PROC. BIOL. SOC. WASH. 103(1), 1990, pp. 248-253

SIX NEW COMBINATIONS IN MOENCH AND "SUJME (: )

Harold Robinson

Abstract.— ThvQQ species, adoensis Schultz-Bip. ex Walp., V. gui- neensis Benth., and V. lasiopus O. HofFm. in Engl., are transferred to the Baccharoides Moench, and three species, Conyza cinerea L., C. patula Ait., and Herderia stellulifera Benth. are transferred to the genus Cyanthillium Blume.

The present paper provides six new com- tinct from the Western Hemisphere mem- binations of Old World Vemonieae that are bers of that genus. Although generic limits known to belong to the genera Baccharoides were not discussed by Jones, his study placed Moench and Cyanthillium Blume. The ap- the Old World Vernonia in a group on the plicability of these generic names to these opposite side the basic division in the genus species groups was first noted by the author from typical Vernonia in the eastern United almost ten years ago (Robinson et al. 1 980), States. Subsequent studies by Jones (1979b, and it was anticipated that other workers 1981) showed that certain pollen types also more familiar with the paleotropical mem- were restricted to Old World members of bers of the Vernonieae would provide the Vernonia s.l., types that are shared by some necessary combinations. A recent study of Old World members of the tribe tradition- eastern African members of the tribe by Jef- ally placed in other genera. The characters frey (1988) also cites these generic names as noted by Jones have been treated by the synonyms under his Vernonia Group 2 present author as evidence of a basic divi- subgroup C and Vernonia Group 4, al- sion in the Vernonieae between groups that though he retains the broad concept of Ver- have included many genera in each hemi- nonia. At this point it is not certain when sphere, and traditional Vernonia s.l. can be anyone will undertake more detailed studies seen as an excessively paraphyletic core ge- of these groups. However, the author does nus that is defined only by what "it is not" need to refer to a few members of these rather than by what "it is." In progressing genera and does not intend to use older, toward definitive generic concepts in the incorrect names. Instead of making isolated Vernonieae, the pattern seen by Jones jus- combinations as needed, as was done with tifies removing at least all of the Eastern Gymnanthemum coloratum (Robinson & Hemisphere members ofthe tribe from Ver- Kahn 1986), an effort is made here to place nonia. This segregation of the Old World needed combinations together in one paper. genera was partially put into practice by the Detailed monographic studies and the ma- use of the names Baccharoides Moench, jority of the combinations are left to future Gymnanthemum Cass., and Cyanthillium workers. In spite of the limited scope of the Blume, by Robinson et al. (1 980) and in the present effort, some explanation is offered. transfers made in the study of

The summary of the Vemonieae by S. B. Cass. (Robinson «fe Kahn 1986). Jones (1977) showed that the Eastern Hemi- At present there is good reason not to sphere species placed in the genus Vernonia transfer all the Old World species of Ver- were all chemically and cytologically dis- nonia out of the genus. Morphological and VOLUME 103, NUMBER 1 249

