Lipochaeta and (: Subtribe Ecliptinae): Establishing Their Natural Limits and a Synopsis Author(s): Warren L. Wagner and Harold Robinson Source: Brittonia, Vol. 53, No. 4, (Oct. - Dec., 2001), pp. 539-561 Published by: Springer on behalf of the New York Botanical Garden Press Stable URL: http://www.jstor.org/stable/3218386 Accessed: 19/05/2008 14:43

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http://www.jstor.org and Melanthera (Asteraceae: Heliantheae subtribe Ecliptinae): establishing their natural limits and a synopsis

WARREN L. WAGNER AND HAROLD ROBINSON

Wagner, W. L. & H. Robinson (Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0166, U.S.A.). Lipochaeta and Melanthera (Asteraceae: Heliantheae subtribe Ecliptinae): estab- lishing their natural limits and a synopsis. Brittonia 53: 539-561. 2001.-We restrict the Lipochaeta to the allopolyploid species of the typical section. Lipochaeta s.str. is interpreted to be the result of an intergeneric hybridization between Melanthera and a presently unknown taxon, perhaps of the genus Wed- elia. Lipochaeta is characterized, in addition to its allopolyploidy (n = 26), by having both flavonols and flavones, disk corollas with 4 lobes, achenes tuberculate at maturity, the disk achenes flattened to slightly biconvex, and ray achenes ob- compressed. Lipochaeta sect. Aphanopappus and Wollastonia are here reduced to synonymy under Melanthera. We transfer 14 Hawaiian Lipochaeta and one New Caledonian species as well as the Asian prostrata to Melanthera. These transfers, along with the species in and , bring the number of species in the genus to 35. Melanthera is delimited by an abruptly narrowed to truncate and flattened top of the achene, (0-)1-15(-20) often unequal, ciliate or barbellate, caducous pappus bristles immediately surrounding the corolla, in- volucral bracts and receptacular paleae with many veins forming longitudinal striations, and n = 15. The florets are 5-merous, the corollas are yellow or white, and rays are absent (in white-flowered species) or present and neutral or fertile. In dealing with species formerly placed in Lipochaeta, the Galapagos L. laricifolia is here transferred from the illegitimate generic name Macraea to Trigonopterum and the Brazilian L. goyazensis is transferred to Angelphytum. We maintain the earlier reduction of Echinocephalum under Melanthera and reduce all three taxa originally described in it to one, M. latifolia. Key words: Asteraceae, Compositae, Heliantheae, Ecliptinae, Echinocephalum, Lipochaeta, Macraea, Melanthera, Trigonopterum, Wedelia, Wollastonia, Ha- waiian Islands, Pacific islands, New World, Africa, .

Taxonomic resolution is necessary for taxa have been placed by Robinson (1981) the group of species that have been placed in the subtribe Ecliptinae of the tribe He- under the various genera Melanthera Rohr, liantheae and have been shown to belong to Lipotriche R. Br., Lipochaeta DC., Ma- the core of the Ecliptinae according to the craea Hook. f., Trigonopterum Steetz ex chloroplast DNA studies of Panero et al. Anderss., and Wollastonia DC. from the Pa- (1999). All the genera share achenes with cific and Palaeotropics. A natural taxonom- phytomelanin in the walls not interrupted ic disposition of these genera is hindered by by striations, style bases not enlarged, style additional problems with some species of branches with stigmatic surfaces as a pair Wedelia Jacq. or taxa related to Wedelia of separated lines, and corolla throats with and with South American species of the ge- a single scarcely colored resin duct. Relat- nus Echinocephalum Gardn. All of these edness of the genera is not in question, but

Brittonia, 53(4), 2001, pp. 539-561. ISSUED: 28 Feb 2002 ? 2001, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A. 540 BRITTONIA [VOL. 53

the precise delimitation of the genera in- tonia, Melanthera, and Lipochaeta. Within volved has not been properly realized. Spe- this clade Wollastonia and Melanthera form cifically, the problems are the narrow inter- a clade that is sister to Lipochaeta. The re- pretation of Melanthera and the overly sults of these two studies show the close broad interpretation of Lipochaeta. There is relationship among these taxa. also the need to formally synonymize the genus Wollastonia with Melanthera and to Restriction of the Concept of Lipochaeta replace the illegitimate name Macraea Hook. f. with Trigonopterum Steetz ex N. The genus Lipochaeta was described by J. Andersson. de Candolle (1836) to include 10 yellow- The present study was initiated to review flowered, opposite-leaved species with fer- the available literature, evaluate morpho- tile ray florets, triangular, triaristate, some- logical characters in putatively related taxa, what winged ray achenes, and biaristate, and provide a natural classification for the somewhat winged disk achenes. The name Hawaiian taxa assigned to Lipochaeta in Lipochaeta was ostensibly inspired by the the most recent treatments (Gardner, 1979; name Lipotriche R. Br. (1817), a name Wagner et al., 1990). To accomplish this we based on a specimen from Africa, but with- have evaluated most of the taxa worldwide out describing any species. Lessing (1831) that are currently treated in the genera in- described two species in Brown's genus, L. cluding Lipochaeta, Macraea, Melanthera, gymnolomoides and L. australis. Both of and Wollastonia, and many of those of these species were transferred from Lipo- Wedelia. Revisionary work (Becker, 1979; triche by de Candolle (1836), the former to Fay, 1979; Strother, 1991; Wild, 1965), gymnolomoides (Less.) DC., coupled with biosystematic studies by and the latter to his new genus Lipochaeta, Gardner and Rabakonandrianina (cited be- but using the oldest name avilable for the low), has brought considerable order to this species, L. lobata (Gaud.) DC. The original complex subtribe and has provided a solid element from Brown's genus was the only foundation for the work presented here. one kept in Lipotriche by de Candolle, and Many recent papers have dealt with as- he used the African specimen cited by pects of the Lipochaeta/Melanthera prob- Brown for the type of the new species Li- lem. Those dealing with Lipochaeta include potriche brownei DC., [= Melanthera scan- Gardner (1976, 1977a, 1977b, 1979), Gard- dens (Schumach. & Thonn.) Roberty]. ner and La Duke (1978), Rabakonandriani- Included in Lipochaeta by de Candolle na (1980), Rabakonandrianina and Carr were a series of American species from (1981), and Carr (1978). Those dealing and the Caribbean, now mostly in with Melanthera include Parks (1973), DC., with one in Wamalchi- Wild (1965), and Cabrera (1970, 1974). Pa- tamia Strother, five Hawaiian species, and pers dealing with Wollastonia include Wild finally there was mention of a single species (1965) and Fosberg and Sachet (1980). The now placed in La Llave. Four of whole series of genera was treated by the five Hawaiian names given by de Can- Stuessy (1977), Robinson (1981), and dolle, including the lectotype of Lipochae- Strother (1991) as members of the Helian- ta, L. lobata (Gaud.) DC., are members of theae subtribe Ecliptinae. Finally, Panero et the distinctive Lipochaeta section Lipo- al. (1999) did a chloroplast DNA study of chaeta having a chromosome number of n the Ecliptinae showing that Melanthera, Li- =26 and the majority of the disk corollas pochaeta, and Wollastonia are more closely with four lobes (Gardner, 1977a). Subse- related to one another than any are to the quent to the original description, during the clades containing Wedelia, Macraea, and remainder of the 19th century, a number of the Wedelia segregate Hi- additional species were ascribed to Lipo- eron. A second more recent study focusing chaeta. Among these was the Galapagos on the Ecliptinae using ITS (Chumley et al., endemic Lipochaeta laricifolia (Hook. f.) 2000) shows a much more robust clade A. Gray. This taxon subsequently has been consisting of , Perymenium, Wollas- resurrected as the genus Macraea (Harling, 2001] WAGNER & ROBINSON: ASTERACEAEASTERACEAB 541

1962; Eliasson, 1984), a comparatively dis- The Broadening of the Concept of tantly related member of the Ecliptinae (Pa- Melanthera nero et al., 1999). By 1935, when Sherff The name of the Melanthera em- revised the genus Lipochaeta, it was delim- genus the black anthers. Such anthers are ited to include only Hawaiian species and phasizes one from New Caledonia. The other not unusual in the Heliantheae, but they are species obvious in the element that had been to the were particularly typical assigned genus of Melanthera where the corollas are white New World and had been trans- primarily and there are no Such white-flowered ferred to a number of other The rays. genera. ge- nonradiate heads have become the accepted nus was again revised by Gardner (1979) to character for the traditional of the 22 and concept comprise shrubby perennial species genus. Other diagnostic features, less often a annual all hav- single species, Hawaiian, emphasized, are the abruptly narrowed to two different basic chromosome num- ing truncate and flattened top of the achene a di- bers, collectively occupying highly with a cluster of many true pappus bristles verse array of habitats ranging from low immediately surrounding the corolla, and coastal zone to steep dry or mesic slopes. involucral bracts and receptacular paleae Gardner also confirmed Harling's (1962) with many veins forming longitudinal stri- exclusion of Macraea of the Galapagos and ations. Such striations are found in species accepted the placement of Lipochaeta li- that have been placed in Wulffia Neck. ex fuana Hochr. from New Caledonia in Wed- Cass., Wollastonia, Echinocephalum, and elia [as W. uniflora (Forst.) Moore, a mis- Lipochaeta, and the character seems to de- applied name now considered to be a syn- limit a related group. Achenes of Melanth- onym of the known as Wollastonia era are not particularly noted for having biflora]. Wagner et al. (1990) reduced the wings, but a narrow, distinct wing is present Hawaiian species to 20. even in the generic type. Among the related Data relevant to developing a monophy- genera to Melanthera only the fleshy-- letic classification for the Hawaiian taxa ed Wulffia seems to lack wings. have been available since the biosystematic Bentham (1873) was the first to broaden and revisionary studies by Gardner (1976, the generic concept of Melanthera. The val- 1977a, 1977b, 1979; Gardner & La Duke, idly named genera he placed in synonymy Psathu- 1978), and experimental hybridization and were Echinocephalum, Lipotriche, rochaeta and Wuerschmittia Sch.- cytogenetic work by Carr (1978), Rabakon- DC., no combinations were made. andrianina (1980), and Rabakonandrianina Bip., although The Wollastonia was in Wed- and Carr (1981). Nevertheless Gardner genus placed did not delineate a elia. The concept included species with yel- (1979) monophyletic low corollas and neutral or fertile & rays. group (Carr, 1978; Fosberg Sachet, 1980; Wild also had an et Smith & (1965) expanded concept Wagner al., 1990; Carr, 1991). of Melanthera in his treatment of the Af- Rabakonandrianina and Carr (1981) pre- rican It was broader sented species. technically cytogenetic evidence that Lipochae- than the of Bentham the inclu- ta sect. arose concept by Lipochaeta by hybridization sion of the generic type of Wollastonia DC., between taxa with Wedelia-like n = 11 an- a species with yellow corollas and fertile and Melanthera-like n = 15 ances- cestry rays. With this concept of Melanthera, Wild This is the same element of Li- try. typical abandoned the use of such generic charac- pochaeta that is distinguished by the 4- ters as rays absent or present or neutral, and lobed corollas of the disk florets. To date, number of pappus bristles. Instead, he re- morphological evidence of this same Wed- lied on achene shape and the strongly striate elia-like ancestor in Melanthera and Wol- paleae. Unfortunately, the concepts of Wild lastonia is lacking. Based on these data, we have not been followed outside of Africa maintain the generic status of Lipochaeta by many authors during the rest of the cen- but restrict it to include only the typical el- tury. ement. Parks (1973) revised the North American 542 BRITTONIA [VOL. 53 and Caribbean species of Melanthera that lieve that at least W. spilanthoides and W. include only those taxa with white corollas, verbesinoides with striated paleae represent black anthers, and no rays. In doing so he products of a hybridization event unrelated dealt with only the narrow concept of the to the one that gave rise to Lipochaeta be- genus. At nearly the same time, Cabrera tween Melanthera-like and Wedelia-like (1974) formally reduced the yellow-flow- taxa. The Australian species differ from Ha- ered and neutral-rayed Echinocephalum of waiian Lipochaeta by their 5-lobed disk co- to synonymy under Me- rollas and the slight necks on the achene. lanthera, with which it shares similar achenes and a pappus of many bristles. Considerations Regarding Wollastonia It remains possible to restrict the concept of Melanthera by the white corollas and In a study dealing with Micronesian lack of rays, by the lack of fertile rays, or and thus geographically well-re- by the presence of many pappus bristles. moved from the Melanthera problem of the However, these delimitations are practical Americas, Fosberg and Sachet (1980) res- in a broad geographical sense. Typical Me- urrected the genus Wollastonia with the pri- lanthera extends into where Echin- mary intent of rescuing the type species, W. ocephalum is found, and African species biflora, from its common misplacement in treated as Melanthera by Wild (1965) in- the genus Wedelia. The uncertain relation- clude some with sterile and some with fer- ship to a more strictly defined Melanthera, tile rays. Even Wollastonia has a single with its more numerous capillary bristles, well-marked pappus bristle of the type seen lack of fertile rays, and white disk corollas, in some Melanthera. We have decided to was of secondary importance. follow Wild's concept in regard to Melanth- Wollastonia was established by Decaisne era, and treat all the species with striated (1834) for three species of the Pacific-In- receptacular bracts and nonfleshy, triangu- dian Ocean regions, one of which, W. sca- lar to tetrangular, truncate, winged achenes briuscula (Gaud.) DC. ex Decne., has been as members of the genus. We have exam- demonstrated by Fosberg and Sachet (1980) ined all possible taxa worldwide that might to be the best choice for the type of Wol- fit into this expanded concept of the genus. lastonia. Fosberg and Sachet also show that The primary exception of taxa we do not W. scabriuscula is both a taxonomic and include in Melanthera is the Hawaiian el- nomenclatural synonym of Wedelia biflora. ement that has 4-lobed corollas and Wede- In the same paper, they discussed the close lia-like elements in its hybrid history. This relationship between Lipochaeta and Wol- is typical allopolyploid Lipochaeta and lastonia, the latter recognized at that time consists of six species. on the basis of a single Pacific-wide spe- A second exception made at present in- cies. They suggested that part of or, better, volves a few species from Australia that do all of Lipochaeta is congeneric with Wol- not actually fit our broadened delimitation lastonia, but held off making the transfers of Melanthera. These species include the because Gardner's revision was recently presently named Wedelia asperrima Benth., completed but not yet published when they W. spilanthoides F Muell., and W. verbes- were writing their paper. Thus, there has inoides F Muell. & Benth. The latter two been an assumption, dating from Fosberg species have striated receptacular paleae as and Sachet (1980), that Wollastonia would in the Melanthera group, but they also have eventually by expanded to include other achenes biconvex and apically narrowed species from the Hawaiian Islands that were but not truncated to a narrow corolla inser- placed in Lipochaeta and perhaps even oth- tion. Wedelia asperrima Benth., which may er Old World taxa. be related to these two species, has heads The comments of Fosberg and Sachet that look Melanthera-like, but has less (1980) ignored the de facto reduction of strongly costate stems, lacks the striations Wollastonia to the synonymy of Melanth- in the paleae, and has much fatter though era by Wild (1965). They instead chose to still biconvex nontruncate achenes. We be- rely on the fertile rays and the single rather 200112001] WAGNER & ROBINSON: ASTERACEAE 543 than many pappus bristles to justify the sep- into synonymy or placed differently than aration from Melanthera. They mentioned where Gardner (1979) placed the name. several times that Wollastonia seemed to be We have included the geographical dis- closer to Lipochaeta than to the typical ray- tribution for each species insofar as we less, white, disk-flowered Melanthera, but know it. We have updated information on they did not further evaluate Wollastonia or the distribution of several Hawaiian, other Lipochaeta to the possible merits of Wild's Pacific, and South American species. We do wider delimitation of Melanthera. this, in part, so that all of the currently ac- cepted taxa and their distributions of both and Melanthera are available in Synopsis of Lipochaeta and Melanthera Lipochaeta one publication. This treatment differs from In this section we provide an account of Wagner et al. (1990) in that we here rec- all of the currently recognized species of ognize the extinct population from the is- Lipochaeta followed by those in Melanth- land of Lana'i included in Lipochaeta sub- era. Within the synopsis of Melanthera we cordata as Melanthera populifolia, and we have the species grouped into two alpha- give subspecies status to the infraspecific betical series: Pacific species with which we Hawaiian taxa. We believe that the geo- worked in greater detail; and the other ad- graphic/ecological distinctions along with ditional species of the genus from North the morphological diagnostic characters America, South America, and Africa. With that distinguish them are best recognized at a few exceptions that are discussed in the the subspecies rather than varietal rank. species notes, we have followed the most Because the achene morphology has in- recent treatments for delimitation of spe- creased significance in our delimitation of cies, including Parks (1973) for the white- Melanthera from Lipochaeta and other flowered, rayless group, Wild (1965) for the closely related genera of the Ecliptinae, we African species, and Wagner et al. (1990) include two plates (Figs. 1, 2) illustrating for the Hawaiian species. We evaluated the the important differences. We have selected taxonomic status of the few additional spe- several species of Melanthera and Lipo- cies from Pacific Islands, Asia, and South chaeta to show the range of variation as America not covered in those revisions. We well as the distinguishing characteristics of also provide here complete nomenclature the paleae, achene, and pappus. For com- and type information for the Pacific, South parison we chose a species each from Per- American species formerly in Echinoce- ymenium and Wedelia. phalum, and the Asian species. For the spe- cies covered in the revisions by Parks Key to Lipochaeta and Melanthera (1973) and Wild (1965) we provide only very abbreviated information, including no- 1. Disk corollas with 4 lobes; achenes tuber- menclature of only the accepted name and culate at maturity, the apex rounded to synonyms only if the name is sometimes sometimes truncate, hispidulous apically; still disk achenes flattened, 2-angled; ray recognized. achenes with a median The reason for a full nomenclatural treat- obcompressed ridge, rarely thickened and weakly 3-an- ment of the Hawaiian species is that Gard- gled; pappus of deciduous connate scales ner (1979) made numerous errors in typi- forming an uneven corona, sometimes fication, especially unnecessary lectotypifi- with short sharp or blunt awns and inter- cations, and additional were vening squamellae; plants with both fla- many species vonols and flavones n = 26 described after Gardner's treatment St. present; by -----.------Lipochaeta John (1984). Although St. John's names 1. Disk corollas with 5 lobes; achenes smooth were placed in synonymy by Wagner et al. to tuberculate at maturity, the apex abrupt- (1990) under the same species as we do ly narrowed, truncate, minutely hispidu- achenes not here, we have resolved in lous apically; disk flattened, problems typifi- achenes cation that could not be examined in the 4-angled; ray 3-angled; pappus usually of 1-15(-20) deciduous short bris- context of a floristic treatment. Moreover, tles; plants with flavonols present; n = 15 some names have been placed either newly Melanthera 544 BRITTONIA [VOL. 53

