“COMPOSITAE & % NEWSLETTER | Number 26 June 1995 ; Scientific Editor: Bertil Nordenstam Technical Editor: Gunnel Wirénius Nohlin Published and distributed by The Swedish Museum of Natural History, Department of Phanerogamic Botany, P.O. Box 50007, S-104 05 Stockholm, Sweden (Director: Professor Bertil Nordenstam) ISSN 0284-8422 CONTENTS Moses A. Isawumi: A new species of Baccharoides (Asteraceae: Vernonieae) from Uganda, East Africa 1 Moses A. Isawumi: Floral microcharacters and taxonomy of the Cyanthillium cinereum complex (Asteraceae: Vernonieae) 11 M. S. Ayodele: Studies on the reproductive biology of Vernonia Schreb. (Asteraceae). III. Floret shapes/number, pappus types and surface structure among different growth habits 26 A. Ouyahya: Etude anatomique preliminaire des akénes de quelques Artemisia dans la bassin méditerranéen occidental 40 Y.R. Ling: Survey of Artemisia and Seriphidium (Asteraceae-Anthemideae) in the Himalayan Mountains and the South Asian Subcontinent 49 I. Hoddélovaé and P. Madrtonfi: Pollen morphology in the Senecio nemorensis Group (Compositae) from the Carpathians 61 Book Announcement: Asteraceae - Cladistics and Classification by K4re Bremer 71 Comp. Newsl. 26, 1995 A NEW SPECIES OF BACCHAROIDES (ASTERACEAE: VERNONIEAE) FROM UGANDA, EAST AFRICA Moses A. Isawumi * Department of Phanerogamic Botany Swedish Museum of Natural History P. O. Box 50007 S-104 05 Stockholm, Sweden Abstract Baccharoides tayloriana formerly including in the Vernonia hymenolepis group is described as a species new to science. The new species is found on the Ruwenzori mountain, Uganda. A short discussion of its systematic position and an illustration are provided. Introduction The genus Baccharoides Moench was resurrected by Robinson (1990) who transferred three Vernonia species to the genus on the basis of its lophate pollen with polar areoles, which is distinctive among Old World tricolporate Vernonieae. Isawumi (1993) transferred four other species and seven infraspecific taxa from the genus Vernonia to Baccharoides based on its strikingly distinctive floral characteristics which separate it from the genus Vernonia s.l. The resurrected genus was found to possess corollas with long, slender, basal tubes, abruptly expanded cylindrical limbs and comparatively short erect lobes; phyllaries are apically appendaged; and the style bases are completely enclosed by long keeled nectaries. During a comprehensive study of the genus Baccharoides (Isawumi & El-Ghazaly in press) using pollen morphology and floral microcharacters to diagnose the species it became obvious that a specimen from Ruwenzori in Uganda, hitherto treated as belonging to Vernonia hymenolepis A. Rich., should be separated as a new species. * Permanent address: i oe a. History Museum, Obafemi Awolowo University, Ile-Ife, Nigeria, after May 31, 1995 Comp. Newsl. 26, 1995 Baccharoides tayloriana Isawumi, sp. nov. (Fig. 1 A1-A3; PL1 A-G) Holotype: Uganda, Ruwenzori, Namwamba valley, Kilembe, c. 2,850-3,150 m, 6.i.1935, Taylor 2934 (S). Frutex erectus tomentosus. Folia alterna petiolata, lamina plana elliptico-lanceolata herbacea, pinnatinervia, margine serrata, 13.5—25 cm. longa et 5—8.5 cm. lata, inferne lanatattomentosa, super sparsim puberula, apice acuminata. Capitula c. 3.5 cm. diametro, homogama, discoidea, in corymbum laxum terminalem disposita. Pedunculus 2—5 cm. longus. Involucrum late campanulatum. Phyllaria 1.7—-2.6 cm. longa, pluriseriata, scariosa, appendicibus apicalibus, apice acuminata. Receptaculum convexum epaleatum. Flosculi disci hermaphroditi, c. 14 mm. longi, corolla e tubo angusto abrupte ampliata lobis quinque erectis. Antherae basi calcaratae; appendicibus apicalibus ovatisanceolatis, pilis unicellularibus tectis; endothecio polarato. Filamentum collum dilatatum; filamenta longa torsiva, nectario connata. Cypselae homomorphae 5-6 mm. longae, 16—20-costatae, sparsim pilosae. Pappi setae pluriseriatae barbellatae caducae. An erect shrub, copiously tomentosely hairy. Leaves alternate; lamina 13.5—25 x 5-8.5 cm, flat, pinnativeined, distinctly serrate at margin with many callose-tipped teeth on each side, elliptic-lanceolate, with cuneate attenuate base tapering into a petiole c. 3-5 cm long, lower surface woolly tomentose with appressed matted T-shaped hairs interspersed with scattered glandular hairs, upper surface sparsely puberulous with scattered patent hairs; apex acuminate. Capitula c. 3.5 cm in diam., homogamous, discoid, in loose terminal flat-topped corymbs. Peduncles 2-5 cm long, with a few lanceolate bracts below the capitula. Involucre broadly campanulate. Phyllaries pluriseriate, with overlapping margins, scarious, apically appendaged; outer phyllaries lanceolate c. 1.7 cm long, glabrescent apically, tomentose basally, with a midvein and usually 2 veins on either side; middle phyllaries c. 2.4 cm long including the appendage, acuminate at tip, broadly lanceolate, tomentose at the junction of the appendage and the base, with a midrib and about 4 veins on either side, acuminate at tip; inner phyllaries c. 2.6 cm long including the appendages, tomentose on the appendage and the margins of the phyllary base, elliptic-lanceolate, gradually tapering to the acuminate apex. Receptacle somewhat convex, epaleate. Disc-florets perfect, c. 14 mm long with long, slender, basal tubes, abruptly expanded cylindrical limbs and 5 lanceolate erect lobes. Veins forming an arch at the corolla lobe without median trace; corolla with capitate glands and biseriate eglandular trichomes concentrated on corolla lobe tips also with many biseriate eglandular trichomes and few capitate glandular hairs on the throat and the tube. Epidermal cells of the adaxial corolla lobe elongate and distinctly wavy. Anthers calcarate; apical appendage ovate—lanceolate, narrower than the thecae, acute at tip, completely covered with unicellular hairs; distal hairs acute/acuminate, proximal hairs obtuse. Endothecial tissue polarized. Filament collar somewhat dilated distally but not thicker than the filament; filament long, coiled and connate to the nectary at the base of the corolla tube. Style base slightly dilated and Comp. Newsl. 26, 1995 enclosed by a long cylindrical nectary; style branches apically tapering, long exserted and curved, with distinctly stigmatic areas throughout the inner portions, outsides covered with unicellular hairs down to the upper portion of the shaft. Cypselas homomorphic, dark brown, 5-6 mm long, narrowly oblong, terete, 16—20-costate, sparsely pilose with short twin hairs, few capitate glands and many idioblasts; testa epidermis ornamented; carpopodium large, annular, the cells heavily thickened with simple pits; ovary crystals broad, lozenge-shaped. Pappus bristles pluriseriate, caducous, flat, acute/acuminate at tip with some dilated distally, basally connate to form an annulus, barbellate on margins, stramineous or somewhat fulvous. Pollen echinolophate, tricolporate, micropunctate with long colpi. Discussion The striking distinguishing characters of the new species are, inter alia, the complete covering of the apical anther appendage with unicellular hairs (Fig.1A2, Pl.1A—B) and the extraordinarily long coiled filaments which are connate to the nectary at the base of corolla tube (Fig.1A3). These characters are autapomorphic for the species, and clearly distinguishes it from the Vernonia hymenolepis group and other species of Baccharoides. The strange apical anther appendage has not been observed in any other species in the family Asteraceae (Karis, Anderberg, Nordenstam, pers. comm.). V. hymenolepis belongs to the genus Baccharoides (Isawumi & El-Ghazaly in press) because it shares with the genus all the diagnostic characteristics mentioned above. The new species is superficially somewhat similar to V. hymenolepis. As a result, most authors might have been lumping it with either V. hymenolepis (Wild 1978, Jeffrey 1988) or V. tolypophora Mattf. (Pope 1992). With the above listed characters, B. tayloriana is, however, distinct, but if it is put as a separate genus, Baccharoides may become paraphyletic. The crystals in the ovary wall of the new species are broad and lozenge-shaped (Dormer 1961, Plate 1), whereas those of V. hymenolepis and other species of Baccharoides are elongate and hexagonal in shape. The observably different macrocharacters of the new species from V. hymenolepis are the larger capitula and larger leaves. The new species belongs to the genus Baccharoides because it shares some mor- phological characters which define the genus. Some of the characters have already been mentioned above and include the elongate corolla tube with abruptly expanded cylindrical limbs and phyllaries apically appendaged. It also shares with the genus some major pollen characters which separate it from the genus Vernonia s.l., some of the characters being lophate pollen with long colpi (P1.1H) and with equatorial lacunae (P1.1G). Its pollen is therefore echinolophate, tricolporate, micropunctate and with large basal columellae, the sort of grains classified by Jones (1981) as 'type C' pollen. The ‘type C' pollen is said to be unique to section Stengelia (now genus Baccharoides) in the Old World and is different from most type C of the Neotropics by the presence 4 Comp. Newsl. 26, 1995 of three equatorial areoles of the intercolpi and strong basal columellae under the crests of the exine (Robinson 1990). Acknowledgements I would like to acknowledge the assistance of Prof. B. Nordenstam and Dr. A. Anderberg for reading the manuscript and offering valuable comments and also Dr. P. O. Karis for discussion of microscopic observations. I am also grateful to Pollyanna Lidmark for the illustration. Financial support from the Swedish Institute is also gratefully acknowledged. References Dormer, K.J. 1961. The crystals in the ovaries of certain Compositae. Ann. Bot. n.s. 25: 241-254. Isawumi, M.A. 1993. New combinations in Baccharoides Moench (Vernonieae: Compositae) in West Africa. Feddes Repert. 104: 309-326. Isawumi, M.A. & G. El-Ghazaly in press. Pollen morphology, floral microcharacteristics and taxonomy of the genus Baccharoides Moench (Vernonieae: Asteraceae). Grana Palyn. Jeffrey, C. 1988. The Vernonieae in East Tropical Africa. Notes on the Compo- sitae:V. Kew Bull. 43: 195-277. Jones, §.B. 1981. Synoptic classification and pollen morphology of Vernonia (Compositae: Vernonieae) in the New World. Rhodora 81:425-447. Pope, G.V. 1992. Compositae. In: Flora Zambesiaca 6, 1:1—264. Robinson, H. 1990. Six new combinations in Baccharoides Moench and Cyanthillium Blume (Vernonieae: Asteraceae). Proc. Biol. Soc. Wash. 103: 248-253. Wild, H. 1978. New and interesting Vernonia species from the Flora Zambesiaca area. Kirkia 11: 1-23. Comp. Newsl. 26, 1995 Fig. 1 Pi. 1 FIGURE LEGENDS Baccharoides tayloriana \sawumi. A: Habit; Al: Disc-floret; A2: Stamen with anther apical appendage covered with unicellular trichomes; A3: Floret with long, coiled filaments which are connate to the nectary at the base of corolla tube. — A—A3: Taylor 2934 (S). Microcharacters of the floret of B. tayloriana Isawumi A: Anther apical appendage covered with unicellular trichomes, x 250. B: Base of anther apical appendage, x 100. C: Corolla lobe showing glandular and unicellular eglandular trichomes, x 100. D: Middle of corolla lobe showing wavy epidermal cells, x 250. E: Polarized endothecial tissue, x 250. F: Ovary crystals lozenge-shaped, x 250. G: LM, polar view of pollen, x 400. H: LM, equatorial view of pollen with long colpus, x 400. — A-H: Taylor 2934 (S). ‘ Comp. Newsl. 26, 1995 Fig. 1 Comp. Newsl. 26, 1995 1A&B Pl Comp. Newsl. 26, 1995 PL1IC&D Comp. Newsl. 26, 1995 PLIE&F Comp. Newsl. 26, 1995 PL1G&H Comp. Newsl. 