anatomical studies show the problem is Vernonia sect. Stengelia (Schultz-Bip. ex

more complex than one Old World genus Walp.) Benth. in Benth. & Hook, f., Gen. versus one New World genus. Several gen- PI. 2: 127. 1873. era should be recognized in both the neo- This group of species has been imper- tropical and paleotropical elements of Ver- fectly recognized in the past, primarily on nonia s.l. More detailed studies of the type the basis ofexpanded foliose appendages on under way in neotropical species (Robinson the involucral bracts. The character is well 1987a, b, c, 1988a, b, c, 1989; Robinson & developed in the genus but is not a reliable Funk 1987) are needed before all transfers basis for distinction. Smith (1971) studied can be made properly. Nevertheless, the po- a group titled the "stengelioid species" of sition of some of the species is fully re- Vernonia, but many species included in the solved, especially of species belonging to study were not Baccharoides. Smith recog- older segregate genera such as Baccharoides nized the heterogeneity in the group he and Cyanthillium. treated, especially in the form of the pollen The lack of total resolution of the tribe is and in the shape of the corolla, but these not a reason to retain under Vernonia s.l. were not detailed for all of the species. names and concepts known to be obsolete. More recently, Kingham (1976), Jones The author believes that Vernonia will ul- (1981), Jeffrey (1988), and the present au- timately be restricted to Western Hemi- thor have studied Baccharoides pollen in sphere species with non-lophate type A pol- more detail. It is lophate and distinctive len and a chromosome number of N = 17 among Old World tricolporate Vernonieae and will perhaps be restricted to only the by the presence of polar areoles. The grains eastern North American element of that fall into Jones' (1981) Type C, although pol- group. lens of all species studied in detail are dif- The detailed variations of pollen in the ferent from most Type C pollen of the Neo- tribe are to be more fully compared and tropics by the 3 equatorial areoles of the illustrated elsewhere. The generic synony- intercolpi and the strong basal columellae mies follow mostly those of Jeffrey (1988), under the crests of the exine. but some additional synonymy is provided Jeffrey (1988) emphasized the pollen, the under Cyanthillium. flattened inner pappus bristles, and the co- rollas with long, slender, basal tubes and Baccharoides Moench abruptly expanded cylindrical limbs and Baccharoides Moench, Methodus. 528. comparatively short erect lobes as definitive

1 794. Type: Conyza anthelmintica L. {=B. characters of Baccharoides within Vernonia

anthelmintica (L.) Moench). s.l.

Ascaricida Cass., Diet. Sci. Nat. 3, suppl. Ignoring some older counts, the genus 38. 1817, nom. superfl. Type: Conyza an- seems to have a consistent chromosome thelmintica L. number of N = 10 (Smith 1971, Jones Candidea Tenore, Atti Reale Accad. Sci. Sez. 1979a). The group is also notable for the

Soc. Reale Borbon 4 (CI. Botan.): 104, t. presence of epoxy resins and has been stud-

