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CHROMOSOME COUNTS of COMPOSITAE from MEXICO and the UNITED Statesl

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CHROMOSOME COUNTS of COMPOSITAE from MEXICO and the UNITED Statesl Amer!_ J. Bot. 64(6): 791-798. 1977. CHROMOSOME COUNTS OF COMPOSITAE FROM MEXICO AND THE UNITED STATESl DAVID J. KEIL AND TOD F. STUESSY Department of Biological Sciences, California Polytechnic State University, San Luis Obispo 93407, and Department of Botany, The Ohio State University, Columbus 43210 ABSTRACT Chromosome counts of Compositae are reported from Mexico and the United States. First generic counts are from Egletes (n == 27), Hydropectis (n == 9), and Pippenalia (2n == ca. 60). First counts also are reported for 22 specific and infraspecific taxa in Baccharis, Erigeron, Flaveria, GnaphaUum, Grindelia, Helenium, Heterotheca, Melampodium, Montanoa, Pery­ menium, Piqueria, Senecio, Stevia, Verbesina, Xanthocephalum, and Zaluzania. Additional counts also are provided for 123 populations of taxa counted previouslY, of which eight are new numbers. Taxonomic implications of certain counts are discussed. THIS PAPER is a continuation of our studies (Keil first or new counts or significant comments. The and Stuessy, 1975) on chromosome numbers in order of commentary follows the sequence of the Compositae principally from the western tribes in the classification of Hoffmann (1890-94; United States and Mexico. As before, we have with the additional recognition of the Tageteae as adopted the policy of reporting new populational a separate tribe [Rydberg, 1915-16]), which is counts of previously documented taxa as well as the same as that used in Table 1. References for first reports for genera, species and varieties. The statements regarding the broad range of chromo­ materials and methods involving conventional somal variation within genera will not be given; acetocarmine squashes of buds for meiotic stages documentation for these counts comes from avail­ are the same as those mentioned in our earlier able chromosomal indices (Darlington and Wylie, paper. All voucher specimens are on deposit in 1955; Cave, 1958-65; Ornduff, 1967-69; Fe­ the herbarium of the Ohio State University (OS). dorov, 1969; Moore, 1970-74). A number of first reports for taxa are con­ RESULTs-The chromosome counts obtained sistent with previous counts for the respective are listed in Table 1. First counts are reported genera. These are listed here and will not be dis­ for three genera, 20 species, and two varieties; cussed further: Baccharis heterophylla, n == 9; 123 additional counts are for taxa counted pre­ Erigeron dryophyllus, n == 9; Flaveria robusta, n viously, eight of which are new numbers. The == 18; Gnaphalium cf. chartaceum and G. stram­ first counts for genera are from Egletes ,Casso (n ineum, n == 14; Grindelia tenella, n == 6; Hetero­ == 27), Hydropectics Rydb. (n == 9), and Pip'pen­ theca chrysopsidis, n == 9; Montanoa pyramidata, alia McVaugh (2n == ca. 60), and first counts for n == 19; Senecio arizonicus, S. flaccidus, and S. species are in Baccharis L., Erigeron L., Flaveria lvootonii, n == 20; Stevia aschenborniana, n == 11; Juss., Gnaphalium L., Grindelia Willd., Helenium S. berlandieri A. Gray var. berlandieri, n == 17; L., Heterotheca Cass., Melampodium L., Mon­ Verbesina tequilana, n == 17; and Zaluzania disco­ tanoa Cav., Perymenium Schrad., Piqueria Cav., idea, n == 18. Senecio L., Stevia Cav., Verbesina L., Xantho­ cephalum Willd., and Zaluzania Pers. Eupatorieae-Our count of ca. 84 I for an un­ determined species of Ageratina is indicative of DISCUSSION-As in our earlier paper (Keil and the cytological complexity of this genus. Based Stuessy, 1975), the discussions are restricted to upon previous reports at the diploid, tetraploid, and apparently pentaploid levels, King and Rob­ 1 Received for publication 30 September 1976; revision inson (1970) postulated a base number of x == accepted 8 February 1977. 17 for Ageratina. Our count probably represents Field work for this study was completed under NSF a pentaploid (5x == 85). Insight into the biological grants GB-30240 and GB-37678. Thanks go to: Marvin­ Roberts and Robert Gardner for assistance during col­ complexity of Ageratina is hindered by the lack of lecting trips; Rogers McVaugh of the University of Mich­ any published revision of the Latin American igan and Lorin Nevling of the Field Museum for courtesy taxa. King and Robinson (1970) listed over 200 and assistance during our research visits to their herbaria; species for the genus (most as transfers from Ted Barkley and Mike Powell for the identification of several 'of our collections of Senecio, Flaveriaand Peri­ Eupatorium L.) but provided no keys, descrip­ tyIe; and Loran Anderson for permission to include an tions, or other means of identification. unpublished mitotic count of Melampodium repens. Our count of n == 20 for Eupatorium sagitta- 791 792 AMERICAN JOURNAL OF BOTANY [Vol. 64 tum is the- first polyploid count for this