PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 115(4):909-922. 2002.

Toward a phylogenetic subfamilial classification for the Compositae ()

Jose L. Panero and Vicki A. Funk

(JLP) Section of Integrative Biology, The University of Texas, Austin, Texas 78712 (VAF) U.S. National Herbarium, NMNH, Smithsonian Institution MRC 166, Washington, D.C. 20013-7012

Abstract. —Molecular studies of the flowering family Compositae (As- teraceae) based on comparative DNA sequence data of chloroplast genes pro- vide new insights into the evolution and radiation of the family. The results support the creation of new groups to maintain a classification that reflects evolutionary history. We are proposing the following new names: subfamilies Corymbioideae, , Gymnarrhenoideae, Hecastocleoideae, and Pertyoideae; tribes Athroismeae, Corymbieae, Dicomeae, Gochnatieae, Gym- narrheneae, Hecastocleideae, Polymnieae; subtribe Rojasianthinae. The totals

now stand at 1 1 subfamilies and 35 tribes. Only one tribe, the , is non-monophyletic having two branches. Most of the new groups are derived from taxa included in tribe Mutisieae, long suspected to be a paraphyletic group. Molecular studies that support these changes are discussed elsewhere

but a summary of their results is presented.

Molecular phylogenetic studies of the of the Compositae using sequence data of family Compositae (Asteraceae) have been the chloroplast gene ndhF and provide ad- instrumental in providing important insight ditional evidence for the paraphyly of sub- into the evolution and radiation of the fam- family . Their data provided ily. Assessments of phylogenetic relation- some support for the recognition of three ships in the family using molecular data be- main groups within subfamily Cichorioi- gan with restriction fragment length poly- deae s.l. splitting sequentially from above morphisms of the chloroplast genome. the Barnadesioideae: 1) Mutisieae, followed These studies identified a 22 kb inversion by 2) Cardueae, and finally, 3) the tribes present in all members of the Compositae Cichorieae, Liabeae, , and Arc- except the Mutisieae subtribe Barnadesieae toteae forming a monophyletic group sister (Jansen & Palmer 1987), which had a chlo- to . The latter group plus Aster- roplast organization similar to the rest of oideae was recognized by Bremer (1994, land . This result was used to elevate 1996) as the vernonioid group. Since the the Barnadesieae to subfamily status (Bre- publication of the molecular phylogeny of mer & Jansen 1992), and the traditional two the Compositae by Kim & Jansen (1995) subfamily system consisting of Cichorioi- only the study by Bayer & Starr (1998) has deae and Asteroideae was expanded to in- provided a competing hypothesis of tribal clude the Barnadesioideae. Kim & Jansen relationship based on noncoding regions of (1995) published what can arguably be the chloroplast genome, but this study did viewed as one of the most important papers not provide good support for any new pat- using DNA sequence data aimed at eluci- terns of relationship. Most studies using the dating the phylogeny of the Compositae. In ndhF data matrix to date have concentrated this paper they identified the major clades on elucidating the placement of a particular 910 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON taxon in the of life of daisies (Karis et argue that eventually they might be part of al. 2001). the Asteroideae and Cichorioideae respec- The goal of our study was to use several tively, but given the amount of data that molecular markers to produce a phylogeny support this cladogram and the fact that the that would clarify the phylogenetic relation- isolated position of these genera has long ships of members of the tribe Mutisieae s.l., been recognized, it is doubtful. Details of and the circumscription of the subfamily the analyses and a discussion of the result- Cichorioideae in general. Our data matrix ing clades will be published elsewhere includes the genes and markers ndhF, trnh- (Funk & Panero, in prep.; Panero et al., in trnF, matK, ndhD, rbcl^, rpoB, rpoCl prep.); we publish these names here so that exonl, 23S-trnl, and ndhl for an approxi- they are available for use in those papers. mate total data matrix of 13,380 bp. Com- The changes to the classification of the bining these chloroplast markers has pro- family proposed herein have been carefully duced a solid phylogeny of the Compositae. reviewed by the authors with the under-

Included here is a strict consensus tree of standing that these changes represent the the relationships of Compositae based on largest overhaul to the classification of the our studies for the Cichorioideae (Panero & Compositae since Bentham (1873). Our Funk, in prep.) and those of Panero et al. confidence in proposing these changes is

(in prep.) for tribe and relatives based on two things, first, an analysis of

(Fig. 1). This phylogeny includes a majority one of the largest data matrices for any of genera of the classical Mutisieae and rep- group of flowering plants produced to resentatives of most other tribes of Com- date—a combination of multiple genetic positae as well as genera deemed, based on markers and a broad sampling of taxa—and their morphology, to be of difficult place- second, an analysis of these data by con- ment (see Bremer 1994). The resulting phy- servative methods (i.e., maximum parsi- logeny has clarified substantially the rela- mony) without the coding of indels or char- tionships of the Compositae and resulted in acter weighting of any type. Bootstrap sup- the need to publish new names to maintain port for nearly all of the clades being rec- a formal taxonomic nomenclature for ognized as new taxa is 100%. Our monophyletic clades. All groups are now experience with this large data matrix monophyletic and reflect the evolutionary shows that additions of new data partitions history of the family except for two branch- only solidifies the phylogenetic signal and es near the base of the tree that constitute branch support. It is clear that past a certain a paraphyletic . It is most like- threshold, unique to each dataset and deter- ly that additional data will provide enough mined by the number of characters and taxa support to recognize the "Stifftia group" as sampled, the phylogenetic signal stabilizes. a new subfamily and tribe, however the It is our view that the addition of new data data are missing several of the genera that partitions in the future will only solidify the should fall into the "Stifftia group" clade branch support reported here. and final resolution of the problem will have to wait until those genera have been Subfamilies collected and analyzed. Two of the mono- Corymbioideae Panero & V. A. Funk, sub- phyletic subfamilies proposed, Corymbioi- fam. no v., Corymbieae Panero & V. A. deae and Gymnarrhenoideae, contain gen- Funk, trib. nov. era whose placement has long been prob- Type: L., Coroll. Gen. 14 lematic. Their isolated placement on the (1737). cladogram combined with their unusual Monotypic (7 species). morphology supports their being named at the subfamily (and tribal) level. One might Plantae in foliis sessilibus et in laminis VOLUME 115, NUMBER 4 911