1, 2. 1839. Type: Candidea senegalensis ied for possible commerical exploitation. Tenore. Neither the chromosome number nor the Vernonia subsect. Stengelia Schultz-Bip. ex chemistry is unique to the genus. Walp., Repert. Bot. Syst. 2: 946. 1843. Many of the species are understood well Vernonia adoensis Schultz-Bip. ex Walp. enough to justify new combinations, but Stengelia Steetz in Peters, Naturw. Reise most of these, and others not known to the Mossambique, Bot. 360. 1864. Type: present author, are left for final resolution Vernonia schimperi DC. by other workers. The species listed by Smith 250 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(1971) are an artificial group and the Smith Cyanthillium Blume should not be used without reference paper Cyanthillium Blume, Bijdr. Fl. Ned. Ind. discussion by Jeffrey (1988). Except to the 889. 1826. Lectotype (designated here): of the Smith paper, the for its discussion Cyanthillium villosum Blume. study itself is limited to East African Jeffrey Isonema Cass., Bull. Soc. Philom. Paris species. It seems that a monograph of the 1817:152. 1817, nom. illeg., non R. Br. genus would be comparatively simple for (1810). Type: Isonema ovata Cass. someone with better resources to study pa- Cyanopis Blume ex DC, Prodr. 5:69. 1836, leotropical , for most of the species nom. illeg. et superfl., non Cass. (1817). occur in . One species was described Lectotype (Jones 1980): Cyanthillium vil- Arabia, and the widely distributed type from losum Blume. species was described from Sri Lanka. Vernonia sect. Tephrodes DC, Prodr. 5:24. The three species combinations needed 1836. Lectotype (Jones 1981): Conyza ci- at present are as follows: nerea L. Claotrachelus Zoll. & Mortiz ex Zoll., Na- BACCHAROIDES ADOENSIS tuur-Geneesk. Arch. Ned Indie 2:263, ex Walp.) (Schultz-Bip. 565. 1845. Type: Claotrachelus rupestris H. Robinson, comb. nov. Zoll. & Moritz ex Zoll. Vernonia adoensis Schultz-Bip. ex Walp., Seneciodes L. ex Post & O. Kuntze, Lex. Repert. Bot. Syst. 2:946. 1843. For ex- Gen. Phan. 2:515. 1903. Type: Conyza tensive synonymy see Smith (1971) and cinerea L. Jeffrey (1988). Distribution is subsaharan Triplotaxis Hutch., Bull. Misc. Inform. Africa with concentration in the east 1 9 14:355. 1914. Lectotype: Herderia stel- (Smith 1971, map p. 19). lulifera Benth. in Hook. Vernonia subsect. Orbivestus S. B. Jones, BACCHAROIDES GUINEENSIS Rhodora 83:61. 1981. Lectotype (Jeffrey 1988): Vernonia karaguensis Oliv. & (Benth. in Hook, f & Benth.) Hiem. H. Robinson, comb. nov. Vernonia subsect. Hilliardianae S. B. Jones, Vernonia guineensis Benth. in Hook, f & Rhodora 83:66. 1981. Lectotype (Jeffrey Benth., Niger Fl. 427. 1849. For extensive 1988): Webbia oligocephala DC. synonymy see Smith (197 1). Distribution Vernonia subsect. Tephrodes (DC.) S. B. is subsaharan Africa from Sierra Leone Jones, Rhodora 83:70. 1981. in the west eastward to western Tanzania The designation of a new lectotype for and south to southwestern Angola and Cyanthillium rejects the earlier choice of C. northern Zimbabwe (Smith 1971, map p. moluccense that seems to date from Jones 48). (1980). Jones' selection was totally arbi- BACCHAROIDES LASIOPUS trary, being the first species listed by Blume without any evident study of the group in- (O. Hoffm. in Engl.) volved, and thus can be rejected under the H. Robinson, comb. nov. Code. The lectotype cited by Jeffrey (1988) Vernonia lasiopus O. Hoffm. in Engl., Pflan- evidently follows Jones. Jones' lectotypifi- zew. Ost. Afrika 403. 1895. For extensive cation is unsuitable because the species is synonymy see Smith (1971). The species virtually unknown, being available to most ranges from western Ethiopia to north- botanists only as a microfiche of the frag- eastern Angola and northeastern Zim- ment in the DeCandolle herbarium at Ge- babwe (Smith, 1971, map p. 38). neva. The fragment C moluccense as seen VOLUME 103, NUMBER 1 251 in the microfiche may be a Cyanthillium. ring don't have the highly distinctive struc- The choice in this paper brings the typifi- ture seen in Triplotaxis but have only a cation into conformity with those of the moderately expanded upper callus on the synonyms, the superfluous generic name achene of a form seen in many Asteraceae. Cyanopis Blume and the homonym Isone- The latter species are not closely related to ma Cass., which both equal Cyanthillium Cyanthillium. Other members of Cyanthil- patula. lium, such as C. cinereum, have achene and It is unfortunate that many Flora formats pappus structures like those of typical Ver- now require types to be selected for all ge- nonia, without unusual rings or fragility. Be- neric names, even those treated as syn- cause of the differences in pappus structure onyms. Although the intention is noble, the listed above, the species now placed in result is too often poorly chosen lectotypes, Cyanthillium were not suspected of forming made by unqualified investigators. a related group separate from Vernonia until

Any study of Cyanthillium should in- the pollen study by Kingham (1 976) and the clude a study of members of recent study by Jeffrey (1988). Nevertheless, Schultz-Bip. (1853). The type, E. plumosa the species seem to share a generally her- Schultz-Bip., of Gabon, is known to the au- baceous habit, herbaceous and usually nar- thor only from its description. Erlangea is rowly petiolate , and broadly to nar- technically distinguished from Vernonia s.l. rowly ovate, slightly acuminate involucral and that part recognized here as Cyanthil- bracts, often with greenish outer surfaces lium by the five, deciduous pappus seg- and purplish margins. Jeffrey (1988) men- ments that have been referred to as plu- tions various forms of T-shaped hairs in mose. The type of Erlangea is described members of the group. with sessile amplexicaule leaves, a feature The pollen type of Cyanthillium is one of that would be unusual in the usually nar- the many paleotropical types in the Ver- rowly petiolate Cyanthillium. As described, nonieae in which the colpus is not evident. the type of Erlangea has a habit similar to Studies by the author show that the polar

