The Lindsaeoid of the Old World II.

A revision of

K.U. Kramer

Botanical Museum and Herbarium, Utrecht

INTRODUCTION

the second the Lindsaeoid Tapeinidium is largest of genera. In the present study Until 17 species are distinguished. Tapeinidium was recognized as a genus its species were

included in Microlepia, where it was originally described as an infrageneric division,

under the or in Davallia. Fée (1852), then Diels (1902), treated it as a genus, but in-

Wibelia which is of Davallia correctly interpreted name Bernhardi, actually a synonym (see Copeland, 1947). The described far have been their leaf species so mainly distinguished by architecture, of especially the degree dissection; see, e.g., van Alderwerelt van Rosenburgh (1909).

is In my opinion this at best one of several useful characters. At least equally important of the other character diat is the structure petiole and the axes of the lamina, a proved

for in the Lindsaea to be very valuable diagnostic purposes neotropical species (Kramer,

1957a) but is much less serviceable in the paleotropical ones. In some cases the rhizome

distinctive. characters the and scales are also These have been grossly neglected in past, the species distinguished by most authors are generally far too widely circumscribed. Diels for when than (l.c.), example, listed three species at a time more twice as many

were known. Accordingly there proved to be a surprisingly large number ofundescribed

of the them species, viz. 8 out 17 recognized here, some of represented by numerous in T. specimens many herbaria and collected long ago but never recognized, e.g., novo- and Lindsaea the guineense T. melanesicum. This contrasts sharply with the situation in in

is same region where the number ofnew species comparatively very small and relatively

have be in many more species to placed synonymy.

GENERAL TAXONOMIC NOTES

is natural and fern and it Tapeinidium a very homogeneous genus, seems unnecessary divide stated to it into subgenera or sections. In my former paper (Kramer, 1957a) I

it is that difficult to separate from Sphenomeris. At present it seems that this is only true the in sense that it is difficult to express the difference in simple words, in a key. During the ofthe doubtwhether preparation present paper there never was any a species belonged the other the leaf of the ultimate divisions to one or genus. In Sphenomeris pattern is

essentially cuneate-dichotomous, with the sori apical on the pinnules (segments); leaf Tapeinidium, on the odier hand, has a pinnate architecture with the sori on lateral

few this in lobes. There are very exceptions to rule; T. tenue apical sori usually occur

beside lateral and for this and other well be the ones, reasons it may most primitive

species in the genus. The spores ofTapeinidium are consistently monolete; in Sphenomeris

545 546 BLUMEA VOL. XV, No. 2, 1967

both monolete and trilete ones occur. The rhizome scales of Sphenomeris also tend to be more acicular than those of Tapeinidium, but the distinction is not very sharp. Much

fact when it closer to Tapeinidium is the genus Xyropteris, a not recognized was first Fertile leaves of like described (Kramer, 1957b). juvenile are surprisingly Tapeini- and the dium, name Tapeinidium bartlettii has been given to such specimens, but the unbroken sori and the auriculate pinnae of adult plants are so different from what is

observed in the that becomes much otherwise genus Tapeinidium a more homogeneous is the latter be entity when Xyropteris excluded, although seems to a relatively recent offshoot of the former.

Tapeinidium is distributed from South and the Ryukyu Islands to Fiji; it is absent

from Ceylon, , New Caledonia (?), and most of the Micronesian islands (map 1). of the Islands The greatest concentrations species are in Borneo (5), Philippine (5),

Celebes and the Moluccas (6), and New Guinea (6). In western Malesia the representation distributed is comparatively weak. Narrowly endemics, according to our present state of knowledge, occur chiefly in New Guinea which has 3 species confined to it and one extending to the Moluccas, but they also occur in western Malesia and elsewhere. but have been Most specimens are from mountain forests, many occasionally reported from lower and that restricted altitudes, it seems few species are actually to higher have been elevations. Tapeinidium seems to be difficult to cultivate; no specimens seen that were fromextratropical botanical gardens and only one or two from tropical gardens.

of the Map 1. Range genus Tapeinidium.

TAPEINIDIUM (Presl) C. Christensen

C. Christensen, Index Filicum (1906) 631. Basionym: Microlepia § Tapeinidium Presl,

Davallia = Epimel. Bot. (1849) 96. Type species; pinnata Cav. Tapeinidium pinnatum

(Cav.) C. Chr.

Wibelia Gen. in Nat. auct. non Bcrnhardi; Fée, Fil. (1852) 331; Diels Engler & Prantl,

4 Pflanzenfam. I (1902) 216. K. U. KRAMER: A revision of Tapeinidiutn 547

Small to medium-sized terrestrial ferns with very short to moderately long-creeping rhizome with at least in the larger species a true solenostele with external and internal strand endodermis and a medullary of sclerenchyma (perhaps lacking in one or two of the smallest species); scales long and narrow, glabrous and non-clathrate. Lamina

Sori of a pinnate type up to the last divisions, these never dichotomously divaricate. terminal uninerval or occasionally binerval (very rarely trinerval), never continuous, on the veins, mostly submarginal. Indusium rigid, attached at the base and at least the of the sides. Pluricellular filiform greater part uniseriate paraphyses present in some

(probably all) species. Spores monolete, ellipsoidal or almost bean-shaped. Gametophyte unknown.

of be the Remarks. Extensive descriptions species that are not new will provided in regional treatments where they are included and where also some illustrations will be

given.

The length of a sorus is measured at right angles to its vein, the widdi parallel to its This but makes the the with the vein. may seem paradoxical, it use of terms consistent much Lindsaea. larger genus

KEY TO THE SPECIES

Lamina and I. simply pinnate with a conform terminal pinna.