26, 1995 FLORAL MICROCHARACTERS AND TAXONOMY OF THE CYANTHILLIUM CINEREUM COMPLEX (ASTERACEAE: VERNONIEAE) Moses A. Isawumi * Department of Phanerogamic Botany Swedish Museum of Natural History P.O. Box 50007 S-104 05 Stockholm, Sweden Abstract In this study of Cyanthillium cinereum (L.) H. Robinson four varieties are recognized, one of which is new and three are new combinations. The floral microcharacters of the species and its varieties are described. Introduction The genus Cyanthillium Blume was resurrected by Robinson (1990) who transferred Vernonia cinerea (L.) Less., V. stellulifera (Benth.) C. Jeffrey and Conyza patula Ait. to the genus. The transfer was based on the pollen characteristics, which are remarkably different from those of the genus Vernonia and other Vernonieae in that the colpus is not evident and the grains have a polar areole surrounded by a tier of five to seven areoles. In other words the polar organization is basically different from most forms in the tribe which have well developed colpi. According to Robinson (1990) the genus Cyanthillium is distinctive within the non-colporate genera of the Old World by the modification of the basal columellae of the exine to form bridges under the crests. However, the genus is substantially different from the genus Baccharoides Moench, which possesses grains with long colpi (Isawumi & El-Ghazaly in press). The pollen study of V. cinerea by Kingham (1976) also pointed out the lack of colpi in the species. *Permanent address: f a —_ Natural History Museum, Obafemi Awolowo University, Ile-Ife, Nigeria, after May 31, 1995 12 Comp. Newsl. 26, 1995 Materials and Methods The material for this study was obtained from the herbarium of the Swedish Museum of Natural History in Stockholm (S). For the micro morphological observations, florets were softened in boiling water to which a drop of detergent was added, dissected under a stereo microscope, mounted in Hoyer's solution (Anderson 1954, King & Robinson 1970) and studied with LM. Results and Discussion MORPHOLOGICAL MICROCHARACTERS Corolla pubescence Pubescence of the corolla lobes is a common feature in the Asteraceae. Uniseriate eglandular trichomes and capitate glands are commonly found on the corolla lobe tips of Cyanthillium cinereum and its varieties in all the geographical areas where they occur (Fig. 1A), except var. ugandense, which has unicellular eglandular trichomes and T-shaped trichomes in addition (Fig. 2E). On the corolla lobe tips of the varieties in Madagascar are found uniseriate and biseriate stalked glands in addition to the ones mentioned above. In all the varieties, the same types of trichomes found on the corolla lobe tips are present on the limbs and tubes with sometimes slight modifications. The uniseriate eglandular trichomes are different from those found in the genus Baccharoides (Isawumi & El-Ghazaly in press) because they are usually very long, about 2-3 cells long, with the apical cells much longer than the others. These trichomes are also different from those present on the corolla lobes of Vernonia s.l. This character can be conveniently used to separate the varieties occurring in Uganda and Madagascar. Anther apical appendage The anther apical appendages are sufficiently different within the species to bring about the delimitation of the varieties. In the varieties that occur in Australia, Tanganyika, Zimbabwe (S. Rhodesia) and Malawi, the apical appendages are distinctly ovate and obtuse at tips (Fig. 2H). Some varieties which occur in Uganda, Jamaica, Trinidad (West Indies), and Philippines, have apical appendages that are slightly dilated proximally, taper towards the apex and are obtuse or sometimes acute at tips (Fig. 31). In Madagascar there are two types of apical appendages. In var. viale, the appendage is more or less lanceolate or ovate-lanceolate and obtuse at tip (Fig. 3J) , while in var. ovatum it is ovate with truncate tip which is slightly notched in the middle (Fig. 3K). Comp. Newsl. 26, 1995 The anther apical appendages are not conspicuously constricted at the base as normally in the tribe Vernonieae (sensu King & Robinson 1970) but they are as wide as the thecae (cf. Anderberg 1991). Furthermore the appendages do not have glands like the neotropical Vernonieae (Robinson & Kahn 1986). Endothecial tissue The endothecial tissue in the species and its varieties is intermediate (Fig. 1B) with the cells thickened on both the horizontal and radial walls (Dormer 1962, Nordenstam 1978). The thickenings are weakly connected across the external surface of the cells (Fig. 1B) as usually found in the tribe Vernonieae (Robinson 1977). This character is autapomorphic for the tribe Vernonieae. The endothecial tissue in the genus Baccharoides is distinctly polarized (Isawumi & El-Ghazaly in press) and therefore different from the one in this species which is placed in the resurrected genus Cyanthillium. Filament collars The filament collars have long been used by synantherologists for taxonomic discrimination (Nordenstam 1978). The collars are the uppermost parts of the filaments. They are narrowly cylindrical, elongate, straight and not wider than the filaments (Fig. 2G) in all the varieties of this species. They are different from those found in the genus Baccharoides which are ovate and sometimes dilated distally (Isawumi & El-Ghazaly in press). The filaments in this species are inserted on the corolla at the limb/tube junction. Base of anther thecae The terminology to describe the basal portion of the anthers, suggested by Robinson (1983), viz., calcarate (spurred) versus ecalcarate, has been used in synantherology by many authors (Bremer 1987, Anderberg 1991, Karis 1993). The anthers in Cyanthillium cinereum are long calcarate because the thecae extend below the point where the filament connects with the anther (Fig. 2G). Ovary wall crystals Ovary crystals have been used successfully in the taxonomic evaluation of several tribes of the family Asteraceae (Dormer 1961, Nordenstam 1978, Nordenstam & EI- Ghazaly 1977, Anderberg 1991). Crystals are usually absent in tribes with carbonized achene walls (Robinson 1977). 14 Comp. Newsl. 26, 1995 In C. cinereum the ovary wall crystals are narrowly elongate och hexagonal in surface view and rectangular in side view (Fig. 2F). Crystals sometimes found in the corolla and style may have various shapes - spindle- shaped, hexagonal, clustered, lozenge. Style base The style base is conspicuously ring-like with the cells well thickened with simple pits (Fig. 1D). The cells are more or less rectangular and range from 1-5 in a row. The ring-like style base is absent in the genus Baccharoides, which has the style base completely covered by the nectary (Isawumi 1993). Cypsela The term cypsela (sensu Stearn 1966, Fahn 1982) is often considered more strictly correct than the term achene (sensu Wagenitz 1976). The cypsela of C. cinereum is subfusiform, terete, more or less ribbed, and narrowed below. The pubescence of the cypselas in the varieties shows some slight variations. The cypselas may have many idioblasts, few capitate glands and many twin hairs whose parallel cells fuse almost to their apices (Fig. 1C). In the varieties occurring in Zimbabwe, Uganda, Malawi, and Madagascar, one of the parallel cells is longer than the other cell. The parallel cells in the varieties in Trinidad, Madagascar (i.e. var. ovatum), British Guiana, and Tanganyika are almost of equal length (Fig. 1C). The cypsela trichomes have been called twin hairs (Hess 1938), but they usually have three cells (Nordenstam 1968, Bremer 1987, Anderberg 1991), one basal-lateral and two elongated parallel cells. At the point'of fusion of the two parallel cells are sometimes found simple pits (Fig. 1C). Testa epidermis Testa epidermis in C. cinereum and its varieties is omamented with pattern rather different from that of Baccharoides (Isawumi & El-Ghazaly in press). Carpopodium The carpopodium of the species and its varieties has cells which are heavily thickened with pits like in the genus Baccharoides. Comp. Newsl. 26, 1995 Pappus The species has an outer pappus of distinct, short, linear, fimbriate scales. The inner pappus bristles are white, terete and readily caducous. Sometimes the outer pappus scales may be absent. The pappus has been a classical source of information at the generic level but less so at higher levels (Bremer 1987), and much emphasis has been given to this structure in the classification of Vernonia s.l. (Jeffrey 1988, Wild 1978, Pope 1992, Isawumi 1989, 1993). TAXONOMY Cyanthillium cinereum (L.) H. Robinson, in Proc. Biol. Soc. Wash. 103: 252 (1990). Conyza cinerea L., Sp. Pl. 2: 862 (1753). Vernonia cinerea (L.) Less. in Linnaea 4: 291 (1829); DC., Prodr. 5: 24 (1836); Oliv. & Hiern in F.T.A. 3: 275 (1877); Eyles in Trans. Roy. Soc. S. Afr. 5: 503 (1916); Mendonca, Contrib. Conhec. Fl. Angol., 1 Compositae: 14 (1943); Adams in F.W.T.A. ed. 2, 2: 283 (1963); Wild in Kirkia 11: 80 (1978); Maquet in Fl. Rwanda, Spermat. 3: 558 (1985); C. Jeffrey in Kew Bull. 43: 224 (1988); G.V. Pope in FI. Zambesiaca 6: 143 (1992); H. Humbert in Fl. Madagascar 1: 18 (1960); Fawcett and Rendle in Fl. Jamaica 7 (5): 162 (1936); Cheeseman, Hill & Burtt in Fl. Trinidad and Tobago: 58 (1940); Bentham in Fl. Australiensis 3: 459 (1866, reprint 1967). Type: Sri Lanka, Hermann (Lectotype BM, Herb. Hermann, vol. 3). (i). var. cinereum Vernonia cinerea (L.) Less. subsp. cinerea, H. Humbert in Fl. Madagascar, Compo- sées 1: 19 (1960). Phyllaries 3-seriate, outer linear, inner lanceolate, c. 4 mm long; cypsela c. 1.5 mm long; outer pappus scale-like, slightly persistent; inner ones terete, barbellate, white, c. 3.5 mm long. Distribution: This variety is pantropical. Madagascar, Majunga, Viguier et Humbert s.n. (S), cited by Humbert (1960); Decary 10592 (S); also in Kenya, Tanzania. A specimen without locality in P (Humblot s.n.) also belongs here. Humblot collected in Madagascar and the Comoro Islands. (ii). var. ugandense (C. Jeffrey) Isawumi comb. nov. Basionym: Vernonia cinerea (L.) Less. var. ugandensis C. Jeffrey, in Kew Bull. 43: 224 (1988); G.V. Pope in Fl. Zambesiaca 6: 143 (1992). Type: Uganda, West Nile Dist. Terego, Hasel 492 (K holotype). Corolla lobe tip with unicellular and T-shaped eglandular trichomes and many capitate and long stalked uniseriate glands. Anther apical appendage lanceolate-ovate and 16 Comp. Newsl. 26, 1995 obtuse at tip; filament collar straight, elongate, with the cells much thickened on the horizontal walls. Cypsela S-costate, 1.5—1.8 mm long, sparsely pilose with twin hairs having parallel cells of unequal length; inner pappus bristles c. 3 mm long. Taxonomic note: This variety, according to Jeffrey (1988) and Pope (1992) can be distinguished by its shorter inner pappus. It has other distinguishing characteristics which include the presence of T-shaped trichomes on the corolla lobe tip (Fig. 2E). This character is found only in this variety. Distribution: Uganda, Ruwenzori, Kilembe, alt. c. 1,200-1,500 m, Taylor 2464 (S); also in Kenya. (iii). var. viale (DC.) Isawumi, comb.nov. Basionym: Vernonia vialis DC., Prodr, 5: 25 (1836). - V. cinerea (L.) Less. subsp. vialis (DC.) H. Humbert in Fl. Madagascar, Composées 1: 20 (1960). Type: Madagascar, Tamatave, Bojer s.n. (BM). Syn. V. arguta Bak. in J. Linn. Soc. 20:175 (1883). Corolla lobe tip with few unicellular eglandular trichomes (like twin hairs), many long uniseriate eglandular trichomes and capitate glands; outer phyllaries linearfiliform, inter phyllaries lanceolate, acuminate/apiculate, 6 mm long. Anther apical appendage lanceolate-ovate and obtuse at tip. Crystals in the style tiny and spindle-shaped. Cypsela c. 1 mm long; innter pappus bristles c. 5 mm long. Taxonomic note: This variety has been found only in Madagascar. Its anther apical appendage is long, lanceolate-ovate (Fig. 3J). It also has many spindle-shaped crystals in the style. Specimen examined: Madagascar, Firingalava, Perrier 597 (S), cited by Humbert (1960). (iv). var. ovatum Isawumil, var. nov. Holotype: "Lecta in Madagascar a D. Thunberg", ex herb. Swartz (S). Herba annua erecta. Folia lanceolata, petiolata, margine serrata; supra pilosa, infra dense pilosa. Receptaculum convexum epaleatum. Phyllaria 3-seriata, dense pilosa; externis linearibus, intimis linearibus-lanceolatis 5 mm. longis, acuminatis apiculatis. Corolla 3.5—5 mm. longa. Antherae basi calcaratae; appendicibus apicalibus ovatis truncatis leviter incisuris. Filamentum collum rectum elongatum. Cypselae 1 mm. longae, 5-costatae, sparsim pilosae. Pappi setae interiores c. 4 mm. longae, barbellatae, albae, caducae. An erect annual herb. Stems branching above, longitudinally ribbed, densely pilose. Leaver lanceolate, petiolate, with serrate margins, pilose above, densely pilose beneath. Receptacle convex, epaleate. Phyllaries c. 3-seriate, densely pilose with very long hairs; outer ones linear, inner ones linear-lanceolate, c. 5 mm long, acuminate- apiculate at tips. Corolla 3.5—5 mm long, very narrowly funnel-shaped; corolla lobe tip with many uniseriate and biseriate stalked glands. Filament collar straight, elongate Comp. Newsl. 