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Home , Amphiachyris, Astereae, Columbiadoria, Croptilon, Ericameria, Ericameria cuneata

AmericanJournal of Botany 83(3): 356-370. 1996.

RECLASSIFICATION OF NORTH AMERICAN (COMPOSITAE: ) COMPLETED: GEN. NOV.1

MEREDITH A. LANE2 AND RONALD L. HARTMAN

R. L. McGregor Herbarium(University of NaturalHistory Museum Division of Botany) and Departmentof Botany,University of Kansas, Lawrence, Kansas 66047-3729; and Rocky MountainHerbarium, Department of Botany,University of ,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 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 , , , Prionopsis, Stephanodoria, and .Phylogenetic analyses of morphologicaland chloroplastDNA restrictionsite data, taken together,demonstrate that these genera are closely related but distinct.

Key words: Astereae; ; 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, , Vanclevea and 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

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thatthese threegenera are not a partof the clade withabruptly ampliate Cis -4 . 0 disk corollas and deltatestyle-branch appendages. U) 1: >~- Certain of the featurespresented in Table 2 are ceded for phyloge- netic analysis. The characteristicsin Table 2 are consistentamong all the species of a genus; we have encapsulatedthe variationwithin the genera in the descriptionsof charactersand in the scoringsused forthe phylogeneticanalysis (e.g., polymorphismsare so coded). Cl, ~ ~ ~~ ~ ~ ~ ~~~~~~~~~~C indicatedin the caption 7Z Greene was chosen as the outgroup(codings as of Table 2) because the cpDNA analysismentioned above indicatedthat Hazardia is a memberof a clade that is sisterto the clade containing the generaof interestin thisstudy. A data matrixwas constructedusing ~~~~~~~~~s~~~~~~~~~~~~~~~~~~~~~i 4OH C MacClade Version 3.0 (Maddison and Maddison, 1992). The matrix -~~~~~~Q~~~~~~~ZZ u K~~~~~~~~1-0 comprisedseven ingroupand one outgrouptaxa vs. the 17 coded char- 0 acters fromTable 2. H H~~~~~~~' Molecular data-The cpDNA data set of Lane and coworkers(1996) containsrepresentatives of all the genera in this studyexcept Olivaea. - -~~~~~~~~~~~~~C ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ A submatrixcomposed of representativesof Grindelia (G. lanceolata Nutt.,Suh 85, : Blanco Co. [TEX]), Isocoma (I. pluriflora[Torrey & A. Gray] Greene, Suh 91, : [TEX]), Prionopsis 0 ( 4~~~~ (P. ciliata [Nutt.]Nutt., Morgan 2084, Texas: SuttonCo. [TEX]), Ra- yjacksonia (Lane 3157, : Pueblo Co. [KANU]), Stephanodoria 0 84~~~~ (S. tomentella[B. L. Robinson] Greenm.,Nesom 6681, Mexico: San Luis Potosi [TEX]), Xanthocephalum(X. humile[Kunth] Benth., Nesom 6769, Mexico: Mexico [TEX]), and the outgroupHazardia (H. detonsa [Greene]Greene, Junak 4158, :Santa Barbara Co.: Santa Bar- bara Botanic Garden [SBBG]) was drawnfrom the large data set using

0 94~ MacClade. Of the 729 charactersin the whole data set, 38 were found to be phylogeneticallyinformative for the taxa of interesthere; they were extractedfrom the largerdata set and used in our analyses.

Combined data-Following the analysis of molecular data, a third to C' matrixwas compiled;this included the 17 morphologicalcharacters plus 4-;~ - the 38 molecularcharacters. Two exhaustivesearches forshortest trees were conductedusing the thirdmatrix: one that excluded Olivaea be- 6O ) ra 'k cause no data were available for the 38 molecular characters,and one 0 ~~~~~~~~~~~~~~~~~~~ that included Olivaea, coded as missing data for the molecular char- acters.

