taxonomic revision of roldana (asteraceae: senecioneae), a genus of the southwestern u.s.a., mexico,...

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    Taxonomic Revision of Roldana (Asteraceae: Senecioneae),a Genus of the Southwestern U.S.A., Mexico, and CentralAmericaAuthor(s): A. Michele FunstonSource: Annals of the Missouri Botanical Garden, 95(2):282-337. 2008.Published By: Missouri Botanical GardenDOI:

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    A. Michele Funston2


    A revision of the genus Roldana La Llave (Asteraceae: Senecioneae) is given, including a key to species and completenomenclature with synonymies, description, distribution, and discussion sections for 48 species; eight varieties are recognizedand 11 new combinations are made, including one species transfer to the genus Psacaliopsis H. Rob. & Brettell. Aphytogeography section for the genus, which is distributed throughout the highlands of Mexico and Central America, with onespecies entering the southwestern-most U.S.A., is also included. New combinations are established for R. acutangula (Bertol.)Funston, R. aliena (B. L. Rob. & Seaton) Funston, R. hartwegii var. carlomasonii (B. L. Turner & T. M. Barkley) Funston, R.hartwegii var. durangensis (H. Rob. & Brettell) Funston, R. hartwegii var. subcymosa (H. Rob.) Funston, R. kerberi var.calzadana (B. L. Turner) Funston, R. kerberi var. manantlanensis (R. R. Kowal) Funston, R. petasitis var. cristobalensis(Greenm.) Funston, R. petasitis var. oaxacana (Hemsl.) Funston, R. petasitis var. sartorii (Sch. Bip. ex Hemsl.) Funston, as wellas for the excluded taxon P. pinetorum (Hemsl.) Funston & Villasenor. Neotypes are designated for Cineraria platanifoliaSchrank, R. ehrenbergiana (Klatt) H. Rob. & Brettell, R. lobata La Llave, R. petasitis (Sims) H. Rob. & Brettell, Senecioacerifolius K. Koch, S. canicidus Sesse & Moc., and S. prainianus A. Berger. Lectotypes are designated for Cacalia nutansSesse & Moc., C. peltata Sesse & Moc., P. pinetorum, R. albonervia (Greenm.) H. Rob. & Brettell, R. aschenborniana (S.Schauer) H. Rob. & Brettell, R. gilgii (Greenm.) H. Rob. & Brettell, R. heracleifolia (Hemsl.) H. Rob. & Brettell, R.heterogama (Benth.) H. Rob. & Brettell, R. kerberi (Greenm.) H. Rob. & Brettell, R. langlassei (Greenm.) H. Rob. & Brettell,R. lanicaulis (Greenm.) H. Rob. & Brettell, R. petasitis var. cristobalensis, R. petasitis var. sartorii, S. brachyanthus Greenm., S.chapalensis var. areolatus Greenm., S. chrismarii Greenm., S. ghiesbreghtii Regel var. pauciflorus J. M. Coult., S. grandifoliusLoes. var. glabrior Hemsl., S. hederoides Greenm., S. hypomalacus Greenm., S. lobatus Sesse & Moc., S. macrobotrys Hemsl., S.rotundifolius Sesse & Moc., and S. schumannianus S. Schauer.

    Key words: Asteraceae, Central America, Compositae, Mesoamerica, Mexico, Roldana, Senecioneae, taxonomy.

    The genus Roldana La Llave (Asteraceae: Sene-

    cioneae) is distributed throughout the highlands of

    Mexico and Central America. It is placed in the

    subtribe Tussilagininae because it has stigmatic

    surfaces united across the inner face of the style

    branches, upper part of stamen filaments (anther

    collar) cylindrical, chromosome numbers n 5 30, and

    principal phyllaries often with the midrib notably

    thickened at the base. The other tussilaginoid genera

    with which it is associated, based on geographic

    distribution, are Digitacalia Pippen, Psacalium Cass.,

    Robinsonecio T. M. Barkley & Janovec, Pippenalia

    McVaugh, Psacaliopsis H. Rob. & Brettell, Villase-

    noria B. L. Clark, Pittocaulon H. Rob. & Brettell,

    Telanthophora H. Rob. & Brettell, Nelsonianthus H.

