SK Pay a 049] “COMPOSITAE © * NEWSLETTER Number 46 30 June 2008 Scientific Editor: Berti. NORDENSTAM Technical Editor: GUNNEL WirENIUS NOHLIN Published and distributed by The Swedish Museum of Natural History, Department of Phanerogamic Botany, P.O. Box 50007, SE-104 05 Stockholm, Sweden ISSN 0284-8422 | CONTENTS TELES, A. M., Sopra, M. & J. R. STEHMANN: Synopsis of Leptostelma (Asteracae: Astereae) ] ANDERBERG, A. A. & A. K. PANDEY: Nanothamnus sericeus THOMSON, a derived species of Blumea 8 KEIGHERY, G.: A new subspecies of Flaveria (Asteraceae) from Western Australia 20 IsawumI, M. A.: The status of generic revision in the African Vernonieae (Asteraceae) ps | ; AveasitE, A. E.: Leaf anatomical studies in some species of the tribe Cichorieae (Asteraceae) in Nigeria 49 VASANTHY, G. & D. DE FRANCESCHI: Studies on Vernonia monosis C. B. CLarKe (Asteraceae): Palynology, Taxonomy and Phytogeography 59 TeLes, A. M. & B. Norpenstam: A new species of Graphistylis (Asteraceae: Senecioneae) from Minas Gerais, Brazil 76 ANDERBERG, A. A., ENGLUND, M. & H. BEENTJE: On the systematic position of Inula rungwensis 83 New taxa and combinations published in this issue 85 7 ave? et 0 ™ aa ae es Po Pat ot j + - \ _ « ' ; 2 ‘4 4 oe aa i F a 7 : age a a 2 5 erat Comp. Newsl. 46, 2008 1 Synopsis of Leptostelma (Asteraceae: Astereae) ARISTONIO M. TELES, MARCOS SOBRAL & JOAO RENATO STEHMANN Universidade Federal de Minas Gerais, Instituto de Ciéncias Biolégicas Departamento de Botanica Av. Antonio Carlos, 6627, Pampulha >_> sls | vin Lt. Mi 31.270-910 Belo Horizonte, Minas Gerais, Brazil LIBRARY aristonio@hotmail.com Alustract NEW YORK There is presented a synopsis of the genus Leptostelma D. Don and four new combinations from Erigeron to Leptostelma are proposed: L. camposportoi (CABRERA) A. TELES & SoOBRAL, L. catharinense (CABRERA) A. TELES & SOBRAL, L. meyeri (CABRERA) A. TELES and L. tucumanense (CABRERA) A. TELES. A key to species of Leptostelma and commentaries about geographical distribution of the species are provided. Resumen Se presenta una sinopse del género Leptostelma D. Don y se propone cuatro nuevas combinaciones de Erigeron para Leptostelma: L. camposportoi (CABRERA) A. TELES & SosrRAL, L. catharinense (CABRERA) A. TELES & SOBRAL, L. meyeri (CABRERA) A. TELES y L. tucumanense (CABRERA) A. TELES. Una clave para las especies de Leptostelma y comentarios sobre la distribucion geografica de las mismas son presentados. Introduction Leptostelma D. Don was described in 1830 to accomodate one single species, Leptostelma maxima D. Don. Later, BENTHAM (1873) considered Leptostelma as a section of Erigeron L. [Erigeron L. sect. Leptostelma (D. Don) BENTH. & Hook.], in which he was followed by Baker (1882) in the Flora Brasiliensis. NESoM (1989) accepted Erigeron as a big genus with 20 sections [including the sections Leptostelma and Trimorpha (Cass.) DC.] and over 400 species, but few years later treated Leptostelma provisorily in the generic rank (NEsom 1994c). Recently Hinp & Nesom (2002) formally reinstated Leptostelma at the generic level with a new combination, Leptostelma tweediei (Hook. & Arn.) D. J. N. Hinp & BOTANICAL GARDEN a WA SEP 7? 2008 Nm Comp. Newsl. 46, 2008 G. L. Nesom. The last generic treatment of the Astereae recognized Leptostelma belonging to subtribe Conyzinae, with five species (NESoM & RoBINsoN 2007). BAKER (1882) considered the Brazilian Erigeron as belonging to two sections, Leptostelma and Caenotus Nutrt., the first section with ray florets with ligules larger than tubular florets (with the same length in Caenotus), usually arranged in one series (several in Caenotus) and receptacle alveolate with short projections (fimbriate) (smooth in Caenotus). Based on these characters Cronquist (1943) transferred all species of sect. Caenotus to Conyza. Erigeron occurs predominantly in continental North and Central America, with some species in the Old World, West Indies and South America (NEsoM 1994c), where it is restricted to the Andes and contiguous regions (NESOM 1994c). Considering this phytogeographical distinction and some morphological differences such as the length of ray florets and receptacle type, all other species of Erigeron sect. Leptostelma need validation under Leptostelma at generic level. These species belong to the subtribe Conyzinae and are closely associated with Apopyros G. L. Nesom, Hysterionica WILLD. and Neja D. Don, forming a natural group marked by a geographic range primarily in southeastern South America, and three-nerved phyllaries (NEsom 1994a, b, c). Noyes (2000) showed from ITS sequence data that the southeastern South American taxa Leptostelma (treated as a section of Erigeron), Apopyros, Hysterionica and Neja are closely related, supporting the hypothesis of Nesom (1994c). According to this author, South American Conyzinae are derived from North American Erigeron, and this subtribe is most closely related to North American Astereae. Taxonomic Treatment Leptostelma D. Don, in Sweet, Brit. Flow. Gard., ser 2. 1: 38. 1831. Type: Leptostelma maximum D. Don, in Sweet, Brit. Flow. Gard., ser. 2. 1: 38. 1831. Perennial herbs. Leaves alternate, basal leaves rosulate or all cauline, entire, serrate, toothed or lobed. Capitulescence corymbiform. Heads pedunculate, heterogamous, radiate; involucre hemispheric or campanulate; phyllaries narrow, |—3-seriate, 3-nervate; receptacle flat or slightly convex, alveolate, and fimbriate; ray florets pistillate, corolla liguliform, white or yellow, 1—2-seriate; disk florets perfect, corolla tubulose, yellow, 5-lobed. Cypselae flattened, 2-ribbed; pappus of numerous capillary bristles, 1-seriate. Comments. Leptostelma comprises six species, occurring in Argentina, Bolivia, Brazil, Paraguay and Uruguay. The genus is closely related to Apopyros, Hysterionica and Neja, differing from Apopyros by the radiate heads (vs. disciform in Apopyros), from Hysterionica by the uniseriate pappus (vs. biseriate Comp. Newsl. 46, 2008 3 in Hysterionica) and from Neja by the 2-ribbed cypselae (vs. 7—10-ribbed in Neja). 1. Leptostelma camposportoi (CABRERA) A. TELES & SosBRAL, comb. nov. Basionym: Erigeron camposportoi CABRERA, Arch. Jard. Bot. Rio de Janeiro 15: 75. 1957. Type: Brazil, estado do Rio de Janeiro, Serra dos Orgaos, in silva, 1800 m alt., CABRERA 12294, 12 Jul 1956 (holotype LP, photo seen; isotypes GH, not seen, RB, not found). Geographical distribution. Brazil, Rio de Janeiro (Serra dos Orgios and Serra do Itatiaia). 2. Leptostelma catharinense (CABRERA) A. TELES & SosBraL, comb. nov. Basionym: Erigeron catharinensis CABRERA, Arch. Jard. Bot. Rio de Janeiro 15: 75. 1957. Type: Brazil, Santa Catarina, Campo dos Padres, Bom Retiro, 2000 m.s.m., 15 Feb 1948, Reitz 2341 (holotype LP, not seen; isotype US, photo seen). Geographical distribution. Brazil, Santa Catarina, in altitudes ranging from 1200 to 2000 m above sea level, in the municipalities of Bom Jardim da Serra, Bom Retiro, Campo Alegre, Lauro Miiller, Orleans and Santo Amaro da Imperatriz. Comments. SoLsric (1962) regarded Erigeron catharinensis as a synonym of Erigeron (=Leptostelma) maximus (D. Don) DC., and commented on the morphological variation of this species. However, we analysed several collections in Brazilian herbaria [Smitru, L. B. 7753 (RB); Reitz, R. 6106 (RB); Reitz, R. 7425 (RB); Reitz, R. 7673 (RB); Reitz, R. 8688 (RB)] and saw many populations in the field, and we could verify that the population found in Santa Catarina (described by Casrera as Erigeron catharinensis) is distinct from those growing in southeast Brazil. The differences are the ovate, petiolate leaves (the petioles winged); involucre 5—6 mm long, glabrous, and ligule 3—5 mm long (vs. lanceolate to oblong-lanceolate, sessile leaves; involucre 12—14 mm long, hirsute, and ligules 12-15 mm long in L. maxima). 3. Leptostelma maxima D. Don, in Sweet, Brit. Flow. Gard., ser. 2. 1: 38. 1831. Aster maximus (D. Don) Less., Syn. Gen. Compos. 182. 1832. Erigeron maximus (D. Don) Orto ex DC., Prodr. 5: 284. 1836. Erigeron maximus (D. Don) Oto ex DC. var. minor Baker, in Martius, Fl. Bras. 6(3): 28. 1882. Erigeron maximus (D. Don) Otto ex DC. var. palustre Baker, in Martius, Fl. Bras. 6(3): 28. 1882. Type: 2 = Erigeron sulcatus DC., Prodr. 5: 284. 1836. Type: In Brasilieae pacois uliginosis, LUND (G-DC?, not seen). 4 Comp. Newsl. 46, 2008 = Erigeron alpestre GARDNER, in Hooker, London J. Bot. 4: 123. 1845. Type: Brazil, Rio de Janeiro, Organ Mountains, GARDNER 5787 (type US, photo seen). = Erigeron palustre GARDNER, 1n Hooker, London J. Bot. 4: 123. 1845. Type: Brazil, Rio de Janeiro, Organ Mountains, GARDNER 507 (type BM, not seen). = Erigeron scaberrimus GARDNER, in Hooker, London J. Bot. 7: 80. 1848. Type: Brazil, in marshy Campos near Villa do Principe, Province of Minas Gerais, Aug. 1840, GARDNER 4923 (type BM, not seen). Geographical distribution. Southeastern and southern Brazil (Minas Gerais, Espirito Santo, Rio de Janeiro, SAo Paulo, Parana, Santa Catarina and Rio Grande do Sul), Argentina (Misiones) and Paraguay (Caaguazu, Paraguari) (SOLBRIG 1962, Soria & ZARDINI 1995). Comments. Leptostelma maxima is a very common species in wetlands of southeastern and southern Brazil. It is morphologically close to L. catharinense but easily distinguished by the shape of the leaves and the size of the heads. In the protologue of the species Davip Don cited: “A native of Mexico; a plant of which was obtained by Mr. HUNNEMANN, in the autumn of 1827 from the Berlin Botanic Garden, for RoperT BARCLAY”. We believe that the type must have been deposited in the herbarium of Berlin (B) and probably destroyed during the second world war. According to SoLBRIG (1962) the reference to Mexico is incorrect, but the same author did not make comment on the native place of the species. In our opinion, the plant is native to South America. 4. Leptostelma meyeri (CABRERA) A.TELES, comb. nov. Basionym: Erigeron meyeri CABRERA, Notas Mus. La Plata, Bot. 19: 198. 1959. Type: Argentina, Prov. Chaco, Meyer 2234 (holotype LP, photo seen; isotype LIL, not seen). Geographical distribution. Argentina, Chaco and northwestern of Entre Rios (CABRERA 1974), 5. Leptostelma tucumanense (CABRERA) A.TELES, comb. nov. Basionym: Erigeron tucumanensis CABRERA, Notas Mus. La Plata, Bot. 19: 196. 1959, “tucumanense’’. Type: Argentina, “Tucuman, Dept. Chicligasta, Las Pavas, 2,500 m”, VeNTuRI 4745 (holotype LP, photo seen; isotypes GH, not seen; LIL, not seen; US, photo seen). Geographical distribution. Known only from the type locality. 6. Leptostelma tweediei (Hook. & ARN.) D. J. N. Hinp & G. L. Nesom, Kew Bull. 57(2): 478. 2002. Basionym: Erigeron tweediei Hook. & ARN., Companion Bot. Mag. 2: 50. 1836. Leucopsis tweediei (Hoox. & ARN.) BAKER, in Martius, FI. Bras. 6(3): 9. 1882. Leucopsis tweediei var. pilosa BAKER, in Martius, FI. Bras. 6(3): 9. 1882. Haplopappus tweediei (Hook. & ARN.) MALME, Ark. Bot. 24(A-6): Comp. Newsl. 46, 2008 5 42. 1931. Type: Uruguay, Maldonado, in boggy ground, TweepiE 1058 (holotype K, not seen). = Erigeron seneciiformis S. F. BLAKE, Proc. Biol. Soc. Washington 36: 51. 1923. Type: Bolivia, hacienda Rosaria, near Reyes, 4 Nov. 1921, WuiTE 1206 (holotype US, photo seen; isotype K, not seen). Geographical distribution. Brazil (Bahia, Distrito Federal, Goias, Mato Grosso, Minas Gerais, Rio de Janeiro, Sao Paulo, Parana, Santa Catarina and Rio Grande do Sul), northeastern Argentina (Chaco, Corrientes, Entre Rios, Formosa, Jujuy and Misiones), Paraguay (Caaguazi and Carapegua), southern Bolivia (Santa Cruz) and Uruguay (Montevideo and San José). (SoLBRIG 1962, CABRERA 1974). Comments. Leptostelma tweediei is morphologically close to L. maxima and differs mainly by the shorter and yellow ray florets. Key to the species of Leptostelma 1. Basal leaves rosulate. DrAWANIMOTC ES ViCMLOW) 2cxtes cceo coscecwettvseceup cause stecstecastes 6. Leptostelma tweediei PPAR AN MEOL EES WAUILE sco 5 cvaveanctdeunn emer tiautecctreras deneemeecese 4. Leptostelma meyeri 1. Basal leaves not rosulate, leaves all cauline. 3. Leaves auriculate; ray florets 1-seriate, ligules yellow..............ee esses SdGOe SSR oe ORO EEE ee ee Ce eee 5. Leptostelma tucumanense 3. Leaves not auriculate; ray florets 2-seriate, ligules white. 4. Involucre glabrous, 5—6 mm; ligules 3—9 mm. 5. Leaves sessile, oblanceolate, attenuate at the base, margin serrate-lobed; ST SS A pl eee Med ea 1. Leptostelma camposportoi 5. Leaves petiolate (petiole winged), ovate, sagittate or auriculate at the base, margin toothed or serrate; ligules 8—9 mm ..................:-cccesccceeeee PRE ies Re oRe Ceci to, SRR a aN 2. Leptostelma catharinense 4. Involucre hirsute, 12—14 mm; ligules 12—15 mm ..................eessecceeseeeees Bee Ae Se Raia Sat aap ee Bisa oct Nas avast Se ee 3. Leptostelma maxima 6 Comp. Newsl. 46, 2008 Acknowledgements We thank SuSANA Freire, Curator of the family Asteraceae of the Herbarium LP for sending digital images of some types and Guy Nesom for encouraging us to publish the combinations. We are also grateful to Emity Woop, Curator of the Herbarium of Harvard University, for bibliographic assistance, and CAPES and CNPq for the scholarship and grant to the first and last author, respectively. References Baker, J. G. 1882. Compositae: Asteroidae, Inuloideae. Jn: C. F. P. MArtius & A. G. EICHLER (eds.), Flora Brasiliensis. Vol. 6. Part. 3. Fleischer, Munich, Vienna, Leipzig. Pp. 1-134. BENTHAM, G. 1873. Compositae. /n: G. BENTHAM & J. D. Hooker (eds.), Genera plantarum. Vol. 2. Lovell Reeve, London. Pp. 163-533. Casrera, A. L. 1974. Compuestas. Jn: A. Burkart (ed.), Flora Ilustrada de Entre Rios (Argentina). Vol. 6. Colleccion Cientifica del INTA, Buenos Aires. Pp. 106-554. Cronoguist, A. 1943. The separation of Erigeron from Conyza. Bull. Torrey Bot. Club 70: 629-632. Hinp, D. J. N & G. L. Nesom 2002. A new combination in Leptostelma D.Don (Compositae: Astereae). Kew Bull. 57: 478. Nesom, G. L. 1989. Infrageneric taxonomy of New World Erigeron (Compositae: Astereae). Phytologia 67: 67-93. Nesom, G. L. 1994a. Apopyros (Asteraceae: Astereae), anew genus from southern Brazil, Argentina, and Paraguay. Phytologia 76(2): 176-184. Nesom, G. L. 1994b. Separation of Neja (Asteraceae: Astereae) from Hysterionica. Phytologia 76(2): 168-175. Nesom, G. L. 1994c. Subtribal classification of the Astereae (Asteraceae). Phytologia 76(3): 193-274. Nesom, G. & H. Rospinson 2007. XI. The tribe Astereae Cass. (1819). Jn: J. W. Kapereir & C. Jerrrey (eds), The families and genera of vascular plants. Vol. 8. Flowering plants, Eudicots, Asterales. Berlin, Springer. Pp. 284-342. Noyes, R. D. 2000. Biogeographical and evolutionary insights on Erigeron and allies (Asteraceae) from ITS sequence data. Plant Syst. Evol. 220: 93-114. Comp. Newsl. 46, 2008 Sotsric, O. T. 1962. The South American species of Erigeron. Contr. Gray Herb. 191: 3-82. Soria, N. & M. Zarpini 1995. Sinopsis de la tribu Astereae (Asteraceae) en Paraguay. Jn: D. J. N. Hinp, C. JEFFREY & G. V. Pope (eds.), Advances in Compositae Systematics. Royal Botanic Gardens, Kew. Pp. 355-378. 8 Comp. Newsl. 46, 2008 Nanothamnus sericeus THOMSON, a derived species of Blumea ARNE A. ANDERBERG! & ARUN K. PANDEY? ‘Department of Phanerogamic Botany Swedish Museum of Natural History P. O. Box 50007, SE-104 05 Stockholm, Sweden ’Department of Botany TM Bhagalpur University, Bhagalpur, 812007 India Abstract Nanothamnus sericeus THOMSON was described as a member of the tribe Mutisieae because of its distinctly bilabiate corolla. Later, the genus has often been treated as a member of the Inuleae but its position there has also been questioned. We investigated the systematic position of this small Indian endemic by jackknife analysis of DNA sequence data from the plastid gene ndhF and from the nuclear ribosomal internal transcribed spacers (ITS). It was concluded that Nanothamnus belongs to the Inuleae-Inulinae, and that it is a member of the genus Blumea, characterized by anumber of autapomorphic character states. The new combination Blumea sericea (THOMSON) ANDERB. & A. K. PANDEY 1s made. Introduction The genus Nanothamnus was described in 1867 by THOMAS THOMSON (THOMSON 1867) on material from the western Indian province Concan. The strikingly two-lipped (4+1 lobes) hermaphroditic flowers made THomson place Nanothamnus in the Mutisieae, the tribe characterized by flowers with bilabiate corolla. The floral morphology has made this genus difficult to place with certainty. BENTHAM (1873) moved the genus from Mutisieae to the Inuleae-Plucheinae and noted that its corolla made it differ from other Plucheinae, but suggested affinities with Thespidium and Epaltes. HooKER (1882) placed Nanothamnus between Pluchea and Epaltes and made the note that: “This curious plant bears some resemblance to a Blumea of the oxyodonta group in habit and foliage”. HorrMANN (1890) followed BENTHAM’ s opinion on the position of Nanothamnus, whereas NAIRNE (1894) erroneously stated that the florets in Nanothamnus were as in Pluchea indica, without mentioning the characteristic two-lipped corolla. Comp. Newsl. 