zy zyxw zyxwvut zyxw Taxonomic studies of the genus Hypoxis in East Africa Inger Nordal, Morten M. Laane, Ellen Holt and Inger Staubo Nordal, I., Laane, M. M., Holt, E. & Staubo, I. 1985. Taxonomic studies of the genus Hypoxis in East Africa. - Nord. J. Bot. 5: 15-30. Copenhagen. ISSN 0107- 055X. East African material of the genus Hypoxis L. has preliminarily been divided into the heterogenous, probably apomictic H . obtusa Burch- complex (2n = 4C50, ca. 75,76, ca. 85, >86, ca. 92, ca. 98, ca. 108, 13Cb135, 16C200) and 5 rather homogenous species: H . angustifolia Lam. (2n = 14,28), H. goetzei Harms (2x1 = ca. 62), H. kilimanjarica Bak., H . malosana Bak. (2n = 14) and H . macrocarpa Holt & Staubo sp. nov. H . kiiimanjarica is divided into ssp. kiiimanjarica and ssp. prostrata Holt & Staubo ssp. nov. zyxwvut I. Nordal, M . M . Laane, E. Holt and I . Staubo, Botanical Institute, Univ. of Oslo, P. 0 . Box 1045, Blindern, Oslo 3, Norway. Introduction The pantropical genus Hypoxis L. presents substantial taxonomic problems both regarding generic and species delimitation. Linnaeus (1759) established the genus based on two species, of which the American H . erectus L. (nomen illegit., = Ornithogalum hirsutum L., Hypoxis hirsutus (L.) Colville) has been selected as type species for the genus (vide Britton & Brown 1913). Later he (Linnaeus 1781) described eight African Hypoxis species, among them H . villosa L. The remaining seven have, however, later been removed from the genus. During the next hundred years several new species were described so that Flora Capensis (Baker 1896) included 43 species and the Flora of Tropical Africa (Baker 1898) included 17 species. The next large contribution was made by Nel(l914). He monographed the genus, reported 83 species from Africa and published a total of 45 new species. Nel applied a set of characters for specific delimitation that were very different from Baker’s. The two classifications was never unified and both are still in use, being applied by different herbaria. Later revisions in connection with different regional flora projects in Africa have all led to a reduction of what appears to be a too large number of species (Andrews 1956, Hepper 1968, Geerinck 1971). Agnew (1974) recognized 5 species in Upland Kenya: H . angustifolia Lam., H . kili- manjarica Bak., H . multijlora Nel., H . obtusa Burch., and H . villosa L. Recently valuable new information has been published in two M. Sc. theses: by Wood (1976) on material from Natal and by Heideman (1979, 1983) on material from Witwatersrand in Transvaal. Lindley (1836) referred Hypoxis to tribus Hypoxideae of Amaryllidaceae, a delimitation later kept e.g. by Bentham (1883), Pax (1889), Nel(1914), and Phillips (1926). Pax & Hoffmann (1930) raised the tribe to subfamily level. Later, a separate family, Hypoxidaceae, has been generally accepted (Hutchinson 1934, Melchior 1964, Geerinck 1968, 1969, Thompson 1976, and Hilliard & Burtt 1978). The generic concept within Hypoxidaceae has varied, and the problem does not appear to be settled. The genus Hypoxis is unambigously characterized within the family by having hairy flowers with 6 stamens, trilocular ovaries without a beak, and free perianth segments. In East Africa only one other genus is represented, namely Curculigo Gaertn., having distinctly beaked ovaries. Hypoxis is, in addition to its widespread distribution in Africa south of Sahara, widely distributed in America, South East Asia and Australia. The genus seems to have a center of variation in South Africa, occurring typically in open, undisturbed grasslands. The cytology of the genus is complex. Wilsenach (1967), Wilsenach & Papenfuss (1967) and Wilsenach & 0 NORDIC JOURNAL OF BOTANY NORD. J. BOT. 5 : 15-u), TROP 092 Nord. J. Bot. 5 (1) 1985 15 z zyxwvutsrqp zyxwvutsr Warren (1967) examined material from South Africa. They suggested x = 8 to be the basic number for some species (H. stellipes 2x, H . cf. zeyheri 2x, H. longifolia 9x, and H . rooperi 12x) and x = 7 for one (H . filiformis 2x). H . acuminata with n = 18 and H. multiceps with 2n = 36 could, however, also indicate a basic number of x = 9. H. rooperi appears to be very variable, in addition to Wilsenach’s 2n = 96, Fernandes & Neves (1962) have reported 2n = 76 and 114, proposing the basic number to be x = 19. Naranjo (1975) has reported H . pusillu from Australia with 2n = 28 and H. decumbens from South America with 2n = 42, both indicating a basic number of x = 7 which thus occurs pantropically. Mehra & Sachdeva (1976) have examined H. aurea from Himalaya which had 2n = 54 and a basic number of x = 6 was suggested. Wilsenach & Warren (1967) concluded that although apomixis has not been proved, everything points to it: “The plants are very polymorphic, there is a great variation in chromosome number, meiosis of pollen mother cells shows characteristic abnormalities in the high chromosome number species, the pollen grains possess different chromosome numbers in suspected apomictic plants, megaspores (or megaspore mother cells) degenerate and twin embryo-sacs are common.” The aim of this work is mainly to survey the morphological variation in order to reach a practical method in species delimitation for the publication of Flora of Tropical East Africa. Whenever possible, we have tried to analyze the cytology and also to procure more data for the discussion of the occurrence of apomixis. Material and methods Morphology In the examined East African material the corm is subglobose to elongated, most often vertical, but sometimes horizontal. In some cases branched corms or corms giving rise to more than one shoot, have been observed. The adventitious roots are fleshy, distinctly contractile, and arranged in a ring, most often above the middle of the corm. Root scars from older roots can be seen below. In a longitudinal section two clearly separated areas can be distinguished: the inner core with vascular bundles encased in fibrous tissue, and the spongy outer part, the storage area, transversed by numerous mucilage canals. The latter contains whitish or yellowish sap, which rapidly darkens when oxidized. The size and shape of the corms seem to vary more or less continuously, from 1 to 9 cm long and from 1 to 8 cm broad, within the examined material. The leaves arise in the apical region of the corm in a phyllotaxis of 1/3. The basal rosette is often surrounded by remnants of vascular strands from older leaves. The leaves have continuous seasonal growth due to intercalary meristems in their bases. This feature will result in a very variable shape and size dependent on stage. Leaves are 5-90 cm long and 0.2-4.0 (-7.0) cm wide. They are linear to lanceolate or ovate, and may be prostrate to erect. They are always keeled, and will in cross-section assume the shape of a V, U or an inverted, flattened W. The number of longitudinal nerves varies from 3 to about 50, depending on leaf width. The leaves are always hairy. The hairs are either double, then often having one arm longer than the other, or in bundles arising from a