26, 1995 with the cells rectangular and uniformly thickened. Anthers calcarate; anther apical appendage ovate with more or less truncate tip, slightly notched in the middle; some of the hairs on the style branches and upper part of shaft short and obtuse at tip; endothecial tissue intermediate with thickenings on both the horizontal and radial walls. Cypselas 1 mm long, 5-costate, turbinate, sparsely pilose with twin hairs having parallel cells of unequal length, few capitate glands and few idioblasts. Pappus without outer scales, inner ones c. 4 mm long, barbellate, white, terete, caducous. Taxonomic note: This variety is different from the other ones because it has anther apical appenage which is ovate with truncate tp slightly notched in the middle (Fig. 3K). The varietal epithet refers to the distinct ovate shape of the apical appendage. Distribution: It occurs probably only in Madagascar, if the location is correct. Swartz’ annotation on the back of the type, "lecta in Madagascar a D. Thunberg” is obviously incorrect, since Thunberg never visited Madagascar. Taxonomic notes on geographically separated populations of C. cinereum var. cinereum (a) Tanganyika Stem branches are densely pilose with long hairs intermingled with long uniseriate ones clustered at nodes and up to 20 cells long. Cypselas are 1.8-2 mm long, densely appressed pilose with twin hairs whose parallel cells are almost equal in length and with acute tips. Pappus with outer scales slightly broader than the inner ones which are terete, barbellate, white, c. 4 mm long, and readily caducous. Specimen examined: Tanganyika, Pangani, Misubugwe forest, Tanner 2190 (S). (b) Malawi The variety in Malawi is differing slightly from the one in Tanganyika by lacking outer pappus scales, and the twin hairs on the cypsela have parallel cells which are of unequal length. These are minor variations which cannot be used to create a new variety. Specimen examined: Malawi, Zomba Dist, Makoka Agric. Resea, Balaka & Usi 1514 (S). (c) Trinidad The twin hairs on the cypsela are similar to the ones in the variety in Tanganyika but different from those found in Malawi because their parallel cells are almost of equal length (Fig. 1C). The anther apical appendage is ovate, much dilated proximally and tapering to the apex and obtuse/acute at tip. Specimen examined: Trinidad, Broadway 7947 (S). 18 Comp. Newsl. 26, 1995 (d) The Philippines This population is similar to the one in Tanganyika. Specimen examined: Philippines, Ramos & Edano 43993 (S). (e) Jamaica The variety in Jamaica is similar to the ones in the Philippines and Tanganyika in all respects. Specimen examined: Jamaica, Yunker 17890 (S). (f) British Guiana Specimen examined: British Guiana, Georgetown, Hitchcock 16595 (S). (g) Zimbabwe The variety in this area is similar to the ones above except that it has no outer pappus and the twin hairs on cypsela have parallel cells which are of slightly unequal length and are acute/obtuse at tips. The cypselas are terete without ribs. Specimen examined: Zimbabwe (S. Rhodesia), Royen 13461 (S). (h) Australia The Australian population is similar to the ones in the Philippines, Jamaica and Tanganyika. Specimen examined: Australia, Eriksson 402 (S). (i) India The population is similar to the one in Zimbabwe with respect to the cypsela which is not ribbed; its cypsela is c. 2 mm _ long. Specimen examined: India, Madras, Keebola' 8197 (S). (j) China The specimen seen is similar to the ones from India, Philippines, etc. Its cypsela is densely pilose, c. 0.6 mm long; inner pappus bristles terete, white, c. 4.5 mm long and readily caducous. Specimen examind: China, Wulsin et al. 12481 (S). Conclusion The species and its varieties are rightly transferred from Vernonia s.l. to the resurrected genus Cyanthillium because of its distinct pollen characteristics and floral microcharacters. Its pollen has no colpus in contrast to the genus Baccharoides (Isawumi & El-Ghazaly in press) and the genus Vernonia s.l. The endothecial tissue is Comp. Newsl. 26, 1995 intermediate unlike the one in Baccharoides which is distinctly polarized (Isawumi & El-Ghazaly in press). The filament collar is elongate and straight unlike the ones in Baccharoides which are more or less shortly cylindrical and sometimes somewhat dilated distally. There are about three varieties in East Africa and three also in Madagascar. The var. lenti (O. Hoffm.) C. Jeffrey reported by Jeffrey (1988) as occurring in Kenya and Tanzania could not be included in this work. The area of greater diversity of the species is therefore probably in East Africa and Madagascar. It is suggested that the species might have originated either in East Africa or Madagascar and spread by introduction to other parts of tropical areas of the World. Hitherto the varieties have been delimited on the bases of the size of inner phyllaries, cypselas and inner pappus bristles (Jeffrey 1988, Pope 1992). It is found in this study that the microcharacters are strongly diagnostic for the varieties and can be easily used along with other characters for their separation and identification. The species is pantropical and found in tropical Asia, India, China, Philippines, Australia, Jamaica, West Indies, South America, and Madagascar. On the African continent it is known from Zimbabwe, Malawi, Mozambique, Zaire, Rwanda, Uganda, Kenya, Tanzania, Tanganyika, Angola, and other parts of Southern Africa, and also in tropical West Africa. It thus occurs throughout the paleotropical region and is widely adventive in the neotropics (Robinson 1990). Acknowledgements I would like to acknowledge the asistance of Prof. B. Nordenstam for reading the manuscript and offering valuable comments. Financial support from the Swedish Institute is also gratefully acknowledged. I am particularly grateful to the staff members of the Botany Department of the Swedish Museum of Natural History for their assistance and cooperation during the course of the study. 20 Comp. Newsl. 26, 1995 References Anderberg, A.A. 1991. Taxonomy and phylogeny of the tribe Gnaphalieae (Asteraceae). Opera Bot. 104. Anderson, L.E. 1954. Hoyer's solution as a rapid mounting medium for bryophytes. Bryologist S57: 242-247. Bremer, K. 1987. Tribal relationships of the Asteraceae. Cladistics 3 (3): 210-253. Dormer, K.J. 1961. The crystals in the ovaries in certain Compositae. Ann. Bot. n.s. 25: 241-254. Dormer, K.J. 1962. The fibrous layer in the anthers of the Compositae. New Phytol. 61:150-156. Fahn, A. 1982. Plant Anatomy, Ed. 3. Pergamon Press, Oxford. Hess, R. 1938. Vergleichende Untersuchungen iiber die Zwillinghaare der Compo- siten. Bot. Jahrb. Syst. 68: 435-496. Isawumi, M.A. 1989. Leaf epidermal studies in the genus Vernonia Schreber tribe Vernonieae (Compositae) in West Africa. Feddes Repert. 100: 335-355. Isawumi, M.A. 1993. New combinations in Bacchariodes Moench (Vernonieae: Compositae) in West Africa. Feddes Repert. 104: 309-326. Isawumi, M.A. & G. El-Ghazaly in press. Pollen morphology, floral micro- characteristics and taxonomy of the genus Baccharoides Moench (Vernonieae: Asteraceae). Grana Palynol. Jeffrey, C. 1988. The Vernonieae in East Tropical Africa. Notes on the Compositae: 5. Kew Bull. 43: 195-277 Karis, P.O. 1993. Heliantheae sensu lato (Asteraceae), clades and classification. Pl. Syst. Evol. 188: 139-195. King, R. M. & H. Robinson 1970. The new synantherology. Taxon 19: 6-10. Kingham, D.L. 1976. A study of the pollen morphology of tropical African and certain other Vernonieae (Compositae). Kew Bull. 31: 9-26. Nordenstam, B. 1968. The genus Euryops. Part II. Aspects of morphology and cytology. Bot. Notiser 121: 209-232. Nordenstam, B. 1978. Taxonomic studies in the tribe Senecioneae (Compositae). Opera Bot. 44. Nordenstam, B. & G. El-Ghazaly 1977. Floral morphology and pollen ultrastructure in some Centaureinae (Compositae) mainly from Egypt. Publ. Cairo Univ. Herb. 7-8: 143-155. Comp. Newsl. 26, 1995 Pope, G.V. 1992. An analysis of the characters and relationships of the tribes Eupatorieae and Vernonieae (Asteraceae). Syst. Bot. 2: 199-208. Robinson, H. 1977. An analysis of the characters and relationships of the tribes Eupatorieae and Vernonieae (Asteraceae). Syst. Bot. 2: 199-208. Robinson, H. 1983. A generic review of the tribe Liabeae (Asteraceae). Smithsonian Contrib. Bot. 54: 1-69. Robinson, H. 1990. Six new combinations in Baccharoides Moench and Cyanthillium Blume (Vernonieae: Asteraceae). Proc. Biol. Soc. Wash. 103: 248- 253: Robinson, H. & B. Kahn 1986. Trinervate leaves, yellow flowers, tailed anthers and pollen variation in Distephanus Cass. (Vernonieae: Asteraceae). Proc. Biol. Soc. Wash. 99: 493-501. Stearn, W.T. 1966. Botanical Latin. Nelson, London. Wagenitz, G. 1976. Was ist eine Achine: zur Geschichte eines karpologischen Begrifffs. Candollea 31: 79-85. Wild, H. 1978. The Compositae of the Flora Zambesiaca area 8—Vernonieae (Vernonia). Kirkia 11: 31-127. Fig.1 A: Fig.2 E: Fig.3 I: - Comp. Newsl. 26, 1995 FIGURE LEGENDS Cyanthillium cinereum var. cinereum in Trinidad - Corolla lobe tip with eglandular trichomes 2-3 cells long and capitate glands, x 230; Endothecial tissue intermediate with both horizontal and radial walls thickened, x 325; Twin hairs with parallel cells almost of equal length, fused almost to the apices and with simple pits at the fused areas; idioblasts on cypsela. x 230; Broadway 7947 (S). C. cinereum var. cinereum in Jamaica —Style base ring-like with thickened cells, x 230; Yuncker 17890 (S). C. cinereum var. ugandense — Corolla lobe tip with T-shaped trichome and capitate glands, x 230 (S); Ovary wall crystals and idioblasts on cypsela, x 230; Taylor 2464 (S). C. cinereum var. cinereum in Australia — Filament collar elongate, narrow and cylindrical, x 230; Anther apical appendage ovate in shape, endothecial tissue intermediate, x 230; Eriksson 402 (S). C. cinereum var. cinereum in the Philippines — Anther apical appendage ovate and tapering towards the apex, x 230; Ramos & Edano 43993 (S); C. cinereum var. viale in Madagascar — Anther apical appendage lanceolate—ovate with obtuse tip, x 80; Perrier de la Bathie 597 (S). C. cinereum var. ovatum in Madagascar — Anther apical appendage ovate with truncate tip slightly notched in the middle, x 230; Thunberg crits): Comp. Newsl. 26, 1995 Comp. Newsl. 26, 1995 sie Comp. Newsl. 26, 1995 25 Fig.3 I—K 26 Comp. Newsl. 26, 1995 STUDIES ON THE REPRODUCTIVE BIOLOGY OF VERNONIA SCHREB. (ASTERACEAE) III. Floret shapes/number, pappus types and surface structure among different growth habits M.S. Ayodele * Botany Department Obafemi Awolowo University Ile-Ife, Nigeria Abstract Floral attributes displayed low variability within and among species of Vernonia, and are thus considered of great diagnostic value in taxonomic evaluation. Floral dimorphism in one of the species studied (V. biafrae) constitutes a noteworthy variant of the usual reports on sexual characters of species in the genus Vernonia. Pappus surface structures give additional evidence as regards their function in achene (fruit) dispersal. The surface structure of pappus (i.e., absence, presence and number of bristles) among the species, is of value recommendable for growth habit delimitation within the genus. Introduction The capitula of the species of the genus Vernonia are reported to show interspecific variation in floret shapes. The florets are reported to be bisexual in species described (Hutchinson & Dalziel 1973, Keay et al. 1964, Jones 1976). The fruits (achenes) are crowned by a pappus composed of numerous bristly hairs. Pappus hairs are either uniform or variable in length in different species (Keay et al. 1964, Jones 1976, Isawumi 1982, Ayodele 1987). The pappus is one of the morphological attributes useful for taxonomic evaluation of the genus (Faust 1972, Jones 1973, Isawumi 1982). The objective of this paper is to report the observed variations in floret shapes and sexual characters of the flowerheads of some Nigerian species of Vernonia. This paper will also report observed variations in the pappus, including details of surface structures, among the different growth forms studied. These will enhance useful deductions on certain aspects of reproductive efficiency in the genus Vernonia. * Current mailing address: P.O.Box 586, Ilesa, Nigeria Comp. Newsl. 