46, 2008 9 MERXMULLER et al. (1977) included Nanothamnus in the Pluchea group of Inuleae- Inulinae where it has remained since, albeit with some doubt. In the first cladistic study of the Inuleae, ANDERBERG (1989) described a variation in the shape and distribution of sweeping-hairs in different groups of the Inuleae sensu MERXMULLER et al. (1977), and also showed that members of the /nu/a group were characterized by having a large oxalate crystal in each cypsela epidermis cell. Apart from its bilabiate corolla, Nanothamnus was found to resemble the Inuleae s. str. in character states such as the large elongated crystal in the epidermis cells, and acute sweeping-hairs on the abaxial surface of the style-branches. Both these features were at the time considered indications of a relationship with the Inuleae s. str. and Nanothamnus was therefore tentatively accepted in that tribe, as a possible relative of B/umea. Ina later paper, ANDERBERG (1991) modified the view, and expressed doubts on the actual position of Nanothamnus, pointing out its bilabiate disc florets, soft anther appendages, and styles with sweeping-hairs situated far below the bifurcation as anomalous character states making the taxonomic position uncertain in the tribe. THoMson (1867) had mentioned that the anthers were very shortly caudate, but ANDERBERG (1991) found them to be calcarate with short tails. Calcarate anthers are more common among taxa of the Cichorioid tribes than among the Asteroideae to which the Inuleae belongs, and this also contributed to the doubts on its systematic position in that tribe. ANDERBERG (1994) once more tentatively included Nanothamnus in the Inuleae because of its elongated cypsela epidermis crystals, but again pointed out its anomalous morphological characters and ANDERBERG & ELDENAS (2007) listed Nanothamnus among the Inuleae genera incertae sedis, possibly being a relative of Blumea, but again they pointed out its morphological differences compared to other Inuleae. Considering the many character states that make Nanothamnus differ from other Inuleae it is now of great interest to test its systematic position by analysis of DNA sequence data. Material and methods DNA was extracted from leaves taken from a herbarium specimen of Nanothamnus sericeus [Voucher: India, SARDESAI 2547 (S)]. To test the tribal position of Nanothamnus in the family, a ndhF sequence was analysed together with the data set (184 taxa) from ANDERBERG et al. (2005) and with an additional number of unpublished sequences, mainly from the Inuleae, a total of 254 sequences of taxa representing all Asteraceae tribes. An ITS sequence was analysed together with other sequences obtained from GenBank (Accession Numbers EF210932-EF210951, EF210953, EF210955-EF210962, EF210967, EF210972- EF210975). 10 Comp. Newsl. 46, 2008 Molecular methods. DNA extraction was carried with QIAGENs DNeasy Plant Mini Kit using the manufacturer’s protocol. PCR reactions were performed with PuReTag Ready-To-Go PCR Beads, 95°C 5 min, followed by 40 cycles of 95°C 30 sec, 50°C 30 sec, 72°C 1 min 30 sec, and finally 72° 8 min. Purification of PCR products was done with Omega Bio-Tek, Inc. E. Z. N. A. Cycle-pure Kit following the manufacturer’ s protocol. Sequencing reactions were made using the BigDye Terminator v3.1 Cycle Sequencing Kit and unincorporated dye terminators were removed using QIAGEN’s DyeEx 96 Kit. Fragments were separated and analysed on an ABI PRISM 3100 Genetic Analyzer. Primers used for PCR and sequencing of ndhF are presented in Table 1, and for ITS in Table 2. The two new sequences have been submitted to GenBank (EU562213 and EU562214). Alignment. Alignment of ndhF was performed with the BioEdit software (HALL 1999) ver. 6.0.5. The aligned ndhF data set included 254 taxa with several representatives of each tribe of the Asteraceae. Alignment of ITS was performed with ClustalX included in the BioEdit package using default settings. The aligned ITS data included 35 sequences from 34 taxa. Phylogenetic analyses. The aligned sequences of ndhF and of ITS were analysed with parsimony jackknifing using the software XAC (Farris 1997) with the following settings: 1000 replications, each with branch-swapping and 10 random- additions of sequences. For the analysis of ndhF, Boopis (Calyceraceae) was used as outgroup (Farris 1972). The ITS data was analysed in the same way using Duhaldea cappa as outgroup. Results The results of the XAC analysis of ndhF placed Nanothamnus in the Inuleae, together with Blumeopsis and species of the genus Blumea (Fig. 1). A jackknife support of 100 % was retrieved for a group comprised of Blumea riparia, B. saxatilis, B. integrifolia, B. psammophila, Blumeopsis flava, and Nanothamnus sericeus (Fig. 2). The sister to this group was Blumea balsamifera (76 %), at the next lower node Caesulia axillaris was sister to the rest (100 %), and then Duhaldea cappa appeared as the sister of all taxa at the base of this clade. The close relationship between Duhaldea and Blumea was discovered by ANDERBERG (1991), and ANDERBERG et al. (2005) found that also the morphologically specialized Caesulia was a member of this clade, another morphologically anomalous genus for which the systematic placement has been discussed. In a recent study, PORNPONGRUNGRUENG et al. (2007) analysed phylogenetical relationships in Blumea and showed that Blumeopsis and Merrittia were ingroups in Blumea, and this is congruent with our present results. In the small sample Comp. Newsl. 46, 2008 1] ndhF tree, Blumeopsis is sister to Nanothamnus, albeit with weak support (65 %), but their position as ingroups in Blumea is strongly supported. The analysis of ITS sequences included a larger sample of Blumea species and gave strong support (100 %) for a position of Nanothamnus in a Blumea clade together with the species B. hieraciifolia, B. mollis and B. oxyodonta (Fig. 3). Discussion We conclude that Nanothamnus is a member of the Inuleae-Inulinae as circumscribed by ANDERBERG et al. (2005), and that it is in fact a derived member of the genus Blumea. Within that genus, it shares a common ancestor with Blumea hieraciifolia, B. mollis, and B. oxyodonta (Fig. 3). It is noteworthy that HOOKER (1882) noted that Nanothamnus resembled a species of the Blumea oxyodonta group. The bilabiate flowers, the epappose fruit, the soft anther appendages, the calcarate anthers, and the sweeping-hairs extending below the bifurcation are apparently autapomorphies that have evolved in Nanothamnus, making it unique within the genus B/umea. Whereas the chromosome number in B/umea is complex with many basic numbers, the meiotic chromosome number in Nanothamnus is n=10 (DARUWALLA 1995), which would indicate a somatic number of 2n=20, and this is fully congruent with that of several Blumea species. Soft wrinkled anther appendages are rare in the family but constitute a synapomorphy of the Arctoteae- Arctotidinae (Karis 2006). The appendages in Nanothamnus are however less soft and wrinkled than those of the Arctoteae and most likely represent an independent character state transformation. Apart from the differences compared to other genera of the tribe, Nanothamnus shares some diagnostic synapomorphies with other genera of Inuleae-Inulinae (except Caesulia), such as the presence of a large oxalate crystal in the cypsela epidermis cells. It also shares the characteristic 3 bp (CCT) insertion in ndhF gene that is found in all Inuleae-Inulinae without exception, and which was described first by ELDENAs et al. (1999). An autapomorphy in Nanothamnus is also a 9 bp insertion in ndhF (AAATTAGAT) that seems to be unique in the Asteraceae. This 9 bp insertion is located 66 bases closer to the 3’ end than the unique CCT insertion. The bilabiate flowers in Nanothamnus are unique in the Inuleae. They have evidently evolved from ordinary tubular hermaphroditic flowers, possibly as an adaptation to a specific pollinator but nothing is yet known about the pollination of this species. Similar odd and sporadic occurrences of bilabiate corollas are found in representatives of many tribes outside the Mutisieae, but they may be somewhat different, and the kind found in Nanothamnus was considered unique 12 Comp. Newsl. 46, 2008 by SMALL (1918). The genetic background to the shift in corolla shape is not known but would be interesting to study. Also the lack of pappus makes Nanothamnus unique in Blumea. The lack of pappus bristles makes it more difficult for the fruits to disperse any longer distances, and that may be one reason why it is confined to a rather restricted area in western India. Nanothamnus cannot be retained as a separate genus as its closest relatives are found within Blumea. Therefore we propose that Nanothamnus is transferred to Blumea to maintain the generic monophyly of the latter. The genus Blumea now includes species from the monotypic genera Nanothamnus, Merrittia and Blumeopsis. Of these Nanothamnus may be the most surprising addition of the three. Blumea sericea (THOMSON) ANDERB. & A. K. PANDEY, comb. nov. Basionym: Nanothamnus sericeus THomson, J. Linn. Soc., Bot. 9: 342 (1867). Illustr.: Fig. 4; THoMson 1867, tab. 3. Low, strongly aromatic, generally much branched herb, branches often prostrate to ascending. Leaves alternate, oblong to ovate-lanceolate, with conspicuously reticulate venation, acutely serrate, sericeous; leaf-teeth with distinct mucro. Capitula aggregated terminally, or axillary, generally heterogamous, disciform, few-flowered, small, ovoid, 3-5 mm long and 2-4 mm wide. Receptacle flat, epaleate. Involucral bracts narrow, imbricate, in 2—3 rows. Outer florets when present, 1-4, functionally female; corolla yellow, c. 2 mm long, 2—3-dentate. Pappus lacking. Central florets 5—6, perfect, corolla yellow, deeply pseudobilabiate with 4+1 lobes. Anthers distinctly but shortly calcarate; thecae shortly caudate; anther appendage soft, somewhat wrinkled; endothecial tissue radial. Style bifid; style-branches with short, triangular, acute sweeping-hairs abaxially, below the bifurcation with longer more obtuse sweeping-hairs; style-base with prominent star-shaped crystal druses. Cypsela ovoid, dark brown with 10 distinct white ribs forming a short five-dentate ring distally; cypsela epidermis cells each with one large elongated oxalate crystal. Pappus missing. Geographic range: Blumea sericea occurs only in western India. It is an endemic of the Western Ghats, growing in the Nasik, Pune, and Shimoga districts of Maharashtra and Karnataka where it grows in dry hills among grasses in forests and forest clearings. Flowers and fruits have been seen from February to April (KumMaAR 1995). Comp. Newsl. 46, 2008 13 Acknowledgements The authors thank Dr. Mitinp Sarpesal, Pune, for providing material of Nanothamnus sericeus, Mattias Myrenas for laboratory assistance, and ELIZABETH Binktewicz for the illustration of style and stamen. Financial support was received as a Swedish Research Council grant (to A.A.) for angiosperm phylogeny. References ANDERBERG, A.A. 1989. Phylogeny and reclassification of the Inuleae (Asteraceae). Can. J. Bot. 67: 2277-2296. ANDERBERG,A.A. 1991.Taxonomy and phylogeny ofthe tribe Inuleae (Asteraceae). PI. Syst. Evol. 176: 75-123. ANDERBERG, A. A. 1994. Tribe Inuleae. /n: Bremer, K., Asteraceae. Cladistics & classification. Timber Press, Portland. Pp. 273-291. ANDERBERG, A. A., ELDENAS, P., BAYER, R. J. & M. ENGLuND 2005. Evolutionary relationships in the Asteraceae tribe Inuleae (incl. Plucheeae) evidenced by DNA sequences of ndhF; with notes on the systematic positions of some aberrant genera. Org. Div. & Evol. 5: 135-146. ANDERBERG, A. A. & P. ELDENAS 2007. Tribe Inuleae. Jn: KapereiT, J. W. & C. Jerrrey (eds.), The families and genera of vascular plants 8. Springer Verlag. Pp. 374-391. ANDERBERG, A. A. & U. Swenson 2003. Evolutionary lineages in Sapotaceae (Ericales): a cladistic analysis based on ndhF sequence data. /nt. J. Plant Sci. 164: 763-773. BENTHAM, G. 1873. Compositae. /n: BENTHAM, G. & J. D. Hooker, Genera plantarum 2(1). Lovell Reeve & Co., London. Pp. 162-533. DaruwaLLa, A. R. 1995. Cytological investigations on the Asteraceae-genus Blumea and related genera Laggera and Nanothamnus. J. Bombay Nat. Hist. Soc. 92: 314-321. ELpEnAs, P., KALLERSJO, M. & A. A. ANDERBERG 1999. Phylogenetic placement and circumscription of tribes Inuleae s. str. and Plucheeae (Asteraceae): evidence from sequences of chloroplast gene ndhF. Molec. Phyl. Evol. 13: 50-58. Farris, J. S. 1972. Estimating phylogenetic trees from distance matrices. Am. Naturalist 106: 645-658. 14 Comp. Newsl. 46, 2008 Farris, J. S. 1997. “Xac”’. Computer program and manual. Swedish Museum of Natural History, Stockholm. HAL, T. A. (ed.) 1999. BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucl. Acids Symp. Ser. 41: 95-98. HoFrMann, O. 1890. Tubuliflorae-Inuleae. /n: ENGLER, A. & K. PRANTL (eds.), Die nattirlichen Pflanzenfamilien 4(5). Engelmann, Leipzig. Pp. 172-210. Hooker, J. D. 1882. The Flora of British India 3. Caprifoliaceae to Apocynaceae. Lovell Reeve & Co., Ashford. Karis, P. O. 2006. Morphological data indicates two major clades of the subtribe Gorteriinae (Asteraceae-Actotidae). Cladistics 22: 199-221. KALLERSJO, M., Bercovist, G. & A. A. ANDERBERG 2000. Generic realignment in primuloid families of the Ericales s.].: A phylogenetic analysis based on DNA sequences from three chloroplast genes and morphology. Am. J. Bot. 87: 1325-1341. Kumar, S. 1995. Inuleae-Plucheinae. Jn: Hasra, P. K., Rao, R. R., Smncu, D. K. & B. P. UnryAt (eds.), Flora of India 13. Asteraceae (Inuleae-Vernonieae). Botanical Survey of India, Calcutta. Pp. 114-163. MERXMULLER, H., Letns, P. & H. ROESSLER 1977. Inuleae - systematic review. Jn: Heywoop, V. H., HARBoRNE, J. B. & B. L. TURNER (eds.), The Biology and Chemistry of the Compositae. Academic Press. Pp. 577-602. Naireng, A. K. 1894. The flowering plants of Western India. W. H. Allen & Co., London. PORNPONGRUNGRUENG, P., GUSTAFSON, M. H. G., BORCHSENIUS, F., ENGLUND, M. & A. A. ANDERBERG 2007. Phylogenetic relationships in Blumea (Asteraceae: Inuleae) based on molecular data. P/. Syst. Evol. 269: 223-243. SMALL, J. 1918. The origin and development of the Compositae IV. The corolla. New Phytol.17: 13-40. Tuomson, T. 1867. On two new genera of Compositae Mutisiaceae, from India. J. Linn. Soc., Bot. 9: 342-344. Comp. Newsl. 46, 2008 15 Table 1 Primer sequences (5’-3’) for ndhF. F = forward, R = reverse. Name Direction Primer sequence RJ] F AGG TAA GAT CCGGTG AAT CGGAAA C Ib lf iGG GACTIE TIC TTL rie “Cc 43] F GAT ACAAAT TTA TAT Tl FIG GG 520 R CAA-ATIG:CTT TIT GACAAG CAP TIG CCG C 5 K GTC TCA ATT GGGTTA TAT GATG 5B F GGA'GCT ACT TIA GCPEEr -G 16 R GIT AAACCT CCCATA AGC ACCATA TIC TGA:C 1260 E TCT TAA TGATAG TTG GTT GTA TTC ACC 1700 R CAT AGTATT ATC TGATTC ATA AGG ATA 1750 R ACT GAAAAA ATT GCATCT TTT 10 R CCC CCT AYA TAL TTG ATA. CEETEL Ce 10B R CCT ACT CCATTT GGAATT CCATC RJ14 R ACC AAGTTC AAT GIT AGC GAGATT AGT C All primers were published in KALLERSJO et al. (2000), except for primers 520 and 1750 that were published by ANDERBERG & SWENSON (2003). Primers RJ1 and RJ14 were designed by K1-JooNG Kim and RosBert JANSEN. Table 2 Primer sequences (5’-3’) for ITS. F = forward, R = reverse. Name ____ Direction _ Primer sequence 18SF FE GAACCTTATCGTTTAGAGGAAGG 26SR R CCGCCAGATTTTCACGCTGGGC 5.8SC F TGCGTTCAAAGACTCGAT 5.8SN R ATCGAGTCTTTGAACGCA Primers 18SF and 26SR were designed by Catarina Rypin, and primers 5.8SC and 5.8SN by YoUNGBAE Suh. 16 Comp. Newsl. 46, 2008 Boopis (Calyceraceae) BARNADESIOIDEAE 96 Adenocaulon Gochnatia Mutisia Onoseris Stifftia Leibnitzia, Gerbera, Piloselloides 001 Nassauvia, Trixis, Perezia 9S Ainsliaea Echinops Tarchonanthus Atractylodes, Carlina 001 Saussurea, Dipterocome, Cephalonoplus, Cirsium, Synurus, Carthamus, Centaurea 69 Gymnarrhena 66 Tragopogon, Cichorium, Lactuca 001 Munnozia, Liabum, Sinclairia £2 | 001 Eremothamnus, Gazania, Arctotis, Haplocarpha SS) Vernonia, Gutenbergia, Lychnophora, Pitocarpha, Stokesia 86 Senecioneae (4 taxa) 001 Calenduleae (3 taxa) 86 Gnaphalieae (4 taxa) 08 001 Anthemideae (4 taxa) 66 86 Astereae (8 taxa) Callilepis, Zoutpansbergia Athroismeae (4 taxa) OL Heliantheae s. lat. (23 taxa) SOqII] SvIPIOIOSW INULEAE - with Nanothamnus Continued in Fig. 3 avaovsa}sy Ul UOIISOd snupYyJoUuDN ¢ “‘SIq UI panuNUOS snumHyjoUDN) YM Iea[NUT IQ AuasojAyd ovaoeiaysy JO aseq ‘exe} p67 - ypu Fig. 1. Parsimony jackknife tree based analysis on ndhF sequences from 254 taxa showing the position of Inuleae with Nanothamnus (Box) in the Asteraceae phylogeny. Support values > 50 % are shown for each clade. Inuleae clade expanded and continued in Fig. 2. 