short common foot cell. We shall denote the latter type stellate. Single hairs are very rare, and glandular hairs never occur. The hairs vary in length, thickness, stiffness, and orientation compared to the lamina. They may be whitish, greyish, yellowish, or red-brown; their colour may vary with stage. The indumentum may be concentrated along the margin andor midrib, or spread all over the surfaces. It is con- zyxwvu zyxwv zyxwvu This is mainly based on East African herbarium material from the following herbaria: B, BM, BR, EA, G, K , L, LISC, M, 0, P, PRE, S, UPS, and WAG. Living material collected by I. Nordal (= I. Bjsrnstad) and A. Bjarnstad has been studied during cultivation in Oslo (referred to in the following as I.N., I.B., and A.B., cfr. Tab. 1). Flowers from herbarium specimens have been boiled in water for morphological analysis. In addition seeds and pollen were examined in a Scanning Electron Microscope in dry conditions after coating with gold-palladium. Pollen grains were stained with cotton-blue solution and pollen stainability was used as an estimate of pollen fertility. In crossing experiments anthers were removed at an early stage, and flowers were isolated by thin clothbags . The fairly small chromosome size in Hypoxis requires special cytological techniques in order to obtain accurate information. Root-tips from cultivated material were fixed in sublimate-picric acid solution as recommended by Qstergren and HenCen (1962). The squashed, Feulgen-stained preparations were placed on a Peltier freezing device, the cover-glass removed by split16 ting-off with a scalpel, then immediately immersed in. pure ethanol. Finally they were embedded in Euparal. Root-tips were, in addition, fixed in Muntzing’s modification of the Navashin-Karpechenko solution, embedded in paraffin and cut on a Minot-type microtome into sections 14 pm thick. The slides were stained in hematoxylin (Laane 1970 p. 70) and embedded in Eukitt. Meiosis was studied by dissecting small buds from specimens that had just started germinating in spring. Developing anthers were fixed in acetic-ethanol 1:3 parts by volume, then the PMCs were dissected and stained in acetic-orcein and squashed. All chromosome drawings are based on tracings from microphotographs. Some of those from high-numbered specimens are reconstructed from a number of photographs at different focal levels. zy zy Nord. J. Bot. 5 (1) 1985 zyxwvutsrqpo zyxwvut zyxw zyxw Fig. 1. Abaxial leaf surfaces showing the indumentum types in Hypoxis A: H. malosuna (Richards 11617, B). B: H. goetzei (Goetze 1416, B). C: H. obtusa-complex (I. Bjernstad 587,O). D: H. obtusa-complex (Perdue & Kibuwa 11469, EA). E) H. obtusu-complex (Schlieben 3506, B). F: H. obtusa-complex (Schultze 238, B). All X 7,5. venient to classify the indumentum types according to the following key (Fig. 1): 1. Hairs stellate, along the margin and abaxial midrib only 2 1. Hairs double or stellate, covering one or both surfaces.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2 . Hairs not appressed ............................... 3 2 . Hairs appressed parallel to the margidmidrib. . . . type D 3. Hairs red-brown, thick, densely clustered. . . . . . . . type B 3. Hairs greyish, not specially thick nor densely clustered . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . type E 4. Stellate hairs along margidmidrib, double hairs of varying density on one or both surfaces . . . . . . . . . . . . . . . . type C 4. Hairs along margin not differing from those on surface 5 5. Varying density, never tomentose . . . . . . . . . . . . . . type A 5. Surfaces tomentose ........................... type F the typical raceme and the corymb (Fig. 2). There is correlation between number of flowers and type of inflorescences, few-flowered are often corymbs, manyflowered are racemes. An acropetal flowering sequence is most common, but a basipetal sequence is found in some taxa (e.g. Fig. 2 D). The axis is often stepwise, not gradual, narrowing towards the apex. The bracts of the upper flowers sometimes appear adnate. The flowers are most often opposite, but sometimes whorls of three or single flowers are intermixed between the pairs (Fig. 2 E). Pedicel length varies from 0.2 to 3.4 cm. The number of flowers per inflorescence varies from 1 to 22, single-flowered being rare, and 2-6 the most common states. The bracts are generally subulate, hairy abaxially and with semitransparent margin. The perianth segments are free, patent and subequal; 5-18 mm long and 2-9 mm broad, the outer often somewhat narrower, longer and more acute than the inner zyxwvu The scape is more or less hairy, and variable in stiffness and robustness. Total length 2-35 (-60) cm. Inflorescence types include all intermediates between zyxwvutsrqponml 2 Nord. I . Bot. 5 (1) 1985 17 d 2 zyx zyxwvutsrqpo zyxwvut zy Fig. 2. Semischematic presentation of types of inflorescence in Hypoxis. Numbers denote flowering sequence. a x : H. angustifolia. d: H. goetzei. e-h: H. obtusa-complex. i-j: H. malosana. ones. Adaxially the segments are plain yellow, abaxially the outer are light green with dense indumentum, the inner yellow and only sparsely hairy along the green, rarely red-brown, dorsal midrib. The perianth persists in an erect position in the fruiting stage. The 6 stamens are inserted at the base of the perianth segments. The filaments are filiform to subulate, short; the outer may be somewhat longer than the inner. The anthers have latrorse dehiscence and are sagittate with the filaments fixed in the cleft. In old stages they may become more or less versatile. The thecae are fused apically (Fig. 3 A-C) or they form a split (Fig. 3 D). Transitional forms occur. Pollen grains of East African Hypoxis spp. are homogenous, ovale, prolate and monocolpate. The exine pattern is semitectate and per-reticulate (Fzgri & Iver- a zyxwvu zyxwv 18 zy Nord. J . Bot. 5 (1) 1985 zyxwv zy zy Fig. 4. Seeds. A: Hypoxis obrusa-complex (I. Bjdrnstad 262, 0),Bm: H. angustifolia (A. Bjdrnstad 2076, 0).All X 27. three lobes joined more or less longitudinally, with receptive surfaces along the sutures and covered by glandular papillae (Fig. 3 E-K). The stigma may be shorter, equally long or longer than the style, the ratio style to stigma length varying gradually from 0.2 to about 2.5 in the examined material. There is a considerable intraspecific variation (cf. Fig. E-J). The capsules are of two different types. The first is a 2-10 mm long pyxis, where the lid, crowned with per- A zyxwvutsr B zyxwvutsrqpo zy Fig. 5. Testa structure. A: Hypoxis obrusa-complex(I. Bjdrnstad 861, 0).B: H. obrusu-complex (Thorold 2752, K). C: H . mafosana (I. Bj0rnstad 814,O). D: H . obrusa-complex(Leedal5202, K). E: H. angustifoh (A. Bjdrnstad 2076,O). F: H . macrocarpa (Bullock 2057, K). All X 200. 