Tribes Subfamilies

Heliantheae |

100 58 Madieae Perityleae

I 68 51 Eupatorieae 72 Millerieae 95 Polymnieae

981 Bahieae Chaenactideae 98 Tageteae

100 Neurolaeneae Asteroideae Coreopsideae 100 Helenieae

100 Athroismeae Plucheeae 100 1 54 Anthemideae Astereae 60 Gnaphalieae 100 ^ Calenduleae Corymbieae Corymbioideae 79 Arctoteae Vernonieae 90 Liabeae Cichorioideae 100 Cichorieae 100 Gundelieae Gymnarrheneae Gymnarrhenoideae

100 Pertyeae Pertyoideae Cardueae 100 100 Dicomeae 53 E Hecastocleideae Hecastocieioideae

Gochnatieae Gochnatioideae

100 Mutisieae (SA) Mutisioideae

iqoj Mutisieae (SA, A) "The Stifftia Group" Bamadesieae Bamadesioideae t ^^^^^—^^^^—^^-^^^^^^ Outgroups (Calyceraceae, Goodeniaceae)

Fig. 1. Strict consensus tree of 12 most parsimonious for 122 representative Compositae taxa based on chloroplast DNA sequence data. Bootstrap values indicated above nodes, SA = South America, A = Asia. Tree is simplified to show only tribe and subfamily placement and arrangement. 912 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON conduplicatis. Capitula uniflora; bracteis in- obtuse or acute, with short sweeping hairs volucri biseriatis, dimorphis; bracteis inter- on abaxial side, sweeping hairs restricted to ioribus 2, involutis; limbis corollarums sal- style and scarcely present below bifurcation verformibus. point on the style shaft. Achenes linear to Scapose, perennial herbs, tufted and elliptic, hirsute or villous, silvery white, forming rosettes. alternate, sessile, pappus of short, crown-like scales variously sometimes conduplicate, blades lanceolate fused or divided into lobes and bristles. to linear, bases attenuate, leaf apices Chromosome number: 2N = 16 (Weitz acuminate, terete, involute or flat in cross 1989). section, margins entire, venation parallel, The phylogenetic position of the South leaf surfaces glabrous to villose or hirsute, African Corymbium has been histor- sometimes with glandular trichomes. Inflo- ically controversial (Bentham 1873) and in rescence of open to congested corymbiform the latest revision of the family at the ge- cymes at the end of scapes that are as tall neric level, the genus is placed in subfamily or taller than the rosette leaves. Heads with Cichorioideae unassigned to a tribe (Bremer one floret, involucre narrowly cylindrical, 1994). Corymbium is unusual in the Com- conspicuously gradated, phyllaries in two positae because of its single-flowered series, chartaceous-coriaceous to herba- heads, heads with two innermost involucral ceous, green with purple tips or suffused bracts enclosing the flower, the broad, pat- with purple when young, outer series of 2—3 ent lobes of the corolla, and the narrowly phyllaries, apex acute to acuminate or lanceolate leaves with parallel venation. rounded, glabrous or pubescent inner series Our studies reveal that the genus occupies of 2 phyllaries, these approximately 2 to 5 a position next to the base of the subfamily times longer than those of outer series, oval Asteroideae. These results together with the to oblong in outline, involute, enclosing flo- distinctive morphology of Corymbium war- ret, bifid or trifid, glabrous or pubescent; rant its inclusion in a new subfamily Cor- receptacle flat. Florets actinomorphic and ymbioideae. bisexual, salverform, corolla mauve, pink, or white, lobes longer than tube with a tuft Gymnarrhenoideae Panero & V. A. Funk, of papillae on the distal end, apiculate at subfam. nov., Gymnarrheneae Panero & apex raised, triangular scler- with a arch of V. A. Funk, trib. nov. ified cells immediately below the lobe apex Type: Desf., Memoires du rendering the tip of the apex apiculate; an- museum d'histoire naturelle, Paris. 4:1. t. thers 5, thecae black, ecaudate, shallowly 1. 1818. calcarate, anther connective of long, fusi- Monotypic (1 species). form, conspicuously sclerified cells, endo- thecium dimorphic, proximal or inner rows Herbae perennes rosettiformes. Synflo- composed of quadratic cells with bipolar rescentiae simplices vel complexae; capi- thickenings and single bridges, these cells tulis subterraneis et aeriae; achaeniis di- separated from distal or outer endothecial morphis. cells by a single cell row of tightly scleri- Perennial amphicarpic herbs. Leaves fied isodiametric cells, distal endothecial simple, forming dense rosettes, sessile, cell rows composed of cells approximately smooth, narrowly lanceolate to nanowly twice as large as those of the inner endo- ovate, apex narrowly acute to attenuate, thecium with bipolar thickenings and single margins denticulate, base truncate, surfaces bridges, anther appendage lacking and re- smooth and glabrous. subter- duced to a series of elongated cells arising ranean or congested in the center of the from the anther connective; styles dissected, leafy rosette. Subterranean heads homoga- color as in corollas, style branches linear. mous and female, corolla vestisial enclosed VOLUME 1 15, NUMBER 4 913 in involucral bracts. Achenes of subterra- nean fruits develop first and the aerial fruits nean heads laterally flattened, blackish, are produced later and only in wet years. sparsely hairy; pappus absent or of short The aerial fruits are wind dispersed whereas bristles that are flattened at the base ap- the subterranean fruits germinate under- proaching scale-like, but bristle-like at the ground and appear to be more drought tol- apex with rounded apices. Aerial heads erant. It is possible that amphicarpy in bearing florets of two types: male florets in Gymnarrhena and other plants is an adap- small groups that are loosely connected on tive mechanism developed in response to very short peduncles and interspersed inconsistent and/or low water-availability among single female florets each of which (Roller & Roth 1964). is strongly enclosed in a stiff white and Gymnarrhena is an unusual member of green bract, with a few rows of chartaceous the Compositae and its history reflects its phyllaries. Florets with whitish corollas, unique morphology. Bentham (1873), Hoff- male florets greatly reduced in size with mann (1894), and Cronquist (1955) placed 3^ lobes and anthers, anther apex abbre- this genus in the Inuleae s.l. and Bentham's viated, filament not elongated into an apical alternate choice was the Astereae. Skvarla appendage, thecae with base short rounded, et al. (1977) pointed out that the did ecalcarate, ecaudate; female florets filiform, not support either of the above placements style branches long with rounded apices. as the genus has Anthemoid pollen and they Achenes obconical, ciliate, with elongate suggested a possible placement in the An- twin hairs, cell walls appear to be thin; pap- themideae. Bremer (1994) listed the genus pus of long, acuminate, laciniate bristles as belonging to the Cichorioideae s.l. but as with acute apices; ovaries of male florets "unassigned to tribe" along with several with a vestigial pappus or lacking. Chro- other problem genera. In our cladograms mosome number: 2N = 20. Gymnarrhena is consistently located by it- A monotypic genus extending from self below the Cichorioideae s.s. and above North Africa to the Middle East. the Pertyoideae ("Asian Mutisieae") clade. Amphicarpy (the ability to produce aerial as well as subterranean fruits) is uncommon Gochnatioideae (Benth. & Hook, f.) Panero in flowering plants being found in only ca. & V. A. Funk, subfam. stat. nov., Goch- 50 species worldwide (Lev-Yadum 2000). natieae (Benth. & Hook, f.) Panero & V. are available that that pre- Data show a high A. Funk, trib. stat. nov. Basionym: ponderance (eight) of those taxa are found Gochnatiinae Benth. & Hook, f.. Gen. PI. in Israel. 168, 216. 7-9 Apr 1873. This taxon has repeatedly been described Type Kunth in HBK, Nov. Gen. as having male central florets and periph- et Sp. 4:19. t. 309. 1820. eral female florets, but the male florets are Includes: Griseb.(2 species), not always grouped in the center of the Gill ex D. Don (1 species), head. In fact groups of florets some male Gochnatia Kunth in HBK (ca. 60 spe- are found at the outer edge of the recepta- cies), Richterago Kuntze (including Ac- cle. It seems more likely that the aerial tinoseris (Endl.) Cabrera, 17 species), heads are actually groups of heads, of some and possibly Chucoa Cabrera (1 species). which are few-flowered male heads and others are single-flowered female heads. A Mutisieis in stylis glabris apice rotun- Thus the monoecious plants bear heads of datis, in appendices antherarum apiculatis et both sexes grouped in a common synflo- in bracteis involucri multiseriatis differt. rescence. The fruits of the two types of Many perennial herbs but also and heads of Gymnarrhena have different de- trees or rcirely dioecious subshrubs or gy- velopmental patterns: the larger subterra- nodioecious shrubs or monoecious rosulate 914 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

