Illinois Natural History Survey bulletin "7^ FEB 1 1990 DEPOSITORY UBRARY DEC ' 1989 UNlvtKiil r ui- ILLINOIS AT URBANA-CHAMPAIGN Aster and Brachyactis in Illinois Almut G. Jones Illinois Natural History Survey Bulletin Volume 34, Article 2 Aster and Brachyactis in Illinois Almut G. Jones Department of Plant Biology University of Illinois at Urbana-Champaign Illinois Natural History Survey Bulletin Volume 34, Article 2 May 1989 Illinois Natural History Survey. Lorin I. Nevling. Chief A Division ot the Illinois Department of Energy and Natural Resources A catalog of the publications of the Illinois Natural History Survev is available without charge from the address below. A price list and an order blank are included with the catalog. Illinois Natural History Survey Distribution Center Natural Resources Building 607 East Peabody Drive Champaign, Illinois 61820 Printing costs for this publication were paid for in part b\ the Department of Plant Biology. I'nivcrsily of Illinois at Llrbana-Champaign. The cover tirawing was executed by Alela Holt. Editor: Audrey S. Hodgins Citation: Jones. A.Ci. \'-W). ,\.\ui and tiiihlntn li^ in Illinois. Illinois Natural Historx Suney Bulletin 34(2):I,W-144. USISSN()07.V49I8 Printed by Authority of the State ol Illinois ( 72724- :M .S-S^)) Contents Introduction 1 39 Generic Description of Aster L. 143 Conspectus of Classification of Illinois /A.svt'/' Species 144 Key 10 Species of Asieidnd Brachyacti.s in Illinois 145 Descriptions of Aster Species 149 Aster aiiiinuiliis Engelm. in Torrey & Gray 149 Aster horeulis (Torrey & Gray) Prov. 150 Aster cdrJifdlltis L. 151 Aster ciliolaius Lindley in Hooker 152 Aster dnimnunnlii Lindley in Hooker 153 Aster diinutsus L. var. striciior Torrey & Gray 1 54 Aster envoules L. 155 Asterfalcatus Lindley in Hooker var. vomiuutunis (Torrey & Gray) A.G. Jones 156 Aster fra^iiUs Willd. 157 Aster fitrvatiis Burgess in Britton & Brown 158 Aster laevisi^. 159 Aster lanceolatus Willd. 1 60 Aster Uiteriflorus ( L. ) Britton 1 62 Aster liiuiriifolius L. 163 Aster nuicrophyllus L. 164 Aster novae-angliae L. 165 Aster X amethystiims Nutt. 1 66 Aster ohlnnf^ifolius Nutt. 166 Aster ontarionis Wieg. 167 Aster oolentangiensis Riddell 168 Aster purviceps (Burgess in Britton & Brown) Mack. & Bush 169 A ster patens A i ton 170 Aster pilosusV^'Md. 171 Aster praealtus Poir. 173 Aster prenantlwides Muhl. ex Willd. 174 Aster puniceus L. 1 75 Aster schreheri Nees 1 77 Aster serkeus Vent. 1 78 Aster shortii Lindley in Hooker 1 79 Aster tatciricits L.f. ISO Aster turhinctlus Lindley in Hooker 181 Aster itmhellatus Miller 1 82 Aster iindiilcitits L. 1 83 Aster urophyllus Lindley in DC. 1 84 Description of Brachyactis Species 1 86 Rrachyaclis ciliatci ( Ledcb. ) Ledeb. 1 86 Glossary of Descriptive Terms 1 87 Literature Cited 190 Index to Scientific Species Names, Including Synonyms 193 Index to Vernacular (Common) Names 194 Cdunly reference map to be used with species distribution maps. Aster and Brachyactis in Illinois Introduction Almut G. Jones The need for a comprehensive study and taxonomic revision of asters in Illinois becomes apparent when one looks at the varied treatments of this genus in the principal tloristic literature of the state (G.N. Jones 1945. \^5{). 1963; Jones and Fuller 1935: Mohlenbrock 1975. 1986; Mohlenbrock and Ladd 197X|. Three factors contribute to the differences found in these works: the number of taxa recorded for Illinois, nomenclatural considerations such as those dictated by the principle of priority, and differences in taxonomic concepts and interpretations. In the first edition of FItira of Illinois (G.N. Jones 1945). .V^ species are treated under Aster, not counting hybrids. Three additional species appear in the third edition (G.N. Jones 1963): A. cliasei G.N. Jones in Jones & Fuller, a species newly described from Illinois: A. lakiricus L.f.. an occasion- ally escaped cultivated species; and A. parviveps (Burgess in Britton & Brown) Mack. & Bush, probably inadvertently omitted from the first two editions but included in Jones and Fuller (1955). In the most recent (iuidc lo llic \ asciilar Flora of Illinois. Mohlenbrock (1986) treats 35 species. A notable change involves the transfer of ,4, piarmicoides (Nees) Torrey & Gray to Solida^o. In this revision, I recognize 3 1 species of Aster in the state of Illinois. My research has concentrated on New World and some Old World species o\ Aster (sensii lata) forever 15 years (A.G. Jones 1974, 1977, 1978a. b, c. I98()a, b, 1982. 1983, 1984, 1987; Jones and Hiepko 1981; Jones and Young 1983; Jones and Lowry 1986; and others), and much additional infomialion has been introduced in this current study. My taxonomic concepts are summarized in the conspectus of classification of Illinois Aster species (p. 144). A recent example of a name change necessitated by application of the principle of priority is that of Aster aziireiis Lindley in Hooker. The name is placed in synon- ymy under /^. oolcntansiiensis Riddell (A.G. Jones 1983). Publication of the latter name preceded that of the former by only seven months. Other examples of name changes are A. junciformis Rydb. to A. horeulis (Torrey & Gray) Prov., and A. panioiriihiis S.F. Blake Xo A. oniurionis Wieg. (Shinners 1949). Most differences among various accounts are attributable to disagreements about taxonomic concepts and interpreta- tions. Changes may be the result of the union of two or more species under one [e.g.. Aster siniple.\ Willd. = A. lanceoki- tus Willd. (Semple 1979; Semple and Chmielewski 1987)|. or they may involve elevation lo ihe rank of species of taxa formerly considered at an interior rank [e.g.. A. ericoiih's L. ( var.) parviceps Burgess in Britton & Brown = A. parv- iceps]. Sometimes names have been misapplied (e.g.. /\. crivoides for A. pilosiis Willd. (cf. Blake 1930)1. and occasionally a species is transferred to another genus [e.g.. A. piarmicoides to Solidai;o (Boivin 1972) and A. hrailiyactis S.F, Blake to Braclna( tis (A.G. Jones 1984)1. The transfer of /4.s7<'//>?(;/7/)/idus (Raf. ) A.G. Jones (.v = 5 chromosomes), have not been widely adopted. Evidence of hybridization between species of that assemblage and members of A. subg. Symphyolricluini (Nees) A.G. Jones (.V = 8 chromosomes) supports a more conservative generic concept (A.G. Jones 1982: Jones and Young 1983: Allen 1985). The following chapters are organized in the fonnat used in most tloristic manuals. The generic description of Aster {sensu strict/)) is followed by a conspectus of classification of the Illinois species and h\ an indented key that includes both Aster and Brachyactis. Ne.xt. the species descriptions are given in alphabetic order by .scientific name, the last being that of Brachyactis ciliata. Within each entry, vernacular names used in literature dealing with the Illinois flora are listed after the scientific name. A new vernacular name is proposed (with some reluctance) in this work for Aster fragilis Willd.: Brittle Aster, a translation of the very appropriate specific epithet. As much as possible, a uniform format is maintained in the descriptions. TTie diagno- sis is followed by a listing of the mitotic (2n) chromosome number! s) and by those synonyms of the specific name that are cited for Illinois in other floristic works, including the bibliographic references. A separate paragraph deals with phenol- ogy, ecology, and general geographic distribution. Standard two-letter postal abbreviations are used for names of the states and of the Canadian provinces, for example. AB (Alberta). NF (Newfound- land). PE (Prince Edward Island), PQ (Quebec), and YT (Yukon Territory). Dot maps show the distribution of native Illinois species at the county level (county names are identified on the preceding map). These maps were based entireh on speci- mens I have examined, annotated, and recorded, and I may v\ell ha\ e missed some counties for which valid records ha\e been published. Previously published maps and lists, however. ma\ include occasional recordings based on niisidentifications. In addition to my own collections. I have examined more than 10.000 herbarium specimens for this study, including all material on deposit in the principal Illinois herbaria: DEK. EIU. F. ILL. ILLS. ISM. KNOX. MOR. MWI. and SIL . Abbrevia- tions correspond to those in Inde.x Herhari- onim (Holmgren et al. 1981 ). GH. MIN. MO. N^\ PH. L'S. and WIS are among the larger herbaria outside the state that yielded distribution data for the asters of Illinois. Some personal collections were borrowed trom P. Shildneck and J. Schwegman. Varieties are treated following the description of the respective sjx'cics. either Mas lysy A',11 H WD HK\( II) \l ll\ IS II I 1M)IS 141 in a shon diagnostic statement or, in cases w here a clear distinction by a combination of characteristics is possible, in the forni of a key. Varietal synonymies are listed only to the extent that they have been cited for or described from Illinois. One reason for the inconsistency in the handling of varieties is that keys are not very meaningful or practical when a high degree of intergrada- tion or overlap in characteristics exists. Moreover. I am convinced that some commonly recognized varieties have acquired their distinctive characteristics via gene flow from other species. Hybrids are mentioned at the end of each entry, and some herbarium collections are listed as examples. Although only a few of the putative hybrids observed in the wild have been experimentally reproduced, the fact that closely related Aster species have a high degree of genetic homology and do occasionally hybridize is now generally accepted (Wiegand 192X. 1932: Cronquist 1947: Semple and Brammall 1982: and others). The classic example of an experi- mentally confirmed hybrid that may survive for several seasons is /A. x amethystinus Nutt.. pro sp. = A. ericoides x A. navae- angliae L. (Wetmore and Delisle 1939: A.G. Jones 1978c). At least some viable achenes are commonly produced, and backcrosses to plants of the parent species have been successful. Because of the relatively frequent occurrence of this hybrid. I have included it in the key and given a compara- tive description after that of one of its parent species {A. novae-angUae). Herbarium specimens of putative hybrid individuals, however, are not entirely uniform, and some are undoubtedly backcrosses to one or the other parent. Intergradation between two species is more evident in some cases than in others, and the key to the species gener- ally does not work well for hybrids or for individuals that exhibit a high degree of introgression from other species. Aside from the hybrid Aster x amethys- tinus. I have included in the key and discussed under their closest relatives two adventive species that may or may not be established in Illinois: A.fakatus Lindley in Hooker (under A. ericoides) and A. ciliola- tiis Lindley in Hooker (under /A. cordifolius L.). I have also included the exotic species A. tcitciriiiis because it is treated in current manuals dealing with the flora of Illinois (Gleason 19.'S2: Gleason and Cronquist 1963: G.N. Jones 1963: Mohlenbrock 1975. 1986). Three species of Aster are included in the list of threatened species of Illinois issued by the Department of Conservation on March 14, 1989: A.furcalus Burgess in Britton & Brown, A. schreheri Nees. and -4. iindulatus L. In my opinion. A. parviceps should be considered for inclusion in the list. The latter species and A.furcalus are true midwestem endemics, and they may be threatened because the plants grow in vulnerable habitats. Although the number of locality records is considerable and some populations are extensive, the.se two species are probably not as common today as they used to be. I have been unsuccessful in my search in many places where they had been previously collected. The other two asters currently listed as threatened may be rare in Illinois, but they are very common in some other areas. Illinois populations represent the western limit of range for these species. Aster schreheri presents a special situation because Illinois and Wisconsin populations are disjunct from the nearest populations to the east and have been named and described as A. chasei (Jones and Fuller 1955). So far I have been unable, however, to find bio- logical evidence that would support taxonomic recognition of these disjuncis as distinct from typical A. schreheri. even at the varietal rank. Other species that ap- proach their limit of range in Illinois and are much more common outside the state are A. horealis. A. dumosus L.. /4. macrophxihis L.. and A. prenanthoides Muhl. ex Willd. To help resolve some of the difficulties of species delimitation that have haunted earlier accounts cif Aster. I have made my keys and descriptions more detailed than 142 Illinois Natlral History Si rvf.v Bllletis Vol. 34 An. 2 those generally found in tloristic works. Many structural features that provide significant taxonomic information are often given short treatment because they are seasonal (e.g., those of mature fruits) or are not always available in herbarium specimens (e.g., rhizomes and basal rosettes). Because I have grown plants of all the species de- scribed here and observed them through the seasonal cycle, I have been able to charac- terize all structural elements. In addition to providing more complete descriptions of the species, this treatment will allow, at least in some cases, the detection of hybrids and populations showing intergradation. The taxonomic method is an ongoing endeavor, and revisions are needed as more and more is learned about the genetics and natural history of the taxa. No doubt, the definitive account of Aster in Illinois will continue to elude us. Acknowledgments The loan of specimens and help received from the curators and collection managers of the following herbaria are acknow ledged: DEK. EIU. F. GH. ILLS. ISM. KNOX, MIN. MO. MOR. MWI. NY. PH. SIU. US. and WIS. Thanks for testing the key and checking some records goes to P. Shildneck (Decatur) and A.C. Koelling (Illinois State Museum. Springfield). I appreciate the constructive criticisms of T.M. Barkley (Kansas State University. Manhattan) and K.R. Robertson (Illinois Natural History Survey. Champaign), who refereed my manuscript. I am especially grateful to W.F. Lamboy (University of Illinois at Urbana- Champaign) for many good suggestions regarding this study and to my assistant Laurel L. McKee for help w ith questions on and proofing of the manuscript. May 1989 AsiF-H AND Bum III Ai IIS in Ii i inois 143 Generic Description of Aster L. Starwort, Wild Aster, Michaelmas Daisy. Frosl-tlower. Herbaceous or somewhat suffrutieose perennials with caudicit'omi or short to long-creeping stolonilonn rhi/ome systems but also including some taprooted annuals (all native Illinois asters are perennial). Stems erect or decumbent, terete in cross section, or slightly ridged at the upper nodes from decurrent leaf bases, single to few at any point of emergence or cespitose, variously branched, glabrous to densely pubescent, sometimes glandular. Leaves simple, alternate, usually polymorphic, those of the basal rosettes and the lower stem often pctiolate. those of the middle or upper stem and of the branches (rameal leaves) commonly subsessile or sessile; blades variously shaped, glabrous to densely pubescent, sometimes glandular, the apex acute, acuminate, attenuate, or obtuse to rounded, the margins entire, crenate, or serrate, never lobed, the base cuneate to rounded or cordate, sometimes sheathing or auriculaie-clasping. Capitulescence paniculifonm. corymbifomi, racemiform, or rarely spicifonn. Heads on short to long peduncles (sometimes subsessile). often secund from leaf axils of the temiinal branchlets. Involucre cylindric. turbinate (often an artifact in pressed and dried specimens), campanulate, or hemispheric, the bracts (phyllaries) in several series. Phyllaries iinbricated and usually gradu- ated, occasionally subequal in length or the outer ones longer than the inner, commonly with a well-delimited apical green areole and a scarious. often indurate or keeled basal portion, or sometimes with green bands extending along the midrib to the base; phyllaries of the outer scries often largely herbaceous. Receptacle flat or slightly convex, alveolate with blunt or sharp (rarely bristle-tipped) teeth or merely pitted in older, fruiting heads. Ray florets several to many, rarely absent or vestigial, mostly in 1 but sometimes in 2 or more series, pistillate and fertile, the strap-shaped or oblong ligule portion of the corollas white, pink, lavender, purple, or blue, never yellow . the tube portion yellowish green when fresh. Disk florets few to many, perfect and fertile, the corollas regular, cylindric or funnelfomi. usually with well-delimited tube and limb portions, the limb .S-lobed. cream- colored or yellow, often appearing brighter yellow before anthesis because of the presence of yellow pollen, commonly aging to pink or purple after the pollen is shed, the tube usually slender, yellow or greenish. Style branches of the disk florets with attenuate, papillate, sterile appendages that are longer than the stigmatic lines. Pappus often simple, composed of a single series of relatively uniform, barbellate, capillary bristles, or if more complex, either with 2 series of capillary bristles more or less uniform in length but with those of the inner series clavellately expanded toward the apex, or with an additional (third) outer series of distinct, very short, attenuate bristles (a feature conventionally referred to as a "double pappus"). Achenes plump or somewhat compressed, 2-5 times longer than wide, gray or straw-colored to purple or brown, glabrous to variously pubescent, with a species-specific, definite number (.'-10) of ribs, these sometimes obscure. Basic chromo- some numbers; v = 5. 8. and 9. Excluded are Aster hiachyactis S.F. Blake [= Brachyaclis ciliala (Ledeb.) Ledeb.] and A. piarmicoides (Nees) Torrey & Gray |= Solidago plarmicoidcs (Nees) Boivin]. Asler can be distinguished from closely related genera (tribe Astereae) only by a combination of characteristics: sterile append- ages of the style branches attenuate, longer than the stigmatic lines; phyllaries in .^ or more usually graduated scries; rays never yellow; principal pappus bristles more or less unifomi in length, not or scarcely overtopping the disk corollas; achenes usually with a definite number of ribs, not densely striated; flowering period typically late summer and fall. Approximately 200 species, worldwide, mostly in the northern hemisphere, with the center of distribution in temperate North America. Taxonomic boundaries well defined in most species but blurred in some, largely as a result of interspecific hybridization, which is sometimes compounded by polyploitly. Moreover, the species exhibit a high degree of phenotypic variability, and similar morphologies may have evolved from different biological origins. Because many questions regarding the taxonomy of \stcr cannot be answered at this time, the species concepts adopted in this treatment are relatively conservative. 144 Illinois Nati ral History Slrvev Bi lletiv Vol. 34 An. 2 Conspectus of Classification of Illinois Aster Species Aster L. subgenus Asier section Biotia DC. ex Torrey & Gray A.funariis Burgess in Britton & Brown A. macrophyllus L. A. schreheri Nees sect. Macrocephali (Kitaniura) A.G. Jones A. taiaricus L.f. subg. Doellingeria (Nees) A. Gray sect. Triplopappus (Torrey & Gray ) A.G. Jones A. umbellatus Miller subg. laiithc (Torrey & Gray) A. Gray A. linariifolius L. subg. Symphyolrichum (Nees) A.G. Jones sect. Concinni (Nees) Torrey & Gray A. laevis L. A. oolentangiensis Riddell sect. CordifoUi G. Don in Loud. A. anonuiliis Engelm. in Torrey & Gray A. ciiiolaius Lindley in Hooker A. cordifalius L. A. dnnnmondii Lindley in Hooker A. shonii Lindley in Hooker A. iindulalus L. A. iinipliyllus Lindley in DC. sect. Dumosi Torrey & Gray A. dumosiis L. A./ra,i,'/7/.vWilld. A. lalerifloiiis (L.) Britton A. onlarionis Wieg. sect. Porlcriani (Rydb.) A.G. Jones A. pcirviccps (Burgess in Bntlon & Brown) Mack. & Bush A. pilosiis Willd. sect. Salicifolii Torrey & Gray A. borealis (Torrey & Gray) Prov. A. Uimcolaliis Willd. A. pracalliis Poir. A. preihiiilhiudcs Mulil. c\ Willd. A. puniceiis L. sect. Turbinelli (Rydb.) A.G. Jones. Stat, and comb, nov.* A. lurhinelhis Lindley in Hooker subg. Virguliis (Raf. ) .A.G. Jones sect. Comolores Torrey & Gray A. sericeus Vent, sect. Eriioidei (Nees) Torrey & Gray A. ericoides L. A.fakatus Lindley in Hooker sect. Ohlongifotii (Rydb.) A.G. Jones A. novae-angliae L. A. ohlongifolius Nutt. sect. Patentes Torrey & Gray A. patens Alton Based on Aster (species group) Tiirhiiwlli Rydb.. Flora of the Prairies and Plains. 803. 804. 1932. Syn.: A. sect. Eucephahis subsect. Turbinelli (Rydb.) A.G.Jones ( 1980a). Type species: A. itirhincllus Lindlcv in Hooker. May \9H9 Ami h \\ii Bkm id \i lis \s h{ wins 145 Key to Species of Aster and Brachyactis in Illinois 1 . Plants annual with a taproot; rays absent or vestigial; pappus conspicuous at anthesis, considerably overtopping the corollas Brai Iixik lis cilialo 1. Plants perennial (Illinois species); rays typically conspicuous; pappus relatively inconspicuous at anthesis, not or barely overtopping the disk corollas \AsU'r\. 2. Basal and lower cauline leaves long-peliolate, the blades well delimited, with mostly cordate to truncate or sometimes rounded bases. .^. Capitulescence corymbifomi or cymilomi; median phyllaries mostly obtuse to rounded at the ape.x; disk corollas with the apparent tubular portion distinctly longer than the Hared or abruptly expanded portion of the limb (includmg the lobes); achenes fusiform, with 7-10 (rarely more) ribs. 4. Plants conspicuously glandular in the capitulescence 1,^. A. macniphyllits 4. Plants typically not glandular (occasionally with a few glands on the peduncles and phyllaries). 