2' Nord. J. Bot. 5 (1) 1985 sisting tepals, comes off in circumscissile dehiscence. The second is a 3-15 (-30) mm long loculicide capsule. There are up to about 50 seeds per fruit. The seeds are subglobose with black or dark brown, very hard testa. Hilum and raphe are distinct, and the micropyle protrudes (Fig. 4). The testa is more or less papillate. The testa structure divides the material into two groups: One with a special folded cuticula coating on the papillae (Fig. 5 E-F), the other lacking this feature (Fig. 5 A-D). Those without the coating may have a more or less distinct secondary structure, making the papillae verrucose (Fig. 5 D). Macroscopically the seeds with the folded cuticula look dull and dark brown, the seeds without the folded structures appear black and often glossy. 36 zyxwvu zyxwv -I 1 zyxwvutsrq zyxwvu zyxw zyxwvuts zyxwv zyxwvutsrq zyxwvutsr / 0 0 0 0 0 0 fm 20 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Morphological delimitation of taxa Grouping of the material according to the two basic works on tropical African Hypoxis of Baker (1898) and Nel (1914) turned out to be impossible. Baker applied mainly quantitative characters which in our material all vary continuously (leaf width, peduncle length, bract size, inflorescence shape). Nel placed special emphasis on the following characters: Shape of anther apex, number of nerves per leaf, leaf shape, ratio style length to stigma length, and inflorescence shape. Except for the anther apex, which is in part a useful character, Nel's characters also show continuous variation. Neither Baker nor Nel used seed and fruit characters, which we have found to give a more distinct basis for delimitation. The testa structure with folded coating on the papillae (Fig. 5 E-F) is very special and complex and justifies a primary division of the material in two groups. Within the group with the special folded seed coating one subgroup has leaves 1.8-7.0 cm broad, another 0.10.8 cm broad. Except for two collections (Bullock 2045 and Davies 742) which will be discussed separately under H. sp. A., the broad-leaved plants are fairly homogeneous. leaves are lanceolate to ovate, with indumentum of type B (Fig. 1 B). Inflorescence is racemeshaped and seems to have a basipetal flowering sequence (Fig. 2 D). The fruit is a pyxis, and the style is always longer than the stigma. The oldest available name for this taxon is H. goetzei Harms. The narrow-leaved subgroup has linear leaves with indumentum of Type A (Fig. 1A). The character of anther apex again divides the subgroup into two: those with a split and those without. The former have on average narrower fruits and longer pedicels compared with the latter (Fig. 6). The plant with apical anther splits have more or less corymbose inflorescences (Fig. 2 A X ) , since the lower pedicels are the longest, ( 0 . 5 ) 1.0-3.2 cm long. The capsule wall is thin, closely fitting the seeds. The opening is loculicidal. Ratio style length to stigma length is very variable, either may be the longer (Fig. 5 E-J). 20 12 16/ 0 0 0 0 0 0 0 / 0 O 0 ;0 4- * * I 2 ' " I " 4 ' 0 I ' . . I . 6 . 8 ' I . . ' I . 10 Capsule width m m Fig. 6. The relation between length of lower pedicel and width of capsule within the group of Hypoxis with folded testa structure. The filled circles represent plants with thecae apically fused. The upper left group H. angustifoliu, the lower right, H. macrocarpa. The type specimen of H. angustifoliu Lam. agrees in all examined characters. The plants with the apically fused thecae have shorter pedicles, the lower only 0.1-0.8 cm long, and the capsules are somewhat broader than in H. ungustifofiu. The opening mechanism is unfortunately not known. There are 1-2 flowers per inflorescence, and the style is always shorter than the stigma. This taxon will be described here as H. macrocarpa Holt & Staubo. The second main group which lacks the special cuticula folding on the testa is large with complex variation patterns. Possible correlations between all characters used by Baker (1898) and Nel (1914) for delimitation have been studied by means of scatter plots. When all material was included, it was impossible to separate subgroups at all. When the material from central and northern parts of the area was analysed separately from material of the southern parts, some clustering trends were obtained (Figs 7, 8). In the montane areas of Kenya and Tanzania occurs a group with shorter perianth segments and narrower leaves than the others (Fig. 7). Within this group a further delimitation is possible. The plants growing at the highest altitudes, Nord. J . Bot. 5 (1) 1985 16 0 0 0 0 0 0 0 0 0 0 0 0 0 0 000 0 0 0 0 0 0 0 0 0 0 4 zy zyxwvuts zyxwvutsr zyxwvutsrq 0 000 0 0 0 000 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 000 timates here are to be regarded as speculative. The H. obtusa-complex with its extensive morphological variation appears to present a corresponding cytological variability. The following approximate chromosome numbers have been counted in mitosis: 40-50, 75, 85 (Fig. 9 D), >86, 92 (Fig. 9 F), 98, 108, 130-135, 160200. On the basis of present information it seems impossible to propose any reasonable basic number. We were able to study meiosis in two representatives of the H. obtusa-complex. One specimen (I.B. 826) has a regular meiotic pattern (Fig. 10 A-J), the other (I.B. 861) was meiotically highly irregular (Fig. 11 A-F). During the first regular meiotic prophase the chromosomes associate in 38 bivalents. In late diakinesis or in the first meiotic metaphase these bivalents are with few exceptions associated with two chiasmata. Failure to pair is rare and multivalent associations were not found in spite of the high chromosome number (Fig. 10 A-C) . zyxwvutsrqp zyxwvutsrqpon zyxwv zyxw leaf width mm Fig. 7. The relation between width and length of perianth segments within the group of Hypoxis lacking cuticula folding on the testa from N Tanzania, Kenya and Uganda. The lower, left group H. kilimanjarica, the rest H. obfusa-complex. i.e. above 2900 m, have reflexed to prostrate leaves and a scape up to 4 cm long. Below, the leaves grow more erectly and the scapes are longer. The type specimen of H. kilirnanjarica Bak. belongs among the latter. The high altitudes taxon will be described here as H. kilimanjarica Bak. ssp. prostrata Holt & Staubo. Correspondingly, for the southern part of the material, clustering trends were best obtained when leaf width was scored against pedicel length in a scatter plot (Fig. 8). The narrow-leaved group also has subterete leaves and a characteristic inflorescence as illustrated (Fig. 2 I, J). The oldest name for this taxon is H. rnalosana Bak. The more or less broad-leaved material distributed over the whole area is too heterogeneous to be included in one species. The variation forms a reticulate pattern, which we up to now have not been able to use for further delimitation. Divisions based on one character alone should be avoided, and no distinct character correlation has been found. Since the variation within the groups by far exceeds the normal variation acceptable for a species, we will include it in what we shall denote as the H. obtusa Burch.