17 Comp. Newsl. 46, 2008 Inuleae-Inulinae . 2. Parsimony jackknife tree (continued from Fig. 1) showing the position Inula, Pulicaria, Schizogyne, Buphthalmum, Vierea, Rhanterium, Carpesium, Telekia, Amblyocarpum, Pentanema, Chrysophthalmum, Ighermia, Asteriscus (38 taxa) Duhaldea cappa Caesulia axillaris Blumea balsamifera Blumea riparia Blumea saxatilis Blumeopsis flava NANOTHAMNUS SERICEUS Blumea integrifolia Blumea psammophila 65 100 100 99 76 100 94 100 99 Continued from Fig. 2 Jasonia, Chiliadenus, Dittrichia, Anvillea, Pallenis, Inuleae-Plucheinae —— Stenachaenium, Antiphiona, Pegolettia, Calostephane, Ondetia, Geigeria, Rhodogeron, Sachsia, Pterocaulon, Iphionopsis, Neojeffreya, Cratystylis, Adelostigma, Allopterigeron Streptoglossa, Coleocoma, Epaltes, Karelinia, Nicolasia, Pechuel-loeschea, Pluchea, Pseudoconyza, Sphaeranthus, Tessaria, Cylindrocline, Doellia, Laggera, Porphyrostemma (108 taxa) 54 Nanothamnus, position in Inuleae-Inulinae ndhF - 254 taxa , a derived representative of Blumea. Support values > 50 % are shown for each clade. of Nanothamnus sericeus in tribe Inuleae Fig 18 Comp. Newsl. 46, 2008 Duhaldea nervosa Pentanema ligneum Pentanema indicum 001 66 O01 Pentanema cernuum Caesulia axillaris ps Blumea aromatica Blumea densiflora 69 001 Blumea lanceolaria $9 Blumea arfakiana 6L Blumea canalensis 6s Blumea balsamifera Blumea virens Merrittia benguetensis Blumea sessiliflora OO €8 Blumea sinuata IL Blumea fistulosa NANOTHAMNUS SERICEUS Blumea hieraciifolia var. macrostachya 001 Blumea hieraciifolia var. hamiltonii SL voulnlq Blumea mollis +9 OO Oo a Blumea oxyodonta Blumeopsis flava Blumea psammophila Blumea saxatilis Blumea clarkei Blumea hieraciifolia 9L Blumea megacephala Blumea riparia Blumea cf. saxatilis avaovloysy Ul UOT}ISOd snuimpyjouDN Blumea lacera Blumea napifolia dvUl[NU]-dvaTNU] JO apejo vawmnyg ‘exe) SE — ST] Blumea paniculata Fig. 3. Parsimony jackknife tree based analysis on ITS sequences from 35 sequences from 34 taxa showing the position of Nanothamnus as an ingroup in Blumea ina clade with Blumea hieraciifolia, B. mollis and B. oxyodonta. Support values > 50 % are shown for each clade. Comp. Newsl. 46, 2008 19 Le are ae <= res Ses, 1.0mm 1.0mm Fig. 4. Blumea sericea. A: Style showing shortly triangular sweeping hairs on the branches and long obtuse sweeping hairs below the bifurcation. B: Stamen showing tailed and calcarate anther base. — A-B: SarpEsAI 2547 (S). 20 Comp. Newsl. 46, 2008 A new subspecies of Flaveria (Asteraceae) from Western Australia GREG KEIGHERY Department of Environment and Conservation Western Australian Wildlife Research Centre P.O. Box 51, Wanneroo, Western Australia, 6065 gregk@calm.wa.gov.au Abstract A new subspecies of Flaveria, F. australasica Hook. subsp. gilgai KEIGHERY is described. The subspecies is confined to cracking clay wetlands in the Pilbara Biogeographical Region of Western Australia. Introduction Flaveria is a genus of 21 species predominantly from North America which was revised by PoweELt (1978). In this revision all Australian populations of the genus were referred to a single species Flaveria australasica Hook. (Yellow Twin Stem). This species is a close relative of the widespread F. trinervia (SPRENG.) C. Mone, which occurs in Africa, India, the Middle East, North and South America and the West Indies. Plants of Flaveria australasica were distinguished from F. trinervia in cultivation by Powe. (1978) on the basis of their taller and more slender habit and narrower, lanceolate leaves. This species is widespread through arid, semi-arid and subtropical Western Australia, Northern Territory, South Australia, Queensland and New South Wales (Fig. 1). The Western Australian Department of Conservation and Land Management is conducting a biological survey of the Pilbara Biogeographical Region of Western Australia. This arid tropical region contains a suite of herb and grass dominated cracking clay wetlands, which flood from summer cyclonic rains and are proving to have a distinctive and highly endemic flora. These endemics are largely from widespread normally tropical genera and include undescribed herbaceous taxa of Boerhavia (Nyctaginaceae), Oldenlandia (Rubiaceae), Salsola (Chenopodiaceae) and Flaveria (Asteraceae). Comp. Newsl. 46, 2008 Herbarium material of Flaveria has been checked at PERTH, MEL and NSW, and collections in NT, AD, HO and BRIS were checked via the Australian Virtual Herbarium. This note describes and illustrates the new subspecies. Taxonomy Key (extract from PoweLL 1978, pp. 606--607) a. Pappus scales present b. Leaves perfoliate, to 4-5 cm wide (Chihuahuan Desert) ................:::ccseeees necteedichec tbat ane heen eens be Atte eines bat ssl F- chlorifolia A. Gray bb. Leaves weakly connate, to 0.7 cm wide (Arizona) ..............::cccssseceeeeneeeees DP Spee Oe a ncncck caidditi's decoded dhins F: macdougalii THEROUX, PINKAVA & KEIL aa. Pappus absent c. Receptacle of glomerule setose d. Achenes 2—2.6 mm long (widespread in North America and elsewhere). Rae ee a ts a iB nase a8 lag Sis da Boas aia Stee te ee F. trinervia dd. Achenes 2.3—4.5 mm long (Australia) .................:eeeeeee F. australasica Key to Flaveria in Australia e, Inflorescences. smnall..8—12 min: GUAM, «.n:2..5c2.5scer2-o soxesen sets seten cee eee eee ee F. australasica subsp. australasica ee. Inflorescences large, 17-25 mm diam. . F: australasica subsp. gilgai Flaveria australasica Hook. subsp. gilgai G.J. KEIGHERY, subsp. nov. (Fig. 2) Herbae annuae, robustae, 40-100 cm altae, erectae vel expansae. Caulis florifer dichotomus, 2—5 cm latus, glaber. Folia 1-5 cm longa, 4-7 mm lata, glabra, margine serrulato vel integro. Capitulum magnum, 17-20 mm latum. Achaenia 3.54 mm longa. Typus: Western Australia, Pilbara Biogeographical Region, 10.5 km. SSE of Mount Bruce, Karijini National Park, 21° 24’S 117° 03’E, 20.VHI.2006, G.J. & B.J. KEIGHERY 970 (Holo PERTH , iso CANB, K, MEL). Erect glabrous robust annual herb, 40—100 cm. tall; older stems smooth, terete, usually red or purple, younger stems smooth, quadrangular, usually green, branching more or less dichotomously towards the inflorescence, roots fibrous. i bo Comp. Newsl. 46, 2008 Leaves opposite, lanceolate to narrowly ovate, 1-5 cm long, 4-7 mm wide at midpoint, green, 3-veined, attenuate, entire or serrulate, glabrous, apex acute. Inflorescence a compound head (capitulescence) of 10—30 partial heads aggregated on a minute, flat, general receptacle. Receptacle with long slender membranous, chaff-like setae. Compound heads solitary, terminal on short branches and in the stem forks, 8-9 mm long and 17—20 mm wide with a common involucre of unequal leafy bracts longer than the head, 31—51 mm long. Heads heterogamous, radiate or disciform, involucral bracts of partial heads 2 or 3, 3-4 mm long, elliptic, obtuse, inrolled. Outermost partial heads comprising a solitary female floret with a yellow ligule ca. 1 mm long, hardly protruding from the bracts; style branches obtuse, glabrous. Inner partial heads with 2—6 tubular bisexual florets; corolla 5-lobed; anthers obtuse at base with ovate apical appendages; style branches truncate with papillose apex. Achenes black, narrowly obovate, 10-ribbed, slightly flattened, ca. 3.5-4.5 mm long by 0.6 mm wide; pappus absent. This subspecies differs from Flaveria australasica subsp. australasica in being a robust plant with broad stems, larger leaves, larger inflorescences, longer leaves and bigger achenes. Since it differs in quantitative characters and occupies a distinct ecological niche (where the two taxa do not co-occur) within the overall range of the nominant subspecies (Figs. | and 3), recognition at subspecific level seems most appropriate. Other specimens examined (all PERTH): Hamersley Railway Line, Dampier to Tom Price, 21° 40’S 117° 14’E, X.1968, J.G. CAMPION s.n.; Hamersley Railway Line, Dampier to Tom Price, 21° 41’S 117° 14’E, X.1968, E.B.J. Smiru s.n.; North of Tampanna Bore, Coolawanyah Station, 21° 37’S 117° 44’E, 4.1X.1996, A.A. MitcHett 1476; 10.5 km. SSE of Mount Herbert, Millstream-Chichester National Park, 21° 24’S 117° 03’E, 20.V.1997, M.E. TRUDGEN 15605. Distribution: Confined to the central Pilbara Biogeographic Region in arid Western Australia. This distribution is entirely within the range of Flaveria australasica subsp. australasica (Fig. 1). Habitat: Recorded as occurring on cracking red clay soils in tussock grasslands of Astrebla pectinata or mixed grasslands/herbfields dominated by Polymeria longifolia, Astrebla elymoides and Dichanthium sericeum. Flowering Period: Flowering occurs from August to September. Mature fruits and seeds are found from October to December. Conservation Status: The subspecies is recorded from Karijini and Millstream- Chichester National Parks, but most of the region is uncleared leased grazing lands. Comp. Newsl. 46, 2008 33 Etymology:The subspecific name denotes the habitat of this taxon. From Ghilgai, an Aboriginal word denoting a soil from inland Australia characterized by a markedly undulating surface with mounds and depressions caused by swelling and cracking of clays during alternate wet and dry seasons. Reference PoweE.LL, A. M. 1978. Systematics of Flaveria (Flaveriinae-Asteraceae). Ann. Missouri Bot. Gard. 65: 590--636. 24 Comp. Newsl. 46, 2008 Fig. 1. Distribution of Flaveria australasica subsp. australasica. Comp. Newsl. 46, 2008 A.B Flaveria australasica subsp. gilgai (G. & B. KeicHery 970, PERTH). A: Flowering branch. B: Achene. C.D Flaveria australasica subsp. australasica (TRUDGEN 494, PERTH) C: Flowering branch. D: Achene. 25 26 Comp. Newsl. 46, 2008 ae 4a] | Fig. 3. Distribution of Flaveria australasica subsp. gilgai. Comp. Newsl. 46, 2008 7 The status of generic revision in the African Vernonieae (Asteraceae) M. A. IsAwuMI Natural History Museum, Obafemi Awolowo University Ile-Ife, Nigeria ayoisawumi@yahoo.com Abstract The tribe Vernonieae is reviewed for Africa and found to consist of 35 genera and 299 species. There is no doubt that more new genera or resurrected genera will be established for the tribe in the Eastern Hemisphere. About eleven subtribes have been established for Neotropical Vernonieae and three subtribes for Paleotropical Vernonieae. There are 10 species and 3 infraspecific taxa which are new combinations in this study. They are found in the genera Orbivestus (3 spp.), Vernoniastrum (4 spp.), Linzia (2 spp., 1 subsp., 1 var.), Gymnanthemum (1 sp., 1 var.). The recent segregation from Vernonia of new genera in each of the two Hemispheres has given a great support for the different phyletic lines that have been proposed for the genus Vernonia in the Old World and the New World. The remaining Vernonia elements in Africa that have not been placed in any new or elevated genera have been estimated to be about 59 taxa. Introduction The tribe Vernonieae is abundantly represented in the New World, particularly in Brazil where the diversity of genera and species is remarkable. In the Old World, numerous species are found in Africa, a secondary centre of distribution, and Madagascar, Southern Asia, and the Malay Archipelago (Jones 1977). In 1977 about 70 genera and 1,456 species were recognized, while 98 genera and 1,300 species were reported in 1994 for the tribe (TURNER 1977, BREMER 1994, NorDENSTAM 1998, IsAwuMI 1996a). However, RoBINSON (1996) recognized 111 genera worldwide with the expectation of a few further additions in the Western Hemisphere. More recently Rospinson (2007) recognized 118 genera disposed in 15 subtribes. There is no doubt that more new subtribes and genera or resurrected genera will be established for the tribe especially in the Eastern Hemisphere (IsawumI 1999), 28 Comp. Newsl. 46, 2008 Apams (1963) reported ten genera including Vernonia Scures. for the tribe in West Africa, whereas Jerrrey (1988) recognized 13 genera in East Africa. However, IsawuMI (1999) recognized 15 genera for the tribe in West Africa including Vernonia, Baccharoides Moencu and Cyanthillium BLUME. RoBINSON (1999, 2007) has established three subtribes for the Eastern Hemisphere Vernonieae based upon the reference works of many authors. He pointed out that the works of KirKMAN (1981), Pope (1983), Koyama (1984), Isawumi (1993, 1995c), and Isawum et al. (1996) have added important information. It has been generally agreed by authors that there are some basic differences between the Vernonieae in the Eastern and Western Hemispheres (Jones 1977, 1981, JerrREY 1988, IsAwumi 1993, 1995c, RoBinson 1999). The Hemispheric trends noted by Jones (1977) are in chromosome numbers and chemistry. The Western Hemisphere species usually have a chromosome number of n = 17, and the Eastern Hemisphere species have mostly n= 9 or 10. RoBiNsoNn & KAHN (1986) and Isawum et al. (1996) briefly noted the apparent restriction of glandular dots on the anthers and their appendages to New World members of the Vernonieae. Rosinson (1999) noted the presence of elongate raphids mostly in the Eastern Hemisphere species while characteristically subquadrate or short raphids are found in the Western Hemisphere Vernonieae. BOHLMANN & Jakupovic (1990) found the distinctive 5-alkyl coumarins only in Eastern Hemisphere genera and species, taxa that were all placed in the new subtribe Erlangeinae by RoBinson (1999). In earlier times the tribe has been the dumping ground for poorly known taxa of the Compositae. Such taxa included Liabum Apvans., Pseudostifftia H. Ros., Moquinia DC., Hoplophyllum DC., and Corymbium L. Liabum and its relatives have now been recognized as members of the distinct tribe Liabeae (Cass.) Ryps. (RYDBERG 1927, RoBINSON & BRETTELL 1973a, b, 1974, NORDENSTAM 1977, BREMER 1994). The two genera Pseudostifftia and Moquinia have been transferred to the new tribe Moquinieae by Rosinson (1994). Karis (1992) has shown that the South African Hoplophyllum is related to Eremothamnus Hore. in the tribe Arctoteae, although these two genera were placed by Ropinson (1994) in the tribe Eremothamneae. The genus Corymbium of South Africa (revised by WeITz 1989) has also been regarded as not belonging to Vernonieae on the basis of chemistry (BOHLMANN & Jakupovic 1990). As a result of molecular studies by PANERO & Funk (2002), Corymbium was placed in a new subfamily Corymbioideae and a new tribe Corymbieae. Some elements added to the tribe Vernonieae or confirmed as members of the tribe are Stokesia L’HEr. and Trichospira KUNTH (ROBINSON 1996). The most interesting additions to the tribe were the various synonyms of Distephanus Cass. (Gongrothamnus StEEvz, Newtonia HorrM. and Antunesia Horro.) that were rather consistently placed in the Senecioneae in the traditional Comp. Newsl. 46, 2008 29 systems of classification (BENTHAM 1873, HOFFMANN 1893) because of their yellow florets and trinervate leaves. Their placement in the Vernonieae was established by Rosinson & KAHN (1986) and others. Subtribal classification of the African Vernonieae The new and re-established subtribes proposed by Rosinson et al. (1980) and RoBINSON (1996) comprise primarily Western Hemisphere Vernonieae. In order to rectify the situation, RoBINSoN (1999) named three subtribes for the Eastern Hemisphere species, and more subtribes can be expected to be recognized in both hemispheres. The two subtribes Vernoniinae and Elephantopinae are also represented in the Eastern Hemisphere (cf. below). Erlangeinae H. Ros. Type: Erlangea plumosa Scu.Bir., Flora 36: 34. 1853. The name, according to RosBinson (1999), is chosen to conform with the already established term “Erlangeoid” (Pore 1983). Characteristically, the subtribe includes Vernonieae with mostly triporate pollen and 5-alkyl coumarins (BOHLMANN & Jakupovic 1990). It also includes genera with 4—5 angled achenes and a herbaceous habit (with few exceptions having 10-ribbed achenes and a woody habit). All the included genera have acicular sweeping hairs of the style. Chromosome number n = 9, 10, 20. The genera included are the New World Acilepidopsis H. Ros. (1989), Mesanthophora H. Ros. (1992) and Telmatophila Marr. ex Bak. (1873), and the Old World genera Acilepis D. Don, Ageratinastrum Mattr., Ambassa StTEE1Z, Bechium DC., Bothriocline OLIv. ex BENTH., Brachythrix WILD & Pope, Cyanthillium BLuME, Decastylocarpus HumBertT, Dewildemania O. HorrM., Diaphractanthus Humpert, Erlangea Scu. Bie., Ethulia L.r., Gutenbergia Sco. Bip. ex Watp., Herderia Cass., Hystrichophora Matrtr., lodocephalus THOREL ex GAGNEP., Kinghamia C. JEFFREY, Lamprachaenium Bentu., Msuata O. Horrm., Muschleria S. Moore, Omphalopappus O. Horrm., Paurolepis S. Moore, Phyllocephalum Buume, Polydora FENzL, Rastrophyllum WiLb & G. V. Porr, and seven genera newly named or elevated by RoBInson (1992, 1999), viz., Cabobanthus H. Ros., Hilliardiella H. Ros., Kovamasia H. Ros., Mesanthophora H. Ros., Orbivestus (S. B. Jones) H. Ros., Oocephala (S. B. Jones) H. Ros., Vernoniastrum H. Ros. Centrapalinae H. Ros. Type: Centrapalus galamensis Cass., Dict. Sci. Nat. ed. D2 382. VENT. The genera of the subtribe are herbaceous or weakly shrubby, and the sweeping hairs of the styles are acicular. The sesquiterpene constituents of the species include elemanolides (BOHLMANN & JAKupovic 1990). The subtribe includes elements with the distinctive Linzia-type pollen (JEFFREY 1988) such as Linzia 30 Comp. Newsl. 