 223. — Acrostichum (Stenosemia) teys-
mannianum Baker in Becc., Malesia 3 (1886)
56. — Stenosemia teysmanniana (Baker) Diels
in E. & P., Nat. Pflanzenfam. I, 4 (1899) 198;
Alderw., Malayan Ferns (1908) 727. — Type:
Teijsmann 10696, Sumba (BO; iso K, L).

Stipe to at least 50 cm long, dark and glossy
above the base, scales on basal part to 10 x 1 mm;
fronds very dimorphous. Sterile frond: lamina to
70 cm or more long; suprabasal pinnae to 21 x 4
cm, lobed to 2—2.5 mm from the costa, lobes
well spaced, mostly entire (rarely slightly crenate),
narrowed distally; basal veins in lobes usually free,
but sometimes forming areoles along costae; veins
above the base forked once or twice, often forming
areoles along costules; erect hairs 0.1 mm long
abundant on lower surface of costae. Fertile frond:
pinnae to 2 cm wide, lobed to less than 1 mm from
the costa, lobes 2 mm wide, widely separate; veins
forming one narrow areole on each side of cos-
tules; sporangia densely arranged along all veins.

Distribution — Malesia: Lesser Sunda Islands
(Sumba, Flores, Babar).

Habitat — A specimen from Babar grew on coral
rock, in shade; no information with type.

17. Heterogonium pinnatum (Copel.) Holt-
tum, Sarawak Museum J. 5 (1949) 163; Revis.
Fl. Malaya 2 (1955) 524, f. 307; Kalikasan 4
(1975) 229. — Stenosemia pinnata Copel.,
Philipp. J. Sci. 1 (1906) Suppl. 146; Philipp.
J. Sci. 2 (1908) Bot. 23; Alderw., Malayan
Ferns (1908) 727; Copel., Fern Fl. Philipp.
(1960) 302. — Type: Copeland 1601, Min-
danao (MICH).

Pleocnemia membranifolia auct.: Bedd., Handb.
Ferns Brit. India, Suppl. (1892) 48, p.p., not
Nephrodium membranifolium Presl.

Fronds strongly dimorphous. Sterile frond: stipe
dark purplish brown, commonly 15—30 cm long,
minutely hairy, later glabrescent on the abaxial
surface (persistent longer hairs near apex), basal
scales c. 8 mm long, less than 1 mm wide; lamina
15—20(—30) cm long, on young plants deltoid, on
older plants elongate; basal pinnae with stalks 2
mm long, above them 1—3 pairs of + adnate pinnae
often with buds at their bases; basal pinnae to c. 12
x 4 cm, lobed to 5—7 mm from the costa, lobes
broad, obtuse, entire; suprabasal pinnae lobed less
than halfway to costa, upper ones much less deep-
ly; apical lamina deltoid, deeply lobed, lobes en-
tire; veins forming areoles along costae of pinnae
and a few along costules in lobes of larger pinnae;
lower surface variably short-hairy on costae, other-
wise almost glabrous; upper surface densely hairy
on costae with a few short thick hairs near sinuses.
Fertile frond: stipe to at least 45 cm long; lamina
to c. 15 cm long, its parts as in sterile fronds but
pinnae lobed to a very narrow wing along costae,
lobes less than 2 mm wide and well-spaced, cover-
ed beneath by sporangia which project beyond the
margin. — Fig. 19d-f.

Distribution — S Thailand; Malesia: Sumatra,
Malay Peninsula, Borneo, Philippines.

Habitat — On limestone in shade, often abun-
dant, also rarely on other rocks in Malaya, more
frequently in Luzon; in alluvial forest in limestone
area of G. Mulu, Sarawak. .

Cytology — The chromosome number 80-82
was recorded by Manton in Holttum (1955), now
been corrected by T.G. Walker (in litt.) to n = 80.

Note — One specimen from Gopeng (Perak) has
basal pinnae of Tectaria form on one frond.

116 Flora Malesiana ser. II, Vol. 2! (1991)

CYCLOPELTIS

Cyclopeltis J. Sm., Companion Bot. Mag. 72 1846) 36; Hist. Fil. (1975) 225; Copel.,
Gen. Fil. (1947) 112; Tryon & Tryon, Ferns & Allied Pl. with spec. ref. to Trop.
America (1982) 484, f. 71: 1, 3, 4; Holttum, Gard. Bull. Sing. 39 (1986) 160. —
Hemicardion Fée, Mém. Foug. 5. Gen. Filic. (1852) 282, pl. 22A. — Type species:
Hemicardion nephrolepis Fée = Cyclopeltis semicordata (Sw.) J. Sm.

Aspidium (§ Polystichum) p.p. auct.: Hook. Sp. Fil. 4 (1862) 16.

Polystichum p.p. auct.: Bedd., Handb. Ferns Brit. Ind. (1883) 201.

Caudex short, erect, its scales narrow, entire or with marginal teeth formed by out-
growths from single cells; fronds simply pinnate; rachis rounded on both surfaces in
living plants, the lower surface bearing small scales and short erect multicellular hairs,
upper surface almost glabrous; pinnae sessile, articulate to the rachis, subentire, their
bases cordate at least on the basiscopic side (hastate on one species); unicellular glands
lacking; veins free, arising from the costae of pinnae in paucipinnate groups, the lower
veinlets not nearly reaching the margin; sori dorsal on lateral veinlets, in one to several
irregular rows parallel to the costa; indusia peltate. — Fig. 20.

Distribution — Malesia and SE Asia (c. 6 species); widely in tropical America (1 spe-
cles).

Habitat — Forest plants of low altitudes; all records indicate a rocky habitat, some spe-
cies on limestone.

Cytology — Base chromosome number 41 (C. semicordata only). The only recorded
cytological observation is of the West Indian species, which is diploid.

Taxonomy — This small genus is not nearly allied to any other in the Tectaria group.
The plants grow on rocks, where their roots would be more quickly affected by periods of
dry weather than those of plants rooted in the soil; the small simple and articulate nature of
their pinnae are well adapted to this habitat. But the reduction of lower pinnae and lack of
any distinctive character in them are peculiar in this group of genera. The nature of the
marginal teeth (where such exist) on scales of the caudex and stipes is also (so far as my
observation goes) peculiar. The nature and distribution of hairs on the rachis resemble
those in Pteridrys.

The distribution of Cyclopeltis is peculiar, also the fact that although quite widely dis-
tributed in the American tropics it is there very uniform, whereas in Malesia there are cer-
tainly several species, though they were not clearly distinguished by the earlier authors,
the result being a complex synonymy.

The West Indian species was the first known and is therefore the type; it was included
in Polystichum by earlier authors on account of its peltate indusia. In 1846 John Smith
had a plant in cultivation at Kew and for the first time recognized the distinctness of the
genus, but he did not clearly distinguish between the West Indian species and Philippine
ones of which he had specimens collected by Cuming.

Holttum — Tectaria Group 19)
KEY TO THE SPECIES

laaeBasesior pinnae, at least-on thei acroscopic side, cordate ..3....2.......- 2

b. Bases of pinnae hastate, their basal auricles acuminate ........ 5. C. mirabilis

Zawkinnaconmature plants 10;cm.ormore longi. 7.2... 2s 3. ee ee es 3

b. Pinnae of mature plants c. 3.5 cm long

Aare as ee eet a lege ane ee .. 4. C. rigida

3a. Few pairs of lower pinnae slightly reduced and more widely spaced; scales bearing

MMAR STMAN ACE LMM cps ene ese 3 01

PANG Wer. cue ey tS et a a 1. C. crenata

b. Many pairs of lower pinnae gradually much reduced; scales lacking marginal teeth 4
4a. Sori exindusiate; acroscopic bases of middle pinnae not cordate

2. C. novoguineensis

b. Sori indusiate; acroscopic bases of middle pinnae cordate ...... 3. C. presliana

1. Cyclopeltis crenata (Fée) C.Chr., Index
Filic. Suppl. 3 (1934) 64. — Hemicardion cre-
natum Fée, Mém. Foug. 5. Gen. Filic. (1852)
283, t. 22A, f. 1. — Aspidium semicordatum
Sw. var. crenatum Hook., Sp. Fil. 4 (1862) 17.
— Cyclopeltis semicordata var. crenata Alderw..,
Malayan Ferns (1908) 163. — Type: Gaudi-
chaud s.n., Tourane (RB, not seen; iso K).

Aspidium semicordatum var. truncatum Hook., Sp.
Fil. 4 (1862) 17. — Type: Parish s.n., Moul-
mein (K).

Cyclopeltis latupana Alderw., Bull. Jard. Bot. Bui-
tenzorg II, 16 (1914) 5; Malayan Ferns Suppl.
(1917) 138. — Type: Rachmat 869, Sulawesi,
Mt Latupa (BO).

Cyclopeltis zamboangana Copel., Philipp. J. Sci.
81 (1952) 21; Fern FI. Philipp. (1960) 250. —
Type: Copeland 1549, 1904, Mindanao (MICH,
teste M.G. Price).

Polystichum semicordatum auct. non (Sw.) Moore:
Bedd., Ferns Brit. India (1865) t. 35; Handb.
Ferns Brit. India (1883) 201, for Asian and Ma-
lesian specimens only.

Scales on caudex to more than 15 mm long,
less than 1 mm wide, bearing (at least distally)
many short teeth formed by outgrowths from mar-
ginal cells (in some Bommean specimens very short);
stipe 1S—40 cm long, scaly throughout when
young; lamina to 60 cm or more long; pinnae com-
monly to 20 pairs, distal ones gradually smaller
and falcate, lamina-apex pinna-like, a few pairs

lower pinnae more widely spaced and slightly re-
duced; middle pinnae almost at right angles to the
rachis, largest sterile ones 13 x 2 cm, almost con-
tiguous, fertile ones often narrower and more wide-
ly spaced, base truncate to subcordate on acroscopic
side, rounded to deeply cordate on basiscopic side,
margins + crisped when living, entire or + crenate,
apex short-acuminate; upper surface of rachis and
costae glabrous or a few hairs present near bases
of costae of lower pinnae; lower surface of rachis
bearing many narrow scales and also short erect
hairs, more slender hairs present on costae; sori
dorsal on veinlets, or sometimes terminal on the
basal acroscopic one of each group, usually in 3
irregular rows on each side of the costa; indusia
small, not covering sorus at maturity, shrivelling
later. — Fig. 20a-c.