5. Rosettes of basal leaves prominent, often numerous forming a dense ground cover; blades of basal and larger cauline leaves broadly ovate to cordate, glabrous or with sparse indument. the petioles mostly much longer than the blades 24. A. schieheii 5. Rosettes of basal leaves not prominent, relatively few; blades of basal and larger cauline leaves oblong-ovate or lanceolate, harshly scabrous above and densely hirsute below, the petioles usually not longer than the blades 8. A. fuiruius 3. Capitulescence variously paniculiform; median phyllaries acute, acuminate, or attenuate, or if obtuse, the apex with a sharp point, never rounded; disk corollas with the tube portion shorter than the expanded limb portion; achens oblong- obovoid. with 4 or .'S(6) ribs. 6. Phyllaries with strongly squarrose or rellexed tips; heads relatively large with (22)25-35 (or more) rays, the involucre hemispherical I. A. anomahis 6. Phyllaries with appressed or only slightly spreading tips; heads relatively smaller with ca K)-25 rays, the involucre campanulale or cylindrical (or turbinate in pressed specimens). 7. Upper stem densely hirtellous or puberulent, the mdument uniformly distributed around the stem; principal leaves copiously pubescent on both surfaces (or the upper surface glabrous in A. shorlii). 8. Cauline leaves sessile and clasping or with auriculate-clasping petioles 30. A. uiuhiUilus 8. Cauline leaves not clasping, the petioles slender or winged, often somewhat decunrent but never clasping. 9. Leaves of the midstem entire or subcntire; rays mostly ( 12)15-25; heads 1.5-2.5 cm in diameter when the rays arc fully extended or pressed 26. A. shorlii 9. Leaves of the midstem sharply serrate or crenate-serrate; rays mostly (8)10-15; heads 1-1.5( 1.8) cm in diameter when the rays are fully extended or pressed 4. A. diumnmndii 7. Upper stem glabrous or puberulent to villous in thin decurrent lines; principal leaves glabrous or variously pubescent. 10. Phyllaries of the 2nd and 3rd series inward with usually broad, short, rhombic green areoles; leaves of the midstem entire or subcntire. 1 1 . Leaves of the midstem auriculate-clasping, glabrous, glaucous 9. A. laevis 1 1 . Leaves of the midstem not clasping, scabrous above and hirsute below, not glaucous 17. A. <>i>lcnhiii'^icn\i\ 146 h.LrNois Natl RAi History Slrvey Bulletin Vol. 34 An. 2 10. Phyllaries of the 2nd and 3rd series inward with obovate or usually narrower, lance-rhombic or oblanceolate to linear green areoles: leaves of the midstem variously toothed (sometimes subentire \nA. iiroplnlliis). 12. Rays typically white (occasionally faintly lavender); phyllaries with a long-attenuate apex, largely scarious, often only the midrib light green, or the green areoles linear-oblanceolale i\. A. umphyllus 12. Rays typically purple or lavender: phyllaries with variously shaped apices, the green areoles well developed, obovate or lance-rhombic to oblanceolate. 1 ?i. Heads 1-1 .."i cm in diameter when the rays are fully extended or pressed: total number of tlorets 16-.30 (Illinois plants); apex of the phyllaries acute or sometimes short-acuminaie 3. A. cordifolius 1 3. Heads 1 .5-2.5 cm in diameter when the rays are fully extended or pressed; total number of florets 30-50; apex of the phyllaries attenuate or narrowly acuminate 3a. A. ciliolatus 2. Basal and lower cauline leaves sessile to subpetiolate or if apparently petiolate. the blades with a cuneate base or gradually narrowed, not well delimited from the petiolar portion. 14. Pappus with an outer whorl of very short bristles in addition to 1 or 2 series of much longer (principal) capillary bristles; leaves more or less uniform in contour except for the lowermost, which are usually bracteifomi; basal rosettes absent. 15. Leaves rigid, l-nerved, lacking secondary nerves, ca 1-3.5 cm long and 0.4 cm or less wide: stems mostly less than 60 cm tall 12. A. linariifoliiis 15. Leaves flexible, with well-developed secondary nerves, ca4— 12 cm long and 1 cm or more wide; stems mostly more than 60 cm tall 29. A. iimhellatiis 14. Pappus simple, the bristles typically in a single series and more or less uniform in length: leaves usually variable in contour but the lowermost not bracteiform; basal rosettes present at least during part of the growing season. 16. Involucres and peduncles, often also the leaves and upper stems, glandular. 17. Rays (40)50-100: outer and median phyllaries narrowly acuminate to long-attenuate, strongly squarrose; cauline leaves auriculate- clasping 14. A. novae-angliae 17. Rays 12-35; outer and median phyllaries obtuse or acute to short-acuminate, appressed or squarrose; cauline leaves variously inserted. 18. Cauline leaves auriculate-clasping: phyllaries strongly graduated, appressed or with slightly recurved tips 19. A. paiens 18. Cauline leaves not or only slightly clasping, rounded at the base, not auriculate: phyllaries only slightly or not at all graduated, strongly squarrose 15. A. ohiongifolius 16. Involucres, peduncles, leaves, and stems not glandular. 19. Achenes with 7-10 ribs, purple or brown at maturity: phyllaries pubescent on the outer (abaxial) surface; stem pubescence unifomils distributed around the stem, never in lines. 20. Leaves and phyllaries appressed silvery-silky on both surfaces; ovaries and achenes glabrous 25. .4. sericeiis 20. Leaves and phyllaries strigillose. hirsute, or cinereous-puberulent; ovaries and achenes pubescent. 2 1 . Principal leaves auriculate-clasping: capitulescence an open. divaricately branched panicle: heads mostly on long, stiff peduncles, not crowded and not secund; ra>s purple 19. A. paiens 21. Principal leaves rounded at the base, onl\ slightK or not at all clasping; capitulescence commonh a racemifomi panicle: heads subsessile or on relatively short peduncles, often crowded and secund; rays white or lavender. 22. Rays lavender; involucre hemispherical; pappus tawny or rose-tinued 14a. .4. x ameth\stinus May I'JSy AsifK ,\\n Bmiii)\(iis IS \iA_i\oii 147 22. Rays typically white: involucre cylindrical to campanulate; pappus white or cream-colored. 23. Heads commonly secund; involucres usually less than 4..') mm high; rays 18 or fewer 6. A. ericoides 2y. Heads commonly not secund; involucres 5 mm high or more; rays 20 or more 6a. A.fakatus var. commuialus 19. Achenes with .-5-5(6) ribs, variously colored; phyllaries mostly glabrous on the outer surface (rarely puberulent in a few species); stem pubescence variously distributed. 24. Leaves clasping, often strongly auriculate. 25. Leaves glaucous; stems glabrous or nearly so; phyllaries strongly graduated, fimi, appressed when fresh, the apical green areole usually broad, rhombic, shorter than the scarious basal portion 9. A. lacvis 25. Leaves not glaucous; stems variously pubescent; phyllaries not at all or only weakly graduated, often flexible and leatlike, at least somewhat spreading or sometimes retlexed, the apical green areole longer than the scarious basal portion, or often the outer phyllaries largely herbaceous. 26. Principal cauline leaves serrate to subentire, gradually tapered toward the base; stem variously hispidulous, as well as hirsute, either over the entire length and sometimes coarsely hispid toward the base, or often with only sparse hispidulous indument in the upper portion of the stem, the lower glabrous or nearly so (depending on the variety); phyllaries long-acuminate or attenuate 2.^. A. pun ii ens 26. Principal cauline leaves sharply serrate, abruptly contracted below the middle into an entire-margined, sometimes apparently subpetiolar basal portion; stem hirsute or villous, not hispidulous; phyllaries acute or short-acuminate 22. A. prciuiiilluniles 24. Leaves not clasping. 27. Involucre (7)8-12 mm high; phyllaries in 6-9 series, strongly graduated, blunt-tipped and rounded on the back, largely scarious, the green areoles small, oblong, in the apical 'A -7' portion 28. A. tiiihinclliis 27. Involucre .^-8( 10) mm high; phyllaries in .^-6 .series, graduated to varying degrees, acute to attenuate or if obtuse, with a sharply pointed tip. not rounded on the back, the green areoles of various sizes and shapes. 28. Basal leaves 3-4 times larger than the principal cauline leaves, the blades 10-30 cm long and 5-10 cm wide, gradually tapered into winged petioles 10-40 cm long; phyllaries with a broad green band along the midrib extending to the base 27. A. uilaricus 28. Basal leaves (often withered and deciduous at flowering time) mostly not larger than the principal cauline leaves, often much smaller; phyllaries usually with well-defined green areoles. 29. Phyllaries and ramcal leaves with a spinulose bristle at the apex; rhi,^ome system caudiciform lacking stoloniform strands; new shoots initiated at the base of old stems or as root sprouts; achenes gray, the ribs obscure. 30. Disk florets 20 or more; involucre somewhat urceolate, (4)5-8 mm high 20. A. pUdsiis 30. Disk florets 6-12; involucre cylindric or turbinate, 3-4 mm high 18. A. parviccp.s 29. Phyllaries and rameal leaves often with a sharp callus point at the apex but lacking a spinulose bristle; rhizome system creeping or if short-branched, the new shoots at least in part mitialed at the ends of stolonifomi rhi/ome strands; achenes variously colored at maturity, the ribs mostly prominent. 3 1 . Reticulate tertiary venation usually conspicuous on the lower leaf surface, the veinlels often brownish, the enclosed green areolae isodiamelric; rays purple or lavender 21. A. praiulliis 148 Illinois Natlral History Si rvey Bi lletin Vol. 34 An. 2 3 1 . Reticulate tertiary venation inconspicuous on the lower leaf surface, the veinlels rarely brownish, or the enclosed areolae oblong; rays variously colored. 32. Ovaries and achenes glabrous, purple or brown at maturity, median phyllaries with mostly broad, short, rhombic (rarely lance-rhombic) green areoles: leaves scabrous above and hirsute below 17. A. oolentangiensis 32. Ovaries and achenes strigillose or puberulent. variously colored at maturity; median phyllaries with linear, oblanceolate, or rhombic-obovate green areoles; leaves variously pubescent or glabrous. 33. Lobes of the disk corollas longer than or as long as the fused portion of the limb (not including the corolla tube); lower leaf surface at least somewhat pubescent: achenes gray at maturity. 34. Lower leaf surface uniformly short-pubescent; heads not secund: rhizomes distinctly long-creeping, stoloniform. the plants forming colonial stands, usually in moist ground 16. A. oniaritmis 34. Lower leaf surface villous to hirtellous along the midrib, otherwise usually glabrous; heads secund; rhizomes short, sometimes tangled but not long-creeping, the plants forming scattered individual clumps, mostly m upland situations 1 1. A. laieriflorus 33. Lobes of the disk corollas shorter than the fused portion of the limb; low er leaf surface usually glabrous (rarely with a few trichomes along the midnb or slightly scabrous toward the margins); achenes variously colored. 35. Flowering heads mostly 1.3-2.5 cm in diameter when the rays are fully extended or pressed. 36. Larger cauline leaves 1-3.5 cm in width, no more than 12 times longer than wide, usually sharply serrate; capitulescence a diffusely branched or elongate panicle; achenes gray 10. A. lanceolattis 36. Larger cauline leaves mostly less than 1 cm in width. 12-15 (or more) times longer than wide, shallowly and remotely serrate to subentire: capitulescence a more or less dichotomously branched round- or flat-topped panicle; achenes purple, or gray with purple streaks 2. A. horealis 35. Flowering heads mostly 0.6-1 .5 cm in diameter w hen the rays are fully extended or pressed. 37. Median phyllaries (3rd or 4th series inward) ca tw ice as wide as those of the outer senes. typically obtuse, the green areoles rhombic-obovate to broadly oblanceolate; rameal leaves relatively unitomi in size; peduncles usually at least I cm long (often much longer) with mans bracteiform leaves; subsessile heads relatively few 5. A. diimosiis 37. Median phyllaries slender, less than twice as wide as those of the outer series, acute or attenuate, the green areoles linear to narrowly oblanceolate: rameal leaves notably unequal in size; peduncles \ ariable in length but the bracteiform leaves typically few: subsessile heads often numerous 7, A.frai;ilis Mav \W-I Asn.N AM) Hh u in u //\ is Ii i isois 144 Description of Aster Species 1. Aster anomalus Engclm. in Torrey & Gray Many-rayed Aster. Blue ,^——I — Aster V-Lr-U_ Herbaceous, with sli)ut branehed-caudicifomi rhizomes that turn woody with age. New shoots arising at or near the base of old stems but also sometimes initiated as root sprouts, the plants forming scattered individual clumps. Stems 1-several. erect or ascending, stout, .'>()-120 cm tall, with ascending or spreading branches mostly from above the middle, uniformly hirtcllous or hirsute. Leaves polymor- phic, the basal and larger cauline ones petiolate. those higher up on the stem gradually reduced in size, changing from subpetiolate to sessile: first leaves of new shoots and of the vernal rosettes with oblong-ovale to lanceolate blades 1^ cm long and 1-2 cm wide, usually purple at least on the lower surface, densely hirtcllous, acute or often with obtuse to rounded tips, the margins few-toothed or subentire; principal cauline leaves with ovate to broadly lanceolate blades 4-9 cm long and ( 1 ..'>|2..^-.'i cm wide, with several pairs of usually arching and anastomosing secondary nerves, hirtcllous or hirsute on both surfaces, the apex acute or acuminate with a short callus point, the margins mostly entire to subentire (rarely crenate-serrale), the base cordate, usually with broad sinuses (rarely truncate), the petioles (2)4-S cm long, slender on the lower leaves, narrowly winged on those of the midstem, densely hirsute to pilose; upper leaves of the main stem lanceolate, entire; rameal leaves abruptly reduced, those of the peduncles bracteifonn. 2-6 mm long, mucronulate. uitergradmg with the phyllanes. Capitulescence a usually ample diffuse panicle, the head-bearing branchlcts sometimes racemiform. Flowering heads 2-.'' cm in diameter when the rays are fully extended, on peduncles O..^-.*!! 10) cm long or rarely subsessile. Involucre hemispherical, 5-\0 mm high, the phyllaries imbricated in 4—6(7) series. Phyllaries graduated, slender, linear-lanceolate, strongly retlexcd or squarrose, grayish green, both surfaces typically hirtcllous, the apex long-attenuate, often with purple tips and with a conspicuous bristly mucro. the margins scabrous; outer phyllaries .^—I mm long, ca half as long as the inner, commonly herba- ceous to near the base: median and inner phyllaries green in the apical '/--"/< portion, the basal portion scurious. Receptacle alveolate with sharp or bluntish teeth. Ray florets (22)23-.^.'i (or more), the corollas 10-1 5( 18) mm long, purple or blue (rarely white), glabrous. Disk florets 20-35 (or more), the corollas (4)4.5-.'S..'i mm long, glabrous or with a few trichomes, the limb only slightly expanded, cream-colored or light yellow turning purple after anthesis. the lobe/limb fraction 0.2-0.25. the tube shorter than the limb. Pappus simple, the bristles as long as the disk corollas or slightly shorter, tawny or rose-tinged, slender, attenuate. Achenes oblong- obovoid, somewhat compressed, 2.5-3.5(3.8) mm long and 0.8-1 .2 mm across, deep purple, or brown with purple speckles, glabrous, with 5 or 6 often irregularly spaced straw-colored ribs. 2// = 1 6; reported chromosome counts of 2« = 18 are probably in error (A.G. Jones 1977, 1980b). Late August-October. Sandy or loamy clay soils underlain by calcareous rocks or sandstone: rocky open woods, woody hilltops, and dry ridges. Occasional near and along major rivers in the western half of the state, north to Peoria and Woodford counties. The range for the species includes MO, AR, s.e. KS. and e. OK. There is good evidence in Illinois of intergradation between this species and Aster shorlii (e.g., Winlenlnf't'r //997(ISM) from Alexander Co.. Rexroai H0H4 (ISM) from Menard Co.. and AXi Jones 6NI7 (ILL) from Union Co.|. 150 Illinois Natlral History Slrvey Bllletin Vol. 34 An. 2 2. Aster borealis (Torrey & Gray) Prov. Rush Aster Herbaceous, with very slender, creeping, stoloniform rhizomes. New shoots mostly initiated from the rhizomes the plants forming clonal stands usually interspersed with other vegetation. Stems 1-few at any point of emergence, slender, erect, 3()-80( 100) cm tall, often somewhat zigzag above, branched only in the capitulescence. sometimes red-tinged to slightly glaucous, nearly glabrous or sparsely puberulent in thin decurrent lines (rarely with uniformly distributed indument on the peduncles). Leaves polymorphic, the basal and lower cauline ones commonly withered and deciduous at flowering time; basal rosette leaves subpetiolate, the blades variously linear- oblanceolate to ovate or suborbiculate. 0.5-1.5 cm long and 2-6 mm wide, the secondary venation obscure, both surfaces glabrous or minutely puberulent toward the margins, the lower surface also usually purple-tinged, the apex obtuse or rounded, the margins shallowly crenate, abruptly narrowed to a slightly winged subpetio- lar portion of variable length, the base dilated and sheathing; principal cauline leaves very slender, linear to linear-oblanceolate, (4)6-15 cm long and 2-6(8) mm wide, with usually only the midrib strongly expressed, glabrous or the upper surface minutely strigillose toward the margins and the lower with a few trichomes along the midrib, the apex attenuate and callus-pointed, the margins usually inrolled, entire or remotely and shallowly serrate, smooth or scabrous, the base sessile and somewhat decurrent; rameal leaves relatively few, ascending or spreading, firm or flexible, entire, similar to the cauline leaves in contour and vestiture, the base somew hat sheathing and decurrent; leaves of the peduncles flexible, not phyllarvlike, linear-lanceolate, 4—12 mm long, often red-tinged, and \k ith a slightly dilated or rounded base. Capitulescence an open, round- or flat-topped, more or less dichotoniously branched, tenninal panicle, rarely also w ith ascending lateral branches initiated in leaf axils of the midstem. Flowering heads ( 1 .5)2-2.5 cm in diameter when the rays are fully extended, not crowded and not secund, typically on slender puberulent or glabrous peduncles 1-5 cm in length. Involucre broadly campanulate, (5.5)6-7 mm high, the phyllaries imbricated in 4—6 series. Phyllaries appressed when fresh or rarely the outermost slightly recurved, subequal or somewhat graduated; outer phyllaries oblong- lanceolate, 3-5 mm long, the innermost slender, linear, 5-6 mm long, the green areoles changing from slenderly oblanceolate to nearly linear, the surfaces glabrous or nearly so, the apex acute, callus-pointed in the outer phyllaries. often red- tipped in the median and inner ones, the margins irregularly ciliolate (rarely villous), orerose- hyaline and with a scarious or reddish rim in the median and inner ph> Maries, the basal h.-i' portion scarious. Receptacle shallowly alveolate w ith sharp teeth. Ray florets 20-30 (or more), the corollas 10-12 mm long, mostly white, sometimes pink or bluish, glabrous or with a few trichomes at the throat and on the tube. Disk florets 25-30 (or more), the corollas funnelform. 5-6 mm long, glabrous or nearly so. the limb abruptly expanded at the throat, cream-colored or pale yellow turning pink after anthesis. the lobe/limb fraction 0.2, the tube shorter than the limb. Pappus simple, the bristles approximately as long as the disk corolla, whitish, soft, slender, and attenuate. Achenes oblong or oblanceolate in contour or often somewhat talcate. plump or slightly compressed, (2.2)2.5-3 mm long. 4-5 times longer than w ide. purple, or gray with purple streaks, thinh strigillose, with 4 or 5 straw-colored ribs. In = 16, 32 (niostlv ), 48. {A.juncciis misapplied, not of Alton—Brendel 1887; Pepoon 1927; Deam 1940. A.junciformis Rydb.—G.N. Jones 1945. 1950. 1963; Femald 19.50; Gleason 1952; Jones and Fuller 1955; Gleason and Cronquist 1963; Swmk 1974; Mohlenbrock 1975; Swink and Wilhelm 1979.) Late August-October. Wet ground: tamarack bogs, sedge meadows, grassy lakeshores, and calcareous fens. Occasional in the counties including and surrounding Chicago, now probabh rare because of destruction of the habitat. The species range is still to be accurately mapped. I have been unable, for example, to locale the specimen cited for Winnebago County by Fell ( 1955: 144) under the name Astir iiiinif(>rniis: the descriptive statement suggests that the plant (found "in boggy places in Coon Mav 14X4 A^im AM) 8K\(in \i ii\ is Ii i imjis 151 Creek bottom") may belong here. According to the floristic literature, the species has been recorded from Anticosli Island. NB. and PQ. westward through ON. MB. SK. s. NT, and s. YT to BC and s. AK, southward to areas surrounding the Great Lakes in NY, PA, OH, and IN. and also including stations in n. lA. ND. SD. n. NE (?). and MT. Excluded are some areas listed in the literature for which the recorded specimens, in my judgment, belong in other species. The plants may be confused with those of some other species, e,g.. Aster dumosus var. strictior Torrey & Gray, but the latter taxon is distinct by having smaller heads and a strict, thyrsiform capitulescence. The characteristics of A. horealis suggest a close relationship in the eastern half of the continent to -A. laiueoUiliis and A. Ii»ii;if(ilins Lam., and in the western half to A. ealonii I A. Gray) Howell and A. ovcidcniaUs (Nutt.) Torrey & Gray. Of these four species only A. Uiiucdialus is an element of the Illinois flora, and good evidence exists of intergradation between that species and /I. horealis (e.g., Bcnkc 527H (F) from McHenry Co.|. 