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Home , Aphanactis, Aphanostephus, Astereae, Astranthium, Chloracantha, Chrysopsis

8 LUNDELLIA DECEMBER, 2008

CLASSIFICATION OF SUBTRIBE CONYZINAE (:)

Guy L. Nesom 2925 Hartwood Drive, Fort Worth, 76109, USA Abstract: Subtribe Conyzinae includes , New World , the North American , and a small group of South American segregated as the genera Apopyros, , , , and Neja. Erigeron is the only genus with species native to regions outside of the New World. About 500 species are included in the subtribe. All North American, Central American, and South American species of Erigeron are included in the present treatment and assigned to one of the 35 sections recognized here. Ten new sections of Erigeron are recognized: sect. Disparipili, sect. Filifolii, sect. Gyrifolium, sect. Lonchophylli, sect. Meridionales, sect Microcephalum, sect. Quercifolium, sect. Radicati, sect. Rhizo- nexus, and sect. Terranea. Conyza is at least biphyletic; each of the groups is represented in the treatment but not all of the South American species are included. Molecular data have made it clear that traditional, North American species of Erigeron form the basal and terminal clades in the evolutionary topology of the subtribe, thus Conyza, Aphanostephus, and the other segregate genera have arisen from within the branches of Erigeron. Erigeron, as currently treated and tentatively maintained here, is paraphyletic. Broad taxonomic alternatives that include only monophyletic taxa are: (1) to treat the whole subtribe as Erigeron or, (2) to recognize Aphanostephus or Aphanostephus, Conyza, and the other South American segregates and at least an additional 5 to 10 new generic-level segregates from species groups traditionally treated as North American Erigeron.

Keywords: Compositae, Conyza, Erigeron.

Subtribe Conyzinae includes Erigeron L., by Nesom (1990a) and confirmed by Noyes New World Conyza Less., the North Amer- and Rieseberg (1999). The genus ican genus Aphanostephus DC., and several Cass. is segregated from Conyza (Zardini, small groups of South American species 1981; Nesom, 1990f, 1990g, 1992d, 2001; sometimes segregated as the genera Apopyros Nesom & Laferriere, 1990). The evolutionary G.L. Nesom, Darwiniothamnus Harling, origin of Aphanostephus now appears un- Hysterionica Willd., Leptostelma D. Don., equivocal—it arose from within the evolu- and Neja D. Don (Nesom, 1994d; Nesom tionary matrix of Erigeron, as indicated by and Robinson, 2006). A molecular-phyloge- the ITS data (Fig. 1) of Noyes (2000a), netic study of Erigeron and relatives (Noyes, confirming earlier research results based on 2000a) essentially confirmed this view of the data from chloroplast DNA (Morgan, 1990). subtribal composition, with a few caveats. Morphology had suggested that Aphanoste- One Mexican species originally described as phus was related to a different subtribe Erigeron (Sundberg and Nesom, 1990) is (Nesom, 1994d). Native species of excluded and now segregated as the mono- previously identified as Erigeron have been typic genus Batopilasia G.L. Nesom & R.D. transferred to Pappochroma Raf. (Nesom, Noyes (Nesom and Noyes, 2000); another 1994e, 1994f). Erigeron rapensis F. Brown, a Mexican species originally described as from the Polynesian island of Rapa, Erigeron now is recognized as the genus now constitutes the monotypic genus Paci- G.L. Nesom et al. (Nesom et figeron Nesom (Nesom, 1994g). al. 1991). Native African species of Conyza In contrast to an hypothesis that the belong in a different subtribe, as suggested Conyzinae arose from ancestors in the

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Southern Hemisphere (Nesom, 1994d), yzinae has species of Erigeron at the base and Noyes and Rieseberg (1999) found that the apex of the phylogenetic topology, with the evolutionary origin of the subtribe was from primarily Brazilian clade including Lepto- within a North American clade. South stelma, Apopyros, Neja, and Hysterionica American Conyzinae are derived from North having originated in a sister relationship American ancestors. with Erigeron species. Conyza appears to be In its current definition, the Conyzinae polyphyletic, at least biphyletic, and there is comprises about 500 species, about 400 of little understanding of the patterns of these generally identified as Erigeron. Eriger- variation among the South American species on is the only genus of Conyzinae with (excluding sect. Caenotus). Morphological species native to regions outside of the New boundaries between Erigeron and Conyza are World. An earlier summary (Nesom, 1989c) not clear and critical species of both have presented an outline of classification of the not been included in the molecular analyses. New World species of Erigeron, recognizing The distinctive and long-recognized genus 17 sections. In the last decade, however, new Aphanostephus apparently has arisen in a species and new sections have been de- sister relationship to an internal clade of scribed in the genus and new ideas regarding Erigeron. Classification at the generic rank infrageneric classification have come to light within the Conyzinae remains tentative. (citations below). In order to provide a If Noyes’s cladogram is accepted as a coherent framework for further studies in reasonable phylogenetic hypothesis for the the systematics and evolution of the Con- Conyzinae, at least in broad strokes and yzinae, a taxonomic update of the subtribe, acknowledging that it is tentative, three including all of the North American and broad taxonomic alternatives for the recog- Caribbean species, is presented here. Bound- nition of genera within the subtribe are aries of some groups of South American possible. Erigeron and Conyza are being clarified, but a complete account of these species is not yet 1. Adopt a strictly clade-based , possible. The South American taxa generally treating cladistically coordinate groups identified as Conyza are represented in the at coordinate ranks, recognizing Apha- present overview by relatively few species. nostephus, Apopyros, Conyza sensu The classification of the Eurasian species of stricto, Darwiniothamnus, Hysterionica, Erigeron has not yet been studied in detail, Leptostelma,andNeja,aswellas but many of them belong with sect. Erigeron sensu stricto (to include Trimorpha (Cass.) DC. and sect. Erigeron Trimorpha Cass. and perhaps Leptilon (see references by Huber and associates), Raf., the horse-weed conyzas) and at some apparently with sect. Fruticosus G. Don least eight other generic-level segre- Taxonomic treatments with keys to gates comprising various species species of New World Erigeron are available: groups traditionally treated within north of (Nesom, Erigeron. Simply the acceptance of 2006); Mexico (Nesom, in ms.); Central Aphanostephus within this conceptual America (Nesom & Pruski, in ms.); and framework would necessitate recogni- South America (Solbrig, 1962). tion of at least five new generic-level segregates, even if Conyza and the OVERVIEW OF SUBTRIBAL TAXONOMY Brazilian segregates all were main- tained within Erigeron sensu stricto— Molecular data indicate that Erigeron and all five of these segregates would and Conyza as currently treated are not come from within North American monophyletic (Noyes, 2000a, his cladogram and Central American Erigeron as reproduced here as Fig. 1). Subtribe Con- currently recognized. 10 LUNDELLIA DECEMBER, 2008

FIG. 1. Strict consensus tree for parsimony analysis of ITS nucleotide sequence data for Conyzinae. Reproduced with permission from Noyes (2000a). Numbers above the branches are bootstrap and decay indices. Branch lengths from one abrbitrarily delected tree are given below branches. Circled numbers identify topologically important nodes. ‘‘Section’’ refers to sections of Erigeron according to Nesom 1989c, 1990a, 1994d. Groups I–VI are groups of taxa, subjectively selected, sharing relationship as determined by the cladogram. E5 Erigeron,C5 Conyza. NUMBER 11 NESOM: REVIEW OF CONYZINAE 11

2. Enlarge Erigeron to include all species Darwiniothamnus Harling of Conyzinae, which would imply the Erigeron L. submergence of all previously recog- Hysterionica Willd. nized segregate genera, including Leptostelma D. Don Aphanostephus.Thisalternative,in Neja D. Don effect, is the opposite extreme of the first alternative. ERIGERON L., Sp. Pl. 2: 863. 1753 [nom. conserv. prop.]. (see Heath, 1994) 3. Maintain monophyletic and morpho- logically well-defined groups currently TYPE: L. (M.L. Green, recognized as genera at generic rank Prop. Brit. Bot. 181. 1929; W. Huber, Taxon (Aphanostephus, Apopyros, Darwi- 41: 563. 1992 – Herb. Linn., No. 994.23, typ. niothamnus, Hysterionica, Leptostelma, conserv. prop.) (also see comments in and Neja) —keeping most of the Nesom, 1989c). remaining species within Erigeron, Infrageneric groups were recognized for which then would be explicitly recog- most of the New World species by Nesom nized as a non-monophyletic (para- (1989c), and others have subsequently been phyletic) genus. The taxonomic rank added (Nesom and Noyes, 1999; Nesom, for treatment of Conyza sensu stricto 1990c, 1994a). The analysis of Noyes (2000a) and other well-defined groups, includ- and recent work on the species-level taxono- ing Leptilon (5 Erigeron sect. Caenotus my of Erigeron for the FNA treatment Nutt.), needs to be considered in detail. (Nesom, 2006) have indicated that some of the groups of Erigeron earlier recognized by Regarding the broader taxonomic ap- Nesom are not monophyletic. This reflects the proach, whether or not to recognize para- misplacement of individual species, but in phyletic taxa with formal rank, the Con- larger part it results from recognition of a yzinae provides an archetypal example. Many number of over-broadly defined groups. For botanists have signed a lengthy statement example, informally treated subgroups of sect. noting that they support formal recognition Asteroidea do not cohere as a single phyletic of paraphyletic taxa (Nordal and Stedje and element, sect. Olygotrichium and sect. Fruti- 148 signatories, 2005), and the Conyzinae cosus include evolutionarily disparate sub- comprises a large group of species where it groups of species, and sect. Geniculactis (sensu may be useful to sustain contrasting philo- Nesom, 1990c) comprises two separate sophical positions. groups. More strictly defined groups convey Definitions of genera within the Con- a better understanding of the evolutionary yzinae clearly need to be adjusted and patterns and distribution of diversity within refined. The present overview represents a the genus. Eventual refinement of the phylo- first step toward that end. genetic arrangement of these groups may suggest or require additional categories and TAXONOMIC OVERVIEW the composition and rank of some groups may be altered, depending on philosophical Astereae subtribe Conyzinae Horan., point-of-view and development of further Char. Ess. Fam. 93. 1847. Type genus, data useful for systematic interpretation. Conyza L. Subtribe Erigeroneae Gren. & Erigeron as treated here includes about Godr., Fl. France 2: 83, 92. 1850. Type genus, 400 species. 304 are New World species Erigeron L. (enumerated below); Old World species are Genera included: estimated to be around 80–100 (most Aphanostephus DC. numerous in eastern and Russia). Apopyros G.L. Nesom Figures 2–7 illustrate various aspects of Conyza Less. habit and floral diversity in the genus. 12 LUNDELLIA DECEMBER, 2008

FIG.4. (sect. Stenactis). Ray corollas are reflexing, heads nodding in bud. Photo by Robert Sivinsky, 2004, from CalPhotos FIG.2. (sect. Fruticosus). database, used with Sivinsky’s permission. Photo by Mrs. W.D. Bransford, 1986, from the Image Gallery of the Lady Bird Johnson E. arenarioides (D.C. Eaton ex A. Gray) Wildflower Center. A. Gray ex Rydb. E. salmonensis Brunsfeld & Nesom 1. Sect. Arenarioides (Rydb.) Nesom, Erigeron salmonensis was recently de- Phytologia 67: 70. 1989. Erigeron [unranked scribed (Brunsfeld and Nesom, 1989); the sp. group] Arenarioides Rydb., Fl. Rocky two species of sect. Arenarioides were Mts. 897. 1918, in clave. TYPE: Erigeron regarded as closely related but isolated arenarioides (D.C. Eaton ex A. Gray) A. Gray within the genus. Molecular data (Noyes, ex Rydb. 2000a) indicate that E. tweedyi Canby (sect. Taprooted, perennial; caudex branch- Spathifolium) is closely related to E. arenar- ing, retaining old bases; glabrous; stems ioides – with this molecular pointer, mor- branched; filiform or linear to linear- phological similarities between sect. Spathi- oblanceolate, not clasping; heads erect in folium and sect. Arenarioides are apparent, bud; rays white, often with an abaxial but sect. Spathifolium, as represented by E. midstripe, not coiling or reflexing; achenes tener, is placed by the molecular data most 1.5–2.2 mm long, 2-nerved; pappus of 10– closely related to sect. Osteocaulis. 16 bristles.

FIG.5. (sect. Dispar- ipili). Heads are nodding in bud. Yellow-rayed FIG.3. Erigeron clokeyi (sect. Stenactis). Ray species have evolved among white- and blue- corollas are reflexing, heads nodding in bud. rayed ones in sects. Disparipili, Erigeron, and Photo by Christopher L. Christie, 2003, from Osteocaulis. Photo by Christopher L. Christie, CalPhotos database. 2004, from CalPhotos database. NUMBER 11 NESOM: REVIEW OF CONYZINAE 13

FIG.7. (sect. Filifolii). Close-up of head. Photo by Steve Matson, 2006, from CalPhotos database.

