Flora of tropical East Africa - Woodsiaceae (2003) (Flora of Tropical East Africa)

LIST OF ABBREVIATIONS A.V.P.=O.Hedberg, Afroalpine Vascular Plants; B.J.B.B.=Bulletin du Jardin Botanique de l’Etat, Bruxelles; Bulletin du Jardin Botanique Nationale de Belgique; B.S.B.B.=Bulletin de la Société Royale de Botanique de Belgique; C.F.A.= Conspectus Florae Angolensis; E.J.=A.Engler, Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie; E.M.=A.Engler, Monographieen Afrikanischer Pflanzen-Familien und Gattungen; E.P.=A.Engler, Das Pflanzenreich; E.P.A.=G.Cufodontis, Enumeratio Plantarum Aethiopiae Spermatophyta; in B.J.B.B. 23, Suppl. (1953) et seq.; E. & P. Pf.=A.Engler & K.Prantl, Die Natürlichen Pflanzenfamilien; F.A.C.=Flore d’Afrique Centrale (formerly F.C.B.); F.C.B.=Flore du Congo Belge et du Ruanda-Urundi; Flore du Congo, du Rwanda et du Burundi; F.D.O.A.=A.Peter, Flora von Deutsch-Ostafrika; F.F.N.R.=F.White, Forest Flora of Northern Rhodesia; F.P.N.A.=W. Robyns, Flores des Spermatophytes du Pare National Albert; F.P.S.=F.W.Andrews, Flowering Plants of the Anglo-Egyptian Sudan or Flowering Plants of the Sudan; F.P.U.=E.Lind & A.Tallantire, Some Common Flowering Plants of Uganda; F.R.=F.Fedde, Repertorium Speciorum Novarum Regni Vegetabilis; F.S.A.=Flora of Southern Africa; F.T.A. =Flora of Tropical Africa; F.W.T.A.=Flora of West Tropical Africa; F.Z.=Flora Zambesiaca; G.F.P.=J.Hutchinson, The Genera of Flowering Plants; G.P.=G.Bentham & J.D.Hooker, Genera Plantarum; G.T.=D.M.Napper, Grasses of Tanganyika; I.G.U.=K.W.Harker & D.M.Napper, An Illustrated Guide to the Grasses of Uganda; I.T.U.=W.J.Eggeling, Indigenous Trees of the Uganda Protectorate; J.B.=Journal of Botany; J.L.S.=Journal of the Linnean Society of London, Botany; K.B.=Kew Bulletin, or Bulletin of Miscellaneous Information, Kew; K.T.S.=I.Dale & P.J.Greenway, Kenya Trees and Shrubs; K.T.S.L.=H.J.Beentje, Kenya Trees, Shrubs and Lianas; L.T.A.=E.G.Baker, Leguminosae of Tropical Africa; N.B.G.B.=Notizblatt des Botanischen Gartens und Museums zu Berlin-Dahlem; P.O.A.=A.Engler, Die Pflanzenwelt Ost-Afrikas und der Nachbargebiete; R.K.G.=A.V.Bogdan, A Revised List of Kenya Grasses; T.S.K.=E.Battiscombe, Trees and Shrubs of Kenya Colony; T.T.C.L.=J.P.M.Brenan, Check-lists of the Forest Trees and Shrubs of the British Empire no. 5, part II, Tanganyika Territory; U.K.W.F.=A.D.Q.Agnew (or for ed. 2, A.D.Q.Agnew & S.Agnew), Upland Kenya Wild Flowers; U.O.P.Z.=R.O.Williams, Useful and Ornamental Plants in Zanzibar and Pemba; V.E.=A.Engler & O. Drude, Die Vegetation der Erde, IX, Pflanzenwelt Afrikas; W.F.K.=A.J.Jex-Blake, Some Wild Flowers of Kenya; Z.A.E.= Wissenschaftliche Ergebnisse der Deutschen Zentral-Afrika-Expedition 1907–1908, 2 (Botanik).

FAMILIES OF VASCULAR PLANTS REPRESENTED IN THE FLORA OF TROPICAL EAST AFRICA The family system used in the Flora has diverged in some respects from that now in use at Kew and the herbaria in East Africa. The accepted family name of a synonym or alternative is indicated by the word “see”. Included family names are referred to the one used in the Flora by “in” if in accordance with the current system, and “as” if not. Where two families are included in one fascicle the subsidiary family is referred to the main family by “with”. Foreword and preface (£3.00)

Glossary (£23.50)

Index of Collecting Localities (£18.50)

PTERIDOPHYTA Actiniopteridaceae (£2.30)

Grammitidaceae

Polypodiaceae (£10.00)

Adiantaceae (£15.00)

Hymenophyllaceae

Psilotaceae (£1.70)

Aspleniaceae

Isoetaceae

Pteridaceae (£8.50)

Azollaceae (£1.70)

Lomariopsidaceae (£10.00)

Salviniaceae (£1.40)

Blechnaceae

Lycopodiaceae

Schizaeaceae (£3.40)

Cyatheaceae

Marattiaceae (£1.70)

Selaginellaceae

Davalliaceae (£1.70)

Marsileaceae (£6.00)

Thelypteridaceae

Dennstaedtiaceae (£6.50)

Oleandraceae (£6.00)

Vittariaceae (£2.95)

Dryopteridaceae

Ophioglossaceae (£7.00)

Woodsiaceae (£8.00)

Equisetaceae (£1.70)

Osmundaceae (£1.70)

Gleicheniaceae (£2.30)

Parkeriaceae (£1.70)

GYMNOSPERMAE (£3.00) Cupressaceae

Cycadaceae

Podocarpaceae

ANGIOSPERMAE Acanthaceae

Aponogetonaceae (£3.90)

Agavaceae

Aquifoliaceae (£1.50)

Aizoaceae (£4.50)

Araceae (£10.20)

Alangiaceae (£1.50)

Araliaceae (£3.00)

Alismataceae (£3.00)

Arecaceae—see Palmae

Alliaceae (£4.00)

Aristolochiaceae (£3.00)

Aloaceae (£14.50)

Asclepiadaceae—see Apocynaceae

Amaranthaceae (£22.00)

Asparagaceae

Amaryllidaceae (£5.10)

Asphodelaceae (£6.00)

Anacardiaceae (£8.50)

Asteraceae—see Compositae

Ancistrocladaceae (£1.85)

Avicenniaceae—as Verbenaceae

Anisophyllaceae—as Rhizophoraceae Annonaceae (£10.50)

Balanitaceae (£6.00)

Anthericaceae (£11.50)

Balanophoraceae (£1.95)

Apiaceae—see Umbelliferae

Balsaminaceae (£11.80)

Apocynaceae

Basellaceae (£1.50)

Part 1 (£20.50)

Begoniaceae

Part 2

Berberidaceae (£1.50)

© 2003 Royal Botanic Gardens, Kew

FLORA OF TROPICAL EAST AFRICA WOODSIACEAE BERNARD VERDCOURT Terrestrial with erect or less often creeping rhizome, sometimes forming a short caudex; stipe not articulate (except in Woodsia), often densely scaly particularly near base, with 2 vascular strands united and U-shaped above. Fronds±spaced or tufted; lamina 1–3 pinnate, the costal groove with raised, often almost winged edges; veins free or anastomosing. Sori round, elliptic to linear or J-shaped or U-shaped with inconspicuous to obvious, entire or fimbriate indusium or indusium sometimes lacking. Paraphyses present or absent. Spores monolete with perispore. A cosmopolitan family (previously called Athyriaceae, but Woodsiaceae has priority by 7 years) included within an extended Dryopteridaceae by Kramer in Kubitzki, Fam. Gen. Vasc. Plants. As treated here, it contains about 15 genera and 500 species, with the Onocleaceae kept separate and Hypodematium Kunze treated under Dryopteridaceae. 1. Sori±round or ovate

2

Sori at least in part elliptic to elongate or J-shaped to U-shaped but some reduced sori towards apices of pinnule-lobes can be ±round

3

2. Fronds large, 0.6–2 m long with pinnules of middle pinnae 4.5–7 cm long; indusium rounded-reniform, sometimes almost absent

1. Dryoathyrium

Fronds smaller, up to 40 cm long with pinnules of middle pinnae 0.5–2 cm long; indusium bladder-shaped

2. Cystopteris (p. 4)

3. At least some sori J-or U-shaped and furthest end crossing the vein to which 3. Athyrium (p. sorus is attached 8) Sori all elliptic to linear, never crossing the vein 4. Trichomes of margins of rhizome-scales distinctly bifid; veins free or anastomosing Trichomes of margins of rhizomes-scales spiniform, not distinctly bifid*; veins free

4 4. Callipteris (p. 13) 5. Diplazium (p. 16)

1. DRYOATHYRIUM Ching in Bull. Fan. Mem. Inst. Biol., Bot. 11:79 (1941)

Rhizome erect, forming a short caudex, with non-peltate rhizome-scales. Fronds tufted; stipes not articulated. Lamina large, bipinnate with pinnules deeply pinnatifid into again pinnatifid lobes so as to appear tripinnate, glabrous except for minute short hairs along the narrowly and evenly winged costules, herbaceous; veins free. Sori superficial, round, with minute rounded-reniform indusia not apparent in mature sori, sometimes almost absent. A tropical genus of about 10 species distributed from tropical Africa and Madagascar to China and Japan; only one species occurs in continental Africa. * Occasionally in Diplazium trichomes can appear minutely bifurcate at extreme apex due to a fine hair below their tips.

D. boryanum (Willd.) Ching in Bull. Fan. Mem. Inst. Biol., Bot. 11:81 (1941); Schelpe, F.Z., Pterid.: 207, t. 59 (1970); Faden in U.K.W.F.: 57, fig. (on 59) (1974); De Vol & Kuo in Fl. Taiwan ed.1, 1:470 (1975); Schelpe & Diniz, Fl. Moçamb.: 223 (1979); Jacobsen, Ferns S. Afr.: 411, t. 308 (1983); Pic. Serm. in B.J.B.B. 55:155 (1985); Schelpe & Anthony, F.S.A., Pterid.: 229, fig. 77/1 (1986). Type: Réunion, Bory de St Vincent (BW 19831, holo., microfiche!) Rhizome erect, ascending or horizontal with upturned apex, forming a short thick fleshy caudex with pale brown concolorous subentire ovate to lanceolate rhizome-scales 5–12(–20) mm long, 5–7 mm wide, acuminate. Fronds tufted, arching, herbaceous, 0.6–2 m tall; stipe matt, greenish brown, up to 1 m long, glabrous except for pale brown scales about the base similar to those on the rhizome. Lamina narrowly ovate in outline, up to 1×0.64 m, acute; basal pinnae hardly reduced, bipinnate with 15–22 pairs of pinnules so deeply pinnatifid as to appear tripinnate; pinnae oblong-lanceolate-acuminate in outline, 13–44 cm long, 7–22 cm wide, attenuate, shortly petiolate, developed equally acroscopically and basiscopically, the basal pinnules somewhat reduced; pinnules oblong-acuminate, 3–12.5 cm long, 0.8–3 cm wide, base truncate, deeply pinnatifid into narrowly oblong obtuse deeply crenate-serrate lobes 4–16 mm long, 2–5 mm wide, with an angular sinus between them, glabrous on both surfaces except for minute short blunt brownish hairs along the costules and veins; veins free; rachis pale brown, glabrous; secondary rachises pale brown, with scattered minute blunt hairs and with a narrow laminar green wing at least about the insertion of the pinnules. Sori circular, 4–13(–16) per pinnule-lobe, ±0.6 mm in diameter; indusium membranous, minute or up to 0.8 mm wide, subentire. Fig. 1 (page 3). UGANDA. Kigezi District: Ishasha Gorge, 6 km SW of Kirima, 21 Sept. 1969, Faden et al. 69/1202!; Toro District: Ruwenzori Mts, Kazinga to Bwamba Pass, 23 Jan 1932, Hazel 124!; Mbale District: Mt Elgon, 24 May 1923, Snowden 777!, 778!