-complex. Cytology The chromosome numbers were counted in root-tip mitoses (Tab. l). Examples of somatic metaphase plates are shown in Fig. 9. H. angustifolia has generally 2n = 14 (Fig. 9 A) (Cameroun, Uganda, Northern Tanzania). In SouthWest Tanzania we found a tetraploid with 2n = 28 (Fig. 9 B). H. rnalosana also has 2n = 14. The basic number is in both cases x = 7. In the robust taxa, H. goetzei and H. obtusa-complex obviously high ploidy levels are present. In H. goetzei a somatic chromosome number of about 62 has been found (Fig. 9 C, E). If the basic number is x = 7, this should indicate a ploidy level of about 9, but any esNord. J. Bol. 5 (1) 1985 481 "1 401 36 1'1 E 0 321 0 0 0 0 0 0 0 A 0 20 0 0 0 0 0 0 0 0 0 0 4 0 0 0 zoo 0 0 zyxwv 0 0 \ o o 0 4 0 0 0 0 O 8 0 12 16 20 24 Pedicel length mm Fig. 8. The relation between pedicel length and leaf width within the group of Hypoxis lacking cuticula folding on the testa from central and southern Tanzania. The filled circles represent H . rnalosana, the open ones represent the H. obmacomplex in the area. 21 A seen (Fig. 11 B). Aggregates of chromosomes occur at this stage near the equatorial plane in a number of cells, typically extended chromosome arms point towards each cell pole. About 25% of the cells appear to pass directly on to the first anaphase. Here, the chromosomes seldom associate in pairs or form a metaphase plate, instead the univalents become irregularly distributed in the first anaphase spindle (Fig. 11 C-D). The forming dyad- nuclei become genetically imbalanced and the chromosomes lost in the spindle appear to form one or more additional nuclei. The same pattern is also reflected in the second meiotic division. Examination of the tetrad cells shows them to contain a variable number of larger and smaller cells (Fig. 11 E). Irregular chromosome separation was sometimes also observed in mitosis of this plant (Fig. 11 F). zyxwv zy zyxwvutsrq zyxwvut zyxwvut Notes on reproduction Representative specimens of H. angustijolia and the H. obtusa-complex were isolated and tested for possible E Fig. 9. Metaphase plates from root tips of East African Hypo8%, 0),2n = 14. B: H. angustifolia (I. Bjmnstad 561, 0),2n = 28. C: H. goetzei (A. Bj~rnstad2261, 0),2n = ca. 61. D) H. obtusa-complex (I. Bj0rnstad 897, 0),2n = ca. 85. E: H. goetzei (A. Bjernstad 2261,0), 2n = ca. 63. F: H. obtusa-complex (I. Bjernstad 262, 0),2n = ca. 92. All X 1OOO. xis. A: H. angustifolia (I. Bj0rnstad Chromosome kinetics are normal at first anaphase and balanced dyad nuclei are formed. The only irregularity found at this stage is a pair of lagging univalents which occur in about 10% of the PMCs. Chromosome behaviour in second meiotic division (Fig. 10 E-I) also appeared normal except for a pair of lagging univalents (Fig. 10 G, I) also at second ana- and telophase. More than 80% of the tetrad cells contained even-sized, probably genetically balanced nuclei. Meiosis in I.B. 861 was highly irregular. U p to the pachytene stage the condensing chromosomes were similar to those in I.B. 826. At later stages most PMCs appeared abnormal. About 10% of the first metaphase nuclei did not form distinct chromosome configurations, instead the chromosomes partly decondensed and irregular restitution nuclei appeared (Fig. 11 A). In some cells (about 5%) a few bivalents were formed besides numerous scattered univalents. At first metaphase a highly variable pattern of chromosome aggregates was 22 ......... ......... Fig. 10. Regular meiosis (n = 38) in the H. obtusa-complex (I. Bjernstad 826, 0).A-C: Metaphase I with regular bivalent formation, mostly two chiasmata. D: Late telophase I. E-F: Second metaphase. G-J: Late second anaphase-telophase,laggard chromosomes seen in G-I. All X 500. zyx zy Nord. J . Bot. 5 (1) 1985 zyxwvutsrqp zyxw zyxwvut zyxwvu zyxwvu zyxwvutsrqpo Tab. 1. Material of Hypoxb in cultivation and chromosome numbers. a: Plants tested for autogamy (all positive) and for apomixis (all negative, except A.B. 554). n: Plants tested for pollen fertility (always high, 75-90%). *: Preparations at the Jodrell Laboratory, Kew. Species Voucher H. angustifolia A. B. 560 H. angustifolia H. angustifolia Habitat Themeda-Brachiaria grassland, alt, lo00 m I.B. 561a,x U2: Ankole Distr. Queen Elizabeth Nat. Park T7:Iringa Distr. Iringa I.B. 743n T3: Pangani Distr. Pan- Grassland on coral outcrop, alt. 20 m Acacia-Commiphora grassland, alt. 1800 m H. angustifolia I.B. 896a.n H. angustifolia Harris 1290 H. angustifolia I.N. 970 H. goetzei A.B. 2261 H. malosana Locality I.B. 814n gani T1: Musoma Distr. Serengeti Nat. Park, Near Lobo T6: Usaramo Distr. Dar es Salaam Cameroon: Adamoua region, Boye. T7:Mbeya Distr. Ruaha Nat. Park 77: Mufindi Distr. 12 H. obtusa-complex H. obtusa-complex Bally 12163 A.B. 554a,n H. obtusa-complex I.B. 262 H. obtusa-complex I.B. 580a,n H. obtusa-complex I.B. 8 2 5 ~ H. obtusa-complex I.B. 826 km S of Sao Hill K4: Machakos Distr. U2: Ankole Distr. Nyabushozi County, Muko Range K4: North Nyeri Distr. Naru Moru T7: Iringa Distr. Mufindi T7:Njombe Distr. 20 km SE of Njombe along the road to Songea as I.B. 825 H. obtusa-complex I.B. 861 T7:Njombe Distr. 13 H. obtusa-complex I.B. 897 H. obtusa-complex (“H. urceolata”) H. obtusa-complex (“H. urceolata”) H. obtusa-complex (“H. urceolata”) H. obtusa-complex (“H. urceolata”) Kew 441:58 (cyt. 62/604) Kew 41554 (cyt. 62J713) Kew 52353 (cyt. 62/1401) Kew 236:62 (cyt. 62/999) km W of Njombe T1: Musoma Distr. Serengeti Nat. Park, Near Lobo. autogamy (Tab. 1). All were able to produce seeds after self-pollination. It was often observed that the anthers in many plants approached the stigma in late anthesis. The same plants were also tested for apomixis, by early demasculation and isolation. Except for o n e case (A. B. 554 from Uganda) n o seeds were produced. The latter did once produce ripe and fully-developed seeds without any pollen present, i.e. apomixis is proved at least in one case. Apomixis is, however, not excluded in the other test cases, as pollen might be required for endosperm production (pseudogami). T h e pollen stainability was in all cases high, i.e. from Degraded miombo 2n 2n = 14 2n = 28 (Fig. 9B) 2n = 14 2n = 14 (Fig. 9a) 2n = 14 Savanna, alt. 940 m Brachystegia-Julbernardia-Pterocarpus Woodland XyrisiPycreus macranthus bog, alt. 1770 m Alt. 1750 m Grassland with Acacia hockii and Themeda triandra, grazed, alt. 1380 Alt. 2000 m 2n = 14 2n = ca. 62 (Fig. 9c, e) 2n = 14 2n = ca. 75* 2n > 86 2n = ca. 92 (Fig. 9f) Alt. 1800-1950 m More or less disturbed 2n = 4G50 grassland along road alt. 1750 m As I.B. 825 n = 38, regular meiosis Grassland alt. 2010 m irregular meiosis Acacia-Commiphora grassland alt. 1800 m zy 2n = 85 (Fig. 9d) 2n = 130-135* 2n = ca. 98* 2n = ca. 108* 2n = 160-200* Preliminary key to the taxa 1. Seed papillae with folded coating (Figs 4B, 5E-F), surface macroscopically dull, brownish ...................... 2 1. Seed papillae without folded coating (Figs 4A, 5A-D), surface macroscopically more or less glossy, black ....... 5 2. Leaves 1%70 mm broad; indumentum variable; flowers 6 3 or more in araceme ............................... 