Sori 2. submarginal, most often binerval; petiole and rachis abaxially sharply bi-angular; scales to

16. T. 5 mm long longipinnulum

Sori and 2. intramarginal, almost always uninerval; petiole rachis abaxially terete or very obtusely

bi-angular, concolorous; pinnae usually obtuse, deeply crenate; scales to i£ mm long.

17. T. melanesicum

2. Sori as in the preceding species; rachis abaxially sharply bi-angular or sulcate, basally dark and

pale-margined; pinnae acuminate, very shallowly sinuate; scales to 2\ mm long.

14. T. acuminatum

Lamina if the confluent into 1. more strongly dissected, or, simply pinnate, upper pinnae a pinnatifid least division lobed base. leaf-apex or at the terminal strongly at its

3. Lamina simply pinnate, or, if more dissected, the primary rachis abaxially sharply carinate; at least

a considerable upper portion of the petiole abaxially sharply bi-angular.

least in the and rachis lamina 4. Petiole, at upper part, dark, pale-angled; pinnate + pinnatifid

or bipinnate.

the 5. Larger pinnae of full-grownplants 20—25 mm wide at widest point; texturesubcoriaceous;

margin often reflexed in dry leaves; lobes of larger pinnae sinuate, the sori not on lobes.

8. T. gracile

5. Larger pinnae of full-grown plants 10—12 mm wide at the widest point; texture herbaceous; of each margin not reflexed; lobes larger pinnatifid or basally subpinnate pinnae lobed, sorus

on a lobe: smaller forms of 1. T. tenue

also and 4. Petiole pale, or, if occasionally darker, the rachis not dark pale-carinate; or lamina simply pinnate.

6. Lamina to 12 cm 1 mm 10. T. long; petiole slender, less than thick. . . . oligophlebium

6. Lamina larger; petiole stouter.

7. Petiole abaxially obtusely bi-angular, dark and dull, ± pale-angled; rachis abaxially sori lamina mostly narrowed-rounded; submarginal, on saw-teeth; simply pinnate.

13. T. prionoides

Petiole least the rachis carinate 7. sharply bi-angular at near apex, nearly always pale; abaxially sori or bi-angular; intramarginal; lamina variously dissected.

8. Lamina pinnate + pinnatifid or more incised *) II. T. luzonicum

8. Lamina simply pinnate or in large leaves the basal pinnae with very few basal lobes *).

For *) intermediates see 12a. 548 BLUMEA VOL. XV, No. 2, 1967

Rachis sori laminal obtuse lobes. 9. abaxiaUy sharply carinate; or on short,

T. 12. pinnatum

Rachis T. 9. abaxially bi-angular; sori intramarginal, on saw-teeth 15. carolinense

Lamina 3. at least pinnate + pinnatifid; primary rachis abaxially terete or bi-angular, or, if obtusely the carinate, petiole abaxially not or only at the apex sharply bi-angular.

rachis rachises the basal 10. Primary atropurpureous; secondary (except sometimes ones) abruptly the basal pale; pinnae pinnatifid, with crenate segments, only pinnae occasionally with some

with their their pinnatifid basiscopic pinnules; most sori greatest extension at right angles to vein. T. calomelanos 9.

rachis least base rachises TO. Primary at at dark; secondary pale; pinnae pinnate + pinnatifid or

sori with their extension their vein. T. subbipinnate; greatest parallel to . . 5. stenocarpum

rachis if 10. Primary pale, or, dark, the secondary rachises not abruptly pale; lamina often more incised.

the 11. All axes, except primary, green-marginedto base or almost so, i.e., lamina only once

fully pinnate, then pinnatifid; secondary axes abaxially rounded. . . 2. T. buniifolium

11. Lamina mostly fully bipinnate; secondary rachises, if marginate, abaxially carinate.

12. Secondary rachises abaxially black, with two pale lateral or one median pale ridge;

lamina bipinnate or almost so, with superficially crenato-lobate pinnules; pinnae not

enlarged at base 7. T. atratum

rachises 12. Secondary abaxially various but not black with pale ridges; lamina often bipinnate -f pinnatifid.

rachis and indusia ultimate lobes with broad and 13. Primary black; abaxially very prominent veins occupying |—J- of their width 6. T. obtusatum

dark 13. Primary rachis and indusia pale to brown; ultimate lobes with immersed, or,

if much veins. slightly prominent, relatively narrower

14. Indusia about twice as broad as long, 0.3 mm long; margin bordering the

apical sorus of the segment usually denticulate; textureherbaceous.

1. T. tenue

14. Indusia longer, or, if only J mm long, ± isodiametric or longer than broad;

margin bordering the apical sorus entire, or no apical sorus; texture firmer.

each lobe with a sorus 15. Larger segments pinnatifid, overtopped by part

of the lobe 3. T. amboynense

for the basal each lobe with 15. Larger segments (except pinnae) crenate, a

terminal or subterminal sorus.

16. Secondary rachises abaxially terete in a considerable basal portion;

basiscopic pinnules of basal pinnae usually enlarged and more dissected

than the others 4. T. novoguineense

16. At least the larger secondary rachises abaxially carinate; basiscopic

of basal and dissected. pinnules pinnae rarely enlarged more

II. T. luzonicum

I. Tapeinidium tenue (Brackenridge) Copeland, B.P. Bishop Mus. Bull. 59 (1929)

69. — Basionym: Microlepia tenuis Brackenridge, U. S. Expl. Exped. (1854) 236. Type:

Sandalwood Bay, Fiji (Vanua Levu?), U. S. Explor. Exped. 3 (US).