herbs; branches rarely spiny. Leaves sim- fined to the Americas. The clade has con- ple, alternate, coriaceous or smooth, lance- sistently appeared in all versions of the olate, oblanceolate, elliptic, ovate, obovate, analysis. The genera in this subfamily are spathulate to narrowly linear; apex acute, characterized by glabrous styles and anthers obtuse, mucronulate to acuminate, and spi- with apiculate appendages. The genus Chu- nose; margins entire, spinose-dentate, den- coa referred by Cabrera (1955, 1977) to the tate to denticulate; base attenuate, truncate, Mutisieae subtribe Gochnatiinae may be- or rounded; surfaces tomentose, glandular long in this group, however, we were un- punctuate, glabrescent or glabrous; subses- able to obtain recently collected material for sile to petiolate, base expanded lat- analysis and the genus differs from other erally, sometimes pubescent; veins pinnate genera of the Gochnatioideae by having pu- or parallel. solitary and ter- bescent style branches. Final placement of minal or few in number to numerous in ra- this genus will have to await further study. cemes or cymes. Heads pedunculate or rare- ly sessile; peduncle bracteate; cylindrical or Hecastocleioideae Panero & Funk, subfam. campanulate; receptacle epaleate, rarely pa- nov, Hecastocleideae Panero & V. A. leate, flat to shallowly convex; discoid and Funk, trib. nov. homogamous or radiate and heterogamous; Type: A. Gray, Proc. Amer. phyllaries in 4—15 series, imbricate, outer Acad. Arts 17:220. 1882. ones ovate-lanceolate to lanceolate, inner Monotypic (1 species) ones subequal to longer than the outer ones, linear-lanceolate to lanceolate, acute, acu- Ad Mutisieas similes sed in synflorescen-