3. Aster cordifolius L. Heart-leaved Aster. Blue Wood Aster Herbaceous, with a branched-caudicifomi or creeping horizontal rhizome system (depend- ing on the variety). New shoots originating at the base of old stems or from rhizome strands, the plants fomiing scattered individual clumps or small colonial patches. .Stems 1-several, erect, (4{))6()-l2() cm tall, often reddish or maroon-colored, commonly somewhat zigzag above, with ascending or divaricate branches from above the middle, typically puberulent in decurrent lines from the upper nodes, in the leaf axils, and on the peduncles, glabrous in the lower stem portion (sometimes entirely glabrous). Leaves polymorphic, the basal and principal catiline ones petiolate, those higher up on the stem gradually reduced in size. changing to subsessile or sessile in the branches; leaves of new shoots and of the vernal rosettes with ovate-cordate to suborbiculate blades 1-8 cm (or more) long and 1-.^ cm (or more) wide, similar to the caulinc leaves in venation and vestiture. purplish below at least during the cool season, the apex acuminate or acute (sometimes obtuse or rounded), the margins coarsely and often irregularly crenale-serrate, the base cordate (rarely rounded), the petioles 1-3 times as long as the blades, ciliate, slender or often winged, dilated toward the sheathing base: blades of principal cauline leaves ovate in contour. 4— 1()( 1.^) cm long and 2—4(6) cm wide, with several pairs of arching and anastomosing secondary nerves, and a weakly expressed reticulum of tertiary veins with irregular areolae, the upper surface sparsely scabrous or glabrous, the lower thinly villous or hirtellous along the major nerves (rarely over the entire surface) or nearly glabrous, the apex acuminate and callus-pointed, the margins sharply serrate with acuminate callus-pointed teeth, the base deeply cordate or truncate to rounded, the petioles progressively shorter and more broadly winged upward on the stem; rameal leaves subsessile to sessile, ovate-lanceolate, nearly glabrous or minutely puberulent above, acuminate and callus-pointed, the margins entire or commonly somewhat toothed and ciliolate; leaves of the peduncles densely spaced, 1 ..'i-.3 mm long, the lower ones Oexible. the upper bracteifonn and intergrading with the phyllaries. Capitulescence a leafy, divaricately much-branched, usually round-topped panicle in the upper '/i-'/: portion of the stem, the head-bearing branchlels rarely racemifomi. Flowering heads 1-1. .S cm in diameter when the rays are fully extended, often crowded, usually not secund. on densely bracteate peduncles 0..3-2 cm long. Involucre narrowly campanulate (sometimes turbinate when pressed and dried), 3.5-5(6) mm high, the phyllaries imbricated in (3)4 or 5 series. Phyllaries strongly graduated, the outer ones lanceolate. ( 1 )l.5-2 mm long, the inner slender, linear-oblanceolatc, 3.5^.5(5) mm long, appressed when fresh, glabrous, the green areoles rhombic to obovate and well delimited in the apical '/i-V- portion, commonly red-tipped, the apex typically acute or obtuse (rarely acuminate) in the outer and median, acuminate to short-attenuate in the inner phyllaries. the 152 Illinois Nati.ral History Si rvly Bi llltin Vol. 34 Art. 2 margins erose-hyaline, irregularly L-iliolate or thinly villous toward the apex, the basal portion scarious, often somewhat rounded on the back and indurate. Receptacle alveolate with sharp teeth. Ray florets (8)10-l.'i. the corollas 7-10 mm long, typically purple (rarely pink), glabrous or nearly so. Disk florets (8)10-12, the corollas 4-^.5(5) mm long, funnelform. glabrous or with a few trichomes at the throat, the limb abruptly dilated, cream-colored or light yellow turning purple after anthesis, the lobe/limb fraction 0.2-0.25, the tube slightly shorter than the limb. Pappus simple, the bristles as long as or slightly shorter than the disk corolla, whitish or faintly rose-tinged, soft, slender, and attenuate. Achenes oblong-obovoid or often slightly falcate, somewhat compressed, 2-2.5 mm long and 0.6-0.8 mm across, dull purple or light brown, glabrous, with 4 or 5 ribs. 2>i = 16. .^2 (often with 1 or 2 B-chromosomes); reported chromosome counts of 2h = 36 are probably in error (A.G. Jones 1977). Including A. sai;itiifolius W^edem. ex Willd. {sensii stricio—A.G. Jones 1980b, 1987; Jones and Hiepko 1981). September-October. Loamy or rocky soils in mesic, mostly wooded habitats: open-wooded slopes and bluffs, woodland edges, also in somewhat disturbed ground at road cuts and along ditches. Occasional throughout the state but more common in the northern part. The range of this variable species extends from PE, PQ. NB, NS, and ME south to GA and AL, westward to MN. lA, e. NE, MO, and AR. Two varieties can be distinguished in Illinois, typical var. cordifoUiis and Aster contifoliiis var. sagitrifolius (Wedem. ex Willd.) A.G. Jones [A. safiitlifoliiis of authors only in part (see also A. dnimmoiulii and -4. iiiophyllus): A. cordifolins subsp. scif;itlifi>liiis (Wedem. ex Willd.) A.G. Jones (1980b)l; 1 . Rhizomes notably creeping, w ilh slender stoloniform strands, the plants colonial; leaves of the midstem with slender or narrowly winged petioles, the blades with a deeply cordate base var. cordifolius 1 . Rhizomes short, branched-caudiciform. the plants forming scattered individual clumps; leaves of the midstem w ilh broadly winged petioles, the blades with a truncate or shallowly cordate base var. sagittifolius Plants of var. safiiitifolius exhibit traits that may have been acquired via gene flow from other species, such as Aster ciUolalus and A. dnim- mimdii. If underground parts are missing. specimens of this taxon can be distinguished only with difficulty from those of typical var. cordifolius. There is evidence of intergradalion between A. coidifoliiis and A. drummondii |e.g.. Henry 2941.2943 (MWI) from Adams Co. and Chase 14906 (ILL) from Peoria Co.], A. laevis [e.g.. Chase 372S (ILL) from Tazewell Co.]. and A. loophylliis [e.g., Evers 5862H (ILLS) from La Salle Co. and Fidler & Fisher 353. 370 (ISM) from Union Co.). 3a. Aster ciliolatus Lindley in Hooker Lindley's .Aster I have included this species in the key on the strength of three herbarium specimens collected in Cook County and located at F [R. Behh 1570. Beverly Hills; F Gates 43. Glencoe: Raddin s.n.. n. Evanstonl. The plants more or less exhibit the traits attributed to Aster ciliolaliis: stems glabrous or nearly so; principal leaves glabrous or sparsely eiliate below along the major nerves: peduncles with no or few bracteiform rameal lea\ es: heads relatively large, the m\olucres 6-7 mm high; phyllaries lanceolate, long-aiienuaie, with lance- rhombic green areoles and indurate, scarious or somewhat discolored bases. In fact, the Gates collection was annotated b\ Shertt as .4. lindleyanus Torrey & Gray, a synonym of A. ciliolatus. Field study is needed to ascertain whether this transcontinental boreal species is really an established element of the Illinois flora. The plants mas be waifs, or lhe\ ma\ be part of a hybrid swami invohmg.A. cordifolius and .4. ciliolatus (or some other species, e.g., A. laevis). Mav \m9 AsnK \M) Rhuiiyutis in Illinois I S3 4. Aster drummondii L\nd\e\ in Hooker Drummond's Aster Herbaceous, u ith stout branched-caudieltomi rhizomes iliat often turn woody with age. New shouts originating at or near the base of old stems, the plants forming scattered individual clumps. Steins 1 -several, erect, stout. 60-120 cm tall, w ith ascending or divaricate branches mostly from above the middle, uniformly soft-hinellous in the branches, coarsely hirsute on the main stem, the indument often in lines, sometimes glabrescent in the lower portion. Leaves polymorphic, the basal and lower cauline ones petiolate. those higher up on the stem gradually or abruptly reduced in size; leaves of new shoots and vernal rosettes with ovate-oblong to suborbiculate blades l-6( 10) cm long and 1-3(3) cm wide, purplish below at least during the cool season, copiously soft-pubescent on both surfaces, the apex acute, obtuse, or rounded, the margins crenate-serrate; principal cauline leaves w ith broadly ovate to ovate-lanceolate blades 6-12(15) cm long and 3-5(6) cm wide, with 5-12 pairs of arching and anastomosing secondary nerves, thinly soft-pubescent to scabrous or rarely glabrous above, copiously hirtellous to hirsute below, the apex acuminate to attenuate and usually callus-pointed, the margins crenate- serrate to sharply (sometimes doubly) serrate, the base cordate or truncate, sometimes oblique; petioles (2)4-8 cm long, usually conspicuously winged and with ciliate margins; upper leaves of the main stem lanceolate, subsessile; rameal leaves oblong-lanceolate, sessile, typically pubescent, callus-pointed, shallowiy toothed to entire and ciliolate; leaves of the peduncles bracteiform. 2-5 mm long, intergrading with the phyllaries. Capitulescence a usually ample panicle, distinctly overtopping the leafy stem portion, the tenninal branchlets often raccmifonm. Flowering heads 1-1. 5( 1.8) cm in diameter when the rays are fully extended, on densely bracteate peduncles 0.2-4 cm long, or sometimes subsessile and secund. Involucre campanulate (or turbinate when pressed and dried). (3.5)4-6 mm high, the phyllaries imbricated in (3)4 or 5 series. Phyllaries strongly graduated, the outer ones lanceolate or subulate, 1.5-2 mm long, the inner linear-lanceolate, ca three times as long as the outer, appresscd or slightly spreading, typically glabrous but occasionally puberulent on the abaxial surface and thinly puberulent on the adaxial surface, the apical green areoles rhombic to oblong or oblanceolate, the apex acuminate or attenuate, often red-tipped, the margins irregu- larly ciliolate toward the apex, and with a scarious rim extending to the tapered portion of the tip. the basal 'A-'/: portion scarious except for the often green midrib. Receptacle shallowiy alveolate with sharp or blunt teeth. Ray florets (8)10-15 (or more), the corollas 7-12 mm long, purple or lavender (sometimes white), glabrous. Disk florets ( 10)13-15 (or more), the corollas 3.5-4.5(5) mm long, funnelform, glabrous, the limb abruptly dilated at the throat, cream-colored or light yellow turning deep purple after anthesis, the lobe/limb fraction O.lK-0.25. the tube slender, slightly shorter than the limb. Pappus simple, the bristles subequal, slightly shorter than the disk corolla, whitish or rose-tinged, soft, slender, and attenuate. Achenes oblong-obovoid, often slightly falcate, somewhat compressed. ( 1 .5)2-3 mm long and 0.6-1 mm across, dull purple or brown, glabrous or often sparsely puberulent in the top portion, with 4 or 5 ribs. 2/1 = 16. 32 (mostly); reported chromosome counts of In = 36 are probably in error ( A.G. Jones 1977). 14. sa^ittifolius Wedem. ex Willd. var. drummondii (Lindley in Hooker) Shinners — Femald 1950; Kihbe 1952; Dobbs 1963; .Swink 1974; Mohlenbrock 1975; Swink and Wilhelm 1979. A. undulaius misapplied, not of L.—Mead 1846; Higley and Raddin 1891; Pcpoon 1927; Kibbe 1952; Dobbs 1963. | August-October. Loamy or rocky soils in mesic. mostly wooded habitats: stream banks, open-wooded slopes, thickets and edges of swamps: also sometimes in disturbed ground along roads and ditches. Common throughout the state. The range of this variable species extends from s. MN east to c. OH. south to w. MS, n. LA. and e. TX, and west to lA, e. NE, e. KS. and e. OK. 154 Illinois Naii ral History Sirvey Bi lletin Vol. 34 An. 2 Plants in Illinois belong mostly in typical var. cinimmondii. including Aster drummondii var. liuidactis Benke. described from Cook County \Benke 4830 (F)| and characterized by rose-colored rays and "rufescent" leaves and stems. Individuals exhibiting phyllaries that are puberulent on the back may fall within the limits of A. finkii Rydb., a species (or more likely a hybrid) described from Iowa and recognized by Shinners ( 1941 ) as an endemic of the driftless area of s.w. WI. I have recorded collections of these plants from 16 counties throughout the state and did not detect geographic or ecological patterns or any consistent correlation of this trait w ith other characters. Occasional hybridization with and gene flow from A. shortii or A. undiila- lus (or both) may in part account for the occur- rence of puberulent phyllaries in A. drummondii. I find it impossible, therefore, to give taxonomic recognition to this morphological variant. There is al.so strong evidence of intergradaiion between A. drummondii and A. urophytlus [e.g.. Evers 2Snfi9 from Jo Daviess Co., 2H300 from Pope Co.. 7I9J7 from Adams Co.. and 77JS3 from Lee Co. (all at ILLS)]. 5. Aster dutnosus L. var. .?/n'ch'or Torrey & Gray Bushy Aster. Rice-button Aster Herbaceous, with a creeping stoloniform rhizome system. New shoots mostly initialed from the rhizomes or sometimes at the base of old stems, the plants forming scattered individual clumps or small colonial patches. .Stems l-few at any point of emergence. slender, erect. 30-70( 100) cm tall, terete, the branches commonly a.scending (this variety), sometimes divaricate, slightly ridged from decurreni leaf bases, sparsely puberulent in lines or nearly glabrous, the indumeni more copious on the peduncles and sometimes uniformly disiributeil. Leaves somewhat polymorphic, the basal anil larger ones mostly \\ ithercd and deciduous at flowering time; basal rosette leaves spatulate. 1-5 cm long, glabrous or minutely scabrous above, usually purple-tinged below. crenate-serrale. abruptly narrowed to a coarsely ciliate subpetiolar portion, dilated and sheathing at the base: principal cauline leaves linear to linear-oblanceolate. 3-10 cm long and 2-3(5) mm wide, with (usually) only the midrib strongly expressed, scabrous or glabresceni above, glabrous below . the apex acute w ith a sharp callus point, the margins usually inrolled. scabrous, shallowly serrulate or rarely entire, the base sessile and often somewhat decurrent but not clasping: rameal leaves numerous, bractei- form, linear-oblong, rather uniform in size at each branch level and with relatively few clusters of smaller leaves in the axils, a.scending or spreading, scabrous-margined, and callus- pointed, those of the peduncles densely spaced, 1-3 mm long, intergrading with the phyllaries. Capitulescence a narrow panicle w ith stiffly ascending racemiform branches above the middle, or sometimes more broadly and diffusely branched from the lower nodes. Flowering heads 0.8-1 .5 cm in diameter when the rays are fully extended, typically on slender, puberulent or glabrous peduncles 1-5 cm (or more) in length (rarely subsessile). Involucre 3-5 mm high, narrowly campanulale (or turbinate when pressed and dried), the phyllaries imbricated in 4 or 5 series. Phyllaries strongly graduated, appressed or slightly spreading, glabrous on the abaxial surface but often with a few trichomes on the adaxial surface, the green areoles well delimited, rhonibic-obovate to broadly oblanceolale, the apex obtuse or acute, the margins irregularly ciliolate, hyaline, and also with a scarious rim. the basal "/< portion scarious except for the often slightly keeled green or brow n midrib: outer phyllaries 1-1.5 mm long, oblong-lanceolate, those largest in surface area (3rd or 4lh series inward) more than twice as long and ca twice as wide as the outer ones, linear or somew hat expanded toward the apex: innemiosl phyllaries very slender. Receptacle shallowly alveolate with sharp teeth. Ray florets 15-25 (or more), the corollas 4-6(8) mm long, typically pink or lavender (sometimes white), glabrous. Disk florets 15-20 (or more), the corollas narrowly funnelfonn. 3.5—4.5 mm long, glabrous, the limb cream-colored or light yellow turning pink after anthesis, the lobe/limb fraction 0.25-0.35. the lube shorter than the limb. Pappus simple, the bristles approximately as long as the disk corolla, w hitish, soft, slender, and attenuate. .-Vchenes oblong-obovoid, plump or slightly compressed, 1 .5-2.5 mm long and 0.5-0.7 mm across, pink, or Mav mxy AsiFJi AM) BkU IIYM IIS IS II 1 IMIIS 155 straw-colored with pink streaks, strigillose and sometimes minutely puncticulate, with 3-5 prominent ribs, the sculpturing conspicuous even in immature ovaries. 2/i = 16. M. sometimes with additional B-chromosomes. Late August-October. Moist or wet ground: bogs and sandy or calcareous Hals. Not common in Illinois; apparently concentrated in the northeastern part of the state, south to Champaign County and west to Lee County. Collections recorded from Hancock and Jo Daviess counties may belong in or near Asia frui;ilis rather than .4, dumostis. Our records mark the northwestern limit for this extremely variable species, the range of which extends from ME to s. ON. Ml. and s. Wl. south to s. FL and LA. and southwest to MS. AR. the s.e. comer of OK. and e. TX. Good evidence suggests intergradation between this taxon and the often sympatric populations of AsU'ifrai;ili.\ [e.g.. AG. Jones 3M)3 (ILL) from Iroquois Co.| and/4, lameolatiis var. interior (Wieg.) Semple & Chmielewski [e.g.. A.G. Jones 3304 (ILL) from Iroquois Co. and EJ Hill 10(1. /.S'.SV (ILL) from Cook Co.|. In fact, all characteristics that, according to Torrey and Gray ( IH41 ) and Wiegand (1928), distinguish 4. diimosus var. strivlior from typical var. diimosus may be traceable to similar characteris- tics in one or both of the above-mentioned taxa. e.g.. the "strict" branching habit, the often racemiform and relatively short-peduncled exposition of heads, and the comparative paucity of indument. These species are closely related and share a basic chromosome number of .v = 8. 6. Aster ericoides L. While Prairie Aster. Heath Aster. Wreath Aster Herbaceous, with a strongly creeping (Illinois plants) or short ' caudiciform rhizome system, depending on the variety. New shoots initiated at the base of old stems or from stoloniform rhizome strands, the plants fomiing colonies or scatiere individual clumps. .Stems 1-few at any poiiil of emergence, erect, the branches ascending or often somewhat fastigiate and arching, .30-80(100) cm tall, uniformly and often harshly pubescent, the trichomes appressed- sirigose. ascending, or spreading to rellexed hirsute (depending on the variety), the lower stem portion sometimes glabrescent. Leaves numerous, firm, sessile, entire, variously pubescent on both surfaces, often with clusters of smaller leaves at the nodes and in the axils of the larger ones, commonly all except the rameal leaves withered and deciduous at flowering time; basal rosette leaves spatulate or oblanceolate, 2-6 cm long and 1 cm or less wide, triple- nerved, i.e.. with Usometimes 2) pair(s) of secondary nerves running from the base more or less parallel to the midrib, the surfaces sparsely pubescent or often glabrous, the apex rounded or obtuse and with a clear spinule. the base sheathing; principal cauline leaves linear to linear-lanceolate. ( 1 )3-6 cm long and 2-7 mm wide, with only the midrib prominent or triple- nerved as in the basal leaves, grayish green, the indument strigose or hirsute (depending on the variety), the apex acute or obtuse and spinulose, the base rounded or slightly clasping; rameal leaves similar in contour, color, and vestilure, abruptly reduced in size, those of the peduncles bracteiform. oblong-ovate, 1.5-5(6) mm long, usually squarrose or sometimes appressed to ascending, intergrading with the phyllaries. Capitulescence a somewhat fasligiately branched panicle or often pyramidal at the top. the head-bearing branchlets arching, racemiform. Flowering heads numerous, small, mostly less than I cm in diameter when the rays are fully extended, crowded and secund. subsessile arising from the leaf axils, or on densely bractcate peduncles 0.5-1(2) cm long. Involucre cylindric or narrowly campanulate. ca 3—4 mm high, the phyllaries imbricated in 3 or 4 series. Phyllaries strongly graduated, firm, squarrose, with a well- delimited oblanceolate or spatulate green areole in the apical portion, the basal 'A-'/; portion scarious and somewhat indurate; outer phyllaries oblanceolate to spatulate. 1.5-2 mm long, at least somewhat pubescent on both surfaces, often conspicuously scabrous-hirsute abaxially, the apex acute to obtuse and with a prominent spinule. the margins scabrous; inner phyllaries linear-lanceolate, more slender than the outer and median, 3—4 mm long, usually glabrous, acuminate or atteiuiate. Receptacle alveolate with sharp teeth. Ra> Horets 8 18(20). the corollas 4-6(7) mm long, while (rarely pink or 156 Illinois Nailrai History Sirvey Bllletin Vol. 34 An. 2 bluish), glabrous or with a few trichomes on the tube. Disk florets 8-12(15), the corollas narrowly funnelform. 2.5-4 mm long, glabrous or with a few trichomes at the throat, the limb gradually expanded, light yellow turning purple after anthesis. the lobe/limb fraction n..^-0..^5. the tube ca half as long as the limb. Pappus simple. the bristles about as long as the disk corolla, whitish, soft, slender, and attenuate. Achenes plump, oblong-obovoid. often somewhat falcate, 1 .2-2 mm long and 0.4-0.6 mm across, deep purple turning brown when weathered, sericeous or densely strigillose, and with 7-9 ribs. 2« = 10 (mostly), 20. Including /\ e.\i,i;iiiis Rydb. \A. mulliflorus Alton—Mead 1846; Brendel 1887; Higley and Raddin 1891; Pepoon 1927; Benke 1928.1 September-October. Unshaded habitats: prairie remnants, dunes, gravelly expo.sed slopes, and open woods: also in somewhat disturbed ground that still supports native prairie vegeta- tion, e.g.. roadsides and railroad rights-of-way, but usually not in severely weedy situations. Widely distributed throughout the state. The species range extends over much of the North American continent, in the East from s. ME to s. VA, in the Midwest from s. ON and s.e. MB to AR, and in the West from SK, AB, and e. BC through e. WA, n.e. OR, ID, and the Great Plains to NM, TX, e. AZ, and n. Mexico. Plants of Illinois belong in subsp. ehcoidvs (A.G. Jones 1978b, c), comprising var. eiicoides and vdT. prosiratiis (Kuntze) S.F. Blake [Aster exiguus—G.