2. Leaves 1-8 mm broad, indumentum of scattered double hairs, rather sparse; flowers 1 4 , in a corymb if more than one. ............................................. 4 3. Lamina glabrous, red-brown stellate hairs along margins and abaxially on midrib.. . . . . . . . . . . . . . . . . 2. H. goetzei 3. Indumentum different . . . . . . . . . . . . . . . . . . . 7. H. sp. A. 4. Leaves lax; flowers 1-6; lower pedicels longer than 10 mm; anther apex with a split; capsule narrow, slender, 2-4 mm broad.. ............................ 1. H. angustifolia zyxwvutsr zyxwvutsrqponmlk 75 to 90%. Nord. J . Bot. 5 (1) 1985 23 exceeding the scape considerably, width (1-) 2-6 (-8) mm. Indumentum sparse with greyish, rarely golden, double hairs (Fig. 1A). Scape 1.5-23 cm long. Inflorescence a corymb (Fig. 2A-C) with 1-5 (-6) flowers. Lower pedicel ( 0 3 ) 1-3.2 cm long. Indumentum of the inflorescence mostly yellowish. Bracts 0.4-1.3 cm long and narrow. Flowers relatively small, diameter 1-2 cm; perianth segments 3.5-9 (-11) mm long and 1.5-3 (-4) mm wide, sometimes with a reddish dorsal line. Filaments filiform 1-4.5 mm long with anthers 1.5-3 (-4.5) mm long, more or less versatile. Thecae apically not fully fused (Fig. 3D). Stigma 0.8-2.6 mm long, most often a little longer than the style (Fig. 3 E-J). Capsule loculicide, 3-15 mm long and 2-4 mm broad. Seeds up to 1.6 mm long. Testa strongly papillate with characteristic cuticula foldings (Figs 4B, 5E). Distribution and habitat. H . angustifolia is distributed throughout East Africa (Fig. 12) and also occurs widely in the rest of Tropical Africa from Guinea to Mauritius, but seems rare in South Africa. It grows in grasslands and open woodland on sand, laterite or clay and has been collected from sea level up to 3000 m. zyxwvut zyxwvuts zyxwvuts zyxwvut zyxwvutsrqp zyxwvutsrqp zyxwvutsr zyxwvu zyxwvutsrq Fig. 11. Irregular meiosis in the H. obtusa-complex (I. Bjarnstad 861, 0).A: Restitution nucleus corresponding to first metaphase. B: First metaphase nucleus showing large chromosome aggregates and a few bivalents. C-D: First anaphases with numerous laggard chromosomes. E: Tetrade cell showing cells formed with nuclei of various sizes. F: Mitotic cell showing lagging chromosomes. All x 500. 4. Leaves erect; flowers 1-2; lower pedicels less than 7 mm; anther apex without a split; capsule 4-6 mm 3. H. macrocarpa broad.. ............................ 5. Leaves less than 3.5 mm wide, with double (rarely triple) hairs ............................................ 6 5. Leaves 3.5-50 mm wide with stellate (rarely only double hairs) along margins and abaxially on midr i b . . ........................... 6. H. obtusa-complex 6. Lower pedicel 0 4 mm long; plants of SW Tanzania to SW Uganda.. ............................ 5. H. malosana 6. Lower pedicel 4 1 0 mm long; mountain plants in Kenya and N Tanzania.. .................. 4. H. kilimanjarica 1. Hypoxis angustifolia Lam. Lamarck 1789: 182. -Type: Commerson s.n., Mauritius (P,holotype). H. luzuloides Robyns & Tournay 1955: 254. - Type: de Witte 1130, Zaire, Tshambi (BR, holotype). Slender plants with a subglobose to cylindrical, most often vertical corm, 0.5-4 cm long with diameter 0.5-2.5 cm. Leaves linear, grasslike, erect or reflexed, length very variable depending on stage, 3 4 5 (-70) cm often 24 Fig. 12. Distribution of H. angustijolia in East Africa. Nord. J . Bol. 5 (1) 1985 zyxwvutsrqp zyx zyxwvutsr zyxwvuts zyxwvu Note. H. angustifolia is a "good" species established at the diploid (rarely tetraploid) level. No indications of apomixis. I"'" ' '.' A , ' I " - ..-.. Selected col1ections.A total of 323 specimens from 202 collections was studied. Uganda: Kerfoot 4958 (EA); Jarrett 55 (EA, K); Wood 354 (EA, K); Lye & Rwaburindore 4440 (EA, K, UPS). Kenya: Gillett 12799 (B, BM, BR, EA, K, P, PRE, S); Napier 1907 (EA); Maas Geesteranus 4573 (BR, K, L, PRE); Glover, Gwynne & Samuel 617 (EA, K); Maas Geesteranus 5220 (BR, G, K, L, PRE, S); Glover, Gwynne & Samuel 122.5 (EA, K, PRE); Magogo & Glover 498 (BR, EA, K, PRE). Tanzania: Greenway & Turner 12007 (EA, K, PRE); Polhill & Paulo 2373 (B, BR, EA, K, LISC, PRE); Drummond & Hemsley 3335 (B, BR, EA, K, LISC); Hornby 2096 (EA, K); Schlieben 324.5 (B, BR, K, LISC, PRE); A. Bj0rnstad 2076 (EA, K, 0); Milne-Redhead & Taylor 7724 (EA, K); Faulkner 2940 (BR, K); Greenway 1441 (EA, K). 2. Hypoxis goetzei Harms Harms 1901: 276. - Type: Goetze 1416, Tanzania, Unyika, Toola (B, holotype; BR, EA, G, L, isotypes). H. esculenta de Wildemann 1913: 537. - Type: Hock s.n., Zaire, Upper Katanga, Dekahou Valley (BR, holotype). H. rubiginosa Nel 1914: 320. - Type: Busse 947, Tanzania, Mgaka Valley (B, holotype). Robust plants with corm 3-15 cm long and diameter 3-10 cm. Leaves lanceolate to ovate, stiff and erect, 1.8-7 cm broad, during anthesis rather short, elongating considerably during growing season; lamina glabrous except for distinct indumentum of orange to red-brown hairs along the margins and abaxial midrib (Fig. IB). Scape 5-15 cm long. Inflorescence a raceme with 4-10 (-14) flowers; flowering seems to proceed basipetally (Fig. 2D); indumentum golden yellowish, rarely greyish. Pedicels 1-7 mm long; bracts 8-20 mm long, 2-4 mm broad. Flowers fairly large, diameter 2.2-3.5 cm; perianth segments 11-17 mm long, 4 . 5 4 mm broad. Filaments subulate, 3-4 mm long with anthers 6-9 mm; thecae apically fused. Style 2.5-5 mm long, i.e. longer than the stigma which never exceeds 2 mm. Fruit a pyxis, 5-9 mm long, 4-6 mm broad. Seeds 1.4-1.7 mm long. Testa structure like in H. ungurfifoliu, papillate, with distinct cuticula folding. Fig. 13. Distribution of H. goetzei in East Africa. zyxwvutsr Distribution and habitat. The main distribution of H. goetzei is in the Southern Highlands of Tanzania (Fig. 13) and adjacent areas of Zambia and Zaire (Katanga). A few specimens from SW Kenya must also be referred to the species. It grows in grasslands or open woodlands from 900 to 2600 m above sea level. It seems to prefer area which are often burnt. Note. The name " H . goetzei" has been used in a much wider sense in herbaria and literature (e.g. Binns 1%8), including also plants with tomentose leaves (cf. Fig. 1F). These lack, however, the special cuticula folding of the testa papillae, and will be referred here to the H . obtusa-complex. H. esculenta De Wild. was wrongly reduced to a synonym of H. subspicata Pax by Geerinck (1971). Its type is in full accordance with H. goetzei. Nel (1914) delimitated H. rubiginosa Nel from H . goetzei by the number of flowers, 4 in the former, which is within the normal variation range for H . goetzei. Selected collections. A total of 30 specimens from 18 collections was studied. Kenya: Symes 285 (EA, K). Tanzania: Richards 11701 (BR, EA, K); Napper 1075 (EA, K, UPS); Greenway & Kanuri 14764 (EA, K, PRE); Leedal 499 (EA); Milne-Redhead & Taylor 8013 (B, BR, EA, K, LISC). 3. Hypoxis macrocarpa Holt 81 Staubo sp. nov. Type: Richards 8768, Tanzania, Ufipa Distr., Sumbawanga, near Mpui (K, holotype). Habitu H. angustifoliae Lam. similis et illi speciei proxima, differt praecipue pedicellis brevioribus, capsulis latioribus et apice antherarum integro. zyxwvut zyxwvutsrqpon zyxwvutsrqponm Nord. J . Bot. 5 (1) 1985 Plants up to 40 cm high with corm 2 . 