Distribution — Southern Burma, Thailand and
Vietnam; Hainan; in Malesia: western Malesia and
Philippines.

Habitat — In forest on rocks, usually but not al-
ways on limestone.

Hybrids — Mr M.G. Price, who has made ex-
tensive field studies of ferns on Mt Makiling,
Luzon, writes that though both C. crenata and C.
presliana grow in the forest on that mountain, he
observed no hybrids, but that Weber 1547 and
Ramos BS 13829, both from Cagayan Province,
Luzon, are probable hybrids. Weber 1547 is repre-
sented at Kew; it is intermediate in frond-form
between the two species and its scales have some
teeth.

118

\

Ss b

CO}

Flora Malesiana ser. II, Vol. 2! (1991)

\S

PJ.Eowarps

Fig. 20. Cyclopeltis crenata (Fée) C. Chr. a. Entire sterile frond, x 0.17; b. base of a fertile pinna, x 2.6;
c. tip of a rhizome scale, showing marginal teeth, x 0.48. — C. presliana (J. Sm.) Berkeley. d. Bases of two
middle pinnae, x 0.17 (a & c: Matthew s.n., 23-11-1912; b: Holttum s.n., 23-2-1949; d: Hose 288).

2. Cyclopeltis novoguineensis Rosenst.,
Feddes Repert. Spec. Nov. Regni Veg. 10
(1912) 329; Alderw., Malayan Ferns Suppl.
(1917) 139. — Type: Bamler 46, New Gui-
nea, Logaueng (orig. not seen; probable isotype
BM).

Aspidium semicordatum vat. biauriculatum Baker,
J. Linn. Soc. Bot. 19 (1882) 294. — Cyclopel-
tis presliana vat. biauriculata (Baker) Alderw.,
Bull. Dép. Agric. Indes Néerl. 18 (1908) 6; Ma-
layan Ferns (1908) 164. — Type: R.B. Comins
s.n., Solomon Islands (K).

Stipe 4-20 cm long, bearing scales through-
out; basal scales to more than 15 mm long, less
than 1 mm wide, thin, not toothed; /amina to 100
cm long; rachis very copiously short-hairy on the
lower surface with some very narrow scales; pin-
nae many pairs, distal and lower ones gradually
smaller; middle pinnae to c. 14 x 1.5 cm, acro-
scopic base truncate with a+ distinct acute auricle,
basiscopic base strongly cordate, margins entire or
irregularly undulate, apex of pinnae of larger fronds

Narrowly acuminate, short-acuminate on small
fronds; basal pinnae c. 1 cm long and wide with
cordate acroscopic base; sori exindusiate, in 1 or 2
irregular rows parallel to the costa, the inner row
terminal on basal acroscopic veinlets, the outer
row mostly not terminal on next veinlets.

Distribution — Malesia: Eastern New Guinea,
Bismarck Archipelago; Solomon Islands.

Habitat — Forest ferns on limestone soils, often
on a steep slope, or on rocks or stream banks.

3. Cyclopeltis presliana (J.Sm.) Berkeley,
Intr. Cryp. Bot. (1857) 513; Alderw., Malayan
Ferns (1908) 165, name only; Copel., Fern Fl.
Philipp. (1960) 249. — Lastrea presliana J. Sm.
in J. Bot. (Hooker) 3 (1841) 412. — Hemicar-
dion cumingianum Fée, Mém. Foug. 5. Gen.
Filic. (1852) 283, nom. nov. superfl. — Poly-
Stichum preslianum (J.Sm.) T. Moore, Index
Fil. (1857) 84; Copel., Polypod. Philipp.
(1905) 16, p.p. —Aspidium preslianum (J.Sm.)
Mett., Farngatt. IV (1858) 36. — Aspidium

Holttum — Tectaria Group

semicordatum Sw. var. preslianum Hook., Sp.
Fil. 4 (1862) 17. — Cyclopeltis cumingiana
(Fée) Morton, Contr. U.S. Natl. Herb. 38
(1967) 46. — Type: Cuming 68, Luzon (BM;
iso K, RB).

Aspidium semicordatum Sw. var. subdimorphum
Christ in Warb., Monsunia 1 (1900) 76. —
Type: Warburg 16587, Celebes (B).

Cyclopeltis favrellii Kjellb., Bot. Jahrb. Syst. 66
(1933) 48. — Type: Kjellberg 3551, Celebes
(BO; iso BM).

Scales on caudex to 15 x 1.5 mm, rather firm,
entire; stipe 3-15 cm long, with rachis bearing
narrower scales on abaxial surface; lamina to 100
cm long; pinnae very numerous, many lower pairs
gradually reduced, the lowest sometimes only 1 cm
long, upper pinnae also gradually reduced and grad-
ing to the frond-apex; middle pinnae commonly 10
x 1.5 cm, largest seen 13 x 1.8 cm, base variably
cordate on acroscopic side, strongly so on basi-
scopic side; margins of larger pinnae distally irreg-
ularly sinuate to crenate; apex of pinnae on small
fronds obtuse, on larger fronds gradually attenuate;
lower surface of rachis densely covered with short
erect hairs (residual narrow scales also present);
lower surface of costae lacking hairs but some-
times bearing small scales; upper surface glabrous;
sori mostly at apices of veinlets, in larger pinnae
in 2 rather irregular rows nearer to margin than to
costa, the inner row on basal acroscopic veinlets;
indusia covering mature sori. — Fig. 20d.

Distribution — Malesia: Borneo, Philippines,
Celebes, Moluccas (Ambon, Ceram), New Guinea.

Habitat — On limestone, in shade.

Note — In 1841 John Smith gave the name Las-
trea presliana to a Cuming Philippine specimen
which he recognized as distinct from the West
Indian Aspidium semicordatum Sw., though Presl
had previously ascribed a Haenke Philippine col-
lection to the latter. Smith referred to Presl’s de-
scription and also described a difference in venation
between the species. Though the latter difference
does not hold, Smith’s name and his cited type are
now accepted. In 1852 Fée published the first good
description of the Philippine species, based on the
same Cuming collection; Morton (1967) preferred
to accept Fée’s name on account of the error in
John Smith’s description.

4. Cyclopeltis rigida Holttum, Blumea 35
(1991) 556. — Type: D.R. Pleyte 997, New
Guinea, Misool Is. (L; iso BO).

Stipe very short, its basal scales c. 5 mm long,
narrow, dentate distally; lamina to 40 cm long;
pinnae rigid, almost entire, narrowed both to base
and apex, apex obtuse, base cuneate on the acro-
scopic side, middle ones 3.5 x 1 cm, lower ones
decrescent, lowest 10-15 mm long; veins distinct
on upper surface, not on lower; abaxial surface of
rachis bearing many very narrow scales and very
short hairs, costae also scaly beneath; sori in one
row on each side of costae, indusiate; indusia firm,
covering mature sori.

Distribution — Malesia: New Guinea (known
only from the type collection).

Habitat — On rock-wall in forest.

Notes — Cyclopeltis rigida agrees in scales with
C. crenata but differs in its much smaller rigid pin-
nae, the lower ones gradually reduced.

Cyclopeltis rigida has pinnae about the same
length as those of C. kingii (Hance) Tagawa from
the Caroline Islands, but the latter has thin pinnae
differently shaped. The scales of the type of C.
kingii are not well preserved but do appear to have
marginal teeth distally.

5. Cyclopeltis mirabilis Copel., Philipp. J.
Sci. 3 (1909) Bot. 346, pl. 4; Alderw., Malayan
Ferns Suppl. (1917) 140. — Type: C.J. Brooks
s.n., Sarawak, Bidi (MICH).

Stipe more than 35 cm long, its scales (not
well preserved on the type) bearing some marginal
teeth; frond-form as C. crenata, a few pairs lower
pinnae slightly reduced; the bases of all pinnae
strongly auricled on both sides, the auricles nar-
rowly acuminate, the acroscopic one much shorter
and narrower than the basiscopic one; largest pin-
nae c. 16 x 2 cm; sori as in C. crenata but in four
TOWS.

Distribution — Malesia: Borneo (known from
the type only).

Note — For much of the above information I am
indebted to Mr M.G. Price who has examined the
type. It appears that the type was a chance mutant
from C. crenata which probably did not become
dispersed and may now be extinct.

120 Flora Malesiana ser. II, Vol. 2! (1991)

LASTREOPSIS

Lastreopsis R.C. Ching, Bull. Fan Mem. Inst. Biol. 8 (1938) 157; M.D. Tindale, Vic-
toria Naturalist 72 (1957) 180; Contr. New South Wales Natl. Herb. 3 (1965) 249. —
Type species: Lastreopsis recedens (J.Sm. ex Moore) Ching = Lastreopsis tenera
(R. Br.) Tindale.

Parapolystichum (Keyserl.) Ching, Sunyatsenia 5 (1940) 239. — Polystichum § Para-
polystichum Keyserl., Polyp. Herb. Bunge (1873) 11, 45. — Dryopteris subg. Para-
polystichum (Keyserl.) C. Chr., Monogr. Dryopteris 2 (1920) 93. — Type species:
Parapolystichum effusum (Sw.) Ching = Lastreopsis effusa (Sw.) Tindale (W Indies).