26, 1995 27 Materials and Methods Capitula at anthesis were harvested randomly from Vernonia plants among field populations, garden and screen house plants. At least five capitula from five plants of each species were used during each investigation of floret morphology and assessment of number of florets per capitulum. Each capitulum was dissected by means of a pair of forceps and a mounted needle to detach the florets from the receptacle. Floret counts were taken, using a tally-counter. The structures of the pappus hairs were observed under a light microscope. A slide mount of pappus hair in water was made and observed under scanning power of a photomicrographic equipment. Photographs of the pappus hair surface structure were taken for each of the species. Drawings were made of floret shapes and fruits with pappus crown. Means of floret counts and coefficient of variation were estimated for the different species. The floret count values were analysed by Duncan's multiple range test for significant differences among the species. Observations/Results There were significant differences among some of the species in the number of florets contained in each capitulum. The 16 species investigated formed 10 groups of non- significant differences in floret counts (i.e. within group). Of the 10 groups however, there were 6 distinctly separable groups, while the other 4 groups overlapped (Table 1). Each of the distinct groups contained mainly one growth form (a shrub or a herb), whereas the overlapping groups contained arboreal, shrubby and herbaceous forms (Table 1). Generally, the species with a large-size capitulum contained higher number of florets, irrespective of the growth form. The overlapping groups contained mainly those species with small-size capitulum from among the different growth forms. The variability in floret counts within each species was observed to be low for the different growth forms. Vernonia florets in all the specimens from the different locations were tubular in shape (Fig. 1). Florets were generally hermaphroditic except in V. biafrae (Fig. 1:8), which had mainly ligulate female florets and a few (1-3) hermaphroditic florets in the capitulum. The style of the florets was bifurcated in all the species. The bifurcated stigma, wrapped in fused anther thecae, was exserted in some species (Fig. 1). The style branches were long and recoiled in some species, and usually dusted with a mass of pollen grains, even at anthesis (Fig. 4, 5 & 6). Among the different species, the pappus hairs showed different orientations (Fig. 2A & B). For example, on broad and heavy fruits (Fig. 2A:4 & 5) there was a 180° 28 Comp. Newsl. 26, 1995 spreading orientation, like a fully-opened hand-wave fan. On other, lighter and narrower fruits, the pappus was either non-spreading (Fig. 2B:13) or showed the reduced-spreading "shuttle-cock" pappus type (Fig. 2B:12). Photomicrographs (100x) of pappus hairs showed projections (teeth) of various shapes and sizes, on the superficially smooth-looking surfaces (Fig. 3 A & B). The number of such teeth or projections and their frequency on each hair varied among the species. The distribution of pappus teeth along each hair also varied among the growth forms of the species investigated. The herbaceous species clearly had more projections on each pappus hair (Fig. 3B). These were spine-like and well distributed over the entire length of the hair (Fig. 3B: 11, 12 & 13). The pappus of the herbs thus was of the plumose type. The shrubby species had fewer teeth (Fig. 3A: 4, 6 & 7) especially located on the distal portions of the hair (Fig. 3A:4). Thus a capillary form of pappus was prevalent among the shrubs. Fig. 3A & B also show the distinct differences in the relative sizes of pappus hair for the species studied. Discussion Although there were significant differences in floret counts among the species studied, the variation of this attribute within each species was relatively low, when compared with other morphological traits. The variation was particularly low for those species with overlapping values (Table 1) irrespective of the growth form. The small-size capitula plants from among all growth forms particularly had low variability in floret counts. The low variability trend in the number of florets per capitulum is noteworthy. Burtt (1977) noted that the constancy of the characteristics of florets, in spite of variation occurring in capitulum character, was one reason for the outstanding success in the Compositae. The attributes of the florets are, therefore, emphasized by plant taxonomists in classification and identification of the Compositae (Burtt 1977). The florets of Vernonia generally have characteristics which complement other reproductive strategies among the species. A good example is the compensatory roles of small-size capitulum/low number of florets, but with larger clusters of capitula on the plants. This compared with the large-size capitulum with numerous florets, but lesser capitula clustering (Ayodele 1992). The floral dimorphism observed in V. biafrae contradicts previous reports of only bisexual florets in Vernonia (Hutchinson & Dalziel 1973, Keay et al. 1964). Heywood et al. (1977) noted a correlation between pappus structure and growth habit of some members of the family Compositae. However, Vernonia was not one of the genera mentioned. They highlighted the increase in the proportions of capillary or plumose pappus in progression from annual to perennial herbs to shrubs. This report Comp. Newsl. 26, 1995 29 by Heywood et al. (1977) is in conformity with the observed variations, as they relate to the growth habits of the species of Vernonia (Fig. 3A & B). The herbs which produced large-size fruits had more projections on the pappus hair than the shrubby species with large fruits. In herbs, the entire length of the pappus hair carries projections. In shrubs, the lower portions of pappus hair have scanty teeth or are totally smooth (Fig. 3B:11 & 16 compared with Fig. 3A: 4, 5 & 6). The presence or absence of projections and distribution along the pappus hair is recommendable as a useful tool for delimitation of growth forms in the genus Vernonia. The role of projections (and indeed the pappus hair on which they are located) in fruit dispersal, can be deduced from the observed interlocking device of pappus hairs on two adjacent fruits on a receptacle (Fig. 4). Only the upper and middle part of the pappus hairs are interlocked in V. tenoreana, a shrub. In V. galamensis, a herb, the pappus hairs are interlocked along the entire length (Fig. 4A & B). Projections are absent in the lower portion of pappus hair in V. tenoreana (Fig. 3A:4). The amount of interlocking was observed to be related to the number of fruits dispersed together when shed from the receptacle. Fruits were dispersed either solitarily or in groups of 2-5 fruits with interlocked pappus hairs, for the large-size fruits. The small-size, light-weight fruits formed a cottony mass of tangled pappus for a larger group dispersal unit of fruits. In some species (e.g., V. biafrae) a sticky oily substance was observed in such a mass of tangled fruits, which are usually dispersed by wind. 30 Comp. Newsl. 26, 1995 References Ayodele, M.S. 1987. Cytological and morphological studies on some species of Vernonia Schreb. in Nigeria. M.Sc. Thesis, Obafemi Awolowo University, Ile-Ife, Nigeria. 98 pp. Ayodele, M.S. 1992. Cytogenetic and reproductive studies on some species of Vernonia Schreb. (Asteraceae) in Nigeria. Ph.D. Thesis, Obafemi Awolowo University, Ile-Ife, Nigeria. 290 pp. Burtt, B.L. 1977. Aspects of diversification in the capitulum. Jn: Heywood, V.H., Harborne, J.B. & B.L. Turner (eds.), The Biology and Chemistry of the Compositae 1: 41-59. Academic Press, London & New York. Faust, W.Z. 1972. A biosystematic study of the Interiores species group of the genus Vernonia (Compositae). Brittonia 24: 363-378. Heywood, V.H., Harborne, J.B. & B.L. Turner 1977. An overture to the Compositae. Jn: Heywood, V.H., Harborne, J.B. & B.L. Turner (eds.), The Biology and Chemistry of the Compositae 1: 1-20. Academic Press, London & New York. Hutchinson, J. & J.M. Dalziel 1963. Flora of West Tropical Africa, 2nd ed. revised by F.N. Hepper, 2: 271-283. Crown Agents, London. Isawumi, M.A. 1982. A numerical taxonomic study of the genus Vernonia and hybrids, Tribe Vernonieae (Compositae), in West Africa. Ph.D. Thesis, University of Ife, Ile-Ife, Nigeria. 203 pp. Jones, S.B. Jr 1973. Revision of Vernonia Section Eremosis (Compositae), in North America. Brittonia 24: 86-115. Jones, S.B. Jr 1976. Cytogenetics and affinities of Vernonia (Compositae) from the Mexican highlands and eastern North America. Evolution 30: 455-462. Keay, R.W.J., Onochie, F.C.F.A. & D.P. Stanfield 1964. Nigerian Trees. 2: 419- 430. Fed.Dept. of Forest Res. Ibadan, Nigeria. Comp. Newsl. 26, 1995 31 Table 1. Number of florets per capitulum in some species of Vernonia Species grouping* from Duncan's i Range | CV%** multiple range test V. galamensis var. ethiopica V. kotschyana V. tenoreana V. stenostegia V. purpurea V. galamensis (= pauciflora) V. ambigua V. perrottetii V. biafrae V. migeodi V. cinerea V. colorata V. nestor V. amygdalina V. conferta V. glaberrima Note: * Means with the same letter are not significantly different ** Coefficient of variation of attribute. + Values are for bisexual tubular and unisexual ligulate female florets respectively. 32 Comp. Newsl. 26, 1995 Table 2. Analysis of variance (ANOVA) in number of florets per capitulum of Vernonia Character | Source of Degree of | Mean variation po fa = or square S Number of 1092388.36 72825.89 | 613.88 * florets per capitulum 45554.80 384; 118.631 in Vernonia 1137943.16 * Significant at 0.05% level. LSD (0.05) = 6.057. Comp. Newsl. 26, 1995 Fig. 1 Drawings showing variation in shape of florets in some species of Vernonia. FIGURE LEGENDS Key: S = stigma; O= ovary; a.l. = anther lobe 1. ‘V. conferta Benth. 2. V. amygdalina Del. 3. V. colorata (Willd.) Drake 4. V. tenoreana Oliv. 5. V. stenostegia (Stapf) Hutch. & Dalz. 6. _ V. kotschyana Sch. Bip. 7. V. glaberrima Welw. ex O. Hoffm. 8. V. biafrae Oliv. & Hiern 9. V. migeodi S. Moore 10. V. cinerea (Linn.) Less. 11. V. galamensis (=pauciflora) (Cass.) Less. 12. V. ambigua Kotschy & Peyr. 13. V. perrottetii Sch. Bip. 14. V. nestor S. Moore 15. V. purpurea Sch. Bip. Fig. 2A & B Fruit shapes and pappus orientation on fruits of some species of Vernonia. A: Tree and shrubby species. B: Herbaceous species. Key to numerals as in Fig. 1. Fig. 3A & B Photomicrographs of surface structure of pappus hairs of some species Fig. 4 of Vernonia. A: Tree and shrubby species. B: Herbaceous species. Key to numerals (1-15) as in Fig. 1, plus 16 = V. galamensis var. ethiopica (Cass.) Less. & Gilb. Fruit dispersal in Vernonia. Pappus hair interlocking device for corporate fruit dispersal. A. V. tenoreana (shrub). B: V. galamensis (herb). C: Cottony mass of fruits in V. biafrae. 33 Comp. Newsl. 26, 1995 35 Comp. Newsl. 26, 1995 Fig. 2B 37 Comp. Newsl. 26, 1995 Fig. 3A 38 Fig. 3B Comp. Newsl. 26, 1995 Comp. Newsl. 26, 1995 39 40 Comp. Newsl. 26, 1995 ETUDE ANATOMIQUE PRELIMINARE DES AKENES DE QUELQUES ARTEMISIA DANS LE BASSIN MEDITERRANEEN OCCIDENTAL A. Ouyahya Département de Botanique et écologie végétale Institut Scientifique Rabat, Maroc Résumé L'auteur décrit l'anatomie des akénes de quelques Artemisia en Méditerranée occidentale et discute spécialement la corrélation qui pourrait exister entre la taille, l'absence ou la présence des cellules épidermiques gélifiées et les conditions climatiques du biotope, ainsi que la valeur taxinomique des cellules gélifiées. Abstract The author describes the anatomic structure of the achenes of Artemisia in the west Mediterranean basin and specially discusses the probable correlation between the size, presence or absence of the mucilaginous epidermic cells and the climatic conditions in the biotope. Besides, the taxonomic value of the mucilaginous cells is discussed. Introduction Cette étude fait suite 4 deux articles antérieurs (Ouyahya & Viano 1984 & 1990), traitant les caractéristiques morphologiques et biométriques des akénes des taxons marocains et a titre de comparaison les taxons affines du bassin méditerranéen occidental. Nous avons entrepris l'étude anatomique des akénes des armoises étudiées en nous basant sur le travail de Korobkov (1973). Celui-ci constitue la meilleure contribution a l'étude carpologique de certaines espéces du genre Artemisia au nord-est de l'ex URSS. Cet auteur a suggéré que, sur la base de la morphologie et de l'anatomie de I'akéne, la différenciation spécifique est tout a fait remarquable. Comp. Newsl. 