46, 2008 and Aedesia, and one element with polar lacunae on its pollen grains and a lack of basal stylar nodes, like Baccharoides (IsAwuM! 1993, IsAwumi et al. 1996). Chromosome number n = 9, 10. The subtribe is typified by Centrapalus Cass. and also includes Adenoon DaAtz., Aedesia O. HorrmM., Baccharoides MoENCcH, Camchaya GaGneP., Lachnorhiza A. Ricu., Linzia Scu. Bip. ex WALP., Neurolakis Matte. and Pleurocarpaea BENTH. Gymnantheminae H. Ros. Type: Gymnanthemum cupulare Cass., Bull. Soc. Philom. Paris 1817: 10.1817. [= G. coloratum (WiLLD.) H. Ros. & B. KAHN]. The subtribe includes all of the true large shrub and tree Vernonieae in the Eastern Hemisphere. The sweeping hairs of the styles often have rather blunt tips. Inner involucral bracts are persistent or deciduous. Stellate hairs are lacking. Chromosome number is n= 9, 10, 15, 20. The subtribe has these genera: Gymnanthemum Cass., Distephanus Cass., Centauropsis Bos. in DC., Oliganthes Cass., and three genera named or elevated by Rosinson (1999), viz., Brenandendron H. Ros., Lampropappus (O. HorrM.) H. Ros. and Myanmaria H. Ros. In addition there is one New World genus, Hesperomannia A. Gray, which is endemic to Hawaii (RoBINsON 2007). Generic classification of the African Vernonieae The first comprehensive work on Vernonieae in West Africa is the key to the genera and species of the tribe by ApAMs (1963). He reported ten genera including Vernonia for the tribe in West Africa, whereas JEFFREY (1988) recognized 13 genera and 126 species in East Africa. IsawuM! (1999) recognized 15 genera in West Africa including the new resurrected genera Baccharoides and Cyanthillium. OLIVER & HIERN (1877) reported 78 species for Africa, and Jones (1977) recorded c. 200 for the continent. ADAMs (1963) reported 60 species for West Africa and this number was reduced to 42 by Isawumi (1996a,b). In the present report about 59 Vernonia species remain untransferred to new, resurrected or elevated genera. Also 35 genera and 299 species are presently recognized for Africa in the tribe. Comp. Newsl. 46, 2008 ull Table 1. List of accepted genera in the tribe Vernonieae in Africa with approximate number of species, general distribution, habit and chromosome number. No. 10. Genera Vernonia SCHREB. Baccharoides MOoENCH Cyanthillium BLUME Elephantopus L. Pseudelephantopus ROHR Sparganophoros VAILL. Ethulia L.f Adesia O. HoFrM. Herderia Cass. Erlangea Scu. Bip. Number of Species ay 25 13 Distribution West & East Africa, Southern Africa West, East & Central Africa, South Africa West Africa, East Africa, South Africa, Madagascar West Tropical Africa, East & Central Africa, Senegal, Macias, Nguema (Fernando Poo) Introduced to East & West Tropical Africa (Native of Neotropics) West, East & Central Africa, Madagascar West, East & Central Africa, South Africa Tanzania, West & Central Tropical Africa West Africa Guinea, Sierra Leone, Tanzania, Ethiopia, Somalia Habit Annual or Perennial Herb Shrub Herb Herb Herb Herb Annual or Perennial Herb Perennial Herb Annual Annual or Perennial Herb Cromosome number n=9or 10 n=10 n=9 n=10 n=10 n=10 32 26. 21. Bothriocline OLIv. ex BENTH. Centratherum Cass. Gutenbergia Scu. Bip. Kinghamia C. JEFFREY Hystrichophora Mattr. Rastrophyllum WILD & Pope Dewildemania O. HoFrM. Distephanus Cass. Ageratinastrum MatTr. Brachythrix WILD & PoPE Msuata O. HorFeM. Stephanolepis S. Moore Bechium DC. Cabobanthus H. Ros. Hilliardiella H. Ros. Orbivestus (S.B. JONEs) H. Ros. Oocephala (S. B. Jones) H. Ros. 31 13 25 West, East & Central Africa, South Africa, Madagascar West Africa West, East & Central Tropical Africa, Southern Africa West Tropical Africa, Zaire Tanzania East Tropical Africa East Tropical Africa South & East Africa, Madagascar Tanzania, Malawi, Zambia Tanzania, Mozambique Tropical Africa Tropical Africa Madagascar Central Africa South Africa, West & Central Tropical Africa West, Central & Southern Africa West, East & Central Africa Comp. Newsl. 46, 2008 Annual or Perennial Herb Annual Herb Annual or Perennial Herb Herb Herb Annual Perennial Small shrubs or vines Perennial Perennial Shrub Perennial Herb Herb Herb Undershrub Shrub n=9, 10 n=9, 20 Comp. Newsl. 46, 2008 33 East, West & Annual < 28. Polydora FENZL 8 GeniAaies Hed n=9 Vernoniastrum Pty ey 29. Hoon 12 Central & Herb n= 10 ; : Southern Africa Cater Southern, West, | Annual or 30. Cis 2 8 East, Central & Perennial ' North Africa Herb West, East, Linzia Central & ™ 31. Scu. Bip. ex WALP. ? South A frica, Herb n— 0 Madagascar South, West, East, Central & 30° 1 ae 30 North Africa, cai ms n= 10, 20 : Africa to Sudan, Madagascar Brenandendron West, East & A 23: H. Ros. : Central Africa Shrub n=? Lampropappus East & Central he (O. Horr.) H. Ros. : Africa Sa Manyonia ; Perennial 355 Fi Bon. i Tanzania Heb In order to redefine the Paleotropical members of Vernonia, RoBINSON & KAHN (1986), Rosrnson (1990a), IsAwumi (1993, 1995a,b), Isawumr et al. (1996) transferred some Vernonia species to the three resurrected genera Distephanus, Baccharoides and Cyanthillium. Rosinson & KAHN (1986) separated Distephanus from Vernonia and recognized 26 species in the genus. The genus is distinct from the genus Vernonia because the species have yellow flowers, with broad sclerified basal appendages on the anther thecae, apical appendages without glands and style base with large abruptly broadened node. Pollen in many species are “Type A” (KEELEY & Jones 1979), with continuous intercolpar perforated tectum and reticulately arranged spines. Some Madagascar species have irregularly lophate pollen grains with perforated tectum restricted to lower sides and bases of crests, with distinct colpi intruded upon by short alternating spurs of reticulate tectum (ROBINSON & KAHN 1986). Baccharoides MorENcH Rosinson (1990a) resurrected the genus Baccharoides and transferred three species of Vernonia to the genus. IsAwumi (1993) transferred twelve species of Vernonia to the genus Baccharoides on the basis of the style base being surrounded by the 34 Comp. Newsl. 46, 2008 nectary and inner involucral bracts with expanded foliose appendages, which may be white or variously coloured. Isawum! et al. (1996) recognized 25 species and 26 infraspecific taxa in the genus Baccharoides on the basis of morphological characters such as the endothecial tissue being “polarized” as described by DoRMER (1962). In the pollen the oral (poral) lacunae (EL-GHAZALY 1980) are joined with the aboral lacunae by broad interlacunar gaps, thus producing the three long colpi running nearly from pole to pole. Each colpus encloses a central compound os with two perpendicular ora and is bounded on either side by the paraoral ridges (IsawumI et al. 1996). This type of aperture with two ora is referred to as colpororate instead of colporate. The columellae in the genus Baccharoides are stout, branched distally and anastomose within the ridges (Iswumi 1995a, IsAwuMI et al. 1996). In this group the oral lacunae are more or less indistinct, and therefore markedly different from the ones found in other groups of Vernonia s.1. In addition, the colpus is long and the ora are not in a well defined constriction. JEFFREY (1988) has pointed out that the Baccharoides group is “one of the most distinct Old World taxa of Vernonieae and the case for its recognition at generic rank is undeniable”. I strongly attest to this fact because it was formerly a distinct entity as Vernonia section Stengelia. The diagnostic characteristics are remarkably different from those of Vernonia s.l. So, the criticism of AYODELE & OLorobE (2005) of the separation of Baccharoides from Vernonia s.1. by ROBINSON (1990a) and Isawumr et al. (1996) could not be sustained mainly on adaptation strategies alone because the group was resurrected as a genus based on many characters. Such re-classification is necessary so that the genus Vernonia may gradually reduce in heterogeneity and size and eventually become a monophyletic genus. As a result, many genera are being separated from Vernonia s.l. in the Neotropical and Paleotropical regions of the World as evidenced in the works of RoBinson (1996, 1999, 2007). Cyanthillium BLUME RoBINsON (1990a) transferred three species of Vernonia to the resurrected genus Cyanthillium on the basis of 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. On the basis of floral microcharacters Isawum! (1995b) accepted the establishment of the genus before transferring four varieties of C. cinereum to the genus. The genus, in contrast to the genus Baccharoides and the genus Vernonia s.l., has no colpus and the endothecial tissue is intermediate unlike the one in Baccharoides which is distinctly polarized (Isawum1 1995a, IsAwuMI et al.1996). The filament collar in Cyanthillium is elongate and straight unlike the ones in Baccharoides which are more or less shortly cylindrical and sometimes somewhat dilated distally (Isawumi 1995a). Comp. Newsl. 46, 2008 35 Genera of African Vernonieae Subtribe Erlangeinae H. Ros. Elevated or Named by Rosinson (1999) and Some New Combinations Bechium DC., Prodr. 5: 70. 1836. Type: Bechium scapiforme DC. There are two species in the genus and both occur in Madagascar. Cabobanthus H. Ros. Type: Vernonia polysphaera BAKER. There are two species and they occur in Zambia, Tanzania and Congo. Cyanthillium Buume, Bijdr. Fl. Ned. Ind. 889. 1826. Type: Cyanthillium villosum BLUME. Seven species are presently placed in the genus and they occur in Africa and Madagascar. Hilliardiella H. Ros. Type: Vernonia pinifolia Less. The genus with eight species is closely related to Cyanthillium, but it differs in its more perennial habit, the non-lophate pollen and the presence of T-shaped hairs of the corolla [except Cyanthillium cinereum (L.) H. Ros. var. ugandense (C. JEFFREY) IsAwuMI which also has T-shaped trichomes on its corolla (IsAWUMI 1995b)]. Chromosome number n= 9, 10 (Jones 1982). Distribution: South Africa, Congo, Tanzania, Malawi, Mozambique, Zambia, Ethiopia, West and Central Tropical Africa. Orbivestus (S. B. Jones) H. Ros. Type: Vernonia karaguensis Outv. & HIERN. The genus is distinct from the related Cyanthillium by the more shrubby habit, the lack of simple hairs on the corolla lobes, and non-lophate pollen (ROBINSON 1999). Chromosome number n = 9, 20 (Jones 1982, MANGENOT & MANGENOT 1962, Menra et al. 1965). Distribution: Angola, Botswana, Kenya, Somalia, Sudan, Mozambique, Nigeria and West Africa. Four species were placed in the genus by Rosinson (1999). Three further species are placed in the genus in this study. They are the following: Orbivestus blumeoides (Hook. f.) IsawuMi, comb. nov. Basionym: Vernonia blumeoides Hook. f. in J. Linn. Soc. Bot. 7: 198. 1864. Type: Cameroon, MANN 1241. 1921 (syntype K). An erect little-branched undershrub. Leaves pubescent with T-shaped trichomes. Capitula campanulate; phyllaries 4-seriate; corolla lobes and tubes with capitate and stipitate glands of various shapes and sizes. Anther calcarate; apical anther appendage ovate—oblong; stylar ring (stylopodium) about 6 cells per row with annular thickenings on the walls. Achene 5-costate, with large carpopodium, 36 Comp. Newsl. 46, 2008 glabrous with idioblasts; pappus caducous, biseriate, outer elements shorter and broader than inner, barbellate setae, glabrous at the base. Distribution: Nigeria, Cameroon. Orbivestus bamendae (C. D. ADAMs) IsAwUMI, comb. nov. Basionym: Vernonia bamendae C. D. Apvams in J. W. Afr. Sc. Assoc. 3: 116. 1957. Type: Cameroon, MAITLAND 1514 (holotype K!). A robust herb with broad sessile leaves. Leaves pubescent with T-shaped trichomes. Capitula campanulate; phyllaries 5—6-seriate; corolla lobes with capitate and stipitate glands. Anther calcarate; apical anther appendage spear- shaped, ovate-oblong; stylar ring about 5 cells per row. Achene 4-costate with idioblasts, glabrescent with biseriate trichomes; carpopodium moderately large; pappus biseriate, caducous, outer elements broader and shorter than inner, barbellate setae, glabrous at the base. Distribution: Nigeria, Cameroon. Orbivestus albocinerascens (C. JEFFREY) IsSAwUMI, comb. nov. Basionym: Vernonia albocinerascens C JEFFREY in Kew Bull. 42(2): 222. 1988. Type: Uganda, Karamoja District, Mt. Moroto, southern foothills, Tweepir 2643 (holotype K). It is similar to Orbivestus cinerascens (Scu. Bip. in SCHWEINF.) H. Ros. as mentioned by Rosinson (1999). Distribution: Uganda, Kenya, Tanzania. Oocephala (S. B. Jones) H. Ros. Type: Vernonia oocephala BAKER. Low much-branched shrubs to | m high, stems with weakly L-shaped simple hairs and with multiseptate simple hairs. Florets about 15 in a head. Anther bases rounded; apical anther appendages glabrous, with thin wall cells; style base with indistinct ring; style branches with acicular sweeping hairs. Achenes weakly 8-ribbed, idioblasts numerous, raphids narrowly elongate; pappus biseriate, outer shorter and broader, inner setiform, subplumose, glabrous near base. Two species. Distribution: Burundi, Congo, Tanzania, Nigeria, Mozambique. Polydora FENZL, Flora 27: 312. 1844. Type: Polydora stoechadifolia FENZL. Mostly annuals; stems with one-armed T-shaped hairs; anthers untailed, pollen triporate; chromosome number n = 9 (Jones 1979, 1982). The genus is credited with eight species (ROBINSON 1999). Distribution: Tanzania, Mozambique, Zambia, Zimbabwe, Congo, Malawi, Angola, Burundi, Botswana, Namibia, Transvaal, Sudan, Ethiopia, South Africa, West Africa to Nigeria. Comp. Newsl. 46, 2008 37 Vernoniastrum H. Ros. Type: Crystallopollen latifolium STEETz in PETERS. The genus is closely related to Polydora FeNz, but differs by the perennial habit, the non T-shaped hairs, the tailed anther bases and the chromosome number of n= 10. The core element of the genus also has the idioblasts of the achene in distinct transverse bands. The genus was credited with eight species by Rosinson (1999). Distribution: Kenya, Tanzania, West Africa to Sudan, South to Angola, Congo, Mozambique, Zimbabwe, Zambia, Burundi, Ethiopia, Somalia, Malawi. Four species are added here as new combinations: Vernoniastrum paraemulans (C. JEFFREY) ISAwUMI, comb. nov. Basionym: Vernonia paraemulans C. JerFREY in Kew Bull. 43(2): 246. 1988. Type: Tanzania, Iringa District, Iringa, MILNE-REDHEAD & TayLor 11150 (holotype K!). It is similar to Vernoniastrum aemulans (VatTKeE) H. Ros. as implied by JEFFREY (1.c.) and mentioned by Rosinson (1999). Distribution: Tanzania. Vernoniastrum migeodii (S. Moore) IsawuMi, comb. nov. Basionym: Vernonia migeodii S. Moore in J. Linn. Soc. Bot. 35: 319. 1902 Type: Nigeria, MiGeop (holotype BM). A perennial herb with sessile leaves. Leaves pubescent with flagelliform trichomes without T-shaped types. Corolla lobes with long uniseriate trichomes, glandular with capitate and biseriate glands. Apical anther appendage spear-shaped; style base peg-like. Achene inconspicuously 5-costate; pappus biseriate, outer elements scale-like and broader than inner, with barbellate setae. Distribution: Nigeria, Benin, Ghana, Cameroon. Vernoniastrum klingii (O. Horrm. & Muscut.) IsawuM1, comb. nov. Basionym: Vernonia klingii O. Horrm. & Muscut. in Mem. Soc. Bot. Fr. 2, 8c: 112. 1910. Type: Guinee, CHEVALIER 15743 (syntype P), KLING 25 (syntype B), BUETTNER 9 (syntype K). Plant perennial; capitula campanulate; phyllaries 4-seriate. Corolla turbinate; corolla lobes glandular with capitate glands, pubescent at tips without T-shaped hairs; stylar ring about 3 cells per row with thin cell walls. Achene 8-costate, glandular on the furrows. Pappus biseriate, outer elements shorter and broader than inner, barbellate setae. Distribution: Guinea, Ghana, Sierra Leone, Benin, Cote dIvoire. 38 Comp. Newsl. 46, 2008 Vernoniastrum camporum (A. Cuey.) IsAwUMI, comb. nov. Basionym: Vernonia camporum A. CueEv. in Mem. Soc. Bot. Fr. 2, 8e: 259. 1917. Type: Guinee, CHEVALIER 20408 (holotype P). Phyllaries 6—7-seriate. Corolla lobes with capitate glands. Stylar ring about 4 cells per row. Achene 8-costate, with idioblasts, pubescent with biseriate trichomes. Pappus biseriate, outer elements shorter than inner ones, persistent with barbellate setae, glabrous at the base. Distribution: Nigeria, Guinea, Sierra Leone, Cameroon, Ghana, Senegal. Resurrected and New Genera of African Vernonieae Subtribe Centrapalinae H. Ros. and Some New Combinations Centrapalus Cass., Bull. Soc. Philom. Paris 1817: 10. 