minate, lanate, sericeous or glabrescent. tiis biordinatis in capitulis solitariis in brac- Florets few-many (5—300); corolla tubes teis spiniferis et in corollas profunde lobatis glabrous, pubescent, and/or glandular. Mar- differt. ginal bilabiate florets (when present) pistil- Stoloniferous shrubs with deep roots, late, (3+1, 3 + 2, 4+1), 10-60, white to growing on vertical cliffs in the cracks of pinkish with or without rose-colored veins; rocks or in very well-drained, sandy soil, staminodia 4-5, free, hyaline (in gynodioe- young stems bright green turning white- cious plants flowers of male plants are her- brown with age, glabrous, old stems ash maphroditic and female plants have flowers gray to brown, trichomes simple, white, re- without anthers). Disc florets hermaphro- stricted to leaf internodes of rapidly grow- ditic or functionally male, actinomorphic ing shoots. Leaves alternate, sessile, blades with 5 deeply divided lobes, yellow, white, narrowly ovate to broadly acicular, leaf ba- rose, or orange-yellow; anther-appendages ses attenuate, leaf apices ending in a ligni- acuminate to apiculate, anther-tails glabrous fied mucro or spine; margins entire with or pubescent; style slender, shortly bilobed, 2-3 spines on each side of the leaf, spines glabrous, apex rounded or obtuse. Achenes mostly confined to basal half; central vein cylindrical, inconspicuously ribbed, villose, lignified with age and turning into a soft sericeous, sometimes glandular; pappus un- spine as leaf blade falls off at senescence; iseriate or biseriate, bristles 25-50, simple venation triplinerved with the two second- (rarely branched), basally free or connate ary veins parallel to the main vein, unarmed into a fleshy ring; setae serrulate, straw-col- leaves produced from short lateral branches ored. Chromosome numbers: Cyclolepis 2N if growing conditions are favorable, leaf ab- = 54, Cnicothamnus 2N = 44, and Goch- axial and adaxial surfaces concolorous and natia 2N = 46. essentially glabrous. Inflorescence of mul- This clade is located above the Mutisieae tiple single-flowered capitula aggregated

s.s. clade and below Hecastocleis and the into compound heads, each aggregation of Carduoideae clade. Its members are con- 1-5 heads associated with a scarious to co- VOLUME 115, NUMBER 4 915 riaceous oval to ovate bract with 3-9 spines connective unevenly thickened and there- on each side, the terminal inflorescence fore producing a "radial" thickening pat- with 4-10 capitula surrounded by 4-5 tern under light microscopy conditions, this bracts. Heads with 1 floret, involucre of 20- pattern gradually changing to polar thick- 30 phyllaries in 4-5 series, phyllaries nar- ened cells with those cells ending the en- rowly ovate, prominently apiculate, herba- dothecium having minimal thickened areas, ceous to scarious, green, densely strigose to pollen tricolpate, smooth. Style glabrous lanose on mid area to basal attachment; re- with no sweeping hairs, dissected into two, ceptacle flat. Zygomorphic or liguliform rounded style branches, each style branch florets absent, actinomorphic florets perfect, ending with a laminar cell proliferation. corollas purple-white changing to white Achenes oval, black, essentially glabrous, with age, deeply lobed, lobes narrowly tri- pappus of a crown of scales, minutely lac- angular and as long as tube, patent at an- erate. Chromosome number: 2N = 16 thesis, tube obtrullate in outline, druses (Powell et al. 1974). present, veins 5, parallel to each other along Hecastocleis was described by Asa Gray the tube and splitting below the sinus of the (1882) and in the protologue of the species lobe, the fiber bundle splitting unequally he remarked on the distinctiveness of the just above the sinus level, producing two genus and of its affinities to the Asian ge- strong veins and a thin vein that dissipates nus . This commentary is proba- in the space between the two main veins bly based on the similarity in corolla mor- and the corolla lobe edge, the two main phology between the Pertyoideae and He- veins diverge from each other and reach the castocleis in which corollas are deeply dis- apex of the lobe where they dissipate into sected and the lobes are spreading. Our the surrounding tissue, the adaxial corolla molecular studies reveal that Hecastocleis lobe strongly sclerified, sclerified cell pro- represents an isolated lineage, possibly the liferation coincides with splitting of pri- survivor of an old radiation of the family mary veins, the sclerified cells appear in the into the North American continent. middle lower middle portion of the lobe and radiate to the edges to approximately cover Pertyoideae Panero & V. A. Funk, subfam. of the lobe surface, additional layers of fj nov., Pertyeae Panero & V. A. Funk, trib. sclerified cells repeat the same overall dis- nov. tribution pattern as the lower layers but ap- Type: Pertya Schultz-Bip., Bonplandia, 10: proximately halfway the length of the lobe. 109. t. 10. 