26, 1995 Matériel et méthode Comme Korobkov (1973), nous avons procédé a des coupes transversales sur plusieurs akénes par taxon. Les coupes ont été effectuées 4 main levée sur les akénes secs, préalablement trempés dans un mélange d'eau, de glycérine et d'alcool éthylique (1/1/1 V). Les coupes ont été ensuite traitées de la fagon suivante: eau de javel (15 min.); ringage a l'eau; eau acétique 4 5 % (5S min.); carmino-vert (10 min.); rincage a l'eau; montage dans la glycérine. Cette méthode a donné de bons résultats pour les akénes dépourvus de cellules gélifiées tels que ceux de A. verlotorum et A. vulgaris. Mais pour les akénes qui ont des cellules gélifiées qui se déchirent en présence d'eau dans les réactifs, nous avons réalisé des coupes sur des akénes secs, puis les coupes ont été montées directement dans la glycérine pure. Résultats et discussion A l'observation au microscope photonique, la paroi des akénes apparait relativement homogéne au sein des taxons analysés. Cette paroi se compose des parties suivantes: - un péricarpe (paroi du fruit), comprenant un épiderme a une seule couche de cellules gélifiées ou non, a cuticule plus ou moins mince et de rares faisceaux libéro-ligneux. En présence de l'eau, les cellules épidermiques se gélifient, se gonflent tout en demeurant trés réfringentes. Ces cellules jouent un réle physiologique important dans l'absorption de l'eau, au moment de la germination, tout en formant une couche de mucilage plus ou moins épaisse (selon le degré de gélification). - un tégument (paroi de la graine) dont l'Epiderme externe (épitesta) est plus ou moins développé, alors que la couche interne est presque complétement résorbée. - un albumen réduit 4 une seule assise de cellules protéiques, allongées dans le sens tangentiel. Les résultats obtenus ont été consignés dans le tableau 1. On y distingue trois groupes, suivant la présence ou l'absence des cellules gélifiées et leur taille. - Le premier groupe comprend les akénes 4 paroi pourvue de grandes cellules gélifiées (> 50 um d'épaisseur). Ce type d'akénes caractérise A. herba-alba (Fig. 2a). - Le second groupe renferme la majorité des espéces étudiées dont les akénes possédent, en effet, des cellules gélifiées de petite taille (< 20 um d’épaisseur) (Figs.la a 4d) ou de taille moyenne (a épaisseur comprise entre 20 et SO um) (Figs. 2b-2c et 3a a 3d). - Les espéces du troisiéme groupe se distinguent facilement des akénes dépourvus de cellules gélifiées (Figs. 4a 4 4c), c'est le cas par exemple de: A. verlotorum, A. vulgaris, A. campestris subsp. lloydii, A. crithmifolia et A. alba subsp. chitachensis. 41 42 Comp. Newsl. 26, 1995 Korobkov (1973) avait établi le méme type de découpage et avait corrélé l'aptitude a la gélification, avec la répartition géographique et les conditions climatiques des armoises russes étudiées. Selon cet auteur les espéces 4 akénes trés gélifiés occupent essentiellement les régions séches (espéces continentales), celles 4 akénes moins gélifiées croissent dans des lieux bien drainés et le troisitme type 4 akénes dépourvus de cellules gélifiées est représenté par des espéces ocAniques ou subocéaniques. Pour les Artemisia marocains, la corrélation est du méme ordre: chaque groupe est corrélé avec l'aridité du biotope. Cepedant, l'adhérence du péricarpe 4 la graine confirme la subdivision du genre Artemisia en deux unités distinctes (Artemisia et Seriphidium) dont le rang sys- tématique est discutable. Ce résultat recoupe ceux que fournissent la nature des capi- tules, d I'indument etc. (Ouyahya 1987). Ce caractére s'ajoute a l'absence ou la pré- sence des cellules gélifiées s'avérent un excellent critére de discrimination (Tableau 1). Comp. Newsl. 26, 1995 Tableau 1. Tableau résumant les données relatives aux caractéristiques anatomiques des akénes des Artemisia étudiés (los coupes anatomiques trans- versales ont été faites sur 30 akénes par taxon). Epaisseur (ép.) en jum. Taxon étudiés Paroi du fruit Cellules gélifiées (péricarpe) nb. de absentes | présentes | présen' présentes couches de ép.>50 | 20<ép.<50} ép.<20 cellules A. absinthium A. alba subsp. chitachensis A. atlantica vat. maroccana A. campestris subsp. glutinosa subsp. lloydii crithmifolia dracunculus flahaultii judaica subsp. sahariensis mesatlantica negrei verlotiorum vulgaris herba-alba ifranensis A. A. A. A. A. A. A. reptans A. A. A. A. A. caerulescens subsp. gallica A. vallesiaca 44 Comp. Newsl. 26, 1995 Bibliographie Benizri, E. 1964. Contribution a l'étude carpologique de l'espéce Ormenis praecox (Link) Briquet au Maroc. D.ES., Fac. Sci. Rabat, 47 pp., 22 fig. Briquet, J. 1916. Etude carpologique sur les genres de Composées Anthemis, Ormenis et Santalina. Ann. Cons. Jard. Bot. Genéve 18-19: 257-313. Korobkov, A.A. 1973. Caractéristiques morphologique et anatomique des akénes de Artemisia sp. du Nord-Est de I'URSS. Bot. Zh. SSSR. 58 (9): 1302-1315. Ouyahya, A. 1987. Systématique du genre Artemisia L. au Maroc. Doctorat d'état és- Sciences, Aix-Marseille III, 433 pp. Ouyahya, A. & J. Viano 1984. Contribution 4 l'étude morphologique et biométrique des akénes de taxones endémiques marocains du genre Artemisia. Lagascalia 12 (2): 223-228. Ouyahya, A. & J. Viano 1990. Etude au MEB du testa des akénes du genre Artemisia. Bol. Soc. Brot., ser.2, 63: 99-113. Légendes des figures Fig. 1 Coupes anatomiques transversales des akénes de Artemisia: a- A. absinthium, b- A. campestris subsp. glutinosa, c- A. mesatlantica, d- A. judaica subsp. sahariensis. I- péricarpe de l'akéne; II- tégument de la graine; IlI- albumen. Faisceaux conducteurs (f.con.); épiderme externe (€p.ext.); épiderme interne (ép.int.) du tégument de la graine; cellules gélifiées (c.gel.). Fig. 2 Coupes anatomiques transversales des akénes de Artemisia. a- A. herba-alba, b- A. negrei, c-A. flahaultii. Fig.3 Coupes anatomiques transversales des akénes de Artemisia: a- A. ifranensis; b- A. atlantica var. maroccana; c- A. dracunculus, d- A. reptans. Fig.4 Coupes anatomiques transversales des akénes de Artemisia: a- A. alba subsp. chitachensis, b- A. verlotiorum, c- A. crithmifolia. I- péricarpe de I'akéne; II- tégument de la graine; IlI- albumen. Faisceaux conducteurs (f. con.); c6te (co.); cuticule (cu.); couche sous-€pidermiques (c.ss.ép.); piderme exteme (€p. ext.); épiderme interne (ép.int.) du tégument de la graine. 45 Comp. Newsl. 26, 1995 Fig. 1 Comp. Newsl. 26, 1995 Fig. 2 Comp. Newsl. 26, 1995 47 Fig. 3 Fig. 4 Comp. Newsl. 26, 1995 Comp. Newsl. 26, 1995 49 SURVEY OF ARTEMISIA AND SERIPHIDIUM (ASTERACEAE-ANTHEMIDEAE) IN THE HIMALAYAN MOUNTAINS AND THE SOUTH ASIAN SUBCONTINENT Ling Yeou-ruenn South China Institute of Botany Academia Sinica Guangzhou, 510650, P. R. China The author has checked specimens cited in Clarke's Compositae Indicae, or in Hooker's Flora of British India vol. 3 and in Materials of Artemisia Linn., sensu lato, in Asia by R. Pampanini. Also collections from South Asia have been studied, either in the Herbaria of Kew Garden (K) incl. the Wallich Herbarium, British Museum of Natural History, incl. Clifford specimens, which were cited by Linnaeus (BM), Linnaeus Herbarium in Linnean Society (LINN), the Botany School of Cambridge University (CGE), Edinburgh Botanic Garden (E), or Paris Museum of Natural History (P), Rijksherbarium, Leiden (L) and Gray Herbarium of Harvard University, USA (GH), and a few from India *) **). Specimens from these herbaria are cited excl. some newly recorded species in Flora Xizangica vol. 4. A re-collation and a revision of this material is presented. I. Artemisia Linn., Sp. Pl. 2: 845 (1753) et Gen. Pl. ed. 5: 367 (1754) sencu stricto, excl. sect. Seriphidium Bess. 1. Subgen. Artemisia 1). Sect. Absinthium DC. A. macrocephala Jacq. ex Bess. in Bull. Soc. Nat. Mosc. 9: 28 (1836); Hook. f., Fl. Brit. Ind. 3: 329 (1881); Nasir et Ali, Fl. W. Pakistan 717 (1972). — A. griffithiana Boiss., Fl. or. 3: 376 (1875). N. India, Kashmir, Nepal, China (N., N.-W., N.-E. et S.-W.), Iran, Mongolia, Afghanistan, Kazakhstan, Kyrgyzstan, Tadjikistan, Russia. (Specimens in BM, E, K, P): A. sieversiana Ehrh. ex Willd., Sp. Pl. 3: 1845 (1800); C.B. Clarke, Comp. Ind.: 165 (1876); Hook. f., Fl. Brit. Ind. 3: 329 (1881); Nasir et Ali, Fl. W. Pakistan 719 (1972). *) Lucknow, Central Institute of Medicinal and Aromatic Plants, Inda. **) Howrah, National Herbarium, India. 50 Comp. Newsl. 26, 1995 — A. persica sensu Schlagint. Cat. [671, 988 (1856), (BM, K)], non Boiss. — A. laciniata sensu Schlagint. Cat.{1116, 1307, 1527, 2285, (BM, E, K, L)] — A. macrantha auct., non Ledeb.: Pamp. in Nuov. Giorn. Bot. Ital. n.s. 34: 677 (1927); p.pquoad Pl. Ind. — Absinthium sieversianum (Ehrh. ex Willd.) Bess. in Bull. Soc. Nat. Mosc. 1 (8): 259 (1829). N. India, Kashmir, N. Pakistan, Nepal, China (N., N.-W., N.-E. et S.-W.), N. Japan, N. Korea, Mongolia, Afghanistan, Kazakhstan, Kyrgyzstan, Tadjikistan, Turkmenistan, Uzbekistan, E. Europ. (BM, CGE, E, GH, K). A. absinthium Linn., Sp. Pl. 2: 848 (1753); C. B. Clarke, Comp. Ind. 164 (1876); Hook. f., Fl. Brit. Ind. 3: 328 (1881); Nasir et Ali, Fl. W. Pakistan 714 (1972). — A. kulbadica Boiss. et Buhse in Nouv. Mém. Soc. Nat. Mosc. 12: 120 (1860). — Absinthium vulgare Lamarck, Fl. Fr. 2: 45. 1778. N. India, Kashmir, N. Pakistan, China (N.-W. & Xizang), Afghanistan, Iran, Kazakhstan, Kyrgyzstan, Russia, Europe, N. et W. Africa, N. America. (BM, CGE, E, K). A. rutifolia Steph. ex Spreng., Syst. Veg. 3: 488 (1826); Nasir et Ali, Fl. W. Pakistan 718 (1972). — A. falconeri C. B. Clarke ex Hook. f., Fl. Brit. Ind. 3: 328 (1881). — A. turczaninowiana Bess. var. falconeri (C. B. Clarke) O. Fedtsch. in Act. Hort. Petrop. 2A: 322 (1905). Kashmir, Nepal, N. Pakistan, China (N.-W. & Xizang), Afghanistan, Iran, Mongolia, Kazakhstan, Kyrgyzstan, Tadjikistan, Russia. (BM, K, P). A. minor Jacq. ex Bess. in Bull. Soc. Nat. Mosc. 9: 22 (1836); Hook. f., Fl. Brit. Ind. 3: 329 (1881); Nasir et Ali, Fl. W. Pakistan 717 (1972). — A. sieversiana Ehrh. ex Willd. var. tibetica C.B. Clarke, Comp. Ind. 165 (1876). — A. tibetica Hook. f. et Thoms. ex Hook. f., l.c. : 329 (1881) pro syn. N. India, N. Pakistan, Kashmir, Sikkim, China (Xizang), Iran, Tadjikistan. (BM, E, GH, K). A. stracheyi Hook. f. et Thoms. ex C. B. Clarke, Comp. Ind. 164 (1876); Hook. f., Fl. Brit. Ind. 3: 328 (1881); Nasir et Ali, Fl. W. Pakistan 720 (1972). Kashmir, N. Pakistan, China (Xizang). (T.T., Type, Tibet Occ., CGE, K). A. younghusbandii J.R. Drumm. ex Pamp. in Nuov. Giom. Bot. Ital. n.s. 34: 708 (1927). China (Xizang). (K, L). A. persica Boiss., Diagn. ser. 1, 1 (6): 91 (1845); C. B. Clarke, Comp. Ind. 165 (1876); Hook. f., Fl. Brit. Ind. 3: 327 (1881); Nasir et Ali, Fl. W. Pakistan 718 (1972). Kashmir, N. Pakistan, China (Xizang et Qinghai), Afghanistan, Iran, Tadjikistan (BM, E, K). Comp. Newsl. 26, 1995 51 2). Sect. Abrotanum Bess. sensu stricto (excl. sect. Artemisia) A. macrantha Ledeb. in Mém. Acad. Sci. Petersb. 5: 573 (1815). — A. macrantha Ledeb. var. turtzchaninoviana (Bess.) Pamp. et var. occidentalis Pamp. in Nuov. Giorn. Bot. Ital. n.s. 34: 677-678 (1927). N. India, China (N.-W. et N.), Mongolia, Tadjikistan, Turkmenistan, Uzbekistan, Russia. A. sacrorum Ledeb. in Mém. Acad. Sci. Petersb. 5: 571 (1815); C. B. Clarke, Comp. Ind. 160 (1876); Hook. f., Fl. Brit. Ind. 3: 326 (1881); Nasir et Ali, Fl. W. Pakistan 719 (1972). — A. annua auct., non Linn.: Pall. Reise 2: 523 (1773). — A. gmelinii auct., non Web. ex Stechm.: Poljak. Fl. URSS 26: 464 (1961). var. 1. sacrorum N. India, Nepal, N. Pakistan, Kashmir, China, Mongolia, Korea, Japan, Afghanistan, Tadjikistan, Russia, Kazakhstan,m Kyrgyzstan. (BM, E, K, GH, Lucknow, Howrah). var. 2. messerschmidtiana (Bess.) Y.R. Ling, comb. nov. — A. messer- schmidtiana Bess. in Nouv. Mém. Soc. Nat. Mosc. 3: 27 (1834). N. India, N. Pakistan, China, Afghanistan, Mongolia, Korea, Japan, Russia. (K). A. gmelinii Web. ex Stechm., Dissert. Artem. 30 (1775); Hook. f., Fl. Brit. Ind. 3: 330 (1881). — A. santolinaefolia Turcz. ex Bess. in Nouv. Mém. Soc. Nat. Mosc. 3: 87 (1834). — A. laciniata auct., non Willd.: Hook. f., l.c.: 326 (1881); Nasir et Ali, Fl. W. Pakistan 716 (1972). — A. sacrorum Ledeb. var. santolinaefolia Pamp. in Nuov. Giorn. Bot. Ital. n.s. 34: 693 (1927), incl. formae et subformae. — A. sacrorum sensu Schlagint., Cat.[11738 (1856), Tibet, (E)], non Ledeb. N. India, Kashmir, Nepal, China (N.-W. et S.