E. garrettii?) or reflexing; achenes 1.5– 3.2 mm long (1–1.2 in E. cavernensis, 3.2– 3.5 mm in E. evermanii), 2-nerved; pappus of 10–35 bristles. Group A. U.S.A.; leaves thin, long- petioled, vestiture relatively sparse. FIG.6. (sect. Osteocaulis). E. cavernensis S.L. Welsh & N.D. At- Ray florets are completely absent. Photo by Steve wood Matson, 2006, from CalPhotos database. E. cronquistii Maguire 2. Sect Asterigeron Rydb., Fl. Rocky Mts. E. evermannii Rydb. 891. 1918. TYPE: Erigeron watsonii Erigeron E. uncialis Blake (incl. E. conjugans sect. Spathifolium Nesom, Phytologia 67: 84. Blake) 1989. TYPE: (A. Gray) A. Gray E. watsonii (A. Gray) Cronquist Perennial, taprooted; caudex simple or Group B. U.S.A.; leaves thick, short- more commonly a multicipital crown or petioled; vestiture closely white-strigillose. with elongating branches, forming mats (E. E. acomanus Spellenb. & P. Knight acomanus, E. tener) connected by systems of E. tener (A. Gray) A. Gray lignescent and caudex branches; E. tweedyi Canby vestiture short-strigillose to villous, hirsute, E. wilkenii O’Kane (O’Kane 1990) or glabrous, hairs white (E. tener, E. Group C. U.S.A.; leaves thick, short- acomanus, E. tweedyi, and E. wilkenii have petioled; glabrous to glabrate; caudex multi- similar vestiture; E. cronquistii, E. everman- cipital. nii, E. subglaber glabrous to glabrate); stems E. garrettii Rydb. branching (simple in E. acomanus, E. wilk- E. subglaber Cronquist enii); leaves spatulate to oblanceolate, entire, Group D. Mexico. with expanded and indurate petiole bases, E. cuatrocienegensis G.L. Nesom often folding, not clasping; heads 1 or 1–3, E. gypsoverus G.L. Nesom nodding in bud (E. acomanus, E. tweedyi), E. unguiphyllus G.L. Nesom behavior unknown in others; phyllaries with Molecular data (Noyes, 2000a) indicate a broad green midband, minutely glandular; that E. tweedyi is closely related to E. rays white to pink, not coiling (or coiling in arenarioides (sect. Arenarioides), but the 14 LUNDELLIA DECEMBER, 2008

closer morphological associates of E. tweedyi 3. Sect. Asteroidea Nutt., Trans. Amer. instead seem to be in sect. Spathifolium. Philos. Soc. ser. 2, 7: 308. 1841. LECTOTYPE): Noyes’s data also suggest that sect. Spathi- Nutt. (Nesom, 1989c) folium (as represented by E. tener) may be Erigeron [unranked sp. group] Decum- closely related to sect. Osteocaulis. The bentes Rydb., Fl. 359. 1906, in vestiture in sect. Spathifolium is similar to clave. TYPE: Erigeron decumbens Nutt. that of sects. Wyomingia, Arenarioides, and Erigeron [unranked sp. group] Asperu- Osteocaulis, but the species in the latter ginei Rydb., Fl. Rocky Mts. 897. 1918, in sections produce conspicuously coiling rays. clave. TYPE: Erigeron asperugineus (D.C. There also are suggestive similarities between Eaton) A. Gray sect. Spathifolium and sect. Asterigeron. Perennial, taprooted; caudex branches Erigeron subglaber has leaves oblanceo- generally slender (-like in E. decum- late to broadly oblanceolate or subspatulate bens); sparsely to densely strigillose or and differs from the other U.S.A. species, strigose to hirsute or hirtellous, stems and which usually have distinctly spatulate involucres often minutely glandular; stems leaves. This species apparently exists only simple or branched, basally ascending to as a single population system on the Elk decumbent; leaves linear to oblanceolate or Mountain Ridge of the Sangre de Cristo subspatulate, entire, usually with indistinct Mountains in San Miguel Co., petiole region, usually 3-nerved, cauline (ESE of Cowles, ca. 10,000–11,500 ft eleva- gradually reduced, not clasping; heads 1– tion). If not a member of sect. Spathifolium, 7+, commonly more than 1, erect in bud; it apparently is isolated geographically as rays white to blue, not coiling or only weakly well as morphologically from other possible so; achenes 1.2–3.5(–4.5) mm long, 2- close relatives (except one) within the genus. nerved; pappus of (6–)12–30 bristles. The identification of the original collection E. asperugineus (D.C. Eaton) A. Gray was E. leiomerus (sect. Asterigeron), to which E. canaani S.L. Welsh (incl. E. higginsii it is similar but which has nodding buds and S.L. Welsh) reflexing rays; the assessment of these E. corymbosus Nutt. behaviors in E. subglaber is based on E. decumbens Nutt. somewhat ambiguous observations of her- E. eatonii A. Gray (incl. E. sonnei barium specimens and online photographs. Greene, E. nevadincola Blake) Erigeron garrettii appears to be very similar E. jonesii Cronquist (incl. E. wahwahensis to E. subglaber, and they are placed here as S.L. Welsh, see Cronquist 1994) closely related despite their considerable E. lassenianus Greene (incl. E. flexuosus separation in geography. Cronquist) The species of Group B are gypsophilic E. maniopotamicus G.L. Nesom & T. endemics of northeastern Mexico (Coahuila, Nelson Nuevo Leo´n, and San Luis Potosı´, respec- E. robustior (Cronquist) G.L. Nesom tively), tentatively placed in sect. Spathifo- Sect. Asteroidea previously was inter- lium emphasizing their short-strigillose ves- preted to include a broader array of species titure and ray behavior. They are and species groups (Nesom, 1989c), but it is geographically disjunct from the more here limited to the strict sense (the ‘‘Erigeron typical members of the section; the closest decumbens group’’ of Nesom, 1989), follow- species of the section sensu stricto is E. ing indications of molecular data (Noyes, acomanus from northwestern New Mexico. 2000a). Studies of this group of species have None of the other species of northeastern recently been published (Strother and Fer- Mexico show features that suggest that they latte 1988; Nesom, 1992b, 1992c, 2004); E. might have been the immediate forebearers maniopotamicus is newly described (Nesom of E. cuatrocienegensis and E. gypsoverus. and Nelson, 2004). NUMBER 11 NESOM: REVIEW OF CONYZINAE 15

Molecular evidence (Noyes, 2000a) sug- are distinctly wider and fewer (45–76) and gests that is most closely the achenes longer (2 mm long) than in the related to species of sect. Asteroidea but it is other species of Cincinnactis. Molecular data closer in morphology to sect. Fruticosus, (Andrus et al., sin prep.), in contrast, suggest especially in its fibrous-rooted rhizomes, that E. stanfordii is phyletically basal-most or clasping leaves, villous involucres, coiling near it in the whole genus, far separated rays, and relatively large, 2(–4)-nerved from the species of sect. Cincinnactis. achenes. Erigeron basilobatus and E. veracruzensis, Erigeron asperugineus has a set of placed in sect. Cincinnactis in 1989, are features common to sect. Asteroidea: slen- transferred here to sect. Quercifolium,as der-taprooted with a simple or branched suggested by the molecular data of Noyes caudex, stems decumbent-ascending, leaves (2000a). The group of Caribbean species mostly basal, spatulate, and 1-nerved or with tightly coiling rays, including E. weakly 3-nerved, heads 1(–2), phyllaries bellioides and E. cuneifolius and placed by often purple, sparsely to moderately hirsu- Nesom (1989) within sect. Cincinnactis,is tulous, densely minutely glandular, rays here treated as sect. Microcephalum. Erigeron blue, not coiling or reflexing, achenes 2.5– exilis has been transferred to sect. Lampro- 3.1 mm, and pappus bristles 20–30. The caules G.L. Nesom & R.D. Noyes (Nesom hirtellous vestiture is more like sect. Pseu- and Noyes, 1999). Erigeron crenatus and E. derigeron; the often strongly petiolate-spat- socorrensis are transferred to sect. Caenotis. ulate leaves are suggestive of Group A of 5. Sect. Disparipili Nesom, sect. nov. sect. Asterigeron. The range of E. asperugi- Vestimento caulino trichomatibus pa- neus in , , , and tentibus in longitudine valde inequali, capi- is more similar to species of section Radicati. tulis in alabastro erectis, et radiis cincinatis 4. Sect. Cincinnactis Nesom, Phytologia distinctus. TYPE: Erigeron disparipilus Cron- 67: 73. 1989. TYPE: Erigeron longipes DC. quist Fibrous-rooted, rhizomatous (taprooted Taprooted, perennial; caudex simple; in E. oaxacanus), perennial (annual in E. cauline vestiture with spreading hairs of narcissus); caudex simple or divided into markedly unequal length (E. chrysopsidis, E. slender, rhizomelike branches; vestiture; disparipilis, E. poliospermus); stems simple or stems simple, rarely branched; leaves ovate- branched; leaves narrowly oblanceolate and lanceolate, ovate-lanceolate, clasping or sub- 1-nerved, entire, not clasping, mostly basal, clasping; heads erect (?) in bud; rays (80–) or cauline well-developed; leaf margins/ 120–350, mostly filiform, white, tightly petioles coarsely ciliate; heads erect in bud coiling at maturity; achenes ca 0.6–1 mm or on arching peduncles; rays present long, 2-nerved; pappus of 12–23 bristles (absent in E. chrysopsidis var. austiniae), (absent or reduced in number in E. narcis- white (yellow in E. chrysopsidis, E. piperia- sus). nus), coiling; achenes 1.6–1.8 mm (E. piper- E. longipes DC. ianus) to 2.1–3 mm to 3.5–4.5 mm (E. latus) E. narcissus G.L. Nesom long, 2-nerved; pappus of 14–30 bristles. E. oaxacanus Greenm. E. chrysopsidis A. Gray E. procumbens (Houston ex Mill.) G.L. E. davisii (Cronquist) G.L. Nesom Nesom (syn 5 E. Small) E. disparipilus Cronquist E. stanfordii I.M. Johnst. ex Nesom E. latus (A. Nelson & J.F. Macbr.) E. tephropodus G.L. Nesom Cronquist Erigeron stanfordii is placed here because E. nanus Nutt. of its tightly coiling rays (see images of E. piperianus Cronquist isotypes at MO, NY), although the laminae E. poliospermus A. Gray 16 LUNDELLIA DECEMBER, 2008

Cronquist (1994) hypothesized that sect. Pauciflori G. Don in Loudon, (and E. rydbergii Cronquist) Hort. Brit. 346. 1830. LECTOTYPE: Aster are closely related to E. untermannii S.L. pulchellus Willd. (5 Erigeron venustus Welsh but the latter is placed here in sect. Botsch.) (Sundberg and Jones, 1987) Wyomingia. The distinction of E. davisii and Erigeron [unranked sp. group] Uniflori its relationship to this group were noted by Rydb., Fl. Colorado 359. 1906, in clave. Nesom (2004). TYPE: Erigeron uniflorus L. 6. Sect. Erigeridium Torr. & A. Gray, Fl. Erigeron sect. Monocephali Vierh., Beih. ECTOTYPE N. Amer. 2(1): 176. 1841. TYPE: Erigeron Bot. Centralbl. 19: 492. 1906. L vernus (L.) Torr. & A. Gray (Nesom 1989c): Erigeron uniflorus L. Fragmosa Raf., Fl. Tellur. 2: 50. 1836. Erigeron sect. Siphonoglossa Botsch., Bot. LECTOTYPE (Nesom 1989c): Erigeron nudi- Mater. Gerb. Bot. Inst. Komarova Akad. caulis Michx. (5 (L.) Torr. Nauk SSSR 16: 393. 1954. TYPE: Erigeron & A. Gray) uniflorus Erigeron [unranked sp. group] Verni Erigeron sect. Platyglossa Botsch., Bot. Small, Man. Southeastern Fl. 1395. 1933, in Mater. Gerb. Bot. Inst. Komarova Akad. clave. TYPE: Erigeron vernus (L.) Torr. & A. Nauk SSSR 16: 388. 1954. TYPE: Erigeron Gray venustus Botsch. Fibrous-rooted, perennial, sometimes Perennial; fibrous-rooted, usually from rhizomatous or stoloniferous; caudex sim- a short, suberect rhizome; caudex simple or ple; glabrous to sparsely strigose or hispid; with short, thick branches; strigose to stems simple or branched; leaves obovate to hirsute or villous; stems simple; leaves spatulate, often thick, entire or shallowly mostly basal oblanceolate to obovate or toothed, not clasping; heads erect in bud; spatulate, entire, not clasping; heads 1, erect rays white, not coiling or reflexing; achenes in bud; rays white or blue (yellow in E. 1.2–1.6 mm, 4-nerved; pappus of 16–30 aureus), coiling (not coiling in E. uniflorus); bristles. achenes 1.6–2.8 mm long, 2-nerved; pappus E. hintoniorum G.L. Nesom of (7–)10–30 bristles (accrescent in E. E. morelensis Greenm. uniflorus). E. palmeri A. Gray North America E. vernus (L.) Torr. & A. Gray E. algidus Jeps. (incl. E. petiolaris E. wellsii G.L. Nesom Greene, see Nesom, 1992b, 1992c) This group of species was earlier includ- E. aureus Greene ed within the primarily western U.S.A. sect. E. alpiniformis Cronquist Fruticosus (Nesom, 1989c) as the ‘‘E. palmeri E. X arthurii Boivin group.’’ The apparent morphological coher- E. denalii A. Nelson ence of these five species, however, and the E. eriocephalus J. Vahl geographic range divided between the east- E. flettii G.N. Jones ern USA and the eastern sierra of Mexico, E. grandiflorus Hooker (incl. E. simplex also characteristic of other groups in various Greene) genera, indicate that sectional status is E. hultenii Spongberg (status uncertain; appropriate for the E. palmeri group. Species Nesom and Murray 2004) of sect. Erigeridium have not yet been E. humilis Graham included in molecular studies. E. hyperboreus Greene (incl. E. alaskanus 7. Sect. Erigeron Cronquist) Erigeron sect. Uniflori G. Don in Lou- E. melanocephalus A. Nelson don, Hort. Brit. 343. 1830. TYPE: Erigeron E. muirii A. Gray uniflorus L. E. porsildii Nesom & D. Murray NUMBER 11 NESOM: REVIEW OF CONYZINAE 17