1862 Anthers 5, calcarate and caudate, purplish Includes: Ainsliaea DC. (50 species), Dias- to pale pink, anther collar with isodiamet- pananthus Miq. (1 species, sometimes ric, thickly and evenly sclerified cells, these placed in synonomy with Ainsliaea), changing abruptly to fusiform narrowly Macroclinidium Maxim. (3 species), My- cells with strongly sclerified walls, the an- ripnois Bunge (1 species), and Pertya ther collar uneven in composition with nu- Schultz-Bip. (15 species). merous sclerified cells abaxially and a band of one to few cells on its adaxial side, an- Ad Mutisieas similes sed in capitulis mi- ther tails approximately % the total length noribus solitariis vel paucis in corollis non of the anther thecae with endothecium tis- bilabiatis; lobis 5 profunde dissectis varia- sue extending approximately '/j to '/, the biliter connatis distinctae. length of the tail, the distalmost end of the Many perennial herbs but also some tail twisted in some anthers, the strongly shrubs or vines and one (Myripnois) gyno- sclerified tissue of the connective gradually dioecious ; glabrous to densely pubes- expands ending in a strongly sclerified an- cent. Leaves simple, scattered and alternate, ther appendage, those cells proximal to the or congested and rosulate at midpoint or 916 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON basal, subsessile to petiolate, lanceolate, ob- duoideae). This is the only group of Mutis- lanceolate, elliptic, ovate, obovate, spathu- ieae s.l. taxa that is located above the Car- late to narrowly-linear or cordate; apex duoideae on the tree and its placement acute, rounded, acuminate, or mucronulate; shows the diversity and variability of the margins entire, pubescent, dentate to den- taxa previously contained in the Mutisieae ticulate; base attenuate or rounded; surfaces s.l. from densely tomentose, to glabrous; peti- The most interesting feature of this group ole sometimes narrowly winged; blades is the deeply but unequally divided actino- mostly with 3 or 5 veins but a few pinnate. morphic corolla. Some corollas appear to be Inflorescence solitary or heads few in num- actinomorphic, some bilabiate and some are ber in spike-like or racemiform to thyrsi- intermediate between the two. All corollas form cymes. Heads discoid or disciform of this group have five equal sized lobes and homogamous, pedunculate or sessile; with 10 veins. The variation occurs in the peduncle bracteate sometimes minutely so; fusion of these lobes and it happens at all involucre cylindrical or campanulate, phyl- levels. Even in the same head where some laries in 3-8 series, imbricate, outer ones corollas have a 3+2 division and others ovate-lanceolate to lanceolate, inner ones have a regular corolla while still others linear-lanceolate to lanceolate, apex acute have 3+1 + 1 or even 3 + 1 . In some species or acuminate, minutely pubescent or more of Ainsliaea the corollas have nearly evenly rarely glabrous. Florets one to few, perfect, divided lobes with all of the lobes skewed corollas actinomorphic with 5 equal sized, to one side or secund which gives a star- deeply divided lobes fused unequally tling appearance to the heads. (sometimes in the same head) with some fulfilling the criteria for bilabiate but always Tribes with 10 veins (most common patterns 3 + 2, Dicomeae Panero & V. A. Funk, trib. nov. 3 + 1 + 1), sometimes secund; white, white Type: Dicoma Cass., Bull. Soc. Philom. 12. suffused with pink, purple, or red, anthers 1817. calcarate, caudate, anther appendages small, Includes: Dicoma Cass. (ca. 40 species), acuminate to acute, anther-tails usually pi- Benth. (13 species), lose and sometimes very long; style slender, Gladiopappus Humbert (1 species), Ma- shortly bilobed, dorsally with short hairs or cledium (Cass.) DC, Cloiselia S. Moore. papillae, apex slightly rounded to acute. (2 species), Pasaccardoa Kuntze (4 spe- Achenes cylindrical, ribbed, glabrous to cies), Pleiotaxis Steetz (26 species). densely villose; pappus of scabrid bristles or in Pertya, plumose bristles. Chromosome Ad Mutisieas similes sed in ramis stylo- numbers: Ainsliaea 2N = 24, 26, 30; Pertya rum base caespitose pilosis et in cellulis su- 2N = 24, 26, 28. perficialibus corollarum typis senecioidis All the members of this clade are found differt. in Asia, from Afghanistan through India to Perennial herbs, shrubs or small trees, China, Taiwan, Japan, and SE Asia. The young stems glabrous or pubescent, tri- only two wholly Asian genera of the Mu- chomes multicellular. Leaves alternate, pe- tisieae not in this clade are dunculate or sessile, blades linear to broad- D.Don and Nouelia Franch., which form a ly ovate, glabrescent to densely pubescent clade only known from southwestern China on abaxial surface, margins entire to ser- and adjacent regions. This China clade is rate, leaf bases various, leaf apices some- found near the base of the tree in the Mu- times terminating in a spine or mucro. In- tisieae s.s. (Fig. 1 ). The Pertya clade is well florescence terminal, solitary, or open thyr- supported and is located above the Car- siform cymes. Heads homogamous or het- dueae plus "African Mutisieae" clade (Car- erogamous, involucre campanulate to VOLUME 1 15, NUMBER 4 917 hemispherical or obconic with innermost rugose, corolla epidermis pattern as op- row of phyllaries producing a strong con- posed to the crested outer walls typical of striction at the end of the involucre, phyl- Mutisieae. However, the occurrence of a