^. Jones 1945, 1950, 1963: Fell 1955; Jones and Fuller 1955; Mohlenbrock and Voigt 1959). Although these two varieties arc widely sympatric and plants with intermediate characteristics have been recorded, the identity and integrity of the taxa seem to be largely maintained as follows: 1 . Pubescence of stems, leaves, and phyllaries often relatively sparse and soft, appressed- strigose, or trichomes on the steins some- times ascending var. I'licoides 1 . Pubescence of stems, leaves, and phyllaries copiously and harshly hirsute, the trichomes on the stems spreading or somewhat retlcxed var. proslruliis In addition, two color forms originally described from Illinois by Benke (1928. 1932b) are occasionally obser\'ed: Asier ericoides f. caeruleus {BenV.e) S.F. Blake (plants with blue to purplish rays), and A. ericoides f. gramsii Benke (plants with "rose-red" rays). Also included here are Illinois collections cited as belonging m the hybrid A. eiicoides var. piosiratus x A. pilosus (Mohlenbrock 1986). The hybrid formula was originally published as A. exiguus x pilosus and attributed to HE. Ahles (Jones and Fuller 1955: 468). These plants are not hybrids but are somewhat atypical individuals of var. prostralus. probably the result of injury, e.g.. from mowing. Hybridization is well documented and rather common between ,4. ericoides and A. mnae- aiigliue. The plants are known under the binomial A. X amellnsliniis Nutt.. pro sp. (see discussion under 14a). 6a. Asterfalcatus Lindley in Hooker var. commutatus (Torrey & Gray) A.G. Jones Western Heath Aster [A. commutatus (Torrey & Gray) A. Gray—Femald 1950] A few specimens collected along a railroad track in Kane County [Sheiif 1798 (F. ILL)] seem to approach the characteristics of and perhaps belong in this taxon. which is of coinmon occurrence from the Dakotas and Nebraska westward, especially in the Great Plains. The plants exhibit a somewhat decumbent sprawling habit, heads that are not secund and are some- s' hat larger than is t\pical for.A.vfc;- ericoides. and an abundance of larger cauline leaves not usually present at tlowering time in plants of A. ericoides. 1 have included this western taxon in the key. but if the Illinois plants really belong here, rather than representing aberrant forms ot A. ericoides that resulted from some kind of injury or other environmental influence, they are almost certainly waifs. 1 do not share the view of Jones and Fuller ( 1955: 468). who suggested that collections of "Aster commutatus sensu Femald (1950) 1433. quoad pi. 111.—Non (Torr. & Gray) .-v. Grav ( 1 884)" are hybrids between A ericoides (as A. c.yiiiuus) and A. pilosus. Ma\ IW4 Asll.K \M) Bk U //) \( ll\ IS ll I INOIS 157 7. Asterfragilis Willd. Brittle Aster (my suggested vernacular name). Sma White Aster Herbaceous, with creeping (sometimes short) hori/ontal rhizomes. New shuols arising mostly from slender stoloniform rhizomes, rarely at the base ol old stems, the plants tbmiing colonial patches or scattered individual clumps. Stems slender, erect. 1-few at any point of emergence. 30-8()( 100) cm tall, commonly branched from below the middle, the branches divaricate or ascending, brittle, often slightly ridged from decurrent leaf bases, minutely puberulent in thin decurrent lines, the main stem usually glabrescent to glabrous. Leaves somewhat polymorphic, with clusters of smaller leaves in axils of the larger ones at most nodes, the principal leaves commonly withered and deciduous at flowering time; basal rosette leaves subpetii)late. spatulate or oblanceolate. 1—1 cm long and less than I cm wide, usually purple-tinged below, with several weakly expressed and anastomosing secondary nerves, glabrous, the apex acute or obtuse, the margins crenate. the subpetlolar portion sparsely ciliate, winged, and somewhat dilated toward the sheathing base: principal cauline leaves linear to linear-oblanceolate. 2-6 cm (or more) long and ( 1 )2-.^ mm wide, usually with only the midrib strongly expressed, both surfaces glabrous or the upper minutely puberulent. the apex acute to attenuate with a sharp callus point, the margins often inrolled. scabrous, entire or shallowly serrulate, the base sessile, usually slightly sheathing and decurrent; rameal leaves numerous, linear or oblong, ascending or spreading, notably unequal in sue. the smaller bracteifomi. those of the peduncles often aciculifonn. 1-2 mm long, glabrous, intergrading with the phyllaries. Capilulescence a diffuse panicle, the head- bearing branchlets typically racemiform and often arching. Flowering heads 0.6-1 cm in diameter when the rays are fully extended, offen crowded but commonly not sccund. subsessile. or on slender peduncles ().2-.1 cm (or more) in length. Involucre 2..S—4 mm high, narrowly campaiuilate or turbinate, the phyllaries imbri- cated in 4 or .S series. Phyllaries strongly graduated, appressed or the outer ones spreading, glabrous, the green areoles linear to linear- oblanceolale. usually extending over the entire length of the midrib, the apex acute, the margins sparsely ciliolate and with a scarious rim extending to near the apex; outer phyllaries slender, linear, 0.8-1 mm long; median and inner phyllaries linear-oblanceolate. the innemiost 2..'i-.3.5 mm long, those largest in surface area (3rd or 4th series inward) typically less than twice as wide as the outer ones. Receptacle alveolate with sharp teeth. Ray florets (12)16-20. the corollas .^—4(6) mm long, while (rarely pink), glabrous. Disk florets 16-20 (or more), the corollas narrowly funnelform, 2.5-3.5 mm long, glabrous, the limb abruptly dilated at the throat, cream-colored or light yellow turning pink after anthesis, the lobe/limb fraction 0.35-0.45. the lobes notably recurved, the tube slightl\ shorter than the limb. Pappus simple, the bristles approximately as long as the disk corolla, whitish, soft, slender, sometimes slightly flattened at the acute apex. Achenes oblong- obovoid. plump or slightly compressed. 1 .0- 1 .8 mm long and ca 0.5 mm across, gray, thinly strigillose. with 4 or 5 often obscure ribs. 2n = 16. 32. [A. vimiiu'Ks misapplied, not of Lam. (ef. Jones and Hiepko 1981; Jones and Lowry 1986), including A. viminciis var. siihdiimosiis Wieg. — Deam 1940; G.N, Jones 1950, 1963; Jones and Fuller 1955; Mohlenbrock and Voigt 1959; Gleason and Cronquisi 1963; .Steyennark 1963; Mohlenbrock 1975, 1986; .Swink and Wilhclm 1979; and others.] Late August-October. Moist or wet ground: bogs, meadows, lakeshores. streamsides, and open bottomlands. Occasional throughout much of the state. The range of this variable species extends from ME south to FL, and at its western limit from s. WI southward through MO and AR to LA and e. TX. Most if not all Illinois populations belong in Asierfitifiilis var. siihJiiniiisiis (Wieg.) AG. Jones, a taxon typified by a collection [Ki(lK»a\ 158 IlLINDIS NaTI RAl. HlSrdRY Si RVhY Bl Ll.ETIN Vol. 34 An. 2 6(S' (GH)) from Olney, Richland County (Wiegand 1928: 171; A.G.Jones 1984). These plants dit'ter from typical var.//u.(,'/7/.s mainly by having relatively long peduncles, i.e., by somewhat resembling plants of A. dumosus. Good evidence suggests occasional hybridization and gene flow between A.fragilis and several other taxa that share the basic chromosome number of .v = 8, such as A. ditnuisus. A. lateriflorus [e.g., A.G. Jones 3I0H (ILL) from Clark Co. and Jelinek 221 (ILL) from Pope Co.|. A. lameolatus var. interior (e.g.. Mead s.n. (ILL) from Hancock Co.|. and A. onuirionis (e.g., Sluldneck /5.SW (ILL) from Cass Co. and Neill 16357 aSM) from St. Clair Co. |. 8. Asterfurcatus Burgess in Britton & Brown Forked Aster Herbaceous, with a creeping, horizontal rhizome system producing fleshy, stoloniform rhizome strands. New shoots arising at the ba,se of old stems or from the rhizomes, the plants forming colonial stands or scattered individual clumps, the few smal rosettes produced later in the season typically vanishing shortly after the first killing frost. Stems 1 -several, erect, often somewhat zigzag in the upper portion, (30)50-120 cm tall, the branches typically confined to the capilulescence, ascending, more or less dichotomously forked, somewhat ridged from decurrent leaf bases, the indument uniformly distributed around the stem, hirtellous or puberulent on the upper stem and in the branches, the lower portion of the stem glabrescent. Leaves polymorphic, those of the basal rosettes and the main stem peliolate: leaves of new shoots and of the autumnal rosettes with oblong to ovate-lanceolate blades 4-10 cm long and 2-5 cm wide, rugose, with an often reddish midrib and several pairs of pinnate secondary nerves, minutely scabrous above and hirsute below, the apex acute or obtuse, the margins serrate, the base rounded or obliquely subcordate, the petioles short, winged, sheathing at the base; cauline leaves gradually reduced in size upward on the stem, those of the midstem with ovate to lance-ovate blades 10-12( 15) cm long and (3)6-8 cm wide, rugose, with (6)8-12 pairs of pinnate secondary nerves, and with a weakly expressed reticulum of tertiary veins with more or less isodiametric areolae, harshly scabrous on the upper surface, densely hirsute over the entire lower surface resulting in a grayish green color, the apex acuminate with a long mucro, the margins sharply serrate with conspicuously mucronate teeth, the base shallowly cordate or truncate to rounded; petioles slender, mostly shorter than the blades, glabrous or irregularly ciliate, the base slightly dilated, sheathing, and somewhat decurrent; rameal leaves relatively few, similar in contour but abruptly reduced in size, subpetiolate to sessile, the base slightly clasping or sheathing, the midrib strongly keeled and decurrent; leaves of the peduncles 0-3, often bracteiform and phyllarylike, depending on the position, sometimes closely subtending the heads. Capilulescence variable, broad-topped, compara- tively few-headed and cymiform in the natural habitat but often ample, much-branched, and compound corymbifomi in disturbed situations. Flowering heads 2.5-3.5 cm in diameter when the rays are fully extended, not crowded and not secund, the peduncles 0.5-3(5) cm long, often leatless or nearly so, densely hirtellous or villous. Involucre campanulate (rarely hemispherical when pressed and dried), 6-8( 10) mm high, the phyllaries imbricated in 5-7 series. Ph\ Maries appressed when fresh, strongly graduated, the outer ones ovate, 1.5-3 mm long, the innermost linear-lanceolate, 5.5-7.5 mm long, those of the outer 2 or 3 series iw ice as w ide as the inner, puberulent on both surfaces, the apex obtuse or rounded, the margins lanate, erose-hyaline, and also often (median and inner phyllaries) with a red rim. the chlorophyllous areoles poorly delimited, w ith fuzzy edges, broadly obovate to rhombic or oblong, in the apical 'A -7' portion or rarely extending to the base in the tomi of bands, the basal 7:-/. portion scarious. indurate, and keeled or rounded on the back. Receptacle shallow ly alveolate with blunt teeth. Ray florets (12)1 5-20, the corollas 12-18 mm long, w hite (sometimes aging to pink), glabrous. Disk florets 25-35 (or more), the corollas tunnelfomi, 6-7(8) mm long, glabrous, the limb partway slenderly tubular, abruptly dilated 1-1.5 mm above the point of insertion of the filaments, cream-colored Mav lysy Aster wo BRAcnrMris in Illinois 159 or lighl yellow luniing purple after anihesis. the fraction of lobe/expanded limb portion 0.4-1X5. the apparent tube (i.e.. including the tubular limb portion) distinctly longer than the expanded limb portion. Pappus slightly shorter than the disk corolla, whitish or somewhat tawny, the bristles in 2 series, those of the inner series firm, w ith a clavellately expanded apex, those of the outer somewhat shorter, more slender, and attenuate. Achenes fusiform, plump. (2.5)3-.''..'i(4) mm long and 1-1 .2 mm across, dull purple or brown, thinly puberulent in lines or glabrescent, with 8-10(12) densely spaced, often straw-colored ribs. 2« = 18. with 0-several B-chromosomes. [A. divaricanis misapplied, not of L.—Mead I846;Kibbe 1952.) (Late July) August-October. Low springy or marshy ground in woody areas, usually underlain by sandstone or limestone: lower (mostly north- facing) slopes and edges on the cutting (eroding) sides of streams and rivers. Rare and possibly threatened, apparently less common now than is indicated by the collection records, although occasional stands are extensive. The populations are restricted to the northern one-third of the state, extending south along the major rivers to Tazewell. Fulton, and Hancock counties. The species range includes scattered stations in s. ML w. IN, s.e. WL e. lA. and s.e. MO. Aster laevis L. Smooth Aster. Smooth Blue Aster Herbaceous, with branched caudiciform or tangled rhizomes that turn woody w ith age but also producing stoloniform rhizome strands. New shoots arising at the base of old stems or from rhizome branches, the plants forming scattered individual clumps or small patches. Stems l-several. erect. (30)50-120 cm tall, with stiff, typically ascending, and commonly somewhat ridged branches above the middle, often reddish in the lower portion, glaucous and essentially glabrous except for sparse hirsute indument on the peduncles and in the leaf axils. Leaves polymorphic, somewhat fleshy, glabrous and glaucous, the basal and larger cauline ones sometimes withered at flowering time: basal rosette leaves commonly with long, winged, and basally dilated petioles, the blades oblong or ovate to lanceolate, 3-10 cm (or more) long and ca 1-3 cm wide, the apex acute or obtuse to rounded, the margins subentire or crenate- serrate. the base rounded or truncate; principal cauline leaves subsessile or mostly sessile. extremely variable in size and shape, oblong- ovate or lanceolate to linear, sometimes abruptly constricted above the base. (4)8-15 cm long and ( 1 )2^.5 cm wide, with l-several pairs of ascending secondary nerves, and a weakly expressed reticulum of tertiary veins with more or less isodiametric areolae, the apex acute or obtuse, usually callus-pointed, the margins entire or shallowly crenate-serrate, and minutely scabrous, the base often strongly auriculate- clasping: rameal leaves similar in contour but abruptly reduced in size, with a clasping or sheathing and somewhat decurrent base; leaves of the peduncles usually densely spaced, bracteiform. lance-subulate. 3-6 mm long, with an often purple callus point, intergrading with the phyllaries. Capitulescence a narrow or often somewhat fiat-topped panicle in the upper 'A -7; of the stem, the branches commonly stiffiy ascending (rarely arching). Flowering heads 1.5-2.5(3) cm in diameter when the rays are fully extended, terminal on somewhat ridged and often sparsely puberulent peduncles varying from 0.2 to 6 cm (or more) in length. Involucre campanulate. 4.5-7(8) mm high, the phyllaries imbricated in 4-6 series. Phyllaries strongly graduated, firm, appressed when fresh, the outer ones subulate or lanceolate. 1.5-2 mm long, the median and inner ones three times as long, linear or often slightly expanded toward the apex, the surfaces glabrous, the apical green areoles well defined, rhombic to lance-rhombic, glaucous, commonly smaller in area than the scarious basal portion, the apex obtuse, acute, or acuminate, with a sharp and often red or purple callus point. the basal portion of outer and median phyllaries usually keeled and indurate. Receptacle alveolate with sharp and typically bristle-lipped teeth. Ray florets 15-25(.^0). the corollas (10)12-15 mm long, blue or purple (rarely white), glabrous. Disk florets 15-25 (or more), the corollas narrowly tunneltorm, (4.5)5-6 inm 160 Illinois Natlral History Slrvey Bllletin Vol. 34 An. 2 long, glabrous, the limb slightly dilated at the throat, yellow turning purple after anthesis, the lobe/limb fraction ca 0.2, the tube slightly shorter than the limb. Pappus simple, the bristles usually somewhat shorter than the disk corolla, tawny or rose-tinged, soft, slender, and attenuate. Achenes oblong-obovoid, plump or somewhat com- pressed, 2-3 mm long and 0.8-1..^ mm across, deep purple at maturity, or brown when weath- ered, glabrous, with 4 or ."i often irregularly spaced ribs. 2n = 48. August-October. Mesic, mostly open habitats: prairie remnants, meadows, lakeshores, stream banks, open woods, and exposed slopes; also in somewhat disturbed ground along roads and railroad rights-of-way. Fairly common throughout most of the state. The range of the species extends from PQ south to GA and AL, northwest to AB, BC, and s. YT. and in the Rocky Mts. and Great Plains southward through c. CO to NM and the Guadalupe Mts. in w. TX. Although .somewhat variable, Illinois plants are assigned to typical var. Icievis. There is evidence of occasional hybridization and intergradation with other species that share the basic chromosome number of v = 8, such as Asier Uinceolalus |e.g.. Benkv 52X0 (F)—cf. Benke 1932b], A. puniceus (e.g.. Fuller 13293a (F) from McHenry Co.], A. oolentan^iensis [e.g., Heitmaim & Heitmann 1027 (F) from Fulton Co.], and A. kitcriflorus ]e.g., Bcnkv 2927 (F) from Kane Co.], 10. Aster lanceolalus Willd Panicled Aster Herbaceous, with a strongly creeping horizontal rhizome system New shoots initialed mostly at the lips stoloniform rhizomes, the extensively clonal but the connecting strands short-l and often disintegrating af two seasons. Stems niosth single at any point of emergence, otlen sunil. erect. 4()-120( 150) cm tall, pubescent in thin decurreni lines from the upper nodes, usually glabrescent or glabrous below the middle, often branching from near the base or producing in midsummer secondary shoots composed of clusters of smaller leaves plus a few heads at most of the nodes and in the leaf axils. Leaves polymorphic, the basal and larger cauline ones often withered or deciduous at flowering time; basal rosette leaves petiolate or subpetiolate. the blades elliptic-oblanceolate or obovate to suborbiculate, 1-8 cm long and 0.5-2 cm wide, often purplish below, with several pairs of anastomosing secondary nerves, glabrous, the apex acute. obtuse, or rounded, the margins coarsely crenate. the petioles winged and with a few long marginal cilia, the base dilated and sheathing; principal cauline leaves sessile or subsessile, elliptic or oblanceolate to linear-lanceolate, (4)6-15 cm long and (0.5)1-2(3.5) cm wide, gradually reduced in size up the stem, with relatively weakly expressed secondary nerves and a reticulum of tertiary veins with oblong areolae. glabrous (rarely scabrellous above), the apex acute to attenuate and callus-pointed, the margins sharply serrate, the base cuneate and somewhat decurrent; rameal leaves similar in contour and texture but mostly entire, those of the peduncles relatively few. flexible. 2-6( 10) mm long, rarely somewhat intergrading with the phyllaries. Capitulescence an ample, leafy , diffuse or often narrow ly elongate panicle. Flowering heads 1-2(2.5) cm in diameter when the rays are fully extended, rarely secund, the peduncles 0.5-5 cm in length. Involucre campanulaie, 3.5-6(7) mm high, the phyllaries imbricated in 3-5(6) series. Phyllaries somewhat or often strongly graduated, comparativeh slender, appressed or slightly spreading, the outer ones 1 .5-3 mm long, the inner 3-6 mm; outer and median phyllaries linear-oblanceolate, glabrous on the abaxial surface, sparseU puberulent on the adaxial surface, the green areoles linear-oblanceolate, the apex acute to acuminate and callus-pointed, the margins somewhat ciliolate. erose-hyaline. and w ith a narrow scarious rim extending to the tip. the basal 7.-7; portion scarious and slighih dilated; innemiost phvllaries linear, very slender, acuminate to attenuate. Receptacle alveolate w iih sharp teeth. Rav florets 20-35 (or more), the corollas 7-IO( 12) mm long, white (rarely pink). Mav 14X4 AsirH AM) Bk \< in m i i\ w Ii i isois 161 glabrous. Disk florets (15)20-30 (or more), ihc corollas tunnellorm, 3-5(6) mm long, glabrous, the limb cream-colored turning purple after anthesis. dilated at the throat, with comparatively long recurved lobes, the lobe/limb fraction 0.35-0.45(0.5). the slender tube somewhat shorter than the limb. Pappus simple, the bristles as long as the disk corolla or often slightly longer, whitish, .soft, slender, and attenuate. Achenes oblong-obovoid, somewhat compressed, 1.5-2 mm long and 0.5-0.8 mm across, gray, thinly strigillose, 4- or 5-ribbed. In = 32, 64 (mostly), rarely 40 or 48. Including /A. sinii^lc.x Willd. and /I. m/cnw Wieg. (Semple 1^^79; Semple and Chmielewski 1987). [A. pcinicuhilus Lam. non Miller; A. rraJescaniii misapplied, not of L.| August-October. Low moist or swampy ground: woods, pastures, bottomlands of rivers and streams, ditch margins, and lakeshores. Very common. LInder the broad taxonomic concept adopted by Semple and Chmielewski (1987), the species has a transcontinental range that includes localities in nearly every state and province of North America. Three varieties can be distinguished in Illinois; 1 ) var. lanceolams [Aslcr paniciilaliis— Brendel 1887; Higley and Raddin 1891; Pepoon 1927; Deam 1940; G.N. Jones 1945. 1950; Fell 1955. A. simplex var. rumosissimus (Torrey & Gray)Cronq.—Femald 1950; Gleason 1952; Dobbs 1963; Gleason and Cronquist 1963; Steyenmark 1963], 2) var. simplex (Willd.) A.G. Jones \A. .swi/j/cv (sensu stricto)—Mead 1846; Femald 1950; Gleason 1952; Kibbe 1952; Jones and Fuller 1955; Mohlenbrock and Voigt 1959; G.N. Jones 1963; Gleason and Cronquist 1963; Swink 1974; Mohlenbrock 1975. 1986; Swink and Wilhelm 1979. A. panUiilulus var. simplex (Willd.) Burgess in Briiton & Brown—Wiegand 1933; Deam 1940. A. lanceolalus subsp. simplex (Willd.) A.G. Jones— 1984), and 3) var. Imcrlor (Wieg.) Semple & Chmielewski 1-4. Inidescanlii sensu auct. non L.—Mead 1846; Brendel 1887; Higley and Raddin 1891; Pepoon 1927; G.N. Jones 1950; Fell 1955. A. vimlneus sensu auct., pro parte, non Lam.—Higley and Raddin 1891; Kibbe 1952. A. interior—Dcim 1940; Ci.N. Jones 1945; Jones and Fuller 1955; Mohlenbrock and Voigt 1959. 4. simplex var. inierior (Wieg.) Cronq.