Lipochaeta truncate and hispidulous apically, with a small central corolla and pappus attach- ment, distal hispidulous surface of little or LIPOCHAETA DC., Prodr. 5: 610. 1836. Mi- no width, angles with narrow wings 0.3-0.6 crochaeta, Nutt., Trans. Amer. Philos. mm, often greatly enlarged at maturity, Soc. (n.s.) 7: 450. 1841, nom superfl. sometimes deeply lobed, sometimes with TYPE: (Gaud.) DC. narrow antrorse projections at upper cor- ners, surface not fleshy; pappus of connate Suffruticose perennial herbs; stems erect, scales forming an uneven corona, some- spreading to prostrate, rounded to quadran- times with short sharp or blunt awns and gular or subhexagonal, costate or ribbed, intervening squamellae. grains ca. sometimes rooting at the nodes. Leaves op- 27 Lmdiam. n = 26. posite, rarely ternate; petioles present and Lipochaeta is a genus of six species en- winged or absent and the bases connate- demic to the Hawaiian Islands. The genus perfoliate; blades chartaceous, subcoria- is here restricted to the allotetraploid taxa ceous, or somewhat fleshy (L. succulenta), with the diploid taxa formerly included here ovate to elliptic, lanceolate or linear-lance- transferred to Melanthera. The genus ap- olate, rarely deltate or spatulate, entire, pears to have had an origin via hybridiza- margins entire to deeply lobed, serrate or tion between Melanthera with n = 15 and irregularly and coarsely serrate, apex acute an unknown n = 11 Wedelia-like colonist, to obtuse, both surfaces subglabrous or perhaps a species of Wedelia. Although we more often appressed pubescent or hispi- can with assurance eliminate the autotetra- dulous and scabrous with glutinous dots. ploidy hypothesis for Lipochaeta (Gardner, Inflorescences terminal with heads solitary 1977a, 1977b; Gardner & La Duke, 1978), or few in cymose clusters. Heads with rays, we cannot be sure if the origin of the genus erect; involucre hemispherical to broadly was in the Hawaiian Islands or in a conti- campanulate; involucral bracts in 2 equal or nental source area, probably North America subequal series of 4-5 bracts each, ap- or South America. Because of the impres- pressed with scarcely spreading tips, ovate sive cytological variation in Wedelia, Carr to oblong, green to tan and sometimes pur- (1978) suggested that the n = 15 and n = ple-tinged near base and along midrib; re- 26 groups assigned to Lipochaeta were the ceptacle paleaceous, the paleae persistent, result of two separate introductions. This rigid, erect or arching over florets, weakly idea continued to be favored after they conduplicate at anthesis, flatter as achenes showed that the n = 26 group was of al- mature, obovate to oblanceolate, obtuse to lopolyploid derivation (Rabakonandrianina, nearly truncate, acute in one species, mu- 1980; Rabakonandrianina & Carr, 1981). cronate to apiculate, narrowly keeled and Without more precise information about the distinctly striate with many longitudinal actual parentage, we cannot determine ribs, sometimes tinged purple toward apex whether the hybridization event took place or along midrib, scabrid or asperulous out- before or after colonization, since both hy- side, especially toward tip; ray florets 6-16 potheses require two colonizations to the per head, pistillate and fertile, rays yellow, Hawaiian Islands. In a mainland origin the entire to 3-toothed; disk florets (15-)25-65 allopolyploid event took place in the con- per head, hermaphroditic, corolla, 4(-5)- tinental source area followed by coloniza- merous, yellow mostly glabrous to scabrid tion of the Lipochaeta ancestor (n = 26). distally, veins without fiber sheath, lobes This hypothesis requires an independent triangular to longer than wide, anther thecae colonization to the Hawaiian Islands by brownish, apical appendage yellow, ovate. Melanthera (n = 15), presumably M. biflo- Achenes tuberculate at maturity, flattened to ra. In the other hypothesis the only differ- slightly biconvex, in rays obcompressed ence is that the Wedelia-like (n = 11) an- with median ridge inside, rarely thickened cestor colonized the Hawaiian Islands and and more triangular, in disk flattened, 2-an- the allopolyploid event took place between gled, in rays and disk rounded or sometimes Hawaiian Melanthera or the colonizing an- 2001] WAGNER & ROBINSON: ASTERACEAE 545 cestor and this colonist, followed at some Sherff in Sherff, Bot. Gaz. (Crawfordsville) 95: extinction of the Wedelia-like col- 96. 1933. TYPE:U.S.A. Hawaiian Islands, Kaua'i, point by 2 mi from 18 Jul 0. 4185 onist. In addition to its Kekaha, 1932, Degener allopolyploid origin, (leg. O. Swezey) (HOLOTYPE:F-659426 [photo F]; species of Lipochaeta differ in having disk ISOTYPE:K-n.v.). corollas with 4 lobes (Gardner, 1977b) and, Lipochaeta alata Sherff var. pulchrior Sherff, Oc- based on another study by Gardner (1976), cas. Pap. Bernice P. Bishop Mus. 20(1): 20. 1949. in both flavonols and TYPE: U.S.A. Hawaiian Islands, Kaua'i, Puehu having flavones, Ridge, along Koke'e rd., 15 Mar 1948, F. Fager- whereas the diploid species here assigned lind & C. Skottsberg 6635 (HOLOTYPE: GB [photo to Melanthera have only flavonols. The F, BISH-4]; ISOTYPES:F). Gardner (1979) erro- achenes differ from those of Melanthera in neously cited this as a nom. nud. that they are tuberculate at maturity, flat- Distribution.-U.S.A. Hawaiian Islands, tened to and in florets slightly biconvex, ray scattered in remnant dry forest, 80-400 m, of in obcompressed. Species Lipochaeta on Kaua'i, and West Maui. general are more polymorphic, occur in of- ten extensive and are found at populations, 1b. connata (Gaud.) DC. lower elevations and in hotter habitats than Lipochaeta acris (Sherff) W. L. Wagner & H. most of Hawaiian Melanthera. subsp. species Rob., comb. et stat. nov. 1. LIPOCHAETA CONNATA (Gaud.) DC., Lipochaeta acris Sherff, Bot. Gaz. (Crawfordsville) connata DC. Prodr. 5: 611. 1836. connata 95: 83. 1933. Lipochaeta (Gaud.) var. acris (Sherff) R. C. Gardn., Rhodora 81: 316. Gaud., Voy. Uranie 464. 1829. Micro- 1979. TYPE: U.S.A. Hawaiian Islands, Kaua'i, chaeta connata (Gaud.) Nutt., Trans. Waimea, 2000-3000 ft, 1864-1865, H. Mann & Amer. Philos. Soc. (n.s.) 7: 452. 1841. W. T. Brigham 540 (HOLOTYPE:F-276925; ISO- TYPES: G-2 F], GH-2, MO, Lipochaeta australis (Less.) A. Gray var. BISH, [photo connata A. Proc. Amer. NY-n.v.). (Gaud.) Gray, Lipochaeta lobata (Gaud.) DC. var. incisior H. St. Acad. Arts. 5: 129. 1861. TYPE: U.S.A. John, Pacific Sci. 13: 185. 1959. TYPE: U.S.A. Hawaiian Islands, s.l., 1819, C. Gaudi- Hawaiian Islands, Ni'ihau, Ka'aliwai, 750 ft, 29 BISH- chaud-Beaupre s.n. (HOLOTYPE:P [photo Mar 1949, H. St. John 23572 (HOLOTYPE: 501492; ISOTYPES:BISH-2, GH, MO-n.v., US). F]). Lipochaeta acris Sherff var. lata H. St. John, Pacific Distribution.-U.S.A. Hawaiian Islands, Sci. 38: 253. 1984. TYPE: U.S.A. Hawaiian Is- bottom of 6 Mar scattered in remnant forest, 20-1160 m, lands, Kaua'i, Ka'aweiki, 1969, dry R. W. Hobdy 94 (HOLOTYPE:BISH-80404). on Ni'ihau, Kaua'i, and West Maui. Distribution.-U.S.A. Hawaiian Islands, la. LIPOCHAETA CONNATA (Gaud.) DC. scattered in remnant dry forest, 20-1,160 subsp. CONNATA m, on Ni'ihau and Kaua'i. Lipochaeta alata Sherff, Bot. Gaz. (Crawfordsville) 95: 81. 1933. TYPE: U.S.A. Hawaiian Islands, 2. LIPOCHAETA DEGENERI Sherff, Bot. Gaz. Kaua'i, along the Hanapepe River, near the Falls, (Crawfordsville) 95: 84. 1933. TYPE: 12 Jul 1895, A. A. Heller 2563 p.p. (HOLOTYPE: U.S.A. Hawaiian Islands, Moloka'i, SW ISOTYPES: NY F-429034; GH, [microfiche, photo near 16 1928, 0. F]). See Wagner & Shannon (1999) for further point, plain sea, May discussion of this type. Degener 4198 (HOLOTYPE: F-659444 Lipochaeta alata Sherff var. acrior Sherff, Bot. Gaz. [photo F]; ISOTYPES:A-n.v., F, G-n.v., (Crawfordsville) 95: 82. 1933. TYPE: U.S.A. Ha- GH-3, K-n.v., MO, NY-2 [micro- waiian U.S. Ex- Islands, Kaua'i, s.l., 1840, Expl. fiche], US, W-2). ped. s.n. (HOLOTYPE:US-57075 [photo F]; ISO- TYPES:GH [photo F], US). Distribution.-U.S.A. Hawaiian Islands, Lipochaeta profusa Sherff, Bot. Gaz. (Crawfords- known from three collections near the ville) 95: 95. 1933. TYPE:U.S.A. Hawaiian Is- only lands, Kaua'i/Ni'ihau, s.l., Jan 1885, 1. Sinclair southwestern point of Moloka'i, near sea s.n. (HOLOTYPE:K [photo F]). Date cited by Sherff level; probably extinct. Last collected in is the date the specimen was received at K, not 1928. the date of collection. Sherff cited island as Ha- wai'i, but the locality was determined to be on Kaua'i or Ni'ihau (St. John, 1954: 144). 3. LIPOCHAETA HETEROPHYLLAA. Gray, Lipochaeta profusa Sherff var. robustior 0. Deg. & Proc. Amer. Acad. Arts 5: 130. 1861. Li- 546 BRITTONIA [VOL. 53