South America (The E. andicola DC. 1967; Spongberg, 1971). In the Noyes complex, see Solbrig, 1962) analysis, the Andean taxa of South America Erigeron andicola DC. are represented by E. cinereus. Erigeron ciliaris Phil. Among the North American taxa, Erig- Erigeron cinereus Hook. & Arn. eron uniflorus alone has accrescent pappus Erigeron incertus (d’Urville) Skottsb. bristles, a feature shared with the species of Erigeron lanceolatus Wedd. sect. Trimorpha. Erigeron leptopetalus Phil. 8. Sect. Filifolii (Rydb.) Nesom, comb. Erigeron myosotis Pers. et stat. nov. Erigeron [unranked sp. group] Erigeron patagonicus Phil. Filifolii Rydb., Fl. Rocky Mts. 897. 1918, in Erigeron pratensis Phil. clave. TYPE: Erigeron filifolius Nutt. ( and Russia not repre- Perennial, taprooted; caudices multicipi- sented) tal, often with relatively short and thin, E. anisophyllus Rech. f. stemlike, lignescent branches; sparsely to E. caucasicus Steven densely strigose with loosely appressed to E. cedretorum Rech. f. ascending, fine hairs, densely white-strigose at E. cilicicus Boiss. ex Vierh. least proximally; leaves linear to filiform, basal E. daenensis Vierh. (not in a rosette) and cauline (gradually or E. eriocalyx (Ledeb.) Vierh. little reduced distally); heads 1–5(–10+)from E. hyrcanicus Bornm. & Vierh. distal branches, in loosely corymbiform E. komarovii Botsch. arrays, erect in bud; rays weakly coiling; E. koraginensis (Komarov) Botsch. achenes 1.4–1.8(–2) mm long, 2-nerved. E. miyabeanus Tatew. & Kitam. Erigeron filifolius Nutt. E. multiradiatus (DC.) C.B. Clarke Erigeron filifolius has features in com- E. petroiketes Rech. f. mon with sect. Wyomingia (but is geograph- E. plesiogeron Rech. f. ically displaced and ecologically incongruent E. pseuderiocephalus M. Popov and lacks the multinerved achenes of the E. silenifolius (Turcz.) Botsch. multi-headed species) and sect. Osteocaulis E. thunbergii A. Gray (but lacks the characteristic indurate-sheath- E. uniflorus L. ing leaf bases) and some features in common E. zederbaueri Vierh. with Group A of sect. Asteroidea.Itis Molecular data (Noyes, 2000a) indicate without obvious near-sister species and that the species of sect. Trimorpha (at least placed here in a monotypic section, with the European species Erigeron acris and E. speculation that its closest relatives are borealis) are more closely related to E. geographically close, perhaps those of sect. uniflorus and to E. grandiflorus (incl. E. Osteocaulis. simplex) than they are to a group of American species including E. flettii, E. 9. Sect. Fruticosus G. Don in Loudon, humilis, E. cinereus, and E. lonchophyllus.In Hort. Brit. 343. 1830. TYPE: fact, sect. Trimorpha and sect. Erigeron sensu Ker-Gawler stricto appear to be more closely related to Woodvillea DC., Prodr. 5: 318. 1836. sect. Rhizonexus than to the latter group of TYPE: Erigeron glaucus Ker-Gawler sect. Erigeron. The implication of this is that Erigeron sect. Phoenactis Nutt., Trans. sect. Erigeron (as outlined above) is biphy- Amer. Philos. Soc. ser. 2, 7: 310. 1841. letic. The amphi-Atlantic E. humilis, howev- LECTOTYPE (Nesom, 1989c): Erigeron specio- er, is hypothesized to have arisen as an sus (Lindl.) DC. allotetraploid offspring of a cross between E. Erigeron [unranked sp. group] Elatiores uniflorus and a species similar to E. melano- Rydb., Fl. Colorado 359. 1906, in clave. cephalus and E. hyperboreus (Engelskjo¨n, TYPE: (A. Gray) Greene 18 LUNDELLIA DECEMBER, 2008

Erigeron [unranked sp. group] Salsugi- E. peregrinus (Pursh) Greene nosi Rydb., Fl. Colorado 359. 1906, in clave. E. potosinus Standl. TYPE: Erigeron salsuginosus (Rich.) A. Gray E. thunbergii A. Gray complex (Asia) (5 E. peregrinus subsp. callianthemus E. yukonensis Rydb. (syn 5 E. glabellus (Greene) Cronq. 5 E. glacialis Nutt.) var. yukonensis (Rydb.) Hulte´n) Erigeron [unranked sp. group] Macranthi Group B. Basal leaves persistent; achenes Rydb., Fl. Colorado 359. 1906, in clave. TYPE: 2-nerved or 2(–4)-nerved; buds nodding (E. Erigeron macranthus Nutt. (5 E. speciosus) coulteri, E. rybius), behavior unknown in Erigeron [unranked sp. group] Glabelli others. Rydb., Fl. Colorado 359. 1906, in clave. E. coulteri Porter TYPE: Nutt. E. rybius G.L. Nesom Erigeron [unranked sp. group] Asperi E. hessii G.L. Nesom Rydb., Fl. Rocky Mts. 897. 1918, in clave. E. kuschei Eastw. TYPE: Erigeron asper Nutt. (5 E. glabellus) Group C. Basal leaves scalelike or not Erigeron sect. Peregrinus Nesom, Syst. strongly differentiated; achenes 2-nerved or Bot. 7: 463. 1982. TYPE: 2(–4)-nerved; buds nodding. (Pursh.) Greene E. elatior (A. Gray) Greene Perennial, fibrous-rooted, rhizomatous, E. speciosus (Lindl.) DC. sometimes with stolons; caudex branched; E. subtrinervis Rydb. glabrate to strigose, hirsute, or villous; stems E. uintahensis Cronquist usually branched; leaves oblanceolate to E. vreelandii Greene (incl. E. platyphyllus spatulate, entire or toothed cauline nearly Greene) even-sized upward, clasping to subclasping; Erigeron yukonensis is rhizomatous and heads 1 (E. howellii) or 1–15(–22) in a fibrous-rooted, although the caudices or loosely corymbiform arrangement, nodding rhizomes sometimes resemble taproots. in bud (apparently erect in E. aliceae, E. Hulte´n (1968) considered it to be closely glaucus, E. glacialis); rays white to blue or related to E. glabellus. pink, weakly coiling or not at all; achenes Molecular evidence (Noyes, 2000a) indi- 1.1–3 mm long; pappus of 14–32 bristles. cates that Erigeron aliceae is most closely Group A. Basal leaves persistent; related to species of sect. Asteroidea, but it is achenes 2-nerved (E. arizonicus, E. cervinus, closer in morphology to sect. Fruticosus, E. formosissimus, E. glabellus, E. yukonensis) especially in its fibrous-rooted and rhizoma- or mostly 2(–4)-nerved (E. aliceae, E. tous habit, clasping leaves often relatively eximius, E. glaucus, E. potosinus) or (4–)5(– large and even-sized up the stem, villous 7)-nerved (E. glacialis, E. howellii, E. pere- involucres, non-coiling rays, and relatively grinus); buds apparently erect (E. aliceae, E. large (2–2.8 mm long), 2(–4)-nerved achenes. cervinus, E. glacialis, E. glaucus), or nodding? Erigeron arizonicus is rhizomatous and (E. formosissimus). fibrous-rooted but the caudices or rhizomes E. aliceae Howell are relatively thick and sometimes resemble E. arizonicus A. Gray a taproot; basal leaves are usually persistent, E. cervinus Greene (incl. E. delicatus spatulate, obovate to ovate, or elliptic, and Cronquist) cauline leaves are subclasping, but they often E. eximius Greene are 3-nerved and have shallowly serrate E. formosissimus Greene margins, features unusual in Group A of E. glabellus Nutt. sect. Fruticosus. E. glacialis Nutt. (syn 5 E. peregrinus var. is relatively isolated, as callianthemus (Greene) Cronquist) early observed by Cronquist (1947, p. 217), E. glaucus Ker-Gawler who noted that ‘‘Aside from E. cervinus,[E. E. howellii A. Gray delicatus] has no very close relatives known NUMBER 11 NESOM: REVIEW OF CONYZINAE 19

to me.’’ For E. cervinus he noted that it is 11. Sect. Geronpternix Nesom & Noyes, ‘‘probably derived from E. peregrinus, but Sida 18: 1163. 1999. TYPE: Erigeron rhizo- the connection is now remote.’’ matus Cronquist Perennial, rhizomatous, with thick fi- 10. Sect. Geniculactis Nesom, Phytologia brous roots, rhizomes or rhizomelike caudex 69: 250. 1990. TYPE: Erigeron coronarius branches creeping-ascending; slender, scale- Greene leaved, without well-defined central axes; Taprooted, annual or short-perennial; strigose to strigose-hirsutulous; stems simple caudex simple; hirsute to hispid; stems or 1–3 branched; leaves cauline, mostly branched; leaves linear to linear-oblong, linear at midstem, entire, not clasping; heads -lanceolate, or -oblanceolate, not clasping, 1(–3), nodding or erect in bud; rays with entire to toothed, rarely lobed; heads erect in abaxial lilac midstripe, not coiling or reflex- bud; disc corollas often prominently inflated ing; achenes 3.5–4.5 mm long, 2–4-nerved above the short tube; rays 40-200, white, (E. lepidopodus) or 5–6-nerved (E. rhizoma- reflexing or (in Erigeron sceptrifer) down- tus); pappus of 25–50 bristles. ward curving-reflexing, barely longer than E. lepidopodus (B.L. Rob. & Fernald) the involucre in E. zacatensis; achenes 0.5– G.L. Nesom 1 mm long, 2-nerved (orange); pappus of 2– E. rhizomatus Cronquist 17 bristles or (in E. reinana) absent. E. These two species, previously hypothe- reinana has relatively few, loosely coiling rays and uninflated disc corollas. sized to be sister species (Nesom, 1989c), were tentatively treated as members of the E. E. arisolius G.L. Nesom foliosus group (sect. Pycnophyllum) in the E. coronarius Greene Nesom (1989c) overview of Erigeron but E. janivultus G.L. Nesom were shown by Noyes (2000a) to be the E. reinana G.L. Nesom phyletically basalmost species of the Con- E. sceptrifer G.L. Nesom yzinae. As noted by Noyes (2000a, p. 106), E. zacatensis G.L. Nesom these two species are characterized by a ‘‘4- The species of sect. Geniculactis are bp insertion that is shared with outgroup restricted to Mexico, except for two that taxa and and with all also occur in southern – E. arisolius other Astereae, but that is absent in all other and E. sceptrifer. Erigeron reinana was Conyzinae.’’ recently added to the group (Nesom, 1998). The E. pumilus group was tentatively 12. Sect. Gyrifolium Nesom, sect. nov. included in the original circumscription of Radice palari crassa, foliis basalibus sect. Geniculactis (Nesom, 1990c) but is here persistentibus spatulati-obovatis, capitulis treated separately as sect. Stenactis Cass. in alabastro erectis(?), et radiis leniter Erigeron reinana is recognized by its cincinatis vel noncincinatis distinctus. TYPE: taprooted habit, coarse vestiture, narrow, A. Gray entire leaves, numerous heads on short Perennial, taprooted (stout); caudex peduncles, erect buds, loosely coiling rays, branches short, thick; hirsute to villous, and much reduced, essentially bristle-less stipitate-glandular or minutely glandular; pappus. It differs from E. coronarius and the stems simple or branched; basal leaves other species of sect. Geniculactis in its fewer, persistent, spatulate-obovate, 1(-3)-nerved, loosely coiling rays and uninflated disc toothed, cauline leaves little reduced upward corollas; it might be placed with less (clasping to subclasping in E. cascadensis, E. technical/diagnostic anomaly in sect. Im- leibergii); heads usually 1 but sometimes 2–4 barba, but the vestiture, leaf morphology, more developed from lower axils, erect? in and capitulescence of E. reinana are more bud; phyllaries relatively broad, erect, and similar to sect. Geniculactis. even-length; rays white, coiling but sometimes 20 LUNDELLIA DECEMBER, 2008

weakly or not at all; disc corollas with long overview: E. macdonaldii (Nesom, 1990d) (1.2–1.5 mm) tubes; achenes 1.2–2.2 mm and E. quiexobrensis (Nesom, 1990e). All long, 2-nerved; pappus of 12–20 bristles. species of the section are restricted to E. cascadensis A.A. Heller Mexico and . Sect. Imbarba E. leibergii Piper and sect. Geniculactis are the closest relatives E. oreganus A. Gray of most South American Conyzinae, accord- The species of sect. Gyrifolium are rare ing to the molecular data of Noyes (2000a). and grow on ledges and cliffs in western 14. Sect. Karvinskia Nesom, Phytologia , , and . 67: 79. 1989. TYPE: DC. They were previously placed within sect. Perennial (annual in E. barbarensis), Fruticosus, emphasizing the relatively well- taprooted (E. barbarensis, E. karvinskianus, developed and subclasping cauline leaves, E. irazuensis, E. pacayensis?) or rhizomatous but the taproot and other features (caudex, (E. aquarius, E. fluens, E. heteromorphus); leaves, ray corollas, and phyllaries) suggest caudex simple; strigose to hirsute; stems that the two groups are not so closely branched; leaves spatulate to oblanceolate or related. The name (‘gyrifolium’) alludes to linear, entire or toothed, not clasping; heads a resemblance of a spinning top to the whorl erect in bud; rays white, becoming lavender of basal leaves atop a thick taproot. at maturity, slightly coiling or not at all; 13. Sect. Imbarba Nesom, Phytologia 67: achenes 1–1.9 mm long, 2-nerved; pappus of 78. 1989. TYPE: Erigeron galeottii (A. Gray ex 10–27 bristles. Hemsl.) Greene E. aquarius Standl. & Steyerm. Perennial (annual in E. strigulosus), E. barbarensis Nesom & T.R. Van rhizomatous (sometimes taprooted in E. Devender strigulosus), sometimes with basal offsets or E. fluens G.L. Nesom producing stolons or runners; caudex simple E. heteromorphus B.L. Rob. or branched; glabrous to sparsely strigose, E. irazuensis Greenm. hirsute, or pilose; stems simple or few- E. karvinskianus DC. branched; leaves spatulate to oblanceolate, E. pacayensis Greenm. entire or toothed, cauline clasping to sub- Two new species have recently been clasping or not clasping; heads erect in bud; added to sect. Karvinskia (Nesom 1998; rays white, weakly coiling; achenes 1–3 mm Nesom and Van Devender, 2007). The long, 2-nerved; pappus a fimbriolate or native range of these species is restricted to scaley crown, bristles absent or occasionally Mexico and Central America; Erigeron a few, short, vestigial bristles present, or karvinkianus has become a worldwide weed pappus in E. macdonaldii of 5–7 short, in tropical and subtropical latitudes. Eriger- persistent bristles 1/2–2/3 as long as the disc on pacayensis appears to be sometimes corollas. taprooted, sometimes rhizomatous. E. astranthioides De Jong & G.L. Nesom 15. Sect. Lamprocaules Nesom, Phytolo- E. forreri (Greene) Greene gia 76: 99. 1994. TYPE: Erigeron pattersonii E. fraternus Greene G.L. Nesom E. galeottii (A. Gray ex Hemsl.) Greene Perennial, fibrous-rooted, slender-rhizo- E. guatemalensis (Blake) G.L. Nesom matous (taprooted in E. tuerckheimii); caudex E. macdonaldii G.L. Nesom divided into short or slender, rhizomelike E. mimus (Blake) G.L. Nesom branches; glabrous to sparsely strigose; stems E. quiexobrensis G.L. Nesom simple or branched, glabrate, shiny; leaves E. strigulosus Greene coriaceous, basal (apparently produced only Two new species have been added to in E. chiangii) spatulate to oblanceolate, entire sect. Imbarba after the Nesom (1989c) to shallowly few-toothed, cauline filiform to NUMBER 11 NESOM: REVIEW OF CONYZINAE 21