1817. Type: Centrapalus galamensis Cass. Annual or perennial, scapose or subscapose herbs; stem hairs simple, multiseptate; involucres hemispherical, bracts 125—150 in about 5—6 series, linear, green, often with small teeth on distal margin; corolla lobes sometimes fringed with long papillae; style base with broad node. Achenes weakly 10-costate, setuliferous; raphids narrowly oblong. Chromosome number n = 9 (Jones 1974, 1979, 1982). Nine species were placed in the genus by Rosinson (1999). Distribution: Sierrra Leone, Nigeria, Congo, South Africa, Angola, Zimbabwe, Sudan, Malawi, Tanzania, Zambia, Mozambique, Uganda, Ethiopia. Linzia Scu. Bie. ex WALP. Rep.2: 948. 1843. Type: Linzia vernonioides Scu. Br. ex WALP. Seven species were credited to the genus by Rosinson (1999). Distribution: Congo, Burundi, Kenya, Tanzania, Angola, Namibia, Mozambique, Swaziland, South Africa (Transvaal, Natal), Madagascar. The following two species and two infraspecific taxa are new combinations : Linzia gerberiformis (O.tv. & HIERN) H. Ros. subsp. macrocyanus (O. HOFFM.) ISAWUMI, comb. nov. Basionym: Vernonia macrocyanus O. Horr. in Bol. Soc. Brot. 13: 20. 1896. Type: Angola, Catumba, WELWITSCH 3883 (isotype BM!). Syn.: Vernonia gerberiformis Ouiv. & HIERN subsp. macrocyanus (O. HOFFM.) C. JEFFREY in Kew Bull. 43: 234. 1988. ROBINSON (1999) regarded V. macrocyanus as a synonym of Linzia gerberiformis. Comp. Newsl. 46, 2008 39 JEFFREY (1988) gave it subspecies status while Isawumi (1995c) maintained its species status. It is different from Linzia gerberiformis because subsp. macrocyanus has (a) appendage-like upper part of phyllary triangular-lanceolate, acuminate, (b) apical anther appendage about 0.6 mm long, ovate, (c) style branches slightly coiled with swollen shaft below, (d) style base without a ring, peg-like (ISAwUMI 1995c, Fig. 9). The synonymy of Linzia gerberiformis subsp. macrocyanus 1s as cited by JEFFREY (1988) and Isawumi (1995c). Distribution: Nigeria, Cameroon, Mali, East and South Africa, Angola. Linzia ituriensis (Muscuu.) H. Ros. var occidentalis (C. D. ADAMS) ISAWUMI, comb. nov. Basionym: Vernonia glabra (StEE1Z) VATKE var. occidentalis C. D. ADAMs in J. W. Afr. Sc. Assoc. 3, 1: 119. 1957. Type: Nigeria, ONocHIE in FHI 34855 (holotype K). Syn.: Vernonia ituriensis MuSCHL. var. occidentalis (C. D. ADAMS) C. JEFFREY in Kew Bull. 43: 231. 1988. Phyllaries 6—7-seriate, pectinate-ciliate at margins, black at tips, shorter than those of Linzia ituriensis (MuscuL.) H. Ros. Corolla 12-14 mm long; corolla lobes with unicellular trichomes sparsely concentrated at the tips and with long stipitate glands. Anther base weakly calcarate with blunt ends; apical anther appendage more or less elliptic with flat end and solid line in the middle. Style branches coiled; stylar ring with 2—3 cells per row. Pappus orange, biseriate, persistent, outer setae shorter and broader than inner, barbellate (see Isawumi 1995c, Fig. 13). Distribution: Nigeria, Cameroon. Linzia nigritiana (Ou1v. & HIERN) [sAwuMI, comb. nov. Basionym: Vernonia nigritiana OLIV. & HIERN in FI. Trop. Afr. 3: 288. 1877. Type: Nigeria, Abeokuta, IRVINE (syntype FHI!). Perennial herb; phyllaries 6—8-seriate, lanceolate, pectinate-ciliate at tips. Corolla lobes funnel-shaped, apically sparsely stiffly hairy, with flagellate, unicellular and uniseriate trichomes. Anther basally shortly calcarate; apical anther appendage about 0.8 mm long, more or less triangular, glabrous; filament collar distinct with thickened cells at cross-walls. Style branches coiled; style base peg-like. Achenes turbinate, strongly 10-costate, hispid with biseriate hairs; carpopodium rather large; pappus biseriate, persistent, orange, the outer shorter, of barbellate setae. 40 Comp. Newsl. 46, 2008 Distribution: Senegal, Gambia, Mali, Sierra Leone, Cote d’ Ivoire, Ghana, Nigeria, Cameroon. Linzia purpurea (Scu. Bip. ex WALP.) ISAwUMI, comb. nov. Basionym: Vernonia purpurea Scu. Bir. ex WALP. in Repert. Bot. Syst. 2: 946. 1843. Type: Ethiopia, ScHimper 1197 (holotype P!), Synonymy as cited by JEFFREY (1988) and Isawumi (1995c), plus Centrapalus purpureus (Scu. Bir. ex WALP.) H. Ros., Proc. Biol. Soc. Wash. 112(1): 236. 1999. Perennial herbs; phyllaries 6—8-seriate, triangular-lanceolate, obscurely pectinate- ciliate at margins, arachnoid-pubescent, sharply apiculate, dark at tips. Corolla funnel-shaped; corolla lobes covered with capitate glands interspersed with unicellular trichomes, with some trichomes concentrated at the tips of the lobes. Anther basally calcarate; apical anther appendage ovate oblong; filament collar distinct with cells thickened on the walls; stylar ring about 24 cells per row. Achenes 10-costate, with numerous idioblasts, pilose with biseriate hairs; carpopodium large; pappus biseriate, yellowish, of persistent barbellate setae. Distribution: Senegal, Mali, Ghana, Nigeria, Cameroon, Guinea Bissau; East, Central and South Tropical Africa. Newly Resurrected or Described Genera of African Vernonieae Subtribe Gymnantheminae H. Ros. and Some New Combinations Distephanus Cass., Bull. Soc. Philom. Paris 1817: 151. 1817. Type: Conyza populifolia Lam. Synonymy as stated by RoBINSON & KAHN (1986). RosBINsON & KAHN (1986) and Rosinson (1999) transferred 27 species into the genus of which 25 species are in Madagascar and continental Africa, one in China and one in Yunnan. Gymnanthemum Cass., Bull. Soc. Philom. Paris 1817: 10. 1817. Type: G. cupulare Cass. (= Baccharis senegalensis Pers. = Gymnanthemum coloratum (WILLp.) H. Ros. & B. KAHN). Synonymy as cited by Rosinson (1999). The genus is credited with 43 species by Rosinson (1999) but 14 of them do not occur in Africa and Madagascar. In this paper one further species and one infraspecific taxon are recognized. So, 30 species and one infraspecific taxon are referred to the genus in Africa and Madagascar. Gymnanthemum auriculiferum (HiERN) IsAwUMI, comb. nov. Basionym: Vernonia auriculifera Hiern in Cat. Welw. Afr. Pl. 1: 539. 1898. Type: Angola, WELWITSCH 3258 (syntypes BM, K, isotypes BR, K). Comp. Newsl. 46, 2008 4] Synonymy as cited by JEFFREY (1988) and IsAwumi (1995Sc). Shrub. Capitula tubular and spindle-shaped with a single floret; phyllaries 6—7- seriate, slightly tomentose at the base of the outermost bracts with one short vein at the tips; outer ovate, hyaline-margined, acute, inner elliptic, apiculate, caducous. Corolla tubular; corolla lobes undulate at margin. Anther basally calcarate; apical anther appendage ovate oblong; filament collar distinct with cells thickened on the walls; stylar ring about 5 cells per row. Achene 10-costate, with idioblasts, pilose with biseriate hairs; pappus biseriate with the outer setae shorter, white, caducous, barbellate. Distribution: Nigeria, Cameroon, East and Central Africa. Gymnanthemum theophrastifolium (ScHweinF. ex Outv. & HIERN) H. Ros. var. richardianum (QO. KUNTZE) IsAwuMI, comb. nov. Basionym: Cacalia richardiana O. Kuntze in Revis. Gen. Pl. 2: 967. 1891. Type: Ethiopia, Gondar, DILLON (holotype P!). Syn.: Vernonia richardiana (O. KUNTZE) P.-SERMOLLI 1n Webbia 7: 340. 1950, V. theophrastifolia ScHwetnr. ex Ovtv. & HIERN var. richardiana (O. KUNTZE) IsawumI, in Advances in Comp. Syst. Kew p. 103.1995. Other synonymy as cited by Isawumi (1995c) and Rosinson (1999). A shrub or a small tree. Capitula turbinate-campanulate, about 10-flowered; phyllaries 8-seriate, glabrous to tomentose at the base of the outermost bracts with the peduncle tomentose; outer ovate, obtuse, successively shorter; inner elliptic, acute, with broad hyaline margins, caducous. Corolla tubular, corolla lobes pubescent. Anther base shortly calcarate, apical anther appendage ovate. Stylar ring about 5 cells per row with the cells heavily sclerified. Achene with idioblasts, obscurely 8—10-costate, pilose with biseriate hairs; pappus biseriate, uniform, dense, of scabrid setae. JEFFREY (1988) recorded this taxon as a synonym of Vernonia theophrastifolia and RoBINsoN (1999) concurred with this view. Isawumi (1995c) regarded it as a variety of V. theophrastifolia. It is still recognized as a variety here because of the following differences: (a) It has simple and biseriate trichomes on the corolla lobes while the corolla lobes of G. theophrastifolium are glabrous. (b) It has broader hyaline margins on the phyllaries. (c) The head is 10- flowered while it is 8-flowered in theophrastifolium. (d) Corolla lobes in richardianum are pubescent with short simple trichomes while they are glabrous in theophrastifolium. 42 Comp. Newsl. 46, 2008 (e) The pappus is biseriate and uniform but in theophrastifolium the outer setae are shorter than the inner ones(see IsawuMI 1995c, Fig. 24). Distribution: Cameroon, Nigeria, Ghana, East Tropical Africa. Brenandendron H. Ros. Type: Vernonia titanophylla BRENAN. Ropinson (1999) pointed out that Brenandendron is closely related to Gymnanthemum but it differs by the distinctive frondiform inflorescence. He placed three species in the genus. Distribution: Sierra Leone, Liberia, Guinea, Cameroon, Sudan, Central Tropical Africa. Lampropappus (O. Horr.) H. Ros. Type: Vernonia lampropappa O. Horr. Rosinson (1999) placed three species in the genus. Distribution: Angola, Congo, Malawi, Zambia. Genera of Eastern Hemisphere Vernonieae Subtribe Elephantopinae Less. (1830) Herbs. Involucre decussate with 4, 6 or 8 bracts; florets 4, corollas often zygomorphic; apical anther appendage thin, glabrous. Achene raphids elongate. Four genera are placed in the subtribe, two of which occur in Africa, viz., Elephantopus L. and Pseudelephantopus Rorr. Elephantopus L. Sp. P1.: 814 (1753) & Gen. Pl., ed. 5: 355 (1754). Type: E. scaber L. (India). The genus is distinguished by having alternate leaves and pappus of 5 to many straight bristles; cypsela with several twin hairs and idioblasts but without glands; endothecial cells intermediate; n= 11. The African species of E/lephantopus were revised by PuiLipson (1938), the East African ones by JEFFREY (1988) and the West African ones by IsAwumi (1999). About twelve species are credited to the genus world-wide while five of them occur in Africa (JEFFREY 1988). Distribution: West, East and Central Africa, Southern Africa, pantropical. Pseudelephantopus Rour corr. C. F. BAKER in Trans. St. Louis Acad. Sci. 12: 55. 1902, nom. cons. Pseudelephantopus Rone in Skr. Naturhist.-Selsk.2(1): 214. (1792), nom. invalid. Type: P. spicatus (AuBL.) C. F. BAKER in Trans. St. Louis Acad. Sci. 12: 45. 1902. Comp. Newsl. 46, 2008 43 Pappus with the bristles flexed or twisted, cypsela with twin hairs and capitate glands but without idioblasts; n = 13 (JEFFREY 1988). Two species are credited to tropical America, and one species to Africa. Distribution: Native of the Neotropics, introduced to West & East tropical Africa and tropical Asia. Genera of Eastern Hemisphere Vernonieae Subtribe Vernoniinae Cass. ex Dumort. (1829) The majority of the genera of subtribe Vernoniinae are found in Western Hemisphere, including Vernonia Scures. Just a few of them occur in Eastern Hemisphere, viz Manyonia H. Ros. and Sparganophoros VAILL. Herbs, weak shrubs or vines. Inflorescence often seriate- or scorpioid-cymose; inner involucral bracts usually persistent. Anther appendages thin-walled, often with glands or hairs. Pollen mostly tricolporate, lophate or non-lophate. Manyonia H. Ros. in Proc. Biol. Soc. Wash., 112 : 224.1999. Type: Vernonia peculiaris Verve. in Kew Bull. 1956: 447. 1956. Only one species was placed in the genus by Rosinson (1999). Distribution: Tanzania. Sparganophoros VAILL. in K6nigl. Akad. Wiss. Paris Phys. Abh. 5: 368. 1754; Boeuo. in Lupwic, Defin. Gen. Pl. ed. Boehm.: 154 (sphalm. ‘Sparganophorus’) 560. 1760. Type: S. vaillantii CRANTz (India). Only one species has been placed in the genus. Sparganophoros sparganophora (L.) C. JEFFREY in Kew Bull. 43: 272. 1988. Synonymy: Ethulia sparganophora L., Sp. Pl. ed. 2: 1171. 1763. Struchium sparganophorum (L.) O. Kuntze in Revis. Gen. Pl. 1: 366. 1891. Distribution: West and East Africa, Madagascar, moist tropics of Malaysia, South India, Sri Lanka, New World, pantropical and widely adventive. Vernonia Scures., Gen. 2: 541. 1791, nom. cons. Type: Vernonia noveboracensis (L.) WILLD. (North America). The new genera and species hitherto elevated to generic rank are those that seem inescapable (RoBINSON 1999) and the remaining elements in Vernonia of the Eastern Hemisphere that do not fit in the genera presently recognized will hopefully be disposed of in future studies. The remaining elements in the Eastern Hemisphere Vernonia will be transferred to other genera because it appears that 44 Comp. Newsl. 46, 2008 the genus Vernonia will be restricted to the Western Hemisphere. About 59 taxa remain in African Vernonia as follows: Vernonia aurantiaca (O. HorrM.) N. E. Br., V. holstii O. Horrm., V. ruvungatundu C. Jerrrey, V. kigomae C. JErFRey, V. tricholoba C. Jerrrey, V. pteropoda Oty. & Hiern, V. nuxiodes O. HorrM. & Muscut., V. bruceae C. Jerrrey, V. stuhlmannii O. HorrM., V. fischeri O. HorrM., V. biafrae Outv. & Hiern var. biafrae, V. biafrae O.tv. & Hiern var. tufnelliae (S. Moore) Isawumi, Vo angulifolia DC., V. syringifolia O. Horrm., V subscandens R. E. Fr., V. brachycalyx O. Horro., V. suprafastigiata Kuatt, V. brachytrichoides C. Jerrrey, V. turbinata Ov. & Hiern, V. homilantha S. Moore, V. vollesenii (’vollensenii’’) C. JerrReEY, V. mikumiensis C. JEFFREY, V. natalensis Scu. ex WALp., V. alticola G. V. Pore, V. wollastonii S. Moore, V. hochstetteri Scu. Bip. ex WALP., V. muelleri WiLb, V. roseoviolacea De Witb., V. wakefieldii Outv., V. kandtii Muscut., V. amblyolepis Bak., V. schweinfurthii Outv. & HIERN, V. luembensis De Witpb. & Muscut., Vv. mbalensis G. V. Pore, V. rhodanthoidea Muscut., V. amoena S. Moore, V. nyassae Ouiv., V. schliebenii Mattr., V. popeana C. Jerrrey, V. tinctosetosa C. JerFreY, V. plumbaginifolia FENZL ex Outv. & Hiern, V. violaceopapposa Dre WiLb., V. parapetersii C. JerrRey, V. violacea OLtv. & Hiern, V. miombicola H. Witb, V. miombicoloides C. JEFFREY, V. acuminatissima S. Moore, V. melanocoma C. JerrreY, V. abbotiana O. HOFFM., V. afromontana R.E. FR. var. pleiocephala Cuiov., V. alboviolacea MUSCHL., V. eminii O. HorrM., V. teitensis O Horrm., V. bauchiensis Hutcu. & DAtz., V. djalonensis A. Cuev., V. jaegeri C. D. Apams, V. nimbaensis C. D. Apams, V. chapmanii C. D. Apams, V. andohii C. D. AbAms, V. mokaensis MILppr. & MattTr. Conclusion The present trend in the tribe Vernonieae has been work towards a full generic revision of the tribe with description of new genera and restoration of some older genera that had been placed in synonymy. Much work has been done with the generic revision of the Neotropical members of the tribe with more work still expected to be done on Paleotropical taxa. RoBiNson (1996) recognized 111 genera in the tribe worldwide, while in this study about 35 genera are recognized for Africa. ROBINSON et al. (1980) recognized eight subtribes in the subtribal classification of the Vernonieae worldwide. Bremer (1994) omitted the two monotypic subtribes of RoBINSON et al. (1980) which are Pseudostifftiinae and Trichospirinae. Later Rosinson (1996) recognized seven subtribes omitting Pseudostifftiinae. All the subtribes are related to the Vernonieae in the Western Hemisphere. ROBINSON (1999) described three new subtribes for Eastern Hemisphere Vernonieae and Comp. Newsl. 46, 2008 4S they are Erlangeinae, Centrapalinae and Gymnantheminae. Recently RoBiNson (2007) recognized 118 genera in 15 subtribes. The number of subtribes and genera are likely to increase with further studies especially on the Eastern Hemisphere members of the tribe. Vernonieae are believed to have originated in the Eastern Hemisphere where the most divergent elements of the tribe, yellow-flowered and trinervate leaved Distephanus occur (BREMER 1994). The Tropics are believed to be the origin of the tribe, since that is the centre of diversity, the area where its primitive species occur, and the region in which the majority of its genera are located (Jones 1977, IsawuMI 1999, Rosinson 1996). According to Jones (1977), 19 endemic genera are found in Africa, and four in Madagascar. Rosinson (2007) accounts for 28 genera endemic to Africa and five genera restricted to Madagascar. The Eastern Hemisphere is also the area where almost all the Vernonieae with x = 9 or 10 occur. The higher chromosome numbers found in the Western Hemisphere Vernonieae follow the pattern of higher numbers in members of group invading new geographical areas (ROBINSON 1996). Acknowledgement I would like to thank Bertit NorpenstaM for carefully revising the paper and for all other useful inputs by him while preparing the manuscript. References Apams, C. D. 1963. Compositae. Jn: HutTcHinson, J. & J. M. DatzieL, Flora of West Tropical Africa, 2°‘ ed. (F. M. Hepper, ed.) 2: 271-283, Crown Agents, London. AyopEeLe, M. S. & O. OLoropE 2005. A biosystematic evaluation of the relationship between three allopatric shrubby species of Vernonia SCHREB. (Asteraceae) in Nigeria. Comp. Newsl. 