pochaeta lobata (Gaud.) DC. var. heter- 400 m, on Ni'ihau, O'ahu, West Maui, and ophylla (A. Gray) Hillebr., Fl. Hawaiian collected once on Hawai'i. Isl. 209. 1888. TYPE:U.S.A. Hawaiian Is- lands, Maui, West Maui, 1840, U.S. Expl. 4a. LIPOCHAETALOBATA (Gaudich.) DC. Exped. s.n. (LECTOTYPE, designated by subsp. LOBATA 1935: ISOLECTOTYPE: Sherff, US-57063; Lipotriche australis Less., Linnaea 6: 510. 1831. Li- GH). Both the lectotype and the isolec- pochaeta australis (Less.) A. Gray, Proc. Amer. totype are mixed collections with L. Acad. Arts 5: 129. 1861. TYPE:U.S.A. Hawaiian rockii Sherff var. dissecta Sherff. The Islands, O'ahu, s.l., 1816-1817, L. C. A. von other material included in L. Chamisso s.n. (LE-n.v.). Sherff correctly cited heterophylla the Chamisso collection as the type, but Gardner by Gray was used as the type for L. rock- (1979) incorrectly (contrary to the ICBN Art. 52 ii Sherff var. dissecta Sherff (US). Gard- Note 1) assumed Lessing's name was based on ner (1979) incorrectly cited a Hillebrand Verbesina lobata because Lessing included the name with a that collection as the type. question mark, merely indicating it was possibly the same. When Gray transferred this taxon he included Verbesina lobata Gaud. Lipochaeta hastata Hillebr., Fl. Hawaiian Isl. 208. which australis 1888, nom. non hastata Kel- (1829), predates Lipotriche illeg., Lipochaeta and therefore he should have (1863). TYPE: U.S.A. Hawaiian Islands, (1831), adopted logg "lobata" as the for this taxon in Lana'i, 1851-1871, W. Hillebrand s.n. epithet Lipo- (HOLO- chaeta. TYPE:B [presumably destroyed; photo BISH, frag Verbesina hastulata Hook. & Arn., Bot. BISH-75694]; ISOTYPES:B-n.v., GH-n.v., MEL Beechey Voy. 87. 1832. Lipochaeta hastulata (Hook. & [photo BISH], US). Am.) DC., Prodr. 5: 611. 1836. Microchaeta lob- Lipochaeta peduncularis Drake, Ill. Fl. Ins. Pacif., ata (Gaud.) Nutt. var. hastulata (Hook. & Am.) part 4, 72, t. 35. 1888. TYPE:U.S.A. Hawaiian Nutt., Trans. Amer. Philos. Soc. II, 7: 451. 1841. Islands, Lana'i, s.l., 1851-1855, J. Remy s.n. (HO- Lipochaeta lobata (Gaud.) DC. var. hastulata LOTYPE:P--n.v.). (Hook. & Am.) Sherff, Bot. Gaz. (Crawfords- heterophylla A. var. molokaiensis Lipochaeta Gray ville) 95: 91. 1933. TYPE:U.S.A. Hawaiian Is- Sherff, Bot. Gaz. (Crawfordsville) 95: 96. 1933. lands, O'ahu, s.l., 1826-1827, G. T. Lay & A. TYPE:U.S.A. Hawaiian Islands, Moloka'i, W end Collie s.n. K Fl). of J. F. Rock 10287 F- (HOLOTYPE: [photo island, 1910, (HOLOTYPE: A. Proc. Amer. Acad. ISOTYPES: GH Lipochaeta calycosa Gray, 659408 [photo F]; BISH-4, [photo Arts 5: 130. 1861. TYPE: U.S.A. Hawaiian Is- F], UC-n.v.). lands, O'ahu, [Diamond Hill], 1840, U.S. Expl. Exped. s.n. (HOLOTYPE:US-57070; ISOTYPES:GH). Distribution.-U.S.A. Hawaiian Islands, Gardner (1979) incorrectly cited the GH sheet as scattered in open areas and margins of lava the holotype. flows, primarily coastal, in grassland, and Lipochaeta australis (Less.) A. Gray var. denticulata alien 0-400 on Lana'i and Wawra, Flora 56: 77. 1873. Lipochaeta lobata shrubland, m, DC. var. denticulata scattered localities on Moloka'i and (Gaudich.) (Wawra) Sherff, Maui. Bot. Gaz. (Crawfordsville) 95: 92. 1933. TYPE: U.S.A. Hawaiian Islands, O'ahu, s.l., 1868-1871, 4. LIPOCHAETALOBATA Prodr. H. Wawra 2294 (HOLOTYPE:W [photo F]). (Gaud.) DC., Ill. Fl. Ins. Pacif. 5: 611. 1836. Verbesina lobata Lipochaeta aprevalliana Drake, Gaud., 71, t. 34. 1888. Lipochaeta lobata (Gaud.) DC. Voy. Uranie 464. 1829. Microchaeta lob- var. aprevalliana (Drake) Sherff, Bot. Gaz. ata (Gaud.) Nutt., Trans. Amer. Philos. (Crawfordsville) 95: 92. 1933. TYPE:U.S.A. Ha- Soc. II, 7: 451. 1841. aus- waiian Islands, O'ahu, s.l., 1851-1855, J. Remy Lipochaeta 272 P-not The tralis A. var. lobata (HOLOTYPE: located). illustration (Less.) Gray (Gaud.) was incorrectly taken to be the holotype by Gard- A. Gray, Proc. Amer. Acad. Arts 5: 130. ner (1979). 1861. TYPE: U.S.A. Hawaiian Islands, Lipochaeta lobata (Gaud.) DC. var. albescens s.l., 1819, C. s.n. Sherff, Bot. Gaz. (Crawfordsville) 95: 92. 1933. Gaudichaud-Beaupre TYPE:U.S.A. Hawaiian Diamond ISOTYPES:G Islands, O'ahu, (HOLOTYPE: P-n.v.; [photo Head, [steep slopes], 28 Mar 1895, A. A. Heller F], B [presumably destroyed; photo F]). 2021 (HOLOTYPE:F-429425 [photo F]; ISOTYPES: Sherff cited this collection as Gaudi- A, GH [photo F], K, MO, NY [microfiche, photo chaud 138 (based on G sheet). F], UC-n.v., US). Lipochaeta lobata (Gaud.) DC. var. grossedentata Distribution.-U.S.A. Hawaiian O. Deg. & Sherff in Sherff, Bot. Gaz. (Crawfords- Islands, ville) 95: 92. 1933. TYPE:U.S.A. Hawaiian Is- scattered and often common in dry coastal lands, O'ahu, in gulch N of middle of ridge be- habitats, sometimes in dry shrubland, 0- tween Pu'u Pane and Pu'u Kamaohanui, 10 Jan 200112001] WAGNER & ROBINSON: ASTERACEAE 547

1932, 0. Degener, K. K. Park & W. Bush 4299 habitats, sometimes in dry shrubland, 0- (LECTOTYPE,designated by Gardner, 1979: F- 400 on West and 666547 ISOLECTOTYPES:F m, Ni'ihau, O'ahu, Maui, [photo F]; [photo F], collected once on Hawai'i. K-n.v.). Lipochaeta lobata (Gaud.) DC. var. hastulatoides 0. Deg. & Sherff in Sherff, Bot. Gaz. (Crawfords- 4b. Lipochaeta lobata (Gaud.) DC. subsp. ville) 95: 93. 1933. TYPE:U.S.A. Hawaiian Is- (0. & Sherff) W. L. SE Pohakea leptophylla Deg. lands, Maui, [West Maui, part], & H. Rob., comb. et stat. nov. Gulch, 11 Jul 1927, 0. Degener 4305 (LECTOTYPE, Wagner designated by Gardner, 1979: F-666545 [photo Lipochaeta lobata (Gaud.) DC. var. leptophylla 0. F]; ISOLECTOTYPES: B-n.v., BM-n.v., F [photo Deg. & Sherff in Sherff, Bot. Gaz. (Crawfords- F], G-n.v., GH, K-n.v., NY-2). ville) 95: 92. 1933. TYPE:U.S.A. Hawaiian Is- Lipochaeta subcordata A. Gray var. membranacea lands, O'ahu, Wai'anae Range, at Kolekole Pass, Sherff, Bot. Gaz. (Crawfordsville) 95: 91. 1933. 1-2 Feb 1915, C. N. Forbes 2024.0 (LECTOTYPE, TYPE: U.S.A. Hawaiian Islands, Maui, western designated here: F-485293 [photo F]; ISOLECTO- Maui, back of Lahaina, s.d., E. Bishop 14 (HO- TYPES:BISH, F-2 [photos F], K-n.v., NY- LOTYPE: B [presumably destroyed; photo F]). n.v., UC-n.v.). Sherff cited "3 type sheets" at F, Since this was the only collection we cannot be but Gardner cited one as the holotype (though he sure what species it represents, but it is likely to did not indicate which one). be L. lobata. Lipochaeta niihauensis H. St. John, Pacific Sci. 13: Distribution.-U.S.A. Hawaiian Islands, 186. 1959. TYPE: U.S.A. Hawaiian Islands, rare, known only from Kolekole Pass and Ni'ihau, Ki'i, 100 ft, 2 Apr 1949, H. St. John Kanehoa, Lualualei, Wai'anae Mountains, 23664 (HOLOTYPE: BISH-501495; ISOTYPES: O'ahu. This distinctive is at risk BISH-2, GH). subspecies Lipochaeta trilobata H. St. John, Pacific Sci. 30: 42. of extinction and has been listed as feder- 1976. TYPE: U.S.A. Hawaiian Islands, Hawai'i, ally endangered. [mtn. slope above Kealakekua], 26-29 Jan 1779, D. Nelson s.n. (HOLOTYPE:BM-n.v., photo OS- 5. LIPOCHAETAROCKII Sherff, Bot. Gaz. n.v.). 95: 100. 1933. TYPE: Lipochaeta christophersenii H. St. John, Pacific Sci. (Crawfordsville) 38: 253. 1984. TYPE:U.S.A. Hawaiian Islands, U.S.A. Hawaiian Islands, Moloka'i, W O'ahu, Wai'anae Mtns., central Lualualei, below end, below Kawela, Mapulou, 22 Mar Kanehoa, head of valley 2, foot of cliffs, 450 m, 1910, J. F. Rock 6156 (HOLOTYPE:GH Jun E. 3691 I 1932, Christophersen (HOLOTYPE: F]; ISOTYPES:BISH-3, F ISOTYPES: Collec- [photo [photo BISH-471459; BISH-2, US). Gardner cited 3 at tion year published as 1922 but specimen labels F]). (1979) isotypes reads 1932. F; only one seen (same as photo). (Fig. Lipochaeta garberi H. St. John, Pacific Sci. 38: 259. 1A-E) 1984. TYPE: U.S.A. Hawaiian Islands, O'ahu, Lipochaeta forbesii Sherff, Bot. Gaz. (Crawfords- right fork of Wailupe Valley, 12 Jan 1920, D. W. ville) 95: 83. 1933. TYPE:U.S.A. Hawaiian Is- Garber & C. N. Forbes 141 (HOLOTYPE: BISH- lands, Maui, [E Maui], Nu'u, S slope of Halea- 75861). Forbes included as a collector on speci- kala, 9 Mar 1920, C. N. Forbes 1916.M (HOLO- men label but not in publication. TYPE: F-659397; ISOTYPES:BISH-2, K-n.v., Lipochaeta kaenaensis H. St. John, Pacific Sci. 38: NY-n.v., UC-n.v., US). 260. 1984. TYPE: U.S.A. Hawaiian Islands, Lipochaeta A. var. malvacea 0. O'ahu, Waialua Distr., Ka'ena, near base of cliff heterophylla Gray Deg. & Sherff in Sherff, Bot. Gaz. (Crawfords- near Manini Gulch, 120 ft, 25 Apr 1978, K. Na- ville) 95: 96. 1933. TYPE: U.S.A. Hawaiian Is- gata, B. Kimura, & G. Spence 1626 (HOLOTYPE: lands, Moloka'i, [W Moloka'i], near Kolo, 5 Apr BISH-421917). 1928, 0. Degener 4199 (LECTOTYPE, designated Lipochaeta lobata (Gaud.) DC. var. hosakae H. St. by Gardner, 1979: F-659443; ISOLECTOTYPES: B- John, Pacific Sci. 38: 263. 1984. TYPE: U.S.A. n.v., BISH, BM-n.v., CAS-n.v., F-3, G-2, Hawaiian Islands, O'ahu, Kipapa Gulch, Waipi'o, GB n.v., GH, K-n.v., MO-2, NY [micro- 24 Feb E. Y. Hosaka 1281 1935, (HOLOTYPE: fiche], UC-n.v., US, W-n.v.). ISOTYPES: BISH-146402; BISH). Lipochaeta kahoolawensis Sherff, Bot. Gaz. (Craw- vittata H. St. Pacific Sci. 38: 278. Lipochaeta John, fordsville) 95: 98. 1933. TYPE:U.S.A. Hawaiian 1984. TYPE: U.S.A. Hawaiian Islands, O'ahu, Islands, Kaho'olawe, s.l., 1851-1855, J. Remy Ka'ena Point, N. Shore Rd., 1/2 mi W of im- 269 (HOLOTYPE: P [photo F]). road 100 from 10 m, 7 Jan proved end, yd. ocean, Lipochaeta lobata (Gaud.) DC. var. maunaloensis R. C. Gardner 281 BISH- 1974, (HOLOTYPE: Sherff, Bot. Gaz. (Crawfordsville) 95: 93. 1933. 419080; ISOTYPES:OS-n.v.). TYPE:U.S.A. Hawaiian Islands, Moloka'i, Maun- aloa, Jun 1912, C. N. Forbes 7.Mo (HOLOTYPE: F- Distribution.-U.S.A. Hawaiian Islands, 659396 [photo F]; ISOTYPE:BISH). scattered and often common in dry coastal Lipochaeta rockii Sherff var. dissecta Sherff, Bot. 548 BRITTONIA [VOL[VOL. 53 - v' I I E .