    Rob. & Brettell, and Barkleyanthus H. Rob. &

    Brettell. It is separated from these genera because it

    arises from a hairy, tuberous caudex with numerous

    fleshy roots; leaves typically palmate or, if pinnate,

    then typically longer than 10 cm and/or variously

    hairy. In this synopsis, 48 species and eight varieties

    are recognized and 11 new combinations are made,

    including one species transfer to the genus Psaca-


    The genus has two centers of diversity; one is along

    the Trans-Mexican Volcanic Belt and Sierra Madre

    del Sur of south-central Mexico and the other is along

    the Sierra Madre de Chiapas and Cordillera Centro-

    americana of Chiapas and Central America. The

    species of Roldana vary in growth form from small

    perennial herbs that seldom exceed 1 m in height to

    freely branching trees that may reach 12 m in height.

    The architecture of the plants is insufficiently well

    understood to apply the rather precise descriptive

    1 If I were to properly recognize all those who helped with this revision, it would surely take as many pages; in memoriam, Ithank my mother Sandra Scharbach Funston and my major professor Theodore Mitchell Barkley for their guidance of this work.Financial support was provided by the Division of Biology, Kansas State University, and the Missouri Botanical Garden.

    2 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. 10.3417/2003151

    ANN. MISSOURI BOT. GARD. 95: 282337. PUBLISHED ON 18 JUNE 2008.

  • terminology of contemporary ecology, e.g., Halle et al.(1978), although the utility of such a classificationscheme is acknowledged. The genus is predominantlymade up of perennial suffruticose herbs that may growto shrubs. There are also four species of acaulescentperennial herbs and nine species that may grow totrees; stems are solitary or in clumps. Plants aretypically distributed in the understory of pine-oakforests at elevations between 1000 and 4000 m.Herbaceous plants are also found along the forestedge in patches of disturbed grasslands. All speciesmay be found in disturbed areas and roadsides. Themain characters used to separate the species arehabit, type and density of pubescence, number ofphyllaries, and, to some extent, the shape of the leafblade and its form of attachment.

    In 1901, J. M. Greenman wrote a revision of SenecioL. s.l. from North and Central America. In hisrevision, he separated the genus into 22 sections.Since that time, there has been considerable debatewhether the genus should remain together or be splitinto segregate genera. Currently, Greenmans conceptof Senecio is split into two subtribes, with 11 generafrom Mexico and Central America placed in thesubtribe Tussilagininae. While Robinson and Brettell(1973a, b, c, 1974) made a comprehensive review ofthese genera that included keys, none was given acomplete revision.


    Information and materials were derived from fieldexperiences in Costa Rica in January 1994, andMexico in March 1996, from consultations in CR,IEB, MEXU, MO, and from herbarium specimens thatwere lent to KSC from A, ASU, CAS, DS, F, GH,KANU, MEXU, MICH, MO, NMC, NY, OS, SD, TEX,UC, UMO, and XALherbarium abbreviations followHolmgren et al. (1990). Herbaria referred to, but fromwhich no loans were requested, are B, C, G, K, P, andUS. Gibson (1969: 97) notes in his dissertation,Correspondence with Dr. G. Wagenitz indicates thatall of the types of the genus Senecio at the BerlinHerbarium were destroyed in 1943. Species arepresented in alphabetical order. Complete lists ofexsiccatae and distribution maps are available fromthe author.

    Microcharacters as reviewed by King and Robinson(1970), Jeffrey (1987, 1992), and Nordenstam (1978)have proven useful in the Senecioneae. Microchar-acter studies of Senecio include Drury and Watson(1965), Jeffrey et al. (1977), Jeffrey (1979), Vincent etal. (1992), and Wetter (1983). For my dissertation(Funston, 1999), microcharacters in Roldana wereinvestigated for consistency within the genus. Not all

    species could be coded; however, those that were

    showed a significant degree of similarity. While

    microcharacters did confirm the placement of species

    in the subtribe Tussilagininae, they were not sufficient

    to include or exclude a species at the generic level. A

    character state matrix of Roldana for Jeffreys (1987,

    1992) analytical descriptors is available from the


    A numerical analysis performed on the species

    Roldana hartwegii (Benth.) H. Rob. & Brettell is also

    available from the author. A data matrix of 64

    collections and five character states was analyzed

    using NTSYS-PC v. 1.8 (Rohlf, 1993). The data matrix

    was standardized using a simple matching coefficient.

    The standardized

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