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Chromosome Studies in Asteraceae from the United States, Mexico, the West Indies, and South Americaamerica'1

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Chromosome Studies in Asteraceae from the United States, Mexico, the West Indies, and South Americaamerica'1 Amer. J. Bot. 75(5): 652-668. 1988. CHROMOSOME STUDIES IN ASTERACEAE FROM THE UNITED STATES, MEXICO, THE WEST INDIES, AND SOUTH AMERICAAMERICA'1 DAVID J. KEIL, MELISSA A. LUCKOW, AND DONALD J. PINKAVA BiologicalSciences Department, California Polytechnic State University, San Luis Obispo, California 93407; Department of Botany, University of Texas, Austin, Texas 78712; Department of Botany and Microbiology, Arizona State University, Tempe, Arizona 85287 ABSTRACT Chromosome counts of Asteraceae are reported from Mexico, the United States, the West Indies, Peru, and Bolivia. First counts are reported for 27 species, eight infraspecific taxa, and three interspecific hybrids in Brickellia,Briekellia, Chrysanthellum, Cirsium, Egletes, Erigeron,Flaveria, Gnaphalium, Heterotheca,Heterotheea, Hieracium, Hymenothrix, Koanophyllon, Layia, Lessingia, Peetis,Pectis, Sclerocarpus, Stuessya, Tagetes and Wedelia. Counts are also reported for 196 taxa or hybrids for which chromosome numbers have been publishedpreviously. Of these, nine are new num­num- bers. Taxonomic implications of certain counts are discussed. THIS PAPER is a continuation of studies (Keil meiotic microsporocytes and were document­document- and Stuessy, 1975, 1977; Keil and Pinkava, ed by camera lucida or free-hand drawings. 1976, 1979, 1981; Keil, 1977; Pinkava and VoucherVoucher specimens, microslides, and drawings Keil, 1977) on chromosome numbers in As­As- are deposited at ASU and OBI. Percent pollen teraceae. VVouchered ouchered chromosome counts rep­rep- stainability, based on 500 grain samples stained resent both hitherto unreportedtaxa and taxa in aniline blue in lactophenol, was calculated for which counts have been published previ-previ­ for some plants. ously. The former counts serve to increase the number of taxa for which cytological data are RESULTS-RESULTS-TheThe chromosome counts ob­ob- available and can provide insights into rela­rela- tained are listed in Table 1. First counts are tionships and patternsof evolution. The latter reportedfor 27 species, eight infraspecific taxa, counts provide information about variation in and three interspecific hybrids; counts are also chromosome complements and broaden the reported for 196 additional taxa or hybrids for geographical sample of the various taxa. which counts have been published previously. Eight of these are new numbers. METHODs-ImmatureMETHODS-Immature capitula were killed DISCUSSION-As in previous papers, the dis­ and fixed in modified Carnoy's fixative [4-6 papers, the dis- cussion is restricted to first or new counts or chloroform: 3 ethanol: 1glacial acetic acid, v:v: significant comments. The sequence of Table v]. Florets were stained in either iron-aceto­iron-aceto- sequence Table 1 and the accompanying commentary is al­ carmine or Snow's stain (Snow, 1963) and an­an- accompanying commentary is al- (Snow, 1963) phabetical by tribe. thers or whole florets were squashed in a drop tribe. of Hoyer's medium in a modification