—Femald 19,50; Gleason 1952; Gleason and Cronquist 1963; Swink 1974; Swink and Wilhelm 1979. A. x interior Wieg., pro sp. — A.G. Jones 1980a, b. A. laneeolaliis subsp. InleriiiriVJkg.) A.G. Jones— 1984|. Plants of these three varieties show a great deal of intergradation. For the most part, however, they may be distinguished as follows: 1. Flowering heads 1-1.5 cm in diameter when the rays arc fully extended; involucres 3^ mm high; disk corollas 3-3.5 mm long; branchlets often racemiform var. inierior 1 . Flowering heads 1 .8-2.5 cm in diameter when the rays are fully extended; involucres (4)4.5-7 mm high; disk corollas typically (4)4.5-5.5 mm long; branchlets usually not racemifomi. 2. Leaves of the midstem linear-lanceolate, 12 times (or more) longer than wide var. lanceolatiis 2. Leaves of the midstem elliptic- lanceolate to oblanceolate, mostly less than 1 1 times longer than wide var. simplex For a different interpretation see Semple and Chmielewski ( 1987). These authors do not regard Aster simplex as sufficiently distinct from typical A. himeolatiis to merit taxonomic recognition. Collections of var. simplex from the central and southern parts of the Midwest, however, consistently seem to differ in several ways from the more northem populations of var. huueolaliis. Aside from the key characters mentioned and the somewhat differential geographic distribution, plants of var. simplex occur more frequently in shady bottomland woods, whereas plants of both var. lanceoUiliis and var. interior are more common in the unshaded habitats of bogs, meadows, lakeshores, and ditch margins. Good evidence suggests that this extremely variable species occasionally hybridi/.es and intergrades with other species of the .v = 8 chromosome number assemblage, e.g.. Aster horealis, A.fragilis, A. laevis, A. lateriflorus, A. punieeiis. and others that are not elements of the Hora of Illinois. 162 Illinois Natural History Slrvey Bulletin Vol. 34 Art. 2 11. Aster laleriflorus (L.) Britton Starved Aster, Calico Aster, Side-flowered Aster, White Woodland Aster Herbaceous, with a short, somewhat tangled rhizome system. New shoots initiated at the base of old stems or from short ascending rhizome branches, the plants fomiing scattered individual clumps. Stems 1-several, .^(>-80(l 20) cm ta comparatively slender, brittle, with ascending or often divaricate branches from the lower nodes resulting in a sprawling or bushy habit, the indumenl villous or hirlellous, usually in decurrent lines from the leaf bases, sometimes uniformly distributed in the upper branches, the main stem often glabrescent or glabrous. Leaves polymorphic, the larger cauline ones at least in part persistent throughout (he flowering period; basal rosette leaves subpetiolate, the blades spatulate or obovate to suborbiculate, the upper surface minutely puberulent or glabrous, the lower surface villous along the midrib (rarely glabrous) and sometimes purplish, the margins crenate-serrate, abruptly narrowed to the winged, ciliate, .somewhat sheathing subpetiolar base; principal cauline leaves sessile or subsessile, mostly elliplic-oblanceolate, rarely linear, (3)5-10(15) cm long and (0.2)1-2(3.5) cm wide. gradually reduced in size up the stem, usually with several pairs of anastomosing secondary nerves, and a weakly expressed reticulum of tertiary veins with oblong areolae, the upper surface typically scabrellous, the lower surface villous or hirtellous along the midrib but otherwise usually glabrous, the apex acuminate to attenuate with a sharp callus point, the margins serrate and minutely scabrous, the base cuneate and somewhat decurrent; rameal leaves similar in contour, more or less glabrous, entire, those subtending the head 1-few, oblong-lanceolate, 1.5-3 mm long, flexible, not phyllarylike. CapUulescence an often ample diffuse panicle. commonly branched from below the middle, head-bearing branchlets slender and wiry. arching, racemifomi. Flowering heads sinal 0.8-1.3 cm in diameter when the rays arc ful extended, often crowded, secund, sessile or subsessile, the peduncles rarely more than I ihc in length. Involucre campanulate (turbinate when pressed and dried). 3.5^.5(5) mm high, the phyllaries imbricated in (3)4 or 5 series. Phyllaries strongly graduated, appressed or slightly spreading, glabrous on the abaxial surface, the apical green areoles obovate- oblanceolate, the basal '/-'/• portion scarious except for the green midrib; outer phyllaries linear-lanceolate, 1-2 mm long, less than half as long as the inner, acute and callus-pointed: median phyllaries linear or often slightly expanded toward the acute to acuminate apex, sparsely puberulent on the adaxial surface, the margins irregularly ciliolate. erose-hyaline, and with a narrow scarious (sometimes reddish) rim extending to the tip; innermost phyllaries very slender, linear. 3.5—1.5 mm long. Receptacle alveolate with sharp or bluntish teeth. Ray florets 10-15(20), the corollas (4)5-8 mm long, white. glabrous. Disk ttorets 10-15(20). the corollas funnelfomi, (2.5)3.5—4.5 mm long, glabrous or with a few trichomes, the limb abrupth dilated at the throat, cream-colored or light yellow before anthesis. soon turning deep magenta, the lobe/limb fraction 0..5-0. 75. the lobes strongly reflexed. the slender tube slightly shorter than the limb. Pappus simple, the bristles about as long as the disk corolla, white, soft, slender, and attenuate. Achenes oblong-obovoid. plump or slightly compressed. (1.3)1.8-2.2 mm long and 0.5-0.7 mm across, gray, sparsely strigillo.se. obscurely 3-5 ribbed. 2n = 16. 32 (mostly). 48. Including A. pendulus Alton. A. horizontalis Desf., and A. lursutkaulis Lindley in DC. \A. vimincus Lam. (see ,A.G. Jones 1984; 379; Jones and Lowry 1986). A. diffusus .Ailon — Brendel 1887; Higley and Raddin 1891. ,A. miser sensu Alton non L.—Mead 1846.] Late August-October. Wooded slopes, high banks of streams and rivers, and edges of bogs; Illinois plants more common in well-drained or upland situalions than in wel ground habitats. Common throughout the state. The range of the species extends in the East from NB. ME. PQ. and s. ON south to PL, and to the west from MN and e. \\ through e. KS to L.\ and s.e. TX. Varietal names can be assigned to collections from Illinois onl\ w iih the greatest difficulty because of a great deal of intergradalion. not only between the \ arieties that have been described (Wiegand 192S) but also in\ol\ing gene flow from other species with a basic chromo.some number of .v = 8, e.g.. Aster dunwsus. A.fragilis. May 1989 ASTEK AND BkmHYM IIS IV ILLINOIS 163 A. lanceokitus . and A. onlarunus. The majority of specimens more or less fit the circumscription and concept of typical var. Uilcrijhinis. Plants of var. horizonlalis (Desf. ) Farw. \A. taterifloius var. pendiilus sensu Wieg. non (Alton) Burgess in Britton & Brownj are not uncommon, especially in the southern half of the state. The characteris- tics that weakly distinguish this variety (very small heads combined with a sprawling habit and a tendency toward relatively moist habitats) may have been introduced in part through gene tlow from A.frafiilis. Illinois collections labelled A. laleriflorus var. hirsiilicaiili.s (Lindley) Porter [sensu Pepoon 1927] are partly referable loA. onlarionis and partly to typical var. Iciteriflorus. The range of var. hiisuticauUs (including var. tenuipes Wiegand ( 192H)| probably does not extend to Illinois; these plants are common in the northeastern states and in Canada west to s. ON. 12. Aster linariifalius L. Fla,\-leaved Aster. Savory-leaved Aster Tufted, often suffruti- cose. with a tangled or cormoid-caudiciform rhizome system that turns woody with age. New shoots arising at the base of old stems but also at the end of ascending, densely bracteate. rhizome strands; basal rosettes absent. Stems cespitose, several to many, slender and wiry. ( 10)2CJ-40(60) cm tal simple, or few-branched only in the capitules- cence (unless injured through grazing or mowing), cinereous, puberulent. sometimes glabrescent toward the base. Leaves numerous, firm, densely spaced, sessile, ascending or spreading to squarrose. more or less uniform in contour, linear or nearly so, entire, those of the main stem 1.5-2..'i(.3..'i) cm long and 1.."1-2.5(4) mm wide, with a keeled midrib, the secondary venation not evident, glabrous except for the scabrous margins and the midrib below, the epidermal cells of both surfaces strikingly bulliform (hand lens needed to see this), the apex acute with a sharp callus point; lowermost cauline leaves bracteiform; rameal leaves abruptly reduced in size, those of the peduncles intergrading with the phyllaries. Capitulescence racemifomi or corymbiform in the upper '/.-/: of the stem. Flowering heads relatively few per stem, commonly fewer than 30. 1 .5-}^ cm in diameter when the rays are fully extended, on leafy peduncles 0.5-.'>( 15) cm in length. Involucre turbinate or campanulate. (6)8-10( 12) mm high, the phyllaries imbricated in 5-8 series. Phyllaries strongly graduated, firm, appressed, often largely scarious on the abaxial surface, the green areoles not well defined, centered in the apical 'A-'/; portion or forming two bands along the whitish midrib, sometimes extending to the base; phyllaries of the outer 2 or 3 series 2.5-4 mm long, lanceolate, notably keeled, the apex acute with a red callus point, the margins ciliate to fimbriate, or lanate toward the apex, those of the 5th or 6th series inward largest in surface area, (6)7-8(10) mm long, linear-lanceolate, keeled or rounded on the back, often with a red rim. the apex obtuse or rounded, the margins conspicuously lanate over the entire length. Receptacle alveolate with sharp teeth. Ray norets 13-17, the corollas (10)12-15( 18) mm long, violet or reddish purple (rarely white), glabrous. Disk florets 20-30 (or more), the corollas narrowly funnelform. 5-6.5(8) mm long, glabrous, the limb gradually expanded, light yellow turning reddish after anihesis. the lobe/limb fraction ca 0.2. the tube slightly shoner than the limb. Pappus tawny, "double" (according to literature references) but often composed of 3 series of bristles, the principal bristles tlmi. in 1 or 2 series, somewhat unequal. 4—7 mm long, as long as the disk corolla or slightly shorter, with at least the innermost clavellately expanded and flattened toward the apex; bristles of the outermost whorl distinctly shorter. I mm long or less, and attenuate. Achenes obconic, plump, 2.5-3(3.5) mm long, ca 1 mm across, dull purple or brown. 3-5 ribbed, the color and sculpturing of the pericarp often obscured by densely sericeous indument. In = 18. \Diplo[Hippiis Unuiiifolius (L.) Hooker—Mead 1846.] Late August-October. Dry sandy, loamy, or rocky soils; exposed slopes and ridges, open woods, dunes, and barrens. Most prominent in the northern half of the state but extending smith to St. Clair County. The range of the species extends from s. PQ and ME south to FL, and to the west from MN through s. MO, s.c. KS, and e. OKloe.TX. 164 Illinois Natural History Sirvey Bllletin Vol. 34 An. 2 13. Aster macrophyllus L Large-leaved Aster. Big-leaved Aster Herbaceous, with a strongly creeping horizontal rhizome system. New shoots arising mostly at the ends of stolonitonn rhizome branches, the plants typically forming extensive colonies, the autumnal basal rosettes numerous, prominent, often forming a dense ground cover. Stems I -few at any point of emer- gence, erect, sometimes slightly zigzag in the upper portion, 30-70(90) cm tall, terete or .somewhat angular above from decurrent leaf bases, often maroon-colored, the branches confined to the capitulescence, a.scending, the upper stem and branches pubescent with white, multicellular, antrorsely curved or spreading trichomes and also invested with sessile or stipitate glands, the lower stem glabrescent or glabrous. Leaves polymorphic, those of the basal rosettes and the lower half of the stem petiolate; basal rosette leaves largest, the blades broadly ovate to cordate, (8)15-25 cm long and (6)8-12( 15) cm wide, with 6-12 pairs of pinnate secondary nerves, and a reticulum of tertiary veins with more or less isodiametric areolae, the upper surface minutely scabrous or glabrous, the lower thinly puberulent, especially along the nerves, the apex acuminate with a sharp mucro, the margins crenate-serrate, each tooth mucronate, the base deeply cordate and some- times slightly oblique, the petioles 1-2 times as long as the blades, with slightly winged and ciliate margins, dilated and sheathing at the base; cauline leaves gradually reduced in si/e up the stem, blades of the larger ones nearly as wide as long, similar to the basal leaves in contour and vestiture except for sparse glandularity along the midribs below, the petioles shorter and progres- sively more broadly winged upward on the stem; upper stem leaves subpcliolate or often sessile and clasping, ovate or obovale to oblanceolate; rameal leaves relatively few, abruptly reduced but variable in size, ovate or oval, the upper surface scabrous and stipitate-glandular, especially along the midrib and the margins, the lower surface more sparsely so. the apex acute or acuminate, the margins crenate-serrate to entire, the base rounded or cuneate. the 0-few leaves of the peduncles ovate-lanceolate. 2-5 mm long. thinly to densely glandular, attenuate, not intergrading with the phyllanes. Capitulescence a dichotomously branched corymbiform panicle in the upper /; - / portion of the stem, the branchlets short and firm. Flowering heads variable, ca 2-3 cm in diameter when the rays are fully extended, sometimes crowded in the individual cymules. the peduncles 0-2.5 cm long. often densely glandular-hirsute. Involucre campanulate or hemispherical. (6)8-10 mm high, the phyllaries imbricated in (4)5-7 series. Phyllaries appressed or somewhat spreading. strongly graduated, those of the outer 2(or 3) series ovate, 2-3 mm long and ca half as wide, acute to obtuse, the green areoles oblong- obovate. in the apical 7-7; portion (rarely extending to the base in the outer phyllaries). the basal portion mostly scarious. indurate, keeled, or rounded on the back, glandular-puberulent over the entire abaxial surface (outermost phyllaries) or only in the chlorophyllous zone and along the midrib (median ph\ Maries), the adaxial surface more or less glabrous, the margins lanate and glandular toward the apex, hyaline toward the base, and with a scarious or often reddish rim; phyllaries of the innermost series slenderly linear-lanceolate. (6)7-9 mm long, ca I mm w ide. acute, largely scarious or sometimes red- rimmed. Receptacle alveolate with sharp teeth. Ray florets 10-15, the corollas 10-13 mm long. varying from white to deep lavender, glabrous. Disk florets 20-30 (or more), the corollas funnelfomi. 7-8 mm long, glabrous, the limb partway slenderly tubular, more or less abruptly tlared ca \.5-2 mm abo\e the point of insertion of the filaments, cream-colored or light yellow aging to purple, the lobes retlexed. the fraction of lobe/expanded limb portion 0.6-0.7. the apparent tube (i.e.. including the tubular limb portion) much longer than the expanded limb portion. Pappus tawny, in 2 series of somewhat unequal bristles, those of the inner series about as long as the disk corolla, finn and with a clavellately expanded apex, those of the outer slightly shorter. mote slender, and attenuate. .Achenes slenderly fusiform or cylindric. 3..'>—4(4.5) mm long and l-l .2 mm across, dull purple or brown, glabrous or w ith a few scattered trichomes near the lop. with S-IO (hick, densely spaced (sometimes doublo-siranded). glossy, golden-brown ribs. 2n = 72. May 1989 AsifR AM) Br uin \( lis in Illinois 1 65 August-October. Moist or swampy ground {sometimes dry soil) in wooded areas. Local, rare in this state, recorded only from Cook and Lake counties, much more common to the north and east of Illinois. The range of this variable species extends in the East from NS, PE, NB, and PQ south to GA. in the Midwest from ON south to s.e. IN. and at the western limit from e. MB through e. MN to n.e. lA. 14. Aster novae-angliae L New England Aster Herbaceous, with a tangled or sometimes caudiciform rhizome system. New shoots arising at the base of old stems or from short, fleshy. horizontal rhizomes, the plants forming scattered individual clumps or small patches. Stems 1-several. 50-1.^0 cm tall, stout, with ' ascending or divaricate branches from above the middle, densely hispidulous-hirsute or pilose, stipitate-glandular in the upper portion, the lower portion less pubescent and usually brown in color. Leaves numerous, polymorphic, the basal and lower cauline ones often withered and deciduous at tlowering time: basal rosette leaves spatulate. ca 2-6 cm long and ()..'i-l .."S cm wide, with 1-3 pairs of anastomosing secondary nerves more or less aligned with the midrib or margins, sparsely hirsute on both surfaces, the margins ciliolate and occasionally with a few shallow teeth; principal cauline leaves sessile and entire, oblong or lanceolate, with several pairs of weakly expressed secondary nerves, and a reticulum of tertiary veins with more or less isodiametric areolae: leaves of the midstem ca ."S-IO cm long and 5-l.'>(2()) mm wide, thinly stngose above, hirsute or hispidulous below, and invested with both sessile and stipitatc glands (more copiously so in the upper leaves), the apex acute and mucronulate, the margins pustulate-scabrous, the base auriculalc-clasping: rameal leaves similar in contour and vestiture. gradually reduced in size, the 1-4 small leaves of the peduncles not phyllarylike. C'apilulescence a leafy, broad, round-topped or somewhat corymbiform panicle in the temiinal ( '/t")'/"-'/" portion of the stem. Flowering heads 2—4. .Si 5) cm in diameter when the rays are lully extended, often crowded but not secund. the peduncles 0.3—1 cm long, glandular and hispidulous. commonly somewhat dilated just below the head. Involucre broadly hemispherical. 5-1,"i mm high, the numerous phyllaries imbricated in 4—6 series. Phyllaries slender, linear-lanceolate, long-acuminate or attenuate, commonly strongly squarrose or reflexed, mostly subequal in length, (5)6-121 15) mm long or the outer ones slightly shorter (rarely longer) than the inner; outer phyllaries largely herbaceous, dark green and often purple-tinged, densely invested with both stipitale and sessile glands: median and inner phyllaries scarious in the basal 7.-7: portion. Receptacle shallow ly alveolate or merely pitted. Ray florets numer- ous, ca 50-100. the corollas 10-20(25) mm long, rose-colored to deep purple (rarely white), with a few trichomes near the throat. Disk 5-15 mm in diameter. Disk florets numerous. 50 or more, the corollas narrowly funnelfomi. (4)5-7 mm long, thinly puberulent in the throat region, the limb only slightly expanded, light yellow turning purple after anthesis at least in the lobes, the lobe/limb fraction 0.2-0.25. the tube ca half as long as the limb. Pappus simple, the bristles somewhat unequal and often slightly shorter than the disk corolla, tawny or rose-tinged, soft, slender, and attenuate. Achenes oblong to obconic. plump, I.S-2.5(3) mm long, ca 0.6-1 mm across, dull purple or brown, obscurely glandular and densely sericeous, with 7-9 ribs. 2;; = 10. 20, occasionally with supernumerary chromosomes. August-October. Unshaded mesic situations: prairie remnants, open woods, streamsides, fens, also in disturbed soils of pastures, old fields, roadsides, and railroad rights-of-way. Common throughout the state; also widely cultivated and often escaped, a fact that partly accounts for the various color forms observed. The range extends from s. PQ, s. ON, and ME south to AL. MS. and AR, west to s. MB, c. ND, c. NE, and e. KS, with scattered stations in WY. CO. OK. and NM. Including f. roseiis (Desf. ) Britlon |var. /v;,vc//.v (Desf.) DC—Higley and Raddin 1891: Pepoon 1927; Benke 1928). plants with rose- colored rays: and f. ficncsecnsis House ( Benke 1932a). a white-rayed form. A short-lived, large- headed, white-rayed individual found in Champaign County (AG. Jones 2604). to which 166 Illinois Natlral History Survey Bllletin Vol. 34 Art. 2 I have applied the latter name, possessed an extra set (genome) of chromosomes (A.G. Jones 1980b). When cross-pollinated from a typical individual, a few viable achenes were produced that grew into typical purple-rayed plants. Hybrids between plants of A.sler novae-angliae and A. ericnides \A. x amelhystinus] are occasionally found in Illinois (see 14aj. Collections of this hybrid have been recorded in habitats shared w ith the parental species from Champaign. Cook. De Witt. Du Page. Fulton. McHenry. Menard. Peoria. Piatt. Richland. Vermilion, and Winnebago counties. The specimens show considerable variability and are probably not all first generation hybrids. 14a. Aster x amethystinus Nutt.. pro sp. \A. ericoides x A. novae-an^liae] This hybrid is readily identifiable (Benke 1930) and has also been produced experimentally (Wetmore and Delisle 1939; A.G. Jones 1978c). The plants occur with sufficient frequency in Illinois to warrant inclusion in the key and a comparative diagnosis: Habit similar to that of Aster ninae-anfiliae. the plants usually with a tangled rhizome system and short stoloniform strands forming small patches. Stems erect, aging to a light brown color, densely hispidulous- hirsute but not glandular. Leaves numerous, sessile, at least some of the larger cauline ones persistent throughout the flowering period and with clusters of smaller leaves produced in the axils; principal cauline leaves linear or oblong to elliptic-oblanceolate, 4—6 cm long and 3-6 mm wide, copiously and harshly hirsute on both surfaces but not glandular, the apex acute or obtuse, mucronulate, the margins entire and scabrous, the base rounded or slightly clasping but not auriculaie; rameal leaves similar in contour and vestiture, those of the peduncles few , oblong-lanceolate, 3-5 mm long, not intergrading with the phyllaries. Capitulescence an ample leafy panicle with ascending branches and a pyramidal or round top. Flowering heads intermediate to the two parent species in most characteristics. 