Davallia denhami Hooker, 2nd Cent. Ferns (i860) pi. 47. — Microlepia denhami (Hooker)

Ind. — Moore, Fil. (1861) 292. Lindsaea denhami (Hooker) Mettenius ex Kuhn, Verh.

Zool. Bot. Ges. 19 (1869) 573. — Wibelia denhami (Hooker) Kuhn, Chaetopt. (1882)

346. — Tapeinidium denhami (Hooker) C. Christensen, Ind. Fil. (1906) 631. — Type: Milne 116, Viti Levu, Fiji (K).

Tapeinidium tenuius Copeland, Philip. J. Sc. 60 (1936) 110, pi. 17. — Type: Brass 3023, San Cristoval, Solomon Is (MICH).

The and the uni- relatively narrow scales, only up to 4 cells wide greater part or and the well biseriate, the denticulate lobes, sori that are often terminal as as lateral. K. U. KRAMER: A revision Tapeinidium of 549

together with the thin characterize this texture, species rather well. In the type the teeth of the lobes are almost, the terminal sori quite lacking, but otherwise it matches the other well. In size and degree of dissection specimens it is quite variable; except for the

two the are smaller and less types Fijian specimens incised than the plants from the western of the part range.

Distribution Seen from Is Bismarck : : Admiralty (Manus), Archipelago (New Hannover,

New Ireland), Solomon Is (San Cristóval, Santa Ysabel, Guadalcanal), New Hebrides

(Aneityum, Banks I.), Fiji (Viti Levu, Vanua Levu, Ovalau). Specimens labelled 'New Caledonia' of and 'Isle Pines' are probably from the New Hebrides (Kramer, 1967).

2. buniifolium Tapeinidium — T. Kramer, spec. nov. moluccanum auct. non (Blume)

C. Chr.; & Univ. Calif. Publ. Wagner Grether, Bot. 23 (1948) 36. — Type: Grether & Wagner 4188, Manus, Admiralty Is, Tjajiak Mts, Mt Dremsl region, in mountainside woods, c. 2000 ft. (MICH; isotype US).

Rhizomatis perbreves solum adsunt; rubro-fuscae, partes squamae pro parte majore uni- vel basi latiores. aciculares, biseriatae, paullo Petioli abaxialiter, ut videtur, basi

subteretes, obtuse Lamina supra biangulares. probabiliter deltoidea, pinnata et quadri- pinnatifida vel basi subbipinnata et tripinnatifida; apex laminae deest, probabiliter pinna tifidus. Pinnae minime primariae 4 pro latere, majores oblongae, breviter petio- lulatae, secundariis latere pinnis majoribus pro circ. 12—15. Rhachis primaria straminea ad brunnea, abaxialiter, ut videtur, biangularis; rhachides ceterae abaxialiter stramineae, ad teretes, anguste viridi-marginatae basim vel basales fere ad basim. Segmenta lanceolata vel linearia, chartacea, basi profunde pinnatifida, apice dentato-lobata. Lobi ultimi ad majores lanceolato-ligulati, 4 X if mm, subacuti, saepe subfalcato-adscendentes, ubi lobi asymmetrici, margine anteriore, sorus, gibboso; minores triangulares, soro

subterminali. Venae simplices vel in lobis majoribus furcatae. Sori uninervii; indusium basim brimnescens, marsupiiforme, versus angustatum, longitudine et latitudine ± aequis. Remarks. known from the Only type collection, as cited above. The most finely

dissected species, except for the larger forms of T. tenue from which it differs, by , i.a., all axes being marginate to the base or almost so.

Tapeinidium Index Filicum 3. amboynense (Hooker) C. Christensen, (1906) 631. — Davallia Hooker, — Basionym: amboynensis Spec. Fil. 1 (1846) 178, t. 56 C. Lindsaea

amboynensis Mett. ex Kuhn, Ann. Lugd. — Wibelia (Hooker) Bat. 4 (1869) 279. amboy-

nensis (Hooker) Kuhn, — A. Smith Chaetopt. (1882) 346. Lectotype: s.n., Ambon (Amboyna), 'from P. B. Webb' (K).

Davallia stenoloba Baker in Malesia Beccari, III (1886) 35. — Tapeinidium moluccanum Chr. (Blume) C. var. stenolobum (Baker) C. Chr., Index Filicum Suppl. Ill (1934) 176.

— & Univ. Calif. Publ. Tapeinidium stenolobum (Baker) Wagner Grether, Bot. 23 (1948)

36. — Type: Beccari s.n., Mt Salhutu, Ambon, 1000 m (FI, fragm. K).

Tapeinidium moluccanum C. Chr., Index Filicum Suppl. Ill (1934) 176, and of nearly all later authors; not Davallia moluccana Blume, Enum. (1828) 237 (= Saccoloma spec.).

Tapeinidium amplum Copeland, Occ. Bishop f. — Papers Mus. 15 (1939) 82, 3. Type: Takamatsu Palau Is. 1572, Garasumao, (MICH; isotypes BISH, K, US). from Distribution: Seen Celebes, the Moluccas (Talaud, Morotai, Halmahera, Ternate,

Ambon, Ceram), the Kei Is, Waigeo, Biak, Western New Guinea, and the Palau Is.

A specimen in BO labelled 'Borneo Nieuwenhuis? 166' is probably from elsewhere.

Remarks. The of Davallia moluccana Blume the type at L was examined by writer; 550 BLUMEA VOL. XV, No. 2, 1967

it is in habit similar T. a Saccoloma, not a Tapeinidium, though very to amboynense.

Ithycaulon, based on D. moluccana, is therefore a of Saccoloma, not ofTapeinidium, , synonym

as was recently stated.