Caudex creeping or suberect, its vascular system a radially symmetrical dictyostele;
stipe grooved on the adaxial surface with linear aerophores along the ridges bordering the
groove; scales on caudex and near base of stipe firm, narrow, consisting mainly of nar-
row cells, not clathrate, scales on rest of frond small and never abundant; frond in almost
all species deltoid-pentagonal, the basal pinnae longest and with elongate basal basiscopic
branches; lamina always much dissected, the ultimate leaflets always small and + deeply
lobed with asymmetric bases, their margins thickened and decurrent to join the margins of
the groove of the pinna-rachis, the prominent costae of the distal parts of pinnae decurrent
to form a raised band between the thickened margins of the groove; veins always free;
hairs almost always present on both surfaces of axes and often on lamina, always consist-
ing of several cells, in some species acute, remaining stiff when dried, in other species not
acute and becoming contorted when dried; unicellular glands sometimes present, ovoid to
elongate; sori orbicular, indusiate or not, the indusia in some species bearing glands like
those of the lamina; sporangia lacking appendages on the body but usually bearing glands
on the stalk; spores winged or with many obtuse protuberances. — Fig. 21.

Distribution — Pantropic; about 35 species, most abundant in Australasia.

Habitat — Forest ferns, mostly of seasonal climates; Malesian species on mountains,
mostly at high altitudes.

Cytology — Base chromosome number 41; Malesian species L. rufescens and L. te-
nera diploid; some Australian species tetraploid.

Taxonomy — In his Index Filicum (1905) Christensen included all species of Lastre-
opsis, as here delimited, in the genus Dryopteris which he recognized as being a complex,
for which he proposed a tentative subdivision on pp. xx—xxii. In attempting a better sub-
division, in his Monograph of Dryopteris (1913 and 1920), he limited himself to the spe-
cies of tropical America. R.C. Ching spent three years (1929-1932) in Europe, studying
ferns with Christensen and visiting the major European herbaria to see the types of Chi-
nese species. He attempted to apply the ideas expressed in Christensen’s Monograph to
the species of Asia. In so doing he recognized that in Asia there is more than one group of
species which accord with Christensen’s definition of Dryopteris subg. Ctenitis. One such
group he separated under the name Lastreopsis (1938) noting that the hairs on the frond
are not contorted when dried; he included only four species (one of them is here excluded).
In a new classification of Leptosporangiate ferns (1940) he raised Dryopteris subg. Para-
polystichum (Keyserl.) C. Chr. to generic rank, placing it, without discussion, in the

Holttum — Tectaria Group 121

family Thelypteridaceae, evidently because the hairs are different from those in his Las-
treopsis, though they are quite different from the hairs shown by Christensen to be
characteristic of Thelypteris. Ching added two species of the Pacific to those of tropical
America included by Christensen in his subgenus.

Dr M.D. Tindale published an account of the Australian species of Lastreopsis in
1957. Then, looking more widely, she discovered that there are species of Lastreopsis in
Africa and America and prepared a full monograph of the genus (1965) recognizing 33
species. In so doing she re-defined the genus and concluded that it comprises species with
different types of hairs, including those placed in Parapolystichum by Ching. With this
conclusion I agree. As an indication of past confusion, it is notable that synonyms in 14
different genera are recorded by Tindale.

Lastreopsis accords with Ctenitis in the presence of glands on various parts of the
frond, on indusia and on the stalks of sporangia. It differs from Crenitis in scales, in the
shape of axes of the frond in relation to the leaflets they bear (not noticed by Ching) and in
the position of linear aerophores on stipe and rachis (mentioned by neither Ching nor Tin-
dale). In scales Lastreopsis is nearer to Tectaria than to Ctenitis.

KEY TO: THE SPECIES

la. Hairs on upper surface of rachis ctenitoid, shrivelled when dried or with obtuse api-

Call CIS OE are eee en eminem nate

b. Hairs on upper surface of rachis persistently terete with acute apical cells

4. L. tenera

2a. Sori exindusiate; scales less than 2 mm long .............. 1. L. rufescens

b. Sori indusiate; scales much longer... .

3a. Frond wholly anadromous
b. Frond catadromous distally

1. Lastreopsis rufescens (Blume) Ching, Bull.
Fan Mem. Inst. Biol. 8 (1938) 160; Tindale,
Victoria Naturalist 73 (1957) 184; Contr. New
South Wales Natl. Herb. 3 (1965) 309; Sledge,
Kew Bull. 27 (1972) 412. — Aspidium rufes-
cens Blume, Enum. Pl. Javae (1828) 168. —
Polypodium rufescens (Blume) Blume, FI. Ja-
vae Filic. (1829) 194, t. 91; Bedd., Ferns S.
India (1864) 78, t. 236. — Phegopteris rufes-
cens (Blume) Mett., Ann. Mus. Bot. Lugd.-Bat.
1 (1864) 223; Bedd., Handb. Ferns Brit. India
(1883) 293; Alderw., Malayan Ferns (1908)
493. — Dryopteris rufescens (Blume) C. Chr.,
Index Filic. (1905) 290; Backer & Posth., Va-
renfl. Java (1939) 48, f. 8. — Type: Blume s.n.,
Java (L 908,338-446, left hand specimen).

2 2 © © © © oe

2 2 eo ee ew

BR te ota Si taine eden a ae Ca eae See 3
EAS cepts 2. L. novoguineensis
Meche aoe Weare Ones 3. L. subsparsa

Caudex short-creeping, covered with persistent
appressed firm scales little over 1 mm long; stipe
to 40 cm long, light castaneous, persistently scaly
near base only; lamina to 45 cm long; basal pin-
nae to 20 cm long, bearing 1 or 2 pairs of free and
1 or 2 pairs adnate pinnules, the rest deeply lobed,
the lobes crenate, basal basiscopic pinnule to 11
cm long with 1 pair of free tertiary leaflets; basal
acroscopic pinnule c. 7 cm long; middle pinnae
deeply obliquely lobed, the largest with a pair of
adnate pinnules; rachis bearing ctenitoid hairs 0.3
mm long on abaxial surface, thicker longer hairs
on adaxial surface; many + appressed cylindric yel-
lowish glands on lower surface, on and between
veins; sori medial on pinna-lobes, on acroscopic
branches where veins are forked, exindusiate, some

122 Flora Malesiana ser. II, Vol. 2! (1991)

P.J.Eowarps

Fig. 21. Lastreopsis tenera (R. Br.) Tindale. a. Base of a distal pinna, showing basiscopic base of lamina
hardly thickened, x 4. — L. rufescens (Blume) Ching. b. Outline of a basal pinna, x 0.5; c. leaflet of a
middle pinna, x 2.2; d. upper surface of rachis and its junction with pinna-rachis, x 12; e. upper surface of
pinna-rachis of a middle pinna, showing the thickened margins of two pinnules joining the raised margin
of the pinna-rachis, x 16 (a: Copeland 2090; b, c: Mousset s.n., Java; d, e: Mousset 376).

Holttum — Tectaria Group

123

cylindric glands present on stalks of sporangia;
spores with inter-connected wings or crests. —
Fig. 21b-e.

Distribution — Sri Lanka; in Malesia: Java and
Lesser Sunda Islands; NE Queensland.

Habitat — On mountains, middle elevations to
2300 m, on shaded slopes.

2. Lastreopsis novoguineensis Holttum, Blu-
mea 35 (1991) 557. — Type: Nakaike 578, Pa-
pua New Guinea, Central Prov., Woitape (K).

Caudex suberect, bearing firm scales to 6 x 1.5
mm; stipes to 60 cm long, scaly as caudex at the
base, covered throughout with very short hairs; /a-
mina to 35 cm long, very firm; basal pinnae to 25
cm long, bearing 4—5 pairs of free pinnules, the
rest connected by a narrow wing along the pinna-
rachis; basal basiscopic pinnule to 13 cm long,
bearing 1 pair of deeply lobed free tertiary leaflets
2.5 x 0.6 cm and many other pairs connected by a
narrow wing; second pair of pinnae also with pin-
nate basal pinnules; upper pinnae all anadromous
at their bases; lower surface of pinna-rachis bearing
many glands and few very short hairs, upper surface
bearing ctenitoid hairs 0.1 mm long; sori medial
on veins, covered with very firm glabrous reniform
indusia.

Distribution — Malesia: Papua New Guinea.

Habitat — Widely on mountains at 1800-3000 m
altitude.

3. Lastreopsis subsparsa (Alderw.) Tindale,
Victoria Naturalist 73 (1957) 184; Contr. New
South Wales Natl. Herb. 3 (1965) 291. — Dry-
opteris subsparsa Alderw., Bull. Jard. Bot. Bui-
tenzorg II, 20 (1915) 14; Malayan Ferns, Suppl.
(1917) 170. — Type: C.A. Backer 9803, E
Java, Besoeki, Mt Hyang (BO).

Dryopteris sparsa auct. [non (Ham.) Kuntze] p.p.:
Backer & Posth., Varenfl. Java (1939) 46.

In size and form of fronds near L. novoguineen-
sis, also in pubescence, but tertiary and other ul-
timate leaflets at most crenate; indusia small, ca-
ducous.

Distribution — Malesia: Java (known only from
the type).

Habitat — At 2000 m altitude.

4. Lastreopsis tenera (R. Br.) Tindale, Victoria
Naturalist 73 (1957) 181; Contr. New South
Wales Natl. Herb. 3 (1965) 316; Sledge, Kew
Bull. 27 (1972) 411; DeVol & Kuo, Fl. Tai-
wan 1 (1975) 334, f. 119; Brownlie, Fl. Nouv.
Caled. 3 Pterid. (1969) 226; Pterid. Fl. Fiji
(1977) 307. — Nephrodium tenerum R.Br.,
Prod. (1810) 149. — Dryopteris tenera (R. Br.)
C. Chr., Index Filic. (1905) 297. — Ctenitis
tenera (R.Br.) Copel., Gen. Fil. (1947) 125.
— Type: R. Brown 23, Queensland (BM, K).