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A " K - L N P FIG. 1. Achenes and paleae of Melanthera, Lipochaeta (Ecliptinae). All specimens deposited in US. A-E. Lipochaeta ro(kii (Hawaiian Islands, Moloka'i, Degener 22207). A. Pale. B. Ray achene. C. Cross section of ray achene. D. Disk achene. E. Cross section of disk achene. F-J. (Caroline Islands, Palau Islands, Fosberg 25Y82). F. Pale. G. Ray achene. H. Cross section of ray achene. I. Disk achene. J. Cross section of disk achene. K-O. M. lavarum (Hawaiian Islands, Hawai'i, Degener 4188). K. Pale. L. Ray achene. M. Cross section of ray achene. N. Disk achene. 0. Cross section of disk achene. P-R. M. nivea (Equador, Balao, Eggers 14339) [note: ray flowers absent in this species]. P. Pale. Q. Disk achene. R. Cross section of disk achene. 20011201IANR&RBNO:ATRCA WAGNER & ROBINSON: ASTERACEAE 5494

Gaz. (Crawfordsville) 95: 101. 1933. TYPE: Islands, Kaho'olawe, alongside jeep rd. from U.S.A. Hawaiian Islands, Maui, E Maui, 1840, Hanakanaea Bay, past dump, just S of concrete U.S. Expl. Exped. s.n. (HOLOTYPE: US-57065 observation post, S side of rd., 90 ft, 21 Apr 1980, [photo F]; ISOTYPE:GH-n.v.). See also Lipo- G. Clarke & C. Corn 374 (HOLOTYPE:BISH- chaeta heterophylla. 475921). Lipochaeta rockii Sherff var. subovata Sherff, Bot. Gaz. (Crawfordsville) 95: 101. 1933. TYPE: Distribution.-U.S.A. Hawaiian Islands, U.S.A. Hawaiian Islands, Moloka'i, s.l., 1851- scattered to common in coastal sites to dry 1855, J. Remy 270 (HOLOTYPE: P [photo F]). forest, often in disturbed areas and succulenta & DC. var. tri- margins Lipochaeta (Hook. Arn.) of lava 15-550 m, on Moloka'i, from fida Sherff, Bot. Gaz. (Crawfordsville) 95: 87. flows, 1933. TYPE:U.S.A. Hawaiian Islands, Moloka'i, scattered localities on Maui, common near Manawai, Aug 1912, C. N. Forbes 397.Mo (LEC- the coast on Kaho'olawe, also a single col- TOTYPE,designated by Gardner, 1979: F-485234 lection presumably from the island of Ha- [photo F]; ISOLECTOTYPES:BISH-3, F-2 [photo wai'i. of "2nd type sheet" F], UC-n.v.). Lipochaeta forbesii Sherff var. sherffii 0. Deg. & Clay, Fl. Hawaiiensis, fam. 344. Lipochaeta for- 6. LIPOCHAETA SUCCULENTA (Hook. & Arn.) besii. 1949. TYPE: U.S.A. Hawaiian Islands, DC., Prodr. 5: 611. 1836. Verbesina suc- Maui, [E Maui, S slope of Haleakala], between culenta Hook. & Bot. and Palaha on a'a 1500 Arn., Beechey Kepuni Gulches, lava, ft, 87. 1832. Microchaeta succulenta 25 Dec 1948, 0. Degener 19292 (HOLOTYPE: Voy. BISH-501024; ISOTYPES:A-n.v., B-n.v., BISH, (Hook. & Arn.) Nutt., Trans. Amer. Phi- F-3 [photo of "sheet 1" F], G-n.v., GB-n.v., los. Soc. II, 7: 451. 1841. TYPE: U.S.A. GH, NY-2 [microfiche], US). The specimen in Hawaiian Islands, Ni'ihau, ("Onee- the F photo ("sheet 1") was remounted after it G. T. & A. Col- was heow"), 1826-1827, Lay photographed. lie s.n. Lipochaeta lobata (Gaud.) DC. var. makenensis 0. (HOLOTYPE: K-n.v.). & Sherff in Brittonia 12: 174. 1960. Deg. Sherff, lanceolata Nutt., Trans. Amer. Philos. TYPE:U.S.A. Hawaiian Lipochaeta Islands, Maui, [E Maui], Soc. II, 7: 451. 1841. TYPE:U.S.A. Hawaiian Is- 40 1 0. I. De- Makena, ft, Apr 1959, Degener, lands, O'ahu, near sea, Oct-Jan 1835-1836, T. & W. 25133 F- gener Fleming (HOLOTYPE: Nuttall s.n. (HOLOTYPE:BM-n.v.). 1537169; ISOTYPES:BISH, F-2, G-n.v., GH- Lipochaeta australis (Less.) A. Gray var. decurrens 2, MEL NY UC- [2 photos BISH], [microfiche], A. Gray, Proc. Amer. Acad. Arts 5: 129. 1861. n.v., US). Lipochaeta connata (Gaud.) DC. var. decurrens scabra H. St. Pacific Sci. 30: 40. Lipochaeta John, (A. Gray) Hillebr., Fl. Hawaiian Isl. 206. 1888. 1976. TYPE:U.S.A. Hawaiian Islands, Hawai'i, Lipochaeta succulenta (Hook. & Arn.) DC. var. 26-29 Jan D. Nelson s.n. s.l., 1779, (HOLOTYPE: decurrens (A. Gray) Sherff, Bot. Gaz. (Craw- BM-n.v.). fordsville) 95: 86. 1933. TYPE:U.S.A. Hawaiian H. St. Pacific Sci. 38: Lipochaeta elliptica John, Islands, Kaua'i, s.l., 1840, U.S. Expl. Exped. s.n. 255. 1984. TYPE: U.S.A. Hawaiian Islands, (HOLOTYPE: US-57076). Superfluously lectotypi- rd. from Hanakanaea Kaho'olawe, alongside jeep fied by Gardner (1979). the and S of concrete obser- Bay, past dump just Lipochaeta connata (Gaud.) DC. var. littoralis Hil- vation post, 90 ft, 21 Apr 1980, G. Clarke & C. lebr., Fl. Hawaiian Isl. 206. 1888. TYPE:U.S.A. Corn 370 (HOLOTYPE:BISH-475923). Hawaiian Islands, Moloka'i, Kalawao, 1851- H. St. Lipochaeta elliptica H. St. John var. artialata 1871, Hillebrand s.n. (LECTOTYPE, designated John, Pacific Sci. 38: 257. 1984. TYPE:U.S.A. here: BISH-146350). Three collections were cited Hawaiian Islands, Kaho'olawe, on coast, W side by Hillebrand. of island, just NW of Hanakanaea Bay, 20-40 ft, Lipochaeta variolosa H. L6v., Repert. Spec. Nov. 22 Apr 1980, L. W. Cuddihy & W. Char 382 (HO- Regni Veg. 10: 122. 1911. TYPE:U.S.A. Hawaiian LOTYPE: BISH-475924). Islands, Kaua'i, Wainiha, Jan 1910, U. Faurie Lipochaeta mauiensis H. St. John, Pacific Sci. 38: 1008 (LECTOTYPE, designated by Gardner, 1979: 265. 1984. TYPE:U.S.A. Hawaiian Islands, Maui, P; ISOLECTOTYPES:BISH, G-2-n.v.). E Maui, Hwy 31, 5.5 mi SE of 'Ulupalakua Lipochaeta succulenta (Hook. & Arn.) DC. var. an- Ranch Office, 500 m, 18 Feb 1975, R. C. Gardner gustata Sherff, Bot. Gaz. (Crawfordsville) 95: 87. 378 (HOLOTYPE:OS-n.v.; ISOTYPE:BISH). 1933. TYPE:U.S.A. Hawaiian Islands, Kaua'i, s.l., Lipochaeta nesophila H. St. John, Pacific Sci. 38: 1909, C. N. Forbes s.n. (HOLOTYPE: F-485175 270. 1984. TYPE: U.S.A. Hawaiian Islands, [photo F]; ISOTYPES:BISH, F [photo F]). Ni'ihau, S half of island, Jan 1912, J. F. G. Stokes Lipochaeta succulenta (Hook. & Arn.) DC. var. bar- s.n. (HOLOTYPE:BISH-475922). The locality is in clayi Sherff, Bot. Gaz. (Crawfordsville) 95: 87. some doubt since Lipochaeta rockii is otherwise 1933. TYPE: U.S.A. Hawaiian Islands, Kaua'i, unknown from Ni'ihau. "Atooi" hills, Jul 1837, G. Barclay 1327 (HOLO- Lipochaeta rockii Sherff var. parva H. St. John, Pa- TYPE:BM [photo F]). cific Sci. 38: 276. 1984. TYPE:U.S.A. Hawaiian Lipochaeta robusta H. St. John, Pacific Sci. 38: 274. 550 BRITTONIA [VOL. 53