5 4 cm long and diameter 2-3 cm. Leaves linear, stifly erect, 2.5-8 mm broad; indumentum sparse, whitish hairs especially along margins and the abaxial midrib. Scape 7.5-12 cm long. Inflorescence with 1-2, rarely 3, flowers and indumentum of greyish to yellowish hairs. Lower pedicels 1 4 mm long. Bracts narrow, up to 1 cm long. Flower 25 lia it is rarely shorter than 10 mm (cf. Fig. 6). The fruits are broader in H. macrocarpa, 3.5-6 mm versus 2 4 mm. H. macrocarpa lacks the typical split anther apex of H . angustifolia, and its flowers are somewhat larger. H. macrocarpa is found in wetter habitats than H. angustifolia. Examined collections. Kenya: Agnew, Kibe & Mathenge 10588 (EA); Rayner 541 (K); Rayner 551 (K); Wiltshire 24 (K). Tanzania: Richards 11621 (K); Richards 11750 (K); Bullock 2057 (K); Richards 8768 (K, holotype); Arasululu in EAH 28737 (EA); Davies 663 (EA, K). 4. Hypoxis kilimanjarica Bak. Baker 1898: 378. -Type: Volkens 781, Tanzania, Kilimanjaro, Marangu towards Mawenzi (B, holotype). H. incisa Nel 1914: 301. -Type: Liebusch s.n. Tanzania, West Usambara, Lutindi (B, lectotype, selected here); Mildbraed 1678, West Usambara, Kiwe-Sabyina-Kathinga-Suttel(syntype, not seen). H. alpina R. E. Fries in Fries & Fries 1948: 78. -Type: R. E. & Th. Fries 2698, Kenya, Aberdare, Kinangop (UPS, holotype). Slender and small plants, never exceeding 20 cm. Corms 5-15 mm long, 4-10 mm broad, growing vertically. Leaves linear, erect reflexed to prostrate, 1-2 mm broad, with sparse indumentum of greyish to yellowish hairs mostly along margins and midrib abaxially. Scape 1-10 cm long. Inflorescence covered with a few whitish to yellowish stellate hairs or almost glabrous. Bracts narrow up to 6 mm long. Flowers small, ca. 1 cm in diameter; perianth segments 4-6 mm long, 1-2.5 mm broad, sometimes with a reddish dorsal line. Filaments filiform 1-3 mm long, with anthers 1-2 mm. Thecae fused or forming a small split apically. Style 0.3-2.5 mm long, stigma 0.8-1.5 mm, i.e. the former may be shorter, equal or longer than the latter. Capsule 3.5-7 mm long, 2-3.5 mm broad; opening mechanism unknown. Seeds about 1 mm long. Testa more or less papillate, but without folded cuticula. zyxwvutsr Fig. 14. Distribution of H. macrocarpa in East Africa. zyxwvu zyxwvutsr zyxwvutsrq diameter 19-23 mm; perianth segments 7-12 (-14) mm long, 3-6 mm broad. Filaments 2.5 mm long with anther 2-3.5 mm; thecae completely fused apically. Stigma 2.5 mm, distinctly longer than the style. Capsule 4-11 mm long, 3.5-6 mm wide; opening mechanism difficult to judge from the herbarium material, but seems to be irregular. Seeds 1.3-1.5 mm long. Testa papillate with distinct cuticula folding (Fig. 5F). Distribution and habitat. The main distribution of H. macrocarpa seems to be SW Tanzania with extensions to SW Kenya (Fig. 14). It has been collected from lo00 to 1850 m above sea level, and it seems to prefer moist, clayish habitats. Once it has been found in a mixed collection with H. malosana, indicating similar ecology. Note. Wilsenach & Warren (1967: 140) have very sensibly pointed out that "Taxonomists shouId refrain from the temptation of describing new species in Hypoxis". We have been very reluctant in establishing the new species. The justification is that its closest relative, H. angustifolia,is a rather homogenous species with a low ploidy level and no indications of apomixis. H. macrocarpa differs in pedicel length, 1-6 mm, in H. angustifo26 H. kilimanjarica Bak. ssp. prostrata Holt & Staubo ssp. nov. Type: Thulin & Tidigs 301, Kenya, N. Cherangani, Chepkotet (UPS, holotype; EA, K, S, isotypes). Ssp. haec ab ssp. kilimanjarica differt foliis prostratis, pedunculis brevioribus, indumento inflorescentiarum sparsiorire. Small plants with prostrate leaves and scape never exceeding 4 cm. Indumentum of inflorescence sparse. Distribution and habitat. H . kilimanjarica ssp. prostrata is known from Kilimanjaro, Aberdare and Cherangani (Fig. 15). It belongs in montane grasslands from 2900 to 3500 m above sea level. zy Examined collections. Kenya: Thulin & Tidigs 201 (UPS, holotype; EA, K, S, isotypes); Mabberley 372 (EA, K); Bogdan 536 (EA). Tanzania: Rauh 129 (EA); Staubo 3 (0). Nord. J. Bot. 5 (1) 1985 zyx zyxw jarica. Nel (1914) justified the species in that it was single-flowered, whereas H. kilirnanjarica is double-flowered. The type specimen of H. afpina R. E. Fries belongs among the most narrow leaved ssp. kilirnanjarica. Examined collections. Kenya: R. E . & Th. Fries 2698 (UPS); Glower, Gwynne & Samuel 951 (EA). Tanzania: Staubo 1 ( 0 ) Volkens ; 781 (B); Drummond & Hemsley 2459 (B, BR, EA, K, S); Liebusch s.n. (B); Volkens 781 (B, holotype). 5. Hypoxis malosana Bak. Baker 1897: 284. - Type: Whyte s.n., Malawi, Mt. Malosa, near Zomba (K, holotype). H. biflora De Wildeman 1913: 537, non Baker 1876: 181. Nom. illeg. - H. dregei Nel var. biflora (De Wild.) Nel 1914: 306. Type: Hock s.n., Zaire, Katanga (BR, holotype). H. dregei sensu Nel 1914: 306, non Baker 1878: 112. H. rniinzneri Nel 1914: 307. - Type: Fromm 127a, Tanzania, Kigoma Distr., Mtembwa Plains (B. holotype). Slender plants up to 38 cm. Corm globose to ovoid 1-4 cm in diameter. Leaves develop after anthesis, stiffly erect, narrowly linear to subterete, 1-3 mm broad with or without distinct nerves and with more or less developed whitish to yellowish indumentum (Fig. 1 A). Scape 4-28 (-34) cm long. Inflorescence 1-4 (-5) flowered (Fig. 21, J), and with indumentum of whitish to yellowish stellate hairs. The lower pedicel is always very short, 0-2 (-4) mm long, the upper somewhat longer, (6) 7-9 (-17) mm long. Bracts narrow, up to 1 cm long. Flowers rather small, diameter 14-18 (-22) mm, with perianth segments 5-10 mm long, 2.5-5 mm broad. Filaments filiform 1.5-4.5 mm long with anthers 1.5-3 (-5.5) mm long. Thecae usually fused apically. Stigma 1-2 (-3.5) mm long, equals or is longer than the style (0.5-2 mm) in length (Fig. 3K). Capsule pyxidal, 1.5-6 mm long, 2-3 mm broad. Seeds up to 1.5 mm long. Testa papillose with dense, rather small papillae, but without folded cuticula (Fig. 5C). zyxwvu zyxwvut zyxwvutsrqpo Fig. 15. Distribution of H. kilimanjarica in East Africa 4 ssp. prostrata. H. kilimanjarica Bak. ssp. kilimanjarica Leaves erect or slightly reflexed. Scape 4-12 cm long. Inflorescence with indumentum of whitish to yellowish stellate hairs. Distribution and habitat. H . kilimanjarica ssp. kifimanjarica belongs in the montane grasslands of Usambara, Kilimanjaro, and Aberdare (Fig. 15) up to about 2900 m, i.e. at somewhat lower altitudes than ssp. prostrata. Note. Superficially H. kilirnanjarica resembles dwarf forms of H. angustifolia. The different testa structures contradict close relationship. H. kilirnanjarica may rather have evolved with its special montane adaptions from the heterogenous H. obtusa-complex. Apomixis cannot be excluded. In addition to the differences between the two