Vicoa {Inula) auriculata Cass, and it may areas of the grains have a polar areole sur- be close to Erlangea schinzii O. Hoffm. of rounded by a tier of five to seven areoles northernmost Namibia, which also has ses- that variously match or straddle or fall be- sile leaves. Examination of material match- tween the pores below them. The polar or- ing the description of the type, and further ganization is basically different from forms evaluation of the pappus distinction, may in the tribe that have well developed colpi. show that the genus falls fully into the syn- In the present group there is not even a onymy of Cyanthillium. consistent single areole above or below the The genus Cyanthillium owes its early pores. The major organizational feature of recognition to the presence of a deciduous the grain is centered on the poles rather than pappus in the species group that includes the pores. SEM studies show that members the lectotype species. The broad achene and of Cyanthillium are distinctive within the short deciduous pappus of C. patula places non-colporate species of the Old World by this species outside of typical Vernonia. the modification of the basal columellae of Other pappus variations occur in Cyanthil- the exine to form bridges under the crests. lium and have been the basis of further ge- Points of attachment of the exine to the neric distinctions. The type species of Trip- footlayer seem to be restricted to positions lotaxis has a very distinctive, collar-like under the intersections of the crests. The outer pappus that persists after the inner pap- precise pollen form is not presently known pus has completely fallen. Other species from outside the genus, although similar forms Africa that are described as having a pappal without modified basal columellae are 252 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON known in species belonging to the group cit- Cyanthillium pubescens Blume, Bijdr. Fl. ed by Jeffrey (1988), for which the name Ned. Ind. 890. 1826. Crystallopollen Steetz in Peters seems to chinense [Lam.] Less., Lin- have priority. naea 4:320. 1829. The few available chromosome numbers Vernonia chinensis (Less.) Less., Linnaea 6: in Cyanthillium fall within the range noted 105, 674. 1831. for paleotropical members of the tribe. Jones Cyanopis madagascariensis DC, Prodr. 5: (1979a) reported N = 20 for C. stelluliferum 69. 1836. (as Triplotaxis). This seems to be based on Vernonia pratensis Klatt, Ann. K. K. Na- X = \Q. Jones reported N = 9, 18, and ca. turhist. Hofmus. 7:99. 1892. 18 for the widespread adventive C. cine- Cacalia patula (Less.) O. Kuntze, Rev. Gen. reum. Jones also reported N = 10 for two PI. 324. 1894. species of the closely related genus Erlan- Cyanthillium chinense (Lam.) Gleason, Bull. gea, including E. remifolia Wild & Pope, Torrey Bot. Club 40:306. 1913. which I suspect is a member of Cyanthil- The species is cited by Gleason (1922) as lium. occurring in tropical Asia, the East Indies, Cyanthillium has problems not seen in and being introduced into Guadeloupe. The Bacchawides, and a monograph will be more species is now also credited to Madagascar difficult. The distribution is throughout the on the basis of the newly synonymized Cy- paleotropical region, and there are many anopsis madagascariensis and Vernonia variations within the group in obvious char- pratensis. acters such as the achene and pappus. The three combinations needed at present are as follows: CYANTHILLIUM STELLULIFERUM (Benth.)

CYANTHILLIUM CINEREUM (L.) H. Robinson, comb. nov. H. Robinson, comb. nov. Herderia stellulifera Benth. in Hook, f &

Conyza cinerea L., Sp. PI. 862. 1753. Benth., Niger Fl. 425. 1849. Vernonia cinerea (L.) Less., Linnaea 4:291. Triplotaxis stellulifera (Benth.) Hutch., Bull. 1829. Misc. Inform. 1914:356. 1914. Senecioniodes cinerea (L.) Post & O. Kuntze, The distribution given by Jeffi"ey (1988) Lex. Gen. Phan. 2:515. 1903. is Uganda, west and central tropical Africa