Flora of tropical East Africa

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KENYA. Naivasha District: Kimakia Forest Station towards South Kinangop, 13 July 1969, Faden 69/908!; Mt Kenya, West Kenya Forest Station, 5 Jan. 1922, R.E. & T.C.E. Fries 742!; Masai District: Narok District, about 4.8 km from Olokurto, edge of Olenguerone Settlement, 16 May 1961, Glover et al. 1167! TANZANIA. Arusha District: 8 km N of Arusha, S slope of Mt Meru, above Eastern sawmills, 29 Feb. 1953, Drummond & Hemsley 1298!; Mpanda District: Mahali Mts, Sisaga, 29 Aug. 1958, Newbould & Jefford 1915! & 1916!; Morogoro District: Uluguru Mts, Mwera Valley, 26 Sep. 1970, Faden et al. 70/586! DISTR. U 2, 3; K 3–6; T 2, 4, 6, 7; Ivory Coast (fide Tardieu), S Nigeria, Cameroon, Bioko, E Congo (Kinshasa), Rwanda, Mozambique, Zimbabwe, South Africa, Madagascar, Reunion and Comoro Is; also widespread in tropical and temperate Asia to NE Himalayas and Malesia (to Luzon) HAB. Evergreen forest often by streams, also in bamboo zone and in mist forest; 1350–2700 m SYN. Aspidium boryanum Willd., Sp. Pl. ed. 4, 5:285 (1810) Lastrea boryana (Willd.) Moore, Ind. Fil.: 86 (1858) Nephrodium catopteron (Kunze) Hook. var. glabrum Hook., Sp. Fil. 4:137 (1862) quoad syntype: Bioko [Fernando Po], Mann s.n. (K!, syn.) N. boryanum (Willd.) Bak., Syn. Fil.: 284 (1867) Aspidium glabratum Kuhn, Fil. Afr.: 133 (1868). Type: Bioko [Fernando Po], Mann s.n. (K, ? holo.)* Nephrodium glabratum (Kuhn) Bak., Syn. Fil.: 300 (1874) Dryopteris glabrata (Kuhn) Kuntze, Rev. Gen. Pl. 2:812 (1891) Aspidium kiboschense Hieron. in P.O.A. C: 85 (1895). Type: Tanzania, Kilimanjaro, Kibosho [Kiboscho], Volkens 1554 (B, holo., BM!, fragment) Dryopteris kiboschensis (Hieron.) C. Chr., Ind. Fil.: 273 (1906) D. boryana (Willd.) C. Chr., Ind. Fil.: 255 (1906) Nephrodium catopteron sensu Peter, F.D.-O.A.: 57 (1929), non (Kunze) Hook. * Tardieu-Blot states that the Mann type is at the BM. Kuhn would not have seen the Kew Mann specimen—but he cites Hooker.

Dryoathyrium

3

FIG. 1. DRYOATHYRIUM BORYANUM—1, pinna,×⅔; 2, fertile pinnule segment,×½. Both from Schelpe 5663. From F.Z. Drawn by Monika Shaffer-Fehre.

Flora of tropical East Africa

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Athyrium boryanum (Willd.) Tagawa in Act. Phytotax. Geobot. 4:144 (1935); Shieh et al. in Fl. Taiwan ed. 2, 1:418 (1994) Ctenitis boryana (Willd.) Copel., Gen. Fil.: 123 (1947) Thelypteris glabrata (Kuhn) Tardieu in Not. Syst. 14:344 (1952) & in Mém. I.F.A.N. 28:120 (1953) pro parte T. glabrata (Kuhn) Tardieu var. hirsuta Tardieu in Not. Syst. 14:344 (1952). Type: Ivory Coast, no specimens cited Athyrium glabratum (Kuhn) Alston in Bol. Soc. Brot. sér. 2A, 30:11 (1956) & Ferns W.T.A.: 64 (1959); Tardieu, Fl. Cameroun 3:232, t. 41/1–2 (1964) Cornopteris boryana (Willd.) Tardieu in Mém. Sc. Madag. 7:30, t. 2, fig. 1–4 (1956) nom. invalid. & in Amer. Fern Journ. 48:32 (1958) & in Fl. Madag. 5(1): 257, fig. 33/1–5 (1958) Parathyrium boryanum (Willd.) Holttum in K.B. 13:449 (1959) Lunathyrium boryanum (Willd.) H.Ohba, Sci. Rep. Yokosaka City Mus. 11:53 (1965) Deparia glabrata (Kuhn) Kato in Bot. Mag. Tokyo 90:36 (1977) D. boryana (Willd.) Kato in J. Fac. Sci. Univ. Tokyo III 13:386 (1984) NOTE. There is considerable argument over the correct generic placing of this fern. Kramer in Kubitzki (Fam. Gen. Vasc. Pl.: 136 (1990)), following Kato’s revision includes Dryoathyrium in Deparia Hook. f. & Grev. but divides the latter into 4 quite distinct sections one of which is Dryoathyrium (Ching) Kato. For the purpose of this flora there seems little point in changing a long used name.

2. CYSTOPTERIS Bernh. in Schrad., Neues J. Bot 1(2): 5, 26 (1805) (‘1806’); Blasdell in Mem. Torrey Bot. Club. 21(4): 1–102 (1963); Kato & Kramer in Kubitzki, Fam. Gen. Vasc. Pl. 1:138 (1990); Fraser-Jenkins, New species syndrome in Indian Pteridology & the ferns of Nepal: 95–100 (1997)

Small to medium-sized ferns, terrestrial or often growing on rocks. Rhizome short and suberect to long-creeping with entire membranous scales. Fronds tufted or spaced; stipe grooved. Lamina rather thin to thinly coriaceous (membranous in one species), usually well-dissected, 1–3-pinnate and pinnatifid; rachis glabrous or scaly, occasionally with bulbils (not in Africa), grooved; veins free. Sori round, at first covered by ovate to oblong, lanceolate or cup-shaped inflated bladder-like indusia attached below the sori; spores echinate or rugose. About a dozen species*. C. fragilis (L.) Bernh. in Schrad., Neues J. Bot. 1(2): 27 (1805); Pirotta in Il Ruwenzori, Parte Scient. 1:428 (1909) (sensu lato); Sim, Ferns S. Afr. ed. 2:88, t. 8 (1915) (sensu lato); A.V.P.: 27 (1957) (sensu lato); Alston, F.W.T.A.; Pterid.: 64 (1959) (sensu lato); Blasdell in Mem. Torrey Bot. Club 21(4): 38 (1963) (sensu stricto); W.Jacobsen, Ferns S. Afr.: 402, t. 302 (1983) (sensu lato); Schelpe & N.C. Anthony, F.S.A., Pterid.: 229, fig. 76/2 (1986) (sensu lato); Derrick, Crabbe & Jermy in Sommerfeltia 6:62 (1987) (sensu stricto); J.Burrows, S.Afr. Ferns: 279, t. 46.5, fig. 65/284, 284a (1990) (sensu lato); Crabbe & Jermy in Fl. Eur. ed. 2, 1:24 (1993) (sensu stricto); Faden in U.K.W.F. ed. 2:33 (1994). Type: Plukenet, Almagestum botanicum: 150, t. 180, fig. 5 (1696) (lecto., chosen by Weatherby in Rhodora 37:376 (1935))** * Schelpe & N.C.Anthony (F.S.A.Pterid.: 229 (1986)) gives about 18. ** Schelpe gives the type as Europe, Herb. Sloane (H.S. 96, Fol. 40) (BM, holo.) but Linnaeus would not have seen the specimen. It could not be a holotype since Linnaeus cites several references. There is also a specimen in the Linnean Herbarium 1251.51 but there appears to be doubt it was available to Linnaeus in 1753.

Rhizome shortly creeping or ±erect with reddish brown glabrous lanceolate scales 3 mm long, 0.6 mm wide, sometimes with marginal glands. Fronds ±tufted or closely spaced; stipe usually shorter than the lamina, straw-coloured to dark brown, 3–20 cm long with lanceolate scales at the base. Lamina lanceolate to ovate-lanceolate, 5–30 cm long, 1.5–14 cm wide, 1–3-pinnate and 3(–4)-pinnatifid; rachis and axes with scattered multicellular glandular hairs; pinnae in 4–20 pairs, ovate to ovate-lanceolate, 0.6–1.8 cm long, 0.5–4 cm wide, basal secondary pinnae shortly petiolate, remainder adnate, ovate to lanceolate, obtuse to acute with apices of the lobes with rounded or acute teeth; veins directed into teeth or sinuses. Sori round, discreet to overlapping at maturity; indusia bladder-like, ovate to oblong or cup-shaped, dentate to lacerate.

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SYN. Polypodium fragile L., Sp. Pl. ed. 1:1091, (1753) (as P.F. fragile)* NOTE. Blasdell records C. fragilis var. fragilis from eastern Africa (op. cit, p. 39) but cites nothing in his list of specimens. Fronds 10–20 cm long, simply pinnate with 4–8 pairs of pinnae; veins mostly running into subsp. lobes or teeth but sometimes into sinuses; spines of spores** ±further apart than length of A spines Fronds 7.5–37 cm long, essentially 2-pinnate with 9–20 pairs of pinnae; veins mostly running into the sinuses; spines of spores densely packed

subsp. B

subsp. A Fronds 10–20 cm tall with stipe 3–10 cm long, very slender. Lamina lanceolate in outline, 5–9.5 cm long, 1.5–3 cm wide, simply pinnate with 4–8 pairs of pinnae and apical lobed segment; pinnae and apical segment ovate or triangular in outline, 0.6–1.5 cm long, 0.4–1.1 cm wide, shallowly to deeply lobed; veins mostly running into lobes or teeth but sometimes into sinuses. Spines on spores more sparse, ±further apart than the length of the spines. Fig. 2/1, 2 (page 6). UGANDA. Mbale District: Mt Elgon, in crater, N slope of Jackson’s Peak, 17 May 1948, Hedberg 979 (photocopy!) TANZANIA. Moshi District: Kilimanjaro, Kikafu R. gorge, 1 Mar. 1997, Hemp 1623! & stream near NW edge of Shira Plateau, 8 Nov. 1968, Bigger 2281! & on same plateau, Simba Camp area, 5 Sep. 1993, Grimshaw 93–616! DISTR. U 3; T 2 HAB. Crevices in rocks, on rocks overhanging streams, hanging from roofs of caves and rock overhangs; 3400–4750 m NOTE. I at first thought this might be Cystopteris nivalis (Pirotta) Pic. Serm. judging by some of Pichi Sermolli’s comments (see note after subsp. B). Discovery of topotypic material in BM (Osmaston 1740, Uganda, Bujuku Valley, Bigo, July 1972) shows this is not so. The morphology of the frond and densely echinate spores are the same as in subsp. B. Although in East Africa this alpine form seems distinctive; similar forms which appear almost identical can be found in S Africa and in the UK. * Linnaeus actually gave the name as Polypodium F. fragile in Sp. Pl. ed. 1:1091 (1753) and in Syst. Veg. ed. 10:1327 (1759) and several authors have suggested that the correct name for the species should be C. filix-fragilis (L.) Borbás, A.Balaton Flórája 2, 314 (1900) (without hyphen); Chiov. in Ann. Bot.: 210 (1903); Becherer in Ber. Schweiz. Bot. Ges. 43 (1): 39 (1934). This was discussed by Merrill in Am. Fern Journ. 25:127 (1935) and by Weatherby in Am. Fern Journ. 27:27 (1937) in which I think he conclusively shows the F. was an error which Linnaeus corrected in manuscript in his own copy of Sp. Pl. ed. 1 and in Fl. Suec. ed. 2:374 (1755) and Sp. Pl. ed. 2:1553 (1763). This needs reiterating since Jones in Fl. Austr. 48:419 (1998) has given the type of the genus as C. filix-fragilis (L.) Bernh. Example 19 to Article 23.3 of the Code of Botanical Nomenclature states the name is to be treated as P. fragile L. ** My thanks are due to P.J.Edwards for preparing scanning electron micrographs from my stubs.

Cystopteris

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FIG. 2. CYSTOPTERIS FRAGILIS— subsp. A: 1, habit,×⅔; 2, fertile pinna, enlarged; subsp. B: 3, habit,×⅔; 4, fertile pinna, enlarged; 1–2 from Bigger 22811; 3–4 from Battiscombe 978. Drawn by Pat Halliday.