0~~~~~~~~~~~r Phylogeneticanalyses-Exhaustive searches for shortesttrees were 0 -~~~~~~~~~~~~~~~~~N made using Phylogenetic Analysis Using Parsimony Version 3.1.1 (PAUP; Swofford,1993). Multistatecharacters were treatedas unor- dered and as representingpolymorphisms; ACCTRAN optimizationwas used in all cases. Trees were drawn for presentationusing MacClade Version 3.0 (Maddison and Maddison, 1992). 00 4 C) toN ~ ~ .u .5* RESULTS The distinctionsamong generapresented in Table 2 are ;-~~~~~~~ ~ ~ 4 those that we have found to be most taxonomicallyin- g ?~~~~~~~~~~~~~~~~~~~~~~~~~~~0P~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~-formative during several years of studyof these members :71 -) ( of Astereae. In addition,the analysis of morphological and cpDNA data provides strongsupport for the segre- gationof the "phyllocephaluscomplex" fromHaplopap- H- - pus sensu Hall (1928). Therefore,we here establish the genus Rayjacksonia to accommodate R. phyllocephala (type species), R. annua, and R. aurea. The formalities of classificationand nomenclatureare presentedbelow. A key to the species of Rayjacksonia, descriptions,and discussion of theirrelationships will appear in a forth- coming thesis and publicationby one of our students. The exhaustive search of the morphologicaldata set resultedin fivefully resolved, equally parsimonioustrees (Figs. 1-5) of 56 steps each witha consistencyindex (CI) of 0.821, a retentionindex (RI) of 0.600, and a rescaled March 1996] LANE AND HARTMAN-RAYJACKsoNiA 365

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1 3 Stephanodoria ~~~~~~~~~~2Grindelia Grindelia

Xanthocephalum { Olivaea Prionopsis Olivaea Prionopsis _ Rayjacksonia 40 0~~ Grindelia Rayjacksonia _ Olivaea - Prionopsis Stephanodoria Xanthocephalum

rL = Rayjacksonia _[ _ Xanthocephalum r Stephanodoria Isocoma Isocoma - socoma

Hazardia Hazardia Hazardia

Prionopsis Prionopsis -Grindelia

rIL Rayjacksonia rf{ Rayjacksonia -Isocoma 0 J Grindelia Grindelia _ Olivaea Olivaea _ Olivaea - Prionopsis ;-~~~~~~~~- Xanthocephalum 1 Xanthocephalum Rayjacksonia Stephanodoria _ Stephanodoria _ Stephanodoria

Isocoma - Isocoma Xanthocephalum

Hazardia Hazardia Hazardia

00 Figs. 1-5. The fivemost parsimonioustrees (56 steps,CI = 0.821, RI = 0.600, RC = 0.493) thatresult from an analysis of scoringsof 17 morphological characters(Table 2) for Grindelia, Isocoma, Olivaea, Prionopsis, Rayjacksonia, Stephanodoria, and Xanthocephalum.The outgroup,Hazardia, was scored as indicatedin the caption of Table 2. 0~~~~ One of these trees (Fig. 5) is equivalent to a 60% majorityrule con- sensus tree. Fig. 6. The strictconsensus tree of the five most parsi- monious trees (Figs. 1-5) derived fromthe analysis of morphological data.

consistencyindex (RC) of 0.493. One of the five trees (Fig. 5) is equivalent to a 60% majorityrule consensus tree. The strictconsensus, however,is completelyunre- solved (Fig. 6). Analysis of thecpDNA data forsix of the seven genera C ) of this clade (excluding Olivaea) also resulted in five most parsimonioustrees (Figs. 7-11) of 37 steps (CI = 0.946, RI = 0.867, RC = 0.820). Again, one of the five 0 ~ ~ ~