Related to T. stenocarpum, T. tenue, and T. luzonicum; the character stated in the key

under heading 15 indentifies this species quite readily.

— Schlechter Torricelli 4- Tapeinidium novoguineense Kramer, spec. nov. Type : 14319, of Mts, Territory New Guinea, 700 m (B, 2 sheets; isotypes BM, BO, K, P). Rhizoma breviter vel pro genere longius repens, squamis brunneis, pro parte majore

ad mm Petiolus vel abaxialiter pluriseriatis, 3 longis vestitum. stramineus obscurior, Lamina teres. bipinnata, vel bipinnata et pinnatifida, basi nonnumquam tripinnata, elongato-triangularis vel subquinquangularis-triangularis, rarius oblonga; rhachis primaria

straminea vel pallide brunnea, abaxialiter teres vel angustato-teretiuscula, ad apicem carinata. Pinnae anteriore tantum majores 15—35 pro latere, basi latere latiores, pinnatae,

cm basales 12—16 longae, pinnulis ca. 20—30 pro latere; pinnulae posteriores pinnarum

basalium plerumque valde protractae, magis incisae. Rhachides secundariae (et ceterae)

stramineae, abaxialiter infra teretes, supra carinatae. Pinnulae ultimae liberae circ.

subcoriaceae vel 20—30 pro latere, coriaceae, nonnumquam statu sicco revolutae,

lanceolatae, obtusae vel subacutae, majores vulgo 3—4 cm longae, 3—4 mm latae,

crenatae vel pinnatifidae. Venae in lobis simplices vel in majoribus furcatae, subpro-

minulae. Pinnae et pinnulae superiores abbreviatae, confluentes. Sori singuli in lobis vel rarius bini vel ultra, uninervii; indusiummarsupiiforme ad subtriangulare, marginem

non attingens, a marginibus lateralibus et apicalis lobi aequidistans.

Distribution: Japen, Western New Guinea, Papua, and the Territory of New Guinea. labels often be Remarks: This is a widespread species in New Guinea, on said to

have almost different and a few doubtful locally common. I seen 50 collections, ones. The T. sheets were usually labelled ‘T. moluccanum’. It seems very close to stenocarpum

which may be only a form ofit; this is difficult to judge as I have seenonly two collections

of the latter. But the obtusely bi-angular petiole and rachis with paler edges, the linear

ultimateand sori of T. be penultimate segments, and the narrower stenocarpum seem to

distinctive enough. T. amboynense is undoubtedly also very closely related.

Guinea 5. Tapeinidium stenocarpum van Alderwerelt van Rosenburgh, Nova 14

— mountain ridge near Idenburg (1924) 52. Type: Lam 1442, W. New Guinea, R., alt. terrestrial in swampy Vaccinium forest, 1420 m (BO; isotypes K, L, SING, U; fragm. US).

Remarks. Except for T. tenue and T. buniifolium this is the most finely dissected species. for It seems to be closest to T. novoguineense; the differences see the key and the note

under the last-named species. from the collection Brass also from Apart type only one seen: 12102 (GH, MICH),

W. New Guinea.

6. Tapeinidium obtusatum van Alderwerelt van Rosenburgh, Nova Guinea 14 (1924)

52. — Type: Lam 1837, W. New Guinea, mountainridge near Doormantop, terrestrial

alt. m SING, in forest, 2500 (BO; isotypes L, U). known Remarks. This distinctive species, only by its type collection, is readily

identified by the characters described in the key. It is the most coriaceous one of the

finely dissected species. Its affinity is uncertain. K. U. KRAMER: A revision of Tapeinidium 551

— Lam W. New J. Tapeinidium atratum Kramer, spec. nov. Type: 1556, Guinea,

mountain ridge near Doormantop, 1420 m (BO, 2 sheets; isotypes L, SING, U). aciculari Rhizoma pro genere longius repens, squamis anguste triangularibus, apice

longo, ad 2 mm longis obtectum. Petiolus atratus, abaxialiter supra acute biangularis,

infra Lamina costis haud pallidioribus, sensim teretiusculus. bipinnata, pinnis pro latere breviter circ. 15—20, remotis, adscendentibus, anguste deltoideis, acuminatis, pinnatis,

supra pinnatifidis; rhachis primaria atrata, abaxialiter acute biangularis, supra costis pallidioribus. Rhachides secundariae obscurae, abaxialiter costis pallidis duabus, lateralibus,

vel singulis, medialibus. Pinnae superiores abbreviatae, confluentes? Pinnulae rigide sicco ad coriaceae, statu fuscae, ad 35 pro latere, angustissime lanceolatae, 17 X 4 mm,

pinnatilobatae, lobis pro latere circ. 9, late rotundatis, ad I X 1 mm, apice obtusae, basi

cuneatae, decurrentes. Costae pallidae, abaxialiter prominulae, obtusae. Venae immersae,

simplices vel unifurcatae. Sori singuli, rarissime bini vel terniin lobis, uninervii, prope

marginem anteriorem, paulum sed manifeste sub apice; indusiumatratum, marsupiiforme. Remarks. The regularly pinnate lamina with only shortly acuminate pinnae and

especially the dark colour of the axes and indusia, with the secondary racbises abaxially

its pale-margined, are distinctive for this species which is only known by type collection.

8. Tapeinidium gracile (Blume) van Alderwerelt van Rosenburgh, Malayan Ferns

(1909) 315. — Basionym: Davallia gracilis Blume, Enum. (1828) 233. — Microlepia

gracilis (Blume) J. Smith, London Jo. Bot. I (1842) 427. — Davallia pinnata Cav. var.

gracilis (Blume) Baker, Syn. Fil. ist ed. (1867) 98, comb, valid. ? — Wibelia gracilis (Blume)

Christ, Ann. Jard. Buitenz. II, 5 (1905) 134. — Type: Blume 1731 or s.n. from Java (L).