of the Astereae-Astereae-Our Our report of 2n = 99II for Egletes methods of Beeks (1955). Chromosome counts n for Egletes prostrata and the only previous report for the and meiotic behavior were ascertained from genus, n = 27 for E. viscosa (L.) Less. var. dissecta Shinners (Keil and Stuessy, 1977), sup­sup- I port a base number of x = 9 for the genus. I Received for publication 17 March 1987; revision ac­ac- Our first report of 2n = 9 for cepted 14 July 1987. report of 2n = 911n for Erigeron bel­bel- Research was supported by NSF Grant DEB 81-04683. lioides is in accord with the established base We thank John Popenoe, Director of the Fairchild Tropical number, x = 9, for Erigeron. The head mor­mor- Gardens, for making facilities available during Dr. Keil's phology of this diminutive plant, however, field work in the West Indies; and George Proctor and suggests that it might better be placed into Co­Co- Peter Bretting for assistance in the field in the Dominican Republic and Jamaica, respectively; Judith M. Canne for nyza, which has the same base number. Cron­Cron- providing bud samples from Bolivian plants; Michael O.0. quist (1943) listed the following features as Dillon for providing seeds of Peruvian plants; and John diagnostic for Conyza: "Central hermaphro­hermaphro- C. Semple for determinations for HeterothecaHeterotheea and Pityopsis dite flowers few; pistillate flowers numerous, species. We particularly thank R. K. Brown for allowing us to publish results from his research on Tragopogon. We with filiform corollas; ligules, if present, in­in- thank Billie L. Turner, John C. Semple, and John L. Stroth­Stroth- conspicuous, shorter than the tubes and scarce­scarce- er for reviewing the manuscript. ly if at all exceeding the pappus." The heads 652 May 1988]1988] KEIL ET AL.-CHROMOSOMEAL. -CHROMOSOME STUDIES ININ ASTERACEAE 653 TABLE 1.1. Chromosome counts of Asteraceae. Symbols: * Indicates first report for a taxon or hybrid combination. ** Indicates previously unreported chromosome countforcount for a taxon. Abbreviations of collectors: MGK = Malcolm G. McLeod, Timothy Gaskin and David Kei/; Keil;K = David Keil; KB = David Keil and Peter Bretting; KL = David Keil and Melissa Luckow; KLL = David Keit,Keil, Melissa Luckow and David Luckow; KPr = David Keil and George R. Proctor; L = Elinor Lehto; MK = Malcolm G. McLeod and David Kei/; Keil;MMK = Malcolm G. McLeod, Henry G. Meyer and David Keil; P = Donald J. Pinkava Tribe Chromosome count Location and voucher Taxon or hybrid combination ANTHEMIDEAE Achillea millefolium L. 2n = 1818,,n UNITED STATES: Virginia: Smyth Co.: White Top Mtn, K12820 pop. (OBI)(OBI) Artemisia californica Less. 2n = 9 11 UNITED STATES: California: Kern Co.: Blue­Blue- stone Ridge below Orchard Peak, MGKl1846MGK11846 (OBI) ASTEREAE = Acamptopappus sphaerocephalus (H. & 2n = 99IIIl UNITED STATES: California: Los Angeles Co.: 5 G.) A. Gray mi W of San Bernardino Co. line on Rt 138, Kl1894KJ 1894 pop. (OBI) Astranthium condimentum DejongDeJong 2n = 33,,n MEXICO: Michoacan: Rt 37,337, 3 mi S ofjctof jct with Rt 15, KL15167 (OBI) Chrysopsis pilosapi/osa Nutt. 2n = 4411u UNITED STATES: North Carolina: Cumberland Co.: Fort Bragg, Kl1650K11650 (ASU) Chrysothamnus nauseosus (Pall.) Britt. 