42: 8-25. BENTHAM, G. 1873. Vernonia.- In: BENTHAM, G. & J. D. Hooker (eds.), Genera Plantarum 2, 1 : 227-231. L. Reeve & Co., Williams Morgate, London. BoHLMANN, F, & J. JAkuPovic 1990. Progress in the chemistry of the Vernonieae (Compositae). Plant Syst. Evolution Suppl. 4: 3-43. Bremer, K. 1994. Asteraceae. Cladistics and classification. Timber Press, Portland, Oregon. Dormer, K. J. 1962. The fibrous layer in the anthers of the Compositae. New Phytol. 61: 150-153. 46 Comp. Newsl. 46, 2008 EL-Guaza_y, G. 1980. Palynology of Hypochaeridinae and Scolyminae (Comp. ). Opera Bot. (Nord. J. Bot. Suppl. Ser.) 58, Copenhagen. HorrMann, O. 1893. Compositae de Africa portugueza. Bol. Soc. Brot. 10: 170—-182(1892”). Isawumi, M.A. 1993. New combinations in Baccharoides MoENCH (Vernonieae: Compositae) in West Africa. Feddes Repert. 104(5-6): 309-326. Isawumi, M.A. 1995a. A new species of Baccharoides (Asteraceae: Vernonieae) from Uganda, East Africa. Comp. News!. 26: 1-10. Isawumt, M.A. 1995b. Floral microcharacters and taxonomy of the Cyanthillium cinereum complex (Asteraceae: Vernonieae). Comp. News!. 26: 11-25. Isawumi, M.A.1995c. Notes on Vernonia (Vernonieae: Compositae) in West A frica. In: Hinp, D.J.N. , JEFFREY, C. & G.V. Pore (eds.), Advances in Compositae Systematics, pp.51—106. Royal Botanic Gardens, Kew. Isawumi, M.A.1996a. Infrageneric classification of Vernonia (Vernonieae: Compositae) in West Africa. Jn: Hinp, D.J.N. & H. Beentse (eds.), Compositae: Systematics. Proceedings of the International Compositae Conference, Kew, 1994 (D.J.N. Hinp, Editor-in-Chief), Vol.1: 531-544, Royal Botanic Gardens, Kew. Isawumi, M.A. 1996b. Trichome types and their taxonomic value at the sectional and species level in West African Vernonia (Compositae). Jn: Hinp, D.J.N. & H. Beentse (eds.), Compositae: Systematics. Proceedings of the International Compositae Conference, Kew, 1994 (D.J.N. Hinp, Editor- in-Chief), Vol.1: 29-39. Royal Botanic Gardens, Kew. Isawumt, M.A. 1999. Floral microcharacters and taxonomy of the tribe Vernonieae: Asteraceae (excluding Vernonia Scures.) in West Africa. Feddes Repert. 110: 359-374. Isawumi, M.A., EL-Guazay, G. & B. Norpenstam 1996. Pollen morphology, floral microcharacters and taxonomy of the genus Baccharoides MOENCH (Vernonieae: Asteraceae). Grana 35: 205-230. JEFFREY, C. 1988. The Vernonieae in East Tropical Africa. Notes on the Compositae:V. Kew Bull. 43: 195-277. Jones. S. B. 1974. Vernonieae (Compositae) chromosome numbers. Bull. Torrey Bot. Club 101: 31-34. Jones, S. B. 1977. Vernonieae — systematic review. /n: HEywoop, V. H., HARBORNE, J.B. & B.L.Turner (eds.), The Biology and Chemistry of the Compositae I, pp. 503-521. Academic Press, London, New York. Comp. Newsl. 46, 2008 47 Jones, S. B. 1979. Chromosome numbers of Vernonieae (Compositae). Bull. Torrey Bot. Club 106: 79-84. Jones, S. B.1981. Synoptic classification and pollen morphology of Vernonia (Compositae: Vernonieae) in the Old World. Rhodora 83: 59-75. Jones, S. B. 1982. Compositae. /n: A. Love, IOPB Chromosome number reports LXXIV. Taxon 3: 119-128. Karis, P. O. 1992. Hoplophyllum DC., the sister group to Eremothamnus O. Horro. (Asteraceae)? Jaxon 41: 193-198. KEELEY, S. C. & S. B. Jones 1979. Distribution of pollen types in Vernonia (Vernonieae: Compositae). Syst. Bot. 4: 195-202. KIRKMAN, L. K. 1981. Taxonomic revision of Centratherum and Phyllocephalum (Compositae: Vernonieae). Rhodora 83: 1—24. Koyama, H. 1984. Taxonomic studies in the Compositae of Thailand 3. Acta Phytotax. Geobot. 35: 49-S8. MANGENOT, S. & G. MANGENOT 1962. Enquete sur les nombres chromosomiques dans une collection de especes tropicales. Revue Cytol. Biol. Veget. 25: 411-447. Meura, P. N., Git, B. S., Menta, J. K. & S. S. Sipnu 1965. Cytological investigations on the Indian Compositae. 1. North Indian taxa. Caryologia 18: 35-68. MeERXMULLER, H. 1954. Compositen-Studien IV: Die Compositen-Gattungen Siidwestafrikas. Mitt. Bot. Staatssamml. Miinchen |: 357-443. NORDENSTAM, B. 1998. Advances in Compositae Systematics. /n: MATHEW, P. & M. Stvatsan (eds.), Diversity and Taxonomy of Tropical Flowering Plants, pp. 113-133. Mentor Books, Calicut. O.iver, D. & W. P. Hiern 1877. Compositae. /n: Otiver, D. (ed.), Flora of Tropical Africa 3: 266—297. Ashford. PANeERO, J. L. & V. A. Funk 2002. Toward a phylogenetic subfamilial classification for the Compositae (Asteraceae). Proc. Biol. Soc .Wash. 115: 909-922. Pore, G. V. 1983. Cypselas and trichomes as a source of taxonomic characters in the Erlangeoid genera. Kirkia 12: 203-231. Rosinson, H. 1989. Acilepidopsis, anew genus of Vernonieae from South America (Asteraceae). Phytologia 67: 289-292. Rosinson, H. 1990a. Six new combinations in Baccharoides MoreNcu and Cyanthillium BLuMe (Vernonieae: Asteraceae). Proc. Biol. Soc. Wash. 103: 248-253. 48 Comp. Newsl. 46, 2008 Rosinson, H. 1990b. Studies in the Lepidaploa complex ( Vernonieae: Asteraceae). VII. The genus Lepidaploa. Proc. Biol. Soc. Wash. 103: 464-498. Rosinson, H. 1992. Mesanthophora, a new genus of Vernonieae (Asteraceae) from Paraguay. Novon 2: 169-172. Rosinson, H. 1994. Notes on the tribes Eremothamneae, Gundelieae and Mogquineae with comparisons of their pollen. Jaxon 43: 33-44. Rosinson, H. 1996. The status of the generic and subtribal revisions in the Vernonieae. /n: Hinp, D. J. N. & H. J. BEENTJE (eds.), Compositae: Systematics. Proceedings of the International Compositae Conference, Kew, 1994. (D. J. N. Hinp, Editor-in- Chief) vol.1. pp. 511-529. Royal Botanic Gardens, Kew. Rosinson, H. 1999. Revisions in paleotropical Vernonieae (Asteraceae). Proc. Biol. Soc. Wash. 112: 220-247. Rosinson, H. 2007. Tribe Vernonieae Cass. (1819). /n: Kuprrzki, K. (ed.), The Families and Genera of Vascular Plants, Vol. 8. Flowering Plants, Eudicots, Asterales (ed. J. W. KADEREIT & C. JEFFREY), pp. 149-174. Springer; Berlin, Heidelberg, New York. Rosinson, H. & R. D. BRETTELL 1973a. Tribal revisions in the Asteraceae. II. The relationship of Trichospira. Phytologia 25: 259-261. Rosinson, H. & R. D. BRETTELL 1973b. Tribal revisions in the Asteraceae. III. A new tribe, Liabeae. Phytologia 25: 404407. Rosinson, H. & R. D. BretTeLL 1974. Studies in the Liabeae (Asteraceae). I].Preliminary survey of the genera. Phytologia 28: 43-63. Rosinson, H. & B. KAHN 1986. Trinervate leaves, yellow flowers, tailed anthers, and pollen variation in Distephanus Cassini (Vernonieae: Asteraceae). Proc. Biol. Soc. Wash. 99: 493-501. Rosinson, H., BOHLMANN, F. & R. M. Kinc 1980. Chemosystematic notes on the Asteraceae. III. Natural subdivisions of the Vernonieae. Phytologia 46: 421-436. Rypperc, P. A. 1927. (Carduales) Carduaceae, Liabeae, Neurolaeneae, Senecioneae (pars.). North American Flora 34(4): 289-360. Turner, B. L. 1977. Summary of the Biology of the Compositae. Jn: HEywoop, V. H., Harsorne, J. B. & B. L. TurNeER (eds.), The Biology and Chemistry of the Compositae I, pp. 1105-1181. Academic Press, London, New York. Weitz, F. M. 1989. A revision of the genus Corymbium (Asteraceae). S. Afr. J. Bot. 55: 598-629. Comp. Newsl. 46, 2008 49 Leaf anatomical studies in some species of the tribe Cichorieae (Asteraceae) in Nigeria A. E. ADEGBITE Department of Biological Sciences University of Agriculture P. M. B. 2240 Abeokuta, Nigeria eadegb@unaab.edu.ng Abstract Leaf epidermal studies were carried out on five species belonging to four genera of the tribe Cichorieae in the family Asteraceae. The species are, Lactuca capensis, Lactuca sativa, Launaea taraxacifolia, Picris humilis and Sonchus oleraceus. All the species possess anomocytic stomatal type and expressed considerable similarities and overlap in their epidermal characters, which prove their genetic and evolutionary relationship. Introduction The family Asteraceae is represented in West Tropical Africa by about thirteen tribes. The tribe Cichorieae is characterized by possession of flowers that are all ligulate and anastomosing lactiferous vessels (LAWRENCE 1951). The tribe Cichorieae (Lettuce tribe) is represented by five genera in West Tropical Africa namely Lactuca, Launaea, Crepis, Sonchus and Picris with only Crepis not reported in Nigeria (HuTCHINSON & DatzieL 1963). The diagnostic features of members of the tribe Cichorieae include: production of milky latex (easily observed when wounded or touched), florets are all ligulate and hermaphroditic, and achenes which may be beaked or shortly beaked or not beaked. The achenes are usually with pappus. HutTcHINsON & DALzicL (1963) reported seven species of Lactuca in West Tropical Africa, but they are now six, with the changing of Lactuca taraxacifolia to Launaea taraxacifolia (JEFFREY 1966). The only two species reported in Nigeria are Lactuca capensis THunB. (Southern and Northern Nigeria) and L. intybacea Jaca. (Northern Nigeria). The genus Launaea, represented by one species in HUTCHINSON & DatziEL (1963), now contains two species which are Launaea nudicaulis L. and L. taraxacifolia 50 Comp. Newsl. 46, 2008 (WILLb.) AMIN ex C. JEFFREY. Only L. taraxacifolia has been reported in Nigeria (Southern and Northern Nigeria). Picris humilis DC. is the only representative of the genus Picris in West Tropical Africa including Nigeria (Southern and Northern Nigeria). The genus Sonchus has ten species in West Tropical Africa, five of which are reported in Nigeria. The five species which are confined to Northern Nigeria are S. oleraceus L., S. angustissimus Hoox., S. chevalieri (O. Horrm. & Muscut.), S. elliotianus HIERN and S. rarifolius OLtv. & HIERN. The species considered in this study are Lactuca capensis, L. sativa L., Launaea taraxacifolia, Picris humilis and Sonchus oleraceus, which are the available representatives of the four genera reported in Nigeria. The choice of these species was greatly influenced by the fact that some of them have no other generic relative in Nigeria as is the case of L. taraxacifolia and P. humilis, or that the related species in the same genera as L. capensis and S. oleraceus could not be located at the time of this study. All the species studied are herbaceous weeds except L. sativa (cultivated lettuce) and they all express at least one of the weedy behaviours that characterize colonising species described by BAKER (1965) and HILL (1977). Lactuca capensis, P. humilis and L. taraxacifolia are perennial herbs, while L. sativa and S. oleraceus are annual herbs. Various characters have been used in describing and delimiting species of the tribe Cichorieae, but not much attention has been given to the leaf epidermal characters. The present study therefore attempts to describe the leaf epidermal characters of the Nigerian species of the tribe in relation to their habitat to throw more light into the taxonomic grouping and the evolutionary relationship of the species. Materials and Methods The propagules of the plants used for this study were collected during field trips in different parts of Nigeria. The plants were raised to maturity in the screen house and the Botanical Garden, Abeokuta, from where leaf samples were collected for the study. Table | shows the sources and other collection data of the accessions. Sizeable portions of the mature leaves of the species were obtained from the standard median level of the leaves. The portions were decolorized by boiling in 90 % ethanol and cleared in 2 % Sodium hydroxide solution. The cleared leaves were then washed thoroughly in water before storing in 50 % ethanol (OLATUNII 1983). Epidermal peeling, sectioning, staining and mounting followed the techniques Comp. Newsl. 46, 2008 5] adopted by OcunpipE & ADEGBITE (1991) and ApeGspite (1995). Abaxial leaf surfaces of the species were studied since the species do not vary much in their adaxial epidermal characters. Stomatal frequency per mm? was estimated from an average of ten counts and guard cell area was calculated from a sample of thirty measurements of guard cells per species. Length and width of guard cells were measured at x400 magnification using an ocular micrometer. The measurements were later converted to microns using a stage micrometer. The guard cell area was estimated by using FRANco’s (1939) formula, which is — Guard cell area = Length x Width x K Where K = FRANCO’s constant = 0.78524 The stomatal index (1) was obtained by expressing the number of stomata per unit area as a percentage of the total number of cells in the same unit area i.e. I=S/E+S ~x 100 Where I = Stomatal Index S = Number of stomata per unit area E = Number of ordinary epidermal cells in the same unit area Results Table 2 shows the data on the epidermal characters of the five species studied while Figure | shows the diagrams of the lower epidermis of the species. There are different arrangements of guard cells (in relation to the subsidiary cells) which produce characteristic stomatal types that can be used for taxonomic purposes at the generic, family or higher levels. Using the classification terms proposed by METCALFE & CHALK (1979), all the species studied possessed anomocytic stomatal type where the stoma Is surrounded by a limited number of cells (usually 3—6) that are indistinguishable in size or form from other cells of epidermis. Intraspecific variations in stomatal density were observed between Northern and Southern Nigeria collections of Lactuca capensis (Fig. 1A and B). The stomatal density was observed to be higher in the Southern Nigeria collections than in Northern Nigeria collections. The guard cell area ranged between 186.41 + 23.04um~? in L. capensis and 204.70 + 19.90um? in S. oleraceus. The guard cell area was found to be comparable in L. capensis, L. sativa, Launaea taraxacifolia and Picris humilis with measurements ranging from 186.41 + 23.04 um’ to 188.80 + 28.52um*. These four species have been reported to be diploids (ADEGBITE & OLORODE 1990). Nn i) Comp. Newsl. 46, 2008 Launaea taraxacifolia showed lowest values for stomatal density and stomatal index. Discussion The variation in stomatal density which often is a reflection of physiological response to a combination of environmental factors could be useful in delimitation at the species level. While stomatal density varies considerably with the age of the leaves, stomatal index is highly constant for a given species (METCALFE & CHALK 1979). The data and the figures revealed variations in stomatal size, stomata index, stomatal density, guard cell area and in the size, shape and density of the epidermal cells of the species studied. These can be of taxonomic importance. The similarities and overlaps observed in the stomatal type, guard cell area and epidermal cell arrangement of the species provide evidence for their genetic and evolutionary relationship and justification for their taxonomic grouping. The intraspecific variations observed in guard cell area and stomatal density of the species are probably due to differential physiological responses to climatic and edaphic factors in the habitat of the various accessions. This is particularly true in view of the differences observed in guard cell area and stomatal density between Southern and Northern Nigeria collections of some of the species. Guard cell area and stomatal density values are usually reduced in the Northern Nigeria accessions in response to the dry weather in the area as a mechanism to reduce water loss through transpiration. Sonchus oleraceus is known to be a tetraploid (StesBins & al. 1953, Hsien & al. 1972, ApEGBITE & OLoropE 1990). The polyploid status accounts for the bigger value of guard cell area recorded for S. oleraceus when compared with the guard cell area values recorded for its diploid relatives. References ApvEGBITE, A. E. 1995. Leaf epidermal studies in three Nigerian species of Aspilia TuHouars (Heliantheae-Asteraceae) and two hybrids. Nig. Jour. Bot. 8: 25-33. ApecsiTE, A. E. & O. OLoropDE 1990. Karyotype and meiotic studies of some Nigerian species of Cichorieae (Asteraceae). [fe Journal of Science 4 (1-2): 84-100. Comp. Newsl. 