7 8 9 GRINDELIA GRINDELIA STEPHANODORIA

PRIONOPSIS PRIONOPSIS XANTHOCEPHALUM

fSTEPHANODORIA ISOCOMA JL ISOCOMA 0~~~~~ .il C0 . _ XANTHOCEPHALUM L STEPHANODORIA GRINDELIA

RAYJACKSONIA XANTHOCEPHALUM PRIONOPSIS ISOCOMA RAYJACKSONIA A O A RAYJACKSONIA

HAZARDIA HAZARDIA - HAZARDIA

10 11 12 4GRINDELIA GRINDELIA GRINDELIA STEPHANODORIA STEPHANODORIA STEPHANODORIA

XANTHOCEPHALUM XANTHOCEPHALUM jL XANTHOCEPHALUM ISOCOMA jL ISOCOMA - ISOCOMA | RAYJACKSONIA 1 RAYJACKSONIA RAYJACKSONIA

HAZARDIA HAZARDIA HAZARDIA

Figs. 7-12. The fivemost parsimonious trees (37 steps,CI = 0.946, RI = 0.867, RC = 0.820) that resultfrom an analysis of 38 cpDNA restrictionsite characters(see Methods) for Grindelia,Isocoma, Prion- opsis, Rayjacksonia,Stephanodoria, and Xanthocephalum(data forOli- vaea were unavailable; the outgroupis Hazardia). One of these trees (Fig. 11) is equivalent to a 60% majorityrule consensus tree. Fig. 12. The strictconsensus tree of the five most parsimonioustrees (Figs. 7- 11) derived fromthe analysis of cpDNA restrictionsite data. March 1996] LANE AND HARTMAN RAYJACKSONIA 367

13 GRINDELIA 15 STEPHANODORIA 9 1 PRIONOPSIS 2 XANTHOCEPHALUM 2 RAYJACKSONIA 1 Olivaea 51 16 STEPHANODORIA 1 GRINDELIA 6 PRIONOPSIS 10 XANTHOCEPHALUM _ fl ~1 3 RAYJACKSONIA ISOCOMA 4 1 ISOCOMA HAZARDIA HAZARDIA Fig. 15. The single most parsimonioustree (73 steps, CI = 0.849, RI = 0.703, RC = 0.597) that resultsfrom an analysis of Grindelia, Isocoma, Olivaea, Prionopsis, Rayjacksonia, Stephanodoria,and Xan- 14 2 GRINDELIA thocephalum(the outgroupis Hazardia) using a combined matrixthat included 17 morphologicaland 38 cpDNA restrictionsite characters. Numbers below branches indicate unambiguous changes that support PRIONOPSIS 1 the indicated clade. Only morphologicalcharacters were available for Olivaea, so lowercase lettersare used forthe Olivaea label, as in Figs. 5 STEPHANODORIA 1-6; genera for which molecular data were available are labeled in 2 uppercase,as in Figs. 7-14. o8 XANTHOCEPHALUM to Fig. 8, except that Olivaea is included as sistergroup to the Stephanodoria-Xanthocephalumclade. 25 7 RAYJACKSONIA ISOCOMA DISCUSSION 2 The 15 trees resultingfrom the analyses conductedin HAZARDIA this study are equally well supported,with CIs > 0.82 and RCs > 0.49 in all cases. Only two of the 15 trees Figs. 13-14. The two most parsimonious trees (71 steps, CI = (Figs. 7, 14) are fullycongruent, which is attributableto 0.845, RI = 0.676, RC = 0.572) thatresult from an analysis of Grin- delia,Isocoma, Prionopsis, Rayjacksonia, Stephanodoria, and Xantho- the largernumber of charactersin the moleculardata set cephalum (Olivaea excluded; the outgroupis Hazardia) using a com- (38) vs. the morphologicalone (17), and the absence of bined matrixthat included 17 morphologicaland 38 cpDNA restriction Olivaea from the analysis that produced Fig. 14. Two site characters.Numbers below branchesindicate unambiguous changes othertrees (Figs. 8, 15) mightbe congruentbut mustbe thatsupport the indicatedclade. questionedbecause Olivaea was excluded fromthe anal- ysis from which Fig. 8 was derived, and because data were missingfor Olivaea for38 of the 55 charactersused trees (Fig. 11) is equivalentto a 60% majorityrule con- to produce Fig. 15. Additional molecular (e.g., ITS se- sensus. In this case, the strictconsensus (Fig. 12) is quence) data mightallow forcomplete resolution of these slightlymore resolved,with Stephanodoria and Xantho- relationships,but all thatthe currentanalyses can show cephalum forminga clade, as do Grindelia and Prion- is thatthese seven genera are closely allied with one an- opsis. other.The lack of resolutionin the strictconsensus trees The first"total evidence" analysis (using six of the (Figs. 6, 12) indicatesthat different pairings of the genera genera and 55 characters)resulted in two equally parsi- share differentcombinations of characteristicsderived monious trees (Figs. 13, 14) of 71 steps (CI = 0.845, RI fromtheir common ancestor.It seems likely that these = 0.676, RC = 0.572). The topologyof Fig. 13, in which groups did not arise in strictlydichotomous fashion. Isocoma is a sistergroup to the Stephanodoria-Xantho- Rayjacksonia is variously allied with Prionopsis, the cephalum clade, while Rayjacksonia is sisterto the Grin- Grindelia-Prionopsis clade, or Isocoma, or it is main- delia-Prionopsis clade, is unlike any of the trees shown tained as a distinctclade (Figs. 1-15). Therefore,in our in Figs. 1-12. The topologyof the tree shown in Fig. 14 opinion, Rayjacksonia should be treatedas an indepen- is equivalentto thatof Fig. 7, which places Isocoma and dent genus and not be unitedwith Isocoma (Jacksonand Rayjacksonia in clades distinctfrom and basal to the Dimas, 1981) or regardedas more closely relatedto Iso- Stephanodoria-Xanthocephalumand Grindelia-Prionop- coma (Hartman,1976) thanit is to any of the othergen- sis clades. The second "total evidence" analysis, which era in this group. Further,Xanthocephalum and Isocoma included all seven genera and 55 charactersalthough the are separatedby several unambiguouschanges (Figs. 13- moleculardata are missingfor Olivaea, resultedin a sin- 15). As we have stated previously(Hartman and Lane, gle most parsimonioustree (Fig. 15) of 73 steps (CI = 1991), theyshould be maintainedas distinctgenera, even 0.849, RI = 0.703, RC = 0.597). This tree is equivalent thoughan intergenerichybrid is known. 368 AMERICAN JOURNAL OF BOTANY [Vol. 83