Flora of tropical East Africa

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subsp. B Fronds 7.5–37 cm tall with stipe 3.5–20 cm long. Lamina lanceolate or ±triangular in outline, 4–25 cm long, 1.5–10 cm wide, essentially 2-pinnate with 9–20 pairs of pinnae, but pinnules narrowly decurrent, deeply pinnatifid; sometimes truly 2-pinnate; veins mostly running into the sinuses. Spores with densely packed spines. Fig. 2/3, 4 (page 6). UGANDA. Toro District: Ruwenzori, Namwamba Valley, 7 Jan. 1935, G. Taylor 2984!; Kigezi District: Virunga Mts, Sabinio, 24 Nov. 1934, G. Taylor 2036!; Mbale District: Mt Elgon, Sasa trail, Dirigana R., 20 June 1997, Wesche 1462! KENYA. Nakuru District: Mau Forest, near Elburgon, H.M.Gardner in F.D. 978!; North Nyeri District: West Mt Kenya Forest Station, 5 Jan. 1922, R.E. & Th. C.E. Fries 762!; Kericho District: SW Mau Forest, along Kiptiget (Chepkoisi) R, ±16 km SSE of Kericho, Salt Lick Falls, 12 June 1972, Faden et al. 72/331! TANZANIA. Masai District: Ngorogoro, near Old Boma, 2 Mar. 1961, Newbould 5683!; Arusha District: Mt Meru crater, 8 Mar. 1971, Richards 26713!; Moshi District: Kilimanjaro, Marangu, R Himo, Nov. 1964, Beesley 58!; Mbeya District: Kikando, 22 Sep. 1956, Richards 6713! DISTR. U 2, 3; K 1, 3–5; T 2, 7; Cameroon Mt, Congo (Kinshasa) (Kivu), Sudan, Ethiopia, Natal; Mediterranean part of Europe, Cape Verde Is., Canary Is. HAB. Mostly on damp rocks, rock faces and cliffs, often near rivers, chiefly in evergreen forests, e.g. Hagenia; Podocarpus; Albizia–Macaranga–Eagara–Syzygium– Croton or Xymalos–Prunus–Maesa –Afrocrania; also on moist loamy banks and in alpine zone; 1400–3800 m SYN. Aspidium viridulum Desv., Mag. Nat. Berl. 5:321 (1811). Type: Tenerife, ?collector (?P) Cystopteris viridula (Desv.) Desv. in Mém. Soc. Linn., Paris 6:264 (1827) Woodsia nivalis Pirotta in Ann. Bot. Roma 7:173 (1908) & in Il Ruwenzori, Parte Scient. 1:478 (1909). Type: Uganda, Toro District, Ruwenzori, Mobuku Valley, Bujongolo, 3800 m (sphalm 3000 m), Cavalli-Molinelli & Roccati s.n. (TO, holo.) [Cystopteris fragilis L. subsp. diaphana sensu Litard in Bull. Géogr. Bot. 21 (No. 255): 20 (9 Jan. 1911)) & in Bull. Soc. Bot. Deux-Sevres: 88 (1911–12); Zeiller in Bull. Soc. Bot. Fr. 59:800 (1912) (abstr.); Fraser-Jenkins, New species syndrome in Indian Pteridology & the ferns of Nepal: 97 (1997), non (Bory) Litard sensu stricto] [C. diaphana sensu Blasdell in Mem. Torrey Bot. Club 21 (4): 47 (1963); Derrick, Crabbe & Jermy in Sommerfeltia 6:62 (1987); Crabbe & Jermy in Fl. Europea ed. 2:24 (1993), non (Bory) Blasdell sensu stricto] C. nivalis (Pirotta) Pic. Serm. in Webbia 23:173 (1968) NOTE. I at first thought this could be identified as subsp. diaphana (Bory) Litard as had been suggested by Blasdell and others; the venation suggests this. Jermy, Flora Europaea ed. 2 states that the spores are densely echinate, their spines almost touching at the base exactly as in the above. There is, however, a photograph of the type of Polypodium diaphanum Bory (Voy. Iles. Mers Afr. 1:328 (1804). Type: Reunion, Plaine des Chicots, Bory St. Vincent (P, holo.)) at the BM and attached to it a SEM photograph of spores showing spore echination which confuses the whole issue. Desvaux’s name cited above probably provides the correct epithet for this subspecies. Spores derived from specimens collected in Tenerife have dense echination.

Cystopteris

9

Crabbe & Jermy (op. cit.) retain this as a species but admit that C. fragilis is a widespread polymorphic polyploid complex and I agree with Fraser-Jenkins in regarding it as a subspecies. Vida in his cytological paper (Act. Bot. Acad. Sci. Hungaricae 20 (1– 2): 181–192 (1974)) also suggest this is the best solution. Fraser-Jenkins states that the difference between the spores of subsp. fragilis and subsp. diaphanum [auctt.] is a 100 % reliable character. However, one specimen of subsp. B: Gardner in F.D. 978 cited above is somewhat intermediate in spore characters but not in frond shape. Pichi Sermolli (who examined the type) gives a number of characters by which he claims C. nivalis differs from typical C. fragilis including stouter rhizome, slightly thicker rhizome scales mixed with hairs and all with red-castaneous pyriform glands; similar hairs occur on the rachis, costae and surface of the lamina; the indusia are wide, broadly cyathiform, attached around one-half of the circumference of the receptacle, coarsely and irregularly dentate-fimbriate, dilacerate when sorus is mature, not glandular; spores with bacula and spinulae much more spaced and shorter (about one third) 1.7–2.5 µ long; the bacula are more numerous than the spinulae and the latter are often hooked. Some hairs can be found on all Cystopteris laminae but the rhizome scales of the Kilimanjaro material cited above do not end in glands. Pichi Sermolli suggested that C. nivalis would occur on other mountains. Fraser-Jenkins (op. cit.) treats C. nivalis as a synonym of C. fragilis subsp. fragilis.

3. ATHYRIUM Roth, Tent. Fl. Germ. 3:31, 58 (1799); Kato & Kramer in Kubitzki, Fam. Gen. Vasc. Pl. 1:132 (1990)

Rhizome branched, creeping or erect, often ±fleshy, with dense non-peltate scales. Fronds tufted or closely spaced. Stipe sulcate, often pink. Lamina 1–3-pinnate to 4pinnatifid, rarely simple (not in Africa), mostly glabrous; veins free (save in one nonAfrican species). Sori superficial, mostly J-shaped or U-shaped (but round or oblong in many European species) and furthest end crossing the vein to which the sorus is attached; indusium present and fimbriate or rudimentary or even lacking (not in Africa). About 180 species, cosmopolitan but mostly in the north temperate zone. 1. Lamina essentially trifid, the apical part pinnate with 12 or more pairs of deeply pinnatifid pinnae

2

Lamina not as above, regularly 2–3-pinnate 2. Lamina appearing ±5-partite since lowest basiscopic pinna of lateral parts very well developed; apices of all parts distinctly narrowly attenuate Lamina strictly tripinnate, the lateral parts without well-developed basiscopic pinnae; apices of all parts not narrowly attenuate 3. Fronds 16–20 cm tall with pinnae up to 2.5 cm long; pinnules oblong-ovate, ±7 mm long with broadly rounded lobes; the stomata beneath appear as dense slightly paler minute spots (T7)

3 4. A. lewalleanum 5. A. rondoense 3. A. sp.

Fronds not as above, usually much larger 4. Rhizome creeping with fronds closely spaced; lamina 2-pinnate, never truly 3pinnate and even when pinnules very deeply pinnatifid, the lobes are decurrent and axis winged; lowest pair of pinnae often somewhat distant and much reduced (often not so noticeable in Southern African material) Rhizome erect with fronds tufted; lamina 2–3-pinnate; lowest pair of pinnae only slightly reduced

4 1. A. schimperi

2. A. scandicinum

1. A. schimperi Fée, Mém. Fougères 5, Gen. Fil.: 187 (1852); V.E. 2:23 (1908); Sim, Ferns S. Afr., ed. 2:133, t. 41 (1915); Tardieu, Mém. I.F.A.N. 28:161, t. 30, fig. 3, 4 (sphalm. 5, 6) (1953); Alston, Ferns W.T.A.: 64 (1959); Tardieu, Fl. Cameroun 3:229, t. 35, fig. 3, 4 (1964); Schelpe, F.Z.Pterid: 202, t. 57/b (1970) & C.F.A., Pterid.: 162 (1977); Kornaś, Distr. Ecol. Pterid. Zambia: 105 (1979); W.Jacobsen, Ferns S.Afr.: 404, fig. 303 (1983); Pic. Serm. in B.J.B.B. 55:155 (1983); Schelpe & N.C.Anthony, F.S.A. Pterid.: 223, fig. 75 (1986); J.Burrows, S. Afr. Ferns: 273, t. 46/1, fig. 65/278, 278a (1970); Schippers in Fern Gaz. 14 (6): 205 (1993); Faden in U.K.W.F. ed. 2:33 (1994). Type: Ethiopia, Debra Eski, “3000 m” Schimper 239 (B, syn., K!, photo., US, iso.) Rhizome creeping, 6 mm in diameter, with reddish brown lanceolate and acuminate scales 7 mm long, 1 mm wide but often some much narrower. Fronds closely spaced, up

Athyrium

11

to ± 1 m tall; stipe dark brown at base, 14–38 cm long, glabrous but with scales similar to rhizome scales at base. Lamina oblong-lanceolate in outline, 23–70 cm long, (6–)10–28 cm wide, acute to acuminate, virtually to quite bipinnate, the pinnules mostly narrowly decurrent; pinnae in 15–20 pairs, lanceolate in outline, 4–18 cm long, 1.5–5 cm wide, attenuate, the basal pair usually markedly smaller and

FIG. 3. ATHYRIUM SCHIMPERI—1, rhizome, stipe and lower part of lamina,×⅔; 2, pinna, ×⅔; 3, lower surface of pinnule showing sori,×4. All from Chase 3774. From F.Z. Drawn by Lura Mason Ripley.

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distant; rhachis straw-coloured; pinnules in 7–18 pairs, narrowly oblong, 5–22 mm long, 2–8 mm wide, sharply serrate-dentate or deeply lobed and lobes serrate. Sori oblong, curved or J-shaped, 1–1.5 mm long, 1–7 per pinnule-lobe and 14–70 per pinnule; indusia pale brown, membranous, erose. Fig. 3 (page 9). UGANDA. Karamoja District: Mt Kadam, Apr. 1959, Wilson 788! KENYA. Trans-Nzoia District: Mt Elgon, Suam Saw Mill track, edge of Kiptogot River Valley, 12 June 1971, Faden et al. 71/452! & Kiptogot waterfall, Aug. 1959, Tweedie 1882! & Oct. 1961, Tweedie 1882A! TANZANIA. Ufipa District: Nsanga Mt, Malonje Plateau, 13 Mar. 1959, Richards 12119!; Mbeya District: Mbeya Range, Nov. 1957, Watermeyer 12!; Rungwe District: Kyimbila, Mbaka, 13 Apr. 1912, Stolz 1220! DISTR. U 1; K 3; T 1 (see note) 4, 7; Ghana, Nigeria, Cameroon, Congo (Kinshasa), Rwanda, Burundi, Sudan, Ethiopia, Angola, Zambia, Malawi, Zimbabwe and eastern S Africa; also India (Himalayas) HAB. Rocky forest margins with Olinia and Juniperus, crevices of rock outcrops in shade; 2100–2400 m SYN. Asplenium schimperi (Fée) A.Braun in Schweinf., Beitr. Fl. Aethiop. 1:224 (1867) NOTE. Many specimens lack rhizomes. The record from U 1 needs confirming from material with a rhizome. Gillman 363 from T 1 (Bukoba District, Kaagya), also without a rhizome is probably this species. 2. A. scandicinum (Willd.) Presl, Tent. Pterid.: 98 (1836) & in Abh. Königl. Böhm. Ges. Wiss. ser. 4, 5:98 (1837); Tardieu, Fl. Madag. 5 (1): 267, fig. 36/9–11 (1958) (as ‘scandinicum’). Type: Réunion, Bory de St. Vincent s.n. (B-W 19832, holo., microfiche!) Rhizome erect, ±fleshy, 5–8 mm wide but the branches forming a caudex up to 2.5 cm wide; rhizome scales brown, lanceolate-attenuate, 7–10 mm long, 1–2 mm wide. Fronds tufted, 0.15–2 m tall; stipe usually reddish or pink when living, 7.5–55 cm tall, glabrous save for scales at base but young stipes of undeveloped fronds often scaly throughout. Lamina ovate-lanceolate in outline, 20–50 cm long, 10–34 cm wide, acute to acuminateattenuate at the apex, 2–3-pinnate with ±18 pairs of pinnae, the basal pair somewhat reduced; pinnae lanceolate in outline, acuminate to caudate at the apex, 1.5–20 cm long, 0.8–8 cm wide, with 10–20 pairs of pinnules; pinnules oblong-ovate to oblong-lanceolate, 1–5 cm long, 0.3–2 cm wide, sometimes cut to the base into secondary pinnules, the lamina then distinctly 3-pinnate but often only 2-pmnate-pinnatifid, the pinnules crenate to divided into narrow lobes; pinnule-lobes or secondary pinnules 3–10 mm long, 1–7 mm wide; soft spinules up to 1 mm long often present where wings of rhachis or rhachilla are interrupted by pinnule-bases and some short indumentum sometimes present. Sori oblong, curved or J-shaped, up to 2 mm long, 5–7 per pinnule-lobe or secondary pinnule; indusium shortly to deeply fimbriate or subentire (fide Schelpe). SYN. Aspidium scandicinum Willd., Sp. Pl. ed. 4, 5:285 (1810) NOTE. I agree with Pichi Sermolli that African material is certainly not identical with typical material from Reunion which has much more delicately cut fronds with the pinnule-lobes much smaller. I have therefore divided it into two subspecies, admittedly not too well defined. subsp. newtonii (Bak.) Verdc, comb. nov. Type:* São Tomé, Newton 2 & Quintas 9 (K!, syn., COI, isosyn.)