-W.), Mongolia, Kazakhstan, Kyrgyzstan, Tadjikistan, Uzbekistan, Ukrain, Russia. (BM, E, GH, Lucknow). A. vestita Wall. ex Bess. in Nouv. Mém. Soc. Nat. Mosc. 3: 25 (1834); C. B. Clarke, Comp. Ind. 160 (1876); Hook. f., Fl. Brit. Ind. 3: 326 (1881. — A. laciniata auct., non Willd.: C.B. Clarke, Comp. Ind. (1878). — A. sacrorum Ledeb. var. minor Ledeb. f. wallichiana Pamp. in Nuov. Giorn. Bot. Ital. n.s. 34: 690 (1927). N. India, Kashmir, N. Pakistan, China (S.-W.). (Wallich Cat. 3301/411, Type, Kumaon, BM, CGE, E, GH, K, L). A. carvifolia Buch.Ham. in Roxb., Hort. Beng. 61 (1814), nom. nud., Fl. Ind. 3: 422 (1832); C.B. Clarke, Comp. Ind. 162 (1876) ("caruifolia”); Hook. f., Fl. Brit. Ind. 3: 324 (1881). —— A. apiacea Hance in Walp., Ann. Bot. Syst. 2: 895. (1852). — A. carvifolia Buch.Ham. ex Roxb. var. apiacea (Hance) Pamp. in Nuov. Giorn. Bot. Ital. n.s. 34: 648 (1927). N. India, Nepal, China (E., C., S. et S.-W.), Burma, Indo-China, Korea, Japan, [Wallich Cat. 3299/409 (1824), Type, Bengal, BM, CGE, K)]. 52 Comp. Newsl. 26, 1995 A. annua Linn., Sp. Pl. 847 (1753); Hook. f., Fl. Brit. Ind. 3: 323 (1881); Nasir et Ali, Fl. W. Pakistan 715 (1972). — A. wadei Edgew. in Trans. Linn. Soc. Bot. 20: 72 (1846). — A. stewartii C. B. Clarke, Comp. Ind. 163 (1876). — A. tournefortiana sensu Schlagint., Cat. [2670. (N.-W. Ind.), (E)] non Reichb. Widely distr. N. Hemisphere and N. Africa. (BM, E, GH, K, Lucknow). A. hedinii Ostenf. et Pauls. in S. Hedin, S. Tibet 6 (3): 41, pl. 3. fig. 1 (1922). — A. biennis auct., non Willd.: Hook. f., Fl. Brit. Ind. 3: 325 (1881); Nasir et Ali, Fl. W. Pakistan 715 (1972). N. India, N. Pakistan, Nepal, Sikkim, Kashmir, China (N.-W. et S.-W.), Tadjikistan (BM, E, GH, K, P). A. pallens Wall. ex Bess. in Nouv. Mém. Soc. Nat. Mosc. 3: 85 (1834). N.-E. India, N. Thailand. (BM, K). A. tournefortiana Reichb., Ic. Exot. Cent. 1: 6, tab. t. (1827); C. B. Clarke, Comp. Ind. 163 (1876); Hook. f., Fl. Brit. Ind. 3: 324 (1881); Nasir et Ali, Fl. W. Pakistan 720 (1972). — A. pinnatifida Jacq. ex DC., Prodr. 6: 120 (1837) pro syn. N. Pakistan, Kashmir, China (N.-W. et S.-W.), Afghanistan, Iran, Mongolia, Kazakhstan. (BM, CGE, J. Gay (1833), Type, GH, K). A. filiformilobulata Y R. Ling et H. S. Puri in Guihaia 5 (1): 1, fig. 1 (1985). N. India, (N.C. Shah 266a, Type CIMAP, Lucknow). 3). Sect Artemisia A. austro-yunnanensis Ling et Y.R. Ling in Bull. Bot. Res. 4 (2): 20, fig. 4: 10-18 (1984). — A. burmanica Pamp. f.latiloba Pamp. in Nuov. Giorn. Bot. Ital. n.s. 33: 455 (1925). — A. dubia Wall. ex Bess. var longeracemulosa Pamp. f. tonkingensis Pamp., Lc. 36: 439 (1930). N.-E. India, China (S.-W.), N. Thailand, N. Burma, Indo-China (K). A. lavandulaefolia DC., Prodr. 6: 110 (1837). — A. vulgaris auct., non Linn.: C.B: Clarke, Comp. Ind. 161 (1876) p.p.; Hook. f., Fl. Brit. Ind. 3: 325 (1881) p.p. quoad syn.— A. dubia Wall. ex Bess. var multiflora Pamp. in Nuov. Giorn. Bot. Ital. n.s. 36: 382 (1929) p.p., quoad pl. Ind. Occ. W. India (L, P). A. verlotorum Lamotte in Mém. Assoc. Franc. Cong. Clerm.—Ferr. 511 (1879). — A. leptostachya auct., non D. Don: DC., Prodr. 6: 113 (1837). — A. vulgaris auct., non Linn.: C. B. Clarke, Comp. Ind. 161 (1876); quoad var. vulgaris p.p.; Hook. f., Fl. Brit. Ind. 3: 325 (1881), quoad syn. — A. leptostachya DC.: Nasir et Ali, Fl. W. Pakistan 720 (1972) p.p. — A. dubia Wall. ex Bess. var. legitima (Bess.) Pamp. f.genuina Comp. Newsl. 26, 1995 53 Pamp. sf. puberula Pamp., f. communis Pamp. sf. intermedia Pamp., var. acuminata Pamp. f. leptostachya (DC.) Pamp. et var. orientalis Pamp. in Nuov. Giorn. Bot. Ital. n.s. 36: 435-438 (1930). — A. dubia auct., non Wall. ex Bess.;Kitam. in Act. Phytotax. Geobot. 23 (1-2): 11 (1968), quoad pl. Nepal. — A. indica auct., non Willd.: Kitam. 1.c. 23 (1-2): 12 (1968), quoad pl. Nepal. et Bhutan. Widely distr. Eurasia (except North) et E. et N. Africa. (BM, E, K). A. velutina Pamp. in Nuov. Gior. Bot. Ital. n.s. 36: 413 (1930). — A. dubia Wall. ex Bess. var. legitima (Bess.) Pamp. f. communis Pamp. sf. intermedia Pamp., 1.c. 32: 449 (1926). Kashmir, N.-E. India, China (S.-W., C. et S.) (BM). A. roxburghiana Wall. ex Bess. in Bull. Soc. Nat. Mosc. 9:57 (1836); Hook. f., FI. Brit. Ind. 3: 326 (1881); Nasir et Ali, Fl. W. Pakistan 718 (1972). — A. hypoleuca auct., non MB.: Edgew. in Trans. Linn. Soc. 20: 71 (1846); C.B. Clarke, Comp. Ind. 164 (1876). — A. revoluta Edgew., |. c. 20: 72 (1846). — A. dubia Wall. ex Bess. var. jJacquemontiana Pamp. in Nuov. Giorn. Bot. Ital. n.s. 33: 454 (1926) p.p., incl formae. — A. eriocephala Pamp., l.c. : 454 (1926). — A. indica Willd. var. exilis Pamp., 1.c.: 33: 459 (1926). — A. roxburghiana Wall. ex Bess. var. acutiloba Pamp. op. c. 36: 29 (1930), incl. f. vestita Pamp. var. 1. roxburghiana India, Nepal, Kashmir, China (S.-W. et N.-W.), Afghanistan, Thailand, (BM, CGE, K, Lucknow). var. 2. grata Hook. f., l.c. : 326 (1881); Nasir et Ali, l.c. 718 (1972). — A. grata Wall. ex Bess. in Mém. Soc. Nat. Mosc. 3: 57 (1834). — A. dubia Wall. ex Bess. var. myriantha (Wall. ex Bess.) Pamp., l.c.: 450 (1926) p.p. N. India, Nepal, N. Pakistan. (K). var. 3. purpurascens (Jacq. ex Bess.) Hook. f., l.c. 3: 326 (1881); Nasir et Ali, lc. 718 (1972). — A. purpurascens Jacq. ex Bess., op.c. 9: 60 (1836). — A. strongylocephala Pamp., op. c. 34: 176 (1927). N. India, Nepal, N. Pakistan, Kashmir, China (S.-W.). (K). A. robusta (Pamp.) Ling et Y.R. Ling, comb. nov. — A. strongylocephala Pamp. var. sinensis Pamp. f. robusta Pamp. in Nuov. Giorn. Bot. Ital. n.s. 34: 178 (1927). — A. dubia Wall. ex Bess. var. myriantha (Wall. ex Bess.) Pamp., op. c. 36: 381 (1929) p.P. N. India, Sikkim, China (S.-W.). (E). A. indica Willd., Sp. Pl. 3: 1846 (1800); Nasir et Ali, Fl. W. Pakistan 716 (1976). — A. leptophylla D. Don, Prodr. Fl. Nep. 182 (1825) p.p. — A. indica Willd. var. multi- flora Wall. ex Bess., var nepalensis Bess. et var. heyneana Wall. ex Bess. in Nouv. 54 Comp. Newsl. 26, 1995 Mém. Soc. Nat. Mosc. 3: 56-57 (1834) et var. legitima Bess. in Bull. Soc. Nat. Mosc. 9: 55-56 (1836) p.p. — A.grata Wall. ex Bess., l.c. 3: 57 (1834) p.p. — A. wallichiana Bess., lc. 3: 69 (1834). — A. vulgaris Linn. var. indica (Willd) Maxim., Fl. Amur. 160 (1859). — A. vulgaris auct., non Linn.: Hook. f., Fl. Brit. Ind. 3: 325 (1881) p.p. quoad syn.; Nasir et Ali, Fl. W. Pakistan 720 (1972) p.p. — A. dubis auct., non Wall. ex Bess.: Pamp. in Nuov. Giorn. Bot. Ital. n.s. 33: 447-452 (1926) et 36: 434441 (1930) incl. var legitima (Bess.) Pamp. cum f. genuina Pamp. et f. communis Pamp., var. acuminata Pamp. f. congesta Pamp., var. multiflora (Wall. ex Bess.) Pamp., var. grata (Wall. ex Bess.) Pamp., var. orientalis Pamp., var. septentrionalis Pamp., var. compacta Pamp. et var. gracilias Pamp., cum formae; Nasir et Ali, l.c. 716 (1972). — A. nilaritica (C. B. Clarke) Pamp., l.c. 33: 452 (1926) p.p., incl. var. lobata Pamp. — A. eriocephala Pamp., l.c. 33: 454 (1926); Nasir et Ali, lc. 721 (1972). — A. dolichocephala Pamp., l.c. 34: 175 (1927); Nasir et Ali, Lc. 721 (1972). — A. myriantha auct., non Wall. ex Bess.: Y.R. Ling in Journ. Nanj. Techn. Coll. For. Prod. 1: 83 (1981). var. 1. indica Widely distr. Asia, except North. (BM, CGE, E). var. 2. elegantissima (Pamp.) Y.R.Ling et C.J. Humphries comb. nov. — A. elegantissima Pamp., |.c. 33: 454 (1926), incl. formae et var. wnaonensis Pamp.; Nasir et Ali, l.c. 716 (1972). N. India, Nepal, Kashmir, China (S.-W.). (BM, K). A. incisa Pamp. in Nuov. Giorn. Bot. Ital. n.s. 33: 456 (1929) incl. var. kunawarensis Pamp.; Nasir et Ali, Fl. W. Pakistan 721 (1972). — A. vulgaris Linn. var. myriantha (Wall. ex Bess.) Pamp. in Nuov. Giom. Bot. Ital. n.s. 33: 450 (1926) p.p. Nepal, N. Pakistan, Kashmir, China (S.-W.). (T.T., Type, Kash., BM, K). A. nilagirica (C. B. Clarke) Pamp. in Nuov. Giom. Bot. Ital. n.s. 33: 452 (1926). — A. vulgaris Linn. var. nilagirica C. B. Clarke, Comp. Ind. 162 (1876). N. India, China (S.-W.), N. Burma, (BM, K). A. amygdalina Decne in Jacq., Voy. Bot. 92. t. 100 (1836); Hook. f., Fl. Brit. Ind. 3: 325 (1881); Nasir et Ali, Fl. W. Pakistan 715 (1972). N. India, N. Pakistan, Kashmir. (BM, K, P). A. moorcroftiana Wall. ex DC., Prodr. 6: 117 (1837); Hook. f. , Fl. Brit. Ind. 3: 327 (1881); Nasir et Ali, Fl. W. Pakistan 717 (1972). — A. wallichiana Bess. in Nouv. Mém. Nat. Mosc. 3: 69 (1834) p.p. — A. hypoleuca auct., non Edgew.: Hook. f., l.c. 3: 326 (1881) p.p., pro syn. — A. strongylocephala Pamp. var. cachemirica Pamp. in Nuov. Giorn. Bot. Ital. n.s. 34: 176 (1927). — A. brunnea Pamp. in Lav. Inst. Bot. Univ. Cagl. 8: 2 (1938). Comp. Newsl. 26, 1995 55 N. India, Bhutan, Nepal, N. Pakistan, Kashmir, China (N.-W. et S.-W.). (BM, CGE, K). A tainingensis Hand.—Mazz. in Act. Hort. Gothob. 12: 277 (1938). — A. campbellii Hook. f. et Thoms. ex C.B. Clarke var. limprichtii Pamp. in Nuov. Giom. Bot. Ital. n.s. 34: 642 (1927) p.p. var. 1. tainingensis China (S.-W., et Qinghai). (E). var 2. nitida (Pamp.) Y.R. Ling comb. nov. — A. moorcroftiana Wall. ex DC. f. nitida Pamp., l.c. 34: 681 (1927). — A. moorcroftiana Wall. ex DC. var. nitida (Pamp.) Ling et Y.R. Ling in Act. Phytotax. Sin. 18 (4): 505 (1980). N.-W. India, Kashmir, China (S.-W.). (K). A. austro-himalayana (YR. Ling et H.S. Puri) Y.R. Ling et H.S. Puri in Guihaia 8 (1): 64 (1988). — A. tenuifolia auct., non Adam ex DC.: Y.R. Ling et H.S. Puri, l.c. 5 (1): 2 (1985). N. India. (N.C. Shah et S.C. Singh, Type, CIMAP, Lucknow). A. tukuchaensis Kitam. in Act. Phytotax. Geobot. 30: 127 (1979). Nepal (BM). A. campbellii Hook. f. et Thoms. in C. B. Clarke, Comp. Ind. 164 (1876); Hook. f., Fl. Brit. Ind. 3: 327 (1881); Nasir et Ali, Fl. W. Pakistan 715 (1972). — A. tibetica sensu Schlagint., Cat. [1399, Tibet. (P)], non Hook. f. et Thoms. ex Hook. f. Bhutan, Sikkim, N. Pakistan, Kashmir, China (Xizang). (BM, CGE, E, GH, K, P). 4). Sect. Viscidipubes Y.R. Ling A. viscida (Mattf.) Pamp. in Nuov. Giorn. Bot. Ital. n.s. 36: 424 (1930); Nasir et Ali, Fl. W. Pakistan 721 (1972). — A. moorcroftiana Wall. ex DC. var. viscida Mattf. in Feddes Rep. Sp. Nov. 22: 247 (1926). N. Pakistan, China (S.-W.). A. vexans Pamp. in Nuov. Giorn. Bot. Ital. n.s. 36: 427 (1930). Bhutan, China (S.-W.). (BM). A. thellungiana Pamp. in Nuov. Giorn. Bot. Ital. n.s. 33: 457 (1926). — A. vulgaris auct., non Linn.: Hook. f., Fl. Brit. Ind. 3: 325 (1881) p.p. N. India, Sikkim, China (S.-W.). (BM, K). A. myriantha Wall. ex Bess. in Nouv. Mém. Soc. Nat. Mosc. 3: 51 (1834). — A. vulgaris Linn. var. myriantha (Wall. ex Bess.) C. B. Clarke, Comp. Ind. 162 (1876) 56 Comp. Newsl. 26, 1995 excl. syn. A. paniculata Roxb. — A. vulgaris auct., non Linn.: Hook. f., Fl. Brit. Ind. 3: 325 (1881) p.p., quoad syn. — A. dubia auct., non Wall. ex Bess.: Pamp. in Nuov. Giorn. Bot. Ital. n.s. 33: 447-454 (1926), 34: 653 (1927), 36: 434-441 (1930), 46: 562 (1939) p.p. incl. var. legitima (Bess.) Pamp. cum f.genuina Pamp. et f. communis Pamp., sf. intermedia Pamp., var. acuminata Pamp. et var. longeracemulosa Pamp. et var. grata Pamp. p.p. — A. burmanica Pamp., I.c. 33: 455 (1926) excl. f. latiloba Pamp. — A. pleiocephala Pamp. var. typica Pamp. f. yunnanensis Pamp., l.c. 36: 446- 448 (1930). var. 1. myriantha N. India, Bhutan, Nepal, Kashmir, China (S.-W. et Qinghai), N. Burma. (BM, E, K). var. 2. pleiocephala (Pamp.) Y.R. Ling, l.c. 42 (2): 446 (1987). — A. vulgaris auct., non Linn.: Hook. f., l.c. 3: 325 (1881) p.p., quoad syn. — A. dubia auct., non Wall. ex Bess.: Pamp., I.c. 33: 449-450 (1926) p.p., incl. var. legitima (Bess.) Pamp. f. communis Pamp. sf. pauciflora Pamp. et var. myriantha (Wall. ex Bess.) Pamp. cum f. meridionalis Pamp. — A. pleiocephala Pamp. var. typica Pamp. cum f. latiloba Pamp. et f. discolor Pamp., l.c. 36: 446-448 (1930). N. India, Bhutan, Nepal, Kashmir, China (S.-W. et Qinghai), N. Burma, N. Thailand. (BM, CGE, K, L). 5). Sect. Albibractea Y.R. Ling A. lactiflora Wall. ex DC., Prodr. 6: 115 (1837). — A. lactiflora Wall. ex DC. f. genuina Pamp., f. septemlobata Pamp. et f. heryana Pamp. in Nuov. Giorn. Bot. Ital. n.s. 34: 674-675 (1927). N.-E. India, China (S., C., S.-W. et S.-E.), Thailand, Indo-China, Indonesia. (E). 2. Subgen. Dracunculus (Bess.) Peterm. 6). Sect. Dracunculus Bess. sensu stricto, excl. sect. Latilobus Y.R. Ling A. wellbyi Hemsl. et Pears. ex Deasy, Tibet Chin. Turk. 397 (1901) nom. nud., in Journ. Linn. Soc. 35: 183 (1902). — A. salsoloides auct., non Willd.: C.B. Clarke, Comp. Ind. 156 (1876) p.p.; Hook. f., Fl. Brit. Ind. 3: 321 (1881) p.p., incl. var. salsoloides Hook. f.; Nasir et Ali, Fl. W. Pakistan 719 (1881) p.p. — A. falconeri C.B. Clarke ex Hook. f., l.c.: 328 (1881) p.p. — A. salsoloides Willd. var. wellbyi (Hemsl. et Pears. ex Deasy) Ostenf. et Pauls. in Hedin, S. Tibet. 