Stephanodoria and Xanthocephalumare sistergroups Annual to perennialherbs or suffrutescentsubshrubs in all of the analyses that include molecular characters, (0.4-) 1.5-8 dm or moretall, generally stipitate-glandular but are not always unitedin the trees resultingfrom the throughout,often densely so. Taproot(3-) 7-25 cm long, morphologicaldata alone (e.g., Figs. 9-11). It is tempting slender to stout,often becoming woody. Stems erect to to suggestthat Stephanodoria should be submergedwith- arcuate-ascending,usually branchingat base or above, in Xanthocephalum(and indeed the combinationexists green to whitishor red-purple,1-7 mm in diam, smooth to accommodatethis interpretation; Robinson, 1893), ex- to irregularlyridged, leafy throughout.Leaves alternate, cept that two of the equally parsimonioustrees include ascending to erect, simple, 1.2-2 times the internode Isocoma in this clade (Figs. 2, 13) and two include Oli- length; linear to oblanceolate, 1-8 cm long, 1-20 mm vaea in it (Figs. 1, 15). No treeincludes all fourof these wide, herbaceous to succulent,uppermost ones much re- genera in a single clade. Therefore,we believe that the duced, marginentire or usually coarsely serratewith 5- distinctstatus of these genera should be maintaineduntil 10 elongate, sometimessubulate, teethgenerally tipped data are obtained thatunequivocally supportone or an- by a white bristle,leaf apices rounded to acute, often otherof the possible mergers. bristle-tipped,midveins usually prominent;secondary Finally, Grindelia and Prionopsis are sistergroups in leaves oftenpresent in axils of primaryones, much re- all of the trees thatresult from analyses thatinclude the duced. Heads 1-3 on primarystems and lateralbranches, molecular data (Figs. 7-15), but this is trueof only one 1-70 or more per , the whole often appearing pa- of the five trees based on morphologicaldata (Fig. 3). niculate or subcorymbiform-cymose,peduncles some- Nesom, Suh, and Simpson (1993) argued that only the times cobwebby; involucreshemispheric, 7-16 mm tall, pappus differsbetween Grindelia and Prionopsis, and 10-20 mm wide, phyllaries50-85, imbricatein four or thatby very little(caducous, distinctawn-like bristles vs. five series, linear,lanceolate, or oblong-lanceolate,each deciduous, united bristles); however,we find that there one-nerved,light green to tan, cartilaginous,sometimes are othercharacters that distinguish the two (Table 2). In narrowedin proximal one-fourthto three-fourths,distal Prionopsis, the phyllariesare abaxially eglandular,and portionherbaceous, apex erect to recurved,often bristle- the leaf apices and marginalteeth are each tipped by a tipped,one-nerved, inner phyllaries with scarious sides; long, soft bristle,while in Grindelia the phyllariesare receptacles 2-9 mm in diam, glabrous, alveolate, walls punctate-glandular,and equivalent bristles are missing of alveoli irregular,often bearing subulate enations0.2- fromthe leaves. The moleculardata stronglysuggest that 1.8 mm long. Ray floretsfertile, 18-46, corollas yellow, Grindelia and Prionopsis are sistergroups, but thereare glabrous,9-15 mm long, tubes 1.8-3.5 mm long, limbs equally parsimonioustrees that suggest that eitherIso- 5.5-12 mm long, style branches 