1984. TYPE:U.S.A. Hawaiian Islands, Maui, East Wollastonia DC. ex Decne., Nouv. Ann. Mus. Paris Maui, Hana, Pu'u Ki [Pu'uki'i], 900 ft, 31 Dec 3: 414. 1834. LECTOTYPE:W. scabriuscula DC. ex 1936, H. St. John & R. J. Catto 17891 (HOLOTYPE: Decne., nom. illeg.; designated by Fosberg, 1980: BISH-471452; ISOTYPES:BISH-2, GH, US). 32 [=Melanthera biflora (L.) Wild]. Psathurochaeta DC., Prodr. 5: 609. 1836. TYPE: Distribution.-Hawaiian Islands, scat- Psathurochaeta dregei DC. [=Melanthera scan- tered along the coasts, usually in a narrow dens (Schumach. & Thonn.) Roberty subsp. dre- band within 30 m of the water, 0-100 m, gei (DC.) Wild]. on all of the main islands Aphanopappus, Endl., Gen. P1. Suppl. 2. 43. 1842. except Lana'i, Schizophyllum Nutt., Trans. Amer. Philos. Soc. and very rare on O'ahu. (n.s.) 7: 452. 1841, non E. M. Fries (1821). TYPE: Aphanopappus nuttallii Walpers, nom. illeg. Melanthera [based on Schizophyllum micranthum Nutt.] [=Melanthera micrantha (Nutt.) W. L. Wagner & H. Rob.]. Wuerschmittia ex 6: 161. MELANTHERARohr, Skr. Naturhist.-Selsk. Sch.-Bip. Walp., Repert. 1846. TYPE: Wuerschmittia abyssinica Sch.-Bip. 2(1):213. 1792. NEOTYPE:Melanthera ni- ex Walp. [=Melanthera abyssinica (Sch.-Bip. ex vea (L.) J. K. Small; designated by Cas- A. Rich.) Benth. & Hook. f.]. sini, J. Phys. Chim. Hist. Nat. Arts 87: Echinocephalum Gardner, London J. Bot. 7: 294. 27. 1818. For a discussion on 1848. LECTOTYPE:Echinocephalum angustifolium typifica- here see D. H. Taxon 30: 491- Gardn.; designated [=Melanthera latifolia tion, Nicolson, (Gardn.) Cabrera, not M. angustifolia A. Rich.]. 492. 1981. The problem with the typifi- cation of this name arises because ICBN Perennial herbs, vines, subshrubs, some- Art 10.1 requires that the type of a ge- times ; stems often pale or maculate, neric name is the type of a name of a quadrangular to subhexagonal, costate, usu- species. When von Rohr published the ally sulcate when dry, appressed strigose to generic name he cited only a specimen asperulate or hispid. Leaves opposite, peti- (von Rohr s.n., C) from Martinique that oles usually conspicuous, less often lacking has never been used as type for the name or winged; blades herbaceous to charta- of any species. Thus, Cassini's mention ceous, ovate or deltate to linear, sometimes of nivea L. as the type of Me- strongly trilobed to pinnately lobed, rarely lanthera can be taken as a neotypifica- ternately compound, margins remotely to tion. Because no species was included closely serrulate or serrate, often with large when the genus was published, typifica- teeth or lobes at broadest part, base sub- tion has been debated by Strother (Taxon truncate to narrowly cuneate or acuminate, 19: 337. 1970), who considered the type apex acute to narrowly acuminate, upper to be M. panduriformis Cass. [=M. ni- surface mixed scabrid and scabridulous, vea], whereas Parks (1973) considered it lower surface appressed strigose to hispid, to be Calea aspera Jacq. [=M. aspera, without glandular dots, usually trinervate which here is a synonym of M. niveal. from near base, rarely penninerved with One could propose a conserved type on short ascending veins. Inflorescences ter- von Rohr's specimen, which, if success- minal or from axils of uppermost leaves, would a ful, provide conserved type vs. with 1-3 heads. Heads with or without rays, a but the neotype; controversy over what erect; involucre hemispherical to broadly name species should be considered type campanulate; involucral bracts indurate would still remain. In of the contro- spite with somewhat herbaceous tips, appressed none of the different selections of versy, with scarcely spreading tips, 10-14 in 1 or has effect on the of type any application 2 subequal series, oblong to narrowly ovate, the name. apex obtuse to acute or attenuate, strigillose P. Browne, Civ. Nat. Hist. Jamaica. 317. outside; receptacle paleaceous, the paleae 1756, nom. rejec. (vs. Amellus L.). TYPE:Santo- persistent, chartaceous with membrana- lina amellus L. (= (L.) J.K. ceous bases, weakly conduplicate and en- Small). the flatter as Lipotriche R. Brown, Observ. Campos 118. 1817. folding achene, becoming TYPE: Lipotriche brownei DC. [=Melanthera achene matures, scarcely to narrowly scandens (Schumach. & Thonn.) Roberty]. keeled, closely multistriated with longitu- 2001] WAGNER & ROBINSON: ASTERACEAE 551 dinal ribs outside, oblanceolate, tips obtuse has been credited with numerous species, to mucronate, or lanceolate, or awned, as- and we here recognize 35 as distinct. Dis- perulous or toothed distally. Ray florets 0- tribution is from eastern North America, 12, neutral or fertile; corollas 5-merous, through , the West Indies, yellow, tube ca. 1 mm long, glabrous, limb South America, Africa, Asia, and many In- oblong, 2.3-13 mm long, apex bidentate, dian and Pacific Ocean islands. Melanthera strigillose below, without glandular dots, appears to be most closely related to the papillose above; style absent or present. neotropical genus Wulffia ( G. Mey.) Disk florets ca. 10-100, bisexual; corollas based on the shared characters of strongly white or yellow, tube glabrous; throat nar- striate paleae and truncate achene apex. rowly campanulate, glabrous outside or sca- Wulffia is distinguished from Melanthera bridulous above and along veins, veins by its specialized fleshy achenes and lack without fiber sheath or with few fibers, of pappus. lobes triangular to longer than wide, dense- All species of Melanthera that have been ly scabridulous outside, without glands, pa- examined thus far have the same haploid pillose inside, papillae often long, often chromosome number, n = 15 (Robinson et only on margin; anther thecae black, apical al., 1981). Examination of chromosome appendage yellow, ovate. Achenes 3-angled number atlases for the past two decades in- (disk achenes 4-angled), abruptly narrowed dicate other species have been counted, and or truncate above, upper surface broad, mi- all but three determinations were n = 15. nutely hispidulous, with small central co- The three exceptions we believe should be rolla and pappus attachment, with or with- verified before being accepted. They are: out wings on angles or with a pseudopap- M. biflora (as Wedelia bifiora) 2n = 45 pus of projections on distal corners and (Ono, 1975, 1977), 2n = 50 (Mathew & edges, surface not fleshy; pappus usually of Mathew, 1978), and M. elliptica n = 16 1-15(-20) deciduous short bristles, mostly (Gill & Omoigui, 1992). 1-2 mm long. Pollen grains 23-25 jtm diam. n = 15. PACIFIC SPECIES OF MELANTHERA Melanthera is delimited by the abruptly narrowed to truncate and flattened top of 1. MELANTHERA BIFLORA Wild, Prodr. 5: the achene with a cluster of 1-15(-20) pap- (L.) 546. 1836. Verbesina bifiora L., PI., pus bristles immediately surrounding the Sp. ed. 2, 1272. 1763. Wollastonia scabrius- corolla, involucral bracts and receptacular with veins cula DC. ex Decne., Nouv. Ann. Mus. paleae many forming longitudi- Paris 3: 414. nom. Stemmo- nal striations, and n = 15. The florets are 1834, illeg. dontia W. F in Saf- 5-merous, the corollas are or white, biflora (L.) Wight yellow Contr. U.S. Natl. Herb. 9: 377. and are absent (in white-flowered ford, rays spe- 1905. TYPE: India cies) or present and neutral or fertile. Typ- (HOLOTYPE:LINN). ical Melanthera has white corollas, lacks (Fig. IF-J) rays, completely lacks fibers in the corolla Lipochaeta ovata R. C. Gardn., Rhodora 81: 321. throat, and has papillae not restricted to the 1979, syn. nov. TYPE: U.S.A. Hawaiian Islands, of the inner surfaces of the disk O'ahu, Honolulu, 1852, N. J. Anderson s.n. (HOLO- margins TYPE:GB [photo OS]). Many of the Anderson col- corolla lobes. The genus is broadened here lections appear to have been labeled incorrectly: to include Echinocephalum of southeastern though stating "Hawaiian Islands," they were ac- South America that has yellow florets, neu- tually collected in southeastern Polynesia. tral disk corolla throats with some fi- For additional synonymy see Fosberg & Sachet, 1980; rays, Smith & Nicolson & ber cells the and inner surfaces Carr, 1991; Fosberg, 1993; along veins, Fosberg, in press. of the disk corolla lobes with papillae only along the margins. Wild (1965) is followed Distribution.-Melanthera bifiora is a in his African treatment that included the widespread coastal species (sometimes oc- fertile-rayed Lipotriche and Wollastonia in curring inland in disturbed sites) occurring the synonymy of Melanthera. We here add from eastern Africa to southeastern Asia, Lipochaeta sect. Aphanopappus. The genus southern south to eastern Australia, 552 BRITTONJABRITTONIA [VOL. 53 and most islands in the Indian and Pacific Lipochaeta integrifolia (Nutt.) A. Gray var. argentea Bot. Gaz. 95: 84. Oceans (but not in the Hawaiian Islands). Sherff, (Crawfordsville) 1933. TYPE:U.S.A. Hawaiian Islands, Maui, on sandy isthmus, 1864-1865, H. Mann & W. T. Brigham 2. Melanthera bryanii (Sherff) W. L. Wag- 371 (HOLOTYPE:F-276879 [photo F]; ISOTYPES: ner & H. Rob., comb. nov. BISH, F, G-n.v., GH-2, MO, NY-n.v., US). Lipochaeta integrifolia (Nutt.) A. Gray var. gracilis Lipochaeta bryanii Sherff, Bot. Gaz. (Crawfords- Sherff, Bot. Gaz. (Crawfordsville) 95: 85. 1933. ville) 95: 97. 1933. TYPE:U.S.A. Hawaiian Is- TYPE: U.S.A. Hawaiian Islands, s.l., Sep-Oct lands, Kaho'olawe, 300 m, 16 Feb 1931, E. H. 1836, C. Gaudichaud-Beaupre 217 (HOLOTYPE: Bryan Jr. 736 (HOLOTYPE:BISH-501028 [photo GH [photo F]; ISOTYPES:B [presumably de- F]; ISOTYPE:BISH). stroyed; photo F], P-n.v.). Lipochaeta integrifolia (Nutt.) A. Gray var. major Distribution.-U.S.A. Hawaiian Islands, Sherff, Bot. Gaz. (Crawfordsville) 95: 85. 1933. known only from the type collection on TYPE:U.S.A. Hawaiian Islands, O'ahu, on the old slopes in pili grassland at 300 m on lava flow back of Diamond Head, 8 Apr 1895, A. Kaho'olawe. Melanthera is A. Heller 2092 p.p. (HOLOTYPE:GH [photo F]; ISO- bryanii appar- TYPES: NY extinct. A-n.v., MO, [microfiche; photo F], ently US). See Wagner & Shannon, 1999, for further discussion of this type. 3. Melanthera fauriei (H. Lev.) W. L. Lipochaeta integrifolia (Nutt.) A. Gray var. mega- Wagner & H. Rob., comb. nov. cephala 0. Deg. & Sherff in Sherff, Bot. Gaz. (Crawfordsville) 95: 86. 1933. TYPE:U.S.A. Ha- Lipochaeta fauriei H. Lev., Repert. Spec. Nov. Reg- waiian Islands, O'ahu, Ka'ena Point, 5 m, 14 Dec ni Veg. 10: 122. 1911. TYPE: U.S.A. Hawaiian 1930, E. Christophersen 1400 (HOLOTYPE:F- Islands, Kaua'i, Holokele [Olokele], Mar 1910, 659415 [photo F]; ISOTYPE:BISH). U. Faurie 1012 (HOLOTYPE:P [photo F]; ISOTYPE: Lipochaeta porophila 0. Deg. & I. Deg., Flora Ha- BM). Sherff labeled the photo of the P sheet as waiiensis Fam. 344. Lipochaeta porophila. 1970. "cotype," but in his monograph (1935) he made TYPE:U.S.A. Hawaiian Islands, Hawai'i, Kau, be- it clear that he did not know where the holotype tween Punaluu and Kamehame Hill, near coast, 7 was and that he was not sure whether the P sheet Jun 1969, 0. Deger,Dege. Degener & Mr. and was a "cotype" or the holotype. BM isotype cited Mrs. Picco 31985 (H()LOTYPE:NY |microfiche]; by Lauener (1980). ISOTYPES:MO-n.v., US). Lipochaeta deltoidea H. St. John, Pacific Sci. 26: 291. 1972. TYPE: U.S.A. Hawaiian Islands, Distribution.-U.S.A. Hawaiian Islands, Kaua'i, Lower Hikimoe Valley, 1800 ft, 18 Apr scattered to locally common along the 1969, R. W. Hobdy 102 (HOLOTYPE:BISH- coasts of Kure Atoll, Laysan, and all of the 468055; ISOTYPES:BISH, GH). Gardner (1979) main islands. superfluously selected one of BISH sheets as the lectotype, and the BISH sheets are annotated as lectotype and isolectotype. St. John illustrated the 5. Melanthera kamolensis (0. Deg. & specimen we take to be the holotype. Sherff) W. L. Wagner & H. Rob, comb. nov. kamolensis 0. & Distribution.-U.S.A. Hawaiian Islands, Lipochaeta Deg. known from a few collections from diverse Sherff in Sherff, Amer. J. Bot. 38: 63. 1951. TYPE: U.S.A. Hawaiian mesic forest, 480-900 m, in Koai'e Can- Islands, Maui, [southernmost central E Maui], yon, Olokele Canyon, Hikimoe Valley, and Kamole Gulch, 21 Dec 1948, 0. Dege- Po'opo'oiki Valley, below confluence with ner, H. F. Clay, R. Bertram 19288 (HO- Ku'ia Valley, Kaua'i. Melantherafauriei is LOTYPE:F F]; ISOTYPES:B-n.v., at risk of extinction and is federally listed [photo BISH, BM-n.v., CM-n.v., CU-n.v., as endangered. F [photo F, BISH], G-n.v., GH-2, K- n.v., LA-n.v., M-n.v., MO-2, NY- 4. Melanthera integrifolia (Nutt.) W. L. n.v., P-n.v., PH-n.v., US-2). Gard- Wagner & H. Rob., comb. nov. Micro- ner's lectotypification (1979) of a GH chaeta integrifolia Nutt., Trans. Amer. sheet is superfluous. Philos. Soc. II, 7: 451. 1841. Lipochaeta integrifolia (Nutt.) A. Gray, Proc. Amer. Distribution.-U.S.A. Hawaiian Islands, Acad. Arts 5: 130. 1861. TYPE: U.S.A. known only from a few collections near one Hawaiian Islands, Kaua'i, ("Atooi"), 4- another in remnant dry forest at ca. 250 m: 26 Jan-Mar 1835, T. Nuttall s.n. (HOLO- southeast of 'Ulupalakua Ranch office, Ka- TYPE:BM). mole Gulch, and west of Kepuni Gulch, 2001] WAGNER & ROBINSON: ASTERACEAE 553 southeastern Maui. Melanthera kamolensis slopes above Manele, Jun 1913, C. N. Forbes is now restricted to one of less 288.L (LECTOTYPE, designated by Gardner, 1979: population F-659394 ISOLECTOTYPES: F than 1000 and has been listed [photo F]; BISH, individuals, [photo F], NY-2-n.v., US, W-n.v.). as federally endangered. Lipochaeta lavarum (Gaud.) DC. var. stearnsii 0. Deg. & Sherff in Sherff, Field Mus. Nat. Hist., Bot. Ser. 17: 581. 1939. TYPE:U.S.A. Hawaiian 6. Melanthera lavarum (Gaud.) W. L. Islands, Lana'i, Kapoho Canyon, 800 ft, Jun Wagner & H. Rob., comb. nov. (Fig. 1936, 0. Degener 11050 (leg. H. Stearns) (HO- 1K-O) LOTYPE:F-954573 [photo F]; ISOTYPES:F, GH, MO-n.v., NY-3 [microfiche]). Verbesina lavarum Gaud., Voy. Uranie 464. 1829. Lipochaeta lavarum (Gaud.) DC. var. scaposa H. St. Lipochaeta lavarum (Gaud.) DC., Prodr. 5: 611. John, Pacific Sci. 38: 263. 1984. TYPE: U.S.A. 1836. Microchaeta lavarum (Gaud.) Nutt., Trans. Hawaiian Islands, Kaho'olawe, E coast of Wai- Amer. Philos. Soc. II, 7: 451. 1841. TYPE:U.S.A. kahalulu Bay, slope below sea cliff, 45 ft, 22 Apr Hawaiian Islands, s.l., 1819, C. Gaudichaud- 1980, G. Clarke & C. Corn 377 (HOLOTYPE: Beaupre s.n. (HOLOTYPE:P [photo F]; ISOTYPE:B BISH-475920). [presumably destroyed; photo F]). Lipochaeta molokiniensis H. St. John, Pacific Sci. Lipochaeta lahainae Wawra, Flora 56: 77. 1873. 38: 267. 1984. TYPE:U.S.A. Hawaiian Islands, TYPE:U.S.A. Hawaiian Islands, Maui, [W Maui], Molokini, W side, N of lighthouse, 90 ft, 23 Apr Lahaina, 1869-1870, H. Wawra 1972 (HOLOTYPE: 1980, G. Clarke 401 (HOLOTYPE:BISH-475925). W-n.v.). Lipochaeta lavarum (Gaud.) DC. var. hillebrandi- Distribution.-U.S.A. Hawaiian Islands, ana Sherff, Bot. Gaz. (Crawfordsville) 95: 89. scattered to sometimes locally common pri- 1933. TYPE:U.S.A. Hawaiian Islands, Maui, La- in coastal habitats but also inland in haina, near sea, 1851-1871, W. Hillebrand s.n. marily (HOLOTYPE:B [presumably destroyed; photo F]). dry forest and rarely in the dry phases of Lipochaeta lavarum (Gaud.) DC. var. longifolia mesic forest, 0-550 m, on Moloka'i, Sherff, Bot. Gaz. (Crawfordsville) 95: 90. 1933. Lana'i, Maui, Kaho'olawe, and northwest- TYPE:U.S.A. Hawaiian Islands, Lana'i, Maunalei ern Hawai'i. Valley, 9 Mar 1915, G. C. Munro 202 (HOLOTYPE: BISH-500972 [photo F]; ISOTYPE:BISH [pre- sumed]). 7. Melanthera lifuana (Hochr.) W. L. Wag- Lipochaeta lavarum (Gaudich.) DC. var. ovata ner & H. Rob., comb. nov. Sherff, Bot. Gaz. (Crawfordsville) 95: 88. 1933. TYPE:U.S.A. Hawaiian Islands, Maui, E Maui, Lipochaeta lifuana Hochr., Bull. New York Bot. below the crater at Kahikinui, Nov 1910, J. F. Gard. 6: 297. 1910. Wedelia lifuana (Hochr.) Hochr. ex Bull. Soc. Bot. France 84: Rock 8674 (HOLOTYPE:GH [photo F]; ISOTYPES: Guillaum., BISH-5, CAS-n.v., F [photo F], K-n.v., 59. 1927. Wollastonia lifuana (Hochr.) Fosb., Al- NY-n.v., UC-n.v.). lertonia 7: 80. 1993. TYPE:New Caledonia. Loy- Lipochaeta lavarum (Gaud.) DC. var. salicifolia alty Islands, Lifou, 1861-1867, E. Viellard 799 Sherff, Bot. Gaz. (Crawfordsville) 95: 88. 1933. (HOLOTYPE:NY; ISOTYPES:GH, K-n.v., NY-2). TYPE:U.S.A. Hawaiian near La- Islands, Maui, Distribution.-New haina, s.d., E. Bishop s.n. (HOLOTYPE: B [presum- Caledonia, including ably destroyed; photo F]). the Loyalty Islands, the Ile des Pins, and Lipochaeta lavarum (Gaud.) DC. var. skottsbergii Grande Terre, along the coasts. Also col- Sherff, Bot. Gaz. (Crawfordsville) 95: 89. 1933. lected at least once on Annatam [Anatom] TYPE:U.S.A. Hawaiian Islands, Maui, s.l., 1833- of the New Hebrides as cited Sherff 1836, F. D. Bennett 43 (HOLOTYPE: B [presumably by destroyed; photo F]). (1935: 63). Lipochaeta lavarum (Gaud.) DC. var. conferta Sherff, Field Mus. Nat. Hist., Bot. Ser. 17: 582. 8. Melanthera micrantha (Nutt.) W. L. 1939. TYPE:U.S.A. Hawaiian s.l., Islands, Lana'i, Wagner & H. Rob., comb. nov. 1864-1865, H. Mann & W. T. Brigham 358 (HO- LOTYPE:F-276873 [photo F]; ISOTYPES:BISH-2, Schizophyllum micranthum Nutt., Trans. Amer. Phi- GH, MO, NY-n.v., US). los. Soc. (n.s.) 7: 453. 1841. Aphanopappus nut- Lipochaeta lavarum (Gaud.) DC. var. lanaiensis tallii Walp., Repert. Bot. Syst. 2: 620. 1843, nom. Sherff, Field Mus. Nat. Hist., Bot. Ser. 17: 582. illeg. Lipochaeta micrantha (Nutt.) A. Gray, Proc. 1939. TYPE: U.S.A. Hawaiian Islands, Lana'i, Amer. Acad. Arts 5: 131. 1861. Aphanopappus Maunalei Gulch, Sep 1917, C. N. Forbes 507.L micranthus (Nutt.) A. Heller, Minnesota Bot. (LECTOTYPE, designated by Gardner, 1979: F- Stud. 1: 915. 1897. TYPE: U.S.A. Hawaiian Is- 485197 [photo F]; ISOLECTOTYPES:BISH, F-2). lands, Kaua'i, ("Atooi"), Kolao [Koloa], 1835, T. Lipochaeta lavarum (Gaud.) DC. var. maneleana Nuttall s.n. (HOLOTYPE: BM-n.v.). Sherff, Field Mus. Nat. Hist., Bot. Ser. 17: 583. 1939. TYPE: U.S.A. Hawaiian Islands, Lana'i, Distribution.-U.S.A. Hawaiian Islands, 554 BRITTONIA [VUL.L-VOL. 53 known only from diverse mesic forest, ca. 10. Melanthera populifolia (Sherff) W. L. 300-400 m, Ha'upu Ridge, Hanapepe Val- Wagner & H. Rob., comb. et stat. nov. Koai'e Olokele and ley, Canyon, Canyon, Lipochaeta subcordata A. Gray var. populifolia Kaua'i. Melanthera micrantha is at risk of Sherff, Bot. Gaz. (Crawfordsville) 95: 91. 1933. extinction and has been federally listed as Lipochaeta populifolia (Sherff) R. C. Gardn., endangered. Rhodora 81: 328. 1979. TYPE:U.S.A. Hawaiian Islands, Lana'i, Maunalei Valley, 18 Jun 1918, G. C. Munro 670 (HOLOTYPE:F-659412 [photo F]; 8a. Melanthera micrantha (Nutt.) W. L. ISOTYPES:BISH-3, US [photo F]). Wagner & H. Rob. subsp. micrantha Distribution.-Hawaiian Islands, known Distribution.-U.S.A. Hawaiian Islands, only from the type collection Maunalei Val- known only from diverse mesic forest, ca. ley, Lana'i, and almost certainly extinct. 300-400 m, Koai'e and Olokele Canyons, and Hanapepe valley, Kaua'i. 11. Melanthera remyi (A. Gray) W. L. Wagner & H. Rob., comb. nov. L. 8b. Melanthera micrantha (Nutt.) W. Lipochaeta remyi A. Gray, Proc. Amer. Acad. Arts Wagner & H. Rob. subsp. exigua (0. 5: 131. 1861. TYPE: U.S.A. Hawaiian Islands, Deg. & Sherff) W. L. Wagner & H. Rob., O'ahu, s.l., 1851-1855, J. Remy 260 (HOLOTYPE: comb. et stat. nov. GH [fragment BISH-501044, photo F]).