6(3): 40 (1922); Nasir et Ali, l.c. 719 (1972). — Oligosporus wellbyi (Hemsl. et Pears. ex Deasy) Poljak. in Mat. Fl. Veg. Kaschg. 11: 169 (1961). N. India, Sikkim, China (Xizang). (BM, E, GH, K, P). Comp. Newsl. 26, 1995 57 A. waltonii J.R. Drumm. ex Pamp. in Nuov. Giorm. Bot. Ital. n.s. 34: 707 (1927). China (S.-W. et Qinghai). (L). A. xigazeensis Ling et Y.R. Ling in Act. Phytotax. Sin. 18 (4): 511 (1980). —A. sal- soloides Willd. var. salsoloides Hook. f. p.p. et var. paniculata Hook. f., incl. syn. halimodendron Ledeb. ex Hook. f., Fl. Brit. Ind. 3: 321 (1881) p.p. China (Xizang). (CGE, GH, K). A. dracunculus Linn., Sp. Pl. 2: 849 (1753); C. B. Clarke, Comp. Ind. 157 (1876); Hook. f., Fl. Brit. Ind. 3: 321 (1881) p.p.; Nasir et Ali, Fl. W. Pakistan 716 (1972). — A. glauca auct., non Pall. ex Willd.: Hook. f., L.c.: 322 (1881); Nasir et Ali. Lc. 721 (1972). — Oligosporus dracunculus (Linn.) Poljak. in Mat. Fl. Veg. Kaschg. 11: 166 (1961). var. 1. dracunculus N. India, N. Pakistan, Kashmir, Sikkim, China (N., N.-E. et N.-W.), Afghanistan, Mongolia, C. & W. Asia, Russia, E., C. & W. Europe et N. America. (BM, CGE, E, KL). var. 2. pamirica (C. Winkl.) Y.R. Ling et C.J. Humphries, comb. nov. — A. pamirica C. Winkl. in Act. Hort. Petrop. 11: 329 (1890) incl. var. aschurbazewi C. Winkl. — A. simplicifolia Pamp. in Nuov. Giorn. Bot. Ital. n.s. 46: 584 (1939); Nasir et Ali, l.c. 720 (1972). — Oligosporus pamiricus (C. Winkl.) Poljak. in Mat. Fl. Veg. Kaschg. 11: 166 (1961). N. Pakistan, China (Xizang, Xinjiang), Afghanistan, Tadjikistan. (P). A. scoparia Waldst. et Kit., Pl. rar. Hung. 1: 66, tab. 65 (1802); C. B. Clarke, Comp. Ind. 158 (1876); Hook. f., Fl. Brit. Ind. 3: 323 (1881); Nasir et Ali, Fl. W. Pakistan 719 (1972). — A. mollissima D. Don, Prodr. Fl. Nep. 182 (1825); Hook. f., I.c.: 330 (1881). — A. elegans Roxb., Fl. Ind. 3: 321 (1832).— A.trichophylla Wall. ex DC., Prodr. 6: 100 (1837); C. B. Clarke, I.c.: 158 (1876). — A. sacrorum sensu Schlagint., Cat. [1352 (1856), Tibet (E)], non Ledeb. — A. kohatica Klatt in Sitzb. Akad. Muench. 91 (1878); Hook. f., I.c.: 330 (1881) (dubia). — A. capillaris Thunb. var. scoparia (Waldst. et Kit.) Pamp. in Nuov. Giorn. Bot. Ital. n.s. 34: 642 (1927) incl. formae. — Oligosporus scoparius (Waldst. et Kit. ) Less. in Linnaea 9: 191 (1834). Widely distr. in Eurasia. (BM, CGE, E, GH, K). A, edgeworthii Balakr. in Journ. Bombay Nat. Hist. Soc. 63: 329 (1967). — A. stricta auct., non Heyne ex DC.: Edgew. in Trans. Linn. Soc. 20: 73 (1846); C. B. Clarke, Comp. Ind. 158 (1876); Hook. f., Fl. Brit. Ind. 3: 323 (1881); Nasir et Ali, Fl. W. Pakistan 720 (1972). — A. stricta Edgew. f. genuina Pamp. in Nuov. Giorn. Bot. Ital. 34: 705 (1927). 58 Comp. Newsl. 26, 1995 var. 1. edgeworthii N. India, Nepal, Sikkim, Kashmir, China (S.-W.). (BM, CGE, E, GH, K). var. 2. diffusa (Pamp.) Ling et Y.R. Ling in Act. Phytotax. Sin. 18 (4): 509 (1980). — A. stricta Edgew. f. diffusa Pamp., l.c. 34: 705 (1927). N. India, Nepal, Kashmir, China (S.-W.). (L). A. demissa Krasch. in Act. Inst. Bot. Acad. Sci. USSR 1(3): 348 (1936). Nepal, China (N.-W. & Xizang). (BM). 7). Sect. Latilobus Y.R. Ling A. desertorum Spreng., Syst. Veg. 3: 490 (1826). — A. desertorum Spreng. var. sprengeliana Bess. in Bull. Soc. Nat. Mosc. 8: 65 (1835). — Oligosporus desertorum (Spreng.) Poljak. in Mat. Fl. Veg. Kaschg. 11: 167 (1961). var. 1. desertorum Bhutan, Sikkim, China (S.-W., N., N.-E.), Japan, Korea, Mongolia, E. Russia. (K, P). var. 2. foetida (Jacq. ex DC.) Ling et Y.R. Ling comb. nov. — A. foetida Jacq. ex DC., Prodr. 6: 98 (1837). — A. desertorum auct., non Spreng.: C. B. Clarke, Comp. Ind. 158 (1878) p.p.; Nasir et Ali, Fl. W. Pakistan 715 (1972) p.p. N. Pakistan, Nepal, N. India, Kashmir, China (S.-W.). (BM). var. 3. tongolensis Pamp. in Nuov. Giorn. Bot. Ital. n.s. 34: 651 (1927). China (S.-W.). (P). A. japonica Thunb., Fl. Jap. 308 (1784). — A. glabrata Wall. ex Bess. in Bull. Soc. Nat. Mosc. 8: 20 (1835). — A. cuneifolia DC., Prodr. 6: 126 (1837). — A. parviflora auct., non Buch._Ham. ex Roxb.: C. B. Clarke, Comp. Ind. 159 (1876); Hook. f., Fl. Brit. Ind. 3: 322 (1881) p.p.; Nasir et Ali, Fl. W. Pakistan 718 (1972) p.p. — A. jacquemontiana auct., non Bess.: Hook. f., l.c.: 321 (1881). — A. dracunculus auct., non Linn.: Hook. f., l.c.: 321 (1881). — A. japonica Thunb. var. myriocephala Pamp. in Nuov. Giorn. Bot. Ital. n.s. 34: 665 (1927) et var. lanata Pamp. in Lav. Inst. Bot. Univ. Cagl. 8: 3 (1938). — Oligosporus japonicus (Thunb.) Poljak. in Mat. Fl. Veg. Kaschg. 11: 169 (1961). Widely distr. in Asia. (BM, CGE, K). A. parviflora Busch.Ham. in Roxb., Hort. Beng. 61 (1814) nom. nud., Fl. Ind. 420 (1832) nom. conserv.: C. B. Clarke, Comp. Ind. 159 (1876) p.p.; Hook. f., Fl. Brit. Ind. 3: 322 (1881) p.p.; Nasir et Ali, Fl. W. Pakistan 718 (1972) p.p. — A. japonica Thunb. var. parviflora (Buch._Ham. ex Roxb.) Pamp. in Nuov. Giom. Bot. Ital. n.s. 34: 665 (1927). — A. subdigitata auct., non Mattf.: Kitam. in Kihara, Fau. Fl. Nep. Comp. Newsl. 26, 1995 59 Himal. 1: 246 (1952-1953). — Oligosporus parviflorus (Buch.Ham. ex Roxb.) Poljak. in Mat. Fl. Veg. Kaschg. 11: 170 (1961). India, Pakistan, Sikkim, Bhutan, Nepal, Kashmir, N. Thailand, China (S.-W.). (BM, E, K, Lucknow, P). A. dubia Wall. ex Bess. in Nouv. Mém. Soc. Nat. Mosc. 3: 39 (1834) (Sect. Abrota- num Bess.). — A. vulgaris auct., non Linn.: Hook. f., Fl. Brit. Ind. 3: 325 (1881) p.p., quoad syn. — A. subdigitata Mattf. in Feddes Rep. Sp. Nov. 2: 243 (1926) p.p. — A. dracunculus Linn. var. inodora auct., non Bess.: Pamp. in Nuov. Giorn. Bot. Ital. n.s. 34: 653 (1927) p.p., quoad pl. C.B. Clarke. — A. dracunculus Linn. var. subdigitata (Mattf.) Pamp. f. thomsonii Pamp. 1.c. 36: 379 (1929) incl. syn. — A. thomsonii C.B. Clarke ex Pamp., cum f. chinensis Pamp. p.p. — A. subdigitata Mattf. var. thomsonii (C.B. Clarke) S.Y. Hu in Quart. Journ. Taiwan Mus. 18 (3-4): 263 (1965). var. 1. dubia N. India, Bhutan, Nepal, China (N., N.-W., C. et S.-W.). (BM, Wallich Cat. 3307a/417, Blinkworth, Isotype, Kumaon, CGE, E, K, L). var. 2. subdigitata (Mattf.) Y.R. Ling, l.c. 42(2): 445 (1987). — A. jacque- montiana Bess. in Bull. Soc. Nat. Mosc. 8: 61 (1835) p.p. incl. syn. — A. glabra Jacq. ex Bess. — A. cannabina Jacq. ex Bess. in DC., Prodr. 6: 97 (1837) pro syn. — A. de- sertorum Spreng. var. jacquemontiana DC., 1.c. 98 (1837). — A. royleana DC., l.c. 115 (1837); Hook. f., 1.c.: 329 (1881). — A. desertorum auct., non Spreng.: C. B. Clarke, Comp. Ind. 158 (1876); Hook. f., I.c.: 332 (1881); Nasir et Ali, Fl. W. Pakistan 715 (1972). — A. glauca auct., non Pall. ex Willd.: Hook. f., l.c.: 332 (1881). — A. dubia Wall. ex Bess. var. jacquemontiana (Bess.) Pamp. I.c.: 451 (1926). — A. sub- digitata Mattf. 1.c.: 243 (1926) p.p., incl. var. falciloba Mattf. — A. foetida auct., non Jacq. ex DC.: Pamp. l.c.: 655 (1927) p.p. — A. dracunculus Linn. var. inodora Bess. f. pinnata Pamp. et var. subdigitata Pamp., incl. f. chinensis Pamp., f. falciloba Pamp. et f. intermedia Pamp. cum sf. oblonga Pamp., l.c.: 379-381 (1929). — Oligosporus jac- quemontiana (Bess.) Poljak. in Mat. Fl. Veg. Kaschg. 11: 168 (1961). N. India, Bhutan, Nepal, Kashmir, China (N., C. et S.-W.). (BM, K, P). II. Seriphidium (Bess.) Poljak. in Mat. Fl. Veg. Kaschg. 11: 171 (1961). — Artemisia Linn. sect. Seriphidium Bess. in Bull. Soc. Nat. Mosc. 1 (8): 222 (1828). — Artemisia Linn. subgen. Seriphidium (Less.) Peterm., Deutschl. Fl. 294 (1848). S. brevifolium (Wall. ex DC.) Ling et Y.R. Ling in Act. Phytotax. Sin. 18 (4): 513 (1980). — Artemisia brevifolia Wall. ex DC., Prodr. 6: 103 (1837); Nasir et Ali, Fl. W. Pakistan 715 (1972). — A. maritima Linn. vera C. B. Clarke, Comp. Ind. 159 (1878). — A. maritima auct., non Linn.: Hook. f., Fl. Brit. Ind. 3: 323 (1881) p.p., quoad syn.; Nasir et Ali, 1c. 717 (1972) p.p. — A. maritima Linn. var. genuina 60 Comp. Newsl. 26, 1995 Pamp. in Nuov. Giom. Bot. Ital. n.s. 34: 679 (1927) incl. f. suberecta Pamp. — A. maritima Linn. var. pauciflora auct., non Ledeb.: Pamp., l.c. 46: 573 (1939). N. India, N. Pakistan, Kashmir, China (W. Xizang). (BM, GH, P). S. thomsonianum (C. B. Clarke) Ling et Y.R. Ling in Act. Phytotax. Sin. 18 (4): 513 (1980). — Artemisia maritima Linn. var. thomsonianum C.B. Clarke, Comp. Ind. 160 (1878). — A. maritima auct., non Linn.: Hook. f., Fl. Brit. Ind. 3: 323 (1881) quoad syn. — A. lehmanniana auct., non Bge.: Hook. f., 1.c.: 323 (1881) pro syn.; Nasir et Ali, Fl. W. Pakistan 717 (1972) p.p. — A. herba-alba auct., non Asso: Nasir et Ali, l.c. 716 (1972). N. Pakistan, Kashmir, China (W. Xizang), Afghanistan. [Schlagint. Cat. 6218 (1856), Tibet, CGE, GH, K)]. S. aucheri (Boiss.) Ling et Y.R. Ling in Act. Phytotax. Sin. 18 (4): 513 (1980). — Artemisia aucheri Boiss., Fl. Or. 3: 367 (1875); Nasir et Ali, Fl. W. Pakistan 715 (1972). — A. maritima Linn. var. b. C. B. Clarke, Comp. Ind. 159 (1878) incl.syn. — A. griffithiana sensu C. B. Clarke, non Boiss. — A. maritima Linn. var. aucheri (Boiss. Pamp. in Nuov. Giorn. Bot. Ital. n.s. 34: 680 (1927). N. Pakistan, China (W. Xizang), Iran, Afghanistan. (BM, CGE, K). S. lehmannianum (Bge.) Poljak. in Mat. Fl. Veg. Kaschg. 11: 175 (1961). — Arte- misia lehmanniana Bge. in Mém. Acad. Sci. Petersb. 7: 340 (1854). — A. maritima auct., non Linn.: Hook. f., Fl. Brit. Ind. 3: 323 (1881) quoad syn. N.-W. India. (E). Comp. Newsl. 26, 1995 61 POLLEN MORPHOLOGY IN THE SENECIO NEMORENSIS GROUP (COMPOSITAE) FROM THE CARPATHIANS Iva Hodélova ) and Pavol MArtonfi 2) 1) Institute of Botany Slovak Academy of Sciences Dubravska cesta 14 SK-842 23 Bratislava, Slovakia 2) Department of Experimental Botany and Genetics Faculty of Science P.J. Safarik University Manesova 23 SK-04 154 Kosice, Slovakia Abstract A description of the external morphology and size of pollen grains in five "pure" and four hybrid taxa of the S. nemorensis group from the Carpathians is presented. The pollen grains of all 9 taxa were examined using image analyser ASBA and SEM. No expressive differences in the size, shape and external morphology of the pollen between the studied taxa were recorded. The length and diameter ranges between 29— 54 x 17-32 lum, the shape is prolate, subprolate to spheroidal. Key words: Senecio nemorensis group, pollen, external morphology, length, diameter, Carpathians. Introduction Reviews of the external morphology of Compositae pollen grains have been made several times (Skvarla & Turner 1966, Mesfin 1984, Otieno & Mesfin 1992 etc.). However, there are no published data about the pollen grains of the Senecio nemorensis group. The aim of the present study is to describe and document the morphology of pollen grains of the S. nemorensis group and is part of taxonomical study of this group in the Carpathians. Five taxa (S. hercynicus Herborg subsp. hercynicus; S. ovatus (P. Gaertn., B. Mey. et Scherb.) Willd. subsp. ovatus; S. germanicus Wallr. subsp. germanicus; S. ucranicus Hodalova nom. prov.; S. nemorensis auct. non L. and four 62 Comp. Newsl. 