linear to narrowly elliptic, not clasping; heads This species of northern North America erect in bud; rays white, weakly coiling or not has been regarded as closely similar to the at all; achenes 0.8–1.8 mm long, 2-nerved; primarily Californian pappus of 11–22 bristles. group, treated here as sect. Pycnophyllum E. chiangii G.L. Nesom Cronquist (Cronquist, 1947; Nesom, 1989c, E. exilis A. Gray 1992a), but E. hyssopifolius was regarded as E. pattersonii G.L. Nesom more isolated and formally segregated as a E. scoparioides G.L. Nesom monotypic section by Nesom and Noyes E. tuerckheimii Urb. (incl. E. ocoensis (1999). Urb. & Eckman?) 17. Sect. Lonchophylli Nesom, sect. Erigeron exilis was recently transferred to nov. sect. Lamprocaules from sect. Cincinnactis Habitu fibrosi-radicato non-rhizomato, (Nesom and Noyes, 1999). caudice simplici, capitulis plerumque 3–12 Erigeron tuerckheimii is endemic to the plerumque in dispositione laxe racemiformi Caribbean island of Hispaniola. The in alabastro erectis, radiis filiformibus non- are small up 20 cm tall, from a woody reflexis noncincinatis involucro parum long- taproot producing thick branches with ioribus distinctus. TYPE: Erigeron lonchophyl- persistent old leaves. The leaves are linear lus Hook. to nearly filiform, with a hard, shiny texture, Biennial or short-lived perennial (some- purple at the base, and are very densely times appearing annual), fibrous-rooted, not arranged on the stems. The heads are solitary rhizomatous; caudex simple; sparsely to on naked or few-bracteate peduncles. Rays densely hirsute, eglandular; stems branched; are white, apparently neither coiling nor basal leaves oblanceolate to spatulate, entire, reflexing. There is no other Caribbean cauline mostly linear, not clasping; heads 1 species similar to it (except the possibly or 3–12 usually in a loosely racemiform conspecific E. ocoensis), and it is placed here arrangement, erect in bud; rays white to with this group of Mexican species on the pink, filiform, slightly longer than the basis of their similarity in leaf morphology. involucre, not reflexing or coiling; achenes Erigeron tuerckheimii is comparable in habit 1.3–1.8 mm, 2-nerved; pappus of 20–30 to species of Neja, but the achenes are flat nonaccrescent bristles. and 2-nerved, in contrast to those of Neja, E. lonchophyllus Hook. which are terete and multi-nerved. is similar to 16. Sect. Linearifolii (G. Don) Nesom, species of sect. Trimorpha in its fibrous- Phytologia 67: 79. 1989. rooted habit, narrow involucres in a racemi- Aster sect. Linearifolii G. Don in Lou- form arrangment, filiform suberect rays, and it was earlier included in sect. Trimorpha don, Hort. Brit. 346. 1830. LECTOTYPE: Michx. (Sundberg (Nesom 1989c). It lacks, however, the inner and Jones, 1987) zone of eligulate pistillate florets character- istic of sect. Trimorpha, and molecular data Perennial, rhizomatous, fibrous-rooted; (Noyes, 2000a) indicate that it is more caudices sometimes branched; strigose to closely related to a group of American strigose-villous; stems branched; basal and species including E. flettii, E. humilis, and proximal cauline leaves much reduced or E. cinereus, placed here in sect. Erigeron. present as scales, cauline blades linear to linear-oblong or oblong-lanceolate, not 18. Sect. Meridionales G.L Nesom & N. clasping; heads erect in bud; rays white, Andrus, sect. nov. not coiling or reflexing; achenes 1.3–1.6 mm Habitu rhizomato fibrosi-radicato, ca- long, 5–6-nerved; pappus of 25–35 bristles. pitulis solitariis in pedunculis scaposis vel E. hyssopifolius Michx. plerumque in dispositione laxe corymbi- 22 LUNDELLIA DECEMBER, 2008

formi in alabastro erectis distinctus. TYPE: erectis, radiis filiformibus arcte cincinatis Erigeron fasciculatus Colla distinctus. TYPE: Erigeron cuneifolius DC. Perennial herbs or , rhizoma- Annual to perennial, fibrous-rooted, not tous, fibrous-rooted; caudex simple or rhizomatous, stoloniferous in some; caudex multicipital; glabrous to strigose or villous; simple; glabrous to sparsely strigose; stems stems branched; leaves oblanceolate to simple, less commonly branched, filiform, lanceolate, entire to toothed or lobed, not bracteate; basal leaves in a rosette, spatulate clasping; heads 1 on scapose peduncles with orbicular to obovate or cuneiform (Erigeron othonnifolius, E. turricola)or blades (oblanceolate-elliptic in E. purpur- mostly in loosely corymbiform arrays, erect ipes), 1-nerved, entire to shallowly crenate or in bud; rays white to purple, not coiling or serrate on the distal half, lobed to pinnatifid reflexing; achenes 1–4 mm long (0.5– in some, cauline bracteate, subclasping or 1.0 mm in D. alternifolius), 2(–3)-nerved; not clasping; heads 1(–few), tiny (involucres pappus of [not counted] bristles. 2–6 mm long, 2–7(–8) mm wide), erect in Darwiniothamnus alternifolius Lawesson bud; rays 24–54, 40–80(–100), white, fili- & Adsersen form, tightly coiling; disc corollas 10–17, Erigeron campanensis Valdeb., Lowrey, fewer than the ray; achenes 0.8–1.4 mm & Stuessy long, rectangular, 2-nerved (orange-resin- Erigeron chionophilus Wedd. ous); pappus of 17–28 bristles. Erigeron ecuadoriensis Hieron. E. bellidiastroides Griseb. (Cuba) Erigeron fasciculatus Colla E. bellioides DC. (Puerto Rico, Cuba, or Erigeron incaicus Solbrig Hispaniola) Erigeron luxurians (Skottsb.) Solbrig syn5 E. semiovalis Urb. (Cuba) Erigeron othonnifolius Hook. & Arn. E. capillipes Ekman & Urb. (Cuba) Erigeron paramensis Aristeg. & Cuatrec. E. cuneifolius DC. (Puerto Rico) Erigeron raphaelis Cuatrec. E. hyoseroides Griseb. (Cuba) Erigeron rosulatus Wedd. E. libanensis Urb. (Cuba) status unknown: E. ocoensis Urb. & Ekman (Hispaniola) Erigeron adscendens Turcz. (Ecuador) E. paucilobus Urb. (Cuba) Erigeron apiculatus Benth. (Ecuador) E. purpuripes Britton & P. Wilson Erigeron (Aster) gilliesii (Hook. & Arn.) (Cuba) Cabrera (Patagonia) E. taylori Britton & P. Wilson (Cuba) Erigeron pauper Benoist (Ecuador) E. thrincioides Griseb. (Cuba) Molecular evidence (Andrus et al., in E. vegaensis Urb. (Hispaniola) prep.) suggests that sect. Meridionales may These species were not fully accounted not be monophyletic. Erigeron ecuadoriensis for in Nesom’s (1989c) overview of Eriger- is indicated by molecular data (Andrus et on, except for the observation that the al., submitted) to be more closely related to group, as characterized by E. cuneifolius E. maximus (Leptostelma)thanothersof and E. bellioides, probably belonged with sect. Meridionales.ThegroupofJuan sect. Cincinnactis. Erigeron bellioides shows Fernandez hexaploids segregated here as as the sister taxon to Aphanostephus in the sect. Terranea probably are derived from molecular analysis by Noyes (2000a). Erig- sect. Meridionales. eron cuneifolius has become a pantropical weed. The group is in need of revisionary 19. Sect Microcephalum G.L. Nesom, study. sect. nov. Habitu fibrosi-radicato non-rhizomato, 20. Sect. Olygotrichium Nutt., Trans. caudice simplici, caulibus simplicibus, foliis Amer. Philos. Soc. ser. 2, 7: 311. 1841. TYPE: basalibus in rosette, capitulis in alabastro Torr. & A. Gray NUMBER 11 NESOM: REVIEW OF CONYZINAE 23

Heterochaeta DC., Prodr. 5: 282. 1836 E. versicolor (Greenm.) Nesom (syn 5 E. (non Besser 1827). LECTOTYPE: Erigeron gilensis Wooton & Standl.; syn 5 E. pubescens Knuth (Nesom, 1989c) mimegletes Shinners) Erigeron [unranked sp. group] Bellidias- E. vicinus G.L. Nesom tra Rydb., Fl. Colorado 359. 1906, in clave. Group B. TYPE: Nutt. E. lobatus A. Nelson Erigeron [unranked sp. group] Diver- E. piscaticus G.L. Nesom gentes Rydb., Fl. Colorado 359. 1906, in E. velutipes Hook. & Arn. clave. TYPE: Erigeron divergens Torr. & A. The three species of Group B (all Gray taprooted annuals) are closely interrelated, Erigeron [unranked sp. group] Flagel- apparently part of sect. Olygotrichium. They lares Rydb., Fl. Colorado 359. 1906, in clave. are characterized by a densely stipitate- TYPE: A. Gray glandular and sparsely hispid-pilose vesti- Annual to biennial or perennial; tap- ture with straight, spreading hairs 0.6–1.5 rooted or fibrous-rooted, not rhizomatous, (–2) mm long. Their distinctions, with sometimes producing runners or stoloni- description of a new species, were discussed form branches; caudex usually simple; vil- by Nesom (1989d). The report of E. velutipes lous to hirsute to strigose or glabrate; stems in Arizona (Nesom and Baker, 1991) notes simple or branched; leaves oblanceolate to variability in habit, suggesting that a taxo- obovate or spatulate, entire to toothed or nomic review of this group would be useful. lobed, not clasping (slightly subclasping in E. Erigeron bigelovii should be investigated calcicola); heads nodding in bud; ray white for possible affinities elsewhere, possibly to pink, often with an abaxial lilac midstripe, with sect. Lamprocaules G.L. Nesom & R.D. not coiling or reflexing; achenes (0.5–)0.7– Noyes. The short-hispid vestiture, wiry, 1.6(–1.8) mm long, 2-nerved; pappus of 6– brittle, highly branched stems, and involu- 18(–19) bristles (16–25 in E. fundus, 23–30 cral morphology are unusual in sect. Olygo- in E. mihianus; bristles absent in E. ono- trichium. Erigeron mihianus also is somewhat frensis, E. versicolor). anomalous in sect. Olygotrichium, especially Group A. in its geography (Mt. Zempoaltepetl, Oax- E. bellidiastrum Nutt. aca) and high number of pappus bristles; on E. bigelovii A. Gray the other hand it produces leafy stolons and E. calcicola Greenm. nonclasping, pinnatifid leaves. E. calvus Coville E. divergens Torr. & A. Gray 21. Sect. Osteocaulis G.L. Nesom, Phy- E. flagellaris A. Gray tologia 67: 82. 1989. TYPE: E. fundus G.L. Nesom (Hook.) Piper E. lemmonii A. Gray Erigeron [unranked sp. group] Lutei E. metrius Blake Rydb., Fl. Rocky Mts. 897. 1918, in clave. E. mihianus Blake TYPE: Erigeron luteus A. Nelson (5 E. linearis) E. modestus A. Gray Taprooted, perennial; caudex branched; E. multiceps Greene stem vestiture of short, white, closely E. onofrensis G.L. Nesom appressed hairs; stems simple or less com- E. pinkavii B.L. Turner monly branched; leaves linear, entire, not E. pubescens Kunth clasping, leaf and stem bases white-indurate; E. religiosus Cronquist heads erect (E. linearis) or nodding (E. E. sionis Cronquist (incl. E. proselyticus bloomeri, E. elegantulus) in bud; rays coiling, G.L. Nesom) yellow in E. linearis, absent in E. bloomeri; E. tracyi (syn 5 E. colomexicanus A. achenes (1.8–)3–4 mm long, 2-nerved; pap- Nelson) pus of 10–50 bristles. 24 LUNDELLIA DECEMBER, 2008