laries in multiple series, herbaceous to scar- particular corolla cell morphology is not an ious sometimes innermost phyllaries her- unequivocal indicator of phyletic relation- baceous with scarious margins, sometimes ship within the family—see for instance with a prominent midrib, patent or spread- genera such as Ainsliaea and Pertya, in- ing; receptacle epaleate, rarely paleate. Flo- cluded by Hansen in Mutisieae but here re- rets actinomorphic or sometimes peripheral ferred to the Pertyoideae.

florets zygomorphic and usually termed bi- The genus Oldenburgia Less, is not in-

labiate with 3 of the lobes greatly expanded cluded in tribe Dicomeae as it appears from and appearing liguliform, corollas, white, present data to be sister to the Tarchonan- yellow, cream or various shades of pink, thus— clade (Tarchonantheae) mauve or lilac, tube narrow variously gla- and collectively the three genera are sister brous to pubescent, lobes conspicuous and to the Cardueae. Ortiz (2000) considered longer than tube, either erect or spreading; Oldenburgia a basal lineage of the tribe and anthers caudate calcarate, tails with 5, and used this taxon along with Gochnatia as or without antrorse ramifications, append- outgroups in his phylogenetic analyses of age apiculate to acute or slightly acuminate, Dicoma and related genera. Bond (1987) pollen smooth or echinate; styles with two considered Oldenburgia not to be closely or four veins, style branches straight or related to any other members of tribe Di- curved, with sweeping hairs restricted to bi- comeae and believed its sister taxon to be furcation point and above, lowermost tri- found among American members of tribe chomes slightly to conspicuously longer Mutisieae subtribe Gochnatiinae. Addition- than those at the distal end of style branch- al data are needed to ascertain the phylo- es. Achenes ellipsoid to obovoid or obcon- genetic position of Oldenburgia. ic, glabrescent to densely pubescent, pubes- The Dicomeae is recognized because it cence on ribs or in the space between them, members are morphologically distinct from carpopodium absent or rarely present, their sister clade the Oldenburgia—Taxchon- achene wall smooth to ribbed, pappus pres- antheae—Cardueae and there are no morpho- ent or absent, isomorphic or dimorphic, logical characters that unify these taxa. sometimes in several rows, persistent or ca- Also, the sister relationship of the Tarchon- ducous. Chromosome numbers: Dicoma 2N antheae and Oldenburgia with the Cardueae = 22; Pleiotaxis 2N = 20. is weak and additional data may show that This tribe contains most genera centered the Tarchonantheae and Oldenburgia are around Dicoma and found in Africa. These sister to the Dicomeae and not the Car- genera were previously classified in Mutis- dueae. ieae subtribe Gochnatiinae. The major ex- ception is the genus Gerbera L. and allies that remain in Mutisioideae. Our molecular Athroismeae Panero, trib. nov. studies support the exclusion of this Dico- Type: Athroisma, DC, Guill. Arch. Bot. 2: ma clade from the Mutisieae as proposed 516. 1833. by Hansen (1991) and their inclusion in Includes: Athroisma DC. (12 species), Ble- subfamily Carduoideae. Hansen (1991) pharispermum Wight ex DC. (15 spe-

stressed corolla cell morphology and col- cies), Leucoblepharis Arnott (1 species). lective patterns derived from these cells as his strongest evidence to support the exclu- Capitula discoidea vel disciformia palea- sion of the African Mutisieae from South ta, floribus peripheralibus 0—2 femineis in American Mutisieae. These genera show a coroUis tubiformibus; thecis antherarum 918 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON base brenter caudatis, parietibus achaenio- chomes with apical diverging cells straight rum phytomelaniniferous. or recurved, ovaries of functionally stami- Perennial herbs, shrubs or small trees, nate flowers in Blepharispermum obconical, young branches glabrous to pubescent, old brown, pappus of fertile flower achenes ei- stems gray to brown. Leaves alternate or ther a crown of twin trichomes sometimes fasciculate in brachyclasts, petiolate or ses- with recurved apices or of deeply lacerated sile, petioles sometimes with a basal spine; fused squamellae. Chromosome numbers: blades linear, lanceolate to ovate, obovate, Athroisma 2N = 20 (Eriksson 1993); Ble- bases attenuate, apices acuminate to retuse, pharispermum 2N = 20 (Eriksson 1992). cuneate or truncate, margins entire, dentate The Athroismeae contains the genera or serrate, venation triplinerved or with a Athroisma DC, Blepharispermum Wight ex single vein, abaxial surface with sessile DC, and Leucoblepharis Arnott centered in glandular trichomes. Inflorescence a con- eastern tropical Africa. These genera were gested glomerule-like cyme solitary or ar- considered to be members of the Inuleae ranged in open, thyrsiform cymes. Heads until they were placed in the Heliantheae disciform, heterogamous, involucre narrow- (Eriksson 1991). Eriksson allied the genera ly campanulate, narrowly conical, phylla- of his Blepharispermum group to members ries 0-2, involute, narrowly elliptical, ovate of subtribe Ecliptinae, more specifically to to oval, apiculate, shallowly to strongly the genera Salmea DC. and Acmella Rich, keeled, apical end entire, trilobed or vari- ex Pers. (Eriksson 1992). This choice of a ously lobed, or ciliate, hyaline, scarious, ab- sister group in America is a logical one giv- axial surface glabrous, pubescent and/or en that these genera share a series of su- glandular; receptacle paleate; pales normal- perficially similar characteristics with the ly narrower and longer than phyllaries or in Blepharispermum group such as a combi- the case of Athroisma wider than the phyl- nation of carbonized achenes, T-shaped laries, commonly with a round apex, rarely twin hairs on the achenes, the suborbicular apiculate. Florets all actinomorphic, female to broadly elliptical shape of the achenes, florets present or absent, 2 to several, co- and the white corollas. The genera Salmea rollas tubular, hermaphrodite florets 2—25, and allies such as Acmella and Spilanthes in Blepharispermum functionally staminate Jacq. differ from the Blepharispermum and only 2-^ present, corolla campanulate, group in having laterally compressed white, greenish-white, purple, tube white or achenes, heads with numerous hermaphro- greenish, glabrous or pubescent mostly with ditic disc flowers with peripheral ray flow- twin hair glands, lobes (4—) 5; anthers 5, ers instead of pistillate, tubular flowers pe- ecalcarate, caudate, thecae white, to pale ripheral to the heads in the Blepharisper- purple, endothecium with polarized thick- mum group, anthers with round and not sag- enings, appendage a continuation of the ittate appendages, and anthers without tails. connective and shallowly constricted above It is indeed this last characteristic that ex- thecae level sometimes with apical gland; cludes the Blepharispermum group from styles divided or filiform, filiform styles as- tribe Heliantheae, as there are no taxa in sociated with functionally staminate flow- Heliantheae with such anther tails. ers, stigmatic area in divided styles is di- Results from molecular studies of He- vided into two rows which are confluent at liantheae sensu lato and related genera (Pa- style branch apex, sweeping hairs present at nero et al. in prep) show that the Blephar- style apex or slightly below rarely present ispermum group (represented by Blephar- below the style branch bifurcation. Achenes ispermum and Athroisma in our studies) is black, radially flattened, ovate to oval, el- basal to the Heliantheae sensu lato. This re- liptic to suborbicular in outline, lateral ridg- sult supports the recognition of this group es ciliate, trichomes fused at base, twin tri- as a new tribe in the Compositae. Molecular VOLUME 1 15, NUMBER 4 919