26, 1995 new hybrids (S. x futakii Hoddlova nom. prov. (S. germanicus subsp. germanicus x S. ovatus subsp. ovatus) (Hoddélova & Marhold, submitted); S. germanicus subsp. germanicus x §. ucranicus; S. germanicus subsp. germanicus x S. nemorensis; S. germanicus subsp. germanicus x S. hercynicus subsp. hercynicus x S. ovatus subsp. ovatus) were recorded. Morphological investigation indicated that great morphological variation observed in many populations of this group is probably caused by introgressive hybridization (Hoddlov4 1994; Hoddlov4 & Marhold, submitted; Hoddlové & Valachovic, submitted). Material and Methods Morphological data were obtained from herbarium specimens (deposited at SAV) collected by us throughout the Carpathians (Slovakia, Ukraine, Romania) during 1990 through 1993. Morphological analysis of pollen grains was based on material from a total of 25 populations (for localities see Appendix) representing all taxa of the S. nemorensis group in the Carpathians. Pollen grains were mounted on slides in glycerine jelly after soaking in glycerine jelly for 3 days. Measurements for morphometrical analysis were made using an image analyzer ASBA (Wild, Heerbrugg, Switzerland) on 120 grains per taxon and computed on a special program made for ASBA - STAT program (Anonymus 1985). Length is defined as the maximum of 4 projections, diameter as area/length. Form factor values characterize the shape of pollen grains (form factor 5.09 for a circular shape, 6 for a square, 7 for a rectangle with a side ratio of 2:1, and 12.4 for a rectangle with a side ratio of 5:1; for more details see BaluSka et al. 1990). The mean and standard deviations were calculated using the STATGRAPHICS program. Scanning electron microscopic (SEM) studies were performed on 2 populations for each taxon. Pollen grains were acetolyzed us follows (modified from Erdtman 1969): tubular flowers were soaked in a 9:1 solution of acetic acid anhydride and sulphuric acid for 15 min, then placed in a boiling water bath for 5 min. They were then centrifuged for 4 min, at 4000 rpm and rinsed with distilled water. Centrifugation and rinsing were repeated three times. The pollen grains were then twice centrifuged and rinsed with 97% ethanol. Acetolyzed pollen was mounted on brass stubs, dehydrated, coated with gold-palladium, and visualized and photographed using a Jeol Scanning Microscope (JSM-840). Unacetolyzed pollen grains of taxa of the S. nemorensis group were studied for comparison (Fig. 10). Comp. Newsl. 26, 1995 Results and Discussion No taxonomically significant differences between the taxa of the S. nemorensis group for length, diameter and external morphology of pollen grains were found. The pollen grains of "pure" and hybrid taxa were well developed (all studied populations have the same chromosome numbers — 2n=40 (unpublished) and the genetic barriers between the taxa are very weak) which could be one of the many reasons that introgressive hybridization between studied taxa occurs very frequently. 63 Table 1. Number of population samples (n) (number of population samples refers to the numbers of localities in Appendix), mean and standard deviation for length (um), diameter (jm) and form factor values for pollen grains of taxa of the S. nemorensis group from the Carpathians = ie S. hercynicus subsp. hercynicus S. ovatus subsp. ovatus S. germanicus subsp. germanicus S. ucranicus S. nemorensis S. germanicus subsp. germanicus x §. ovatus subsp. ovatus S. germanicus subsp. germanicus x §. ucranicus S. germanicus subsp. germanicus x §. nemorensis S. germanicus subsp. germanicus x S. hercynicus subsp. hercynicus x S. ovatus subsp. ovatus 1-3 Aes 8-13 length diameter mean/(range 37.01/ (29-46) 47.64/(18-28) 38.52/(29-48) 49.78/(20-31) 37.15/(31-54) 47.56/(19-30) 37.42/(28-47) 48.32/(19-32) 37.95/(25-47) 48.90/(17-28) 37.99/(29-46) 49.06/(20-30) 35.00/(29-43) 44.50/(17-29) 38.53/(32-46) 50.46/(21-30) 39.14/(31-48) 50.52/(19-29) standard deviation 3.00 2.07 3.82 2.68 4.93 2.31 4.13 Peig)s) 5.85 2.18 3.04 1.98 2.66 252 2.24 155 4.07 1.82 factor 5.47 5.46 5.53 5.48 5.48 5.47 5.49 5.45 5.45 64 Comp. Newsl. 26, 1995 Size Variation in pollen grain length and diameter is relatively great within all studied taxa. The length ranges between 29-43 jum, the diameter between 17-32 uum. Shape and Apertures The pollen grains are radially symmetrical and isopolar. No taxonomically significant differences between the taxa in shape were recorded, however, the shape of the pollen grains within each studied taxon is variable (Figs. 4-5). The shape is characterized by form factor values (Tab. 1). According to these values the pollen grains could be described as prolate, subprolate to spheroidal, similar to most pollen grains in Compositae (Mesfin 1984: 28; Otieno & Mesfin 1992), also the pollen grains in the studied taxa are tricolporate (Fig. 2) with colpi having lalongate ora (Figs. 1, 3, 5, 9). Spines The sculpture of the grains is echinate, with the perforations concentrated obviously at the spine bases. The length of spines ranges between 3.0 — 5.2 um. Perforations The perforations at the spine bases and on the tectal surface between spines are irregular in their size and distribution. The holes on the tectal surface between spines are usually smaller than the perforations at the spines. The largest cavities are located away from the extreme base of spines (Figs. 1, 2, 4, 5, 6, 7, 8). Distal part of the spines is obviously without perforations (Fig. 7). Infraspecific variation in the number and distribution of holes was observed. Nonperforate pollen grains were also rarely found (Fig. 9). Acknowledgements We are obliged to M. Stankovic of the Geological Institute of Dionyz Stir, Bratislava, M. Hauskrecht and V. Matisovd, for their help with the preparation of pollen material. We also thank J. Compton, Reading, for language correction. This work has been supported by the Ministry for Science and Research, Vienna, Austria, through the East-West Program of the Austrian Academy of Sciences (Project No. OWP 58) and by the Grant Agency for Science, Bratislava, Slovakia (Grant No. 2/41). Comp. Newsl. 26, 1995 References Anonymus 1985. ASBA Software Anleitung Wild & Leitz A.G. Baluska, F., Kubica, §. & M. Hauskrecht 1990. Postmitotic 'isodiamteric’ cell growth in the maize root apex. Planta 181: 269-274. Erdtman, G. 1969. Handbook of Palynology. Munksgaard, Copenhagen. Hodalova, I. 1994. Poznamky k problematike komplexu Senecio nemorensis agg. v. Karpatoch. Zpravy Ceskolov. Bot. Spolecn. 28 (1993): 1-14. Hodalova, I. & K. Marhold 1995. Sympatric populations of Senecio ovatus subsp. ovatus, S. germanicus subsp. germanicus (Compositae) and their hybrid in the Carpathians and the adjacent part of Pannonia. I. Multivariate morphometric study. (Submitted). Hod4lové, L & M. Vallachovic 1995. Sympatric populations of Senecio ovatus subsp. ovatus, S. germanicus subsp. germanicus (Compositae) and their hybrid in the Carpathians and the adjacent part of Pannonia. II. Synecological differentiation and distribution. (Submitted). Mesfin, T. 1984. The genus Bidens (Compositae) in Tropical Africa. Symb. Bot. Upsal. 2A (1): VII+138 pp. Otieno, D.F. & T. Mesfin 1992. Pollen morphological studies in Senecio (Compo- sitae-Senecioneae) from Ethiopia. Comp. Newsl. 20/21: 22-28. Skvarla, J., Turner, B.L., Patel, V.C. & A.S. Tomb 1977. Pollen morphology in the Compositae and in morphologically related familes. Jn: Heywood, V.H., Harborne, J.B. & B.L. Tuer (eds.), The Biology and Chemistry of the Compositae, 1: 141- 248. New York/London: Academic Press. Appendix List of the localities of the population samples. 1. Slovakia, Nizke Tatry, Mt. Chopok (Hoddlovd 1990).— 2.Slovakia, ZApadné Beskydy, Such4 Hora (Hoddlova 1990). — 3. Poland, Tatry, Dolina Kondratowa, Pieklo (Hoddlova 1990). — 4. Slovakia, Bukovksé vrchy, Nova Sedlica (Hodélova 1989). — 5. Slovakia, Turcianska kotlina, Valca (Hoddélova 1993). — 6. Slovakia, Slovenské rudohorie, Roznava (Hoddlova 1993). — 7. Slovakia, Mala Fatra (Krivanska Fatra), Terchova (Futak, Jasicova & Zahradnikova 1964). — 8. Ukraine, Prikarpattja, Knjaz Dvor (Hoddlova & Kagalo 1990). — 9. Ukraine, Zakrpats"ke peredgir’ja, Ugelja (Hoddlové & Kagalo 1991). — 10. Slovakia, Bukovské vrchy, Nova Sedlica (Hodd4lova 1989). — 12. Ukraine, Gorgany, Kremenci (Hoddlova & Kagalo 1990). — 13. Romania, Muntii Cindrel, Sibiu, Paltinis (Hoddlova 1992). — 14. Ukraine, Cyvcyno-Grynjavski gory. Mt. Cyvcyn (Ginda 1990). — 15. Ukraine, Comp. Newsl. 26, 1995 Cornogora, Mt. Coma gora (Hoddlova et Kagalo 1991). — 16. Romania, Muntii Retezat, Cimpu lui Neag (Hod4lova and Valachovic 1993).— 17. Slovakia, Provazsky Inovec, near the chalet Chata na Inovci (Hoddlovd 1992). — 18. Slovakia, Biele Karpaty, Mt. Velka Javorina (Hoddlova 1993). — 19. Slovakia, Slanske vrchy, between the villages Herlany and Banské (Hoddlova 1989). — 20. Ukraine, Cornogora, Mt. Goverlo (Hoddlova et Kagalo 1990). — 21. Ukraine, Comogora, Mt. Dancer (Hoddlova et Kagalo 1990). — 22. Romania, Muntii Retezat, Cimpu lui Neag (Hod4lova et Valachovic 1993). — 23. Romania, Muntii Cindrel, Sibiu, Paltinis (Hod4lova 1992). — 2A. Slovakia, Vysoké Tatry, Mt. Solisko (Hoddélova 1989). — 25 Slovakia, Vysoké Tatry, Strbské Pleso (Hod4lova 1993). Figs. 1-9 Fig. 10 FIGURE LEGENDS Scanning electron micrographs of acetolyzed pollen grains of S. nemorensis agg. 1. §. hercynicus subsp. hercynicus (magnification x 2700) 2. — S. ovatus subsp. ovatus (magnification x 1900) 3. S. germanicus subsp. germanicus (magnification x 2000) 4-5. S. ucranicus (magnification x 1500, 2000) 6. S§. nemorensis (magnification x 2300) 7. S. germanicus subsp. germanicus x S. ovatus subsp. ovatus (magnification x 6500) 8. S. germanicus subsp. germanicus x S. ucranicus (magnification x 2500). 9. S. germanicus subsp. germanicus x S. nemorensis (magnification x 2000) Unacetolyzed pollen grain of S. germanicus subsp. germanicus x S. hercynicus subsp. hercynicus x S. ovatus subsp. ovatus (magnifica- tion x 1990. Comp. Newsl. 26, 1995 Fig. 1 —3 67 68 Fig. 4—5 Comp. Newsl. 26, 1995 Comp. Newsl. 26, 1995 Fig. 6 —8 69 70 Fig. 9 — 10 Comp. Newsl. 26, 1995 Comp. Newsl. 26, 1995 71 BOOK ANNOUNCEMENT ASTERACEAE Cladistics and Classification by Kare Bremer 752 pp. 89 figures (including 179 detail drawings), 24 tables, 70 cladograms, 6 x 9", hardcover, ISBN 0-88192-275-7.© 1994 Available from Timber Press, Inc. The book may be ordered from either of two locations: -—US. office: 133 S.W. Second Ave, Suite 450, Portland, OR 97204-3257. Telephone Nos. (800) 327-5680, (503) 227-2878, Fax (503) 227-3070. Price: U.S. $79.95 Book orders from the USA, Canada, or worldwide other than the U.K. and Europe: Please add, for each book, $9.50 shipping and handling, $6.00 for overseas surface delivery, or 35% for overseas air delivery. — U.K. office: 10 Market St., Swavesey, Cambridge, CB4 5QG, United Kingdom. Telephone (0954) 232959. Price: U.K.£52.50 Book orders from the United Kingdom and the Europe: Please add, for each book, £4.25 shipping and handling. This volume is a comprehensive documentation of the evolutionary relationships of Asteraceae (Compositae) and the groups within it. All available information on the family's phylogeny is assembled, with chapters on cladistics, classification, morphology, evolution, and each of the subfamilies and tribes. All genera are described, and there are 179 detail drawings and 70 cladograms. This volume is the most complete treatment of the family in more than a century. Kare Bremer, professor of systematic botany and dean of biology at Uppsala University, Sweden, has researched the family Asteraceae for more than 25 years. A leading synantherologist, he is one of the first and foremost proponents of cladistics in plant systematics. He has authored numerous articles in English and Swedish for international scientific journals and is coauthor or coeditor of a number of other books and monographs. 72 Comp. Newsl. 26, 1995 Here's what reviewers had to say: "_..Bremer’s book will doubtless come to function as a bible, perhaps a new testament, for at least the current generation of students of composites." — J.L. Strother, UC, Taxon 43, November 1994. "This book will be a valuable resource to synantherologists, plant taxonomists, and systematists." — Michael O. Dillon, Taxon 43, November 1994. "All of us interested in composites should thank K4re Bremer for putting together a tremendous amount of information on the phylogeny of the family. He has produced a book of which he should be justifiably proud...if you are doing research in Asteraceae, no doubt in the next few years (or months) the pages of this tome will become well worn because you cannot ignore the tremendous amount of information between the covers." — Dan Crawford, Ohio State University, Columbus, Plant Science Bulletin, Vol 40, Number 2, Summer 1994. "Overall, I recommend this book based on its scholarly production and provocative content. Depending on one's need, this may be a "must have" reference." — SIDA 16 (2): 388, 1994. "For any particular genus, Bremer’s book provides summary information and references to monographic and other literature. Nearly 600 of the approximately 1100 references cited date from 1976 or later." — John L. Strother, University Herbarium, UC, The Quarterly Review of Biology, March 1995. "One of the most useful features of the book is the inclusion of generic descriptions with information about distribution and number of species. This is the first recent book on the Asteraceae to attempt this task." — Timothy K. Lowrey, Department of Biology, University of New Mexico, Albuquerque, ASPT Newsletter, April 1994.