Athyrium

13

* It is clear from Baker’s description that he saw both sheets although he merely cites ‘Island of St. Thomas, Newton’. Exell (Cat. Vasc. Pl. S.Tomé: 73 (1944)) gives Newton 2 and Newton 88 as being at Kew but there is no Newton 88, it is Quintas 9 collected on ‘6.88’ (June 1888]. I am certain Baker had also misread it as Newton and am treating the two sheets as syntypes. Exell adds “also from Ruwenzori”.

Pinnule-lobes or secondary pinnules ovate or oblong, typically shortly and obtusely crenate. UGANDA. Toro District: Ruwenzori, Mijusi Valley, 31 Mar. 1948, Hedberg 615! & Mihunga, 14 Jan. 1939, Loveridge 365!; Ankole District: Kalinzu Forest, 4 km NW of Saw Mill, W of Rubuzigye, 19 Sep. 1969, Faden et al. 69/1140! KENYA. Aberdare Mts, Ramsden s.n.!; Fort Hall District: Kyama R., below Mbugiti School, 13 July 1969, Faden & Evans 69/890!; Kericho District: 8 km NW of Kericho, Kimugung R., 10 June 1972, Faden et al. 72/292! TANZANIA. Moshi District: Kilimanjaro, ravine E of Maundi Crater, 12 Oct. 1993, Grimshaw 93783!; Lushoto District: W Usambaras, Mangula, 18 Sep. 1981, Mtui & Sigara 64!; Iringa District: Kilombero, Mwanihana Forest Reserve, above Sanje village, 10 Oct. 1984, D.W. Thomas 3864! DISTR. U 2, 4; K 3–5; T 2–4, 6, 7; São Tomé, Sudan, Malawi, Mozambique, Zimbabwe, Swaziland, South Africa (the Madagascan material probably belongs here) HAB. Forest e.g. Parinari excelsa, Polysdas–Macaranga–Fagara–Albizia– Neoboutonia, extending into ericaceous belt, often by streams on marshy ground; 1150– 3500 m SYN. Athyrium newtonii Bak. in Ann. Bot. 5:307 (1891); Exell, Cat. Vasc. Pl. S. Tomé: 73 (1944); Schippers in Fern Gaz. 14:205 (1993) A. laxum Pappe & Raws., Syn. Fil. Afr. Austr.: 16 (1859), non Schumach. Type: South Africa, Natal, Gueinzius s.n. (ubi?) [Athyrium scandicinum sensu Hieron. in P.O.A. C: 84 (1895); V.E.: 23 (1908); Sims, Ferns S. Afr. ed. 2:133, t. 42 (1915); Schelpe, F.Z.Pterid.: 204 (1970); Schelpe & Diniz, Fl. Moçamb. Pterid.: 220 (1979); W.Jacobsen, Ferns S. Afr.: 404 (1983); Pic. Serm. in B.J.B.B.: 55:154 (1985); Schelpe & N.C.Anthony, F.S.A.Pterid.: 223 (1986); Schippers in Fern Gaz. 14:205 (1993); Faden in U.K.W.F. ed. 2:33 (1994) pro parte, non (Willd.) Presl sensu stricto] Athyrium scandicinum (Willd.) Presl. var. scandicinum sensu Schelpe in F.Z.Pterid.: 204 (1970); Schelpe & Diniz, Fl. Moçamb.: 22, (1979); Jacobsen, Ferns S. Afr.: 405, fig 304a, 304b (1983); Burrows, S. Afr. Ferns: 273, t. 45/6, fig. 65/279, 279a, 279b (1990) NOTE. R.B. & A.J. Faden 74/412 (Morogoro District: Uluguru Mts, Bondwa summit, 2120 m, in Podocarpus–Allanblackia–Polysdas–Balthasaria–Syzygium forest) has erect rhizomes but the frond architecture is similar to that of A. schimperi. Attempts to divide the mainland subspecies further have been unsuccessful. It was at first thought that the southern African population might form a taxon equivalent to A. laxum but the variation overlaps too much with that of material further north to make it practical to do so. Clair Thompson mentions (on a label) it can be epiphytic but there is no confirmation of this. Schelpe described A. scandicinum var. rhodesianum (Bol. Soc. Brot, Sér. 2, 41:211 (1967); F.Z.Pterid.: 204, t. 57, fig. a (1970). Type: Zimbabwe, Pungwe Gorge, Schelpe 5722 (BOL, holo., BM!, iso.)) but it seems doubtfully different and he makes no mention

Flora of tropical East Africa

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of it in the F.S.A.Pterid. account. Burrows (op. cit.) and Jacobsen (op. cit.) keep it distinct. Without really adequate rhizome data A. schimperi and A. scandicinum can be difficult to distinguish particularly in southern Africa and some field notes mention creeping rhizomes yet say fronds are tufted. 3. A. sp. Rhizome erect with spreading roots; scales chestnut, lanceolate, 3 mm long, 1 mm wide, attenuate. Fronds tufted, 16–20 cm tall; stipe 5–6 cm long, scaly at base. Lamina oblong-lanceolate in outline, 9–12 cm long, 4–5 cm wide with 10–15 pairs of pinnae; pinnae oblong in outline, 1–2.5 cm long, 0.3–1.3 cm wide, subacute and not attenuate at apex, with 4–5 pairs of pinnules and short apical lobed part; pinnules oblong-ovate, up to about 7 mm long, 5 mm wide, 3–5-lobed and ±crenate, with sparse flattened trichomes beneath. Sori one per pinnule, about 1 mm long; indusia deeply fimbriate. TANZANIA. Iringa District: Luhomero Massif, South Msisimwana ridge, 19 Aug. 1985, Rodgers & Hall 4558! DISTR. T 7 HAB. Rock outcrop; 2300 m NOTE. It was suggested that this was a young plant of A. scandicinum but it is fertile. No other material has been seen which resembles it and until more is collected in the same area its status is uncertain. 4. A. lewalleanum Pic. Serm. in Webbia 27:440, f. 18 (1973) (‘1972’). Type: Burundi, Bubana, road towards Mabaye, R.Nyamagana, Lewalle 3433 (Herb. Pic. Ser. 24934, holo., Herb. Lewalle, iso.) Rhizome long-creeping, slender; scales pale brown or dark-marked, triangular, 0.2–1.2 mm long, 0.3–0.4 mm wide, acuminate, cordate at the base, mixed with multicellular hairs. Fronds few, paired or solitary, 15–40 cm tall; stipe 12–24.5 cm long, minutely pubescent or ± glabrous or with few scales at base. Lamina ± triangular-hastate in outline, 11.5–22 cm long, 12–26 cm wide, with scattered flattened white multicellular hairs, essentially trifid, the apical part pinnate with 12 or more pairs of pinnae; pinnae lanceolate, up to 5 cm long, 1.8 cm wide, attenuate at the apex, deeply pinnatifid with ±10 pairs of pinnatifid segments about 2–7 mm wide; lateral parts similar to apical but with lowest basiscopic pinna well developed, up to 10 cm long; rachis shortly pubescent. Sori round, ±1 mm in diameter, 1–10 per segment; indusium sub-reniform, fixed on one side only, persistent or ± disintegrating and scarcely visible. TANZANIA. Kigoma District: Kasye Forest, 18 Mar. 1994, Bidgood et al. 2788! & same locality, 19 Mar. 1994, Bidgood et al. 2806! DISTR. T 4; Burundi, ? Zambia (see note) HAB. Evergreen forest along streams and rivers and on valley slopes, Baikiaea– Julbernardia– Monopetalanthus–Parkia–Tessmannia–Maesopsis–Pycnanthus; 900 m NOTE. Kornaś (in K.B. 33:101 (1978)) has described a very similar plant A. annae (Type: Zambia, Kalungwishi R. gorge below Lumangwe Falls, Kornaś & Medwecka– Kornaś 3746 (KRA, holo., K!, Herb. Pic. Serm., iso.) differing in a number of small points. I suspect it is conspecific. Frazer-Jenkins has annotated the Kew isotype suggesting the centro-basally attached indusium is wrong for Athyrium and that the species might be a Ctenitis or Hypodematium but it does not seem to fit well in either.

Athyrium

15

5. A. rondoense Verdc., sp. nov. affinis A. lewalleani Pic. Serm. lamina stricte tripartita, pinnis basiscopicis partium lateralium haud anguste caudatis, pinnis, pinnulis lobisque pinnularum majoribus differt. Typus: Tanzania, Rondo Plateau, Bidgood et al. 1550 (K!, holo., BR, C, DSM, EA, MO, NHT, P, WAG, iso.) Rhizome creeping with blackish or black-veined clathrate scales ±1.5 mm long; fronds single, 15–40 cm tall; stipe slender, 10–20 cm long with few scales and dense very short hairs. Lamina up to 20 cm long, 15 cm wide, tripartite, the parts ± confocal, each simply pinnate but pinnae very deeply divided so as to appear bipinnate; main parts broadly lanceolate in outline up to 18 cm long, 6 cm wide, acuminate but not long-caudate at the apex; each part with 6–9 pairs of pinnae up to ±5 cm long, 2 cm wide, the apical ones reduced, toothed only at the apex, running together and narrowly decurrent into each other resembling a series of fish-tails; lower pinnae deeply divided into ±6 elliptic lobes, the largest ±15 mm long, 8 mm wide, ±entire to bluntly or subacutely lobed or toothed; main lobes of largest pinnae decurrent, separated by narrowly winged midrib 1–2 mm wide. Sori 1–4 per pinnalobe, ±1 mm wide; indusium fimbriate and hairy. TANZANIA. Lindi District: Rondo Plateau, Rondo Forest Reserve, 14 Feb. 1991, Bidgood et al. 1550! DISTR. T 8; only known from the type HAB. Evergreen forest of Milicia–Albizia–Dialium in small gully and amphitheatre around well on escarpment; 650 m NOTE. This had been confused with Arachniodes foliosa (C. Chr.) Schelpe (Dryopteridaceae) and is superficially similar but the pinnule-lobes are not aristatedentate.

4. CALLIPTERIS Bory, Voy. Quatre princip. Iles Afrique 1:282 (1804); Pacheco & Moran in Brittonia 51:343–388 (1999)

Rhizome erect, stout, usually mucilaginous, scaly; scales brown eventually with dark brown to black edges with marginal trichomes bifid. Fronds large. Stipe often tuberculate to spinulose, scaly at the base. Lamina 1–2-pinnate (entire in one S American species), thin to fleshy, the distal portion often only pinnatifid, often proliferous; lateral veins parallel with lateral veinlets anastomosing regularly with those of adjacent groups or free (see note below). Sori elongate, along the veinlets, numerous, often in U-shaped pairs; indusium elongate, usually membranous. Genus often restricted to 3 species in Africa to Malesia, Australia and Polynesia but as redefined by Pacheco & Moran (op. cit.) with many species also in the Neotropics and others in the Old World which need transferring to the genus, which may then contain over 30 species. I am not completely convinced of this so far as the Old World species are concerned. Pacheco & Moran (op. cit.) have revised some of the New World species. They define Callipteris by its rhizome scales having dark chestnut to black margins and bifid marginal teeth; not all species have anastomosing veins but the type does and the group of 15 species they have revised. They further indicate that several old world species, apart from the type, should be transferred to Callipteris. I have found difficulties with this, however, apart from the fact that most specimens including nearly all types do not have rhizomes preserved; there is variation in some species where trichomes can be minutely bifid or uniformly undivided and scales are not black-margined. They mention that Bory de Saint-Vincent’s original conception of the genus contained only species with anastomosing veins but this is not so. Fronds simply pinnate; veins anastomosing

1. C. prolifera

Fronds bipinnate; veins not anastomosing

2. C. ulugurica

1. C. prolifera (Lam.) Bory, Voy. Quatre princip. Iles Afrique 1:283 (1804); D.L. Jones in Fl. Australia 48:422, figs. 88, 139/d–e. Type: La Reunion, Commerson s.n. (P–LAM!, syn., microfiche 742/3–4!, BM!, photo., Morton Neg. 2765) Rhizome ascending to erect, fleshy; scales pale brown, lanceolate, 7–10 mm long, attenuate, 1–2 mm wide at base, pale to eventually black at margins and with stiff bifid trichomes. Fronds tufted, 1.5–2.1 m tall; top of stipes and rachis sometimes with short stiff spinules or in some forms strongly muricate but usually ±smooth in East Africa; bulbils in upper 3 pinna axils and sometimes below in 7th, producing young plants in situ; stipe 15–90 cm tall. Lamina oblong-lanceolate, 15–120 cm long, 26–44 cm wide, with terminal segments 10–21 cm long, 7–10 cm wide with teeth or lobes up to 2.5 cm long, 1.5 cm wide; pinnae alternate, oblong, 6.5–24 cm long, 3–6.5 cm wide, tapering at apex, truncate, emarginate or subcordate (in young plants) at the base, the lowest the smallest,

Callipteris

17

coarsely toothed or crenate, the teeth or crenae 4×6 mm; venation anastomosing. Sori brown, numerous, narrow, appearing like contiguous branched trees from each side of the midribs of the pinnae in each area bounded by a lobe with 7–8 pairs of sori per tree; the whole gives a series of parallel zig-zag lines along the length of the pinna; indusia linear. Fig. 4 (page 14). UGANDA. Bunyoro District: Budongo Forest Reserve, nature reserve close to Sonso R., 31 Dec. 1995, Poulsen et al. 910!; Toro District: Bwamba, Kabanga, 21 Nov. 1935, A.S.Thomas 1499! & Bwamba Forest Reserve, 28 July 1960, Paulo 614! TANZANIA. Lushoto District: Amani, July 1913, Grote in A.H. 5792! & Kwamgumi Forest Reserve, NW of Mhinduro peak, 11 Nov. 1986, Farkas & Pócs 86606! & Mwesini [Mwezi] to Wugu [Iwugu] near Bwiti, June 1917, Peter 20487! DISTR. U 2, 4; T 3; Guinea to Nigeria, Cameroon, São Tomé, Principe, Bioko, Gabon, Congo (Kinshasa), Sudan, Angola, Mascarene Is.; Malesia, Queensland, Polynesia HAB. Riverine forest of Cynometra alexandri, Khaya anthotheca etc., swamps with Raphia, sometimes in shallow water; rain-forest, rock-faces (in Usambaras); 400–1200 m

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FIG. 4. CALLIPTERIS PROLIFERA— 1, rhizome,×⅔; 2, frond, 3, part of frond with bulbils, enlarged; 4, plantlet,×⅔; 5, scale,×8; 6, hairs on scale margin, enlarged. 1 from Cheek 5864; 2, 5, 6, from Poulsen 910; 3 from Leeuwenberg 2336; 4 from Paylo 614. Drawn by Pat Halliday.