E. barbellatus Greene Diplemium Raf., Fl. Tellur. 2: 50. 1836. E. bloomeri A. Gray LECTOTYPE (Nesom, 1989c): Erigeron strigo- E. elegantulus Greene sus Muhlenb. ex Willd. E. linearis (Hook.) Piper Erigeron [unranked sp. group] Ramosi Molecular data (Noyes, 2000a) suggest Rydb., Fl. Colorado 359. 1906, in clave. that sect. Osteocaulis, sect. Spathifolium, and TYPE: Erigeron ramosus (Walter) B.S.P. (5 E. sect. Arenarioides may be closely interrelated. strigosus Muhlenb. ex Willd) Species of sect. Arenarioides have white and Erigeron [unranked sp. group] Annui highly sclerified leaf bases similar to those of Small, Man. Southeastern Fl. 1395. 1933, in sect. Osteocaulis. All have similar vestiture, as clave. TYPE: E. annuus (L.) Pers. also does sect. Wyomingia. Annual to biennial or short-lived peren- 22. Sect. Pauciflori G. Don in Loudon, nial, fibrous-rooted, rhizomatous in some forms of E. strigosus; caudex simple; strigose Hort. Brit. 343. 1830. LECTOTYPE (Nesom 1989c): Erigeron bellidifolius Willd. (5 to sparsely hirsute; stems branched; leaves Michx.) oblanceolate, toothed, not clasping; heads Musteron Raf., Fl. Tellur. 2: 50. 1836. loosely paniculiform or corymbiform, nod- ding in bud; rays white, not coiling or TYPE:‘‘Erigeron bellidifolium’’ Raf. (probably 5 E. bellidifolius Willd. 5 E. puchellus reflexing; achenes (0.5–)0.8–1.2 mm long, Michx.) 2(–4)-nerved; pappus (disc florets) of 8–15 Erigeron [unranked sp. group] Pulchelli bristles, bristles absent (in E. annuus and E. Small, Man. Southeastern Fl. 1395. 1933, in strigosus) on ray florets. clave. TYPE: Erigeron pulchellus Michx. E. annuus (L.) Pers. Perennial, rhizomatous, fibrous-rooted, E. strigosus Muhlenb. ex Willd. primary rhizomes relatively slender, produc- E. tenuis Torr. & A. Gray ing slender, herbaceous, scale-leaved, stolo- Sect. Phalacroloma has been restricted niform rhizomes; caudex simple; villous; to two species that have pappose disc stems branched; leaves oblanceolate, achenes but that lack the series of bristles toothed, clasping; heads nodding in bud; on the ray achenes— and rays white, coiling tardily at tips; achenes E. annuus (Cronquist, 1947; Nesom, 1.3–1.8 mm long, 2(–4)-nerved; pappus of 1989c). Molecular data (Noyes, 2000a) (22–)28–36 bristles. place E. tenuis and E. strigosus as sister E. pulchellus Michx. species, with E. annuus sister to this pair. Erigeron pulchellus was earlier placed Occasional plants are encountered that within sect. Fruticosus (see commentary in suggest hybridization occurs between E. Nesom, 1989c) and the molecular analysis tenuis and E. strigosus. by Noyes shows it in an unresolved The reproductive biology and evolution position close to species of that section. of Erigeron strigosus and E. annuus are In view of its distinctive morphology and undergoing detailed study (Noyes and isolated geography, however, it seems Rieseberg, 2000; Noyes, 2000b; Noyes, reasonable to recognize the monotypic sect. 2005; Noyes and Allison, 2005; Noyes, Pauciflori, especially since it is not clear 2006a, 2006b; Noyes et al. 2007). Formal that sect. Fruticosus, as delimited here, is variants have recently been described at monophyletic. varietal rank within E. strigosus (Allison and Stevens, 2001; Noyes et al., 2006). 23. Sect. Phalacroloma (Cass.) Torr. & A. Gray, Fl. N. Amer. 2(1): 175. 1841. 24. Sect. Polyactis (Less.) G.L. Nesom, Phalacroloma Cass., Dict. Sci. Nat. 39: 404. Phytologia 66: 416. 1989. Polyactis Less., Syn. 1826. TYPE: Erigeron strigosus Muhlenb. ex gen. Comp. 188. 1832. TYPE: Erigeron Willd. delphinifolius Willd. NUMBER 11 NESOM: REVIEW OF CONYZINAE 25

Stenactis Cass., Dict. Sci. Nat. 37: 485. 25. Sect. Pseuderigeron Torr. & A. Gray, 1825 (non Erigeron sect. Stenactis Torr. & A. Fl. N. Amer. 2(1): 177. 1841. LECTOTYPE: Gray). TYPE: Erigeron delphinifolius Willd. Nutt. (Nesom, 1989c) Polyactidium DC., Prodr. 5: 281. 1836. Erigeron [unranked sp. group] Caespi- TYPE: Erigeron delphinifolius Willd. tosi Rydb., Fl. Colorado 359. 1906, in clave. Achaetogeron A. Gray, Mem. Amer. TYPE: Erigeron caespitosus Nutt. Acad. Arts, n. ser. 4 (Pl. Fendl.): 72. 1849. Perennial, taprooted; caudex branches TYPE: Erigeron wislizeni (A. Gray) Greene relatively thick and short, retaining old leaf Perennial and fibrous-rooted or tap- bases; stems simple, erect to decumbent- rooted and annual; caudex simple or rarely ascending; stems and leaves eglandular, branched, rhizomatous in some species; involucres glandular or eglandular; leaves hirsute to strigose; stems branched, less basal (persistent) and cauline, narrowly commonly simple; leaves mostly oblanceo- oblanceolate to spatulate (1–)3-nerved, cau- late to oblong, toothed or lobed to pinnat- line gradually reduced, not clasping; heads ifid, less commonly entire, clasping or not 1–4, erect? in bud; rays white to blue, clasping; heads usually in loosely corymbi- coiling, often weakly; achenes 1.5–2.2 mm form to paniculiform arrays, solitary in a few long, 2-nerved; pappus of 12–25 bristles. species, arching-nodding in bud; rays white E. abajoensis Cronquist to blue, filiform, reflexing; achenes 1– E. caespitosus Nutt. 1.3 mm long, 2–4-nerved; pappus bristles E. goodrichii S.L. Welsh 8–12 (to 17 in E. podophyllus), basally is a segregate of E. caducous, absent or greatly reduced in some caespitosus, separated from the latter only by species. its strigose (versus hirsute) stems and leaves E. annuactis G.L. Nesom and greater tendency for 1-nerved leaves. E. basaseachensis G.L. Nesom 26. Sect. Pycnophyllum Cronquist, Brit- E. caulinifolius G.L. Nesom tonia 6: 141. 1947. TYPE: Erigeron foliosus E. circulis G.L. Nesom Nutt. E. coroniglandifer G.L. Nesom Perennial, taprooted; short caudexlike E. dactyloides (Greenm.) G.L. Nesom branches or relatively slender, rhizomelike E. delphinifolius Willd. caudex branches producing numerous, ba- E. eruptens G.L. Nesom sally ascending stems, sometimes with slen- E. griseus (Greenm.) G.L. Nesom der, lignescent offsets; hirsute to strigose, E. inoptatus A. Gray puberulous, or glabrous; stems few- E. jenkinsii G.L. Nesom branched, rarely simple; leaves filiform to E. mayoensis G.L. Nesom linear-oblanceolate, entire, not clasping; E. nacoriensis G.L. Nesom heads 1 or 2–5(–10) in loosely corymbiform E. neomexicanus A. Gray arrays, erect in bud; rays white to blue, E. oreophilus Greenm. weakly coiling, sometime only at the tips, E. podophyllus G.L. Nesom sometimes apparently not coiling, absent in E. polycephalus (Larsen) G.L. Nesom some species; achenes 1.8–3 mm long, 2–4 E. rhizomactis G.L. Nesom (–5)-nerved; pappus of 17–45(–61 in E. E. seemannii (Schultz-Bip.) Greene reductus) bristles. E. subacaulis (McVaugh) G.L. Nesom E. aequifolius Hall E. wislizeni (A. Gray) Greene E. angustatus Greene Two species have been described since E. biolettii Greene the published revision of this section (Ne- E. blochmaniae Greene som, 1989a): E. jenkinsii (Nesom, 1993a) E. breweri A. Gray and E. mayoensis (Nesom, 1993b). E. elmeri (Greene) Greene 26 LUNDELLIA DECEMBER, 2008

E. foliosus Nutt. dryophyllus, E. veracruzensis; caudex simple; E. inornatus A. Gray strigose-pilose to hirsute, often deflexed on E. klamathensis (G.L. Nesom) G.L. the proximal portions of stem; stems Nesom branched; leaves oblanceolate, to elliptic, E. mariposanus Congdon entire or crenately toothed to pinnately E. miser A. Gray lobed, cauline clasping to subclasping or E. oxyphyllus Greene not; heads 1 or 2–35(–50) in diffuse arrays E. petrophilus Greene from distal branches, nodding or arching- E. reductus (Cronquist) G.L. Nesom nodding in bud; rays not coiling or reflexing, E. sanctarum S. Watson white to blue; achenes 0.6–1.8 mm long, 2- E. serpentinus G.L. Nesom nerved; pappus of 8–32 bristles. E. supplex A. Gray North American This group (as sect. Linearifolii) was E. basilobatus Blake (ne Mexico) revised by Nesom (1992a), bringing a E. cieloensis G.L. Nesom (ne Mexico) number of changes in circumscriptions of E. dryophyllus A. Gray (ne Mexico) taxa; one new species, E. serpentinus, was E. geiseri Shinners (central Texas and s added. Erigeron klamathensis was raised in ) rank (Nesom, 2004). Erigeron hyssopifolius E. heleniae G.L. Nesom (ne Mexico) was excluded by Nesom and Noyes (1999) to E. philadelphicus L. (e USA) constitute the monotypic sect. Linearifolii. E. quercifolius Lam. (e USA) and E. sanctarum are placed E. tenellus DC. (ne Mexico, s Texas) here in sect. Pycnophyllum for the first time, E. turnerorum G.L. Nesom (ne Mexico) although Cronquist (1947) earlier noted the E. veracruzensis G.L. Nesom (ne Mexico) resemblance of E. supplex to this group of Caribbean species and to a putative close relationship E. caeruleus Urb. (Hispaniola) between E. supplex and E. sanctarum. Sect. E. darrellianus Hemsl. (Bermuda) Pycnophyllum is centered in , E. dissectus Urb. (Hispaniola) extending northward to Washington and E. domingensis Urb. (Hispaniola) southward into ; E. oxyphyllus E. fuertesii Urb. (Hispaniola) is the southeasternmost species, occuring in E. jamaicensis L. (Jamaica) Sonora and Arizona. syn5 E. earlei Britton & P. Wilson (Cuba) 27. Sect. Quercifolium G.L. Nesom, syn5 E. polycladus Urb. (Martinique) sect. nov. syn5 E. rivularis Sw. (Jamaica, Hispa- Duratione annuo vel breviviventi, cau- niola) dice simplici, foliis serratis vel lobatis E. pinetorum Urb. (Hispaniola) (5 E. amplexicaulibus vel subamplexicaulibus, buchii Urban?) vestimento caulino deflexo, et radii non- E. polycladus Urb. (Martinique) cincinatis distinctus. TYPE: Erigeron querci- E. psilocaulis Urb. (Hispaniola) folius Lam. Some species of this group (Erigeron Erigeron [unranked sp. group] Philadel- quercifolius, E. philadelphicus, E. caeruleus, E. phici Rydb., Fl. Colorado 359. 1906, in clave dryophyllus, E. turnerorum, and E. tenellus) (non Small 1933). TYPE: Erigeron philadel- were placed by Nesom (1989c) within sect. phicus L. Olygotrichium, while another related group Annual or short-lived perennial, fi- (including E. darrellianus, E. jamaicensis, brous-rooted, taprooted in E. geiseri, E. and others) was noted as probably belonging tenellus, E. turnerorum, stoloniferous in E. in the same section. A new species was basilobatus, E. cieloensis, E. domingensis, E. described (Nesom, 1989e)—E. cieloensis, polycladus, E. psilocaulis, rhizomatous in E. putatively the sister species of E. basilobatus, NUMBER 11 NESOM: REVIEW OF CONYZINAE 27

which is transferred here from sect. Cincin- (except in E. rydbergii); stems simple, less nactis (sensu Nesom, 1989c). Erigeron hele- commonly branched; leaves oblanceolate to niae is recently described (Nesom, 2007). spatulate, entire, not clasping, abaxial faces Molecular data of Noyes (2000a) indicate glabrous or glabrate, adaxial faces strigose to that sect. Quercifolium is phyletically coor- villous (both faces hairy in E. parryi); heads dinate with the clade that includes Aphano- 1(–3), behavior in bud not known, phyllaries stephus and Erigeron sect. Microcephalum. minutely glandular; rays coiling (not coiling Sect. Quercifolium, as delimited here, is in E. parryi); achenes 1.8–2.8 mm, 2-nerved; the only section of Erigeron with a distribu- pappus of 12–20 bristles (caducous in E. tion divided between the Caribbean, Mexico, parryi, E. radicatus). and the . The Caribbean species E. lackschewitzii G.L. Nesom & Weber (type localities in parenthesis) are closely E. ochroleucus Nutt. (incl. E. scribneri similar to E. quercifolius: E. caeruleus, E. Canby ex Rydb.) dissectus, E. fuertesii (perhaps conspecific E. parryi Canby & Rose with E. caeruleus), and E. jamaicensis. E. radicatus Hook. (incl. E. huberi S.L. Erigeron darrellianus apparently is the only Welsh & N.D. Atwood) species of the genus E. rydbergii Cronquist to have developed a distinctly shrubby habit. Erigeron parryi differs from the other Erigeron domingensis is remarkably similar in species in its apparently non-coiling rays and habit to E. flagellaris (of sect. Olygotrichium, leaves equally strigose to hirsute-strigose on with herbaceous stolons, tendency for a both faces; like most of the others, however, single head on the erect stems), but E. it produces hairs with purplish cross walls domingensis has more herbaceous phyllaries, and the pappus bristles are basally caducous, white to blue or purple rays without an as also in E. radicatus (Nesom, 2004). This abaxial midstripe, and up to 3 capitula per group was placed in sect. Asteroidea in 1989 stem, probably derived from its relatives as the ‘‘E. radicatus group.’’ with a corymbiform capitulescence. 29. Sect. Rhizonexus G.L. Nesom, sect. , previously regarded as nov. closely related to E. geiseri (Nesom, 1987, Distinctus habitu fibrosi-radicato rhizo- 1989e), is placed in the present treatment mato, caudex systemata diffusa ramorum with sect. Phalacroloma. Erigeron pinetorum tenuium rhizomiformium formans. TYPE: Urb. was previously placed as a close relative D. Eaton of the species related to E. quercifolius but is Perennial, rhizomatous, fibrous-rooted, now known to be conspecific with Conyza forming diffuse systems of slender, rhizome- primulifolia (Lam.) Cuatrec. & Lourteig (5 like caudex branches; glabrous or sparsely Conyza chilensis Spreng.). strigose; stems simple or few-branched; 28. Sect. Radicati (Rydb.) Nesom, leaves narrowly oblanceolate to oblong, comb. et stat. nov. entire, not clasping; heads 1(–3), erect in Erigeron [unranked sp. group] Radicati bud; rays blue to pink or purple, not Rydb., Fl. Colorado 359. 1906, in clave. reflexing or coiling or sometimes coiling TYPE: Hook. slightly at the tips; achenes 1.5–2 mm long, Erigeron [unranked sp. group] Laetevir- 2-nerved; pappus of 10–21 bristles. entes Rydb., Fl. Rocky Mts. 897. 1918, in E. gracilis Rydb. clave. TYPE: Erigeron laetevirens Rydb. (5 E. E. ursinus D. Eaton ochroleucus) The morphological similarity of Erigeron Perennial, taprooted; caudex multicipi- gracilis and E. ursinus and their apparent tal or with short, thick branches; hirsutulous morphological isolation as a group were to villosulous, hairs with purplish cross walls noted by Cronquist (1947) and by Nesom 28 LUNDELLIA DECEMBER, 2008