2n = 9911I1 UNITED STATES: California: Los Angeles Co.: 5 ssp. hololeuca (A. Gray) Hall & mi W of San BernardinoBemardino Co. line on Rt 138, Clem. Kl1895K11895 pop. (OBI) Chrysothamnus viscidiflorus (Hook.) 2n = 99,,11 UNITED STATES: Wyoming: Natrona Co.: Al­Al- Nutt. cova, K12870 (OBI) Conyza canadensis (L.) Cronq. n = 9 UNITED STATES: Arizona: Maricopa Co.: McDowell Mtns, Lane 1737 (ASU); California: Monterey Co.: 6 mi N of Castroville, K10945 (OBI, ASU); Montana: Carbon Co.: Devil's Canyon area, R. K. Brown 491-1 (ASU) ConyzaConyzafilaginoidesfilaginoides (DC.) Heiron. 2n = 991111 MEXICO: Mexico: 4.5 mi W of Santa Maria del Monte, KL15413A (OBI) Conyza aff. sophiifolia H.B.K. n = 9 MEXICO: Durango: Rt 40, 19.5 mi E of La Ciu­Ciu- dad, P13399 (ASU) CorethrogyneCorethrogynefilaginifoliafilaginifolia (H. & A.) 2n = 55,111 UNITED STATES: California: San Luis Obispo Nutt.Nutt. var. robusta Greene Co.: Morro Bay, MMK11856MMKl1856 (OBI) **EgletesEgletes prostrata (Sw.) Ktze. 2n = 9911I1 ANTIGUA AND BARBUDA: Antigua: St. George's Parish: Windthorpes Bay, K16554 pop. (OBI) Ericameriaarborescens (A. Gray) 2n = 911911 UNITED STSTATES:ATES: California: San Luis Obispo Greene Co.: Cuesta Grade, K11855Kl1855 pop. (OBI) **ErigeronErigeron bellioidesbel/ioides DC. 2n = 9 11 PUERTO RICO: Isla de Cabra, K16503 pop. (OBI) Erigerondivergens T. & G. 2n = 3x = 27 UNITED STATES: Arizona: Santa Cruz Co.: Ari-Ari­ zona Rt 289 S of PefiaPeiia Blanca Lake, L20379 (ASU) [pollen stainability 23.8%; macropollen and micropollen present]; Yavapai Co.: 1-17,I-17, 0.5 mi N of Sunset Point, L19918 (ASU) [pol-[pol­ len stainability 16.4%; macropollen and micro-micro­ pollen present] ErigeronflagellarisErigeron flagellaris A. Gray 2n = 9119n UNITED STATES: Arizona: Coconino Co.: San Francisco Peaks, K11516Kl1516 (ASU) = *Erigeron*Erigeron foliosusfoliosus Nutt. var. blochman- 2n = 9,,9 11 UNITED STATES: California: San Luis Obispo iae (Greene) Hall Co.: MontafiaMontana de Oro State Park, MMK1MMKl1861 1861 (OBI) ErigeronformosissimusErigeron formosissimus Greene var. 2n = 91,9 11 UNITED STATES: Arizona: Coconino Co.: Mt. formosissimusformosissimus Elden, K11515Kl1515 (ASU) Erigeronplatyphyllus Greene 2n = 9119I1 UNITED STATES: Arizona: Coconino Co.: San Francisco Peaks, P13718P13718 (ASU); West Fork, Oak Creek L20218 (ASU) Erigeronstrigosus Muhl. 2n = %9 11 UNITED STATES: Virginia: Smyth Co.: White Top Mtn K12823 (OBI) 2n = 3x = 27 UNITED STATES: North Carolina: Cumberland Co.: Fort Bragg, K11636Kl1636 (ASU). Pollen stain- 654 AMERICAN JOURNAL OF BOTANY [Vol.[Vol. 75 TABLE 1.1. Continued Tribe Chromosome count Location and voucher Taxon or hybrid combinationcombination ability 25.9%; macropollen and micropollen present Grindelia inuloides Willd. 2n = 66,,11 MEXICO: Jalisco: Ajijic, KL15602 (OBI)(OBI) Grindelia procera Greene 2n = 101110lo + 1llvIv + 2 UNITED STATES: California: Kern Co.: Blue­Blue- fragments stone Ridge below Orchard Peak, MGKl1842MGKJ 1842 (OBI) Gundlachia corymbosa (Urb.) Britt. 2n = 991111 ANTIGUA AND BARBUDA: Barbuda: along sand road to Palmetto Pt, K16562A (OBI) Gutierrezia california (DC.) T. & G. 2n = 88,,11 UNITED STATES: California: Kern Co.: Keck Rd, 4.3 mi SE of jct with Baker Rd and Twis­Twis- selman Rd, MGKl1839MGK11839 (OBI) Gutierrezia wrightii A. Gray 2n = 44,,II UNITED STATES:
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