1-1.2 mm long; disk coma (Fig. 8) or Rayjacksonia (Figs. 3-5, 13) should be floretshermaphroditic, 60-150 or more,corollas yellow, included in this clade. Nesom and Suh (in Nesom, Suh, glabrous,5-7 mm long, tubes 2.5-3.8 mm long, throats and Simpson, 1993) have provided the combinationto abruptlyampliate, 1.7-2 mm long, lobes 0.6-1.1 mm accommodatePrionopsis withinGrindelia, but, again, we long, erect to slightlyspreading, anther bases rounded, believe thatthe distinctstatus of these genera should be antherappendages narrowlyovate, stylebranches 0.9-1.7 maintained until data are obtained that unequivocally mm long, theirappendages more or less deltate.Achenes supportone or anotherof the possible mergers. tan, covered with silky pubescence, the trichomes0.2- 0.8 mm long, achenes of ray floretsasymmetric, broadly ellipsoid to obovoid, obscurelythree-angled, 1.4-2.1 mm long, angles often stronglyribbed, faces with 0-4 ribs, RAYJACKSONIA R. L. Hartman& M. A. Lane walls thick,indurate, achenes of disk floretsbroadly el- lipsoid to clavate, somewhatcompressed radially, 1.4-3 Herbae nunc annuae nunc perennes vel suffrutices mm long, angles stronglyribbed, faces with 5-9 ribs, usque ad plus quam 8 dm alti, generatimglandibus stip- walls thin,membranous; pappus persistent,whitish to tan, itatisobsitae ubique. Caules erecti ad arcuate ascenden- bristles40-70 in threeor fourgraduated series, each bris- tes, foliacei. Folia laminis linearibus vel oblanceolatis, tle tangentiallyflattened, tapering from base to apex, bar- integrisvel plerumquedentibus 5-10 elongatis interdum bellate,of ray florets2.4-3.5 mm long, of disk florets3- subuliformibus,saepe setis albis terminatisgrosseserratis, 6 mm long; x = 6. Gulf of Mexico coast of western suprema reducta. Capitula 1-3 in caule primario et Florida, extreme western Louisiana, Texas, and north- quoque ramo laterale,una tota saepe aspectu paniculae eastern ; the area of Houston, Texas; and vel cymae subcorymbosae;involucrum hemisphaericum, northwesternTexas, , Kansas, and to phyllariisimbricatis in 4 vel 5 seriebus,apicibus erectis eastern , Colorado, and Wyoming. Soils ad recurvatis,saepe setiferis;receptaculum alveolatum sandy,well-drained, alkaline and/orsaline. Type species: parietibusalveolarum extensionesirregulares saepe sub- Rayjacksonia phyllocephala (DC.) R. L. Hartman& M. ulatas facientibus.Flosculi radii 18-46, corollis flavis; A. Lane. flosculi disci 60-150, corollis flavis,fauce abrupteam- The genus is named forDr. RaymondC. Jackson,who pliata, appendicibusramorum styli plus minusvedeltatis. firstreported the correlationin the Astereae between a Achenia brunneolapubescentia, flosculorum radii asym- base chromosomenumber of x = 6 and the abruptlyam- metricaet late elliptica ad obovata, obscure 3-angulosa, pliate or "goblet-shaped" disk floretcorollas. Further- flosculorumdisci late elliptica ad oblanceolata, aliquan- more, he has spent much of his professionalcareer in- tum lateralitercompressa; pappus setis ancistrispersis- vestigatingspecies of Haplopappus sensu lato. tentibusin 3 vel 4 seriebus imbricatis.Chromosomatum numerus basalis: x = 6. Rayjacksonia annua (Rydb.) R. L. Hartman & M. A. March 1996] LANE AND HARTMAN-RAYJACKSONIA 369

