LIST OF ABBREVIATIONS A.V.P.=O.Hedberg, Afroalpine Vascular Plants; B.J.B.B.=Bulletin du Jardin Botanique de l’Etat, Br...
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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)
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This edition published in the Taylor & Francis e-Library, 2005. “ To purchase your own copy of this or any of Taylor & Francis or Routledge’s collection of thousands of eBooks please go to http://www.ebookstore.tandf.co.uk/.” © 2003 Royal Botanic Gardens, Kew ISBN 0-203-97114-0 Master e-book ISBN
ISBN 90-5809-412-X (Print Edition)
FLORA OF TROPICAL EAST AFRICA CALLITRICHACEAE INGA HEDBERG & OLOV HEDBERG
CALLITRICHE L., Gen. Pl., ed. 5:5 (1754); Schotsman in B.J.B.B. 55:291–296 (1985) & in Bull. Mus. Hist. Nat. Paris, ser 4, 10:3–17 (1988); O.Hedberg & I.Hedberg, Tropical African Callitriche, Biol. Skr. 54:19–30 (2001) Creeping or submerged, monoecious, aquatic herbs with filiform stems and opposite, entire exstipulate leaves. Stems with axillary glandular scales; both stem and leaves with peltate hairs. Flowers axillary, either solitary or with one male and one female flower in the same axil; perianth absent; bracteoles 2, crescent-shaped or absent; male flower consisting of one stamen with a slender filament and a reniform anther; female flower consisting of a single carpel with a 4-celled ovary containing a single anatropous ovule in each locule; styles 2, filiform. Fruit schizocarpic, separating into 4 mericarps. Seeds with a fleshy endosperm. A family with a single genus. Description as for the family. An almost cosmopolitan genus with about 30 species, of which 4 are found in the Flora area. The leaf shape and arrangement vary considerably with the habitat of the plant. Submerged plants often have lanceolate or linear leaves and long internodes, while in plants where the shoots reach the surface the internodes tend to be shorter and have rosettes of elliptic or spathulate leaves. Terrestrial plants tend to have ovate or suborbicular leaves. The adaptation of the genus to aquatic life has resulted in the reduction of several morphological features, so that only a few reliable characters remain which can be used for identification. The most useful character for identification is that of
Flora of tropical east africa callitrichaceae
2
ripe fruits. Other taxonomically useful characters, albeit unsuitable for use in species’ keys, are the morphology of pollen and the number of chromosomes. In pollen, the exine surface shows the Croton pattern, which is a reticulate pattern with processes (clavae) at the intersections of the walls (muri) (Erdtman, G., 1952, Pollen morphology and plant taxonomy: Angiosperms, p. 173). This pattern is present in all tropical African species. Chromosome counts from European material shows the somatic numbers between 6 and 40. The four tropical African species fall well within the range of variation. All tropical African material was earlier classified as C. stagnalis Scop., but recent taxonomic work has shown that several species can be recognized based on morphological, pollen and chromosomal differences. 1. Fruit distinctly winged
2
Fruit not winged 2. Wings symmetrical around the mericarp; petiole mostly 3-veined Wings asymmetric; petiole 1-veined at base 3. Larger leaves 10–16 mm long, 5 (−8)-veined; petiole 1(−3) -veined Larger leaves usually 6–8 mm long, usually 3-veined; petiole 1-veined
3 1. C. oreophila 2. C. anisoptera 3. C. keniensis 4. C. vulcanicola