Lastrea recedens J.Sm. ex Moore, Gard. Chron.
(1855) 708 with fig.; Bedd., Ferns S. India
(1863) t. 98; Handb. Ferns Brit. India (1883)
260. — Nephrodium recedens (J.Sm.) Hook.,
Sp. Fil. 4 (1862) 135, t. 265. — Dryopteris
recedens (J.Sm.) Kuntze, Revis. Gen. PI.
(1891) 813; Alderw., Malayan Ferns (1908)
198. — Lastreopsis recedens (J. Sm.) Ching,
Bull. Fan Mem. Inst. Biol. 8 (1938) 161. —
Ctenitis recedens (J. Sm.) Copel., Gen. Fil.
(1947) 124; Fern Fl. Philipp. (1960) 242. —
Type: Cuming 96, Luzon (BM; iso K).

Aspidium gardnerianum Mett., Farngatt. IV (1858)
71. — Type: Gardner 60, Sri Lanka (B?; at K
Gardner 1110).

Dryopteris simozawae Tagawa, Acta Phytotax. Geo-
bot. 4 (1935) 135. — Lastrea simozawae (Ta-
gawa) Tagawa, Acta Phytotax. Geobot. 8
(1939) 169. — Type: Simozawa 768, Taiwan
(KY, seen by Tindale).

Caudex short-creeping with very closely placed
stipes; scales thin, to c. 10 x 1 mm, persistent;
Stipe to SO cm long, light castaneous, bearing
slender terete hairs to 0.5 mm long; lamina to 75
cm long and wide but fronds 20 cm long, some-
times fertile; basal pinnae to 30 cm long (includ-
ing stalk 3.5 cm), basal basiscopic pinnules to 15
x 8 cm bearing tertiary leaflets to 4 x 1.5 cm lobed
almost to their costae, the lobes crenate, the lowest
tertiary leaflets free, the rest decurrent or joined by
a narrow wing, basal acroscopic pinnules to 10 x
5 cm bearing tertiary leaflets to 2.5 x 1.2 cm; up-
per pinnae catadromous; rachis bearing terete hairs
0.5 mm long both sides, those on the abaxial
side slender, spreading, on adaxial side antrorse,
thicker, sometimes longer; lower surface of leaflets
bearing short erect hairs on and between veins,

124 Flora Malesiana ser. II, Vol. 2! (1991)

Sumatra, Bali, Philippines (Luzon, Leyte); Taiwan;
Queensland, New Caledonia, Fiji.
Habitat — In Luzon at 250-700 m, in forest.
Note — The margins of decurrent bases of leaf-

lets are hardly thickened in this species.

also some almost spherical red glands, upper sur-
face with fewer hairs; sori at or near ends of veins;
indusia small, bearing red spherical glands. — Fig.

21a.
Distribution — S India & Sri Lanka; in Malesia:

Index to scientific plant names

INDEX TO SCIENTIFIC PLANT NAMES

125

Suprageneric epithets have been entered under the family name to which they belong preceded by the indi-
cation of their rank (subfamily, tribe, etc.). Infrageneric epithets have been entered immediately under the
generic name to which they belong, preceeded by the indication of their rank (subgenus, section, etc.).
Infraspecific epithets have been entered under the specific name to which they belong preceded by the indi-
cation of their rank (subspecies, variety, forma, etc.).

Synonyms have been printed in italics. Page numbers in bold type denote main treatment; an asterisk
behind a page number denotes the presence of a figure of the concerned taxon.

Acrostichum 101, 105
aptifolium (Kunze) Hook.
101
auritum Sw. 49
hollrungii (Kuhn) Baker 38
quercifolium Retz. 75
semipinnatum Roxb. 78
stenosemioides Baker 112
taccifolium (Fée) Hook. 89
teysmannianum Baker 115
Aenigmopteris Holttum 3, 102
dubia (Copel.) Holttum 102,
103
elegans Holttum 103, 104*
katoi Holttum 103, 104
mindanaensis Holttum 102,
103
pulchra (Copel.) Holttum
102, 103
Alsophila subglandulosa Hance
35
Arcypteris Underw. 8, 9, 39
brongniartii (Bory) Holttum
15
difformis (Blume) Underw. 13
gigantea Holttum 15
irregularis (Presl) Ching 12
mecrodonta (Fée) Holttum 13
Aspidiaceae 2, 3
tribe Aspidieae Ching 1, 2
tribe Dryopterideae Ching 2
Aspidiaceae auct. 1
Aspidium Sw. 4, 8, 39, 40,
105, 118
sect. Euaspidium Hook. 39
sect. Polystichum auct. 116
alatum Hook. & Grev. 92
amplifolium Alderw. 72
angilogense Christ 17
angulatum (Willd.) Mett. 85
angulatum auct. 96
bamlerianum Rosenst. 52
beccarianum Baker 52
beccarianum (Cesati) Diels 93

(Aspidium)

biseriatum Christ 85
bolsteri Copel. 93
brachiatum Zoll. & Moritzi

70
brongniartii (Bory) Diels 15
brooksii (Copel.) C. Chr. 98
bryantii Copel. 93
burchardii Rosenst. 87
calcareum Pres 70
cesatianum C. Chr. 52
cicutarium auct. 58
coadunatum Wall. ex Hook.

& Grev. 58
coadunatum auct. 56
conjugatum Blume 10, 16
dahlii (Hieron.) Diels 20
decastroi Alderw. 83
decurrens Presl 90

var. mamillosum (Moore)

Alderw. 90
devexum Kunze ex Mett. 57
difforme Blume 12
divergens Rosenst. 87
durvillei Bory 67
evenulosum Alderw. 52
everettii (Baker) C. Chr. 91
falcipinnum Alderw. 85
ferrugineum Sw. 58
fissum Kunze 87
fuscipes Bedd. 48
gardnerianum Mett. 123
giganteum Blume 114

var. minor Hook. 57

grande Mett. 81
grandifolium Presl 81, 91
heptaphyllum Baker 81
heterodon Copel. 90
immersum Hook. 51
intermedium Blume 30, 33
intermedium Mett. 57
irregulare (Presl) C. Chr. 12
irriguum Presl 88
Johannis-winkleri C. Chr. 93

(Aspidium)

Juglandifolium (Baker) Christ
89
kawakamii Alderw. 81
keckii Luerssen 72
kehdingianum (Kuhn) Christ
99
kuhnii C. Chr. 52
kunstleri Bedd. 52
kwarenkoense Hayata 67
labrusca auct. 98
ledermannii Brause 52
leuzeanum (Gaudich.) Kunze
19
var. alsophiloides Christ
19)
lobbii Hook. 73
macrophyllum auct. 90
maingayi (Baker) Holttum
78
malayense Christ 54
melanocaulon Blume 66
melanorachis (Baker) C. Chr.
69
membranaceum Hook. 57
membranifolium (Presl)
Kunze 49
membranifolium auct. 108
menisciicarpon (T. Moore)
Mett. 85
menyanthidis Presl 88
microsorum Pres| 66
microthecium Fée 7
multicaudatum (Clarke)
Bedd. 54
nebulosum (Baker) C. Chr.
66
novo guineense Rosenst. 93
nudum auct. 80
obtusilobum auct. 32
oligophyllum Rosenst. 87
pachyphyllum Kunze 81
palmatum Mett. 95
pandurifolium C. Chr. 100

126 Flora Malesiana ser. II, Vol. 21 (1991)
(Aspidium) (Aspidium) (Ctenitis)
papuanum (Copel.) Alderw. subtripinnatum Miq. 36 balabacensis (Christ) Copel.
67 sumatranum C. Chr. 78 47
papyraceum Alderw. 85 syrmaticum Willd. 5 boholensis Holttum 22, 27*,
pentaphyllum Alderw. 97 © teratocarpum Alderw. 76 28, 31

pentaphyllum Willd. 85
persoriferum Copel. 80
f. latipinna Alderw. 79,
80
f. paucipinna Alderw. 79
platanifolium Mett. 96
platyphyllum Presl 80
pleiosorum Alderw. 79
pleocnemioides Alderw. 52
polymorphum Wall. ex
Hook. 87
preslianum (J. Sm.) Mett.
118
profereoides Christ 112, 113
prominens Alderw. 51
psilopodum C. Chr. 75
pteropus Kunze 90
pulvinuliferum auct. 33
quinquefoliatum C. Chr. 85
rarum Alderw. 100
repandum Willd. 80, 87
ridleyanum Alderw. 90
rufescens Blume 121
rufinerve (Hayata) Hayata 17
sagenioides Mett. 108
Sagenioides auct. 47
sanctum Blume 85
saxicola Blume 114
semibipinnatum Hook. 73
semicordatum Sw. 119
var. biauriculatum Baker
118
var. crenatum Hook. 117
var. subdimorphum Christ
119
var. truncatum Hook. 117
stifolium (Willd.) Mett. 84
var. teysmannianum
Christ 85
simonsii (Baker) Bedd. 67
singaporeanum Hook. &
Grev. 100
spectabile Blume 6
stenophyllum (Baker) Diels
73
subaequale Rosenst. 113
subcaudatum Alderw. 77
subdecurrens (Luerssen)
C. Chr. 78
subdigitatum Blume 75

terminale Rosenst. 85
ternatense Alderw. 80
ternifolium Alderw. 85
teysmannianum Hook. 85
trichotomum Fée 46
tricuspe Bedd. 91