Distribution: Indo- (one coll. from Nhatrang, Annam), West Java, Bali, Borneo

(one coll. each from Brunei and Sarawak), Celebes, Ceram, and the Philippine Is (Luzon,

Sulu and The to be Negros, Mindanao, Archip., Leyte, Mindoro). species seems most in and absence from Peninsula is common Java Luzon. Its Sumatraand the Malay rather

surprising.

Remarks. Rather easily recognized by the pinnate + pinnatifid lamina and dark, carinate abaxially pale-margined axes, the bi-angular petiole, and the primary rachis. be that the for this lineare It may correct name species is Tapeinidium (Cav.) C. Chr.; I have the Dicksonia linaeris not seen type of Cav. On a photograph kindly sent from Madrid the by Dr E. Paunero the specimen looks most like T. ‘biserratum’, but character

of the axes which I regard as crucial is not visible.

calomelanos — Korthals from 9. Tapeinidium Kramer, spec. nov. Type: s.n. G.

Sakoembang, S.E. Borneo (L, 2 sheets).

Rhizoma breviter vel (pro genere) longius repens, squamis badiis elongato-triangulari-

bus longe acuminatis ad 2 mm longis vestitum. Petioli pullo-atropurpurei, abaxialiter

teretes. Laminapinnata et profunde pinnatifida vel basi bipinnata (et pinnatifida), elongato- vel vel subcoriacea vel triangularis oblonga subquinquangularis, 10—35 cm longa, Rhachis vel obscure abaxialiter coriacea. primaria atropurpurea rarius badia, teres.

Rhachides secundariae (et ceterae) abrupte pallidae (interdum praeter basales), abaxialiter

omnino teretes. Pinnae majores 12—20 pro latere, lanceolatae, inferiores vulgo trian- vel interdum gulares, subsessiles, acuminatae, profunde pinnatifidae basi pinnatae et

tunc pinnulae majores basiscopicae interdumpinnatifidae, ceteraepinnatilobatae; segmenta

(pinnulae) majores ad 18 pro latere, adscendentia, lanceolata, serrata vel crenata, basi

inaequaliter cuneata, obtusa vel subacuta, ad apicem mox decrescentia. Pinnaead apicem laminae confluentes. Venae simplices vel in lobis majoribus furcatae, immersae vel 552 BLUMEA VOL. XV, No. 2, 1967

paulum prominulae. Sori uninervii, singuli in lobis; indusium obscurum, vulgo longius quam latum, marginem non attingens. Sumatra Borneo the Korthals Distribution: (locality perhaps incorrect), (beside type s.n. without locality, B, BM, L), Celebes ( Elbert 3113 BO, K, L, SING), and Luzon ,

(Alcasid & Edaño 5083 and Merrill 933, both MICH). hesitated before Remarks. 1 have long describing this as new; it is known from a few scattered, mostly rather old collections. Its leafarchitecture is rather like that of certain forms of T. luzonicum, but the axes are entirely different. The dark, abaxially terete primary and the abruptly pale secondary axes are unique in the genus. The Philippine collections are smaller than the other ones.

10. Tapeinidium oligophlebium (Baker) C. Christensen, IndexFilicum (1906) 631. —

Basionym: Davallia oligophlebia Baker, Jo. Bot. 26 (1888) 323. — Wibelia oligophlebia

— Ch. (Baker) Christ, Ann. Jard. Buitenz. II, 5 (1905) 134. Type: Hose 220, Sarawak, Laupi (K).

Protolindsaya brooksii Copeland, Philip. J. Sc. 5, Bot. (1910) 283. — Tapeinidium brooksii Index Filicum Brooks (Copel.) C. Chr., Suppl. Ill (1934) 176. — Type: 47, Sarawak, with the Gunong Bengkaim (SAR, holotype?; isotype BM, 2 sheets; specimens same number but different dates in BM, K, and P).

small like form This species looks suspiciously a depauperate but fertile of another luzonicum. species, e.g., T. But the almost 20 collections I have seen are all from Borneo which is from whereas T. luzonicum (Sarawak, except one Kalimantan), occurs throughout western and central Malesia. This leads to the conclusion that it is a taxon that deserves recognition, although it may not be a distinct species. The of brooksii has almost linear laminas with short type Protolindsaya very pinnules, but two collections by Richards ( K, SING, U, US, and K) comiect it with 2024, 1747, typical T. oligophlebium.

— Davallia 11. Tapeinidium luzonicum (Hooker) Kramer, comb. nov. Basionym: luzonica Fil. I — Gen. Hooker, Spec. (1846) 174, t. 60 B, f. 2, 3, 5. Wibelia bipinnata Fee,

Fil. — (1852) 331, nom. superfl. Type: Cuming 139 (K ?; isotypes B, L).

Lindsaea pinnata (Cav.) Mett. ex Kuhn var. bipinnata Mett. ex Kuhn, Ann. Mus. Lugd.

Bat. 4 (1869) 279. — Lectotype: Zollinger 1305, Java (HBG, L, isotypes).

Davallia Lond. — philippinensis Harrington, J. Linn. Soc. 16 (1877) 27. Microlepia

philippinensisTapeinidium(Harrington) Copeland, Polypod. Philipp. (1905) 56. — Index philippinense (Harrington) C. Chr., Filicum Suppl. Ill (1934) 176. — Type: Steere s.n. Luzon, Mt Mahayhay (K).

