The Lindsaeoid ferns of the Old World II. A revision of Tapeinidium K.U. Kramer Botanical Museum and Herbarium, Utrecht INTRODUCTION the second the Lindsaeoid fern 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 plants 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 India and the Ryukyu Islands to Fiji; it is absent from Ceylon, Australia, 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.