Callipteris

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SYN. Asplenium proliferum Lam., Encycl. Méth. 2:807 (1786); Hieron. in P.O.A. C: 83 (1895) A. decussatum Sw. in J. Bot. (Schrad.) 1800 (2): 51 (1800) & in Syn. Fil.: 76, 260 (1806); Willd., Sp. Pl. ed. 5, 5:310 (1810). Type: Mauritius, coll. ignot.? (? S, B-W! 19877 iso., microfiche) Diplazium proliferum (Lam.) Kaulf.*, Enum. Fil.: 182 (1824); Hieron. in V.E. 2:24, fig. 20/d–f (1908); Bonap., Notes Ptérid. 1:78 (1915); Alston, J. Bot. 72, Suppl. Pterid.: 5 (1934) & Ferns W.T.A.: 65 (1959); Schelpe, C.F.A. Pterid.: 163 (1972); Johns in Kew Mag. 8(3): 128, t. 178, fig. 5, (1991); Schippers in Fern Gaz. 14:205 (1993) D. incisum Schum., Beskr. Guin. Pl.: 458 (1827) & in K.Danske Vid. Selsk. Nat. Mat. Afh. 4:232 (1829). Type: Ghana, Akwapim, Isert s.n., Thonning s.n. (C, syn.)** D. stratum Schum., Beskr. Guin. Pl.: 459 (1827) & in K. Danske Vid. Selsk. Nat. Mat. Afh. 4:233 (1829). Type: Ghana, Akwapim, ?Thonning s.n. (C, holo.)** Athyrium proliferum (Lam.) Milde in Bot. Zeit. 1870:353 (1870); Tardieu, Mém. I.F.A.N. 28:163, t. 31, figs. 3–4 (1953) Diplazium decussatum sensu Peter, F.D.-O.A. 1:67 (1929) (Peter attributes this to (Sw.) J. Sm. in Bot. Mag. 72, comp.: 28 (1846) but Smith does not mention Swartz but a Wallich specimen from Nepal) NOTE. The species has an extensive synonymy. I have excluded some specimens from the description which appear to be different e.g. Ceylon material with pinnae 33– 42×8–11 cm, the lowest part with lobes cut almost to the midrib, 4.5–7×1.5–2.5 cm. 2. C. ulugurica Verdc, sp. nov., Diplazio arborescenti (Bory) Sw. similis, squamis rhizomatis nigromarginatis, trichomatibus marginalibus nigris distincte apice bifurcatis; a C. prolifera (Lam.) Bory frondis bipinnatis, pinnulis pinnarum inferiorum usque 2.5 cm longis, 2.2 cm latis, venatione haud anastomosanti differt. Typus: Tanzania, Morogoro District, NE Uluguru Mts, Kinole, Faden et al. 70/681 (EA!, holo., K!, EA, iso.) Rhizome erect; scales with black margins, linear-lanceolate, 1.6–2.4 cm long, 0.8–2 mm wide at the base, very attenuate, dull, the marginal trichomes black and nearly all bifid. Fronds tufted, 0.5–1.5 m tall, somewhat fleshy; stipe straw-coloured, 40–45 cm long, densely scaly near base or for some distance. Lamina oblong-lanceolate, mostly bipinnate, 21–60 cm long, 32(–?60) cm wide, the apical part 7–11 cm long, 5.5–6 cm wide, long-attenuate, divided into oblong lobes 1–3 cm long, 0.6–1 cm wide, crenate; upper pinnae in 6–7 pairs, narrowly oblong-triangular, 4.5–13 cm long, 1.2–3.3 cm wide, ±truncate at base, narrowly attenuate and crenate at the apex, the upper ones sessile, shallowly lobed, the lower stalked, with up to 12 pairs of oblong lobes up to 2 cm long, 0.9 cm wide; lower pinnae in 4 pairs, oblong-attenuate, 13–28 cm long, 6–11 cm wide with (3–)8–9(–11) pairs of pinnules up to 5.4–7.5 cm long, 1.4–2.2 cm wide, very shallowly lobed or crenate; pinnae apices similar to frond apices; venation not anastomosing. Sori brown, linear, curved, 4–7 mm long, diverging from midrib, appearing different according to the rank of the segment involved; in the lobes of apices of fronds and pinnae and the ultimate pinnules there are 2–7 pairs of sori; in the upper pinnae where lobes are not divided to the base the lobes have 1–3 pairs according to position; the main sori are on veins ending in sinuses of the marginal divisions but occasionally there are much shorter sori on lateral veins; capitate paraphyses± numerous, often with dark heads; indusia very narrow, unilaterally attached.

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TANZANIA. Morogoro District: NE Uluguru Mts, Kinole, 28 Sept. 1970, Faden et al. 70/681! & 30 Apr. 1970, Pócs & Harris 6165P! & E slope of Lupanga, 11 Oct. 1971, Pócs & Mwanjabe 6470A!; Ulanga District: SW of Mahenge, Muhulu Mts, 24 Feb. 1932, Schlieben 1829! DISTR. T 6; not known elsewhere HAB. Intermediate rain-forest with Allanblackia, Parinari, Cylicomorpha, Myrianthus; 900–1050 (–?1500) m * Sometimes attributed to Thouars in Fl. Trist. d’ Ac.: 35 (1804) but I can find no mention in that work. ** The specimens are confusing—see Hepper, W. Afr. Herb. Isert & Thonning: 160 (1976).

SYN. Diplazium sp. B; Schippers in Fern Gaz. 14:205 (1993) NOTE. Pócs & Chambuko 6223X (N Uluguru Mts, below Manga peak, 16 Aug. 1970) is the same but many of the trichomes are not bifid. Pócs et al. 87201/N (Ulanga District, Mahenge Plateau, Mawenge Forest Reserve above Isongo village, 10 Oct. 1987 at 1150–1250 m) is probably also the same but I have not seen the rhizome and the lamina architecture differs. The specimen seen is not adequate but a main pinna appears to be ±40 cm long with at least 10 pairs of pinnules about 10×2.5 cm divided for ⅔–¾ distance to costa into ±12 crenate lobes each with 8 sori in 4 angled pairs. Black-headed paraphyses are numerous. More material is needed.

5. DIPLAZIUM Sw. in J. Bot. (Schrad.) 1800 (2): 61 (1801); Kato & Kramer in Kubitzki, Fam. Gen. Vasc. Pl. 1:133 (1990)

Mostly rather large ferns with creeping or erect rhizome sometimes forming a trunk-like caudex up to 1 m tall; scales non-peltate, dark, entire or with spine-like or tooth-like emergences. Lamina simple or 1–4-pinnate; veins free. Sori superficial, elliptic to linear and elongate, the indusium either attached along one side or all round or where a pair of sori are on either side of a vein the linear indusia are set back to back; indusium rarely small and fugacious. Pantropical with about 370 species. The species are difficult and this account is no more than a framework for further work. It is very important to assess characters from the same part of the frond architecture which is sometimes difficult to ascertain in poor herbarium specimens when the apex of a frond can be mistaken for a pinna and undeveloped pinnae for true pinnules; when the junction of the pinna with the main rhachis is preserved there is no difficulty. The rhizome should always be collected or at least a portion showing scales. 1. Fronds simply pinnate; pinnae 4–16 cm long, 0.8–1.8 cm wide, crenatetoothed to shallowly squarely pinnatifid (T 6, 7)

1. D. pseudoporrectum

Fronds bipinnate or almost tripinnate; pinnae distinctly divided into pinnules

2

2. Fronds with gemmae at and/or below apex

3

Fronds without gemmae 3. Pinnae 3.5–42×14–18 cm; pinnules 4–10×1.2–5 cm with strongly pinnatifid lobes 0.8–2.5 cm×3–7 mm (U 2; T 4) Pinnae up to 25×10 cm; pinnules 1.5–5×0.7–1.7 cm with very shallowly crenate almost subentire lobes 5–8×5–6 mm (T 6)

4 6. D. humbertii 5. D. ulugurense

4. Indusium narrow, attached to one side of sorus*, ‘asplenioid’, sometimes difficult to see

5

Indusium wider, attached all around the sorus, ‘allantodioid’, bursting irregularly, across top or sometimes along one edge, often leaving fragments all around the sorus but very obvious before dehiscence

6

* Contiguous sori on either side of a usually basal vein have their indusia back to back and this can resemble a single sorus appearing to have a centrally split indusium). 5. Sori 3–4(–5) mm long, usually 1–6 per pinnule-lobe* (see note after species) often only 1 (on lowest acroscopic vein of the venation in each pinnule lobe); pinnules shallowly to deeply cut but lamina not appearing tripinnate; pinnulelobes mostly crenulate only at the broadly rounded apex; rhizome-scales 2. D. nemorale

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broader and shorter, 9×2–5 mm; paraphyses present Sori 1–3(–4 on basal vein) mm long, 7–(11–23) per pinnule-lobe (see note after generic description); pinnules very deeply cut so that lamina appears almost tripinnate; pinnule-lobes crenate-lobed to serrate; rhizome-scales long 3. D. and narrow, up to 3.2 cm long and 0.5–1.5(–2) mm wide; paraphyses absent zanzibaricum** 6. Lamina bipinnate with pinnules divided into 6–9 pairs of lobes for only ½ to ⅔ of width, not cut to near base, and lobes ±entire all round or toothed or crenate at apex; paraphyses short and few (K 5, Kakamega Forest)

8. D. sp. B

Lamina virtually tripinnate with pinnules divided into 10–15 pairs of lobes almost or quite cut to the base but still decurrent and not entirely free; lobes mostly deeply crenate all round; paraphyses mostly obvious and numerous (U 2, K 5, T 4, including Kakamega Forest) (forms of D. humbertii without gemmae will key here—they are even closer to being tripinnate with the 7. D. lobes more separate as well as more deeply cut) velaminosum

1. D. pseudoporrectum Hieron. in E.J. 28:342 (1900) & in V.E. 2:24 (1908); F.D.-O.A.. 1:67 (1929). Types: Tanzania, Morogoro District, SE Uluguru Mts, Nghweme [Ng’heme], Stuhlmann 8796, 8810 (B, syn.)*** Rhizome erect or ascending, woody; scales brown, lanceolate, 5–7 mm long, 1.2–2 mm wide, reticulate, the cell walls darker and strongly raised, margins with short blunt emergences. Fronds tufted, 50–90 cm tall, simply pinnate; stipe 22–40 cm long, scaly at base. Lamina oblong-lanceolate, 33–68 cm long, 10.5–29 cm wide, with 14–31 pairs of pinnae and apical triangular segments 8.5–12 cm long, 4 cm wide, toothed at apex and pinnatifid beneath with ±5 lobes on each side; pinnae stipitate, ±falcate, narrowly oblonglanceolate, 4–16 cm long, 0.8–1.8 cm wide, forwardly directed crenate-toothed to shallowly squarely pinnatifid, the lobes ± crenate; up to 30 lobes on each side and pinnule apex narrowly attenuate, crenulate; stipes of pinnae 4–5 mm long. Sori 1–3 on each side of pinna costa for each lobe, diverging, unequal, the longer collateral pair 6–8 mm long, the shorter 3 mm long; indusium elongate. Fig. 5 (page 18). TANZANIA. Kilosa District: Ukaguru Mts, NNE slope of Mamiwa ridge, below Mnyera peak, 30 July 1972, Pócs & Mabberley 6739A!; Morogoro District: Uluguru Mts, Morogoro to Lupanga Peak track, 16 Aug. 1951, Greenway & Eggeling 8615!; Iringa District: Mwanihana Forest Reserve, above Sanje village, 10 Oct. 1984, D.W.Thomas 3857! * See note after generic description. ** A specimen from the Sese Is. swamp forests will key here but has a more developed indusium— see species 4, D. sp. A. *** A fragment of type material is preserved at the BM! (C.Christensen Herb 4889) but it is not stated from which Stuhlmann sheet it came.