1. C. oreophila Schotsman in B.J.B.B. 55:222, t. 7 (1985); Hedberg & Hedberg in Fl. Ethiopia 2, 1:427, fig. 52/1–8 (2000). Type: Rwanda, Gitovu, Bequet 780 (BR!, holo.) Stem 10–35 cm long in water or 3–20 cm long on mud. Leaves submerged, floating or aerial, elliptic to spathulate, up to 11 mm long and 5 mm wide but often much smaller, 3– 7-veined; petiole usually 3-veined. Flowers solitary, in the axils of leaves; bracteoles 2, falcate; stamens at dehiscence 1–2 mm, after dehiscence up to 7 mm long; anthers 0.5– 0.6 mm long; styles up to 2 mm long, erect or spreading. Fruit suborbicular, 1.6–1.8 mm long and 1.6–2 mm wide, with a distinct, symmetrically winged mericarp. Pollen reticulum not as wide as the clavae; clavae irregularly triangular, surface uneven. Fig. 1/1–4 (p. 3). UGANDA. Toro District: E slope of Ruwenzori, 14 Apr. 1948 Hedberg 802!; Kigezi District: Bukinda, Feb. 1948, Purseglove 2580! & ?Narosanje, Apr. 1945, Purseglove 1653! KENYA. Aberdare, Kinangop, Chandler 2322!; Nanyuki District: West Mt Kenya Forest Station, 26 Dec. 1921, Fries & Fries 319!; Narok District: Enesambulai Valley, 15 Aug. 1970, Greenway & Kanuri 14557! TANZANIA. Moshi District: Kilimanjaro, above Marangu, Feb. 1894, Volkens 1834!; Lushoto District: W Usambaras, 21 June 1953, Drummond & Hemsley 2968!; Mbeya District: Mbeya Mt, 13 Dec. 1962, Richards 17044! DISTR. U 2; K 1, 3, 4, 6; T 2, 3, 7; W Tropical Africa, Cameroon, Congo (Kinshasa), Rwanda, Burundi, Sudan, Ethiopia HAB. In stagnant or running water in ponds or small streams and on mud or moist soil; 1150–3250 m SYN. [C. stagnalis sensu auctt. mult. e.g. De Wild., Pl. Bequaert. 2:58 (1923) p.p.; Sam. in Notizbl. Bot. Gart. Berlin 85:322 (1925) p.p.; W.Robyns, Fl. Spermat. Pare Nat. Albert 1:484 (1948) p.p.; A.V.P.: 127 (1957) p.p.; A.Robyns, F.C.B. 7:351 (1958) p.p.; U.K.W.F.
Callitriche
3
ed. 2:82 (1994) p.p., non Scop.]
2. C. anisoptera Schotsman in Bull. Mus. Hist. Nat. Paris, sér. 4, 10:7 (1988). Type: Uganda, Toro District, Ruwenzori, Bujuku Valley near Bigo camp, 3400 m, Hedberg 398 (BR!, holo., BM, EA, K!, S!, UPS!, iso.) Stem 5–25 cm long, prostrate, mat-forming. Leaves spathulate to elliptic, 4–6 mm long and 2–2.5 mm wide, 3–5-veined; petiole 1-veined. Flowers solitary, the male with 2 bracteoles; stamens 2–6 mm long, anthers ± 0.5 mm long; style 1.5–3.5 mm long. Fruit ± 1.3–1.5 mm long and 1.6–1.8 mm wide, variously and unequally winged with best developed wings in the distal part. Pollen reticulum meshes more than twice as wide as the clavae; clavae with a globular surface. Fig. 1/5–6 (p. 3). UGANDA. Toro District: Ruwenzori, Bujuki Valley, March 1948, Hedberg 398! & Mobuku Valley, July 1952, Ross 537! & Ruwenzori, June-July 1968, Hamilton 739! KENYA. Nanyuki District: Mt Kenya, Jan. 1971, Hedberg 4853! DISTR. U 2; K 4; Congo (Kinshasa) HAB. Along paths and small streams, on mud in bogs; 2900–4000 m SYN. [C. stagnalis sensu Hedberg, A.V.P.: 127 (1957) p.p.; A.Robyns, F.C.B. 7:351 (1958) p.p.; U.K.W.F. ed. 2:82 (1994), non Scop.]
3. C. keniensis Schotsman in Bull. Mus. Hist. Nat. Paris, sér. 4, 10:10 (1988). Type: Kenya, Turkana District, Murua Nysigar Peak, Paolo 977 (BR!, holo., EA, K!, PRE, iso.) Robust aquatic herb, with stems up to 40 cm long. Larger leaves spathulate to elliptic, up to 15 mm long and 4 mm wide, 5–8-veined; petiole 1 (−3)-veined. Flowers either solitary or one male and one female in the same axil; bracts 2, about 1.5 mm long; stamens 3–4 mm long, after dehiscence up to 5 mm long; anthers 0.8 mm long. Fruit ± orbicular, ± 1.5 mm, unwinged. Pollen reticulum meshes not as wide as the clavae; clavae irregularly triangular with sharp angles. Fig. 1/7–8 (p. 3). UGANDA. Mbale District: Elgon, Sasa trail in crater, Oct. 1996, Wesche 116! KENYA. Turkana District: Murua Nysigar Peak, Sept. 1963, Paulo 977!; Mt Elgon, in the crater, May 1948, Hedberg 939!; Mt Kenya, Teleki Valley, July 1948, Hedberg 1763! DISTR. U 3; K 2–4; not found elsewhere HAB. In small ponds and streams; 2150–4250 m SYN. [C. stagnalis sensu Hedberg, A.V.P.: 126 (1957) p.p.; U.K.W.F. ed. 2:82 (1994), non Scop.]