46, 2008 53 Baker, H. G. 1965. Characteristics and mode of origin of weeds. /n: Baker, H. G. & G. L. Stepsincs (eds.), The Genetics of colonising species. Academic Press, New York. FRANCO, C. 1939. Relation between chromosome number and stomata in Coffea. Bot. Gaz. 100: 817-827. Hit, T. A. 1977. The Biology of Weeds. The Institute of Biology’s Studies in Biology, No. 79, Edward Arnold (Publishers) Ltd. London. Hsien Tsun-Suin, SCHOOLER, A. B., ALLYN BELL & J. D. NALEWAJA 1972. Cytotaxonomy of three Sonchus species. Amer. Jour. Bot. 59 (8): 789-796. HuTCcHINson, J. J. & J. M. DAuLzieL 1963. Jn: F. N. Hepper (ed.), Flora of West Tropical Africa, Vol. 2. (2° ed). Crown Agents, London. JEFFREY, C. 1966. Notes on Compositae |: The Cichorieae in East Tropical A fria. Kew Bull. 18 (3): 427-486. Lawrence, G. H. 1951. The Taxonomy of Vascular Plants. The Macmillan Company, N. Y. MetTcatre, G. R. & L. CHALK 1979. Anatomy of the Dicotyledons. Vol. | (2™ed.). Clarendon Press, Oxford. OcunpirPkE, O. T. & A. E. Apecsite 1991. The leaf epidermal studies of some species of Aspilia THouars (Asteraceae). Fedde '’s Repertorium 102 (7-8): 597-594. Ouatunst, O. A. 1983. Practical manual for plant Anatomy. (2™ ed.). (Unpublished). STEBBINS, G. L., JENKINS, J. A. & M. S. WaLrers 1953. Chromosomes and phylogeny in the Compositae tribe Cichorieae. Univ. Calif: Publ. Bot. 26 (6): 401-430. 54 Comp. Newsl. 46, 2008 Table 1. Sources and distinguishing characters of the materials used Ibe ; Ace. Location/ 5a Species a ene Pats hs Distinguishing Characters : Leaves toothed, spathulate or 13 ae 14 ; ea oblanceolate, sessile, alternate, : nem ras SO ONES | high seed set, ligule purple; Derived Savanna. e een, ki pappus white. ame ee Leaves similar to 1-3, ligule 13-16 Oghamoso,.0ya Gea deep purple, pappus yellow, Derived Savanna. nigtise dae Abandoned farmland 31-22 along Ipetumodu TTC Similar to 13 but with deep ; Ode Omu road, Osun lobing of leaves. State, Guinea Savanna. Roadside along Ifon- Leaves Eee ADS) | Frroad Ovo state toothed, low floret number per Lactuca capensis Coe ete St . head (5—9), ligule mauve or THUNB. deep purple. Disturbed land at Dele Adeymo Avenue, lies 26 Cubemasey Oe CS Similar to 13. Derived Savanna. Rest house vicinity 28 Igbeti Oyo State, Similar to 13. Guinea Savanna. beak? Leaves relatively narrow. Waste place, Universit : : Me 493/10 | of a State. ‘ nignhy ae Heule lent jc adl canis Coen purple, pappus white, poor seed ; sesete Leaves longer and broader than 493/12 | Same as 493/10. 493/10 and comparable to 13, but deeply lobed. Leaves obovate, glabrous, Lactuca sativa 0 Seon Bes sessile, truncate apex, wavy Savanna. leaf margin, ligule yellow, high seed set. Comp. Newsl. 46, 2008 55 Cultivated area at the foot of Hill 1, Obafemi Leaves deltoid-pinnatifid, sessile, alternate, glabrous with 4-7 | Awolowo University eS a ae eee Campus, Osun State. Dea ee : seed set. Derived Savanna. Undeveloped plot along Olugbode St. Ile-Ife, tl a? Osun State. Derived/ euleatele Guinea Savanna. Roadside along Armoko-Ado Ekiti Leaves relatively smaller and 10-11 sts ; bap road, Ekiti State. Guinea | narrower than 4. taraxacifolia Savanna. (WILLD.) AMIN Undeveloped land, ex C. JEFFREY Apake area, Ogbomoso, | .._. iH Oyo State. Derived ee Savanna. Farmland, Ahoro-dada, 27 Ogbomoso, Oyo State. | Similar to 4. Guinea Savanna. Stadium area, Sil Ogbomoso, Oyo State. | Similar to 4. Guinea Savanna. Waste place, University Hose \agitaiciieant Heme Leaves deeply lobed and finely toothed. State. Dry Savanna. Trans motel area, Leaves basal rosette, sessile, 12 Ogbomoso, Oyo State. | nearly linear spathulate, Guinea Savanna. multiple scapes, ligule yellow. Disturbed area, Ede- Ejigbo road, Osun Leaves typically spathulate, 18-19 ; : : : au a State. Guinea/Derived single scape, ligule yellow. Picris humilis Sanyaniia. DE: Abandoned land, 23 Oluponna, Oyo state. Same as 12. Derived Savanna. Rest house vicinity, 29-30 | Igbeti, Oyo State. Same as 12. Derived Savanna. Comp. Newsl. 46, 2008 pinnatifid, auriculate, sessile; 56 ; Waste places, Re es lyrate or deltoid- 493/2 | of Jos, Plateu State. limclc vee = Upland/Sudan Savanna. oe ee » COPIOUS Se production. Sonchus me oleraceus L. 493/3 | Same as 493/2 Similar to 493/2 493/4 493/5 493/11 Comp. Newsl. 46, 2008 57 Table 2. Data on epidermal leaf characters of the Cichorieae species studied Species Stomatal type Guard cell area (um?) Stomatal density (mm’) Stomatal index (1) Epidermal cell shape Lactuca capensis Anomocytic 186.41+23.04 652+33 23.81 Highly wavy cell outline, irregularly packed Lactuca sativa Anomocytic 188.41419.29 | 620436 2213 Highly wavy cell outline, irregularly packed Launaea taraxacifolia Anomocytic 188.80428.52 | 312+30 10.71 Straight cell outline, fairly large, irregularly packed Picris humilis — Anomocytic 187.87423.86 484430 28 Less wavy cell outline, irregularly packed. Sonchus oleraceus + Anomocytic 204.70+£19.90 528+25 29.09 Less wavy cell outline, irregularly packed 58 Comp. Newsl. 46, 2008 Fig. 1. Diagram of the abaxial leaf surface of the Cichorieae species. A. Lactuca capensis (Northern Nigeria collection) B. Lactuca capensis (Southern Nigeria collection). C. Lactuca sativ D. Launaea tar fol E. Picris humil F. Sonchus ol Comp. Newsl. 46, 2008 59 Studies on Vernonia monosis C. B. CLARKE (Asteraceae): Palynology, Taxonomy and Phytogeography GEORGE VASANTHY & Dario DE FRANCESCHI* Dept. of Ecology Institut Francais, P.O. Box 33 11. St Louis Street Pondicherry-605 001, India vasanthi.g@ifpIndia.org (author for correspondence) *MNHN, Département Histoire de la Terre UMR 5143 Paléobiodiversité et Paléoenvironment 8. rue Buffon 75005 Paris, France dariodf@mnhn.fr Abstract Merging the South Indian tree Vernonia monosis C.B. CLARKE with the Indo- malayan and Southeast Asiatic tree Vernonia arborea Bucu.-Ham. as done ina few Indian floras needs to be corrected mainly because of their pollen morphological dissimilarity. Nevertheless the pollen morphological similarity of the three South Indian tree Vernonia taxa and the Indo-Burmese V. volkameriifolia DC. underscores their potential inter-relationship. Our survey (about 80 specimens) of V. monosis has revealed an interesting phenomenon: a large majority of specimens from the Western Ghats have |-flowered capitula whereas heads with a variable number (few to one) of florets occur in V. monosis specimens from the Babaduban-Coorg- Hassan sector. The taxonomic and phytogeographical issues emerging from our findings are discussed. Introduction The trees of Indian Asteraceae belong to Vernonia Scures. The tree Vernonia taxa of South India are: V. monosis C. B. CLARKE, V. shevaroyensis GAMBLE and V. travancorica Hook. f. (GAMBLE 1921). In recent taxonomy they have been treated in the genus Gymnanthemum Cass. (ROBINSON 1999) and even more recently transferred to the resurrected genus Monosis DC. together with three other taxa 60 Comp. Newsl. 46, 2008 (RoBINsON & SKvARLA 2006). In the present paper the taxa discussed are referred to Vernonia in the traditional broad sense. Based on many macro-morphological characters Vernonia monosis has been merged with the Indo-malayan and Southeast Asiatic V. arborea Bucu.-Ham. (e.g. RAMAMOORTHY 1976, MATTHEW 1983, MANILAL 1988, KesHAVA Murtuy & YOGANARASIMHAN 1988). The pollen micro-morphology of the South Indian tree Vernonia taxa is identical but dissimilar to that of V. arborea. Our present work, a follow-up of VASANTHY (1985) and VASANTHY & De FRANCESCHI (1992), may help researchers to appreciate the diagnostic value of pollen morphology of the tree Vernonia species. We also report here the occurrence of capitula with few florets in some specimens of V. monosis (Fig. 4; Table 2). This deviation from the 1-flowered character state [described by CLARKE (1876) as “Capitula 1-flora’’| raises issues related to phylogeny and phytogeography of the tree Vernonia taxa of South India. For nomenclatural references of the South Indian taxa, the reader is referred to CLARKE (1876), GAMBLE (1921) and RAMAMOoorTHY (1976) and those of V. arborea to Koster (1935, 1966). The following list of taxa studied for the present work is furnished with notes on their distribution and ecology. 1) Vernonia monosis C. B. CLARKE (syn. Monosis wightiana DC. in BEDDOME 1972; Plate 226) (Figs 1:5; 2 & 3). Distribution: See Fig. 6 and the tentative key in this work. Ecology: Small, evergreen, heliophilous trees 10-30 m tall, growing in areas of subequatorial climate (c. 200 m to 2500 m) along forest fringes, ecotonal areas and in disturbed parts of forests as secondary vegetation. 2) Vernonia shevaroyensis GAMBLE (Fig. 1:A) Distribution: See Fig. 6 and the tentative key in this work. Ecology: On forest-fringes; highly endangered by human activities (Rao & Drxir 1996). 3) Vernonia travancorica Hook. f. (Fig. 1:C) Distribution: See Fig. 6 and the tentative key in this work. Ecology: In evergreen forests, along forest fringes. 4) Vernonia arborea Bucu.-HaoM. Type specimen in the herbarium of the Royal Botanic Garden, Edinburgh (KOsTER 1935). Comp. Newsl. 46, 2008 61 Distribution: India, Malay Peninsula, southern China, Tonkin, Vietnam, Laos, Philippines, Malay Archipelago (KostTER 1966) and Sri Lanka. Of the two varieties occurring in Sri Lanka, V. arborea var. javanica (BLUME) C. B. CLARKE is more widespread than var. arborea (GRIERSON 1972). Ecology: Small trees (up to 30 m). Vernonia arborea and its varieties are prevalent in the primary and secondary forests and the montane rain-forests (KOSTER 1935). Materials and Methods Specimens of the South Indian tree Vernonia taxa housed in the following herbaria were examined for this study. BSI-WC: Botanical Survey of India, Western Circle, Pune, India. CAL:Botanical Survey of India, Central National Herbarium, Sibpore, Howrah, India. HIFP: The herbarium of Institut Francais, Pondicherry-605 001, India. JCB: St. Joseph’s College, Bangalore, India. K: Royal Botanic Gardens, Kew, Richmond, Surrey, U.K. PCM: Presidency College, Dept. of Botany, Madras (Chennai-600 005), India. RHT: Rapinat Herbarium, St. Joseph’s College, Trichy, Tamilnadu, India. Acetolysed (see ErRpTMAN 1952) pollen grains were micro-photographed with Scanning Electron Microscopes. Results A. Common characters of the selected tree Vernonia taxa a) Habit: trees with differential trunk. b) Phyllotaxy: alternate. c) Leaf venation: pinnate. d) Heads: homogamous with tubular floret(s), actinomoprhic 5—lobed corollas. e) Inflorescence: + primarily compound corymbose-paniculate. f) Capitulum: oblong-campanulate or cup-shaped. g) Involucre: 4— to 5—seriate. h) /nvolucral bracts: persistent. i) Receptacle surface: naked or glabrous. j) Style base: expanded. k) Achenial ribs: 10 (8-12). 62 Comp. Newsl. 46, 2008 B. Pollen types i) K monosis type (Fig. 4) 3-colpororate [with colpus (ecto-aperture) + os A (meso-aperture) + os B (endo- aperture)],+ radio-symmetric, suboblate to oblate spheroidal; polar lacunae 3x2, confluent with the apertural lacunae; echinulo-lophate (crest-like elevations bearing spinules); lacunate [39 lacunae: 12 para-apertural, 6 inter-apertural, 3 ororal, 6 aboral, 6 equatorial and 6 polar (see VASANTHY et al. 1993: Fig. 5)]; spinules blunt or truncate; tectum bilaterally and partly (distally) covering columellae, tectal exterior imperforate; columellae simple to composite, well-fused with the foot- layer; endexine present. Remark: This type of pollen characterizes not only V. monosis but also the two other tree Vernonia taxa of South India: V. shevaroyensis and V. travancorica [Plates 5 & 13-18 in VASANTHY 1985; VasantHy & GiRARD 2007: Digital images (10) and SEM (1) of K travancorica] and the Himalayan-Indo-Burmese V. volkameriifolia (our unpubl. SEM result of specimen from Assam: N. Cachar Hills, T. YMDELL-74, K) ii) V. arborea type (Fig. 5) 3-colpororate [with colpus (ecto-aperture) + os A (meso-aperture) + os B (endo-aperture)| + radio-symmetric, suboblate to oblate spheroidal; polar area with variable sub-lacunar patterns; sub-lophate (with elevated crest-like walls; not forming network); spines prominent; tectal exterior micro-reticuloid (microperforate); columellae of sublophos simple to composite, well-fused with the foot-layer; endexine present. Remark: This type of pollen is present (VASANTHY 1985) in some of the South Indian Vernonia taxa: V. anaimudica BENtTH., V. bourdillonii GAMBLE, V. bourneana W. W. So.,V. comorinensis Scu.-Bip., V. malabarica Hook. f., V. salviifolia WiGHT (shrubs), and V. ramaswamii BENTH. (undershrub). C. Population(s) of V. monosis with few - to one-flowered capitula Varying number of florets per capitulum has been observed by us in the V. monosis specimens from the Bababudan-Coorg sector (Fig. 3 & Table 2). D. Phytogeography of specimens examined (Fig. 6) a) With capitulum more than 1-flowered in Vernonia “monosis”: 10 specimens collected from the Mysore Plateau (between Bababudan Hills and Coorg-Hassan Districts: 13°31’32”N to 12°12’13”N and 75°48’49”E to 75°6’E). b) The typical 1-flowered capitulum state in Vernonia monosis: about 65 specimens Comp. Newsl. 46, 2008 63 from the Western Ghats, B. R. Hills (4 specimens) and the Eastern Ghats: Kolli Hills (1 specimen). c) V. shevaroyensis (\-flowered capitulum): 2 specimens from Yercaud, Salem (ic45 50 7N2 78° 11-25" B). d) K travancorica: 16 specimens collected from the Travancore region (09°38’39”N to 08°22’23”N and 77°07’09”E to 77°29’30”E). Discussion Vernonia monosis vs V. arborea: Differences Vernonia in the traditional sense is a large genus comprising about 1000 species (WILLIS 1988), but in recent generic classification reduced to 22 species confined to the New World (Rosinson 2007). Vernonia s. lat. is characterized by many pollen types (e.g. Jones 1979, 1981; VAsANTHy 1985) and this genus is eurypalynous (as defined by ErpTMAN 1952). The pollen types present in the Vernonia of the Old World have been classified into six types (A—-F; Fig. | in Jones 1981). In his classification of subsection Strobocalyx (BLUME ex DC.) S. B. JoNEs (JONES 1981: pp.64—65), the pollen of the type species V. arborea is placed under type A [3-colpor(or)ate, echino(sub)lophate, sublacunate] and that of V. volkameriifolia (resembling the pollen type of V. monosis) under type B. The pollen with a complete tectum (A type sensu JONES 1981) is the least specialised (BoLIck & KEELEY 1994). Hence the pollen type in South Indian tree Vernonia taxa (hereafter V. monosis type; see Table 1) having incomplete tectum is evolutionarily specialised. A few other character states such as spinulate sculpture, lophote-lacunate surface and imperforate tectum of V. monosis pollen type are considered more specialised than the spiny sculpture, sublophate-sublacunate surface and micro-reticuloid tectum of V. arborea pollen type. Thus we underscore the diagnostic value of pollen morphology in differentiating the southern Indian tree Vernonia taxa particularly V. monosis from the widely distributed Indo-malayan and Southeast Asiatic V. arborea. 64 Comp. Newsl. 46, 2008 Table 1. Pollen morphological dissimilarities between V. arborea and Ve monosis SES V. arborea type (Fig. 5) V. monosis type (Fig. 4) 1. Polarity Sub-isopolar (sub-lacunate) Isopolar,’ with tieradiate polar ridge Within the well-developed 2. Aperture complex | Within the confluent sub- aperting| agin eaaaeiede (lacunar pattern) lacunae : inter-lacunar gaps, 3. Type of wall Sub-lophos Lophos ; : Echinulate [spin(ule h.: 4. Sculpture Echinate [spine h. >3.0 um] hone 5. Tectal Exterior Micro-reticuloid Imperforate fee oalacinne Sub-lacunae (surrounded by ents sub-lophos) 7 ; : Imperforate bilateral tectum Tectal micro-reticulum of | .. : : 7. Lacunar Floor and SI Guh santa cece distally restricted exposing Tectal Descension i P the bases of columellae and continuous the lacunae Note: The definitions of the following terms are after WoDEHOUSE (1935). Lophos: Crest-like ridges of pollen-wall. Lacuna(e): Large pits or depressed angular spaces in the exine partitioned by lophate walls. The South Indian Vernonia monosis has often been merged with V. arborea (e.g. RAMAmMoorTHY 1976) regardless of the latter’s extra-regional distribution (KOSTER 1966). Despite this nomenclatural change, these two taxa appear to be so distinct in respect of characters pertaining to the capitulum, pollen, corolla, anther, achene and pappus as to be taxonomically distinct on species or even higher level (cf. RoBINSON & SKVARLA 2006). Vernonia monosis (mostly 1-flowered) has echinulo- lophate grains (Table | & Fig. 4), corolla with biseriate vesiculate trichomes, anthers with triangular bases (not tailed), obovoid achenes and one row of pappus (our obs.). On the other hand, in V. arborea the capitulum is 3—6 flowered (never 1-flowered); pollen grains are echino-sublophate with microreticuloid exterior (Fig. 1: Ain Jones 1981; Fig. 5 in this work), the other differences being corolla with spreading trichomes, anthers with tailed bases, achenes sub-triangular and pappus sub-biseriate (KosTER 1935). In the light of these observations V. monosis and V. arborea are to be considered as two distinct species, at least. Though the former being a later homonym, it is to be conserved not only for its specific Comp. Newsl. 