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FIG. 5. DIPLAZIUM PSEUDOPORRECTUM—1, rhizome,×⅔; 2, frond showing both surfaces, ×⅓; 3, pinna,×1; 4, venation, enlarged and diagrammatic; 5, indusium, enlarged and diagrammatic. All from Faden 74/386. Drawn by Pat Halliday.

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DISTR. T 6, 7; not known elsewhere (see note) HAB. Montane forest including mist forest, Allanblackia–Cussonia–Dasylepis– Ocotea, Cyathea–Ocotea–Podocarpus and Parinari–Newtonia–Ocotea–Macaranga; 1400–2100 m SYN. D. pseudoporrectum Hieron. var. angustipinnatum Reimers in N.B.G.B. 11:922 (1933); Schippers in Fern Gaz. 14:205 (1933). Type: Tanzania, Morogoro District, NW Uluguru Mts, Schlieben 3108 (B, holo., BM!, iso.)* NOTE. In Schlieben 3108 all the pinnae have retained the very shallowly toothed margins of the uppermost pinnae but new material shows variations in this respect. Some sheets of the species had been determined as D. silvaticum (Bory) Sw. and some Burmese specimens of that are very similar. It would be foolish to make suggestions without a revision of the Old World species. 2. D. nemorale (Bak.) Schelpe in Bol. Soc. Brot., sér. 2, 41:212 (1967) & in F.Z. Pterid.: 205, t. 58a (1970); Schelpe & Diniz, Fl. Moçamb., Pterid.: 221 (1979); W. Jacobsen, Ferns S. Afr.: 408, t. 306 (1983); J. Burrows, S. Afr. Ferns: 278, t. 46/3, fig. 66/282, 282a (1990); Faden in U.K.W.F. ed. 2:33 (1994). Type: Madagascar, Antananarivo, Pool s.n. (K!, holo.) Rhizome erect or ascending forming caudex, up to 20(–30) cm long, 7–20 cm in diameter; scales shiny bronze-brown, 9 mm long, 2–5 mm wide, the edges not or narrowly dark and with few undivided trichomes. Fronds tufted, 0.5–1.8 m tall; stipe brown, 18–90 cm long, grooved above, swollen and slightly scaly at the base. Lamina ±ovate, 45–95 cm long, 30–70 cm wide, bipinnate-pinnatifid; apical segment 7–12 cm long, 3.5–5 cm wide, attenuate and crenate at the tip, lobed beneath; pinnae in 10–15 pairs, the ± 5 below the apical segment pinnatifid, 9 cm long, 2 cm wide, the lobes crenate; rest of pinnae oblong to oblong-ovate, pinnate, 20–50 cm long, 3.5–22 cm wide, with (7–)10–16 pairs of triangular-lanceolate pinnules 2.5–11.5 cm long, 1–2.7 cm wide, shallowly to very deeply pinnatifid according to position, narrowly acuminate and crenulate at the apex; pinna-stalks 0–1.5(–3) cm long; venation not anastomosing. Sori brown, linear, 3–4(–5) mm long, usually one on the lowest acroscopic vein of the branching system in each pinnule-lobe but there are usually other shorter ones present, 1– 6(–9) in the basal lobes but usually only 1 in the upper lobes; the appearance of the son varies according to which part of the frond is examined; paraphyses present; indusium very narrow, unilaterally attached. KENYA. Embu District: SE Mt Kenya, Irangi Forest Station, 30 Apr. 1972, Faden et al. 72/193! TANZANIA. Lushoto District: Shagayu Forest, valley N of Shagein, along tributary to Umba R., 21 Oct. 1986, Schippers 1589!, Morogoro District: Uluguru Mts, Ululu Falls, near Bunduki fishing camp, 27 Nov. 1974, Balslev 356; Iringa District: Mwanihana Forest Reserve, above Sanje Village, 10 Oct. 1984, D.W.Thomas 3863! DISTR. K 4; T 3, 5–7; São Tomé, Malawi, Mozambique, E Zimbabwe HAB. Montane and lower montane forest e.g. Ocotea–Ensete–Dracaena–Cyathea– Parinari– Strombosia–Syzygium–Myrianthus–Ficalhoa, in swampy places and on steep slopes; (1100–)1350–1900m** SYN. Asplenium nemorale Bak. in J.L.S. 15:417 (1876) A. hylophilum Hieron. in P.O.A. C.: 84 (1895). Type: Tanzania, Lushoto District, W Usambaras, Mbaramu, Holst 2480 (B, holo., BM!, fragment & photo, K!, iso.) (see note)

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25

Diplazium hylophilum (Hieron.) C. Chr., Index Fil.: 233 (1905); Faden in U.K.W.F. ed. 1:57 (1974); Schippers in Fern Gaz. 14:205 (1993) D. stolzii Brause in E.J. 53:381 (1915). Type: Tanzania, Rungwe District, Rungwe, Stolz 1233 (B, holo., BM!, fragment & photo., K!, iso.) D. arborescens (Bory) Sw. var. nemorale (Bak.) C. Chr. in Perrier, Cat. Fl. Madag., Ptérid.: 36 (1932) * Schippers gives the type as Stuhlmann 8796 but this is not correct. ** It is difficult to sort out the exact ecology of the collected material from field notes which give general ranges for a large area.

NOTE. I at first agreed with Christensen that D. nemorale was not more than a local variety of D. arborescens and that D. hylophilum could be looked on as another variety but since the situation is not resolved I have not disturbed Schelpe’s treatment of the species. Two sheets of the eight making up the EA gathering of Faden et al. 72/193 are a young plant with the largest pinnules only 2.5×1 cm and not deeply pinnatifid. The isotype of D. hylophilum preserved at Kew consists of the top of a frond with similar small pinnules so I believe it has been interpreted correctly but mature specimens with rhizomes are needed from the Usambaras. The type of D. nemorale consists of a single pinna and it does not seem certain it is a pinna with pinnatifid pinnules rather than the apex of a frond with pinnatifid pinnae. The somewhat cordate base of the pinnules is a minor variation not matched elsewhere. D. arborescens (Bory) Sw., Syn. Fil.: 92 (1806); Willd., Sp. Pl. ed. 5, 5:354 (1810) (Type: Reunion, by R. St. Denis, Bory de St Vincent s.n. (P, holo., B-W 19950, iso. (microfiche!)) differs in having 6–15 sori in all the pinnules and they are joined basally in pairs at the mid vein. Material from the Comoro Is. is particularly characteristic. There are, however, specimens from Zimbabwe which are deceptively similar e.g. B.S.Fisher 1331 from Umtali, Vumba, 18 July 1947. Bory describes it as having a big trunk. A further problem is the identity of material from West Africa cited by Alston (F.W.T.A. Pterid.: 65 (1959)) Schippers 1130 (Iringa District, Mufindi, slope towards Kigogo R., 1630 m, 9 Nov. 1985) is probably only a form of D. nemorale with rather different pinna architecture and more sori; a lower pinnule is 40×19 cm with largest pinnule-lobes up to 20×9 mm with up to 14 sori. More material with rhizome is needed to decide its status. This is D. sp. C of Schippers in Fern Gaz. 14:205 (1993). 3. D. zanzibaricum (Bak.) C. Chr., Ind. Fil.: 241 (1905); Schelpe in F.Z., Pterid.: 205, t. 58b (1970); Schelpe & Diniz, Fl. Moçamb., Pterid.: 222 (1979); W.Jacobsen, Ferns S.Afr.: 410, t. 307 (1983); Schelpe & N.C.Anthony in F.S.A., Pterid.; 227 (1986); J.E.Burrows, S.Afr.Ferns: 276, t. 46/41, fig. 66/281, 281a (1990); Schippers in Fern Gaz. 14:205 (1993); Faden in U.K.W.F. ed. 2:33 (1994). Type: Tanzania, mainland W of Zanzibar, probably Nguru Mts, Last 261 (K!, holo.) Rhizome large, erect; scales pale chestnut brown, linear to linear-lanceolate, 1.5–3.2 cm long, 0.5–1.5(–2) mm wide, finely attenuate, not black-margined or rarely partly so (see note), with numerous stiff spiniform trichomes, very narrowly striate with raised walls of the elongate reticulation; trunk 0.4–1.2 m long, 15 cm wide. Fronds tufted, 1–2.5 m tall; stipes up to 1.5 m tall, with 2 C-shaped bundles, very densely covered with scales similar to rhizome scales at base. Lamina ovate-lanceolate, 1.3–1.6 m long, 0.9–1.1 m

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wide, bipinnate, with about 15 pairs of pinnae; pinnae (5–)42–68 cm long, (1–)11–25 cm wide, stalk up to 2.5 cm long; pinnules in 12 to 27 pairs, oblong-lanceolate to oblongtriangular, (1–)7–13 cm long, (0.3–)1.5–4 cm wide, attenuated to a crenate apex, deeply cut into 13–15 pairs of lobes so that lamina is almost tripinnate; lobes narrowly oblong, 0.5–2 cm long, 2–8 mm wide, crenate-lobed to serrate, including the apex; costae often with multicellular ±serrate hairs up to 1.5 mm but sometimes entirely glabrous; venation free, the veins often pale above towards margin. Sori 1–3 mm (–4 mm on basal vein) long, (7–)11–23 per pinnule-lobe; indusium narrow, on one side only, or when two sori are paired and contiguous on basal vein, the two indusia are persistent between them; young sori not covered with wide indusium all round; paraphyses absent. UGANDA. Toro District: E Ruwenzori, Bwamba Pass, July 1940, Eggeling 4007! (see note) & Bwamba Pass, 3 Feb. 1934, Longfield 39!; Kigezi District: Kinaba Gap, Dec. 1938, Chandler & Hancock 2540! KENYA. Meru District: NE Mt Kenya, Ithanguni Forest, road along base of volcanic cone Kirui and 15 km from Nkubu, 22 June 1969, Faden et al. 69/764! & Nyambeni Hills, above Kiegoi, 1 June 1969, Faden et al. 69/665! TANZANIA. Pare District: N Pare Mts, Kindoroko Forest Reserve, 24 Feb. 1987, Schippers 1771!; Morogoro District: Uluguru Mts, Bondwa, 2 May 1970, Pócs 6183/E!; Rungwe District: Rungwe Forest, Tukuyu, June 1958, Watermeyer 39! DISTR. U 2; K 4; T 2, 3, 6, 7*; W Congo (Kinshasa), Zambia, Malawi, Mozambique, Zimbabwe & South Africa; also recorded from Madagascar and Comoro Is. HAB. Evergreen forest, e.g. Podocarpus–Lasianthus–Psychotria–Pauridiantha– Xymalos,Cyathea forest, bamboo forest, occasionally in swampy areas; 1450–2550 m SYN. Asplenium zanzibaricum Bak. in Ann. Bot. 5:311 (1891); F.D.-O.A. 1:80 (1929) Aspidium sulcinervium Hieron. in P.O.A. C: 85 (1895). Type: Tanzania, Kilimanjaro, Marangu, 2300 m, Volkens 1492 (B, holo., BM!, iso. (pro parte)) Diplazium sulcinervium (Hieron.) C. Chr. in Perrier, Cat. Pl. Madag. Pterid.: 37 (1932) D. sp. D; Schippers in Fern Gaz. 14:205 (1993) NOTE. In Eggeling 4007 a few areas of the scale margin are black with black trichomes and a very few have a minute bifurcation at the apex. The indusium is almost invisible and flattened into the lobe surface. More material is needed. 4. D. sp. A Scales on stipe-base black-edged with trichomes showing a trace of bifurcation. Fronds up to 1.2 m tall; stipe 56 cm long, slender. Lamina narrowly triangular, 60 cm long, 45 cm wide with ±13 pairs of pinnae up to 20 cm long, 8 cm wide with long crenate apex; pinnules up to 15 pairs, lanceolate, serrate at the tip, 1.5–6.5 cm long, 0.4–1.8 cm wide, pinnatifid for about ⅓–¾ into up to 10 pairs of oblong lobes, subentire or slightly crenate, 3–8 mm long, 2–5 mm wide. Sori linear to oblong, 1–3 mm long, 1–7 per lobe, those nearest the pinnule costule being the longest; paraphyses lacking; indusium wider than in D. zanzibaricum but splitting and remaining on one side of sorus, with dark reticulation. UGANDA. Masaka District: Sese Is., Towa Forest, 30 June 1935, A.S.Thomas 1350! DISTR. U 4 HAB. Primary forest; 1200 m NOTE. This is close to D. zanzibaricum in the lack of paraphyses but the indusium is more developed and the pinnule-lobes less crenate than in typical material. It is the only