Flora of tropical east africa callitrichaceae
4
FIG. 1. CALLITRICHE OREOPHILA—1, habit; 2, leaf; 3, node with leaves and stamen; 4, fruit. C. ANISOPTERA—5, leaf; 6, fruit. C. KENIENSIS—7, leaf; 8, fruit. C. VULCANICOLA—9, leaf; 10, fruit. 1– 2 from Hedberg 802, 3 from Polhill
Callitriche
5
437, 4 from Bally 9879; 5–6 from Mabberley 452 and after Schotsman; 7 from Townsend 2281, 8 from Wesche 116 and after Schotsman; 9–10 from Hedberg 604 and after Schotsman. Drawn by Henk Beentje. 4. C. vulcanicola Schotsman in B.J.B.B. 55:294 (1985). Type: Kenya, Nanyuki District, Mt Kenya, W slope along the Burguret track, Hedberg 4403 (UPS!, holo., BR, EA, K!, iso.) Prostrate and often mat-forming, sometimes submerged herb, with stems 5–15 cm long. Leaves spathulate or elliptic, up to 8 (−10) mm long and 2–5 mm wide, usually 3veined; petiole 1.5–3 mm long, 1-veined. Flowers solitary, bracteoles 1–2, ± 2 mm long, narrow; stamens 4–8 mm long, anthers ± 0.5 mm long; styles up to 1.5 mm long. Fruit 1.2–1.5 mm long and 1.6–2 mm wide, unwinged, but with distinctly edged margins. Pollen reticulum meshes as wide as the clavae; clavae obtusely triangular, surface almost smooth. Chromosome number: 2n=18 (counted on Hedberg 435, from Aberdare Mts and Hedberg 4403 from Mt Kenya (Hedberg & Hedberg 1988:16)). Fig. 1/9–10 (p. 3). UGANDA. Kigezi District: Muhavura, Oct. 1948, Hedberg 2117! & Mgahinga, Nov. 1954, Stauffer 789!; Toro District: Ruwenzori, Mijusi Valley, Mar. 1948, Hedberg 604! KENYA. Aberdare Range, Kinangop, July 1948, Hedberg 1650!; Kiambu District: near junction of S Kinangop and Thiba roads, Apr. 1978, Gilbert & Thulin 1047!; Nanyuki District: Mt Kenya, Nov. 1967, Hedberg 4403! DISTR. U 2; K 3, 4; Congo (Kinshasa) HAB. Moist depressions in bog and grassland, along streams; 3000–4050 m SYN. [C. stagnalis sensu Hedberg, A.V.P.: 126 (1957) p.p.; A.Robyns, F.C.B. 7:351 (1958) p.p.; U.K.W.F. ed. 2:82 (1994), non Scop.]
Callitriche deflexa Hegelm., collected from Tanzania: Kilimanjaro, Layamungu Coffee Research Station, 4200 ft., M.Bigger 1236, is most likely an introduction. The species is native to South America and has been recorded from Portugal, Morocco and southern Africa, usually associated with cultivated areas (see Schotsman, op. cit.). It differs from the African species in being very small, with small leaves (2–4 mm long) and small fruits (0.5–0.7 mm).
INDEX TO CALLITRICHACEAE
Callitriche L., 1 Callitriche anisoptera Schotsman, 2 Callitriche deflexa Hegelm., 4 Callitriche keniensis Schotsman, 2 Callitriche oreophila Schotsman, 2 Callitriche stagnalis Scop., 2, 4 Callitriche stagnalis sensu auctt., 2, 4 Callitriche vulcanicola Schotsman, 4
No new names validated in this part
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