(1989c), where they were treated as the ‘‘E. The Noyes analysis (2000a) suggests that ursinus group’’ of sect. Asteroidea. Erigeron E. pinnatisectus is closely related to sect. leiomerus (treated here within sect. Aster- Scopulincola, where its pinnatifid leaves are igeron) was placed by molecular data (Noyes, similar to those of E. pringlei, rather than to 2000a) in a sister relationship to E. ursinus, the 3-parted ones of sect. Tridactylia. but E. gracilis was not included in the , closely similar to E. molecular sampling. pinnatisectus, is also transferred here from sect. Tridactylia. Both species also have 30. Sect. Scopulincola Nesom, Phytologia folding leaves and reflexing rays like others 67: 84. 1989. TYPE: G.L. of sect. Scopulincola. Nesom & Roth Perennial, taprooted; caudex branches 31. Sect. Stenactis Torr. & A. Gray, Fl. N. long, slender, rhizomelike; glabrous or Amer. 2(1): 172. 1841. LECTOTYPE (Nesom nearly so; stems simple or branched; leaves 1989c): Nutt. oblanceolate to obovate or spatulate, entire Erigeron [unranked sp. group] Pumili or pinnatisect in some, folding along the Rydb., Fl. Colorado 359. 1906, in clave. midvein, not clasping; heads 1–4, nodding in TYPE: Erigeron pumilus Nutt. bud, phyllaries graduate and densely glan- Perennial, taprooted; caudex branches dular but otherwise glabrous to sparsely not thickened, leaf bases not marcescent; strigose, often purplish; rays white, reflexing petiole bases ciliate; heads commonly more (absent in E. mancus); achenes 1.2–2.3 mm than 1 per stem, nodding in bud; leaves long, 2-nerved; pappus of 11–30 bristles. linear-oblanceolate to narrowly oblanceolate Group A. Leaves spatulate, entire; Ar- or oblanceolate-spatulate, 1-nerved, cauline izona, New Mexico, Colorado, Utah, Ne- well-developed (except var. condensatus); vada, Montana, . rays reflexing (laminae reduced in E. apha- E. kachinensis S.L. Welsh & Moore nactis, E. calvus, E. concinnus; ray florets 0 in E. leiomerus A. Gray E. ovinus); disc corolla throat white-indurate E. nauseosus (M.E. Jones) A. Nelson and inflated (not inflated in E. pygmaeus); E. scopulinus Nesom & Roth achenes 1.2–2 mm long (2.3–2.8 mm in E. Group B. Leaves mostly toothed to pygmaeus); pappus bristles 7–27. pinnatifid; Arizona. E. aphanactis (A. Gray) Greene E. anchana G.L. Nesom E. calvus Coville E. heliographis G.L. Nesom E. clokeyi Cronquist E. pringlei A. Gray E. concinnus (Hook. & Arn.) Torr. & A. Gray E. saxatilis G.L. Nesom E. engelmannii A. Nelson Group C. Leaves pinnatisect; high ele- E. maguirei Cronquist vation in New Mexico, Colorado, Utah, E. pumilus Nutt. (incl. E. zothecinus S.L. Wyoming. Welsh) E. mancus Rydb. E. pygmaeus (A. Gray) Greene E. pinnatisectus (A. Gray) A. Nelson E. vetensis Rydb. Sect. Scopulincola (especially Group A) This group was placed by Nesom is distinguished from sect. Asterigeron pri- (1989c) as the ‘‘Erigeron pumilus group’’ of marily on the basis of the reflexing ray sect. Asteroidea. It was later treated with behavior. The toothed to pinnatifid leaves of other species with sharply reflexing rays as Groups B and C are a further distinction. A an informal subgroup within sect. Genicu- recent study of the Erigeron. pringlei group lactis (Nesom, 1990c). Molecular data, in (Nesom, 1990b) resulted in the description contrast, indicate that the E. pumilus group of three additional species related to E. is phylogenetically removed from sect. pringlei, all separated here as Group B. Geniculactis sensu stricto, and the distinctive NUMBER 11 NESOM: REVIEW OF CONYZINAE 29

morphology of the group suggests that it be Sect. Terranea comprises a group of six separately and formally recognized. The apparently closely related species endemic to species occur at low to mid-level elevations the Juan Fernandez Islands of Chile and are in the western U.S.A. Erigeron pygmaeus hypothesized (based on an analysis of grows at higher elevations and is anomalous morphological, chromosomal, and flavonoid in several morphological features. data) to have radiated from a single The narrow Utah endemic Erigeron introduction to the island more distant from maguirei fits within sect. Stenactis in features the mainland (Isla Masafuera; Valdebenito of habit as well as ray and bud behavior. et al., 1992). The epithet ‘‘stuessyi’’ was Among the other species, it is most similar applied to one of the species in the in its subspatulate leaves to the rare E. calvus publication by Valdebenito et al. (1992), from the Inyo Mountains of California. but the name has never been validated. The placement here of Among South American mainland spe- emphasizes the tendency for reduction or cies included in a survey for possible loss of rays prevalent in the section. Bud ancestors, the Valdebenito et al. (1992) study behavior is not known, and the 3-nerved placed Erigeron leptorhizon DC. (2n 5 18) leaves and deflexed-hirsute cauline vestiture (see Conyza group A 5 Erigeron sect. suggest that it may instead be related to E. Caenotus) as the species most similar to the caespitosus (sect. Asteroidea, Group B) (Blake E. fernandezianus group, but the molecular first described E. ovinus as E. caespitosus var. analysis of Noyes (2000a) instead places E. anactis). It also is similar in some respects to fernandezianus as closely related to the other E. nauseosus (sect. Scopulincola, Group A). group of species commonly placed in Conyza Erigeron ovinus is endemic to the Sheep and (see Conyza group B), distantly related to E. Groom ranges and Mount Irish in Clark and leptorhizon. As noted by Valdebenito et al. Lincoln cos., Nevada (1992, p. 477), ‘‘Continental species with a chromosome number of n 5 27 would be 32. Sect. Terranea (Colla) Nesom, more probable as an ancestral taxon.’’ Plants comb. et stat. nov. of the mainland species E. ecuadoriensis and Terranea Colla, Mem. Accad. Sci. Tor- E. lanceolatus (see sect. Meridionales) are ino 38 (Pl. Rar. Chil., Fasc. 4): 11, t. 23. 1835 known to be hexaploid, and it seems likely [reprints 1834]. TYPE: Terranea fernandezi- that the Juan Fernandez endemics are derived ana Colla from sect. Meridionales. Perennial herbs to subshrubs or (Erig- The Noyes (2000a) molecular analysis eron fernandezianus) woody shrubs or small included two samples of Erigeron rosulatus: trees, rhizomatous; caudex simple; glabrous the mainland sample is closely related to to strigose; stems branched; leaves oblan- other mainland species (here in sect. Mer- ceolate, to lanceolate, entire to toothed or idionales), while the sample from the Juan pinnatifid, not clasping; heads 1 or in a Fernandez Islands clusters with the species of loosely corymbiform array, erect in bud; sect. Terranea. rays, white, not coiling or reflexing; achenes 1–2 mm long, 2-nerved; pappus of [not 33. Sect. Tridactylia Nutt., Trans. Amer. counted] bristles. All species are hexaploid, Philos. Soc. ser. 2, 7: 310. 1841. TYPE: 2n 5 54 (Valdebenito et al. 1992). Pursh. E. ingae Skottsb. Erigeron series Multifidi Rydb. ex J.F. E. luteoviridis Skottsb. Macbr. & Payson, Contr. Gray Herb. 49: 73. E. fernandezianus (Colla) Solbrig 1917. Erigeron [unranked sp. group] Multif- E. rupicola Phil. idi Rydb., Fl. Colorado 359. 1906, in clave. E. ‘‘stuessyi’’ (ined.) LECTOTYPE: Erigeron compositus Pursh. (Ne- E. turricola Skottsb. som, 1989c) 30 LUNDELLIA DECEMBER, 2008

Erigeron [unranked sp. group] Compo- E. lanatus at the base of the caudex, but this siti Rydb., Fl. Rocky Mts. 896. 1918, in clave. may be a rhizome; the diffuse system of TYPE: Erigeron compositus Pursh. relatively long, slender, rhizomelike, branch- Taprooted, perennial; caudex divided es is similar to morphology in sect. Tridac- into slender, rhizomelike branches (variable tylia. in Erigeron compositus); glabrous to strigose or hispid; stems simple (few-branched in E. 34. Sect. Trimorpha (Cass.) DC., Prodr. basalticus and E. allocotus); leaves oblanceo- 5: 290. 1836. Trimorpha Cass., Bull. Sci. Soc. late to spatulate 3-lobed or 3-toothed near Philom. Paris 1817: 137. 1817. Erigeron subg. the apex, not clasping; heads erect in bud; Trimorpha (Cass.) Popov, Acta Inst. Bot. rays white, weakly coiling or not at all Acad. Sci. URSS, Ser. 1, Fasc. 7: 10. 1948. (sometimes distinctly in E. compositus); TYPE: Erigeron acris L. achenes 1.5–2.7 mm long (3.5–4 mm in E. Trimorpha sect. Brachyglossae Vierh., lanatus), 2-nerved; pappus of 10–20 bristles Beih. Bot. Centralbl. 19: 423. 1906. Erigeron (25–35 in E. lanatus). subsect. Brachyglossae (Vierh.) Pawlowski, E. allocotus Blake Fragm. Florist. Geobot. 16: 256. 1970. E. basalticus Hoover LECTOTYPE (Nesom, 1989b): Erigeron acris L. E. compositus Pursh Trimorpha sect. Macroglossae Vierh., E. denalii A. Nelson (incl. E. mexiae K. Beih. Bot. Centralbl. 19: 424. 1906. LECTO- Becker) TYPE (Nesom, 1989b): L. E. flabellifolius Rydb. Tessenia P. Bubani, Fl. Pyrenaea 2: 264. E. lanatus Hook. 1899. LECTOTYPE: Erigeron alpinus L. (Ne- E. pallens Cronquist som, 1989b) E. purpuratus Greene Erigeron [unranked sp. group] Acres E. salishii Douglas & Packer Rydb., Fl. Colorado 359. 1906, in clave. E. trifidus Hook. TYPE: Erigeron acris L. E. vagus Payson Fibrous-rooted, perennial, biennial, or Recent studies have explored the nature annual; caudex usually simple; stems usually of variation in sexual and agamospermic branched; leaves mostly entire, not clasping; populations of E. compositus (Noyes and heads erect in bud, in a corymbiform to Soltis, 1996; Noyes et al., 1995). Erigeron broadly racemiform cluster; inner involucral pinnatisectus was shown by Noyes (2000a) to long-attenuate to caudate; ray (pistil- be distantly related to sect. Tridactylia; E. late) florets in 2 zones, those of the outer pinnatisectus and its close relative E. mancus, zone with erect, filiform, white to pink or both of which have pinnately dissected purplish lamina, not coiling or reflexing, leaves (vs 3-lobed or 3-toothed), are here those of inner zone without lamina; achenes transferred to sect. Scopulincola. 2–2.4 mm long, 2-nerved; pappus bristles A possible relationship of Erigeron 12–35, accrescent (elongating in fruit to ca. 2 lanatus to sect. Erigeron needs to be times involucres). investigated; the leaves are entire or weakly E. acris L. * (incl. E. kamtschaticus DC., 3-toothed at the apex, and the species was E. angulosus Gaudichaud*, E. droeba- speculated to be involved as a parent in the chiensis O.F. Mu¨ll.*, E. elongatus hybrid origin of E. trifidus (Packer, 1983). Ledeb., and E. politus Fries*) The origin of E. trifidus through hybridiza- E. alpinus L.* tion between E. lanatus and E. compositus E. atticus Villars* was confirmed by Burke (2005), although as E. baicalensis Botsch. currently recognized, E. trifidus probably is E. borealis (Vierh.) Simmons* polyphyletic. An ambiguous, slender tap- E. canariensis O. Sida rootlike structure sometimes is observed in E. caucasicus Steven* NUMBER 11 NESOM: REVIEW OF CONYZINAE 31