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27

specimen of Diplazium seen from the Sese swamp forest and may be distinct. It resembles much W African material probably misnamed D. hylophilum. 5. D. ulugurense Verdc, sp. nov. affinis D. humbertii (C. Chr.) Pic. Serm., pinnis, pinnulis et lobis ultimis pinnularum minoribus, lobis ipsis magis quadratis breviter crenato-dentatis vel subintegris haud profunde pinnatifidis differt. Typus: Tanzania, Morogoro District, Uluguru Mts, E slope of Bondwa, Faden et al. 70/635 (K!, holo., iso., EA, iso.) Rhizome thick and fleshy, erect to ascending; scales pale chestnut, narrowly lanceolate, 8–12 mm long, 1.2–2 mm wide, attenuate, not black-margined and with few simple trichomes, prominently reticulately nerved. Fronds tufted, 1.2 m tall; stipe 46 cm long, scaly at base. Lamina triangular-ovate, ±70 cm long, 48 cm wide, bipinnate, gemmiferous at apices of some pinnae and probably from near apex of frond also; pinnae in ±13 pairs, oblong-lanceolate, up to 25 cm long, 10 cm wide, deeply pinnatifid at apex and with ±6–12 free pinnules beneath, according to position; pinnules oblong, 1.5–5 cm long, 0.7–1.7 cm wide, shortly narrowed at the apex, crenate to distinctly pinnatifid; ultimate lobes of largest pinnae in ±7 pairs, ± square, up to 8 mm long, 6 mm wide, crenate-toothed. Sori elliptic to linear, 1.5–3 mm long, one on acroscopic basal nerve of some ultimate lobes but only about 2–5 per pinnule, the upper sori quite small; indusium breaking irregularly, spotted and streaked inside with brown; paraphyses evident. * A sheet labelled Zanguebar, Sir John Kirk recd. 10/[18] 82’ is undoubtedly not from Zanzibar but from somewhere in T 6.

TANZANIA. Morogoro District: Uluguru Mts, E slope of Bondwa, 26 Sept. 1970, Faden et al. 70/635! DISTR. T 6 HAB. Montane evergreen rainforest; 1650 m SYN. D. sp. A; Schippers in Fern Gaz. 14:205 (1993) NOTE. Material has been annotated with a specific epithet based on ‘Uluguru’ but this has not been published (see Johns, Pterid. T.E.A.: 84 (1991). I have therefore taken up the name. 6. D. humbertii (C. Chr.) Pic. Serm. in Webbia 27:444 (1973) (‘1972’). Type: Congo (Kinshasa), mountains W of Lake Kivu, Humbert 7497 (BM!, lecto.*, fragment and photo) Rhizome ascending, thick, fleshy, up to 7 cm in diameter, elongate, with scales dark chestnut, linear, 7–23 mm long, 1.5–2 mm wide, reticulate, mostly black-edged, entire, not filiform at apex. Fronds tufted, 1.3–1.5 m tall, sometimes gemmiferous in upper axils; stipe greyish straw-coloured, 30–112 cm long, deeply trisulcate above, with dense scales at base but ±glabrous. Lamina triangular-ovate, up to 1 m long, 82 cm wide, bipinnatepinnatifid to virtually tripinnate with decurrent secondary pinnules; rachis strawcoloured; pinnae in ± 16 pairs, oblong-lanceolate, (8–)35–42 cm long, (3–)14–18.5 cm wide, with stalks 1–2 cm long and 10–16 pairs of pinnules; pinnules oblong, (1–)4–10 cm long, (0.5–)1.2–5 cm wide with 9–12 pairs of narrowly oblong-lanceolate to oblong lobes 0.8–2.5 cm long, 3–7 mm wide; lobes themselves mostly deeply crenate-lobed (up to ±3 mm) but those on apical pinnules and towards apex of basal pinnules are ±entire. Sori almost round to elliptic, 1–1.5 mm long, 1–18 per pinnule lobe (one at base of each

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secondary lobe of the lobe); paraphyses dense; indusium covering and attached all round the sorus, bursting irregularly into lobes and leaving a cup and remnants or dorsally dehiscent. UGANDA. Toro District: Ruwenzori, Nyamgasani Valley, Jan. 1935, Synge 1470!; Ankole District: Kalinzu Forest, 4 km NW of saw mill, W of Rubuzigye, 19 Sep. 1969, Faden et al. 69/1135! & 69/1136! & 69/1135! & Kasyoha-Kitomi Forest Reserve, NE of Kyambura R, 9 June 1994, Poulsen et al. 560! (see note) TANZANIA. Mpanda District: Mahali Mts, Sisaga, 29 Aug. 1958, Newbould & Jefford 1930! DISTR. U 2; T 4; Congo (Kinshasa) (Kivu) HAB. Lower montane forest, Parinari excelsa–Carapa–Polyscias, bamboo forest and mist forest, in swampy places and by rivers; 1300–1450 m (Uganda) and 2250–2500 m (Tanzania) SYN. Athyrium humbertii C. Chr. in Dansk Bot. Arkiv 9 (3): 54, t. 6 /7–9 (1937) NOTE. Poulsen et al. 560 cited above, and Poulsen et al. 607 from the same forest appear not to be gemmiferous. The Tanzanian specimen also has no gemmae but is closely similar to typical D. humbertii. Where some characters differ from the description above the extra information is available as follows—rhizome horizontal (fide collector) with scales up to 23 mm long, attenuate; fronds up to 2 m long with lamina 1 m long and 82 cm wide; pinnae in ± 16 pairs, 8–42 cm long, 3–15 cm wide; pinnules in 10–15 pairs; sori 0.5–1.2 (-2) mm wide. It is quite possible that this will prove only a form of D. velaminosum. Populations of both need examining for detailed frond architecture and presence of gemmae. 7. D. velaminosum (Diels) Pic. Serm. in Webbia 27:443 (1973) (‘1972’); Faden in U.K.W.F.: 57, 59 (1974). Type: Cameroon, Buea, ‘in Unter-Buea, nahe dem Ost-Ende’, Preuss 910 (B, syn., BM!, isosyn.)** * The two sheets mentioned by Pichi Sermolli could not be found at the BM. ** I do not know why Alston did not find the type material at Berlin, only an empty cover—it may have been on loan or misplaced in post-war chaos—but even more inexplicable is the fact that a duplicate of Preuss 910 was in the BM and was overlooked.

Rhizome ascending to erect, stout, to 40 cm long, 10 mm diameter, with mostly blackedged linear-lanceolate scales up to 3 cm long, 1–2 mm wide, scales with fine-tipped marginal trichomes. Fronds tufted 1.2–1.8 m tall, not proliferous; stipe 64–100 cm long. Lamina up to 90(–140) cm wide (not seen complete), bipinnate; pinnae triangularlanceolate in outline, 14–44(–70) cm long, 11–21 cm wide, with attenuate crenate apex and 9–23 pairs of pinnules; pinnules triangular-lanceolate, 3.5–11 cm long and 1–3 cm wide, with attenuate crenate apex and divided almost to the base (in lower pinnules) into 10–15 oblong lobes 0.5–2.5 cm long, 3–8 mm wide which are subentire to shallowly pinnatifid; venation often with scattered long multicellular hairs. Sori elliptic, 1–12 per lobe, 0.5–3(–4) mm long, the 2 lowest often contiguous lengthwise; paraphyses mostly numerous and obvious but few and short in some specimens; indusium wide and obvious before dehiscence, attached all round the sorus and often bursting irregularly across the top or sometimes only along one edge, or leaving fragments all around the sorus. UGANDA. Toro District: Kibale National Park, near Kanyawara, 13 July 1994, Poulsen et al. 656! & Kibale [Kebale] Forest, 14 Dec. 1957, B.E.G.Allen 3711!; Kigezi

Diplazium

29

District: Bwindi National Park, N Sector [Kayonza], Ishasha Gorge, 28 Mar. 1995, Poulsen et al. 818! KENYA. N Kavirondo District: Kakamega Forest, Kibiri Block, S side of Yala R., 20 Jun. 1970, Faden et al. 70/10! DISTR. U 2, 3; K 5; Guinea, Liberia, São Tomé, Cameroon HAB. Forest, Croton megalocarpus–Bequaertiodendron–Dracaena etc., often in swampy places; 1200–1600 m SYN. ?Hemitelia velaminosa Diels in E. & P. Pf. 1 (4): 132 (1899); Alston in F.W.T.A. Pterid.: 29 (1959) Cyathea velaminosa (Diels) Domin, Pterid.: 264 (1929) [Diplazium zanzibaricum sensu Alston, F.W.T.A. Pterid.: 64 (1959) pro parte; Tardieu, Fl. Cam. 3 Pterid.: 236 (1964) pro parte, non (Bak.) C. Chr.] 8. D. sp. B Rhizome erect, fleshy; scales lanceolate-attenuate, 16 mm long, 1.5–2 mm wide, darkedged with few trichomes. Fronds tufted, ±1 m tall, bipinnate, not gemmiferous; stipe green, grooved, black and swollen at base, 25–35 cm tall; pinnae in 11–15 pairs, oblonglanceolate, up to 28 cm long, 9.5 cm wide with pinnatifid apical segment and 11–15 pairs of pinnules; pinnules oblong-lanceolate, 2.5–7 cm long, 0.8–2.8 cm wide, with crenate apex and divided into 6–9 pairs of lobes for about ½–⅔ of width; free part of lobes oblong, 3–8 mm long, 3–6 mm wide, often falcate near apex, ±entire all round or toothed or crenate at apex. Sori elliptic to oblong, 1.5–3 mm long, 1–5 per lobe; paraphyses short and few; indusium covering and fixed all round the sorus, breaking irregularly but sometimes little or none left in mature sori. KENYA. N Kavirondo district: Kakamega Forest, NE of Forest Station, 24 Nov. 1969, Faden et al. 69/1976! & along Yala R., about 4.8 km SE of Forest Station, 25 Nov. 1969, Faden et al. 69/2023! DISTR. K 5 HAB. Evergreen forest of Celtis–Croton and Sapotaceae, by stream; 1650–1700 m SYN. [D. velaminosum sensu Faden in U.K.W.F. ed. 1:57 (1974) pro parte] D. sp. sensu Faden in U.K.W.F. ed. 2:33 (1994) NOTE. Faden (1994) points out that this taxon has the broader less divided lobes of D. nemorale but small sori. UNIDENTIFIED WOODSIACEAE Rhizome not seen. Fronds about 1 m tall; stipe not seen. Lamina ±ovate, up to 48 cm long, 24 cm wide, 2-pinnate with pinnules deeply pinnatifid so that frond appears 3pinnate but costulae are very narrowly winged by the decurrent ultimate segment-bases; pinnae in at least 15 pairs, lanceolate, up to 14 cm long, 6 cm wide with 15–17 pairs of pinnules; pinnules oblong-lanceolate, up to 4 cm long, 1–3 cm wide with 5–9 pairs of segments up to 7 mm long, 4 mm wide; segments with 1–2 bifid lateral lobes on each side and terminal 2–4-fid lobe, the teeth subacute; costulae with rather sparse multicellular hairs up to ±1mm long. Sori not seen. TANZANIA. Lushoto District: W Usambaras, Shume-Magamba Forest Reserve, 2 May 1987, Kisena 577 pro parte! DISTR. T 3 HAB. Presumably forest; 1680m

Flora of tropical East Africa

30

NOTE. Sheets 3, 4 & 5 out of 5 are this species, 1 & 2 being a Dryopteris species. Sheets 3–5 had been named Athyrium scandicinum but are not that species and show more resemblance to Dryoathyrium but are not a good match for the species treated in this part. Possibly this also is a Dryopteris.