Studies of members of tribe Inuleae (El- on abaxial surface with long tapered tri- denas et al. 1999) have revealed that the chomes reminiscent of those on tube of ray genus Anisopappus Hook. & Arn. is a basal corolla, glandular trichomes either sessile or element in the tribe and possibly another stipitate, pales with several resin tubes, res- member of tribe Athroismeae. Further mo- in yellowish. Ray florets 3-8, pistillate and lecular studies will be necessary to eluci- fertile, corolla white, tube densely pubes- date the relationships of Anisopappus to cent, limb deeply trilobed with central lobe members of the Athroismeae as circum- longer than flanking ones, adaxial surface scribed here. along margins with a few multicellular tri- chomes with tapered, terminal cells, abaxial surface with scattered glandular and multi- Polymnieae (H. Rob.) Panero, trib. stat. cellular trichomes. Disc corollas 10-85 + nov., Basionym: Polymniinae H. Rob. , Phytologia 41:43, 1978. functionally staminate, 5-lobed, corolla white to yellowish white or yellow, tube Type: L., Diss. Chen. 22. 1751; Amoen. Acad. 3:15. 1756. narrowly cylindrical, approximately half as Monotypic, 3 species long as throat, glabrous, throat broadly campanulate, lobes with a few, broadly Perennial herbs or annuals, stems sparse- based trichomes on abaxial surface; anthers ly to densely pubescent with a strigose to 5, ecaudate and ecalcarate, anther thecae softly hispid pubescence, internodes terete pale brown or tan, endothecium cells bi- to shallowly angular, yellowish green to polar with 2—^ bridges, connective and an- green with scattered purple spots, trichomes ther collar thickenings radial, anther ap- white to slightly brown, some trichomes up pendage conspicuously deltate with glan- to 2.5 mm long, glandular trichomes with dular trichomes on abaxial surface; pollen terminal brown cells, the base formed from 25-27 micrometers in diameter. Style of ray two rows of trichome cells. Leaves oppo- flowers divided into two separate branches site, petiolate, blades mostly ovate to sub- each with prominent stigmatic lines, sweep- orbicular in outline, bases attenuate to trun- ing hairs reduced to small papillate protu- cate, round, apices acuminate, margins en- berances on distal portion of abaxial surface tire to shallowly lobed or deeply dissected, of the tapering stigmas, stigmas of disc venation pectinate, abaxial surface densely flowers deltate and fused with no stigmatic covered with sessile glandular trichomes, surfaces with prominent sweeping hairs on tapered trichomes scattered along major abaxial surface of distal half. Achenes sub- veins, adaxial surface with a few sessile orbicular in outline and asymmetric in cross glandular trichomes and scattered, multicel- section, the abaxial half round and larger lular trichomes with raised multicellular ba- than the flattened adaxial half compressed ses. Inflorescence an open, paniculiform in its development by adjoining pale walls, cyme. Heads radiate, involucre campanu- with a ridge on each lateral edge and two late to hemispherical, subequal, phyllaries additional ridges on the adaxial surface that in two series, outer series herbaceous, inner can be variously distinct or fused, brown to series stramineous, chartaceous, with sev- black with a minutely coroniform pappus eral resin ducts, resin dark yellow; phylla- probably derived from the basal cells of the ries sparsely to densely pubescent on ab- corolla tube, with no striation on the achene axial surface with both tapered and glan- wall. Chromosome numbers: N = 15 II dular trichomes; receptacle slightly convex, with 1 fragment (Smith et al. 1992, Free- paleaceous; outermost pales broadly ovate man and Brooks 1988); other numbers in- to oval with irregularly lacerate distal ends, clude N = 16, 33+1 and 2N = 60, >50. this grading into filiform pales in the center The genus Polymnia was revised by of the head, distal end sparsely pubescent Wells (1965) to include 19 species from 920 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

North and South America with yellow and color. Leaves opposite, petiolate, blades white ray corollas. Robinson (1981) provid- palmate, suborbicular in outline with 2-7 ed a list of characters including the pale col- lobes on each side of the leaf, base shallow- ored anther thecae, the lack of a differen- ly hastate, apex acuminate, margins broadly tiated involucre, and a distinctive set of tri- serrate to crenate, each lobe ending with a chomes on the corolla lobes to justify the prominent mucro, venation triplinerved, ab- transfer of most species of Polymnia into axial surface paler green than adaxial sur- the genus Smallanthus Mackenzie. The face, sessile glandular trichomes moderate- three species presently recognized in Po- ly distributed across blade interspersed with lymnia are endemic to wet, riverbanks in a few tapered multicellular trichomes es- mesic, hardwood deciduous forests of the pecially along veins, adaxial surface bright eastern United States and Canada. green, sparsely pubescent with scattered, Molecular phylogenetic studies of mem- thickened multicellular trichomes, with bers of Heliantheae s.l. (Panero et al., un- prominent bases. Inflorescences terminal, published data) support Robinson's concept open cymes of 3-18 heads. Heads radiate, of the genus Polymnia and clarify its phy- nodding, involucre hemispherical, gradate, logenetic position. The genus Polymnia is imbricate, glabrous, phyllaries suborbicular a single lineage, basal to the tribes Helian- to round, green rimmed with purple, in 3 theae, Perityleae, Madieae, Eupatorieae, series, receptacle paleaceous, slightly con- and Millerieae. As suggested by Robinson vex; pales conduplicate, keeled, margins (1981) Polymnia is not related to members lacerated with 5-6 sclerified awns on each of subtribe Melampodiinae. There two rea- side, without a terminal awn but rather the sons for elevating the Polymniinae to tribal distalmost awns on each side of the pale

status: 1) the phylogenetic position of the joined and diverging from the pale apex, genus in the molecular study of the tribe glabrous, pales growing with age, spreading

Heliantheae (Panero et al. in prep.), and 2) and turning the fruiting head into a spiny the decision by Baldwin et al. (2002) to ball. Ray florets 9-15, neutral, corolla white split the traditional Helenieae. This decision suffused with purple or pink on abaxial reflects the desire of most workers to main- side, densely pubescent on abaxial surface tain the Eupatoriae as a tribe, and not a sub- with a few scattered glandular trichomes, tribe of a very large tribe Heliantheae. essentially glabrous on adaxial surface.