Davallia hosei 26 — Baker, J. Bot. (1888) 323. Type: Ch. Hose 219, Sarawak, Lambur (K).

Tapeinidium sumatranum van Alderwerelt van Rosenburgh, Bull. Jard. Bot. Buitenz.

HI. 2 (1920) 174. — Type: Brooks 332/S, Sumatra, Benkulen (BO, fragm.). biserratum Flora and of other Tapeinidium sensu Holttum, Rev. Malaya II (1954) 339, authors; not Davallia biserrata Blume.

Distribution: (1 coll.), Malaya (Pahang, Kelantan, Trengganu, Perak, Johore),

Natuna Is, Lingga Is, Banka, Sumatra, West Java (few coll.), Borneo (Sabah, Brunei, Sarawak, Kalimantan), Celebes, and the Philippine Is (Luzon, Mindanao, and Polillo).

Remarks. This widespread species is readily identified by the pale petiole which is least the the abaxially bi-angular at in upper part, pale, abaxially sharply carinate primary and secondary rachises and costa, and by the at least pinnate + deeply pinnatifid lamina. K. U. KRAMER: A revision of Tapeinidium 553

T. and there intermediates It may only be a deeply incised form of" pinnatum, are some

(see under ‘T. biserratum’, after the following species), but, as already noted by Holttum

of T. luzonicum and of T. are (I.e., p. 340), typical specimens pinnatum so overwhelmingly more numerous than intermediates that I prefer to treat them as distinct.

The above applies to var. luzonicum. Although it seems dangerous to describe varieties in differences the a genus where many species are distinguished by rather slight only, of luzonicum following two forms which are distinct from the bulk of the specimens T. differ by such superficial characters that I prefer to call them varieties; they are also geographically much more restricted.

a. — Elmer Urdaneta var. leptophyllum Kramer, var. nov. Type: 14103, Mindanao, Mt (L; isotypes BO, HBG, MICH).

Differt a varietate luzonica petiolis gracilioribus, basi laminae maxime 1 mm crassis,

basi saepe brunncis; lamina graciliore, subtripinnata, habitu earn Pityrogrammae calo- melaneos simulante, segmentis ultimis liberis minoribus, anguste lanceolatis, plurimis

ultra in lobis non 1 cm longis, 1—2 mm latis; venis magis conspicuis, saepe simplicibus; soris vel in singulis binis lobis saepe regulariter rotundatis; indusiis ca. 0.3 x 0.3 mm.

PHILIPPINES. Edaño Negros: Whitford 1498 (MICH); 21429 (MICH), id. 5952 (GH,juvenile, doubtful).—

Mindanao: Elmer Loher 1117 (K, US); Mendoza & Convocar 8711 (MICH, SING); Warburg 14157 (B);

14103 (type, L, BO, HBG, MICH). — Camaguin: Ramos 14878 (B, BO, K, L, SING, US). — Panay:

Ramos — —Luzon: & Edaño 50666 (B, BM, BO, L, US). Leyte: Wenzel 551 (GH), 777 (MICH). Micholitz s.n. (GH, K); Vidal s.n. (K); Weiss 4107 (L); Elmer 17852 (K, p.p. mai.).

This unlike Remarks. variety is not T. tenue but differs in some important characters like the non-dentate soriferous lobes and the structure of the rachis. The differences from

and therefore better typical T. luzonicum are quantitative rather than qualitative it seems to treat it as a variety.

b. — var. thelypteridoides Kramer, var. nov. Type: W. M. A. Brooke 81 go, Sarawak,

Mt Santubong, 2000 ft, on damp rock in forest (L; isotypes SING, US). varietate luzonica solum vel Recedit a lamina pinnata + pinnatifida pinnatilobata, leviter pirniis regulariter incisis, infimis maxime ad f, lobis oblongo-ligulatis, serrato- crenatis; textura chartacea; venis manifestis; soris saepe magis inframarginalibus. Habitu speciei Thelypteridis, e.g., Th. dentatae, sat similis.

BORNEO. Sarawak: W. M. A. Brooke 8190 (type). North Borneo: W. Meijer 560 (BO, L), 562 (BO,

id. K, L); SAN 25295 (K, L).

almost Remarks. This may be a species in its own right, but a few specimens bridge makes the gap between it and var. luzonicum which treatment as a variety preferable.

12. Tapeinidium pinnatum (Cav.) C. Christensen, Index Filicum (1906) 631. —

Basionym: Davallia pinnata Cavanilles, Descr. (1802) 277 ( non Mett. ex Kuhn, 1869). —

Saccoloma pinnatum (Cav.) Presl, Tent. Pteridogr. (1836) 126. — Microlepia pinnata

Hooker's — Wibelia Bernhardi (Cav.) J. Smith, Jo. Bot. 3 (1841) 416. pinnata (Cav.)

Fil. bis — Lindsaea ex Fee, Gen. (1852) 331, t. 27 B. pinnata (Cav.) Mett. ex Kuhn, Ann.

Lugd. Bat. 4 (1869) 279. — Type: L. Née s.n., Philippine Is (also incorr. cited 'Chile')

(MA, not seen; photogr. U).

Davallia flagellifera Hooker & Greville, Ic. Fil. (1831) t. 183, ex char. — Type: Wallich s.n., Pulu Penang, not seen. 554 BLUMEA VOL. XV, No. 2, 1967

Davallia serrata Roxburgh ex Griffith, Calc. Jo. 4 (1844) 514 (non Willdenow, 1810),

— of Wales' Island ex char. Type: Roxburgh s.n., Prince (Pulu Penang), not seen.