species. Counts have· been published previously for 27 Diploid counts of n = 10 have been r~ported p~e­ of the 37 recognized species of Melampodium viously (Turner and Flyr, 1966; Ked and Pln­ (Stuessy, 1970, 1971; Keil and Stuessy., 1975). kava, 1976), also from Sinaloa where this taxon Of particular interest, therefore, are the fIrst chro­ is common (Robinson, 1926). mosome counts for two additional species, M. Five species of Piqueria have been counted be­ repens (2n = 54) and M. tepicense (n = 12). fore as n = 10, 11, 12, and 24. Our first report The former taxon and the related M. bibractea­ of n = 12 for P. cf. triflora conforms to the tum S. Wats. constitute section Bibractiaria known chromosomal diversity. Our voucher dif­ (Steussy, 1971), which has been the ?nly section fers from the typical P. triflora in having glabrous of the genus still unknown cytologIcally. The rather than hirtellous phyllaries and stems. relationships of this section with other sectio~s are difficult to assess due to the extreme reductIon of Astereae-Achaetogeron, a genus of about floral features in both species. The chromosome chromosom~l lev~l.s eight species, has had two re­ number of 2n = 54 seems probably based on x = ported previously: n = 9 from A. plnnatlfldum 9, which would place this group closest to section A. Gray (Turner and Flyr, 1966) and A. griseus Zarabellia with the same chromosomal base Greenm. (Dejong and Longpre, 1963); and n (counts of n = 9, 18, and 27 have been reported). = 27 from A. forreri Greene (Dejong and Long­ Morphologically, M. longifolium Cerv. ex Cav. forre~i pre, 1963). Our count of n = 36 for A. cf. would be the species possibly linking the two sec­ is not only a new chromosomal .level for th.IS tions, although the Iconnection is .distant. The species, but also the first report of the octoploid high ploidy level (probably hexaploId) of M. re­ level for the genus. pens corresponds with its specialized prostrate , .Our count of n = 27 for Egletes viscosa is the, growth habit and reduced floral features. On mor­ first for the genus. An ancestral base number of beca~se phological grounds, M. tepicense is most closely x = 9 for Egletes is suggested, especially related to M.divaricatum (Rich. in Pers.) DC. two related genera (Hoffmann, 1890-94; ShIn­ and M. dicoelocarpum Robins. of section Serra­ ners, 1949), Grangea Adans. and Dichocephala tura (Stuessy, 1972). The first chromoso~e re­ DC., are known also as n = 9. port for M. tepicense of n = 12 fits well wIth the Our two populational counts of n = 8 for sectional base number of x = 12 and corroborates Gutierrezia glutinosa are the first reports of a the placement of the species on morphological tetraploid chromosomal level for this species, criteria. which has been counted before as n =4 (e.g., Our two chromosome counts of n = 18 for Powell and Turner, 1963; Reveal and Spellen­ Parthenice mollis confirm two recent reports from berg, 1976). Infraspecific euploidy is not un­ this species of the same number (Urbatsch, 1975; common in Gutierrezia, and it has been reported Sauck 1975). The firm establishment of a chro­ (Solbrig, 1960a) from G. bracteata Abrams (n moso~e number for the monotypic Parthenice is = 8 and 12), G. microcephala (DC.) A. Gray of particular interest to the junior author, because (n = 8 and 16), and G. sarothrae (Pursh) Britt. in a previous paper (Stuessy, 1973) the genus & Rusby (n =4 and 8). was transferred from the subtribe Melampodiinae Two chromosomal base numbers are known in to the Ambrosiinae near Parthenium based on Xanthocephalum: x = 4 and 6. Our first report overall morphological resemblance. The chromo­ of n = 6 for X. benthamianum fits well with its some counts of n = 18 for Parthenice fit well with close morphological relationship to x.. gymno­ the presumptive base of x = 18 for the entire Am­ spermoides (A. Gray) B. & H. (SolbrIg, 1961) brosiinae (Payne, Raven, and Kyhos, 1964). which is also n = 6 (e.g., Solbrig, 1960b). How­ Recent pollen studies on Parthenice (Bolick and ever, X. benthamianum also bears a strong .over­ Skvarla, 1976) do not contradict this placement all morphological resemblance to some speCIes of in the Ambrosiinae. Grindelia (x = 6). The possibility that the spe­ The previous chromosome counts (n = 15, 30, cies of Xanthocephalum with a base of x = 6 are ca. 45, ca. 86) in Perymenium are based clearly more closely related to Grindelia than to the x on x = 15. Our first report of n = 30 for P. = 4 species of Xanthocephalum needs to be ex­ buphthalmoides (var. tenellum) suggests a tetra­ amined critically. ploid species. Two previous counts of n = 15 H eliantheae-Many chromosome counts have (Turner, Beaman, and Rock, 1961) and n = 45 been made in Bidens, and the genus is clearly on (Strother, 1976) have been reported for Pery­ a base of x = 12. Bidens reptans var. urbanii has menium mendezii. Our count of n = 30 com­ been counted before at the tetraploid level of n = pletes the euploid series within this taxon.
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