Disc florets 200+ , perfect, lobes deeply dis- Subtribes sected and spreading, distal section of co- rolla black, basal half white, tube densely Rojasianthinae Panero, subtrib. nov. pubescent, trichomes multicellular, tapered, Type: Rojasianthe Standley & Steyermark, blackish-green, throat sparsely to moderate- Publ. Field Mus. Nat. Hist. Bot. Ser. 22: ly pubescent, pubescence decreasing toward 311. 1940. distal end and lobes, lobes 5, black, essen- Monotypic (1 species) tially glabrous; anthers 5, ecaudate and Ad subtribustet Montanoinas similae sed ecalcarate, black, filament densely pubes- in paleis margine spiniferis, in corollis disci cent on its distal end immediately below an- bicoloratis, et in setis pappi pluribus cadu- ther collar, trichomes multicellulai; terminal ceis differt. cell shallowly rounded, black, those on bas- Shrubs or small trees, young stems terete, almost area of filament hyaline, apical ap- purplish green to brown, densely villous on pendage deltate, black; style white, style

leaf internodes and growing shoots, tri- branches white rimmed with black on outer chomes multicellular, tapered with a few edge, densely pubescent abaxially towards sessile glandular trichomes interspersed, old distal third of length and then abruptly gla- stems and trunk smooth, coppery-brown in brous on distalmost tip, black. Achenes ob- VOLUME 115, NUMBER 4 921 ovate, purple-raspberry in color, turning Acknowledgments brownish when mature, sparsely pubescent We thank M. Bonifacino, M. Dias de- to glabrous when mature, pappus of multi- Moraes, M. Koekemoer, P. Simon and T. ple caducous awns, awns golden yellow. Tinder-Smith for collecting plant material Chromosome number: 2N = 38 (Strother and S. Keeley for sending material of Hes- 1983). peromannia A. Gray. We thank Tetsukazo The monotypic genus Rojasianthe is en- Yahara and Aiko Ohtsuka of Kyushu Uni- demic to the mountainous regions of Me- versity, Japan, for providing DNA of Pertya soamerica on the border between Mexico and Ainsliaea. We are grateful to the Hesler Guatemala. range of the species is and The Fund, Department of Botany, University of In it more extensive in Guatemala. Mexico Tennessee, Knoxville, for providing funds is found in open cloud forests or natural to cover costs associated with sequencing clearings in the forest especially along wa- of the rpoB gene. We are grateful to B. Cro- terways. To the casual observer with some zier for assistance in the lab and to H. Rob- experience with tropical American sunflow- inson for providing the Latin diagnoses, ers Rojasianthe is reminiscent of a species reading the manuscript, and moral support, of Montanoa Cerv., especially M. liebman- as well as for his reminding us that now we nii S. F. Blake. It shares with this genus the have enough DNA strands of data to hang same chromosome number, opposite phyl- ourselves. We also thank G. Nesom for lotaxy, accrescent pales, blackish disc co- helpful comments on the manuscript. Funds rollas (e.g., Montanoa pteropoda S. F. from the National Science Foundation (to Blake), and neutral ray flowers with white JLP) and the Mellon Foundation (via the corollas. It is however, the accrescent pales Smithsonian's Fellowships and Grants pro- that are probably the most interesting mor- gram; to VAF) supported the molecular stud- ies; a Scholarly Studies grant phological feature shared by these two gen- Smithsonian funded some of the field work (to VAF). era. No other American sunflower has this Miscellaneous chromosome numbers were characteristic. Rojasianthe, like Montanoa, obtained from the website: index to chro- produces conspicuous fruiting balls that, as mosome numbers in Asteraceae, http:// in some species of Montanoa, are very www-asteraceae. cla.kobe-u.ac.jp/index. spiny to the touch once the pales become html. lignified and dry up. Results from molecular studies of the Literature Cited Heliantheae and relatives (Panero et al., in prep.) reveal that Montanoa and Rojasian- Baldwin, B. G., B. L. Wessa, & J. L. Panero. 2002. the constitute the basal lineages of the two Nuclear rDNA evidence for major lineages of different clades of Heliantheae sensu stric- helenioid Heliantheae (Compositae). Systematic Botany 27:161-198. to. Montanoa is sister to the species of sub- Bayer, R. J., & J. R Starr. 1998. Tribal phylogeny of tribe Ecliptinae as emended by Panero et al. the Asteraceae based on two non-coding chlo- (1999). Rojasianthe is basal to a clade con- roplast sequences, the trnL intron and the trnL/ taining members of the Verbesiniinae, Rud- trnP intergenic spacer. Annals of the Missouri Botanical Garden 85:242-256. beckiinae, Zaluzaniinae, Zinniinae, Helian- Bentham, G. 1873. Notes on the classification, history, thinae, Engelmaniiinae, Ambrosiinae. and and geographical distribution of Compositae. Montanoa, with its 25 species, and the Journal of the Linnean Society, Botany 13:335- monotypic Rojasianthe are each sister to a 577. Bond, R 1987. A revision of Oldenburgia (Astera- radiation in the American continents of a ceae—Mutisieae). South African Journal of total of approximately 1500 species of sun- Botany 53:493-500. flowers with mostly yellow corollas. Bremer, B., & R. K. Jansen. 1992. A new subfamily 922 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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