1 .3-2 cm in diameter when the rays are fully extended, often crowded, some- times secund. Involucre hemispherical, 4—6 mm high. Phyllaries strongly retlexed. somewhat graduated or often subequal, scabrous-puberulent but not glandular, mostly with a conspicuous, scarious basal portion. Ray florets 20-30 (or more), the corollas ."i-lO mm long, lavender. Disk 4-7(8) mm in diameter. Disk florets 20-30 (or more), the corollas 3-4 mm long. Pappus tawny or rose-tinged. Achenes fusifonn or obconic. 1.5-2 mm long, dull purple or brown, densely sericeous but not glandular, 7-9 ribbed. 2/i = 10. 15. Asler oblongifolius Nutt Aromatic Aster Herbaceous, with a tangled, often stoloni- form or sometimes cau- diciform rhizome system New shoots usually arising from rhizome strands ( rarely at the base of old stems ), the plants forming scattered sprawling clumps or small colonial stands. Stems 1-several, erect or de- cumbent, brittle, I5-70(I(X)) cm tall, with ascending or divaricate branches from below the middle, variably hispidulous-hirsute or hirtellous and also glandular, more copiously so on the branches, the lower stem usually lacking glands. Leaves numerous, at least some of the larger cauline ones persistent throughout the flowering period; basal rosette leaves oblanceolate or spatulate. 2-5(7) cm long and 0.5-1.5 cm wide, with 1-3 pairs of secondary nerves aligned with the midrib or the margins, hirsute, often also glandular (rarely glabrous or nearly so), the apex rounded and with a short mucro, the margins entire and coarsely ciliate (sometimes remoteK toothed); principal leaves sessile, oblong or linear-lanceolate, (1.5)3-10 cm long and 0.5-1.5(2) cm wide, usually with 1-3 pairs of weakly expressed and anastomosing secondan. nerves, and a reticulum of tertiary veins w ith more or less isodiametric areolae, the upper surface usually scabrous and often glandular, the lower copiously hirsute, the apex acute or obtuse, spinulose-mucronulate. the margms entire and scabrous-ciliolaie, the base rounded or slightly clasping but not auriculate. often with axillary clusters of smaller leaves; upper cauline and rameal leaves graduallv reduced in size, similar in contour, copioush invested with both sessile Mav 1989 ASIF.R AND Bum HIM IIS IN ILLINOIS 167 and stipitate glands; leaves of the peduncles bracteifonn. densely spaced, several to many, linear or oblong, .'-K nun long, spiniilose, usually spreading or squarrosc, rarely ascending, more or less intergrading with the phyllaries. Capitules- cence an often ample, leafy, diffuse panicle commonly branching from the lower nodes. Flowering heads 1 .5-3 cm in diameter when the rays are fully extended, usually not secund, on glandular-hispidulous peduncles 0.5-5 cm in length. Involucre campanulate (or hemispherical when pressed and dried), 5-7(8) mm high, imbricated in -4-6 series. Phyllaries not strongly graduated, sometimes subequal. 3.5-7(8) mm long, densely glandular on both surfaces and along the margins, those of the outer 2 or 3 .series with strongly squarrosc lips; outermost phyllaries sometimes largely herbaceous, oblanceolate, acute; median phyllaries green in the apical '/i-7' portion, linear-lanceolate, long-acuminale or attenuate, the margins erose-hyaline or ciliolate. the basal '/:-"/< portion scarious; innermost phyllaries very slender. Receptacle alveolate with sharp teeth. Ray florets (20)25-35. the corollas lavender to deep purple (rarely white), 10-15 mm long, glabrous or nearly so. Disk florets 30-40 (or more), the corollas narrowly funnelform to nearly tubular. 4.5-6 mm long, glabrous or nearly so. the limb weakly delimited, deep yellow before anthesis aging to reddish purple, the lobe/limb fraction 0.18-0.2. the tube ca half as long as the limb. Pappus simple, the bristles somewhat shorter than the disk corolla, tawny or commonly rose-tinged, soft, slender, and attenuate. Achenes fusiform or obovoid. often slightly falcate, 2-2.5 mm long. 0.5-0.8 mm across, dull purple at maturity or brown when weathered, sirigillose or sericeous, with 7-l()( 1 1 ) straw-colored ribs. 2n = 10. 20. Late August-October. Dry sandy, loamy, or rocky soils in mostly open habitats; dunes, hill prairies, open-wooded blulfs, and sandstone or calcareous cliffs. Locally abundant in uplands associated w ith larger rivers. The range of the species extends from PA south to AL. west to w. ND. s.c. CO. n.e. NM. and n.c. TX. Most if not all the plants in Illinois belong in typical var. ohloni^ifoims [including Asier ohliiiis^ifoliiis var. aniiiisunus Shinners (1941 ) — Gleason 1952; Cileason and Cronquist 1963; Steyermark 1963). Occasional plants seem to verge toward the southwestern var. iii;iclulus A. Gray \A. Kiinileini (sic) Fries ex A. Gray ( 1884), pro syn.j. which is distinct by being compara- tively low-stemmed with numerous firm, relatively small leaves and by having a notably fastigiate branching habit. 16. Aster ontarionisV^ieg Ontario Aster Herbaceous, with a creeping horizontal rhizome system. New shoots initiated at the tips of stolonifomi rhizomes, the plants fomiing colonies but the connecting strands short- lived and mostly disintegrating after one or two seasons. Stems single (rarely 2 or 3) at any point of emergence. 40-120 cm tall, erect, with ascend- ing or divaricate branches from near or below the middle and also in midsummer with short secondary shoots composed of smaller leaves plus a few heads arising from the leaf axils, the indument villous or hirsute, uniformly distrib- uted on the branches, often in decurrent lines in the middle portion of the stem, the lower portion glabrescent. Leaves polymorphic; vernal rosette leaves with spatulate to oblanceolate-obovate blades, sometimes purplish below, minutely puberulent or rarely glabrous on both surfaces, the apex acute to rounded, the margins crenate- serrate. abruptly narrowed to a winged, ciliate, somewhat sheathing subpetiolar base; principal cauline leaves usually persistent throughout the (lowering period, sessile or subsessile. oblanceo- late or elliptic-lanceolate. 2-8( 12) cm long and 0.5-3.5 cm wide, gradually reduced in size upward on the stem, with several pairs of anastomosing secondary nerves, and with a weakly expressed reticulum of tertiary veins w ith oblong areolae, scabrellous above, softly (often minutely) puberulent over the entire lower surface, and sometimes densely villous along the midrib below, the apex acute or acuminate to short-alleiuiale, usually callus-poinlcd, the margins creiiale-scrrate and ciliolate. the base cuneale and somewhat decurrent; ramcal leaves similar in contour and veslilurc but entire and notably unequal in si/e, those sublentling ihe 168 Illinois Natlral Historv Si rvev BLLLETrs Vol. 34 Art. 2 head 1-few. linear-lanceolate, 1-3 mm long, flexible, not phyllarylike. Capitulescence an often ample, diffuse or elongate panicle above the middle of the stem, the head-bearing branchlets ascending or spreading but not strongly arching and typically not racemitbnm. Flowering heads 0.8-1.5 cm in diameter when the rays are fully extended, often crowded but typically not secund, subsessile or on peduncles 0.2-1(2) cm in length. Involucre campanulate (turbinate when pressed and dried), 3^.5(5) mm high, the phyllaries imbricated in 3-5 series. Phyllaries appressed or somewhat spreading, strongly graduated, the outer ones 1-2 mm long, less than half as long as the inner, slenderly linear, the median ones somewhat expanded toward the tip, the green areoles linear-oblanceolate, often extending to the base, both surfaces slightly puberulent, the apex acute or acuminate and callus-pointed, the margins irregularly ciliolate, hyaline, and with a .scarious rim extending to the tip; innermost phyllaries glabrous, very slender and attenuate. Receptacle alveolate with sharp teeth. Ray florets 15-25, the corollas 4-8 mm long, white, glabrous or nearly so. Disk florets 12-20 (or more), the corollas funnelform, (2.5)3—4.5 mm long, glabrous, the limb strongly flared, cream- colored or light yellow before anthesis turning magenta or purple, the lobes long and retlexed, the lobe/limb fraction 0.5-0.6, the slender tube slightly shorter than the limb. Pappus simple, the bristles about as long as the disk corolUi whitish, soft, slender, and attenuate. Achenes oblong-obovoid, plump or slightly compressed, 1.2-1.8(2) mm long, 0.4—0.6 mm across, gray, strigillosc. and often puncticulate (caused by the pustulate trichome bases), with 3-5 ribs. 2ii = 32. \A. missdurieiisi.s Britton in Britton & Brown non (Nutt.) Kuntze, includmg var. llnrsoick's (A. Gray) Wieg.—Deam 1940.-4. lateriflonis var. h'nsiiucaiilis misapplied, not of (Lindley) Porter—Pepoon 1927, at least in part. A. paiito- irichus S.F. Blake (see Shuiners 1949)—C..N. Jones 1945.) September-October. Moist grouiul habitats: river bottoms, creek margins, bogs, and marshes, frequently in wooded areas. Common throughout the state. The species range extends m the Fast from s.w. PQ, s.w. ON, and W^' south to AL. and at the western limit from MN and n.e. SD through e. NE and s.e. KS to e. TX and n. LA. Some of the specimens examined suggest intergradation with other species that share the basic chromosome number of .v = 8, e.g.. Aster fragilis. A. lanceiiUilus. and .4 lateriflorus. A considerable number of herbarium specimens have been misidentified as A. lateriflorus. If rhizomes are lacking, the two species are sometimes not readily distinguishable. 17. Aster oolentangiensis Riddell Sky-blue Aster, Azure Aster Herbaceous, with short horizontal rhizomes or a branched-caudicifomi system. New shoots arising at the base of old stems or from rhizome branches the plants forming scattered individual clumps. Stems 1 -several, erect, (30)60-100 cm typically with stiff, ascending branches above the middle, nearly glabrous or with hispidulous lines in the leaf axils and along decurrent leaf bases, the head-bearing branchlets slightly ridged, often uniformly scabrous-puberulent. Leaves polymorphic, the basal and lower cauline ones petiolate. commonly persistent throughout the flowering period; blades of basal rosette and larger cauline lea\es ovate to ovate-lanceolate. 4— 15( 18) cm long and 1—4(6) cm wide, w ith 2-several pairs of anastomosing secondary nerves, and a reticulum of tertiary veins with more or less isodiametric areolae, the upper surface scabrous, the lower softly hirsute, the apex acute to acuminate, the margins subentire to shallov\ ly .serrate, the base rounded, cordate, or truncate, the petioles twice as long (or more) as the blades, slightly winged, ciliatc. dilated toward the sheathing base; leaves of the midstem subpetiolate or sessile, ovate-lanceolate (in Illinois plants), gradually or somewhat abruptly reduced, acute or attenuate, and callus- pointed, the margins entire and scabrous, the base cuneale and soniewhal decurrent; rameal leaves abruptly reduced in size, linear-lanceolate or subulate, appressed or closely ascending, those of the peduncles bracteifomi. densely spaced. 1.5-3(4) mm long, scabrous-marguied. inter- grading with the ph\llanes, Capitulescence lypicalK a naixow tinrsifomi panicle in the upper Mav 14X9 AsrcR AM) Bk\ciiy\< ii\ \\ Ii 1 rsois 169 'A -7; portion ot ihc stem, the branches ascend- ing, sometimes raeemilomi. Flowering heads 1.5-2 cm in diameter when the rays are tulK extended, sometimes secund. the peduncles variable in length to 6 cm, densely bracteate. Involucre cylindric to campanulate (often turbinate when pressed and dried). 5-1 mm high, the phyllaries imbricated in 4 or .5(6) series. Phyllaries strongly graduated, appressed or those of the outer series slightly spreading, glabrous, the outer ones 1 .5-2 mm long, less than half as long as the inner, linear-lanceolate to subulate. the median and inner ones linear or often slightly expanded toward the apex, the apical green areoles rhombic, usually shorter than the indurate, scarious basal portion of the phyllaries, the apex sharply acute, acuminate, or obtuse, and mucronulate, the margins erose-hyaline and .somewhat ciliolate. Receptacle alveolate with sharp teeth. Ray florets ( 12)13-20. the corollas 8-12( 14) mm long, commonly blue or violet- purple (rarely white), glabrous. Disk florets 15-25 (or more), the corollas narrowly fun- nelform, 4-4.5 mm long, glabrous, the limb light yellow turning deep purple after anthesis. the lobe/limb fraction 0.2-0.25. the tube considerably shorter than the limb. Pappus simple, the bristles about as long as the disk corolla, cream-colored or slightly rose-tinged, soft, slender, and attenuate. Achenes oblong-obovoid. somewhat compressed. 1 .8-2 mm long and less than 1 mm across, dull purple, or straw-colored with purple streaks, glabrous or nearly so, with 4 or 5 ribs. 2/1 = .^2; reported chromosome counts of 2ii = .^6 (under the name A. azureus) are probably in error (A. G. Jones 1977). [A. azureus Lindley in Hooker—most authors prior to this publication (see A.G.Jones 1983).! August-October. Dry sandy, loamy, or rocky soils in relatively open or unshaded situations: prairie remnants, open woods, bluffs, dunes, and barrens. Locally common throughout the state. The species range extends from s. ON and w. NY west to MN and e. .SD and south through the central slates to n.c. FL. LA. and n. TX. Plants in Illinois belong in typical var. ook-nkiiii^ii'nsis (cf. AG. Jones 1983). There is some suggestion of intergradation with and gene flow from other species that share the basic chromosome number of .» = 8, e.g., Asier Jruminoiulii and A Uievis. 18. Aster parviceps (Burgess in Britlon & Brown) Mack. & Bush ^_^_^ Small-headed Aster ^ s i i ^^ Short-lived herbaceous perennial with a caudiciform rhizome. New shoots originating at the base of old stems or often from root sprouts produced in autumn of the previous season, the plants forming small scattered individual clumps. Stems 1-few, slender, erect, 20-70(90) cm tall, with ascending branches in the upper '/i-'/; portion, variably pilose to hirsute, the indument unifomily distributed or sometimes in decurrent lines from the leaf bases, the lower stem glabrescent. Leaves polymorphic, the basal and larger ones commonly withered and deciduous at flowering time: basal rosette leaves subpetiolate. oblanceolate or spatulate, 1—I cm long and 3-7 mm wide, with weakly expressed anastomosing secondary nerves, glabrous or sparsely pubescent, and often purplish on the lower surface, the apex obtuse and bristle-tipped, the base sheathing; cauline leaves sessile, often with clusters of smaller leaves in the axils, those of the midstem linear- oblanceolate to lanceolate, 4-8 cm long and 2-5 mm wide, thinly pilose above, puberuleni to hirsute below or sometimes nearly glabrous, the apex acute to attenuate and usually armed with a hyaline bristle, the margins entire or shallowly serrate, ciliolate; rameal leaves copious, mostly subulate and spinulose, those of the peduncles bracteiforni, appressed or ascending, 2—1 mm long, scabrous-margined, intergrading with the phyllaries. C'apilulescence a narrow panicle, with a pyramidal or Hat top. in the upper Vi-V- portion of the stem, the branches ascending or sometimes arching, typically racemiform. Flowering heads small. 7-10 mm in diameter when the rays are fully extended, secund. the peduncles densely bracteate. 0.3-2(4) cm long. Involucre cylindric to narrow ly campanulate (turbinate when pressed and dried), 3—4.5 mm high, the phyllaries imbricated in 3-5 scries. Phyllaries graduated, appressed when fresh, glabrous, the apical green areoles lance-rhombic; outer phyllaries subulate. 1 .5-2 mm long, ca half 170 Illinois Natlral History Slrvfy Bllletin Vol. 34 An. 2 as long as the inner; median and inner phyllaries linear-lanceolale. attenuate or acuminate with marginally inroUed spinulose lips, the margins erose-hyaline and commonly slightly ciliolate, the basal '/:-/> portion scarious. Receptacle alveolate with sharp teeth. Ray florets 10-16(18), the corollas ca .'i mm long, white, glabrous. Disk Horets 8-l()( 15). the corollas narrowly funnelform. (2)2.5-3 mm long, glabrous, the limb light yellow turning purple after anthesis, the lobe/limb traction 0.25, the tube shorter than the limb. Pappus simple, the bristles about as long as the disk corolla, white, soft, slender, and attenuate. Achenes oblong, plump, O.S-l .5 mm long and 0.4-0.6 mm across, whitish or gray, puberulent, obscurely 2^ ribbed. 2n = 16, 32. [A. lenuifolius misapplied. not of L.—Mead 1846. A. pilosiis Willd. subsp. parviceps (Burgess in Britton & Brown) A.G. Jones (1984).] August-October. Sandy or loamy soils in mostly unshaded habitats: barrens, hill prairies. rarely in somewhat disturbed ground that still supports prairie vegetation. Not common: mostly occurring in the western half of the state but also recorded from Cook and Will counties. As delimited herein (i.e., not including A.srer depauperatus (Porter) Fern.], the species range extends from lA and IL south to MO. n. AR, and s.e. KS. with one station recorded from n.e. OK. Aster parviceps is very closely related to A. pilosiis and was originally described as a variety of that species (under the name A. ericoides parviceps Burgess in Britton & Brown). For the time being and pending further study, I have reversed my decision ( 1984) to combine the two species, mainly because their habitats and reproductive vigor are strikingly different. Furthermore, there seems to be a difference in the predominant ploidy levels. Based on v = 8 chromosomes, plants of A. parviceps are recorded as being mostly diploid or sometimes tetraploid (Semple and Chmielewski 1985). whereas those of A. pilosiis are generally hexaploid, at least in my experience. 19. Ax/er patens Alton Spreading Aster. Late Purple Aster Herbaceous, with a branched-caudiciform. often tangled or some- times cormoid rhizome system, but also producing stoloniform rhizome strands. New shoots initiated at the base of old stems or from the rhizomes, resulting in scattered 1-several stemmed individuals or small patches. Stems erect, brittle, often stout. 50-100(120) cm tall, with stiff divaricate or ascending branches above the middle, the indument uniformly distributed around the stem. variously scabrous-hirsute to cinereous-pube- rulent, or villous on the branches and peduncles. Leaves polymorphic, the basal and lower cauline ones withered and deciduous at flowering time: basal rosette leaves spatulate. rugose, with 1—4 pairs of arching and anastomosing secondary nerves, variously scabrous-hirsute, the apex acute to rounded, the margins entire or often w ith a few teeth, the narrowed subpetiolar base winged and sheathing: principal cauline leaves ovate- lanceolate to oblanceolate. rarely spatulate. often constricted above the strongly auriculaie-clasping base, 2-6( 10) cm long and 1-2(2.5) cm wide, grayish green, rugulose on both surt'aces. with usually pinnate and anastomosing secondary- nerves, and a reticulum of tertiary veins \s ith isodiametric areolae, variously pubescent, more densely so along the midrib below, the apex acute or obtuse and uith a short mucro, the margins entire or appearing minutely denticulate from the pulvinate bases of coarse scabrous cilia: rameal leaves abruptly reduced in size, those of the peduncles bracteifomi. 1-3(5) mm long, appressed or sometimes squarrose. intergrading with the phyllaries. Capitulescence an open, divaricately branched panicle. Flowering heads 2-3.5 cm in diameter when the rays are fully extended, not crowded and not secund. typically at the ends of stiffly ascending or spreading, densely bracteate peduncles 2-10(15) cm in length. Involucre campanulate to turbinate. 5-8( 10) mm high, the phyllaries imbricated in May 1989 AsifK AND Bk.U in UT/S IN ILLINOIS 171 5-8 series. Phyllaries strongly graduated, linear to ovate-lanceolate, appressed or often slightly recurved-spreading. strigillose or cinereous- puberulent on the abaxial surface and near the lip on Ihe adaxial surface, the apical portion also frequently invested with both sessile and short- slipilale glands: outer phyllaries 1.5-2.5 mm, the inner (4)6-7 mm long; median phyllaries acute or acuminate to obtuse, the chlorophyllous areoles grayish green, rhombic-oblanceolate, poorly delimited in the apical ('A)7<-7.' portion, mostly shorter than the scarious basal portion. Recep- tacle shallowly alveolate with sharp or rounded teeth. Ray florets (12)15-25 (or more), the corollas typically blue or deep purple (rarely white), 1()-15( 17) mm long, glabrous or with a few scattered trichomes. Disk florets 20-50. the corollas narrowly funnelfonn, 4.5-6 mm long, glabrous or nearly so, the limb light yellow turning purple after anthesis. the lobe/limb fraction 0.18-0.2. the tube shorter than the limb. Pappus simple, the bristles about as long as the disk corolla or slightly shorter, tawny, soft, and attenuate. Achenes oblong-obovoid or fusiform, plump, 2. 5-.^. 