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SERIES II — PTERIDOPHYTA Ferns and Fern Allies
Volume 2, part 1: Tectaria Group
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LIBRARY THE NEW YORK BOTANICAL GARDEN
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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
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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.
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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
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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,
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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).
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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).
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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.
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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).
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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).
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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,