var. glabrum Alderw. 87
trifoliatum (L.) Sw. 39
trifolium Alderw. 85

var. compitale Alderw. 85,

86
variolosum Wall. ex Hook.
515,525.70
vastum Blume 92
f. latior Alderw. 93
vile Kunze 30
vile Raciborski 30
vitis (Racib.) C. Chr. 97
weinlandii Christ 67
wendlandii 69
whitfordii Copel. 13
wigmanii Racib. 109
zippelianum C. Chr. 58
zollingerianum Kunze 70
zollingerianum auct. 51
Asplenium alatum Ridley 90
ambiguum Sw. 113
Athyrium 2
loheri Christ 45
Campylogramma pteridiformis
Alderw. 97
trollii Goebel 78
Cardiochlaena alata Fée 90
laevis Fée 81
menyanthidis (Presl) Fée 89
sinuosa Fée 81
Chlamydogramme Holttum 3,
37
elata Holttum 38, 39
hollrungii (Kuhn) Holttum
37, 38*, 60
Ctenitis C. Chr. 2, 3, 21, 26,
32, 41, 42, 101, 106,
121
aciculata (Baker) Ching 22,
24*, 25
alteroblumei Holttum 23,
27*, 30, 33, 34*
angusta Holttum 23, 35
atrorubens Holttum 23, 34

bulusanica Holttum 23, 29
costulisora Ching 36
croftii Holttum 23, 32
decurrentipinnata (Ching)
Ching 23, 28, 33
dissecta (G. Forster) 49
dubia (Copel.) Copel. 103
eatonii (Baker) Ching 35
elata Holttum 23, 31
erythradenia Holttum 23, 28,
35
fijiensis (Hook.) Copel. 32
humilis Holttum 23, 30
kinabaluensis Holttum 22,
26
var. crassisquama Holttum
26
var. kinabaluensis 26
kjellbergii (C. Chr.) Ching
22, 25
koordersii Holttum 23, 32
laxa Copel. 46
loheri (Christ) Copel. 45
manii auct. 31
manilensis (Presl) Holttum
45
mesodon (Copel.) Copel. 45
minutiloba Holttum 23, 31
muluensis Holttum 22, 25
obscura (Hook.) Copel. 108
paleolata Copel. 23, 35
pallens (Brackenr.) M.G.
Price 22, 26, 27*
propinqua (Presl) Copel. 22,
27*, 28
pulchra (Copel.) Copel. 103
ramosii Copel. 46
recedens (J. Sm.) Copel. 123
thodolepis 36
sagenioides (Mett.) Copel.
108
seramensis Holttum 23, 29*
setosa (Presl) Holttum 23,
33
setosa auct. 23, 35
silvatica Holttum 23, 28, 30
subconnexa (Christ) Holttum
22, 28
var. alstonii Holttum 28
var. subconnexa 28

Index to scientific plant names 127

(Ctenitis) Dennstaedtiaceae (Dryopteris)
subglandulosa (Hance) Ching subfam. Tectarioideae Holt- aciculata (Baker) C. Chr. 25
23, 27*, 34*, 35, 36 tum 1 balabacensis Christ 47
submarginalis (Langsd. & Dictyocline Presl 39 decurrentipinnata Ching 33
Fisch.) Ching 21 Dictyopteris Presl 8, 9, 10 dissecta (G. Forster) Kuntze
subobscura (Christ) Holttum ambigua (Hook.) Alderw. 113 41, 49

23, 31

subtripinnata (Miq.) H. Ito 36

sumbawensis Holttum 23,
32

tabacifera (Alderw.) Ching
22, 26

tenera (R. Br.) Copel. 123

trichotoma (Fée) Copel. 47

vilis (Kunze) Ching 23, 24*,

30, 33
vilis auct. 23, 34
wenzelii Copel. 111
Ctenitis auct. 42, 105
Ctenitopsis Ching 39, 41, 42,
105, 106
dissecta (Forster) Ching 49
fuscipes (Bedd.) C. Chr. 48
membranifolia (Presl) Ching
49
obscura (Hook.) C. Chr. 108
sagenioides (Mett.) Ching
108
setulosa (Baker) C. Chr. 47
trichotoma (Fée) 46
Cyatheaceae 17
Cyclopeltis J. Sm. 1, 2, 3, 116
crenata (Fée) C. Chr. 117,
118*, 119

cumingiana (Fée) Morton 119

favrellii Kjellb. 119
kingii (Hance) Tagawa 119
latupana Alderw. 117
mirabilis Copel. 117, 119
novoguineensis Rosenst.
117, 118
presliana (J. Sm.) Berkeley
117, 118*
var. biauriculata (Baker)
Alderw. 118
rigida Holttum 117, 119
semicordata (Sw.) J. Sm.
116
var. crenata Alderw. 117
zamboangana Copel. 117
Davallia 2
Dendroglossa subquinquefida
(Fée) Fée 99
taccifolia (Fée) Fée 89
Dennstaedtia sp. 104

andaiensis (Baker) Alderw. 15

attenuata Presl 8

barberi (Hook.) Bedd. 52

beccariana (Cesati) Alderw.
93

bolsteri (Copel.) Alderw. 93

brooksii (Copel.) Alderw. 98

bryantii (Copel.) Alderw. 93

carinata Alderw. 92

compitalis Alderw. 19

dahlii (Hieron.) Alderw. 20

difformis (Blume) T. Moore
8, 13

distincta Alderw. 114

elliptica (Copel.) Alderw. 114

ferruginea (Mett.) Alderw. 58

hancockii (Baker) Alderw. 78

hemiteliiformis (Racib.)
Alderw. 19

heterosora (Baker) Bedd. 78

irregularis (Presl) Presl 8, 12

labrusca auct. 98

macrodonta Fée 12, 13, 15

me galocarpa (Hook.) J. Sm.
114

olivacea (Copel.) Alderw. 19

pentaphylla Alderw. 85

petrophya (Blume) T. Moore
114

polycarpa (Mett.) Bedd. 52

pteroides Presl 15

saxicola (Blume) Alderw. 114

subaequalis (Rosenst.)
Alderw. 113

vitis (Racib.) Alderw. 97

Digrammaria Presl 105

ambigua auct. 112

Diplazium 2
Dryomenis Fée

menisciicarpon T. Moore 85
phymatodes Fée 85

Dryopolystichum 2
Dryopsis Holttum & Edwards 2,

32

Dryopteris Adans. 1, 2, 40, 41,

105, 106, 120
subg. Crenitis C. Chr. 21
subg. Parapolystichum
(Keyserl.) C. Chr. 120

dissecta auct. 48
dubia Copel. 103
escritorit Alderw. 47
gigantea (Blume) Kuntze 114
gurupahensis (C. Chr.)
C. Chr. 109
intermedia (Blume) Kuntze 29
var. microloba Christ 34
kjellbergii C. Chr. 25
laokaiensis C. Chr. 108
leuzeana (Gaudich.) Kuntze
19
mesodon Copel. 45
mettenianum Christ 7
var. novo guineensis
Brause 8
microthecia (Fée) C. Chr. 7
millettii C. Chr. 30
obscura (Hook.) Christ 108
obtusiloba (Baker) Kuntze 32
pulchra 103
pulvinulifera auct. 33
raciborskii Alderw. 47, 49
recedens (J. Sm.) Kuntze 123
rhodolepis (Clarke) C. Chr.
36
rizalensis Christ 26
rufescens (Blume) C. Chr.
121
rufinervis Hayata 17
Sagenioides (Mett.) Kuntze
108
f. centracta Alderw. 109
subsp. obscura C. Chr.
108
var. gurupahensis C. Chr.
109
schizoloma (Alderw.) C. Chr.
108
setosa (Blume) Kuntze 30
simozawae Tagawa 123
Sparsa auct. 123
squamulifera Alderw. 31
subarborea
var. glabrior Alderw. 31
subconnexa (Christ) C. Chr.
28
subglandulosa (Hance) Hayata
35

128 Flora Malesiana ser. II, Vol. 2! (1991)

(Dryopteris) (Heterogonium) (Lastrea) |
subsagenioides Alderw. 7 giganteum (Blume) Holttum recedens J. Sm. ex Moore |
subsparsa Alderw. 123 107, 114 123 |

syrmatica (Willd.) Kuntze 6
var. petiolosa Christ 6
tabacifera Alderw. 26
tenera (R. Br.) C. Chr. 123
ternata (Baker) Kuntze 82
vangenderenstortii Alderw. 7
vilis (Kunze) C. Chr. 30
wigmanii (Racib.) 109
zeylanica Alderw. 32
Gymnogramma Desv. 105
ambigua Hook. 112
maingayi Baker 78
palmata Baker 97
philippinensis Fée 113
Gymnopteris decurrens Hook. 78
hollrungii Kuhn 38
latifolia Goldm. 99
quercifolia (Retz.) Bernh. 75
taccifolia (Fée) Presl 89
trilobata J. Sm. ex Hook. 99
Hemicardion Fée 116
crenatum Fée 117
cumingianum Fée 118
nephrolepis Fée 116
Hemigramma Christ 38, 39, 59,
60
decurrens (Hook.) Copel. 78
grandifolia Copel. 38
hollrungii (Kuhn) Copel. 38,
39
latifolia (Goldm.) Copel. 99
latifolia auct. 98
Slifolia (Rosenst.) Copel. 76
taccifolia (Fée) Copel. 89
zollingeri (Kurz) Christ 39,
98
var. major Christ 99
Hemionitis L.
gymnopteroidea Copel. 99
f. major Copel. 99
maingayi (Baker) Ridley 78
zollingeri Kurz 59, 98, 99
Heterogonium Presl 1, 3, 10,
36, 41, 43, 105
alderwereltii Holttum 107,
112
aspidioides Presl 105, 107,
112, 114
aspidioides auct. 113
calcicola K. Iwatsuki &
M. Kato 107, 111
ceramense Holttum 106, 107