5 mm long and 0.8-1.2 mm across, dull purple or brown, sericeous or strigillo.se, with 7-10 ribs. 2n = 10, 20 (Illinois plants). Including A. pcilenussimus Lindley in Hooker (see R.L. Jones 1983). September-October. Dry sandy or rocky soils: edges of Ozarkian forest lands, open- wooded bluffs, sandstone and limestone ridges; also in disturbed soils of clearings and old fields. Occasional in the southern one-third of the state, north to Jersey, Montgomery, and Crawford counties. The range of the species extends from New England to n. FL. west to s. IL. s. and c. MO, s.e. KS, and the eastern half of OK and TX. Two sympatric varieties have been recorded from Illinois with approximately equal frequency, var. luilcns and var. pulotussimus (Lindley in Hooker) Torrey & Gray. The laxa intergrade freely but may be distinguished as follows: I . Involucre 8-10 mm high, broadly campanu- late or turbinate; median phyllaries ovate- lanceolate, 1.2-1.5 mm in widih. obtuse, appressed. copiously cinereous-sirigillose on the back, not conspicuously glandular, or toward the apex with minute sessile glands thai are obscured by the indument \M. paieniissimus I . Involucre usually less than 8 mm high, slenderly campanulate or turbinate: median phyllaries linear-lanceolate, mostly less than 1.2 mm in width, acute or acuminate, somewhat recurved-spreading, sparsely strigillose on the back mainly along the midrib, conspicuously glandular with both sessile and short-stipitate glands var. patens The var. pawnlissimiis was originally described ias Aster pcili'iilissinnis) from a Missouri collection and is, according to Steyer- mark ( 1963), the more common form of A. patens in the Ozarks. Plants of typical var. patens are more prevalent to the east. Occasional herbarium collections seem to verge morpho- logically toward the small-leaved and slender- stemmed southern var. f>iacUis Hooker. However, plants of the latter variety are largely diploid |2/i (= 2.V) = 10], and that ploidy level has not been recorded for Illinois collections (A.G. Jones 1980b; R.L. Jones 1983). Specimens forming the basis for records from the Chicago region (Higley and Raddin 1891; Pepoon 1927), including A. patens |var.l phli)i;ifiiliiis sensu Higley &Raddin and Pepoon non (Muhl. ex Willd.) Nees, do not belong in this species. 20. Aster pilosus VJMd. Hairy Aster. Frost-weed Aster Herbaceous, with a caudicifomi rhi/ome. New shoots arising at the base of old stems or from root sprouts produced in Ihc previous season, the plants rorniing scattered individual clumps. Stems I -several, stout, erect. 30-120 cm tall, with ascending or divaricate branches from near or below the middle, variously pilose lo hirsute, the indument in decurrcnt lines from the nodes or often uni- formly distributed, sometimes the stems glabrous or nearly so. Leaves polymorphic, the basal and larger cauline ones commonly withered and deciduous at flowering lime; rosette leaves 172 Illinois Natlral History Slrvey Bi lletin Vol. 34 An. 2 spatulate. the blade 1-6 cm long and ()..'i-l .5 cm wide, abruptly narrowed to a winged subpetiolar portion, with 1 or 2 pairs of secondary nerves arching toward the apex, glabrous, or the lower surface thinly pilose along the midrib and often purplish, the apex rounded or obtuse, the margins softly ciliate or scabrous and often shallowly serrate, the base sheathing; cauline leaves sessile or subsessile, often with clusters of smaller leaves in the axils, the principal ones clliptic- oblanceolate to linear-lanceolate, 4-12 cm long and 0.5-2.5 cm wide, gradually reduced in size upward on the stem, with a prominent midrib and 1-several pairs of anastomosing secondary nerves, glabrous to variously pilose or hirsute (more so along the nerves below), the apex attenuate and usually armed with a hyaline spinule, the margins serrate or often entire, softly ciliate, the base cuneate, somewhat sheathing and decurrent; rameal leaves copious, subulate, those of the peduncles bracteiform, often densely spaced, appressed to ascending or sometimes spreading, ,^-6( 10) mm long, ciliolate, more or less intergrading with the phyllaries. Capitules- cence an ample, leafy, usually diffuse panicle, the branches commonly racemiform. Flowering heads 1 .5-2(2.5) cm in diameter when the rays are fully extended, often crowded and secund, subsessile, or on densely bracteate peduncles 0.5-3(5) cm in length. Involucre campanulatc or broadly urceolate (i.e.. somewhat constricted below the slightly recurved phyllary tips). .3.5-5(6) mm high, the phyllaries imbricated in 4 or 5 series. Phyllaries commonly at least somewhat graduated (rarely subequal in length), appressed or slightly spreading, glabrous except for a few marginal cilia near the apex; outer phyllaries subulate, 2-.^(4) mm long, sometimes largely herbaceous, the median and inner ones linear-oblanceolate. 3-5(5.5) mm long, with broadly rhombic to lance-rhombic green areoles in the apical V^-V- portion, the scarious basal portion somewhat indurate and rounded on the back, the apex with sharply acute or marginally inrolled acuminate tips, spinulose, the margins erose-hyaline. Receptacle shallowly alveolate with shaqi teeth. Ray florets 15-30, the corollas 7-4( 10) mm long, white (rarely pink), glabrous. Disk florets 30-40 (or more), the corollas narrowly funnelform, (3)3.5-4.5 mm long, glabrous or nearly so, the limb cream-colored or light yellow turning lavender or reddish puiple after anthesis. the lohc/limb fraction 0.25-0.3, the tube much shorter than the limb. Pappus simple. the bristles about as long as the disk corolla, white, soft, slender, and attenuate. Achenes oblong, plump. 1-1 .5 mm long and 0.5-0.7 mm across, whitish or gray, minutely puberulent. and obscurely 2-4 ribbed. 2n = 32. 40 (rarely), 48 (mostly). Including A. prinftlei (A. Gray) Britten in Britton & Brown. [A. ericoides misapplied, not of L.—Brendel 1887; Higley and Raddin 1891; Pepoon 1927; Benke 1938; Kibbe 1952. in part. A. villosus Michaux non Thunb. A. ericoides var. villo.sus (Michaux) Torrey & Gray—Higley and Raddin 1891; Pepoon 1927; Benke 1928. A. polxphylltis Willd. non Moench—Higley and Raddin 1891; Pepoon 1927.] August-October. Disturbed ground in mostly unshaded habitats; roadsides, railroad rights-of- way, open woods, and pastures. Very common, weedy, recorded from every county. The range extends from s. PQ and s. ON south to G.A and n. FL and at the western limit from e. SD through e. NE, e. KS. and e. OK to LA. Several varieties of Aster pilosiis recognized in widely circulated tloristic manuals (e.g.. Deam 1940; Femald 1950; Gleason 1952; Gleason and Cronquist 1963; Steyermark 1963; and Cronquist 198(1) occur in Illinois; 1 ) var. pilosus. 2) var. plarvphxllus (Torrey & Gray) S.F. Blake [A. ericnulcs var. plai\ph\lh(.s Torres & Gray — Pepoon 1927|. 3) var. demotus S.F. Blake, and 4) var. priiifik'i (A. Gray) S.F. Blake \A. ericoides var. priiiiilei A. Gray (1884). A. pringlei—Jones and Fuller 1955; G.N. Jones 1963; Mohlenhrock 1986]. The type collection iorA. pilosKS {scnsii stricio. i.e., for var. pilosus) is from Illinois (Jones and Hiepko 1981): it was originally named bv Michaux and is also the t\pe for .4 villosus (Jones and Lowry 1986). The relati\ely broad- leaved, densely villous to pilose plants of var. picinphyllus (accepted by Deam 1940; Dobbs 1963; Stevemiark 1963) can be. in my opinion, produced under the influence of cenain environ- mental conditions. For the most part, they tu quite well in typical var. pilosus. a view also taken by others (A. Gray 1884; Gleason 1952: Gleason and Cronquist 1963; Cronquist 1980). Recent studies suggest that \ars. prinf:lci and dcnuniis perhaps should be combined under the tbimcr name (Semple and Chmielewski 1985; see also Mohlcnbrock 1975). My own obser%ations lend to support this concept (including here also A. pohphylhis). The variety is characterized as being nearly glabrous and having relatively Mav 19X4 Asifn am)Bki \i us is Illinois 173 narrow leaves: these plants are fairly common throughout the state. When taken in the narrow sense, however, var. piinfili'i is restricted in Illinois to Lake Count) (S\\ink 1974; Mohlen- brock and Ladd 197S; Swink and Wilhelm 1979). A great deal of intergradation between the above varieties can be found in herbarium specimens and observed in the field. 1 have not attempted, therefore, to elaborate on keys in the literature dealing with varieties of this variable species (see Deam 1940: Femald 1950: Gleason 1952: Gleason and Cronquist 1963: Steyermark 1963: andCronquisl 1980). 21. Aster praealtus Poir. Willow-leaved Aster, Willow Aster Herbaceous, with a strongly creeping rhizome system, the connecting strands often persistent for several seasons. New shoots mostly originating at the tips of fleshy stoloniform rhizome strands, the plants forming extensive colonies. Stems commonly single at any point of emergence, stout, erect, sometimes slightly reddish or glaucous. (40)60-150 cm tall, with ascending branches above the middle, variously hispidulous to hirsute, the indumeni in decurrent lines from the nodes or sometimes uniformly distributed around the stem, especially on the branchlets. the lower stem portion glabrescent. Lea\es polymorphic, the basal and lower cauline ones commonly withered and deciduous at llowering time: basal rosette leaves fleshy, spatulate. 4—7 cm long and 1-2.5 cm wide, deep green above and often purplish below , with several pairs of anastomosing secondary nerves, minutely scabrous above, glabrous below, the apex rounded, the margins entire or shallowly serrate, ciliate on the narrowed subpetiolar portion, the base dilated and sheathing; principal cauline leaves firm, sessile, elliptic to linear- lanceolate. 4-12(15) cm long and 0.3-1.5 cm wide, gradually reduced in size upward on the stem, rugulose, with indistinct secondary nerves but a conspicuous reticulum of brownish tertiary veins (notable on the lower surface), enclosing isodiametric areolae, the upper surface com- monly scabrous at least near the margins, sometimes strongly so. rarely glabrous, the lower surface mostly glabrous (in Illinois plants) or sometimes puberulent. the apex acute to attenuate w ith a sharp callus point, the margins frequenlls inrolled, entire or shallowly serrate, and scabrous, the base gradually tapered, often slightly rounded, decurrent: leaves of the upper stem and branches usually with clusters of smaller leaves (plus a few heads) in the axils, therefore numerous and notably unequal in size, oval to linear-lanceolate, acute or obtuse, callus- pointed, scabrous-margined: leaves of the peduncles relatively few. similar in contour, ascending or recurved-spreading, (2)-l-l() mm long, fiexible. not intergrading with the phyllar- ies but often closely subtending the head. Capitulescence a mostly ample, leafy, slenderly thyrsiform or broadly diffuse panicle in the terminal '/> -7; stem portion, the head-bearing branchlets rarely racemiform. Flowering heads 1.5-2(2.5) cm in diameter when the rays are fully extended, usually not secund. on peduncles 0.3-2 cm (or more) in length. Involucre campanulate. (4)5-7(8) mm high, the phyllaries imbricated in 4 or 5(6) .series. Phyllaries appressed or with slightly recurved tips, at least somewhat or often strongly graduated, the outer ones 2-3(3.5) mm. the inner (4)5-6.5 mm long: outer and median phyllaries oblanceolate. somewhat constricted above a slightly dilated base, the green areoles oblanceolate or lance- rhombic, or sometimes the outer phyllaries largely herbaceous, the abaxial surface glabrous. the adaxial surface sparsely puberulent, the apex acute to acuminate and often w ith a reddish callus point, the margins ciliolate. erose-hyaline, and with a narrow scarious rim extending nearly to the tip. the basal '/:-'/> portion scarious: innermost phyllaries very slender, linear, with linear-oblanceolate. light green areoles, the apex acuminate or attenuale. Receptacle alveolate with sharp teeth. Ray florets ( 15)2()-.3() (or more), the corollas 7-l()( 12) mm long, typically lavender to rose-purple (rarely white), glabrous. Disk florets 25-35 (or more), the corollas tubular or narrowly funnelfonti, 4-6.5 mm long, the limb cream-colored or light yellow turning purple after anthesis, the lobe/limb fraction 0. 1 S-0.2, the tube much shorter than the limb. Pappus simple, the bristles often slightly longer 174 Illivois Natiral History Slrvf.v Bi lletin Vol. 34 An. 2 than the disk corolla, whitish, soft, slender, and attenuate. Achenes oblong-obovoid. plump or slightly compressed. 1.5-2 mm long and 0.5-f).8 mm across, purple, or straw-colored with purple .streaks, thinly strigillose, with 4 or 5 ribs. 2n = 32. Including A. suhaspcr Lindley in Hooker (see Wiegand 1933). \A. salicifoHus Alton and sensu auct. non Lam.—Brendel 1X87; Higley and Raddin 1891; Pepoon 1927; G.N. Jones 1945, 1950; Fell 1955. A. c aniens misapplied, not of Nees—Mead 1846, and many herbarium sheets so labelled.] September-October. Low moist or swampy ground: woods, thickets, meadows, banks of streams and rivers, ditch margins, and lakeshores. Common throughout the state. The range of the species extends in the eastern half from ON, ML and PA to n. FL and in the western half from s. MB through ND, SD, NE, and OK to TX and n. Mexico. The plants are variable, but attempts to distinguish the varieties that have been described (Wiegand 1933) are not very successful. The reason is a high degree of morphological intergradation and an apparent lack of convincing geographic range separation. The majority of Illinois collections can be identified with typical var. praeallus [including var. angnslior Wieg., pro parte (and in the sense of some authors and collectors)—Deam 1940; Steyemiark 1963]. The var. suhaspcr (Lindley in Hooker) Wieg., originally described (as Asler suhasper) from the St. Louis area, has been recognized by several authors (Deam 1940; Steyermark 1963; Swink 1974; Swink and Wilhelm 1979). Plants of this variety are fairly common; they are more readily identifiable by their copiously scabrous leaves and stems than by the characters used to define the taxon in the keys of Deam (1940) and Steyermark (1963), namely cauline leaves that are relatively short and elliptic, and rameal leaves that are mostly oval and obtuse. Good evidence exists of occasional hybridization and intergrada- tion with ,4, lancealaius (e.g., I H. Chase IJ7 (F) from Stark Co,, Evers I09()W (ILLS) from Johnson Co., AG. Jones 4616 (ILL) from De Witt Co.) and with A. piinleeits \ ar. firmus (Nees) Torrey & Gray (e.g., Winreniiificr 16261 (ISM) from Cook Co., Wolf 170 (F) from Fulton Co., Wade & Wade 2.*,S'.? (F) from Ogle Co., and others]. 22. Aster prenanthoides Muhl. ex Willd. Crooked-stemmed Aster, Crooked Aster Herbaceous, with a creeping rhizome system. New shoots produced at the tips of stoloniform rhizome strands, the plants typically colonial. .Stems mostly single at any point of emergence, erect, (40)60-120 cm tall, the older ones often dark purple, branched near or commonly above the middle, the branches zigzag, somewhat angled in cross section from decurrent leaf bases, villous or hirsute in decurrent lines, the indument some- times uniformly distributed on the peduncles, the lower portion of the stem glabrescent or glabrous. Leaves polymorphic, those of the basal rosettes subpetiolate, the blades obovate to oblanceolate, 1.5-7 cm long and 1-2 cm wide, with several pairs of anastomosing secondary nerves, scabrellous above or glabrous on both surfaces, acute or obtuse, crenate to crenate-serrate. abruptly narrowed to a slender or slightly winged petiolelike portion, w ith long marginal cilia and an often reddish, dilated, sheathing base; principal cauline leaves mostly p)ersistent throughout the flowering period, subsessile or sessile, oblanceolate in outline but abruptly and strongly contracted near or below the middle. 8-16(20) cm long and 1.5-5.5 cm wide, gradually reduced in size upward on the stem, the blades with 6-10 (or more) pairs of ana.stomosing secondary nerves; nenes in the w ider. apical portion pinnate, those in the narrower, basal portion running parallel to the midrib, the reticulum of tertiary veins weakly expressed with irregular areolae, the upper surface scabrellous. the lower minutely strigillose or glabrous but commonly hispidulous or villous along the midrib, the apex long-acuminate with a callus point, the margins sharply serrate, w ith callus- pointed teeth on the wider blade portion but commonly entire on the narrowed ponion. the base dilated and stronglv auriculate-clasping. the midrib extended on the stem as a decurrent ridge: Mav lysy Aster anu Bumid m lis \\ Illinois 175 rameal leaves similar in most aspects but otieii only slightly contracted near the middle, glabrous or nearly so. the margins less sharply toothed or often entire, the base sessile and clasping; leaves of the peduncles relatively few. lanceolate. 3-12 mm long, not intergrading with the phyllaries. Capitulescence a broad, flat- or round-topped, dichotomously branched panicle in the upper 74-7; portion of the stem, the branches slender. sometimes purplish. Flowering heads ( 1 . 8)2-2. .'i cm in diameter when the rays are fully extended, not crowded and not secund. on slender peduncles I—I cm in length. Involucre campanu- late or hemispheric. 5-i) mm high, the phyllaries imbricated in 4 or 5(6) series. Phyllaries often only slightly graduated, flexible, with recurved or sometimes retlexed tips, the outer ones mostly lanceolate and slightly constricted near the middle. 2.5—4 mm long, the inner and median ones linear-oblanceolate. 4—5 mm long, the green areoles linear-oblanceolate. or sometimes the outer phyllaries largely herbaceous, the abaxial surface glabrous, the adaxial surface slightly puberulent. the apex acute and callus-pointed, the margins irregularly ciliolate. erose-hyaline. and with a narrow scarious rim extending nearly to the tip. the basal 7' -7; portion scarious. Recep- tacle alveolate with blunt teeth. Ray florets 18-25 (or more), the corollas 10-15 mm long, typically lavender or bluish (rarely white). glabrous. Disk florets 20-25 (or more), the corollas funnelform, (3.5)4—5 mm long, often thinly puberulent near the throat, the limb cream- colored or light yellow turning purple after anthesis at least in the lobes, the lobe/limb fraction 0.2-0.25. the tube slightly shorter than the liinb. Pappus simple, the bristles as long as or slightly shorter than the disk corolla, some- what discolored, sott. slender, and attenuate. Achenes cylindric-oblanceolate or obovoid. slightly compressed. 2-3(3.5) mm long and 0..5-0.8 mm across, dull purple or straw-colored, thinly strigillose. 4- or 5-ribbed. 2« = 32 (mostly), 48. Late August-October. Moist or swampy ground; woods, thickets, meadows, seeps, and stream banks. Occasional, recorded by me from only seven counties in Illinois, much more common to the east. The range of the species extends from s. ON. NY. and PA south to NC and TN and west to MN and lA. 23. Aster punkeus L. Purple-stemmed Aster Red-stemmed Aster, Swamp Aster Herbaceous, with a horizontal, short or often strongly creeping rhizome system, the con- necting strands persistent for several seasons. New shoots mostly originating at the tips of fleshy sloloniform rhizome strands, the plants sometimes forming extensive colonies. Stems mostly single at any point of emergence, stout, erect, commonly purplish or maroon-colored. 50-150( 180) cm tall, with ascending or divari- cate branches in the upper half, variously hirsute to hispidulous (or both) in decurrent lines from the nodes, or the indument nearly unifomily distributed around the stem, the basal portion sometimes coarsely hispid. Leaves polymorphic, the basal and lower cauline ones commonly withered and deciduous at flowering time, but the head-bearing branchlets often very leafy (MUX. firmils)\ basal rosette leaves spatulate or oblanceolate. 3-10 cm (or more) long and 0.3-2 cm (or more) wide, dark green and somewhat rugulose above, often purplish below, the venation and vestiture similar to that of the principal leaves, the apex acute to rounded, the margins remotely crenate-serrate to subenlire. the lower subpetiolar portion of the blade winged, dilated, and sheathing at the base; principal cauline leaves variable, finn, sessile, (6)10-15(20) cm long and ( 1 )2-5 cm wide, gradually reduced in size upward on the stem, with .several pairs of pinnate and anastomosing secondary nerves, and a reticulum of tertiary veins with isodiametric or oblong areolae, the upper surface scabrous or glabrous, often glossy (resulting from the presence of bulliform epidcmial cells), the lower surface glabrous or minutely strigillose and often with scabrous, hispidulous. or villous indument along the midrib, the apex acute, acuminate, or attenuate, with a callus point, the margins shallowly serrate to subentire. the base typically clasping, often strongly auricled in the lower leaves, sometimes merely rouiidcil ami decurrent in those higher up 176 Illinois Natural History Si rvey Bllletin Vol. 34 An. 2 on the stem; rameal leaves often numerous, elliptic-lanceolate to linear-lanceolate, gradually reduced, acute to attenuate, callus-pointed, the vestiture similar to that of the larger leaves, the margins subentire to entire and often inrolled, the midrib and larger nerves commonly extending downward on the stem as decurrent or almost winged ridges; leaves of the peduncles relatively few. 0..'i-2 cm long, flexible, somewhat con- stricted above a dilated base, often closely subtending the head and appearing to be a part ot the involucre, but rarely bracteiform. Capitulescence an often ample, leafy, broad, round- or flat-topped, more or less dichotomously branched panicle in the upper 'A-'/; stem portion, the branches ascending or divaricate. Flowering heads 1 .5-3..'i(4) cm in diameter when the rays are fully extended, typically not secund. subsessile and overtopped by the subtending rameal leaves, or often on densely hirsute to villous peduncles 0.2-3 cm (or more) in length. Involucre campanulate. (6)8-l2{ 15) mm high, the phyllaries imbricated in 4-6 series. Phyllar- les typically not or only somewhat graduated, 6-10( 15) mm long, flexible, leaflike, the outer ones often appreciably longer than the inner and largely herbaceous; phyllaries of the 2nd and }rd series inward lanceolate to linear-oblanccolate. often spreading or retlexed. somewhat constricted above a slightly dilated base, glabrous (rarely slightly scabrous) on the abaxial surface and thinly puberuleni on the adaxial surface, th- ;.pex attenuate or acute to obtuse (depending on the variety), and callus-pointed, the margins irregularly ciliolate or lanate, the scarious basal portion typically much smaller than the green portion and often indurate or slightly keeled; innermost phyllaries very slender, linear, with linear-oblanceolate. light green areoles in the apical '/ -'/. portion, the margins erose-hyaline and with a scarious rim. Receptacle alveolate with sharp teeth. Ray florets 20-;t) (or more). the corollas 12-18(20) mm long, lavender to deep bluish purple (rarely white), glabrous or slightly puberuleni on the throat and tube. Disk florets (20)30-50 (or more), the corollas narrowly funnelfonn, abruptly dilated at the throat, (4.5)5-6 mm long, glabrous or with a few trichomes, the limb cream-colored or yellow lumiiig pink or puiple after anlhesis. the lobe/limb fraction 0.2-0.3. the tube shorter than the limb. Pappus simple, the bristles about as long as the disk corolla, whitish or slightly discolored, soft, slender, and attenuate. Achenes oblong or oblanceolate in contour, often somewhat falcate, plump or slightly compressed, (2)2.5-3.5(4) mm long and 1 mm or less across. purple at maturity or brown when weathered. thinly puberulent or glabrescent, 4- or 5-ribbed. 