Lane, comb. nov.-Sideranthus annuus Rydb., Bull. Manatee Co.: Palmetto,S shore of Sneed's Island, TorreyBot. Club 31: 653. 1904.-Haplopappus phyl- near mouth of Manatee River, 21-23 Aug 1895, locephalus DC. subsp. annuus (Rydb.) H. M. Hall, G. V. Nash 2432 (Holotype: NY!; Isotypes: G, Publ. Carnegie Inst. Wash. 389: 58, 1928.-Aplopap- GH!, K, MICH!, ND-G, PH [photo: RM!], US! pus annuus (Rydb.) Cory, Rhodora 38: 407. 1936.- [photo: RM!]). phyllocephala (DC.) Shinners var. Eriocarpum floridanum Gand., Bull. Soc. Bot. annua (Rydb.) Shinners,Field & Lab. 18: 40. 1950. France 65: 41. 1918.-TYPE: U.S.A.: Florida: Haplopappus phyllocephalusDC. var. annuus (Rydb.) [Escambia Co.]: Pensacola, S. M. Tracy 8515 Waterf., Rhodora 62: 321. 1960.-Machaeranthera (Holotype: LY; Isotypes: GH!, ND-G, US!). annua (Rydb.) Shinners,Sida 1: 378. 1964.-TYPE: Eriocarpum glaucum Gand., Bull. Soc. Bot. France U.S.A.: Colorado: Lincoln Co.: Platte Valley, Jules- 65: 41. 1918.-TYPE: U.S.A.: Texas: [Galveston burg, 28 Aug 1891, P. A. Rydberg 145 (Lectotype Co.]: Galveston Island, S. M. Tracy 7365 (Holo- [here designated]:NY!). type: LY; Isotypes: G, GH!, US). Rayjacksonia aurea (A. Gray) R. L. Hartman& M. A. Eriocarpumtracyi Gand., Bull. Soc. Bot. France 65: Lane, comb. nov.-Haplopappus aureus A. Gray, 41. 1918.-TYPE: U.S.A.: Florida: Manatee Co.: Mem. Amer. Acad. Arts, ser. 2, 4: 76. 1849.- Palmetto:Sneed's Island, S. M. Tracy6354 (Holo- aureus (A. Gray) Kuntze, Revis. Gen. P1. 1: 317. type: LY; Isotypes: GH!, US!). 1891.-Sideranthus aureus-(A. Gray) J. Small, Fl. S.E. U.S. 1186. 1903.-Machaeranthera aurea (A. Gray) EXCLUDED TAXON Shinners,Field & Lab. 18: 41. 1950.-TYPE: U.S.A.: Texas: [HarrisCo.?]: low prairiesnear Houston, Sep- Haplopappus phyllocephalus DC. subsp. primitivusH. Oct, C. Wrights.n. (Holotype: GH! [photo: UC!]; Iso- M. Hall, Carnegie Inst. Wash. Publ. 389: 56. 1928. = types: GH!, PH!, US!). Machaeranthera gymnocephala(DC.) Shinners,Field Rayjacksonia phyllocephala (DC.) R. L. Hartman& M. & Lab. 18: 40. 1950 (Hartman,1976, 1990). A. Lane, comb. nov.-Haplopappus phyllocephalus DC., Prodr. 5: 347. 1836.-Aplopappus rubiginosus LITERATURE CITED Torr. & A. Gray var. phyllocephalus (DC.) A. Gray, Syn. Fl. N. Amer. 1(2): 130. 1884.-Aster phylloceph- ANDERSON, L. C., D. W. KYHOS,T. MOSQUIN,A. M. POWELL,AND P H. alus (DC.) Kuntze, Revis. Gen. P1. 1: 316. 1891.- RAVEN 1974. Chromosomenumbers in Compositae. IX. Haplo- pappus and otherAstereae. American Journal of Botany 61: 665- Eriocarpumphyllocephalum (DC.) Greene,Erythea 3: 671. 15. 1895.-Eriocarpum rubiginosum(Torr. & A. Gray) BROWN, G. K., AND W. D. CLARK. 1981. Chromosome numbers in Brittonvar. (DC.) A. Heller, Contr. South AmericanHaplopappus Cass. (Compositae). AmericanJour- Herb. 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