Lipochaeta exigua 0. Deg. & Sherff in Sherff, Distribution.-U.S.A. Hawaiian Islands, Amer. J. Bot. 28: 30. 1941. Lipochaeta micrantha occurring in locally moist sites in dry forest (Nutt.) A. Gray var. exigua (0. Deg. & Sherff) R. to Leucaena shrubland, 30-180 m, north- C. Gardn., Rhodora 81: 325. 1979. TYPE:U.S.A. western Wai'anae Mountains, O'ahu. Hawaiian Islands, Kaua'i, Nawiliwili, 0.75 mi SW of Hokunui, on summit ridge, 8 Jan 1940, 0. Degener & E. Ordonez 12610 (LECTOTYPE, des- 12. Melanthera subcordata (A. Gray) W. ignated by Gardner, 1979: F-1014051; ISOLECTO- L. Wagner & H. Rob., comb. nov. TYPES:B-n.v., BISH, F-2, G-n.v., GH, MO- 2, NY-5 [microfiche], UC-3-n.v., US-2). Lipochaeta subcordata A. Gray, Proc. Amer. Acad. Arts 5: 130. 1861. TYPE: U.S.A. Hawaiian Is- Distribution.-U.S.A. Hawaiian Islands, lands, Hawai'i, on the coast, 1840, U.S. Expl. Ex- known from diverse mesic forest, ca. ped. y.n. (HOLOTYPE:US-57067). only Lipoclhaeta flexuosa Drake, Ill. Fl. Ins. Pacif., part 300-400 m, Ha'upu Ridge, Kaua'i. 4, 72, t. 35. 1888. TYPE:U.S.A. Hawaiian Islands, Hawai'i, s.l., 1851-1855, J. Remy 265 (tHl)IOTYPE: 9. Melanthera perdita (Sherff) W. L. Wag- P). ner & H. comb. nov. Lipochaeta intermedia 0. Deg. & Sherff in Sherff, Rob., Bot. Gaz. (Crawfordsville) 95: 102. 1933. TYPE: U.S.A. Hawaiian Hawai'i, Kona Lipochaeta perdita Sherff, Bot. Gaz. (Crawfords- Islands, Distr., 0. 4254 A. ville) 95: 99. 1933. TYPE:U.S.A. Hawaiian Is- Hu'ehu'e, May 1932, Degener (leg. F-668723 F, lands, [probably Ni'ihau], s.l., 26-29 Jan 1779, Meebold) (HOLOTYPE: [photo frag D. Nelson s.n. (HOLOTYPE:BM [photo F]). GI). Lipochaeta kawaihoaensis H. St. John, Pacific Sci. Distribution.-U.S.A. Hawaiian 13: 181. 1959. TYPE: U.S.A. Hawaiian Islands, Islands, Ni'ihau, Kawaihoa Point, head of steep gully, 300 scattered in dry forest and alien grassland, ft, 31 Mar 1949, H. St. John 23611 (HOLOTYPE: 550-1800 m, North Kona District and Pu'u BISH-500977; ISOTYPES:BISH-2, MO-n.v., Kanalopaka Nui, and South Kohala District, US-2). Hawai'i. Distribution.-U.S.A. Hawaiian Islands, 13. Melanthera tenuifolia W. L. known only from the types, listed above. (A. Gray) & H. comb. nov. Although habitats on Ni'ihau have been ex- Wagner Rob., tensively altered by ranching, it is not Lipochaeta tenuifblia A. Gray, Proc. Amer. Acad. known definitively whether this species is Arts 5: 131. 1861. TYPE: U.S.A. Hawaiian Is- extinct. Suitable habitats, including the type lands, O'ahu, Ka'ala Mtns., 1840, U.S. Expl. Ex- are on land and are not ac- ped. s.n. (LECTOTYPE,designated by Sherff, 1935: locality, private GH; ISOLECTOTYPES:NY-n.v., US). Sherff indi- cessible for surveys. The species is thus cated that Gray had two collections available, the presumed to be extinct. one he selected as the lectotype and Remy 276 2001]20011 WAGNER & ROBINSON: ASTERACEAE 555