gurupahense (C. Chr.)
Holttum 109
gurupahense auct. 108
lobulatum Holttum 106, 109
nieuwenhuisii (Racib.)
C. Chr. 112
novoguineense Holttum 107,
111
obscurum (Hook.) Holttum
108, 109
pinnatum (Copel.) Holttum
106, 107, 110*, 112,
115
pinnatum x H. sagenioides
112
pluriseriatum Holttum 107,
115
profereoides (Christ) Copel.
106, 107, 110*, 113
rufescens Holttum 107, 109
sagenioides (Mett.) Holttum
107, 108, 110*
saxicola (Blume) Holttum
114
stenosemioides (Baker)
C. Chr. 106, 107, 112
subaequale (Rosenst.) Holt-
tum 113
subglabrum Holttum 106,
109
teysmannianum (Baker) Holt-
tum 107, 115
teysmannianum auct. 108
wenzeliu (Copel.) Holttum
107, 111
wigmanii (Racib.) Holttum
107, 109, 110*
Lastrea Bory 4, 105
barclayi Carr. 67
blumei Bedd. 33
blumei T. Moore 30, 33
chupengensis Ridley 45
dissecta (G. Forster) Carr. 49
intermedia
var. blumei auct. 31
var. rhodolepis Bedd. 36
manilensis Presl 45
melanopus (Hook.) Bedd. 108
padangensis Bedd. 30
pallens Brackenr. 26
presliana J.Sm. 118, 119
propinqua Presl 28, 33

setosa Presl 33
simozawae (Tagawa) Tagawa
123
spectablis auct. 113
syrmatica (Willd.) T. Moore 5
Lastreopsis R.C. Ching 2, 3,
120, 121
effusa (Sw.) Tindale 120
manilensis ( Presl) Price 45
novoguineensis Holttum 121,
123
parishii (Hook.) Ching 45
var. chupengensis (Ridley)
Holttum 45
recedens (J. Sm. ex Moore)
Ching 120, 123
rufescens (Blume) Ching
120, 121, 122*
subsparsa (Alderw.) Tindale
121, 123
tenera (R. Br.) Tindale 120,
121, 122*, 123
Leptochilus hilocarpus Fée 99
hollrungii (Kuhn) C. Chr. 38
latifolius (Goldm.) C. Chr.
99
rumicifolius Ridley 87
slifolius Rosenst. 76
subquinquefidus Fée 99
taccifolius Fée 89
zeilanicus (Houtt.) C. Chr. 75
Lomariopsis 1, 2
Luerssenia 36
kehdingiana Kuhn 99
Macrothelypteris setigera
(Blume) Ching 30
Maxonia 2
Microbrochis Presl 39, 42
apiifolia (Schk.) Presl 42
Microsorium trifidum Fée 88
Nephrodium Rich. 4, 43, 105
sect. Pleocnemia 8, 10
sect Sagenia Hook. & Baker
39, 42, 43
aciculatum Baker 25
barclayi (Carr.) Baker 67
blumei auct. 33
chrysotrichum Baker 17
cicutarium auct. 58
var. coadunatum 58
everettii Baker 91
giganteum (Blume) Baker 114

Index to scientific plant names

(Nephrodium)
giganteum auct. 15
griffithii Baker 54
hosei Baker 73
juglandifolium Baker 89
leuzeanum (Gaud.) Hook. 19
leuzeanum auct. 17
melanopus Hook. 108
melanorachis Baker 69
membranifolium Hook. 48
membranifolium Pres| 49,
115
morsei Baker 87
multicaudatum Clarke 54
nebulosum Baker 66, 85
nudum Baker.73
obscurum (Hook.) Diels 108
obtusilobum Baker 32
oligodictyon Baker 114
parishii Hook. 45
preslii Baker 28
propinquum R.Br. 28
recedens (J. Sm.) Hook. 123
rhodolepis Clarke 36
sarawakense Baker 30
setosum Baker 30
setulosum Baker 47
simonsti Baker 67
spectabile (Blume) Hook.
6,7
stenophyllum Baker 73
subdigitatum Baker 75
subpedatum Harr. 87
tenerum R. Br. 123
ternatum Baker 82
varium Pres] 20
vitis Racib. 97, 98

Nephrolepis 40
Ophioglossum zeilanicum
Houtt. 75
Parapolystichum (Keyserl.)
Ching 120, 121
effusum (Sw.) Ching 120
Phegopteris (Presl) Fée 105

brongniartii (Bory) Mett. 15

dahlii Hieron. 20

difformis (Blume) Mett. 12

ferruginea Mett. 58

irregularis (Presl) Mett. 12

macrodonta (Fée) Mett. 13

megalocarpa (Hook.)
Salomon 114

obscura Christ ex Hook. 108

obscura Fée 109

petrophya (Blume) Mett. 114

(Phegopteris)

philippinensis (Fée) Mett. 113
philippinensis auct. 108
polycarpa Mett. 52

rufescens (Blume) Mett. 121
schizoloma Alderw. 108, 109
subconnexa Christ 28
subdecurrens Luerssen 78
subobscura Christ 31

Phlebiogonium impressum Fée

a

Pleocnemia Presl 1, 3, 4, 8, 9,

10, 39, 105
acuminata Holttum 12, 16
acutissima 4, 7
andaiensis (Baker) Holttum

11, 14*,15
australis 4, 7
brongniartii (Bory) Holttum

11,15
chrysotricha (Baker) Holttum

17
confertiloba 4
conjugata (Blume) Presl 10,

12, 16, 18*

var. elatior Holttum 16’
cumingiana Presl 11, 12, 13,

17, 18*, 20
dahlii (Hieron.) Holttum 12,

18*, 20
devexa (Kunze ex Mett.)

Alderw. 57
dimidiolobata Holttum 18*,

20
fimbrillifera Alderw. 16
gigantea (Blume) Pres! 114
hemiteliiformis (Racib.)

Holttum 10, 17, 19
x intermedia Holttum 9, 11,

15
irregularis (Presl) Holttum

10, 11, 12, 13, 14*,

15, 16, 20
javanica Presl 16
kingit (Copel.) Alderw. 55
leuzeana (Gaud.) Presl 8, 9,

10, 11, 12, 16,19

var. angilogense (Christ)

Alderw. 17

var. echinocarpa Rosenst.
20

var. hemiteliiformis
Racib. 17

var. lobato-crenata
Rosenst. 20

129

(Nephrodium)

leuzeana auct. 16, 17, 20

macrodonta (Fée) Holttum 9,
D113, 14%, 20

megalocarpa (Hook.) Bedd.
114

megaphylla Holttum 12, 19

membranacea (Hook.) Bedd.
57

var. novoguineensis

Rosenst. 58

membranifolia auct. 112, 115

microthecia 4, 8

olivacea (Copel.) Holttum 5,
6, 10, 12, 6, 16, 18*, 19

pleiotricha Holttum 12, 16,
18*

porphyrocaulos Alderw. 12,
16

presliana Holttum 11, 12, 20

profereoides (Christ) Alderw.
113

rufinervis (Hayata) Nakai 17

seranensis Holttum 12, 17

stenosemioides Alderw. 112

syrmatica 4

trimenii Bedd.

f. dissecta Alderw. 54
tripinnata Holttum 20

Pleopeltis grandidentata (Cesati)

Alderw. 92
schultzei (Brause) Alderw. 91

Polybotrya Humb. & Bonpl.

100, 105
apiifolia (J. Sm.) Kunze 101
aurita (Sw.) Blume 51
cicutaria Blume 51
nieuwenhuisii Racib. 112

var. brookii Alderw. 112

orientalis Blume 51
stenosemioides (Baker)

Copel. 112

Polydictyum heterophyllum

Presl 89
menyanthidis (Presl) Pres] 89

Polypodiaceae 2, 60

subfam. Dryopteridoideae 2
section A 1, 2

Polypodium L. 2, 10, 105

andaiénse Baker 15
angulatum Willd. 85
asplenioides Bory 30
barberi Hook. 52
beccarianum Cesati 93
brongniartii Bory 15

130

Flora Malesiana ser. II, Vol. 2! (1991)

(Polypodium)
christovalense C. Chr. 97
confertum Roxb. 13
cumingianum Hook. 13, 15
difforme (Blume) Blume 12
dilatatum
var. grandidentatum Cesati
92
dissectum G. Forster 49
eximium Kunze 13
grandidentatum (Cesati) Baker
92
hancockii Baker 78
heterosorum Baker 78
hirtum Presl 33
ingens Brause 93
involucratum Roxb. 80
irregulare Presl 12
labrusca Hook. 97, 98
latifolium Forst. 66
leuzeanum Gaud. 10, 19
macrodon Reinw. ex Baker
13, 15
maingayi (Baker) Diels 78
megalocarpum Hook. 114
multiflorum Roxb. 13
nigripes Hassk. 66
obscurum Hook. 108, 109,
115
oldhamii Baker 35
oxyodon Baker 6
palmatum Blume 97
petrophyum Blume 114
pteroides Presl 15
rufescens (Blume) Blume 121
schultzei Brause 91
semipinnatum Roxb. 78
stifolium Willd. 84
viscosum C.H. Wright 108
Polystichum Roth 2, 40, 116
sect. Parapolystichum
Keyserl. 120
preslianum (J. Sm.)
T. Moore 118
semicordatum auct. 117
Polystichum auct. 116
Proferea Presl 39
Psomiocarpa Pres! 3, 100, 101
apiifolia (Kunze) Presl 100,
101*
Pteridrys C. Chr. & Ching 2,
3,4
acutissima Ching 4, 6*, 7
australis Ching 4, 7
cnemidaria (Christ) C. Chr. 4

(Pteridrys)
confertiloba Holttum 4, 6*, 8
microthecia (Fée) C. Chr. &
Ching 4, 6*, 7
olivacea (Rosenst.) Copel. 5,
6*,8
syrmatica (Willd.) C. Chr. &
Ching 4, 5*
Quercifilix Copel. 39, 59
zeilanica (Houtt.) Copel. 39,
75
Rumohra 2
Sagenia Pres 39, 40, 41, 42,
43, 106
alata (Hook. & Grev.) Bedd.
92
coadunata J. Sm. 58
decurrens (Presl) T. Moore 90
gigantea (Blume) T. Moore
114
var. minor Bedd. 57
griffithii (Baker) Bedd. 54
latifolia Pres] 42
lobbii (Hook.) Christ 73
longicruris Christ 67
macrodonta Fée 58
mamillosa Moore 90
melanocaulis (Blume)
T. Moore 66
pteropus (Kunze) T. Moore
90
subdigitata (Baker) Christ. 75
vitis (Racib.) Christ 97

Selliguea maingayi (Baker) Bedd.