The species is the most widely distributed and Angola. member of the genus. It occurs throughout the paleotropical region and is widely ad- Literature Cited ventive in the Neotropics. Gleason, H. A. 1922. Tribe 1. Vemonieae.— North American Flora 33:47-1 10. CYANTHILLIUM PATULUM (Ait.) Jeffrey, C. 1988. The Vemonieae in East Tropical H. Robinson, comb. nov. Africa.-Kew Bulletin 43:195-277. Jones, S. B. 1977. Vemonieae— Systematic review. Conyza patula Ait., Hortus Kew. 3:184. Pp. 503-521 in V. H. Heywood et al., eds., The 1789. biology and chemistry of the Compositae. Ac-

Isonema ovata Cass., Bull. Soc. Philom. ademic Press, London, New York, 1 189 pp.

Paris 1817: 152. 1817. . 1979a. Chromosome numbers of Vemonieae (Compositae). — Bulletin ofthe Torrey Botanical Conyza chinensis Lam., Encycl. 2:83. 1786, Club 106:79-84. hom. illeg., non L. . 1979b. Synopsis and pollen morphology of Cyanthillium villosum Blume, Bijdr. R. Vernonia (Compositae: Vemonieae) in the New Ned. Ind. 889. 1826. World. -Rhodora 81:425-447. VOLUME 103, NUMBER 1 253

. 1980. Tribe Vemonieae. In J. F. Macbride, . 1988c. Studies in the complex

ed.. Flora of Peru. — Fieldiana, Botany n. s. 5: (Vemonieae: Asteraceae). VI. A new genus, Ay- 22-69. «/a. — Proceedings of the Biological Society of

. 1981. Synoptic classification and pollen mor- Washington 101:959-965.

phology of Vernonia (Compositae: Vemonieae) , & V. A. Funk. 1987. A phylogenetic analysis in the Old World. -Rhodora 83:59-75. of Leiboldia, Lepidonia, and a new genus Stra- Kingham, D. L. 1976. A study of the pollen mor- mentopappus (Vemonieae: Asteraceae). — Bo- phology of tropical African and certain other tanische Jahrbiicher fiir Systematik, Pflanzen-

Vemonieae (Compositae). — Kew Bulletin 31:9- geschichte und Pflanzengeographie 108:2 1 3-228.

26. , & B. Kahn. 1986. Trinervate leaves, yellow

Robinson, H. 1 987a. Studies of the Lepidaploa com- , tailed anthers, and pollen variation in

plex (Vemonieae: Asteraceae). I. The genus Distephanus Cassini (Vemonieae: Astera- Stenocephalum Sch.Bip. — Proceedings of the ceae).— Proceedings of the Biological Society of Biological Society of Washington 100:578-583. Washington 99:493-501.

. 1987b. Studies in the Lepidaploa complex , F. Bohlmann, & R. M. King. 1980. Che- (Vemonieae: Asteraceae). II. A new genus, mosystematic notes on the Asteraceae. III. Nat- Echinocoryne. — VTOceedingi, of the Biological ural subdivisions of the Vemonieae. — Phyto- Society of Washington 100:563-589. logia 46:421-436.

. 1987c. Studies in the Lepidaploa complex Schultz-Bip., C. H. 1 853. Triga novorum Cassiniace-

(Vemonieae: Asteraceae). III. Two new genera, arum generum. — Flora 36:33-38. Cyrtocymura and Eirmocephala. — Froceedings Smith, C.E. 1971. Observations on stengelioid species of the Biological Society of Washington 100: of Vernonia. Agriculture Handbook no. 396. 844-855. U.S.D.A., Agricultural Research Service.

. 1988a. Studies in the Lepidaploa complex (Vemonieae: Asteraceae). IV. A new genus, Les- Department of Botany, National Mu- singianthus. — Froceedings of the Biological So- seum of Natural History, Smithsonian In- ciety of Washington 101:929-951. stitution, Washington, D.C. 20560. . 1988b. Studies in the Lepidaploa complex (Vemonieae: Asteraceae). V. A new genus, Chrysolaena. — Proceedings of the Biological Society of Washington 101:952-958.