E. cyanactis Rech. f. (Vierh.) Pawl., E. neglectus, E. orientalis Boiss., E. elatus Hook. and E. rhodopaeus (Vierh.) Hayek. E. epiroticus (Vierh.) Hala´csy* Sida (1988) formally divided sect. Tri- E. glabratus Hoppe & Hornsch. ex Bluff morpha into three series, based primarily on & Fingerh.* form of the capitulescence and vestiture: ser. E. granatensis Lippert Trimorpha O. Sida, ser. Politae O. Sida, and E. hispanicus (Vierh.) Maire ser. Macrophyllae O. Sida. E. lalehzaricus (Rech.) O. Sida As suggested by Nesom (1994c) and as E. leptocladus Rech. f. discussed in detail and experimentally con- E. macrophyllus Herbich firmed by Huber and Nilsson (1995) and E. mesatlanticus Maire Noyes (2000a), autogamous breeding sys- E. nannogeron Rech. f. tems apparently have arisen independently E. neglectus Kerner* in a number of Astereaean groups, including E. nivalis Nutt. (5 E. debilis (A. Gray) Trimorpha and Conyza. In these, the pistil- Rydb.; E. scotteri Boivin) late florets of a head are greatly increased in E. orientalis Boiss. number (often outnumbering the bisexual E. podolicus Besser florets), in several series, the inner some- E. pseudoelongatus Botsch. times reduced to filiform eligulate florets, E. semibarbatus DC. and the outer with reduced lamina. E. serotinus Weihe E. uralensis Less. 35. Sect. Wyomingia (A. Nelson) Cron- E. venustus Botsch.* quist, Brittonia 6: 140. 1947. (*asterisks denote Eurasiatic species Wyomingia A. Nelson, Bull. Torrey Bot. studied by Huber and Nilsson, 1995) Club 26: 249. 1899. TYPE: Erigeron pulcher- Nesom (1989b) hypothesized that sect. rimus A.A. Heller Trimorpha (as the genus Trimorpha)is Erigeron [unranked sp. group] Cani separate from Erigeron, more closely related Rydb., Fl. Colorado 359. 1906, in clave. to Conyza, and provided nomenclatural TYPE: A. Gray transfers for North American taxa. Studies Erigeron [unranked sp. group] Tetra- by Walter Huber and colleagues and by pleuri Rydb., Fl. Rocky Mts. 897. 1918, in Noyes (2000a), however, have made it clear clave. TYPE: Erigeron tetrapleuris (A. Gray) that species of sect. Trimorpha are closely Heller (5 E. utahensis) related to those of sect. Erigeron and that Taprooted; perennial; caudex simple or both sections are relatively recently derived with short, thick branches; vestiture strigil- within the genus (Huber, 1993; Huber and lose of short, closely appressed hairs; stems Leuchtmann, 1992; Huber and Nilsson, simple or branched; leaves linear to narrowly 1995; Huber and Zhang, 1991; Oberha¨nsli oblanceolate, not clasping; heads erect in and Huber, 1993; Utelli et al., 1995). bud (nodding? in ); rays The North American species are Erigeron white to blue, coiling (reflexing in E. canus, acris (circumboreal), E. elatus,andE. nivalis. E parishii); achenes (1.5–)2–3.4 mm long, 2- The Eurasiatic species are diploids with nerved, (2–)4(–6)-nerved, or 6–8-nerved, identical karyotypes and their high genetic (8–)10–14-nerved (E. canus), strigose to identities suggest very recent evolutionary strigose-sericeous or with velutinous-ciliate differentiation. Some of these are among nine margins and glabrous faces; pappus of 15–50 species listed by Sida (1988) as ‘‘Species bristles (9–10 in E. katiae). excludendae’’—he gave no comment about Group A. Stems branched; cauline leaves the reason for their exclusion or their suggested prominent; heads numerous, erect in bud; disposition: E. alpinus, E. atticus, E. borealis, E. rays coiling; achenes (2–)4(–6)-nerved, epiroticus, E. gaudinii Bru¨gger, E. hungaricus sparsely strigose. 32 LUNDELLIA DECEMBER, 2008

E. sparsifolius Eastw. in discerning relationships among the sub- E. utahensis A. Gray groups also were pointed out in the original Group B. Stems simple, heads 1; leaves description of E. sivinskii (Nesom, 1991). mostly basal and linear, cauline reduced; Studies by Tonne (1999) indicate that heads 1; achenes 2-nerved, faces glabrous, Erigeron bistiensis is a synonym of E. margins pilose-sericeous. pulcherrimus and support the hypothesis E. compactus Blake (Nesom and Hevron, 1995) that E. unter- E. consimilis Cronquist mannii and E. carringtonae are conspecific. E. sivinskii G.L. Nesom Neel and Ellstrand (2001) studied patterns Group C. Stems simple or branched, of allozyme diversity in E. parishii, which is heads 1–4; leaves basal and cauline; achenes threatened by mining. Distinctions between 2-nerved, villous-hirsute; pappus of 9–10 (E. E. utahensis and E. sparsifolius were noted by katiae) or 15–25 (E. nematophyllus) bristles. Nesom (2004). E. katiae N.D. Atwood and S.L. Welsh (2007) APHANOSTEPHUS DC., Prodr. 5: 310. E. nematophyllus Rydb. 1836. TYPE: Aphanostephus ramosissimus Group D. Stems simple; leaves mostly DC. basal and linear, cauline reduced; heads 1; Keerlia DC., Prodr. 5: 309. 1836. LEC- achenes 2-nerved ()or TOTYPE (designated here): Keerlia ramosa 4–8-nerved, faces strigose-sericeous. DC. 5 Aphanostephus ramosissimus var. E. argentatus A. Gray ramosus (DC.) B.L. Turner & Birdsong E. pulcherrimus A.A. Heller (incl. E. Leucopsidium Charpentier ex DC., bistiensis G.L. Nesom & Hevron) Prodr. 6: 43. 1838. TYPE: Leucopsidium E. untermannii S.L. Welsh (incl. E. arkansanum DC. 5 Aphanostephus skirrho- carringtonae S.L. Welsh) basis (DC.) Trel. Group E. Stems simple or branched; Aphanostephus subg. Euaphanostephus leaves linear, mostly basal (Erigeron canus) Blake, Contr. Gray Herb. 53: 24. 1918. TYPE: or basal and cauline (E. parishii); heads 1–4 A. ramosissimus DC. (E. canus) or 1–5(–10) (E. parishii), nodding Aphanostephus subg. Pappophanus in bud; rays reflexing (also slightly coiling at Blake, Contr. Gray Herb. 53: 24. 1918. TYPE: tips in E. parishii); achenes (8–)10–14- A. skirrhobasis var. hallii (A. Gray) Blake nerved, terete, glabrous (E. canus)or4- Aphanostephus sect. Pappophanus (Blake) nerved, angled, sparsely strigose to pilose (E. Shinners, Wrightia 1: 101. 1946. parishii). Aphanostephus sect. Pappopecus Shin- E. canus A. Gray ners, Wrightia 1: 108. 1946. LECTOTYPE: A. E. parishii A. Gray ramosissimus DC. (designated here) Sect. Wyomingia, as outlined here, is a Annual or perennial, taprooted; caudex heterogeneous group united by strigillose simple, stems branched; leaves oblanceolate, vestiture, linear leaves, and coiling rays slightly clasping or not clasping; heads (except in Group E). Achenes tend to be nodding in bud; rays with abaxial midstripe, multi-nerved in Groups A and E and densely not coiling or reflexing; achenes thick- hairy in Groups B, C, and D. The leaves of ribbed, columnar, 4-angled, often bearing Erigeron untermannii are often spatulate and duplex trichomes with a coiled or recurved glabrous to glabrate abaxially, features more apex; pappus a low, fimbriate- or ciliate- similar to E. radicatus and its close relatives; rimmed corona (in A. skirrhobasis var. rays of E. untermannii apparently tend to kidderi (Blake) B.L. Turner, the pappus both coil and reflex. The two species of members may appear as relatively slender, Group E are unique in their combination of barbellate bristles with expanded bases). The nodding heads and reflexing rays. Problems genus is distinguished by its combination of NUMBER 11 NESOM: REVIEW OF CONYZINAE 33

ovate-lanceolate, thin-margined phyllaries, with 3 parallel nerves, entire, not clasping; conical receptacles, achene morphology, heads 1–4 or 15–75 in a thyrsoid-cormbiform pappus, and base chromosome number of arrangement, erect in bud; rays absent (pe- x 5 5. ripheral florets pistillate, fertile, tubular with 3– A. pilosus Buckley 4 apical teeth); achenes 1.8–4 mm long, A. ramosissimus DC. (incl. A. arizonicus subcylindric, with (4–)5(–6) orange-resinous A. Gray, A. humilis (Benth.) A. Gray, nerves; pappus of 28–40 bristles. A. jaliscensis Shinners, A. pachyrrhizus A. corymbosus (Hook. & Arn.) Nesom Shinners, A. potosinus Shinners, and A. warmingii (Baker) Nesom A. ramosus (DC.) A. Gray) Apopyros includes two species mostly in A. riddellii Torr. & A. Gray Brazil but reaching Paraguay and northeastern A. skirrhobasis (DC.) Trel. (incl. A. Argentina. Nesom (1994c) observed that kidderi Blake) Apopyros is similar to Neja in its parallel- Aphanostephus includes four species veined leaves with shiny-indurate surfaces, (including eight varietal taxa) restricted to multinerved and nearly terete achenes, and the south-central USA and the northern half tendency to produce stiffly pilose-hirsute of Mexico, as recognized by Turner (1984). vestiture, but Noyes (2000a) found that Turner’s phylogenetic reconstruction of the Apopyros is more closely related to Lepto- genus, based on morphology, was largely stelma. corroborated by Elisens et al. (1992). CONYZA [group A] Compared to Aphanostephus, other spe- Erigeron sect. Caenotus Nutt., Gen. cies of Conyzinae are conservative in involu- . 2: 148. 1818. Conyza sect. Caenotus cral morphology (phyllaries linear-lanceolate, (Nutt.) Cronquist ex Cuatrec., Webbia 24: without wide, thin-scarious margins), achene 211. 1969. TYPE: L. morphology and vestiture (thin-walled, flat, Conyzella Fabr., Enum. (ed. 1) 86. 1759. mostly 2-nerved but subterete and multi- TYPE: Erigeron canadensis L. nerved in a few taxa, the duplex hairs never Leptilon Raf., Amer. Monthly Mag. 268. glochidiate), pappus (of barbellate bristles or 1818. TYPE: Erigeron divaricatus Michx. reduced), and base chromosome number (x 5 Erigeron sect. Multiflori G. Don in 9) (see Nesom, 2000a). The genus was Loudon, Hort. Brit. 343. 1830. LECTOTYPE hypothesized to be closely related to Town- (Nesom 1989c): Erigeron canadensis L. sendia, , and a group of Southern Suffrutescent perennial (Erigeron crena- Hemisphere genera (subtribe Brachyscomi- tus, E. socorrensis) or annual; taprooted; nae) on the basis of morphological features caudex simple; hirsute to nearly glabrous; (Nesom, 1994d). In contrast, molecular data stems branched; leaves linear to lanceolate or (cf. Morgan, 1990; Noyes, 2000a; Noyes and oblanceolate, entire or toothed, not clasping; Rieseberg, 1999) indicate that Aphanostephus heads erect in bud, in a columnar to has arisen from within the Conyzinae, appar- pyramidal or loosely corymbiform arrange- ently closely related to sect. Microcephalum and ment; rays white, filiform, lamina slightly sect. Cincinnactis. longer than the involucre, not coiling APOPYROS Nesom, Phytologia 76: 177. (apparently coiling in E. socorrensis)or 1994. TYPE: Apopyros warmingii (Baker) G.L. reflexing; achenes 0.8–1.5 mm long, 2- Nesom nerved; pappus of 11–20 bristles. Perennial, from a thick, ligneous, some- Erigeron canadensis L. (5 Conyza cana- times tuberous rhizome; caudex simple; gla- densis (L.) Cronquist) brate to sparsely pilose-hirsute; stems simple or Erigeron crenatus Eastw. branched; leaves all cauline, the proximal Erigeron divaricatus Michx. (5 Conyza scalelike, linear-lanceolate to elliptic above, ramosissima Cronquist) 34 LUNDELLIA DECEMBER, 2008

Erigeron leptorhizon DC. (5 E. gaudi Conyza microcephala Hemsl. chaudii DC.) Erigeron pazensis Schultz-Bip. ex Rusby Erigeron socorrensis I.M. Johnston Erigeron pinnatus Turcz. 1851 (non L. Erigeron subalpinus Urb. 1753) (5 Conyza cardaminifolia Variation in Erigeron crenatus and E. Kunth; E. cardaminifolia (Kunth) socorrensis, from Isla Socorro of the Revilla- Wedd.; E. turczaninowii Wedd.) gigedo Islands, Mexico, was studied by Erigeron spiciformis Griseb. (5 Conyza Nesom (1990h). Erigeron subalpinus is spiciformis (Griseb.) Zardini) endemic to Cuba and was described as very Erigeron spiculosus Hook. & Arn. (5 similar to Erigeron canadensis. Conyza spiculosa (Hook. & Arn.) Zardini) CONYZA [group B] Erigeron stenophyllus Hook. & Arn. (5 Conyza Less., Syn. Gen. Comp. 203. Conyza gayana Phil.) 1832. Erigeron sect. Conyza (Less.) Baill., Retz. (if distinct Hist. Pl. 8: 143. 1882. TYPE: Conyza chilensis from E. floribundus; 5 Conyza suma- (Spreng.) d. Don ex D. Don 5 Conyza trensis (Retz.) E. Walker) primulifolia Lourteig & Cuatrec. Erigeron trihecatactis Blake (5 Conyza Astradelphus E.J. Remy [nom. nov.], Ann. trihecatactis (Blake) Cuatrec.) Sci. Nat. Bot. ser. 3, 12: 185. 1849. Gusmania E.J. Erigeron tunariensis Kuntze (5 Conyza Remy in C. Gay, Hist. Chile Bot. [Fl. Chilena] 4: tunariensis (Kuntze) Zardini) 12. 1849 (non Guzmania Ruiz & Pavon 1802). Erigeron uliginosa Benth. (5 Conyza TYPE: Astradelphus chilensis (Spreng.) E.J. Remy 5 Gusmania chilensis (Spreng.) E.J. Remy 5 uliginosa (Benth.) Cuatrec.) 5 Conyza chilensis Spreng. 5 Conyza primulifolia Erigeron variifolius Blake ( Conyza (Lam.) Cuatrec. & Lourteig 5 Erigeron chilensis coronopifolia Kunth) (Sprengel) D. Don ex G. Don in Loudon 5 Erigeron pazensis usually is not included Erigeron remyanus Wedd. among these species, but the habit and Annual to perennial; taprooted; caudex general aspect suggest that it belongs here. simple; hirsute to strigose, often strongly Solbrig (1962, p. 37) noted that it is ‘‘A glandular; stems simple or branched; leaves species morphologically similar to Conyza,it lanceolate to oblanceolate or spatulate, entire to is undoubtedly an Erigeron due to the toothed or lobed, clasping or not clasping; heads preponderance of tubular over ligulate erect in bud; rays white, lamina absent to slightly , and the ligules, which are clearly longer than the involucre; achenes 0.8–2 mm longer than the tubes.’’ long, 2-nerved; pappus of ca. 10–25 bristles. An accurate estimate of the number of Conyza arabidifolia E.J. Remy South American species generally treated as L. (5 Conyza bo- Conyza has not been developed, but there nariensis (L.) Cronquist) probably are between 60 and 100 (e.g., see Zardini, 1976). Relatively few of the species Conyza burkartii Zardini are included here. Erigeron chilensis (Spreng.) D. Don ex G. Cronquist (1943), Cuatrecasas (1969), Don in Loudon (5 Conyza primuli- Zardini (1976), Nesom (1990a), and The´baud folia (Lam.) Lourteig & Cuatrec. 5 and Abbott (1995) have provided suggestions Conyza chilensis Spreng.) toward a distinction between Conyza and Conyza deserticola Phil. Erigeron. Group B of Conyza is heterogeneous Erigeron floribundus (Kunth) Schultz- and likely will prove to be non-monophyletic. Bip. (5 Conyza floribunda Kunth) Erigeron laevigatus Rich. (5 Conyza laevi- DARWINIOTHAMNUS Harling, Acta Horti gata (Rich.) Pruski 5 Conyza apurensis Berg. 20: 108. 1962. TYPE: Darwiniothamnus Kunth) tenuifolius (Hook. f.) Harling NUMBER 11 NESOM: REVIEW OF CONYZINAE 35