subspecies mentioned above, we have found that ssp. prostrata seems to have lower testa papillae and a smaller ratio style length to stigma length than ssp. kilimanjarica. More material is needed to judge these characters. The type material of H. incisa Nel represents a typical H. kilimanjarica ssp. kiliman- Distribution and habitat. The main distribution of H. rnalosana is the Southern Highlands of Tanzania and adjacent areas of Zaire (Katanga), Zambia and Malawi. It extends through western part of Tanzania to SW Uganda (Fig. 16). It grows in different types of grasslands, dry to fairly humid. Flowering seems related to fire. It has been collected from 960 to 2900 m above sea level. Note. H. rnalosana seems to be a “good” species, established at diploid level. In literature (e.g. Moriarty 1975) and on herbarium specimens “H. dregei Nel” is the common name for the species. Nel(l914) cited Baker’s (1878: 112) H. sericea Bak. var. dregei Bak. in the protologue. This variety is based on Drege 8525 (K) and Cooper 1811 (not seen) from Cape. The former has been examined and belongs to a species not represented in Tropical Africa at all. The type of the illegitimate H. biflora De Wild. and H. dregei Nel var. biflora (De zyxwvutsrqpon Nord. J. Bot. 5 (1) 1985 27 cial cuticula folding of the testa (Fig. S), by larger flowers and broader leaves than H. kilimanjarica (Fig. 7) and by different inflorescence and broader leaves than H. malosana (Fig 2,s). Taxa preliminary included in the complex: H. obtusa Burchell in Ker-Gawler 1816: Tab. 159. -Type Burchell s.n., Botswana, Pellat Plains (K, holotype). H. fiheri Pax 1893: 143. -Type: Fisher 611, East Africa, sine loc. (B, holotype). H. gregoriana Rendle 1895: 408. - Type Gregory s.n., Kenya, Kikuyu Escarpment, Kedong (BM, holotype). H. laikipiensis Rendle 1895: 407. -Type: Gregory s.n., Kenya, Laikipia (BM, holotype; B, isotype). H. aculeolara Nel 1914: 327. - Type: Miinzner 59, Tanzania, Msamoia (B;holotype). H. apiculara Nel 1914: 327. - Type: Hildebrandt 2542, Kenya, Taita Hills (B, holotype; BM, K , isotypes). H. araneosa Nel 1914: 310. - Type: Endlich 238, Tanzania, Kilimanjaro (B, holotype; M, isotype). H. arenosa Nel 1914: 325. - Type: Holst 93, Tanzania, Usambara (B, holotype). H. campanulata Nel 1914: 314. - Type: Janensch & Hennig s.n., Tanzania, Lindi (B, holotype). H. crispa Nel 1914: 334. - Type: Volkens 360, Tanzania, Kilimanjaro (B, holotype). H. cryprophylla Nel 1914: 316. - Type: von Trotha 140, Tanzania, Karagwe (B, holotype). H. demissa Nel 1914: 328. - Type: Prittwitz & Gaffron 171, Tanzania, “reise nach Uluguru, Uhehe und Usanga” (B, holotype). H. infausta Nel 1914: 319. - Type: Stuhlmann 9161, Tanzania, Uluguru (B, holotype). H. ingrata Nell914: 311. -Type: Thomson s.n., Tanzania, N of Lake Nyassa. (K, holotype; B, isotype). H. mulirfroru Nel 1914: 317. -Type: Evan James s.n., Uganda, Elgon Distr. (K, holotype). H. probara Nel 1914: 317. - Type: Stolz 537, Tanzania, Kyimbila (B, holotype; G, K, L, M, S, isotypes). H. protrusa Nel 1914: 336. - Type: Merker 18, Tanzania, Kilimanjaro (B, lectotype, selected here); Jaeger 243, Tanzania, between Akida Maussa and Mangati (B, syntype). H. retracfaNel 1914: 312. -Type: Whyte s.n., Kenya, near Eldama (B, lectotype, selected here, K,isotype). H. urceolata Nel 1914: 336. - Type: Whyte s.n., Kenya, near Nairobi, (K,lectotype, selected here); Obst s.n., Tanzania, sine loc. (B, syntype); Dawe 103, Uganda, Ohagwe (K,syntype); Dawe 231, Uganda, Buddu (K, syntype). H. zernyi Schulze 1939: 375. - Type: Zerny 20, Tanzania, Lupembe Hill, WSW of Songea (B, holotype). H. matangensis Schulze 1939: 376. - Type: Zerny 374, Tanzania, Uyanga-yenga t o Lindi Road, WSW of Songea (B, holo- zyxwvutsr zyxwvu zyxwv z zyxwvutsrq zyxwvutsrqp zyxwvutsrqp zyxwv Fig. 16. Distribution of H. malosana in East Africa. Wild.) Nel from Katanga corresponds to the Tanzanian material of H. malosana in every aspect. H. bifolia Bak. (Baker 1876) is based on material from Transkei (not seen), and it was reduced to H. angustifolia Lam. by Baker (1878). The holotype of H. miinzneri Nel belongs in the outskirts of the variation range of H. malosana, having slightly larger flowers, longer leaves and scapes. In addition it has been collected in the periphery of its distribution area (Kigoma Distr.). One specimen from Tabora Distr. (Dautrique 101, BR) is more or less intermediate and links the H. miinzneri type to the rest of H. malosana. Selected collections. A total of 51 specimens belonging to 37 collections was studied. Uganda: Eggeling 1106 (K); Tanzania: Bullock 3293 (K); Richards 11617 (B, BR, K,LISC, S); Bruce 751 (K); I Bjcjrnstad 794 (0);Richards 14146 (BR, K);MilneRedhead & Taylor 8058 (B, BR, E A , K, LISC). 6 . Hypoxis obtusa Burch.eompkx This group certainly includes apomictic taxa. The group is extremely heterogenous, but at the moment we are not able to suggest further delimitation. The group is more or less negetatively defined by the lack of the spe28 type). Plants small and slender to very robust. Corm 1-8.5 cm long, 0.7-7 cm broad. Leaves linear to lanceolate, erect to almost prostrate, 0.4-5 cm broad. Indumentum variable (Fig. 1 D-F), d o u r most often whitish. Scape 2-45 (-60 cm) cm long. Inflorescence with (1-) 2-20 flowers, those with few flowers shaped more or less like a corymb, those with many flowers like a raceme (Fig. 2 E-H). Flower diameter 1.5-3.8 cm; perianth segments 5-18 mm long, 2.5-9 mm broad. Filaments subulate, 1.5-4 mm long with anther 3-8 mm long; thecae fused, rarely with an indistinct split. Style 0.5-3.2 mm long, usually shorter than the stigma, 1.3-3.2 m m long. Fruit a pyxis, 3-10 mm long, >7 mm broad. Seeds subglobose 1-2 mm long. Testa pattern variable, usually paNord. J. Bot. 5 (1) 1985 pillate, always without cuticula folding. The group needs further study. Distribution and habitat. Plants of the H. obtusa-complex are distributed over the whole of Tropical Africa. They belong in different types of grassland and woodland, at altitudes from 300 to almost 3000 m above sea level. Note. The oldest African taxon in the genus is H. villosa L., a name also used for East African material here referred to the H. obtusa-complex (Agnew 1974). The original collection is from the Cape and deposited in LINN. It is represented by one leaf and a single-flowered inflorescence. Until the genus is revised in South Africa, and H. villosa is further characterized and circumscribed, we recommend the name excluded in Tropical Africa. LISC, M, 0, P, PRE, S, UPS, and WAG. Prof. K. Jones kindly allowed us to study his preparations of Hypoxis root-tips in Jodrell Laboratory, Royal Botanic Gardens, Kew. References Agnew, A. D. Q. 1974. Upland Kenya wild flowers. -London. Andrews, F. W. 1956. The flowering plants of the Sudan 3. Abroath. Baker, J. G . 