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Reclassification of North American Haplopappus (Compositae: Astereae) Completed: Rayjacksonia Gen

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Reclassification of North American Haplopappus (Compositae: Astereae) Completed: Rayjacksonia Gen AmericanJournal of Botany 83(3): 356-370. 1996. RECLASSIFICATION OF NORTH AMERICAN HAPLOPAPPUS (COMPOSITAE: ASTEREAE) COMPLETED: RAYJACKSONIA GEN. NOV.1 MEREDITH A. LANE2 AND RONALD L. HARTMAN R. L. McGregor Herbarium(University of Kansas NaturalHistory Museum Division of Botany) and Departmentof Botany,University of Kansas, Lawrence, Kansas 66047-3729; and Rocky MountainHerbarium, Department of Botany,University of Wyoming,Laramie, Wyoming82071-3165 Rayjacksonia R. L. Hartman& M. A. Lane, gen. nov. (Compositae: Astereae), is named to accommodate the "phyllo- cephalus complex," formerlyof Haplopappus Cass. sect. Blepharodon DC. The new combinationsare R. phyllocephalus (DC.) R. L. Hartman& M. A. Lane, R. annua (Rydb.) R. L. Hartman& M. A. Lane, and R. aurea (A. Gray) R. L. Hartman & M. A. Lane. This transfercompletes the reclassificationof the North American species of Haplopappus sensu Hall, leaving that genus exclusively South American.Rayjacksonia has a base chromosomenumber of x = 6. Furthermore,it shares abruptlyampliate disk corollas, deltatedisk style-branchappendages, and corolla epidermalcell type,among other features,with Grindelia, Isocoma, Olivaea, Prionopsis, Stephanodoria, and Xanthocephalum.Phylogenetic analyses of morphologicaland chloroplastDNA restrictionsite data, taken together,demonstrate that these genera are closely related but distinct. Key words: Astereae; Asteraceae; Compositae; Haplopappus; Rayjacksonia. During the past seven decades, taxonomic application lopappus sensu Hall (1928) are reclassifiedand are cur- of the broadly defined Haplopappus Cass. sensu Hall rentlybeing treatedas indicatedin Table 1. (1928) has given way to recognitionof several segregate genera in NorthAmerica, despitepleas for conservatism Taxonomic background-When Hartman (1976, fromsome authors(e.g., Cronquist,1994). The polyphy- 1990) transferredH. sect. Blepharodon DC. into Mach- letic nature of Haplopappus sensu Hall has been dem- aeranthera Nees (Table 1), he noted that the "phyllo- onstratedby studiesof pollen morphologyand flavonoid cephalus complex" did not belong with sect. Blepharo- chemistry(Clark, Brown, and Mayes, 1980; Clark et al., don. Several morphologicaland cytologicalstudies (Hart- 1980), by studies of chromosomenumber (Anderson et man and Lane, 1984, 1991; Lane and Hartman, 1984, al., 1974), and by morphological investigations(e.g., 1985; Lane, Hartman,and Brown, 1987) have provided Shinners,1950, 1951; Johnston,1970; Turnerand San- evidence (sharedbase chromosomenumber of x = 6, disk derson, 1971; Turner,1972; Urbatsch, 1976; Hartman, corollas thatare abruptlyampliate, style-branch append- Lane, and Brown, 1987; Nesom 1989, 1990, 1991a, b; ages thatare deltate,and corolla epidermalcell type) for Nesom and Morgan, 1990; Nesom et al., 1990). the relationshipof the "phyllocephalus complex" with Withthe breakupof Haplopappus sensu lato, all of the Isocoma (Hartman,1976), and also withGrindelia Willd. taxa recognizedby Hall (1928) have been accommodated (Steyermark,1937; Lane, 1982), Olivaea Schultz-Bip.ex withinpreviously named genera, except for H. phyllo- Benth. (DeJong and Beaman, 1963), Prionopsis Nutt. cephalus DC. and its variety(at which rank Hall treated (Nesom, Suh, and Simpson, 1993), Stephanodoria H. annuus [Rydb.] Cory) and H. aureus A. Gray, also Greene (Lane, 1980), and XanthocephalumWilld. (Lane, called the "phyllocephalus complex" (Venugopalan, 1983). Those studies indicated thatthe "phyllocephalus 1966; Hartman,1976). In thispaper, we presentevidence complex" is a coherentassemblage of species thatshould forthe genericstatus of thisgroup, formally describe the be recognized at the genericlevel, althoughJackson and new genus, and presentan analysis of its phylogenetic Dimas (1981) concluded otherwise based on a green- relationshipswith certain other Astereae. With the estab- house hybridizationbetween a species of the complex lishmentof this genus, all of the taxa formerlyof Hap- and one of Isocoma. 