(GH, P), with the material of both collections 15. Melanthera venosa (Sherff) W. L. mounted on the same sheet at GH. Gardner Wagner & H. Rob., comb. nov. (1979) superfluously lectotypified with the same specimen as Sherff. Lipochaeta venosa Sherff, Bot. Gaz. (Crawfords- ville) 95: 100. 1933. TYPE:U.S.A. Hawaiian Is- Distribution.-U.S.A. Hawaiian Islands, lands, Hawai'i, Waimea, Nohonaohae Crater, Jun rare in diverse mesic forest, 700-900 m, 1910, J. F. Rock 8349 (HOLOTYPE: F-659409 [pho- only in the central Wai'anae Mountains, to F]; ISOTYPES:BISH-4, GH, UC-n.v., US). O'ahu. Lipochaeta pinnatifida H. St. John, Pacific Sci. 38: 272. 1984. TYPE:U.S.A. Hawaiian Islands, Ha- wai'i, Waimea, several hundred meters N of Pu'u 14. Melanthera tenuis (0. Deg. & Sherff) Pa'a, near tank rd., 2400 ft, 1 Feb 1981, F. R. W. L. Wagner & H. Rob., comb. nov. Warshauer 3160 (HOLOTYPE:BISH). St. John con- sidered Warshauer 3160, which was collected to Lipochaeta tenuis 0. Deg. & Sherff in Sherff, Bot. demonstrate variability within a population of Gaz. (Crawfordsville) 95: 102. 1933. TYPE: Melanthera venosa (Warshauer, pers. comm. U.S.A. Hawaiian Islands, O'ahu, Wai'anae Val- 1984), a mixed collection, and used different parts ley, on lateral spur leading to summit ridge be- of it as the types of two taxa, the other sheet being tween Ka'ala and Kalena [up toward Mt. Ka'ala], used as the type of L. setosa. Holotype missing 24 Apr 1932, 0. Degener, K. K. Park & W. Bush 8/1987; not found in general collection at BISH 4258 (LECTOTYPE, designated by Gardner, 1979: in June 1997. F-666542 [photo F]; ISOLECTOTYPES:F [photo F], Lipochaeta setosa H. St. John, Pacific Sci. 38: 276. K-n.v., MO, NY [microfiche]). Sherff cited the 1984. TYPE: U.S.A. Hawaiian Islands, Hawai'i, collector only as O. Degener and gave no collec- Waimea, several hundred meters N of Pu'u Pa'a, tion number; MO sheet lists all three collectors near tank rd., 2400 ft, 1 Feb 1981, F. R. War- and gives a number, and was annotated by Sherff shauer 3160A (HOLOTYPE:BISH). St. John seg- "Cotype!" Both sheets at F have labels prepared regated this portion of a collection and added the by Sherff, with no collector number and only De- "A." He named the other part Lipochaeta pin- gener given as the collector. natifida. The two sheets collected were made to Lipochaeta dubia 0. Deg. & Sherff in Sherff, Field show the population variation of Melanthera ven- Mus. Nat. Hist., Bot. Ser. 17: 580. 1939. TYPE: osa (Warshauer, pers. comm., 1984). Holotype U.S.A. Hawaiian Islands, O'ahu, NE ridge [slope] missing 8/1987; not found in general collection of Pu'u Hapapa, 7 May 1939, 0. Degener, E. in Jun 1997. Ordonez, J. Foster 12331 (HOLOTYPE:F-1004397 Lipochaeta warshaueri H. St. John, Pacific Sci. 38: [photo F]; ISOTYPES:A-2-n.v., F-2, GB-n.v., 280. 1984. TYPE:U.S.A. Hawaiian Islands, Ha- GH-3, MO, NY-3 [microfiche], US-3). wai'i, Kohala Dist., Waimea Plain, near base of Sherff changed the number on the labels of the F Kohala slopes, gulch next to Waikamali Gulch, sheets from 12332 to 12331; NY sheets all have 1680 ft, 31 Jan 1981, F. R. Warshauer 3159 (HO- correct number of 12331 on Degener's original LOTYPE:BISH). Present disposition is based on label. the description and illustration as the holotype has Lipochaeta tenuis 0. Deg. & Sherff var. sellingii 0. been missing since Aug 1987. Deg. & Sherff, Fl. Hawaiiensis, fam. 244. Lipo- chaeta tenuis. 1940. TYPE:U.S.A. Hawaiian Is- Distribution.-U.S.A. Hawaiian Islands, lands, O'ahu, NE slope of Pu'u Happapa, near scattered in dry shrubland, 730-915 m, contour trail, 3 Sep 1938, 0. H. Selling 3336 (HO- known only from South Kohala District, LOTYPE:GB; ISOTYPE:BISH). Cited as a Selling and Heihei cin- collection, but labels on BISH and GB sheets Nohonaohae, Holoholoku, have all three collectors (Cranwell, Selling, and der cones, Pu'upa, and an unnamed cinder Skottsberg) listed. Gardner (1979) erroneously cone northeast of Nohonaohae, Hawai'i. cited Degener et al. 12253 as the type of this Melanthera venosa is at risk of extinction collected from the same name; although plant. and has been federally listed as endangered. Lipochaeta minuscula 0. Deg. & Sherff in Sherff, Bot. Leafl. 9: 9. 1954. TYPE:U.S.A. Hawaiian Is- lands, O'ahu, on ridge N of Kolekole Pass, 15 16. Melanthera waimeaensis (H. St. John) Jun 1947, 0. Degener 21660 (leg. M. Kerr) (HO- W. L. Wagner & H. Rob., comb. nov. LOTYPE:F [photo F]; ISOTYPE:BISH). Gardner (1979) superfluously lectotypified on BISH sheet, Lipochaeta waimeaensis H. St. John, Pacific Sci. 26: but Sherff clearly cited F sheet as the type. 293. 1972. TYPE: U.S.A. Hawaiian Islands, Kaua'i, Waimea Canyon, upper slope of W side, Distribution.-U.S.A. Hawaiian Islands, 1200 ft, 17 Apr 1967, R. W. Hobdy 101 (HOLO- uncommon in diverse mesic forest, on TYPE:BISH-501058; ISOTYPES:BISH, GH, US). or in sites such as St. John wrote "Holotype" on both BISH sheets, slopes, open streambeds, but illustrated 501058 and caption says it is from 700-900 m, only in the central Wai'anae the holotype. We therefore consider it to be the Mountains, O'ahu. holotype and the other sheet to be an isotype. 556 BRITTONIA [VOL. 53

Distribution.-U.S.A. Hawaiian Islands, TYPE: Africa. N Rhodesia [Zambia], known only from the type locality in di- Cruse 224 (HOLOTYPE: K-n.v.). verse mesic forest, ca. 350-400 m, on the Distribution.-Africa (Congo and Zam- upper slope, west side of Waimea Canyon, Kaua'i. Melanthera waimeaensis is at risk bia). of extinction and has been listed federally 19. MELANTHERAANGUSTIFOLIA A. Rich. in as endangered. Sagra, Hist. Isla 11: 54. 1850. TYPE: Cuba, La Sagra s.n. (LECTOTYPE, OTHER SPECIES OF MELANTHERA designated by Parks, 1973: P-n.v.). 17. MELANTHERAABYSSINICA (Sch.-Bip. ex Distribution.-U.S.A. (southern ), A. Rich.) Benth. & Hook. f., Gen. P1. 2: western Cuba, Hispaniola, Dominican Re- 377. 1873. Wuerschmittia abyssinica public, Mexico (Chiapas, Quintana Roo, Sch.-Bip. ex A. Rich., Tent. Fl. Abyss. 1: Tabasco, Vera Cruz, and Yucatan), Central 413. 1848. TYPE: Africa. , America (Guatemala, , and Pana- Schimper 334 (LECTOTYPE, designated ma). here: P-n.v.; ISOLECTOTYPES:BM-n.v., BR-n.v., K-n.v., M-n.v.). Two ad- 20. MELANTHERA ELEGANS C. D. Adams, J. ditional syntypes were cited in the pro- W. Afr. Sci. Assoc. 8: 135. 1964. TYPE: tologue. Africa. , C. D. Adams 12728 (HO- LOTYPE: K-n.v.). Distribution.-Africa (, Congo, Eritrea, Ethiopia, Kenya, , Su- Distribution.-Africa (Guinea). dan, , Togo, and Zambia). 21. MELANTHERAELLIPTICA O. Hoffm., in 18. MELANTHERA ALBINERVIA O. Hoffm., Engl., Bot. Syst. Jahrb. 24: 474. 1898. Bol. Soc. Brot. 13: 30. 1896. TYPE:Af- TYPE: Africa, Kling 47 (SYNTYPE: B [pre- 162 rica. Angola, Anchieta 119 (HOLOTYPE: sumably destroyed]); Kling (SYNTY- COI; ISOTYPE:LISU). PE: B [presumably destroyed]); Kling 163 (SYNTYPE: B [presumably destroyed]); Distribution. Africa (Angola, Congo, Kling 47 (SYNTYPE:B [presumably de- Kenya, Malawi, Mozambique, Namibia, stroyed]); Bittner 34 (B [presumably de- Transvaal, Tanzania, Uganda, Zimbabwe, stroyed]); Bittner 678 (B [presumably and Zambia). destroyed]).

18a. MELANTHERA ALBINERVIA O. Hoffm. Distribution.-Africa (Cameroon, Cen- tral African and subsp. ALBINERVIA Republic, Ghana, Nigeria). Distribution.-Africa (Angola, Malawi, 22. MELANTHERAFELICIS C. D. Adams, J. W. Mozambique, Namibia, Transvaal, Tanza- Afr. Sci. Assoc. 8: 135. 1964. TYPE:Af- nia, Zimbabwe, and Zambia). rica. Guinea, 24 Jan 1936, Jacques-Felix 1868 (HOLOTYPE:K-n.v.). 18b. MELANTHERA ALBINERVIA O. Hoffm. Distribution.-Africa (Guinea). subsp. ACUMINATA (S. Moore) Wild, Kir- kia 5: 12. 1965. Melanthera acuminata 23. MELANTHERAGAMBICA Hutch. & S. Moore, J. Linn. Soc., Bot. 35: 344. Dalz., Fl. W. Trop. Afr. 2, 1: 146. 1931. TYPE: 1902. TYPE: Africa. Kenya, Scott Elliot Africa. Gambia, Hayes 586 (HOLOTYPE: 7052 (HOLOTYPE: BM-n.v.; ISOTYPE: K-n.v.). K-n.v.). Distribution.-Africa (Gambia and Guin- Distribution.-Africa (Kenya, Uganda, ea-Bissau). and Tanzania). 18c. MELANTHERA ALBINERVIA O. Hoffm. 24. MELANTHERALATIFOLIA (Gardn.) Ca- subsp. CAUDATAWild, Kirkia 5: 12. 1965. brera, Darwiniana 16: 411. 1970. Echin- 2001]20011 WAGNER & ROBINSON: ASTERACEAE 557

ocephalum latifolium Gardn., London J. Wedelia prostrata Hemsl., in F Forbes & Hemsl., J. Bot. 7: 294. 1848. TYPE:Brazil. Ceara Linn. Soc., Bot. 23: 434. 1888, nom. nov. Ver- besina Hook. & Bot. in cane fields near Oct prostrata Arn., Beechey Prov., Crato, Voy. (5), 195. 1837, non L. (1753). Wollastonia 1838, G. Gardner 1728 (LECTOTYPE, des- prostrata Hook. & Am., Bot. Beechey Voy. (6), ignated here: BM; ISOLECTOTYPES: K 265. 1838, non DC. (1836). TYPE:, Macao [photo US], NY-2, US-frag). One ad- and adjacent islands, Apr 1827, Lay & Collie s.n. (HOLOTYPE:K-n.v.). ditional syntype was cited (G. Gardner 3848). (Fig. 2A-C) Distribution. Coastal areas from Bonin Echinocephalum lanceolatum Gardner, London J. Islands, , China, Japan (Honshu), Bot. 7:295. 1848, syn. nov. TYPE:Brazil, Ceara Ryukyu Islands, , and . Prov., near Aracaty, Aug 1838, G. Gardner 1729 Melanthera prostrata is often subdivided (HOLOTYPE:BM-n.v.; ISOTYPES:K-n.v. [photo into two varieties. Melanthera prostrata US], US-frag). var. rosbusta Makino is now to Echinocephalum angustifolium Gardner, London J. thought rep- Bot. 7:295. 1848, syn. nov. TYPE:Brazil, Goyaz resent a hybrid between M. prostrata and Prov., near Sape, Feb 1840, G. Gardner 3848 M. biflora (Peng et al., 1998), which we do (bis) (HOLOTYPE:BM-n.v.; ISOTYPES:K-n.v. not recognize formally with a binomial. [photo US], US-frag). Distribution.-South America, wide- 29. MELANTHERAPUNGENS Oliv. & Hiern, spread in the east from Brazil, , Fl. Trop. Afr. 3: 382. 1877. TYPE:Africa. and , 40-300(-1100) m, Sudan, Schweinfurth 1990 (HOLOTYPE: usually along rivers or in inundated savan- K-n.v.; ISOTYPE:BM-n.v.). na, sometimes in mesic forest. Distribution. Africa (Central African Republic, Congo, , Sudan, and 25. MELANTHERALIGULATA Small, Bull. Uganda). New York Bot. Gard. 3: 439. 1905. TYPE: U.S.A. Florida, Small & Wilson 1775 30. MELANTHERA RHOMBIFOLIAO. Hoffm. & (HOLOTYPE: NY-n.v.). Muschler, Mem. Soc. Bot. Fr. 2, 8: 117. 1910. TYPE: Africa. Chevalier 999 Distribution.-U.S.A (Florida). Mali, (HOLOTYPE: P--n.v.). 26. MELANTHERANIVEA (L.) Small, Fl. SE Melanthera tithonioides Assi, Adansonia 4: 338. U.S. 1251, 1340. 1903. Bidens nivea L. 1964. TYPE: Africa. Ivory Coast, Mangenot & Assi 1A 4142 Sp. P1. 2: 833. 1753. TYPE:Dillenius, J. (HOLOTYPE: P). 1732 Hort., Elth. tab. 47 (LECTOTYPE,des- Distribution.-Africa (Ivory Coast, Mali, ignated by Parks, 1973). (Fig. 1P-R) and Nigeria). Melanthera aspera (Jacq.) Small, Bull. Torrey Bot. Club 36: 164. 1909. Calea aspera Jacq., Collec- 31. MELANTHERARICHARDSIAE Wild, Kirkia tanea 2: 290. 1788. TYPE:Jacquin (HOLOTYPE:BM 5: 9. 1965 (as M. richardsae). TYPE: Af- [photo US]). rica. N Rhodesia [Zambia], Richards Distribution. and southeastern 1491 (HOLOTYPE: K--n.v.; ISOTYPES: United States to northern South America BR-n.v., E-n.v.). (, , Peru, and ), Distribution. Africa (Rwanda, Tanza- and the Greater and Lesser Antilles. nia, Uganda, and Zambia).