78
Stenosemia Presl 39, 42, 43,
101, 105
aurita (Sw.) Presl 42, 51,
105, 109
aurita auct. 108, 113
cicutaria (Blume) Presl 51
pinnata Copel. 42, 43, 115
teysmanniana (Baker) Diels
42,115
Stenosemia auct. 105
Syngramma zollingeri (Kurz)
Diels 98
Tectaria Cav. 1, 2, 4, 8, 10, 21,
36, 38, 39, 41-43,

105, 106, 121

sect. Sagenia Holttum 3, 9,
10, 39, 42

sect. Tectaria 3, 40, 42, 59,
60

subg. Heterogonium 105

(Tectaria)
adenophora Copel. 45, 56
ambigua (Hook.) Copel. 113
amplifolia (Alderw.) C. Chr.
Ve
andaiensis (Baker) C. Chr. 15
andersonui Holttum 44, 48
angulata (Willd.) Copel. 62,
63, 66, 81, 85
apiifolia (Schk.) Copel. 42
artinexa (Clarke) Ching 39
aspidioides (Presl) Copel. 112
athyriosora M.G. Price 60,
69
aurita (Sw.) S. Chandra 42,
44, 46, 49, 53*, 101,
106
var. minor Christ 51
balabacensis (Christ) Price 47
bamleriana (Rosenst.) C. Chr.
44, 52, 53*
barberi (Hook.) Copel. 44,
52, 53*
barclayi (Carr.) C. Chr. 67
beccariana (Cesati) C. Chr.
64, 65, 93, 94*
brachiata (Z. & M.) Morton
52, 60, 68*, 70
brauniana (Karst.) C. Chr. 41
brevilobata Holttum 62, 82
brooksii Copel. 65, 98
bryantii (Copel.) Copel. 93
burchardii (Rosenst.) C. Chr.
87
calcarea (Pres!) Copel. 60, 70
calcicola Copel. 114
carinata (Alderw.) C. Chr. 92
celebica C. Chr. 93
cesatiana (C. Chr.) Copel. 52
cherasica Holttum 60, 71
christii Copel. 44, 56
christovalensis (C. Chr.)
Alston 65, 88, 94*, 97
chrysotricha (Baker) C. Chr.
7
clemensiae Copel. 91
coadunata (J. Sm.) C. Chr.
43, 45, 57, 58
var. coadunata 58
var. minor Holttum 58,
59
compitalis (Alderw.) C. Chr.
19
craspedocarpa Copel. 62, 64,
719

Index to scientific plant names

0202 7108
ais

(Tectaria)

crenata Cav. 62, 80, 81, 91
crenata auct. 80
croftii Holttum 63, 86
cumingiana (Hook.) C. Chr.
13
cumingiana Tard. & C. Chr.
13, 15
cuneifolia (Bonap.) Love &
Love 87
curtisii Holttum 61, 68*, 70
var. curtisii 70
var. hendersonii Holttum
70, 71
dahlii (Hieron.) C. Chr. 20
danfuensis Holttum 60, 69
decastroi (Alderw.) C. Chr.
62, 83
decurrens (Presl) Copel. 64,
65, 80, 90, 91
decurrens hybrid 64
decurrenti-alata Ching &
Wang 92
devexa (Kunze ex Mett.)
Copel. 44, 50*, 57
var. devexa 57
var. minor (Hook.) Holt-
tum 57, 58
var. novoguineensis Holt-
tum 57, 58
dimorpha St John 80
dissecta (G. Forster) Lellinger
44, 49, 50*
diversisora Copel. 97
dolichosora Copel. 62, 74*,
719
durvillei (Bory) Holttum 60,
62, 67, 68*
elliptica Copel. 114
evenulosa (Alderw.) C. Chr.
52
everettii (Baker) C. Chr. 91
exauriculata Holttum 62, 81
falcipinna (Alderw.) C. Chr.
85

fauriei Tagawa 61, 64, 74*,
77

ferruginea (Mett.) Copel. 45,
58

filisquamata Holttum 65, 96

fimbrillifera (Alderw.)
C. Chr. 16

fissa (Kunze) Holttum 63, 87

fuscipes (Bedd.) C. Chr. 41,
44,48

(Tectaria)

gemmifera Ching & Wang 77

gigantea (Blume) Copel. 114

grandidentata (Cesati) Holt-
tum 64, 92

grandifolia (Presl) Copel. 81

grandifolia auct. 91

griffithii (Baker) C. Chr. 44,
54

var. amplissima Holttum
44,54, 55
var. griffithii 54, 55
var. singaporeana Holttum
54, 55
gymnocarpa Copel. 58
gymnosora Holttum 77
hainanensis Ching & Wang
el
hemiteliiformis (Racib.)
C. Chr. 19
heptaphylla (Baker) C. Chr.
81
herpetocaulos Holttum 63,
89
hilocarpa (Fée) Price 39, 59,
60, 63, 65, 80, 99
holttumii C. Chr. 60, 66
hosei (Baker) Copel. 73
impressa (Fée) Holttum 44,
S152
ingens (Atk.) Holttum 41
ingens auct. 47
inopinata Holttum 65, 94*,
98
irregularis (Presl) Copel. 12,
39
var. brongniartii Copel. 15
var. macrodon Copel. 13
irregularis auct. 13
irrigua (Presl) Copel. 88
isomorpha Holttum 62, 81
jacobsii Holttum 61, 74*, 75
johannis-winkleri (C. Chr.)
C. Chr. 64, 93
Juglandifolia (Baker) C. Chr.
89
kawakamii (Alderw.) C. Chr.
81
keckii (Luerssen) C. Chr. 61,
68*, 72
kehdingiana (Kuhn) M.G.
Price 36, 65, 94*, 99
kingii Copel. 44, 55, 58
kwarenkoensis (Hayata) C.
Chr. 67

(Tectaria)

labrusca (Hook.) Copel. 65,
97, 98
latifolia (G. Forster) Copel.
99
laxa (Copel.) Price 43, 46,
53%
ledermannii (Brause) 52
leuzeana (Gaudich.) Copel.
19, 39
leuzeana auct. 16, 17
lobbii (Hook.) Copel. 61,
73,75
var. allosora Holttum 73,
75
var. denticulata Holttum
73,75
var. lobbii 73
loheri Copel. 91
lombokensis Holttum 61, 76
macrodonta (Fée) C. Chr. 58
macrodonta auct.59
macrodus (Baker) C. Chr. 13
macrota Holttum 61, 72
maingayi (Baker) C. Chr. 78
malayense (Christ) Copel. 54
manilensis (Presl) Holttum
40, 43, 45, 53*
var. chupengensis (Ridley)
Holttum 45
var. manilensis 45
megalocarpa (Hook.) C. Chr.
114
melanocaula (Blume) Copel.
50*, 60, 66
melanorachis (Baker) Copel.
60, 68*, 69
menyanthidis (Presl) Copel.
63, 88
mesodon (Copel.) Price 43,
45
mexicana (Fée) Morton 42
microchlamys Holttum 65,
96
microlepis Holttum 63, 88
minuta Copel. 65, 97
modesta C. Chr. 73
moussetii Holttum 44, 48
multicaudata (Clarke) Ching
54
murudensis Copel. 82
nabirensis Holttum 44, 54
nebulosa (Baker) C. Chr. 60,
63, 66
nesiotica Holttum 44, 55

polymorpha (Wall. ex Hook.)

Copel. 63, 86, 87
var. cuneifolia Bonap. 87

prominens (Alderw.) C. Chr.
51

pubescens Copel. 45, 59

ramosii (Copel.) Holttum 43,
46, 101

ramosii x Psomiocarpa apii
folia Presl 43, 46

ramosil x Tectaria aurita
(Sw.) Chandra 43, 46

rara (Alderw.) C. Chr. 65,
100

65, 100
squamipes Holttum 45, 56
stalactica M.G. Price 65, 95
stenosemioides (Alderw.)
C. Chr. 112
subaequalis (Rosenst.) Copel.
113
subcaudata (Alderw.) C. Chr.
61, 64, 77
subcordata Holttum 63, 83
subdigitata (Baker) Copel. 61,
15
subpedata (Harr.) Ching 87
subtriloba Holttum 65, 96