Perennial shrubs, fibrous-rooted; caudex sometimes branched at the base; leaves mostly simple or branched; glabrous to strigose; oblanceolate, entire to pinnatisect, not clasping; leaves linear to oblanceolate, entire to mu- heads 1-several, erect in bud; rays yellow or cronate, in condensed terminal whorls; stems white, not coiling or reflexing; achenes 1–2 mm branching; heads in a condensed-corymbi- long, 2-nerved; pappus of scabrid bristles. form capitulescence mostly within the leaves; H. aberrans (Cabrera) Cabrera rays white, strongly coiling; achenes dimor- H. bakeri Hicken phic, ray 2(–4)-nerved, disc 3–5(–6)-nerved. H. cabrerae Espinar D. tenuifolius (Hook. f.) Harling (5 H. glaucifolia (O. Kuntze) Solbrig Erigeron tenuifolius Hook. f.) H. jasionioides Willd. D. lancifolius (Hook. f.) Harling (5 H. montevidensis Baker Erigeron lancifolius Hook. f.) H. nebularis Deble, Oliveira & Marchiori Darwiniothamnus, an endemic of the H. pinnatiloba Matzenb. & Sobral Gala´pagos Islands, has been treated at H. pinnatisecta Matzenb. & Sobral generic rank by various authors (e.g., Har- H. pulchella Cabrera ling, 1962; Cronquist, 1971; Lawesson and Overviews of Hysterionica have been pro- Adsersen, 1987). Infraspecific taxa have been vided by Cabrera (1946), Ariza Espinar (1980), described in each of the two species. and Nesom (1993c, 1994b). Three species from Darwiniothamnus alternifolius,the Brazil have recently been added (Matzenbacher most recently described species (Lawesson and Sobral, 1996; Deble et al., 2004). and Adsersen, 1987), does not have signif- LEPTOSTELMA D. Don in Sweet, Brit. Fl. icant features of the two species upon Gard. ser. 2. 38. 1830. TYPE: Erigeron maximus which the generic concept was based: the Erigeron sect. Leptostelma (D. Don) plants lack the shrubby ‘‘rosette’’ habit, the Benth. & Hook., Gen. Pl. 2: 280. 1873. leaves are not in condensed terminal Perennial herbs, rhizomatous and fibrous- whorls, the heads are not in a condensed- rooted; caudex simple; glabrous to puberulent corymbiform capitulescence mostly within or hispid; stems branched; leaves ovate to the leaves, and the achenes are not dimor- oblanceolate, net-veined, entire or toothed, phic. Further, the ray floret lamina do not clasping; heads erect in bud; rays yellow to coil and molecular evidence indicates that white, not coiling or reflexing; achenes 2(–3)- its closest relationships are elsewhere (An- nerved, glabrous; pappus of 30–40 bristles. drus et al., in press); it belongs among the Erigeron camposportoi Cabrera species of Erigeron sect. Meridionales, Erigeron catharinensis Cabrera where it is listed here. Leptostelma maximum D. Don (5 Nesom (1989c) suggested that Darwi- Erigeron maximus (D. Don) DC.) niothamnus is related to the primarily Erigeron meyeri Cabrera Mexican group Erigeron sect. Cincinnactis. Erigeron tucumanensis Cabrera Recent molecular evidence (Noyes, 2000a; Leptostelma tweediei (Hook. & Arn.) Hind Andrus et al., in press.) shows that the two &G.L.Nesom(5 Erigeron tweediei Darwiniothamnus species are most closely Hook. & Arn.; Erigeron seneciiformis related to sect. Cincinnactis and the Carib- Blake) bean sect. Microcephalum. The remaining combinations to Lepto- stelma are being made by Aristoˆnio Teles in HYSTERIONICA Willd., Ges. Naturfr. connection with his synopsis of the tribe Freunde Berlin Mag. 1: 140. 1807. TYPE: Astereae in Brazil. Hysterionica jasionioides Willd. Annual or perennial herbs to subshrubs, NEJA D. Don in Sweet, Hort. Brit. (ed. taprooted; caudex simple; glandular and oth- 2) 299. 1830. TYPE: Neja gracilis D. Don 5 erwise glabrous to hirsute or hispid; stems Neja filiformis (Spreng.) Ness 36 LUNDELLIA DECEMBER, 2008

Neja sect. Podoneja DC., Prodr. 5: 325. aceae: Astereae)—Molecular evidence for multiple ´ 1836. TYPE: Neja gracilis 5 Neja filiformis origins for the endemic flora of the Galapagos Islands. J. Biogeography. (Spreng.) Ness Ariza Espinar, L. 1980. Las especies centroargentinas de Neja sect. Monogyria DC., Prodr. 5: 325. Hysterionica (Compositae). Darwiniana 22: 537–549. 1836. LECTOTYPE (Nesom, 1993b): Neja linear- Atwood, N. D. and S. L. Welsh. 2007. New taxa of ifolia DC. 5 Neja pinifolia (Poir.) G.L. Nesom Camissonia (Onagraceae); Erigeron, Hymenoxys, Perennial herbs; taprooted; caudex and Tetradymia (Compositae); Lepidium and Physaria (Cruciferae) from Arizona, New Mexico, branched; glabrous to glabrate to sparsely and Utah. Rhodora 109: 395–414. pilose; stems usually branched; leaves mostly Brunsfeld, S. J. and G. L. Nesom. 1989. Erigeron basal, filiform to lnear-oblanceolate, entire, not salmonensis (Asteraceae), a rare new species from clasping; heads solitary, long-pedunculate or Idaho. Brittonia 41: 424–428. on sparsely bracteolate stems, erect in bud; rays Burke, J. L. 2005. The evolutionary origins of , a rare plant in , Ph.D. diss., yellow or white, not coiling; achenes fusiform- University of Lethbridge, Alberta, . cylindric, with 7–10 raised, thick, orange- Cabrera, A. L. 1946. El ge´nero Hysterionica en el resinous nerves; pappus of scabrid bristles. Uruguay y en la Republica Argentina. Notas Mus. N. dianthifolia (Griseb.) Nesom La Plata 11(Bot. 53): 349–358. N. filiformis (Spreng.) Nees Cronquist, A. 1943. The separation of Erigeron from Conyza. Bull. Torrey Bot. Club 70: 629–632. N. marginata (Griseb.) Nesom ———. 1947. A revision of the North American species of N. nidorelloides DC. Erigeron, north of Mexico. Brittonia 6: 121–302. N. pinifolia (Poir.) Nesom ———. 1971. Compositae. Pp. 300–367 in I. R. Wiggins N. pulvinata (Cabrera) Nesom and D. M. Porter, eds. Flora of the Gala´pagos Islands. Stanford Univ. Press, Stanford, Calif. Nesom (1994b, 1994c) suggested that ———. 1994. Intermountain flora. Vol. 5. . Neja might be most closely related to The Botanical Garden, Bronx, NY. Apopyros but molecular data (Noyes, Cuatrecasas, J. 1969. Conyza. Prima flora Colombiana. 2000a) indicate that it is probably the sister 3. Compositae – Astereae. Webbia 24: 198–228. group to Hysterionica, the genus from which Deble, L. P., A. Silveira de Oliveira, and J. N. C. Marchiori. 2004. Hysterionica nebularis, espe´cie it has been segregated. The six species of nova de Astereae – Asteraceae para o Estado do Neja occur in southeastern Brazil and Rio Grande do Sul (Brasil). Cieˆncia Florestal Santa adjacent Uruguay, Paraguay, and Argentina, Maria 14: 9–11. except for the Cuban endemic N. marginata Elisens, W. J., R. D. Boyd, and A. D. Wolfe. 1992. (Nesom, 1993c, 1994b, 2000). Genetic and morphological divergence among varieties of Aphanostephus skirrhobasis (Asteraceae – Astereae) and related species with different ACKNOWLEDGMENTS chromosome numbers. Syst. Bot. 17: 380–394. Engelskjo¨n, T. 1967. Contribution to the cytotaxon- I’m grateful to Rick Noyes for facilitat- omy of Grah., E. uniflorus L., and ing permission to reproduce the cladogram their hybrid. Nytt Mag. Bot. 14: 77–85. Harling, G. 1962. On some Compositae endemic to the from his 2000 publication in Plant System- Gala´pagos Islands. Acta Horti Berg. Bd. 20, 3: 63–120. atics & Evolution, Javier Francisco-Ortega Heath, P. V. 1994. Commentary on the proposal to for providing color photos of the three conserve Erigeron Linne. Calyx 4(3): 108–109. Gala´pagos species, and Bob Sivinski for Huber, W. 1993. Biosystematisch-okologische Untersu- permission to use a photo of his. chungen an den Erigeron-Arten (Asteraceae) der Alpen. Vero¨ffentl.Geobot.Inst.ETH,Zurich114:1–143. ——— and A. Leuchtmann. 1992. Genetic differen- LITERATURE CITED tiation of the Erigeron species (Asteraceae) in the Alps: a case of unusual allozymic uniformity. Pl. Allison, J. R. and T. E. Stevens. 2001. Vascular Flora of Syst. Evol. 183: 1–16. Ketona Dolomite Outcrops in Bibb County, ——— and O. Nilsson. 1995. Close genetic affinity of . Castanea 66: 154–205. northern and other Eurasiatic Erigeron species. Andrus, N., A. Tye, G. Nesom, D. Bogler, C. Lewis, R. Pp. 197–222 in D. J. N. Hind, C. Jeffrey, & G. V. Noyes, P. Jaramillo, and J. Francisco-Ortega. 2009 Pope, eds. Advances in Compositae Systematics, (in press). Phylogenetics of Darwiniothamnus (Aster- Royal Botanic Gardens, Kew. NUMBER 11 NESOM: REVIEW OF CONYZINAE 37

——— and H. Zhang. 1991. Morphologische und ———. 1992b. Taxonomic notes on Erigeron (Aster- chemotaxonomische untersuchungen an den Erig- aceae: Astereae) of California, Nevada, and Arizona. eron-Arten der Alpen. Ber. Geobot. Inst. ETH, Phytologia 73: 186–202. Zurich 57: 116–164. ———. 1992c. Erigeron and Trimorpha (Asteraceae: Hulte´n, E. 1968. Comments on the flora of and Astereae) in Nevada. Phytologia 73: 203–219. . Ark. Bot., Bd. 7, 1: 1–147. ———. 1992d. Laennecia spellenbergii (Asteraceae: Lawesson, J. E. and H. Adsersen. 1987. Notes on the Astereae), a new species from , Mexico. endemic genus Darwiniothamnus (Asteraceae-Aster- Phytologia 73: 267–269. eae) from the Gala´pagos Islands. Opera Bot. 92: 7–15. ———. 1993a. Erigeron jenkinsii (Asteraceae: Aster- Matzenbacher, N. I. and M. Sobral. 1996. Duas novas eae), a new species from the Rio Mayo region of espe´cies de Hysterionica Willd. (Asteraceae – Astereae) Sonora, Mexico. Phytologia 75: 118–120. no sul do Brasil. Comun. Mus. Cienc. Tecnol. – ———. 1993b. Erigeron mayoensis (Asteraceae: Aster- PUCRS: Se´r. Bot., Porto Alegre 2(1): 15–21. eae), a new species from northwestern Mexico. Morgan, D. R. 1990. A systematic study of Machaer- Phytologia 75: 218–220. anthera (Asteraceae) and related groups using ———. 1993c. A Cuban endemic: Hysterionica mar- restriction site analysis of chloroplast DNA and a ginata (Asteraceae: Astereae) rather than Aster taxonomic revision of sect. Psilac- grisebachii. Phytologia 75: 163–165. tis, Ph.D. diss., Univ. of Texas, Austin. ———. 1994a. Erigeron pattersonii (Asteraceae: Aster- Neel, M. C. and N. C. Ellstrand. 2001. Patterns of eae), a new species from Nuevo Leon, Mexico. allozyme diversity in the threatened plant Erigeron Phytologia 76: 96–100. parishii (Asteraceae). Amer. J. Bot. 88: 810–818. ———. 1994b. Separation of Neja (Asteraceae: Aster- Nesom, G. L. 1987. Erigeron turnerorum, a new species eae) from Hysterionica. Phytologia 76: 168–175. from Mexico and its relatives. Sida 12: 287–292. ———. 1994c. Apopyros (Asteraceae: Astereae), a new ———. 1989a. Taxonomy of Erigeron sect. Polyactis genus from southern Brazil, Argentina, and Para- (Compositae: Astereae). Phytologia 66: 415–455. guay. Phytologia 76: 176–184. ———. 1989b. The separation of Trimorpha (Composi- ———. 1994d. Subtribal classification of the Astereae tae: Astereae) from Erigeron. Phytologia 67: 61–66. (Asteraceae). Phytologia 76: 193–274. ———. 1994e. Taxonomic dispersal of Australian Erigeron ———. 1989c. Infrageneric taxonomy of New World (Asteraceae: Astereae). Phytologia 76: 143–159. Erigeron (Compositae: Astereae). Phytologia 67: 67–93. ———. 1994f. Pappochroma Rafin. is the correct ———. 1989d. A new species of Erigeron (Compositae: generic name for Erigeron pappochroma Labill. Astereae) from Arizona. Phytologia 67: 304–306. Phytologia 76: 426 p. ———. 1989e. A new species of Erigeron (Asteraceae: ———. 1994g. Pacifigeron (Asteraceae: Astereae), a new Astereae) from Tamaulipas, Mexico. 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