27. MELANTHERAPARVIFOLIA Small, Fl. SE 32. MELANTHERAROBINSONII Wild, Kirkia 5: U.S. 1251, 1370. 1903. TYPE:USA. Flor- 15. 1965. 1965. TYPE: Africa. N Rhode- ida, Blodgett s.n. (HOLOTYPE:NY-n.v.; sia [Zambia], Robinson 2099 (HOLOTYPE: ISOTYPE:GH-n.v.). SRGH-n.v.; ISOTYPE:K-n.v.). Distribution.-U.S.A. (southern Florida). Distribution.-Africa (Congo, Tanzania, and Zambia). 28. MELANTHERAPROSTRATA (Hemsl.) W. L. Wagner & H. Rob., comb. nov. 33. MELANTHERASCABERRIMA Hiern, Cat. \

558 BRITTONIA [VOL. 53

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FIG. 2. A-C. M. latifolia (Argentina, Entre Rios, Dpto. Concordia, Parque Rivadavia, Burkart et al. 25502) [note: ray flowers neutral in this species]. A. Pale. B. Disk achene. C. Cross section of disk achene. D-H. Wedelia calycina (Virgin Islands, St. Croix, Fosberg & Ogden 55325). D. Pale. E. Ray achene. F. Cross section of ray achene. G. Disk achene. H. Cross section of disk achene. I-M. Perymenium jelskii (Peru, Prov. Otuzco, Dpto. La Libertad, Chanchacap, Saga'stegui et al. 11669). I. Pale. J. Ray achene. K. Cross section of ray achene. L. Disk achene. M. Cross section of disk achene. 200112 WAGNER & ROBINSON: ASTERACEAE 559

Afr. P1. Welw. 1, 3: 582. 1898. TYPE:Af- Klatt, Bull. Herb. Boiss. 4: 839. 1896. rica. Angola, Welwitsch 3556 (HOLOTYPE: TYPE: Africa. Mozambique, Menyharth BM-n.v.; ISOTYPES:K-n.v., LISU- 735 (HOLOTYPE:GH-n.v.). n.v.). Distribution.-Africa (Angola, Botswa- Distribution. Africa (Angola). na, Mozambique, Namibia, Transvaal, Zambia, and Zimbabwe). 34. MELANTHERA SCANDENS(Schumach. & Bull. I.EA.N. 16: 68. Thonn.) Roberty, EXCLUDED NAMES 1954. Buphtalmum scandens Schumach. & Thonn., Beskr. Guin. P1. 392. 1827. 3: TYPE: Africa. Ghana, s.n. (Ho- Lipochaeta amazonica Poepp., Nov. Gen. Sp. 49. Thonning 1845. = Wedelia rudis Benth. ex H. Rob. LOTYPE: (Baker) C-n.v.). Lipochaeta costaricensis (Benth.) Benth. & Hook. f., 2: = Lasianthaea Distribution.-Widespread in Africa. Gen P1. 373. 1873. fruticosa (L.) K. Becker var. fasciculata (DC.) K. Becker. Lipochaeta asymetrica H. Leveille, Fedde Repert. 10: 34a. MELANTHERASCANDENS (Schumach. & 122. 1911. = Bidens asymetrica (H. Leveille) Sherff Thonn.) Roberty subsp. DREGEI (DC.) Lipochaeta fasciculata DC., Prodr. 5: 610. 1836. = Wild, Kirkia 5: 8. 1965. Psathurochaeta Lasianthaea fruticosa (L.) K. Becker var.fasciculata K. Becker. DC., Prodr. 5: 609. 1836. TYPE: (DC.) dregei Lipochaeta goyazensis Hook., Lond. J. Bot. 7: 406. Africa. [Republic of South Africa], Natal 1848. = Angelphytum goyazense (Hook.) H. Rob. & [KwaZulu-Natal], Drege s.n. (HOLOTYPE: W. L. Wagner, comb. nov. G-DC [microfiche]). Lipochaeta hastata Kellogg, Proc. Calif. Acad. Sci. 2: 106. 1863. = Verbesina hastata (Kellogg) Kellogg Distribution.-Africa (Mozambique, Re- ex Curran public of South Africa, and Swaziland). Lipochaeta lantanifolia Schauer, Linnaea 19: 729. 1847. = lantanifolia (Schauer) Strother (Hook. f.) A. Proc. Amer. MELANTHERASCANDENS & Lipochaeta laricifolia Gray, 34b. (Schumach. Acad. Arts 5: 131. 1862. = Trigonopterum larici- Thonn.) Roberty subsp. MADAGASCARIEN- folium (Hook. f.) W. L. Wagner & H. Rob., comb. SIS (Bak.) Wild, Kirkia 5: 7. 1965. TYPE: nov. Macraea laricifolia Hook. f., Trans. Linn. Soc. London 20: 210. 1847. In the of Africa. Madagascar, Baron 2344 p.p. process determining the present disposition of this taxon we discovered (K-n.v.). that Macraea Hook. f. (Trans. Linn. Soc. London Macraea J. Distribution.-Africa (Angola, Congo, 20: 209. 1847) was a later homonym of Lindl. (Quart. J. Sci. Lit. Arts ser. 2, 1: 104. 1828), Ethiopia, Kenya, Malawi, Mozambique, a synonym of Viviania Cav. in the Geraniaceae. The Sudan, Tanzania, Uganda, Zambia, and next and only available generic name is Trigonop- Madagascar). terum Steetz ex N. J. Andersson (Kongl. Vetensk. Acad. Handl. 1853: 183. 1854. TYPE: T. pontenii N. J. Andersson). We here make the correct com- 34c. MELANTHERASCANDENS (Schumach. & bination for the Galapagos Islands endemic (for ev- Thonn.) Roberty subsp. SCANDENS idence for generic recognition see Harling, 1962; Distribution. Africa Came- Eliasson, 1984). (Angola, Lipochaeta macrocephala Hook. & Arn., Bot. Beech- roon, Congo, Gabon, Ghana, Nigeria, and ey Voy. (10) 436. 1841. = Lasianthaea macroce- Sierra Leone). phala (Hook. & Arn.) K. Becker Lipochaeta monocephala DC., Prodr. 5: 610. 1836. Lasianthaea K. Becker var. 34d. MELANTHERASCANDENS (Schumach. & fruticosa (L.) fruticosa. scaberrima Benth., J. Bot. 11: 43.1840. SUBSIMPLICIFOLIALipochaeta Thonn.) Roberty subsp. scaberrima (Bentham) Blake Wild, Kirkia 5: 6. 1965. TYPE: Africa. Lipochaeta? serrata (Llave) DC., Prodr. 5: 611. 1836. Cameroon, Mildbraed 10752 (K-n.v.). = Zexmenia serrata Llave Lipochaeta strigosa DC., Prodr. 5: 610. 1836. = Wam- Distribution.-Africa (Cameroon, Con- alchitamia strigosa (DC.) Strother go, Ghana, Kenya, Malawi, Sierra Leone, Lipochaeta tagetiflora G. Don in Sweet, Hort. Brit. ed. Sudan, Tanzania, Uganda, Zimbabwe). 3: 360, nom. nud. Lipochaeta texana Torr. & A. Gray, Fl. N. Amer. 2: 357. 1842. = Kunth var. his- 35. MELANTHERA TRITERNATA (Klatt) Wild, pida (Kunth) Strother Kirkia 5: 9. 1965. Wedelia triternata Lipochaeta umbellata DC., Prodr. 5: 610. 1836. = Las- 560 BRITTONIA [VOL. 53

ianthaea ceanothifolia (Willd.) K. Becker var. cean- tematic implications in Lipochaeta (Compositae: othifolia Heliantheae). Amer. J. Bot. 64: 810-813. Lipochaeta umbellata DC. var. conferta DC., Prodr. 5: . 1977b. Observations on tetramerous disc flo- 610. 1836. = Lasianthaea ceanothifolia (Willd.) K. rets in the Compositae. Rhodora 79: 139-146. Becker var. ceanothifolia . 1979. Revision of Lipochaeta (Compositae: Heliantheae) of the Hawaiian Islands. Rhodora 81: 291-343. Acknowledgments & J. C. La Duke. 1978. Phyletic and cladistic relationships in Lipochaeta (Compositae). Syst. We are most grateful to Alice Tangerini Bot. 3: 197-207. for her superb pencil illustrations of the Gill, L. S. & D. I. Omoigui. 1992. Chromosome num- achenes and paleae presented in Figures 1 bers in some Nigerian Compositae. Comp. Newsl. and 2. We the curators at 20/21: 12-15. acknowledge G. 1962. On some endemic to and NY for for Harling, Compositae BISH, F, MO, loaning types the Galapagos Islands. Acta Horti Bergiani 20: 63- study and for helping with various queries 120. about their collections. We thank Derral Heller, A. A. 1897. Observations on the ferns and Herbst for help with information of some flowering plants of the Hawaiian Islands. Minne- collections at BISH. We the sota Bot. Stud. 1: 760-922. appreciate Lauener, L. A. 1980. Faurie's Hawaiian types at the helpful comments by reviewers Gerry Carr, British Museum. Notes Roy. Bot. Gard. Edinburgh Jose Panero, and Tod Stuessy, which im- 38: 495-497. proved the paper in numerous ways. Lessing, C. F. 1831. Synanthereae rich. In: A. de Chamisso & D. de Schlechtendal, editors. De plan- tis in Literature Cited expeditione Romanzoffiana. Linnaea 6: 83- 170, 209-258 (addenda), 510-526 (continuatio). A. & P. M. Mathew. 1978. In IOPB chro- K. M. 1979. A of the genus Las- Mathew, Becker, monograph mosome LX. Taxon 27: 223-231. ianthaea (Asteraceae). Mem. New York Bot. Gard. reports 1-64. Nicolson, D. H. & F. R. Fosberg. In press. Forsters 31(2): and of the second Cook Bentham, G. 1873. In: Bentham. G. and botany expedition (1772- Compositae. Veg. J. D. Hooker, Genera Plantarum. 2(1): 163-533. 1775). Reg. M. 1975. Chromosome numbers of some endem- Cabrera, A. L. 1970. Novedades sinanterologicas en- Ono, trerrianas. Darwiniana 16: 409-411. ic species of the Bonin Islands I. Bot. Mag. Tokyo 88: 323-328. 1974. 158. Compositae. Flora Ilustrada de En- . 1977. studies on the tre Rios (Argentina) 6: 106-538. Cytotaxonomic flowering endemic of the Bonin Islands I. Candolle, A. P. de. 1836. Compositae. Prodr. 5: 4- plants species 706. Mem. Nat. Sci. Mus. 10: 63-80. R. K. & A. Carr, G. 1978. Chromosome numbers of Hawaiian Panero, J. L., Jansen J. Clevinger. 1999. of subtribe flowering plants and the significance of cytology in Phylogenetic relationships Ecliptinae selected taxa. Amer. J. Bot. 65: 236-242. (Asteraceae: Heliantheae) based on chloroplast DNA restriction site data. Amer. J. Bot. 86: 413- Chumley, T. W., J. L. Panero, S. C. Keeley & R. 427. K. Jansen. 2000. A phylogeny of the Ecliptinae (Asteraceae: Heliantheae) as inferred from internal Parks, J. C. 1973. A revision of North American and Caribbean Melanthera Rhodora transcriber spacer (ITS) sequences, and the origin (Compositae). of Lipochaeta. Suppl. Amer. J. Bot. 87(6): 119. 75(802): 169-210. K.-F. Decaisne, J. 1834. Herbarii timorensis descripto. Peng, C.-I., Chung & H.-L. Li. 1998. 144. Nouv. Ann. Mus. Hist. Nat. 3: 333-501. Compositae. Flora of Taiwan, Ed. 2, 4: 807-1102. Eliasson, U. H. 1984. Chromosome number of Ma- Rabakonandrianina, E. 1980. Infrageneric relation- craea laricifolia Hook. fil. (Compositae) and its ships and the origin of the Hawaiian endemic genus bearing on the taxonomic affinity of the genus. Bot. Lipochaeta (Compositae). Pacific Sci. 34: 29-39. J. Linn. Soc. 88: 253-256. & G. D. Carr. 1981. Intergeneric hybridiza- Fay, J. J. 1979. Revision of Perymenium (Asteraceae- tion, induced polyploidy, and the origin of the Ha- Heliantheae) in Mexico and Central America. Al- waiian endemic Lipochaeta from Wedelia (Com- lertonia 1: 235-296. positae). Amer. J. Bot. 68: 206-215. Fosberg, F. R. 1993. The Forster Pacific islands col- Robinson, H. 1981. A revision of the tribal and sub- lections from Captain Cook's resolution voyage. tribal limits of the Heliantheae (Asteraceae). Smith- Allertonia 7: 41-86. sonian Contr. Bot. 51: 1-102. & M.-H. Sachet. 1980. Systematic studies of , A. M. Powell, R. M. King & J. F. Weedin. Micronesian plants. Smithsonian Contr. Bot. 45: 1- 1981. Chromosome numbers in Compositae, XII: 40. Heliantheae. Smithsonian Contr. Bot. 52: 1-28. Gardner, R. C. 1976. Evolution and adaptive radia- Rohr, J. von. 1792. Plantae-Slaegter poa St. Croix. tion in Lipochaeta (Compositae) of the Hawaiian med tilfoiede Anmaerkninger af Vahl. Skriv. Nat. Islands. Syst. Bot. 1: 383-391. Selsk (Kiobenhaven) 2: 205-227. . 1977a. Chromosome numbers and their sys- St. John, H. 1954. Review of Mrs. Sinclair's "Indig- 20011I WAGNER & ROBINSON: ASTERACEAE 561

enous flowers of the Hawaiian Islands." Hawaiian Zexmenia, and (Compositae-Heliantheae- Plant Studies 23. Pacific Sci 8: 140-146. Ecliptinae). Syst. Bot. Monogr. 33: 1-111. . 1984. Novelties in Lipochaeta (Compositae). Stuessy, T. 1977. Heliantheae: systematic review. Pag- Hawaiian Plant Studies 119. Pacific Sci. 38: 253- es 673-697 In: V. H. Heywood, J. B. Harborne & 282. B. L. Turner, editors. The biology and chemistry of the Academic London. Sherff, E. E. 1935. Revision of Tetramolopium, Li- Compositae. Press, W. L. & R. K. Shannon. 1999. Dubautia, and Railliardia. Bernice P. Wagner, Angiosperm pochaeta, from the Hawaiian collections of A. A. Hell- Mus. Bull. 135: 1-136. types Bishop er. Brittonia 51: 422-438. A. C. & G. Carr. 1991. Asteraceae. In. A. C. Smith, , D. R. Herbst & S. H. Sohmer. 1990. Manual Flora Vitiensis Nova: a new flora of Smith, of the flowering plants of Hawai'i. Bishop Museum 5: 254-320. National (Spermatophytes only) Trop- Special Publication 83. University of Press ical Botanical Garden, Lawai, Hawaii. and Bishop Museum Press, Honolulu. Strother, J. L. 1991. of Complaya, Ela- Wild, H. 1965. The African species of the genus Me- phandra, logeton, Jefea, Wamalchitamia, Wedelia, lanthera Rohr. Kirkia 5: 1-17.