Gen. Fil. char. — Wibelia javae Fée, (1852) 331, ex Type: Kollmann s.n., Java, not seen.

Davallia — firmula Baker, Ann. Bot. 8 (1894) 123. Microlepia firmula (Baker) C. Chr.,

Index Filicum (1906) 426. — Tapeinidium firmulum (Baker) C. Chr., Index Filicum

Ill — W. Hancock Barisan Luva Suppl. (1934) 176. Type: 72, Sumatra, Range, near (?) (K). Distribution: India S. (Kerala, one coll.), the Ryukyus, , Thailand, the Malay

Peninsula (Pahang, Kelantan, Selangor, Malacca, Negri Sembilan, Johore, Perak, P.

Penang), Singapore, the Riau Is, Sumatra, the Lingga Is, Banka, W. Java, Borneo

(Sabah, Sarawak, Kalimantan), Celebes, and the Philippine Is (Sulu Archip., Leyte,

Basilan, Polillo, Mindanao, Bucas Grande, Palawan, Biliran, Mindoro, Panay, Luzon). This Remarks. is the most common and widespread species in the genus, distinguished

by pale to medium brown, abaxially at least above bi-angular petiole, abaxially sharply lamina with the confluent carinate rachis, simply pinnate upper pinnae into a pinnatifid

leaf-apex, serrate or crenate linear pinnae, and laminal sori, i.e., not so close to the margin

as to be in the lobes.

12a. Tapeinidium biserratum (Blume) van Alderwerelt van Rosenburgh, Malayan

Davallia — Ferns Suppl. (1917) 509. — Basionym: biserrata Blume, Enum. (1828) 232.

biserrata — Blume Microlepia (Blume) Presl, Epimel. Botan. (1849) 97. Type: s.n., Java (L). Remarks. of Davallia between Blume's type biserrata agrees with a series ofintermediates

T. luzonicum and T. There between and of these pinnatum. are 25 30 intermediates,

relatively very few in comparison with the large series of typical specimens of both their species. They occur throughout common range: Malaya, Sumatra, West Java, and the have the dissection often found Borneo, Philippine Is. Some irregular pattern Scortechini and Kunstler in hybrids (Wagner, 1962; Meyer, 1965), e.g. s.n. (BM) 2144

(SING), both from Perak. In these and other intermediates I failed to find anything

but what seemed to be normally developed spores. There is thus no direct evidence

of and hybrid origin, if the two species are so closely related as to have fertile hybrid wonders barrier that offspring one what keeps them largely apart. My impression is

biserratum a aberrant form of but from a T. is phenotypic, T. pinnatum, apart specimen with leaves of the of those of biserratum rhizome shape T. pinnatum with T. on one

I have for Field studies (Micholitz s.n., Philippines, K), no proof this. might help to

find solution for the — also the the end of the a problem. See note at present paper.

13- Tapeinidium prionoides Kramer, spec. nov. — Type: Bünnemeijer 1910, Banka, G. Siang, Sungai Liât (L; isotype BO). differt rhachide Tapeinidio pinnato simile, abaxialiter haud carinata, saepe angustato- dentibus basali teretiuscula, petiolo obscuro; pinnis serratis, adscendentibus, saepe majore, soris dentes insidentibus, indusio marginem fere attingente.

CHINA Henderson SOUTH SEA ISLANDS. Anambas Is: 20445 (BO, SING). — Riau Is: Bünnemeijer 7886

(BO, L), 7887 (BO, L); Fox s.n. (SING); Schlechter 13635 (B). — Lingga Is: Bünnemeijer6632 (BO,L,U).— Banka: Bünnemeijer 2135 (BO, L), 1910 (type, L, BO).

Remarks. The I the specimens saw had been determined as T. pinnatum, but features described the in Latin diagnosis exclude them. The species has a remarkable distribution, of occurring on some small islands to the East Malaya and Sumatra.

acuminatum 14. Tapeinidium Kramer, spec. nov. —Type: Escritor 21173, Philippine _ Is, Luzon, Isabela Prov., Palanan Bay (L; isotypes BO, BRJ, GH, MICH). K. U. KRAMER: A revision of Tapeinidium 555

Tapeinidio pinnato simile, differt lamina imparipinnata, scilicet pinna terminali con- formi; rhachide abaxialiter biangulari, costis duabus lateralibus, pallidis, nec carinata,

ut in T. pinnato.

Remarks. known from the but two collections from North Only type collection; also this robust and have Borneo may belong to species. They are, however, more Wood Lahad the paler axes; one, 2007, Datu Distr. (K), is incompletely fertile, other, W. SAN Meijer 20989, Ranau Distr. (K), is sterile. These two collections may even the Wood collection have abortive represent another, undescribed species. The type and indicationof what the spores, an hybrid origin, but I am unable to suggest parent species have been. may

carolinense — Kanehira Caroline 15. Tapeinidium Kramer, spec. twv. Type: 1615,

Is, Ponape (US, 2 sheets).

Tapeinidio pinnato simile, differt rhachide abaxialiter bi-angulari nec carinata, sorisque serriformes rhizomatis margine plus approximatis, dentes insidentibus; squamae 5 mm

longae; sporae perbreviter ellipsoidales.

CAROLINES. Takamatsu Ponape: Finsch 29 (B); Kanehira 79s (BISH), 1349 (K), 1613 (type, US); 934 (BISH, BO, MICH, US); Ledermann 13436 (B, K), 13678 (B, K), 13800a (B, K); Stone 3384 (U).

Remarks. In spite of its similarity to T. pinnatum this species is easily recognized by the

above-mentioned characters. It also shares some features with T. longipinnulum. It seems

be have from to fairly common in Ponape; surprisingly I not seen any specimen Kusaie.