INDEX TO WOODSIACEAE Arachnoides foliosa (C. Chr.) Schelpe, 12 Aspidium boryanum Willd., 2 Aspidium glabratum Kuhn, 2 Aspidium kiboschense Hieron., 2 Aspidium scandicinum Willd., 10 Aspidium sulcinervium Hieron., 21 Aspidium viridulum Desv., 7 Asplenium decussatum Sw., 15 Asplenium hylophilum Hieron., 19 Asplenium nemorale Bak., 19 Asplenium proliferum Lam., 15 Asplenium schimperi (Fée) A.Braun, 10 Asplenium zanzibaricum Bak., 21 ATHYRIUM Roth, 8 Athyrium annae Kornaś, 12 Athyrium boryanum (Willd.) Tagawa, 4 Athyrium glabratum (Kuhn) Alston, 4 Athyrium humbertii C. Chr., 22 Athyrium laxum Pappe & Raws., 11 Athyrium lewalleanum Pic. Serm., 12 Athyrium newtonii Bak., 11 Athyrium proliferum (Lam.) Milde, 15 Athyrium rondoense Verdc., 12 Athyrium scandicinum sensu auct, 11 Athyrium scandicinum (Willd.) Presl, 10, 11, 12 subsp. newtonii (Bak.) Verdc., 10 var. rhodesianum Schelpe, 11 var. scandicum sensu Schelpe, 11 Athyrium schimperi Fée, 8, 9, 11 Athyrium sp., 11 CALLIPTERIS Bory, 13 Callipteris prolifera (Lam.) Bory, 13, 14, 15 Callipteris ulugurica Verdc., 15 Cornopteris boryana (Willd.) Tardieu, 4 Ctenitis boryana (Willd.) Copel., 4 Cyathea velaminosa (Diels) Domin, 23 CYSTOPTERIS Bernh., 4 Cystopteris diaphana (Bory) Blasdell, 7 Cystopteris filix-fragilis sensu auctt., 5 Cystopteris fragilis (L.) Bernh., 4, 6 subsp. diaphana (Bory) Litard, 7

Index to woodsiaceae subsp. diaphana sensu auct., 7 subsp. A, 5 subsp. B, 7 var. fragilis, 5 Cystopteris nivalis (Pirotta) Pic. Serm., 7, 5 Cystopteris viridula (Desv.) Desv., 7 Deparia boryana (Willd.) Kato, 4 Deparia glabrata (Kuhn) Kato, 4 DIPLAZIUM Sw., 16, 1 Diplazium arborescens (Bory) Sw., 15, 20 var. nemorale (Bak.) C. Chr., 19 Diplazium decussatum sensu auct., 15 Diplazium humbertii (C. Chr) Pic. Serm., 22, 21 Diplazium hylophilum (Hieron.) C. Chr., 19, 20, 21 Diplazium incisum Schum., 15 Diplazium nemorale (Bak.) Schelpe, 19, 20, 23 Diplazium proliferum (Lam.) Kaulf., 15 Diplazium pseudoporrectum Hieron, 17, 18 var. angustipinnatum Reimers, 19 Diplazium serratum Schum., 15 Diplazium silvaticum (Bory) Sw., 19 Diplazium stolzii Brause, 19 Diplazium sulcinervium (Hieron.) C. Chr., 21 Diplazium sp., 21, 23 Diplazium sp. A of Schippers, 22 Diplazium sp. B of Schippers, 16 Diplazium sp. C of Schippers, 20 Diplazium sp. D of Schippers, 21 Diplazium sp. sensu Faden in U.K.W.F., 23 Diplazium ulugurense Verdc., 21 Diplazium velaminosum (Diels) Pic. Serm., 22 Diplazium velaminosum sensu Faden, 23 Diplazium zanzibaricum (Bak.) C. Chr, 20, 21, 23 Diplazium zanzibaricum sensu auct., 23 DRYOATHYRIUM Ching., 1 Dryoathyrium boryanum (Willd.) Ching, 2, 3 Dryopteris boryana (Willd.) C. Chr., 2 Dryopteris glabrata (Kuhn) Kuntze, 2 Dryopteris kiboschensis (Hieron.) C. Chr., 2 ?Hemitelia velaminosa Diels, 23 Lastrea boryana (Willd.) Moore, 2 Lunathyrium boryanum (Willd.) H.Ohba, 4 Nephrodium boryanum (Willd.) Bak., 2 Nephrodium catopteron (Kunze) Hook., 2 var. glabrum Hook., 2 Nephrodium catopteron sensu auct., 2

32

Index to woodsiaceae

33

Nephrodium glabratum (Kuhn) Bak., 2 Parathyrium boryanum (Willd.) Holttum, 4 Polypodium fragile L., 5 Polypodium diaphanum Bory, 7 Thelypteris glabrata (Kuhn) Tardieu, 4 var. hirsuta Tardieu, 4 Woodsia nivalis Pirotta 7

New names validated in this part Athyrium rondoense Verdc. Athyrium scandicinum (Willd.) Presl subsp. newtonii (Bak.) Verdc. Callipteris ulugurica Verdc. Diplazium ulugurense Verdc.

GEOGRAPHICAL DIVISIONS OF THE FLORA

Index to woodsiaceae

Bignoniaceae Bischofiaceae—in Euphorbiaceae Bixaceae (£1.50) Bombacaceae (£3.90) Boraginaceae (£14.80) Brassicaceae—see Cruciferae Brexiaceae (£1.50) Buddlejaceae—as Loganiaceae Burmanniaceae (£3.00) Burseraceae (£13.30) Butomaceae (£1.50) Buxaceae (£1.50) Cabombaceae (£1.50) Cactaceae (£1.50) Caesalpiniaceae—in Leguminosae Callitrichaceae (£4.00) Campanulaceae (£4.50) Canellaceae (£1.50) Cannabaceae (£1.50) Cannaceae—with Musaceae Capparaceae (£7.50) Caprifoliaceae (£1.50) Caricaceae (£1.50) Caryophyllaceae (£3.00) Casuarinaceae (£2.00) Cecropiaceae—with Moraceae Celastraceae (£13.00) Ceratophyllaceae (£1.50) Chenopodiaceae (£3.00) Chrysobalanaceae—as Rosaceae Clusiaceae—see Guttiferae Cochlospermaceae (£1.50) Colchicaceae Combretaceae (£8.90) Commelinaceae Compositae Part 1 (£32.00) Part 2 (£35.00) Part 3 Connaraceae (£3.00) Convolvulaceae (£13.00) Cornaceae (£1.50) Costaceae—as Zingiberaceae Crassulaceae (£11.00) Cruciferae (£11.00) Cucurbitaceae (£13.00)

37

Index to woodsiaceae

Cyanastraceae—in Tecophilaeaceae Cyclocheilaceae (£1.75) Cymodoceaceae (£4.00) Cyperaceae Cyphiaceae—as Lobeliaceae Dichapetalaceae (£3.70) Dilleniaceae (£1.50) Dioscoreaceae (£3.00) Dipsacaceae (£3.00) Dipterocarpaceae (£3.90) Dracaenaceae Droseraceae (£1.50) Ebenaceae (£11.00) Elatinaceae (£1.50) Ericaceae Eriocaulaceae (£6.50) Eriospermaceae (£2.00) Erythroxylaceae (£3.00) Escalloniaceae (£1.50) Euphorbiaceae Part 1 (£31.50) Part 2 (£22.00) Fabaceae—see Leguminosae Flacourtiaceae (£5.90) Flagellariaceae (£1.50) Fumariaceae (£1.50) Gentianaceae (£15.00) Geraniaceae (£3.00) Gesneriaceae Gisekiaceae—as Aizoaceae Goodeniaceae (£1.50) Gramineae (£74.00) Part 1 (£13.00) Part 2 (£38.00) Part 3 (£54.50) Gunneraceae—as Haloragaceae Guttiferae (£4.50) Haloragaceae (£3.00) Hamamelidaceae (£1.50) Hernandiaceae (£3.00) Hippocrateaceae—in Celastraceae Hugoniaceae—in Linaceae Hyacinthaceae (£6.00) Hydnoraceae (£4.00) Hydrocharitaceae (£3.25) Hydrophyllaceae (£1.85)

38

Index to woodsiaceae

Hydrostachyaceae (£3.00) Hymenocardiaceae—with Euphorbiac Hypericaceae (£3.00)—see also Guttif Hypoxidaceae Icacinaceae (£3.00) Illecebraceae—as Caryophyllaceae Iridaceae (£15.00) Irvingiaceae—as Ixonanthaceae Ixonanthaceae (£3.00) Juncaceae (£3.00) Juncaginaceae (£1.50) Labiatae Lamiaceae—see Labiatae Lauraceae (£4.00) Lecythidaceae (£1.50) Leeaceae—with Vitaceae Leguminosae (£74.00) Part 1, Mimosoideae (£13.00) Part 2, Caesalpinioideae (£18.50) Part 3 Papilionoideae (£59.00) Part 4 Papilionoideae (£59.00) Lemnaceae (£3.00) Lentibulariaceae (£3.00) Limnocharitaceae—as Butomaceae Linaceae (£3.00) Lobeliaceae (£8.30) Loganiaceae (£4.50) Loranthaceae (£12.75) Lythraceae (£11.20) Malpighiaceae (£3.00) Malvaceae Marantaceae (£3.00) Melastomataceae (£9.00) Meliaceae (£11.00) Melianthaceae (£1.50) Menispermaceae (£3.00) Menyanthaceae (£2.00) Mimosaceae—in Leguminosae Molluginaceae—as Aizoaceae Monimiaceae (£1.50) Montiniaceae (£1.50) Moraceae (£14.25) Moringaceae (£2.75) Muntingiaceae—with Tiliaceae Musaceae (£3.90) Myricaceae (£3.00)

39

Index to woodsiaceae

Myristicaceae (£2.70) Myrothamnaceae (£1.80) Myrsinaceae (£4.30) Myrtaceae (£17.00) Najadaceae (£3.90) Nectaropetalaceae—in Erythroxylaceae Nesogenaceae (£1.50) Nyctaginaceae (£4.00) Nymphaeaceae (£3.90) Ochnaceae Octoknemaceae—in Olacaceae Olacaceae (£3.00) Oleaceae (£3.00) Oliniaceae (£1.50) Onagraceae (£3.00) Opiliaceae (£1.50) Orchidaceae Part 1, Orchideae (£18.50) Part 2, Neottieae, Epidendreae (£22.00) Part 3, Epidendreae, Vandeae (£24.00) Orobanchaceae (£1.50) Oxalidaceae (£3.00) Palmae (£8.50) Pandaceae—with Euphorbiaceae Pandanaceae (£3.90) Papaveraceae (£1.50) Papilionaceae—in Leguminosae Passifloraceae (£6.00) Pedaliaceae (£3.00) Periplocaceae—see Apocynaceae Phytolaccaceae (£1.50) Piperaceae (£5.00) Pittosporaceae (£3.00) Plantaginaceae (£1.50) Plumbaginaceae (£3.00) Poaceae—see Gramineae Podostemaceae Polygalaceae Polygonaceae (£4.50) Pontederiaceae (£1.50) Portulacaceae (£10.00) Potamogetonaceae Primulaceae (£3.00) Proteaceae (£3.25) Ptaeroxylaceae (£2.00) Rafflesiaceae (£2.00)

40

Index to woodsiaceae

Ranunculaceae (£3.00) Resedaceae (£1.50) Restionaceae Rhamnaceae (£4.50) Rhizophoraceae (£3.00) Rosaceae (£6.00) Rubiaceae Part 1 (£22.00) Part 2 (£31.50) Part 3 (£23.70) Ruppiaceae (£1.95) Rutaceae (£7.75) Salicaceae (£2.05) Salvadoraceae (£3.00) Santalaceae Sapindaceae (£15.00) Sapotaceae (£7.50) Scrophulariaceae Scytopetalaceae (£1.50) Selaginaceae—in Scrophulariaceae Simaroubaceae (£3.40) Smilacaceae (£1.85) Solanaceae Sonneratiaceae (£1.50) Sphenocleaceae (£1.50) Sterculiaceae Strychnaceae—in Loganiaceae Surianaceae (£2.00) Taccaceae (£1.50) Tamaricaceae (£1.50) Tecophilaeaceae (£1.50) Ternstroemiaceae—in Theaceae Tetragoniaceae—in Aizoaceae Theaceae (£1.50) Thismiaceae—in Burmanniaceae Thymelaeaceae (£4.50) Tiliaceae (£20.50) Trapaceae (£1.50) Tribulaceae—in Zygophyllaceae Triuridaceae (£1.85) Turneraceae (£3.00) Typhaceae (£1.50) Uapacaceae—in Euphorbiaceae Ulmaceae (£3.00) Umbelliferae (£14.80) Urticaceae (£11.00)

41

Index to woodsiaceae

42

Vacciniaceae—in Ericaceae Vahliaceae (£1.50) Valerianaceae (£3.00) Velloziaceae (£3.00) Verbenaceae (£17.80) Violaceae (£6.30) Viscaceae (£6.20) Vitaceae (£17.80) Xyridaceae (£6.20) Zannichelliaceae (£1.40) Zingiberaceae (£5.75) Zosteraceae (£1.70) Zygophyllaceae (£3.15) Adviser on Linnaean types: C.Jarvis Assistant editor: Jonathan Stansbie Typeset by Margaret Newman Parts of this Flora are obtainable from: A.A.Balkema, P.O. Box 825, 2160 SZ Lisse, The Netherlands, http://balkcma.ima.nl Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, England (older parts only). www.kew.org