1876. On new bulbous plants from the Eastern Provinces of Cape Colony. - J. Bot. 14: 181-184. - 1878. A synopsis of Hypoxidaceae. - J. Linn. SOC.17: 93126. - 1896. Amaryllideae. - In: Thiselton-Dyer, W. T. (ed.), Flora Capensis, Vol. 6, London, pp. 171-246. - 1897. Diagnoses Africanae. - Kew Bull. 10: 243-300. - 1898. Amaryllideae. - In: Thiselton-Dyer, W. T. (ed.), Flora of Tropical Africa, Vol. 7, London, pp. 377-382. Bentham, G. 1883. Amaryllideae. - In: Bentham, G. & Hooker, J. G., Genera Plantarum, Vol. 3. London, pp. 71 1-740. Binns, B. 1968. A first check list of the herbaceous flora of Malawi. - Zomba. Britton, N. I. & Brown, A. 1913. An illustrated flora of the Northern United States (2. ed.) Vol. 1. - New York. De Wildeman, E. 1913. Decades novarum speciorum Florae Katangensis. - Feddes Repert. 11: 535-547. Fernandes, A. & Neves, J. B. 1962. Sur la caryologie de quelques monocotyledones Africaines. - Comptes rendu de la IVe reunion de 1'A.E.T.F.A.T.- Lisboa. Fries, R. E. & Fries, Th. C. E. 1948. Phytogeographical researches on Mt. Kenya and Mt. Aberdare, British East Africa. - Svensk Vet.-Akad. Handl. Ser. 3, 25, No. 5. Fzgri, K. & Iversen, J. 1964. Textbook of pollen analysis (2. ed.). - Copenhagen. Geerinck, D. 1968. Considerations taxonomiques au sujet des Haemodoraceae et des Hypoxidaceae (Monocotyledones). - Bull. SOC.Bot. Belg. 101: 265-278. - 1969. Genera des Haemodoraceae et des Hypoxidaceae. Bull. Jard. Bot. Nat. Belg. 39: 47-82. - 1971. Hypoxidaceae in Flore du Congo, du Rwanda et du Burundi. - Bruxelles. Harms, H. 1901. Amaryllidaceae. - In: Engler, A. (ed.), Beitrage zur Flora von Afrika 22, Bot. Jahrb. Syst. 30: 239445. Heideman, M. E. 1979. Taxonomic studies in the genus Hypoxis L. (Hypoxidaceae) on Witswatersrand. - M. Sc. Thesis. Univ. of Witwatersrand (Unpubl.). - 1983. Studies of diagnostic features in the genus Hypoxis L. (Hypoxidaceae R. Br.) on the Witwatersrand. - Bothalia 14: 889-893. Hepper, F. N. 1968. Hypoxidaceae. - In: Hepper, F. N. (ed.), Flora of West Tropical Africa (2. ed.) Vol. 3,1, London. Hilliard, 0. M. & Burtt, B. L. 1978. Notes on some plants from southern Africa, chiefly from Natal 7. - Notes Roy. Bot. Gard. Edinb. 36: 43-76. Hutchinson, J. 1934. The families of flowering plants. Vol. 2. London. Ker-Gawler, J. B. 1816. Hypoxis obtusa. - Bot. Reg. 2: 159. Laane, M. M. 1970. HQndboki kromosomteknikk for biologer og medisinere. - Oslo. Lamarck, J. B. P. A. M. 1789. EncyclopCdie Methodique. Vol. 3. - Paris. Lindley, J. 1836. A natural system of botany (2. ed.). London. Linnaeus, C. 1759. Systema naturae (10. ed.). - Stockholm. Linnaeus, C. fil. 1781. Supplementum Plantarum. - Braunschweig. Mehra, P. N. & Sachdeva, S. K. 1976. Cytological observations on some W. Himalayan monocots IV. - Cytologia 41: 3153. zyxwvutsrqp zyxwvuts Selected collections. A total of 473 specimens belonging to 306 collections was studied. Uganda: Eggeling 3597 (K); Wilson 997 (EA, K); Hazel 441 (BR, K, S); Lye, Faden & Evans 4328 (EA, K); Wood 992 (B, BR, K, P, PRE, S); Snowden 822 (BM, K); Langdale-Brown 2065 (EA, K). Kenya: Gillett 13902 (BR, EA, K); Napier 1942 (EA, K); Dummer 5082 (K); Bogdan 3691 (B, EA, K); Symes 51 (EA, K); Faden in EAH 67224 (EA); Gilbert Rodgers 479 (BR, E A , K, S); Maas Geesteranus 5193 (BR, G , K, L, PRE, S); Glover, Gwynne & Samuel 614 (BR, EA, K, PRE). Tanzania: Greenway 10780 (EA, K, PRE); Bally 2424 (G, K); Greenway 6941 (EA, K); Faulkner 1420 (B, BR, K, LISC, S); Richards 7207 (BR, K); Burtt 1064 (K); Hornby 2097 (EA, K); Drummond & Hemsley 1565 (EA, K); Schlieben 3506 (B, BR, K, LISC, PRE); I. Bj~irnstad401 (K, 0);Polhill & Paulo 1394 (B, BR, E A , K, LISC, P, PRE); Richards 17006 (B, BR, K, UPS); Semsei 2593 (EA, K); Milne-Redhead & Taylor 7733 (B, BR, EA, K, LISC). zyxwvutsrqpo zyxwvu zyxwvutsrqp 7. Hypoxis sp. A Two specimens with seeds of the H. angustifolia-type (Fig. 4b) have been impossible to refer to any of the taxa presented. They are both very robust and thus resemble H. goetzei. They deviate, however, with respect to indumentum and inflorescence, compared to the rather homogenous H. goetzei.The latter is highly polyploid, and apomixis might be expected. Davies 742 (K) from South-West Tanzania, Mbeya Distr., differs from H. goetzei by having more than 20 flowers in a dense raceme and by the indumentum of long white hairs along leaf margins and midribs. Bullock 2045 (K) from Ufipa Distr. also of SW Tanzania, differs by being extremely robust, and strongly tomentose with a complete covering of red-brown hairs. Until more material is known we propose that they are preliminarily kept separate under the heterogenous Hypoxis sp. A. Acknowledgements - The living material studied has been collected on expeditions to East Africa financially supported by the Norwegian Research Council for Science and the Humanitites and NORAD. We are also grateful to Directors and Curators of the following herbaria: B, BM, BR, EA, G, K, L, zyxwvutsrqpon Nord. J. Bot. 5 (1) 1985 29 zyxwvu zyxwvuts zyxwvutsr zz Melchior, H. 1964. Engler's Syllabus der Pflanzenfamilien. Berlin. Moriarty, A. 1975. Wild flowers of Malawi. - Cape Town. Naranjo, C. A. 1975. Chromosomal studies in Hypoxis L. Karyotype of H . decumbens L. - Phyton 33: 45-49. Nel, G. 1914. Studien iiber die Amaryllidaceae-Hypoxideae, unter besonderer Beriicksichtigung der afrikanischen Arten. - In: Engler, A. (ed.), Beitrage zur Flora von Afrika 43, Bot. Jahrb. 51: 234338. Ostergren, G. & HenCen, W. K. 1962. A squash technique for chromosome morphological studies. - Hereditas 48: 332341. Pax, F. 1889. Amaryllidaceae. - In: Engler, A. & Prantl, K. (eds.), Die Natiirlichen Pflanzenfamilien. Part 2. sect. 5, Leipzig, pp. 97-125. - 1893. Amaryllidaceae africanae. - In: Engler, A., Beitrage zur Flora von Afrika 2, Bot. Jahrb. 15: 140-144. - & Hoffman, K. 1930. Amaryllidaceae. - In: Engler, A. & Prantl, K. (eds.). Die Natiirlichen Pflanzenfamilien (2. ed.) Vol. 15a, Leipzig, pp. 391-430. Phillips, E. P. 1926. The genera of South African flowering plants (1 ed.). - Cape Town. Rendle, A. B. 1895. A contribution to the flora of Eastern Tropical Africa. - J. Linn. SOC.30: 373-435. Robyns, W. & Toumay, R. 1955. Monocotyltes nouvelles ou critiques de la region du Parc National Albert (Congo Belge). - Bull. Jard. Bot. Nat. Belg. 25: 239-260. Schulze, G. M. 1939. Hypoxis zernyi Schulze. -In: Mildbraed, J. (ed.), Neue Arten aus dem Matengo-Hochland, Siidwestliches Tanganyika-Territ, leg. H. Zemy 2, Notizbl. Bot. Cart. Berlin 14: 376-378. Thompson, M. F. 1976. Studies in the Hypoxidaceae. 1. Vegetative morphology and anatomy. - Bothalia 12: 111-117. Wilsenach, R. 1967. Cytological observations on Hypoxis. 1. Somatic chromosomes and meiosis in some Hypoxis species. - J. S. Afr. Bot. 33: 75-84. - & Papenfuss, J. N. 1967. Cytological observations on Hypoxis. 2. Pollen germination, pollen tube growth and haploid chromosome numbers in some Hypoxis species. - J. S. Afr. Bot. 33: 111-116. - & Warren, J. L. 1967. Cytological observations on Hypoxis. Embryo-sac development in H . rooperi and H . filiformis. - J. s. Afr. Bot. 33: 133-140. Wood, S. E. 1976. A contribution to the knowledge of the genus Hypoxis L. (Hypoxidaceae) in Natal, South Africa. M. Sc. Thesis. Univ. of Pietermaritsburg (Unpubl.). zyxwv 30 zy Nord. J. Bot. 5 (1) 1985