16. Tapeinidium longipinnulum (Cesati) C. Christensen, Index Filicum Suppl. Ill

— Rendic. Accad. Fis. (1934) 176. Basionym: Davallia longipinnula Cesati, R. Sei. e Mat.

16 — Napoli (1877) 29; Beccari, Malesia III (1886) 35. Type: Beccari s.n., New Guinea,

Ramoi (FI).

Davallia I.e. — intramarginalis Cesati, (1877) 29. Type: Beccari s.n., New Guinea,

'Mt Arfak a Putat' (FI).

Tapeinidium marginale Copeland, Philipp. J. Sc. 6, Bot. (1911) 82. — Type: King 283

New Guinea, Papua (MICH; isotype P).

Remarks. This species is often confused with T. pinnatum, but it seems to be much from closer to T. melanesicum. The conform terminal pinna distinguishes it at once the

first-named species; for the differences with T. melanesicum see the key. have T. longipinnulum is apparently not uncommon in New Guinea; I seen many

collections from W. New Guinea and smaller series from the Territory of New Guinea

and Papua, furthermore from Woodlark and Rossel Islands off eastern Papua, and three from Buwalda and collections Ceram: 5907 (BO, K, L) andKornassi 67 (BO) 712 (BO, L).

melanesicum 17. Tapeinidium Kramer, spec. nov. —Type: Stone 2312, Solomon Is, Santa Ysabcl, Horara Islet, Tatamba Lagoon (U; isotypcs BISH, US). Rhizoma brevitcr rcpcns, squamis pullo-brunncis, pro parte majore uni-biseriatis, Petiolus brunncus ad 11 mm longis vestitum. ad purpureo-castaneus, abaxialiter teres

vel ad apicem breviter obtuseque biangularis, ecanaliculatus et concolor. Laminaoblonga, terminali simpliciter pinnata, pinnis plerumque 10—15 pro latere et impari conformi;

rhachis straminea vel fusca, abaxialiter teres vel obtuse biangularis. Pinnae coriaceae,

angustissime lanceolatae, acutae vel breviter acuminatae, 10—25 cm longae, 6—15 mm

latae, leviter et regularitcr crenatae vcl bis crenatac, lobis valdc adsccndentibus. Costa 556 BLUMEA VOL. XV, No. 2, 1967

abaxialiter prominens, obtusa; venae prominulae, maxime obliquae, uni- vel rarius vel sub ad bifurcatae. Sori singuli bini, lobis eosque leviter ingredientes vel apicem pinnae lobos insedientes, uni- vel rarius binervii. Indusium pallidum vel brunneum, marsupiiforme, marginem non attingens.

T. T. Remarks. This species was formerly confused with longipinnulum or pinnatum; the differences are stated in the key.

I have seen 28 collections which will be cited in the account of the Lindsaeoid ferns of the smaller Islands. from Solomon Pacific They are the Is (Santa Ysabel, Santa Cruz,

San Cristobal), the New Hebrides, the Santa Cruz Islands (Vanikoro), and Fiji (Viti

Levu, Vanua Levu, and Ovalau).

SPECIMENS OF UNCERTAIN IDENTITY

often work number of As in monographic a problematic specimens remain. Note-

these a of from the Islands that resemble worthy among is series plants Philippine T. gracile in most characters, notably those of the axes, but have much less incised pinnae, not unlike those of T. ‘biserratum’. but the to be They may represent hybrids, spores seem normally developed. It is not impossible that they belong to an undescribed, endemic

which the of T. lineare also The fact that Philippine species to type may belong. they share characters of two species makes me refrain from describing them here as a species.

Good Elmer from and Elmer examples are: 18107 (BO, GH, MICH, US) Luzon, iogi2 and and Sulit all from Mindanao. (BO, L, MICH, US) 7P73 (BO, L) 8732 (MICH, SING),

EXCLUDED SPECIES

Bot. stortii Tapeinidiumbartlettii Copeland, Univ. Calif. Publ. 14 (1929) 376, pi. 60 =Xyropteris

(v. A. v. R.) Kramer.

Ill C. Index Filicum = Saccoloma Tapeinidium moluccanum (Blume) Chr., Suppl. (1934) 176 sp. (see under T. amboynense).

Fournier. Tapeinidiummoorei (Hooker) Hieronymus, Hedwigia 62 (1920) 13 = Lindsaea moorei (Hooker) See Kramer 1967, p. 569.

REFERENCES

ALDERWERELT VAN ROSENBURGH, C. R. W. K. VAN. 1909. Malayan Ferns. Batavia.

COPELAND, E. B. 1947. Genera Filicum. Waltham, Mass.

4 DIELS, L. 1902. Polypodiaceae, in: Engler & Prantl, Die natiirlichen Pflanzenfamilien I . F£E, A. L. A. 1852. Genera Filicum. Paris & Strasbourg.

KRAMER, K. U. A revision of the Lindsaea in the New World with notes allied 1957a. genus on genera.

Acta Bot. Neerl. 6: 97—290.

A of Lindsaeoid ferns. Ibid. 1957b. new genus 6: 599—601.

1967. The Lindsaeoid ferns of the Old World. I. New Caledonia. Ibid. 15: 562 —584.

D. E. MEYER, 1965. Zum morphogenetischen Prinzip der Irregularitatvon Artbastarden und Bastardarten,

voin Blickpunkt der Systematik. Willdenowia 4: 63—73. WAGNER, W. H. Jr. 1962. Irregular morphological development in hybrid ferns. Phytomorphology

12: 87 —100.