132 Flora Malesiana ser. II, Vol. 2! (1991)
(Tectaroa) (Tectaria) (Tectaria)
nitens Copel. 64, 95 repanda (Willd.) Holttum 62, sulitii Copel. 64, 91
novoguineensis (Rosenst.) 80, 81, 91 suluensis Holttum 62, 74*,
C. Chr. 64, 65, 93 repanda x Tectaria hilocarpa 77
nuda (Baker) Copel. 73 63 sumatrana (C. Chr.) C. Chr.
oligophylla (Rosenst.) rheophytica Holttum 63, 88 62, 78
C. Chr. 87 ridleyana (Alderw.) C.Chr. 90 tabonensis M.G. Price 63,
olivacea Copel. 19 rigida Holttum 44, 48 82
palmata (Mett.) Copel. 65, rufescens Holttum 63, 83 taccifolia (Fée) M.G. Price
95 rumicifolia (Ridley) C. Chr. 87 63, 89
var. dimorpha Holttum 64, schmutzii Holttum 63, 86 teratocarpa (Alderw.) C. Chr.
95, 96 schultzei (Brause) C. Chr. 64, 61, 76
var. palmata 95, 96 84*, 91 terminalis (Rosenst.) C. Chr.
var. platanifolia (Mett.) semibipinnata (Hook.) Copel. 85
Holttum 65, 95, 96 61, 65, 73 ternata (Baker) Copel. 62, 65,
var. sumatrana Holttum semipinnata (Roxb.) Morton 82
95 62, 64, 78 ternifolia (Alderw.) C. Chr.
pandurifolia (C. Chr.) C. Chr. setulosa (Baker) Holttum 42, 85
65, 100 43,47 translucens Holttum 61, 71
papuana Copel. 67, 69 var. raciborskii (Alderw.) trichotoma (Fée) Tagawa 43,
papyracea (Alderw.) C. Chr. 85 Holttum 47 46
pentaphylla (Alderw.) C. Chr. var. setulosa 47 tricuspis (Bedd.) Copel. 64,
85 siifolia (Willd.) Copel. 63, 84*, 91
peralata Copel. 90 76, 80, 84* trifida (Fée) M.G. Price 63,
platanifolia (Mett.) Copel. 96 simonsii (Baker) Ching 60, 88
pleiosora (Alderw.) C. Chr. 67 trifolia (Alderw.) C. Chr. 85,
36, 74*, 79, 92 simulans Ching 78 86
pleocnemioides (Alderw.) singaporeana (Hook. & trifoliata (L.) Cav. 39
C. Chr. 52 Grev.) Copel. 36, 50*, variolosa (Hook.) C. Chr. 51

variolosa auct. 70
vasta (Blume) Copel. 64, 92,
93
villosa Holttum 61, 72
weinlandii (Christ) Copel. 67
zeilanica (Houtt.) Sledge 39,
61, 75
zollingeri (Kurz) Holttum 59,
65, 98
Tectaria auct. 8, 105
Tectaridium Copel. 3, 36
macleanii Copel. 36, 37*
Thelypteridaceae 1, 2, 43
Thelypteris 2, 40, 121

Index to revised families in Series I (Spermatophyta)

Aceraceae 4; 3, 592
Actinidiaceae s.s. 4; 37
Aizoaceae 4; 267
Alismataceae 5:317; 6: 915
Alseuosmiaceae 10: 335
Amaranthaceae
4: 69,593; 6:915; 8: 549
Anacardiaceae 8: 395
Ancistrocladaceae 4; 8
Aponogetonaceae 4: 11; 7: 213
Araliaceae—I 915593
Araucariaceae 10: 419
Aristolochiaceae 10: 53
Balanophoraceae
IOS 92554
Basellaceae 5: 300
Bat(id)aceae 5; 414
Betulaceae 5; 207; 6: 917
Bignoniaceae 8: 114; 9: 554
Bixaceae s.S. 4; 239
Burmanniaceae
AMS 92= 539593) 9s D4
Burseraceae 5; 209, 567
6: 917; 7: 820; 9: 555
Butomaceae 5: 118
Byblidaceae 18 NBS)
Callitrichaceae 4; 251
Campanulaceae
6: 107, 928; 9: 556
Cannab(in)aceae 4: 222
Cappar(id)aceae 6:61; 7: 822
Caprifoliaceae
4: 175; 6: 928; 9: 556
Cardiopteridaceae 7 8B)
Celastraceae 6: 227, 389, 930
Centrolepidaceae 5: 421
Ceratophyllaceae 4: 41
Chenopodiaceae
4: 99,594: 6: 932; 9: 557
Chloranthaceae 10: 123
Chrysobalanaceae 10: 635
Clethraceae 7: 139
Cochlospermaceae 4: 61
- Combretaceae
— 4:533; 5: 564; 6: 932
Coniferales 10: 337
Connaraceae
5: 495; 6: 933; 9: 557
Convolvulaceae 4: 388, 599

5: 558; 6: 936; 7: 823: 9: 558
Cornaceae 8: 85
Corynocarpaceae 4: 262; 5: 557

Dichapetalaceae 5: 305; 6:
Dilleniaceae 4: 141; 7:
Dioscoreaceae 4:
Dipsacaceae 4:
Dipterocarpaceae 2h
Droseraceae
Oe Se Isepie 4 te
Elaeagnaceae 10:
Elatinaceae 4:
Epacridaceae 6:
Ericaceae 6: 469,
8: 549; 9: 562; 10:
Erythroxylaceae 5: 543; 8:
Fagaceae 13 Aes | Ye
Flacourtiaceae See dil
6: 943; 7: 827; 9:
Flagellariaceae 4: 245; 9:
Geraniaceae 6: 445; 9:
Gnetaceae 4: 336; 6:
Gonystylaceae 4:
Goodeniaceae Sp 3)3)5)-
6: 949; 7: 827; 9
Haemodoraceae 5: 111; 10
Haloragaceae U8 228)
Hamamelidaceae 5
Hippocrateaceae 6:
Hydrocharitaceae 5:
6: 952; 7: 828; 9: 566; 10:
Hydrophyllaceae 4:
Hypericaceae 8: 1; 10:
Icacinaceae Melee Qs
Iridaceae 8: 77; 10:
Ixonanthaceae 10:
Juglandaceae 6:
Juncaceae 4: 210; 9:
Juncaginaceae 4;
Labiatae 8: 301; 9:
Leeaceae 18
Lemnaceae 18
Lentibulariaceae 8:
Liliaceae s.s. 9;
Linaceae 10:
Loganiaceae
6: 293,953; 9:
Lophopyxidaceae as
Magnoliaceae 10:
Malpighiaceae 3:
Martyniaceae 4:
Menispermaceae 10:
Monimiaceae 10:
Moringaceae 4:
Myoporaceae 4:
Myricaceae 4:
Najadaceae 6:
Nyctaginaceae 6:
Nyssaceae 4:
Ochnaceae is
Olacaceae 10: 1,
Onagraceae 8:

Index to revised families in Series

Crassulaceae 4: 197; 9: 558
Cruciferae 10: 541
Crypteroniaceae 8: 187
Ctenolophonaceae 10: 629
Cupressaceae 10: 442
Cyperaceae f: 4352) 93107
Datiscaceae 4: 382
Cyatheaceae 1: 65
Gleicheniaceae baal
Isoetaceae 1: 62

Lindsaea group ifs
Lomariopsis group 1:
Schizaeaceae 1:

941 Opiliaceae 10:51
824 Oxalidaceae 7: 151, 829
293 Papaveraceae 5: 114
290 ~~ Passifloraceae 7: 405
237 Pedaliaceae 4: 216; 7: 829
Pentaphragmataceae 4: 517
562 Pentaphylacaceae yy 1721 |
151 Philydraceae Ae aS
203 Phytolaccaceae 4; 228
422 Pinaceae 10: 447
943 ‘Pittosporaceae 5: 345; 6: 960
716 Plumbaginaceae 4: 107
549 Podocarpaceae 10: 351
563 Podostemaceae 4:65; 6: 963
565 Polemoniaceae 4: 195
563 Polygalaceae 10: 455
564 Pontederiaceae 4: 255
565 Portulacaceae 7 SPA
944 Primulaceae 6: 173
349 Proteaceae 5: 147
567 Punicaceae 4: 226
566 Restionaceae 5: 416
717 ~+Rhizophoraceae
828 5: 429; 6: 965; 8: 550
363 Sabiaceae £0: 351
389 Salicaceae 5: 107
381 Salvadoraceae 4: 224
717 + Sarcosperma(ta)ceae re 32
207 Saururaceae 4: 47
717 — Scyphostegiaceae 5: 297; 6: 967
566 Simaroubaceae 6: 193, 968
717 ~~ Sonneratiaceae
621 L) sas 280551326: 973
143. Sparganiaceae 4: 233; 10: 718
566 Sphenocleaceae 4: 27
57 Sphenostemonaceae 10: 145
566 Stackhousiaceae AS 535
755 . Staphyleaceae 6: 49
219 Stylidiaceae 4: 529; 6: 976
275 Styracaceae 4: 49; 9: 568
189 Symplocaceae
607 8: 205; 9: 569; 10: 718
Taccaceae 7: 806
567 Taxaceae 10: 347
89 Thymelaeaceae
561 4: 349: 6: 1,976; 7: 830
125 = Trapaceae 4: 43
216 Trigoniaceae 4: 59
157 Trimeniaceae 10: 327
255‘ Triuridaceae 10: 109
45 Turmeraceae 4: 235
265 Typhaceae 4: 242; 6: 982
276 Ulmaceae Sa
157. ~Umbelliferae 4: 113, 595
450 5: 555; 6: 983; 7: 830; 9: 569
29 Valerianaceae 4: 253
97 Violaceae 7: 179, 831; 10: 720
717 + Xyridaceae 4: 366,598; 9: 571
98 Zygophyllaceae 4: 64
II (Pteridophyta)
177 _‘ Tectaria group F smaaell |
255‘ Thelypteridaceae 1-331

Biel

ISBN 90-71236-11-0

FLORA MALESIANA
Founded by C.G.G.J. van Steenis (1901-1986)

Compiled and published under the auspices of

Foundation Flora Malesiana

Board of the Foundation:
Aprilani Sugiarto, Jakarta, chairman

P. Baas, Leiden, vice-chairman

M.C. Roos, Leiden, secretary / treasurer
K. Larsen, Aarhus

A. Latiff Mohamad, Bangi

G.LIl. Lucas, Kew

D.H. Nicolson, Washington

M.A. Rifai, Bogor

P.F. Stevens, Cambridge, USA

Editors:

W.J.J.O. de Wilde (Series I)
H.P. Nooteboom (Series II)
C. Kalkman

Requests for scientific information to be addressed to the Editors,
at the address as mentioned below.

Subscription orders and requests for sample copies to be sent to:

Publication Department

Rijksherbarium / Hortus Botanicus

P.O. Box 9514

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The Netherlands Fax (31) (71) 273511