1 Manuscriptreceived 4 May 1995; revisionaccepted 27 June1995. The authorsthank Billie L. Turner,Guy Nesom, and herbarium TEX- MATERIALS AND METHODS LL forloan of specimens;John Semple and GregoryBrown for valu- able discussionof Haplopappus and related genera; David Morganand Morphological data-In Table 2 we comparemorphological features YoungbaeSuh forthe use of cpDNA restrictionsite data; David Keil of our new genus and the six othergenera thatwe considerto be most forassistance with Latin; Stacie Kawaguchifor her assistance in the closely related. Even though we have hypothesized(Hartman, 1990; constructionof Table 1 and JohnStrother for his donationof its first Lane and Li, 1993) thatthey mightbe part of the "x = 6" group of incarnation;Christopher Haufler, Gregory Brown, John Strother, and Astereae, Pyrrocoma, Vanclevea and Xylorhiza are not included be- TheodoreBarkley for critical readings of the manuscript; and Raymond C. Jacksonfor the use ofhis name. This study was supportedby grants cause phylogeneticanalysis of a large cpDNA data set that contains to MAL fromNSF (BSR 8508631and BSR 8908963)and theUniver- 163 taxa of Astereae and 729 restrictionsites obtained from digests sityof KansasGraduate Research Fund. with 16 enzymes (M. Lane et al., 1996; mutationlists on file at the R. 2Author for correspondence(Fax: 913-864-5093;e-mail: MLA- L. McGregorHerbarium, Division of Botany,University of Kansas Nat- [email protected]). ural HistoryMuseum [KANU] with specimens of our genus) showed 356 March 1996] LANE AND HARTMAN-RAYJACKSONIA 357 00 C0104 10 10 cl 0 Cid 0 4-i --q --q -4 C) z z z ob g r. -Q, cis CN C ON C . cis ON ON N CN Ch ON ON 4 4 CA cn CN (ON Ch ON 5N 0 0 C\ C\ C 4 C\ x 0 to 0 z cl C cd O\C 0 r-- 4 4 0 C) z z z C43 Cldr- Itt 0 0 C) ON ON C ON C C C CY\ CN CN ON C CIA ON ON ON t) --( ON C\ ON CIA ON ON ON CN CN ON C CN ON le,0 ON 1-4 r, cis -,;j o Z o" (ON aN 0 0 C\ CN t -4 I-q 1-4 -4 -4 W cl ON ON cis ON ON ON cis (u -4 ON w ON ON CIS r--( r--( - -4 0 0 t 04 : C's: -- o, lo C'sN 0-4 'A rA C's co rA ;3 ON CN - g - cl, 0 IL) (L) (L) (L) IL) -W o 0 0 0 o o w o o 0 0 10 rA w w w w w w w w (L) (L) (L) 0 0 0 0 0 0 t3 0 M0 C"i z u z z z z z z z z u u u Z Z t4 0 7:1 4 CA 4! 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C t (5 0 4 cn 0 CO) Z! it to Y 358 AmERICAN JOURNAL OF BOTANY [Vol. 83 co 0 rA0 rA0 0 w0 rA0 rA z z z z Z Z Ch CN 4 4 4 4 4 4 C Cf) Clf) cn Cf) cf) Cf) I-q 1--q 0 C C ON C ON ON C C ON _q I-q cis cis 0 t-c) CIS c \C \C \C \0 C>CN \Ct- C> \C OIA (0, C ON CN oo 00 10 ON CIA C ON 0 C (7N d" ON ON -q . --q ON CIN cl W q cis I-q 0 -q 0 --q cis CZ cl cl CIS 0 cd 0 \C C> C) (::> 0 W ;111-rA co u co &0 u rb 0 0 CN C,CIA cli 4) 0) 0) 4) 0 0 q) ,, O-, r- c-, C\ O., 0 O O ON O CN 5N c-, O ,- CN CN ON 0 0 0 0 -4 -4 0 0 1 C>131 1 C11 C', 2 0 a 2 $--4 Z "O z Z') Iz, m 40--,U z m -um PO z z Cd > 'o, n 0 4- rA rA (2, > > 0 0 Z rA -S -St 4 4 C1 z zi Z3 t3 T3 C's U rA d > > 4-) 4 0 ;> Cld C13 75 u u z C.) IL) .10 C 64 i 111.) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 (A ; cqs C's cis 11-1 >-, >lb C;3 rA ;-4 rA ai t4 Z 62 Cd C U U) 0 Z 3 < cd z Z (-Z) C 6; Q (Z -3 ; CA Z 0 CZ, 2" 4 1 110, r-n IS "i ti ct,Q 1:4Q Q Q to cd O r.
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