46, 2008 65 distinction but also for its long usage in standard taxonomic works (e.g. CLARKE 1876, GAMBLE 1921). Furthermore, the type specimen of V. arborea (in the herbarium of the Royal Botanic Garden, Edinburgh) has been studied by C. B. CLARKE (Koster 1935: pp. 384-385) but CLARKE (1876: pp. 23-24) has not merged V. monosis with V. arborea. Variations in number of florets (6> 1) per capitulum of Vernonia monosis Enormous variation in the number of florets per head (from numerous to one) characterizes the family Asteraceae. The specific epithet of V. monosis has originated from the generic name Monosis DC. in Wicur [capitula single- flowered as described by BEppome (1872)]. It characterizes the single-flowered capitulum state of this taxon (CLARKE 1876, GAMBLE 1921). RAMAMOoRTHY (1976) has described the heads of a specimen of V. “arborea” from the Hassan District, Karnataka (C. J. SALDANHA -16277) as few-flowered, and we have observed 3-flowered character state in the heads of the same specimen (see Table 2). The number of flowers in a head ranges fom one to many in the genus Vernonia s. lat. The one-flowered capitulum state is apomorphic (derived) whereas the many-flowered capitulum is plesiomorphic [least specialised (KEELEY & TURNER 1990, VasANtTHy & De FRANCHESCHI 1992). Koster (1935: p. 307) har discussed the evolutionary tendency showing reduction in the number of flowers per head in the tribe Vernonieae and its taxonomic significance. The few-to one-flowered character states of V. monosis of the Western Ghats (Table 2) seem to corroborate the above evolutionary hypothesis. 66 Comp. Newsl. 46, 2008 Table 2. Variability in the number of florets per capitulum (6g1) in Vv. monosis from the Bababudan-Coorg-Hassan Sector (Fig. 6) hei wedi Collection Data of Specimens examined per capitulum Bababudangiri Hills: Kemmangundi, S. ArAvasy, Feb. 1992, 674 HIFP. Ley, Bababudangiri Hills: Shankar Shola, B. R. RAMesu, April 1991, HIFP. 4>3 Coorg: Viranjendrapet, P. F. Fyson, s.n., Feb. 1916, PCM. 4 Coorg: Kadamakal Reserve Forest near Gallibedu, J.-P. PascaL -841, March 1977. HIFP. Hassan: Mudigere road, C. J. SALDANHA -16277, Feb. 1970, 3 JCB. 3 Bababudangiri Hills: Dattarayapet, N. RAMAA Rao, Feb. 1980, K. 2 Coorg, Hooker & THompson, s.n. & s.d., K. 2 Coorg: Mercara, G. S. Puri, March 1958, BSI-WC. 21 Coorg: Mercara, PARKER -2144, Jan. 1924, K. ns Chickmagalur: Mulainagiri road, C. J. SALDANHA & RAMESH, April 1980, JCB. | Coorg: Abbey falls road, C. J. SALDANHA et al., Dec. 1988, JCB. Suggestions for Nomenclatural Changes and Revision The fragile species delimitation of the South Indian tree Vernonia taxa has become evident on the analysis of pollen morphology, achenial character (10-ribbed), our observation of variation in the number of floret per capitulum in V. monosis and the vegetative characters described by GAMBLE (1921). The following arrangement (a tentative key) highlights the need for nomenclatural changes requiring future emendation of the description of these three species and validation of the following taxonomic changes i. Splitting of V. monosis into its varieties or subspecies. ii. Merging V. shevaroyensis with V. monosis as a variety or a subspecies. Comp. Newsl. 46, 2008 67 A Tentative Key Leaf margin wavy, leaf shape ovate, leaf surface glabrous on both sides. IC AGS 65. NOW ELEG cos aaicschontee ess ccsaonsant ane neectns Vernonia travancorica (Distrib. Travancore-Tineevelly Hills) Leaf margin entire or rarely serrate, leaf shape obovate, leaf surface pubescent on the nerves above, densely brown tomentose beneath. Heads 61 -flowered (Table 1).................... Vernonia “monosis” - Group A. (Distrib. Bababudan-Coorg-Hassan sector) egdS l= flowered: (F100) xccc-sceacssantacse-t vases Vernonia monosis - Group B. (Distrib. Biligiri Rangan Hills, W. Ghats, Nilgiris, Palnis, Anamalais Hills of Travancore, High Wavys; E. Ghats: Kolli Hills) Leaf margin entire, leaf shape oblanceolate, leaf surface glabrous above, scabrid pubescent beneath. CES Ag Sele NLOMIEREG) 25 i sc nen sabe seseeccsioncsevte coseeeuey a Vernonia shevaroyensis (Distrib. E. Ghats: Shevaroys) It is conjectural that the populations of Vernonia “monosis” (Group A) represent an intermediate stage in capitulum evolution (6~1 transition of floret number per capitulum) between the typically 1-flowered heads of V. monosis (Group B) and V. shevaroyensis complex and the 63 flowered heads of V. travancorica. We conclude this report with the question: “Should the Group A of Vernonia “monosis”’ be treated as a subspecies of V. monosis?” Acknowledgements We offer our thanks to Drs. J.-P. MULLER, F. BorNeE and P. CouTeron for support. We are grateful to (late) Dr. S. A. J. Pocock for the SEM pictures of V. monosis pollen. We thank Prof. B. Norpenstam for helpful suggestions, K. RAMESH Kumar for electronic plate preparation (Figs. 1, 2, 4 & 5) and photography (Fig. 3), R. Srvarasan for drafting Fig. 6 and G. MuruHu SHANKAR and N. RAVICHANDRAN for technical assistance. 68 Comp. Newsl. 46, 2008 References Beppome_, R. H. 1872. The Flora sylvatica for Southern India, Vol. 2: 225 & 226. Reprinted 1978 by J. K Printers, Delhi-110008, Periodical Export Book Agency, Vivek Vihar, Delhi-110032. Bouick, M. R. & S. C. KEELey 1994. Pollen morphology and classification of the Vernonieae (Compositae). Acta bot. Gallica 141 (2): 279-284. CuiarkE, C. B. 1876. Compositae indicae; Descriptae et secus genera Benthamii ordinatae. Published by M/S Bishen Singh Mahendra Pal Singh, 23-A, New Connaught Place, Dehra Dun. ErpDTMAN, G. 1952. Pollen morphology and plant taxonomy - Angiosperms. Almgvist & Wiksell, Stockholm. GamMBLE, J. S. 1921. Vernonia Scures. In: Flora of the Presidency of Madras. Vol. 2: 470-475. 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Rosinson, H. & J. SKVARLA 2006. Studies on the Gymnantheminae (Vernonieae: Asteraceae): restoration of the genus Monosis. Proc. Biol. Soc. Wash. 119 (4): 600-607. VaSsANtuy, G. 1985. Palynology and certain aspects of histology of South Indian Vernonieae (Asteraceae). Ph. D. Dissertation, Madras University, Madras, India. The US Library of Congress (Code Number I.E. 58158, and Card Number 88-910069). An electronic version (CD-ROM-IFP Publ.) in prep. VasANTHY, G. & D. DE FRrancescui 1992. A cladistic analysis of the Indo- malesian tree Vernonieae of the Asteraceae. Hennig XI, Paris, France, Aug. 25—28 (Poster presentation). Abstracts: p. 84. VANSANTHY, G., De FRANCESCHI, D. & S. A. J. Pocock 1993. Geometric aspects of pollen: Polyhedrons (Vernonieae-Asteraceae), Rotated symmetry (Trichanthereae-Acanthaceae) and Pyramidal to prismatic spinules (Nothapodytes-\cacinaceae). ErpTuAN Commemoration Volume, Grana 52 (Suppl. 2): 37-43. VasANTHy, G. & P. Grarp 2007. Pollen grains of South Indian trees. V. 1. 0. A user-friendly multimedia identification software [CD-ROM]. Collection Ecologie N° 45. IFP (with a user manual of 20 pages). For sale at the IFP Library (library@ifpindia.org). Wiis, J. C. 1988. A dictionary of the flowering plants and ferns. Cambridge University Press, 8" edition (revised by H. K. Airy SHAW). Wopbenouse, R. P. 1935. Pollen grains. Their structure, identification and significance in science and medicine. Reprinted in 1959 by Hafner Publishing Co., New York. 70 Comp. Newsl. 46, 2008 Fig. 1. A. Vernonia shevaroyensis (Shevaroy Hills: Yercaud, 1500 m, G. T. -1330, HIFP). B. K. monosis (Palnis: Top Station, 1900 m, Feb. 1972, F. Blasco -1150, HIFP). C. Vv. travancorica (Agasthiamalai, 1100 m, Feb. 1978, J.-P. PAscAL -688, HIFP). Note the phyllotaxy, leaf shapes and capitulum types (1-flowered heads in A & B and 6—3-flowered heads in C). Comp. Newsl. 46, 2008 7] Fig. 2. A flowering branch of Vernonia monosis characterized by the typical 1-flowered heads in terminal compound corymbose-panicle [from Nilgiris, Coonoor 1,500 m. (Tamilnadu); Photo and collection (DDF-472, HIFP) by D. De FRANcescui. Note the actinomorphic, tubular and 5-lobed, purple coloured florets. 72 Comp. Newsl. 46, 2008 Fig. 3. A flowering branch of Vernonia “monosis” with 6>4 flowered heads (ex. a & b) from Bababudangiri Hills: Kemmangundi, alt. 810 m, S. ARAvAsy, Feb. 1991, HIFP (see Table 2). The leaf characters (margin entire, shape obovate, surface pubescent on the nerves above, densely brown tomentose beneath) of this specimen are similar to those of 1-flowered V. monosis (cf. GAMBLE 1921). Comp. Newsl. 46, 2008 73 Fig. 4. Scanning Electron Micrographs (SEM) of pollen of Vernonia monosis (Nilgiris: Gudalore, Nadughani, 900 m., S. Aravasy, Feb. 1980, HIFP): A. Equatorial view: one of the three apertures in view. B. Polar view: the tri-radiate polar ridge delimiting the three polar lacunae. C. A close-up of figure B. D. Fractured exine showing the details of lophate exine. 74 Comp. Newsl. 46, 2008 Fig. 5. Scanning Electron Micrographs (SEM) of pollen of Vernonia arborea vat. javanica DC. (Sri Lanka, Madulkelle, TBW-1999, K): A. Equatorial view of echinate (spiny), sub-lophate and sub-lacunate grain with one of the three colpororate apertures in view (cf. Fig. 1: Ain Jones 1981). B. Polar view. Comp. Newsl. 46, 2008 75 . Vernonia species [ 1200 - 1800 m ‘ : V._ monosis - | fld heads be] Soe ems a 4. V monosis - 6 to | flds heads (00 ea arly 6S. V shevaroyensis - | fld heads V. travancoria -6 to3 fld heads Fig. 6. Distribution of the South Indian tree Vernonia taxa. The inter-montane hypothetical link indicates the spread of 1-flowered capitulum Vernonia monosis in the Western and Eastern Ghats and the “bridging” B. R. Hills. 76 Comp. Newsl. 46, 2008 A new species of Graphistylis (Asteraceae: Senecioneae) from Minas Gerais, Brazil ArisTonio M. Teves! & BertiL NORDENSTAM? ' Departamento de Botanica, Instituto de Ciéncias Biolégicas Universidade Federal de Minas Gerais, Av. Antonio Carlos 6627 — Pampulha — Caixa Postal 486 — 31270-901, Belo Horizonte MG, Brazil aristonio@hotmail.com ’Department of Phanerogamic Botany Swedish Museum of Natural History Box 50007, SE-104 05 Stockholm, Sweden Abstract A new species of Graphistylis (Senecioneae - Asteraceae) is described from State Park of Rio Preto, Minas Gerais, Brazil. Graphistylis riopretensis A.TELES & B.Norp. is illustrated, and compared with the related species Graphistylis dichroa (Bona.) D.J.N.Hinp. A key to the species of Graphistylis from Minas Gerais state is provided. Introduction The genus Graphistylis was described by NorDENSTAM (1978) to accomodate the species of Senecio sect. Dichroa CABRERA. All the species of the genus are exclusive of southern and southeastern Brazil, and are characterized by the suffrutescent habit, leaves petiolate to subpetiolate, capitulescence corymbose-paniculate, heads commonly radiate or rarely discoid (G. serrana (ZARDINI) B.Norb.), ray florets normally 5, and style branches apically with a conspicuous elongated hair pencil and lateral shorter hair tufts (NoRDENSTAM 1978, 2007). Another particularity of the genus can be observed in plants “in vivo”, viz. the wine colour present in stem, leaves and phyllaries. Frequently in the same plant the stem and abaxial leaf face vary in colour from green to wine. The genus was described initially with six species. Hinp (1993) combined Cacalia dichroa Bona. in Graphistylis as G. dichroa (BonG.) D.J.N.Hinp, and later NoRDENSTAM (1994) proposed the combination of Senecio serranus ZARDINI in Graphistylis (G. serrana) and without having knowledge of the publication Comp. Newsl. 46, 2008 7 of Hinp (1993) proposed again the combination of C. dichroa in Graphistylis. In the same year, HIND (1994) proposed the same combination as published by NorDENSTAM (1994) of the species described by ZARpINI (1992). However, this time the valid combination was the one proposed by NorpEnstAmM (1994), which was published earlier in the same year and has priority. The species of Graphistylis are characteristic of campos de altitude (high-altitude grasslands), a cool humid grass-dominated formation restricted to the highest summits (altitudes higher than 1,500 m above sea level) of the southeastern Brazilian highlands (SAFForD 1999, Garcia & PiRANI 2003), often associated with populations of Sphagnum and Chusquea, forming small clusters of shrubs (TELES 2008). Up to now the genus comprised eight species (G. argvrotricha (DUSEN) B.Norp., G. cuneifolia (GARDNER) B.Norb., G. dichroa (BonG.) D.J.N.HIND, G. itatiaiae (DusEN) B.Norp., G. oreophila (DUSEN) B.Norb., G. organensis (CASAR.) B.NorD., G. serrana (ZARDINI) B.Norp. and G. toledoi (CABRERA) B.Norp.), but in this paper we describe a new species, which so far has been collected only in Minas Gerais, Brazil, increasing the number of species in the genus to nine. Description and Discussion Graphistylis riopretensis A.TELES & B.Norb., sp. nov. G. dichroae valde affinis, sed foliis oblanceolatis margine serratis (in illa foliis lanceolatis ad obovatis margine dentatis) et flosculorum radii laminis majoribus (13 mm longis et 6 mm latis; in G. dichroa 4-9 mm longis et 1.5—3 mm latis). Type: Brazil. Minas Gerais: Sao Gongalo do Rio Preto, Parque Estadual do Rio Preto, Pico Dois Irmaos, 10.VIII.2004, P. L. VIANA & N. O. F. Mota 1825 (BHCB holotype). Fig. 1. Perennial suffrutex, erect, ca. 1.5 m high, rhizomatous. Stem simple, multisulcate, medullated, foliose throughout, glabrous. Leaves alternate, oblanceolate, 3—11 cm long, 0.7—3 cm wide, flat, concolorous, pinnately veined, glabrous adaxially, sparsely hirsute abaxially, petiolate, apex acute, base obtuse, margin regularly serrate with small teeth; petioles 0.1—0.7 cm long, sparsely hirsute. Capitulescence broadly paniculiform. Heads heterogamous, radiate, pedunculate; peduncles 2.54 cm long, bracteolate, hirsute; bracteoles lanceolate, 3-4 mm long, hirsute. Involucre campanulate, 9-11 mm long, 7-9 mm wide, calyculate; calycular bracts ca. 5, lanceolate, 3-4.5 mm long, ciliate; phyllaries 13, oblong, glabrous dorsally, with apex acute, wine-coloured and penicillate, margin scarious-hyaline; receptacle flat, alveolate. Ray florets 8, pistillate; corolla liguliform, yellow; tube 78 Comp. Newsl. 46, 2008 5.5 mm long; lamina 13 mm long, 6 mm wide, 6-nervate, apex 3-dentate; style 9 mm long, style branches 4 mm long. Cypselas terete, 2.5 mm long, glabrous; pappus bristles 8 mm long, persistent. Disk florets ca. 20, perfect, corolla tubular, 10 mm long, 5-lobate; lobes 1.5—2 mm long, anthers 2.5 mm long, base obtuse, connectival appendage oblong, 0.5 mm long; style 11.5 mm long, style branches 2.5 mm long, apex papillose, obtuse, with a conspicuous elongated hair pencil and lateral shorter hair tufts. Cypselas terete, 2.5 mm long, 10-ribbed, glabrous; pappus bristles 9 mm long, persistent. Geographical distribution and habitat: Graphistylis riopretensis is presently known only from a single collection in the municipality of Sao Gongalo do Rio Preto, in State Park of Rio Preto, Minas Gerais state (Fig. 2). Like the other species of the genus, G. riopretensis occurs in campos de altitude (high-altitude grasslands), in altitudes ranging from 1,800 to 1,830 m above sea level. Conservation status: Based on the presently available information, the new species can be scored, considering IUCN conservation criteria (IUCN 2001, TELEs & NAKAJIMA 2006), as critically endangered (CR), since its area of occurrence is smaller than 100 km”, its area of occupancy is smaller than 10 km’, and it is known from only one locality (criteria Blabi,11, B2abi,11). Fenology: Flowers and fruits were collected in August. Notes: Graphistylis riopretensis is closely related to G. dichroa, but is distinguishable inter alia by its leaves oblanceolate with margins serrate (vs. leaves lanceolate to obovate and with margins dentate), as well as the larger ray florets (13 mm long, 6 mm wide, as compared to 4-9 mm long and 1.5—3 mm wide in G. dichroa). For further differences, cf. the key below. Key to species of Graphistylis from Minas Gerais state 1. Leaves with the abaxial face densely white-toment0Se ...............:::secseseceseeeees Aen eral iith Len Cte ree. sees rge celebs . bank 429 Rt Graphistylis argyrotricha 1’. Leaves glabrous on both faces or only hirsute on the veins of the abaxial face. 2. Leaves more than four times longer than wide. 3. Leaves lanceolate to obovate, margin dentate; peduncles to 2.5 cm long; phyllaries 8; ray florets 5—7, tube 0.3—5 mm long, ligule 4-9 mm long, 1.5—3 mm wide; disk florets with corolla 4-9 mm long ...... Graphistylis dichroa 3’. Leaves oblanceolate, margin serrate; peduncles more than 2.5 cm long; phyllaries 13; ray florets 8, tube 5.5 mm long, ligule 13 mm long, Comp. Newsl. 46, 2008 79 Gamm wide: disk florets with:corolla 10 mir LONG, 0.......2.c