2n = 16 (typically). 32. Including -4. /irmMs Nees and A. hicididus (A. Gray) Wieg. (see A.G. Jones 19X4, 1987). \A. novi-helgii misapplied, not of L.—Brendel 1887; Higley and Raddin 1891; Mohlenbrock 1975. \9H6.A. loni;ift>lius misapplied, not of Lam. — Pepoon 1927: Fell 1955; Jones and Fuller 1955. .4 parens misap- plied, not of Alton—Higley and Raddin 1 89 1; Pepoon 1927.) (Late August) September-October. Low moist or swampy ground: woods, thickets, meadows, banks of streams and rivers, ditch margins, and lakeshores. Common in the northern two-thirds of the state, extending south to St. Clair. Bond. Fayette, and Lawrence counties. The range of the species [sensii lau>) extends in the East from NF and LB south to f^. and in the West from SK and AB to ND, with a few scattered records from SD, e. NE, LA. MO. and from Van Zandt and Smith counties in n.e. TX. The plants are extremely variable, but two morphologically well-delimited varieties can be distinguished in Illinois; \ix.puimeus (including f. dcmissHs (Lindley) Fern.—Dobbs 1963) and vdiT.firmiis (Nees) Torrey & Gray [Aslerfirmus— Mohlenbrock 1986. A. piiniieiis var. Iiicidultis A. Gray—Higley and Raddin 1891: Pepoon 1927; Mohlenbrock 1 975..4. ///(/is 177 The ranges ot these two varieties in lllniois are overlapping, but plants of\ar.firmus seem to be more common than those of var. piiniccus. Attempts to separate the two ta\a at the level of species have been frustrated because of a high degree of intergradation and inconstancy in the character states. There is also evidence of intergradation between Aslci piiniccus and such other members of the \ = 8 chromosome number assemblage as A. kinceolalus [e.g., Behh sil IF 17355) from Winnebago Co. and Wade & WmIc 1744 (ISM) from Ogle Co.], /A. lalerifldriis [see Steyermark 1963), and A. pniealliis (e.g.. Smith 6A7 and 6liii\ belong mostly in A. piiniceiis vM.finmis. Although G.N. Jones (1950. 1963) in his keys treated A. lonfiifolius in accordance with Lamarck's type and circumscription, the species probably does not get into Illinois. The collec- tions cited by Pepoon ( 1927) ioi A. pcilens \Raddin (F) and Umhach (F)] belong in A. piiniceiis vht. firnuis. not in A. laevis as suggested by SwinkandWilhelm(1979). 24. Aster schreberi Nees Schreber"s Aster Herbaceous, with a strongly creepmg horizontal rhizome system. New .shoot.s arising mostly at the ends of stolonifonii rhizome branches, the plants forming extensive colonies; basal rosettes typically numerous, even in midseason far outnumbering the flowering stems and often fomiing a dense ground cover. Flowering stems single at any point of emergence, erect, 30-80 cm tall, terete or. at the upper nodes, somewhat angular in cross .section from decurrent nerves, the ascending or divaricate branches mostly confined to the capitulescence, the main stem sometimes reddish, glabrous, the branches sparsely scabrous or puberulent. sometimes in decurrent lines, but the indument usually unifomily distributed on the peduncles. Leaves polymorphic, those of the basal rosettes and the lower half of the stem petiolate; basal rosette leaves largest, the blades broadly ovate to cordate. (8)10-25 cm long and (6)8-16 cm wide, with 5-8 pairs of pinnate secondary nerves, and a weakly expressed reticulum of tertiary veins with mostly isodia- mctric (rarely oblong) areolae, the upper surface glabrous or thinly scabrous, the lower w ith a few irichomes along the nerves, the apex acuminate with a sharp mucro, the margins coarsely and often irregularly crenate-serrale. each tooth w ith a clear or green to purplish mucro. the base deeply cordate, the lobes sometimes overlapping or the sinuses often broad, the petioles 1-2 times as long as the blades, with slightly winged and glabrous or sparsely ciliate margins, somewhat dilated and sheathing at the base: cauline leaves gradually reduced in size upward on the stem, blades of the larger ones often nearly as wide as long, similar to the basal leaves in contour and indument except for the progressively shorter and gradually more broadly winged petioles; upper cauline leaves subpetiolate or sessile, elliptic or obovate. with a truncate or cuneate base; rameal leaves relatively few. abruptly reduced, variable in size, ovate or oval, subglabrous or the indument similar to that of the larger leaves, the apex acute or acuminate, the margins finely and sharply serrate and ciliate. the base rounded or cuneate; leaves of the peduncles 0-few, ovate-lanceolate, 2-3 mm long, fiexible, attenuate, entire, ciliate, not intergrading with the phyllaries. Capitulescence a terminal corymbi- form panicle, but often also with lateral flowering branches from the upper nodes of the main stem, the cymule-bearing branchlets stiffiy divaricate or dichotomous, the peduncles (if developed) usually ascending at a wide angle, 0.3-2 cm long, slender, typically not glandular (rarely with a few stipitate glands). Flowering heads variable. 2-2.5 cm in diameter when the rays are fully extended, sometimes crowded and sessile in the individual cymulcs. Involucre campanulate (turbinate when pressed), 5-7 mm high, the phyllaries imbricated in 4 or 5 series. Phyllaries strongly graduated, appressed when fresh (loosely spreading in fruiting heads and when pressed and dried), those of the outer 2 series oblong-ovate, 1-2(2.5) mm long, glabrous or slightly puberulent on the back, not glandular (rarely with a few minute glands at the tips), the green areolcs oblong to oblanccolate (rarely 178 lLLl\<)rS NaTI RAL HlSTORY SlRVF.V BlLLETIV Vol. 34 An. 2 emending to the base as a broad band), the margins usually densely lanate and with a scarious rim, the basal (7')'/' -7' portion of the phyllaries scarious. indurate, and keeled or rounded on the back; phyllaries of the inner 2 series linear-lanceolate, 4.3-6(7) mm long, glabrous, largely scarious or only the midrib green, the apex often reddish. Receptacle alveolate with sharp teeth. Ray florets 6-X( 12), the corollas (8)IO-12( I.S) mm long, white or cream-colored, glabrous. Disk florets 15-25 (or more), the corollas funnelform, (5)5.5-7 mm long, glabrous, or thinly puberulent on the slender, tubular portion of the limb, the limb strongly flared ca 1 .5 mm above the point of insertion of the filaments, cream-colored or light yellow aging to purple, the lobes reflexed, the fraction of lobe/expanded limb portion 0.4—0.5. the apparent tube (i.e., including the tubular limb portion) distinctly longer than the expanded limb portion. Pappus tawny, the bristles in 2 series, those of the inner series about as long as the disk corolla, firm, with a clavellately expanded apex, those of the outer somewhat shorter, more slender, and attenuate. Achenes slenderly fusiform, 3-5 mm long and 1-1.2 mm across, dull purple or light chocolate brown, glabrous or with a few scattered trichomes near the top. with (7)8-10 thick (sometimes double-stranded) straw-colored ribs. 2n = 54. Including A. cluisei G.N. Jones in Jones & Fuller ( 1955). Late July-September. Mesic but usually well-drained loamy or gravelly soil in wooded areas: plants in Illinois found mostly on north- facing slopes of ravines or along streams and rivers ca 30-60 m (or more) above the water level. Occasional in the northern one-third of the state, south to Tazewell County and west to Rock Island, Henry, and Knox counties. Illinois (and Wisconsin) records represent the western outpost of this species, disjunct by ca 450 km from the nearest populations to the east. The species range extends from ME west through s.e. ON to s.e. WI and south through PA. WV, and s.e. OH to VA, e. KY, e. TN, and n. AL. Axler schreheri has been included in the Illinois list of threatened species, but the plants are of rather common occurrence in the piedmont and mountain woods of the eastern and southeast- em states. I have so far been unsuccessful in a search for consistent characters whereby the midwestem populations |sometimes treated as Asler chdsci—G.N. Jones 1%3] can be distin- guished from those to the east, at least at the varietal level. The plants also share the same hexaploid chromosome number of 2/? (= 6.v) = 54. 25. Aster sericeus \eni. Silky Aster s May IWy AsllK AM) Bk U H) \t /A l\ II IISOIS 179 (4)5-6 mm long, often largely herbaceous, ovate, acute, mucronulate. the median ones 6-S( 10) mm long, ovate-lanceolate, acuminate or attenuate, green in the expanded apical 7.-7' portion, the abruptly narrowed basal portion scarious. indurate, and rounded on the back; innermost phyllaries very slender, attenuate, often reddish at the base. Receptacle strongly alveolate with sharp teeth. Ray florets ( 10)1.5-25. the corollas 12-15( IX) mm long, deep purple (rarely white), with a few trichomes near the throat. Disk florets 20-30 (or more), the corollas narrowly funnel- form, (5)6-7 mm long, thinly puberulent on the tube and throat, the limb bright yellow turning reddish purple after anthesis. the lobe/limb fraction 0.18-0.2. the tube much shorter than the limb. Pappus simple, the bristles about as long as the disk corolla, discolored or tawny, relatively firm, attenuate. Achenes fusiform, plumpish. 2-.'< mm long and 0.7-1 mm across, purple at maturity or brown when weathered, glabrous, prominently 7-10 ribbed. 2n = 10. Late August-October. Dry sandy, loamy, or rocky soils in unshaded situations: sand barrens, dunes, hill prairies, and open-wooded bluffs. Local, mostly in the northern half of the stale, but extending southward along the Mississippi River to Randolph County. The range of the species extends from s. ON and Ml to TN. and in the West through the eastern half of the Great Plains from s.e. MB toTX. Illinois plants belong in var. seiiceiis. 26. Aster shortii L\r\d\ey in Hooker Short's Aster Herbaceous, with branched-caudiciform or short horizontal rhi/omcs thai turn woody with age. New shoots initiated at or near the base of old stems resulting in scattered indiv clumps. .Stems 1 -several, erect (40)80-120 cm tall, much branched and bushy, densely and unifomily sofl-hirtellous on the upper stem and in the branches, the indument in decurrenl lines farther down, or the stem glabrescent in the lower portion. Leaves polymorphic, the basal and principal caulinc ones petiolatc; leaves of new shoots and of the vernal rosettes with the blades mostly ovate, 1-6 cm long and 1-.1.5 cm wide, dark green above, purplish below at least during the cool season, with I or 2 pairs of anastomos- ing secondary nerves curving forward from the ba.se and additional pinnate nerves emanating from the midrib, the apex acute or obtuse, the margins crenate. the base cordate or rarely truncate; petioles twice as long as the blades, slender, not at all or only narrowly winged, often densely pilose or hirsute: principal cauline leaves largely persistent throughout the flowering period, regularly spaced, the internodes 2-.^(4) cm long, the blades ovate to lanceolate, some- times falcate, 5-10( 15) cm long and 2-7 cm wide, often conspicuously rugulose. with several pairs of pinnate and anastomosing secondary nerves, and a reticulum of tertiary veins with more or less isodiametric areolae, the upper surface glabrous, or slightly scabrous along the midrib, the lower surface copiously hirtellous or hirsute, the apex acute to attenuate or acuminate and with a sharp callus point, the margins mostly entire and .scabrous, or shallowly crenate-serrate in the lower leaves, the base cordate, truncate, or rounded, sometimes oblique, the petioles half as long as the blades or less, slender, not sheathing or clasping; rameal leaves more or less abruptly reduced in size, lanceolate to ovate, entire, sessile, or sometimes with short petioles, densely hirtellous. acute or obtuse with a sharp, often purple callus point; leaves of the peduncles minute, bracteiform, 1-1.5 mm long, inter- grading with the phyllaries. Capitulescence an ample, diffuse, broad-topped panicle, the head-bearing branchlets often arching. Flowering heads 1 .5-2.5 cm in diameter when the rays are fully extended, usually not crowded and not secund. the peduncles densely bracteate, 0.2-.^(5) cm long, hirtellous. Involucre campanulate (or hemispherical when pressed and dried). 4.5-6 mm high, the phyllaries imbricated in 5 or 6 series. Phyllaries appressed when fresh, strongly graduated, the outer triangular or lanceolate. 1-1 .5 mm long, the inner linear-lanceolate, 4-5.5 mm long, puberulent on both surfaces, the green areoles oblanceolate to rhombic in the apical '/--'/A'h) portion, usually much shorter than the scarious basal portion (rarely the outer phyllaries with a green band to the base), the apex acute or 180 iLLrSOlS NaTI RAL HlSTORY SlRVEY BULLETIN Vol. 34 Art. 2 acuminate, the margins erose-hyaline and irregularly ciliolate. Receptacle alveolate with sharp teeth. Ray florets (lSllX-23. the corollas 10-15 mm long, typically deep blue or purple, glabrous. Disk florets 20-25 (or more), the corollas funnelform, 5-6 mm long, glabrous or nearly so. the limb abruptly dilated at the throat, light yellow turning reddish purple after anthesis, the lobe/limb fraction 0.18-0.2, the tube shorter than the limb. Pappus simple, the bristles about as long as the disk corolla, tawny or rose-tinged, soft, slender, and attenuate. Achenes oblong- obovoid, somewhat compressed, 2.5-3 mm long and ca 1 mm across, dull purple at maturity or brown when weathered, glabrous, with 4 or 5 ribs. 2n = 16; reported chromosome counts of 2/i = 18 are probably in error (A.G. Jones 1977). September-October. Well-drained soils; edges of upland woods, thickets, wooded river banks, and open-wooded slopes. Common nearly throughout the state.The species range extends from s. ON, w. PA, and n.w. MD, south to GA, n. fT., and AL. west to e. MN and e. lA with scattered records from AR (but apparently none from MO). Plants in Illinois belong in var. shorlii. including f. gronemanii Benke, described as a plant with rose-red rays from a collection made near Elgin, Kane County [Bcnke 4S72 (F)]. There is evidence of occasional hybridization and intergradation with Aster anomalus and A. urophylhis (e.g.. Fell 53-1005 (ILL) from De Kalb Co. and Winleninger 14794 (ISM) from Jersey Co.], and with other related species. 27. Aster tataricus L.f. Tartarian Aster Herbaceous, with a branched comioid- caudiciform rhizome system but also producing short, fleshy, stolonifomi rhizome strands. New shoots arising from nodes and tips of rhizomes and also from many winter buds just below the ba.se of old stems, the plants aggressiveU spreading by vegetative reproduction. Stems 1-several, stout, erect, 80-150 cm tall, somewhat angled and ridged from dccurrent leaf bases, branched only in the capitulescence, uniformly hirtellous or scabrous with antrorsely hooked trichomes. Leaves polymorphic, the basal ones most prominent tomiing convoluted clusters rather than rosettes, very coarse, 3-4 times as large as the largest cauline leaves, the blades oblanceolale, 10-30 cm long and 5-10 cm wide, strongly rugose, with 6-12 pairs of pinnate and anastomosing secondary nerves, the upper surface minutely scabrous, the lower surface densely scabrous or puberulent especially along the nerves, the apex acute, obtuse, or rounded, and mucronate, the margins crisp, crenate-serrate, each tooth with a conspicuous mucro, the base of blades gradually tapered, the petioles winged. 10-40 cm long, as long as the blades or longer, sheathing at the base; cauline leaves gradually reduced upward on the stem. (8)10-15(18) cm long and 2-5 cm wide, relatively densely spaced with intemodes 2—1 cm long, the lower leaves oblanceolate and subpetiolate, the upf)er lanceolate and sessile, similar to the basal leaves in surface, venation, and indument, the apex acute or acuminate and mucronulate, the margins serrate or entire, densely ciliolate. the base cuneate, sheathing, and decurrent; rameal leaves abruptly much reduced in size, lanceolate. 0.5-1 cm long, puberulent, acute or attenuate and mucronulate, those of the peduncles few. bracteiform but tlexible, not phyllarylike. Capitulescence an elongate or broad-topped panicle in the upper 'A -'A of the stem, the terminal branchlets short with few-headed corvmbiform clusters. Flowering heads 2-2.5(3) cm in diameter when the ra\s are fully extended, sessile in the axils of subtending leaves or on puberulent and somewhat ridged peduncles 0.5-1.5(2.5) cm long. Involucre campanulate or hemispherical. 6.5-8( 10) mm high, the phyllaries imbricated in 5 or 6 series. Phyllaries appressed or somewhat spreading, graduated, those of the outer series triangular or lanceolate, 3—4 mm long, those of the inner series linear-lanceolate, 6-8 mm long, glabrous or minutely puberulent. with green bands along the midrib to the base, or the outer phyllaries sometimes largely herba- ceous, the apex acute in those of the outer 2 series, attenuate in those farther mward. the margins entire or erose-hyaline and w ith a red rim. Receptacle alveolate with sharp teeth. Ray florets 15-18(20). the corollas l(V15(18)mm long, lavender, glabrous. Disk florets 20-30 (or more), the corollas funncltomi, (4.5)5-6 mm long, glabrous or with a few trichomes at the throat, the limb abruptK dilated, light yellow luminu la\ender after anlhcsLs at least in the May mX9 Asirii AM) Bh u in \< lis i\ Ii i isdis 1X1 lobes, the lohc/limb traction 0.2. the lobes notably rceurved. the slender tube slightly shorter than the limb. Style branches soniev\hat atypical tor the genus in having acutish rather than attenuate sterile appendages. Pappus simple, the bristles shorter than the disk corolla, white or cream-colored, soft. slender, and attenuate. Achenes obconic-obovoid. slightly compressed or plump. \.5-2 mm long and ca 1 mm across, light brown, thinly strigillose, with 4 or 5{(^) ribs. 2/i = .'i4. October. Disturbed ground: roadsides, thickets, and waste places. Occasionally escaped from cultiyation. Introduced from n.e. Asia. Recorded by me from Champaign. Montgomery. Sangamon. St. Clair, and Vermilion counties. 2&. Aster turbinellus Lindlev in Hooker Herbaceous, with stout, branched, caudiciform rhizomes that turn woody with age. New shoots initiated at or near the base of old stems resulting in scattered individual clumps. Stems 1 -several, 5()-10() cm tall, with many ascending or divaricate branches from near or below the middle, glabrous or hirtellous in thin decurrent lines. Leaves polymorphic, notably variable in size, the lower ones soon withered and decidu- ous; basal rosette and lower cauline leaves subpetiolale, the blades oblong-oblanceolate. glabrous, the apex acute, obtuse, or rounded, the margins shallowly crenate and ciliolate, the subpetiolar portion coarsely ciliate, winged, and slightly dilated toward a sheathing base: principal leaves gradually reduced upward on the stem, sessile, firm, linear-oblanceolate to elliptic- lanceolate, gradually tapered at both ends. 4-12 cm long and ()..'S-2 cm wide, with a prominent midrib and weakly expressed anastomosing secondary nerves, the upper surface glabrous, the lower surface scabrous only along the midrib, the apex acute with a sharp callus point, the margins mostly entire, ciliolate, the base cuneate or rounded and hugging the axillary branchlets; rameal leaves similar iii contour, those of the peduncles often densely spaced, bracteiform. appressed or ascending, oblong to subulate. 1.5—4 mm long, continuous with the phyllaries but distinct in having a pointed apex with a brown or purple mucro. Capitulescence an open, broad, round-topped panicle. Flov*erinj> heads 2 3 cm in diameter when the rays are tully extended, nt)t crowded and not secund, usually at the ends of wiry, ascending or divaricate, few- to many- bracted peduncles ( 1 )4-10(2.'i) cm in length (rarely sessile). Involucre slenderly campanulatc or turbinate, 7-12 mm high, the phyllaries imbricated in 6-9 series, inserted on a prominent obconical rachis 2—t mm in height. Phyllaries strongly graduated, appressed or somewhat spreading, indurate, rounded on the back or slightly keeled, the green areoles well delimited, oblong to rhonibic-oblanceolate. centered in the apical 'A -7' portion, usually much shorter than the scarious basal portion, the apex obtuse. rounded, or almost truncate, sparsely lanate. including the top (areolar) portion of the adaxial surface, the margins with a comparatively broad scarious rim extending to the tip; outer (lower- most) phyllaries ovate. 1.5 mm long, the median and inner ones linear, to 7 mm long. Receptacle alveolate with sharp teeth. Ray florets 15-20. the corollas 10-15 mm long, blue or purple, glabrous. Disk florets 15-20 (or more), the corollas narrowly funnelform, 4.5-6(7) mm long, glabrous, the limb yellow turning purple after anthesis at least in the lobes, the lobe/limb fraction 0.18. the tube sightly shorter than the limb. Pappus simple, the bristles about as long as the disk corolla, tawny or somewhat rose-tinged, comparatively firm, attenuate. Achenes oblong in contour, somewhat compressed. ( 1.8)2-2.8 mm long and ca 1 mm across, light brown or gray, puberulent or minutely strigillose, often puncticu- late. with .(-5 ribs. 2/i = 96. sometimes with additional B-chromosomes. Late August-October. Dry loamy or rocky soils: edges of upland woods, open-wooded slopes and bluffs, and also in somewhat disturbed ground of pastures and roadsides. Local, mostly in the southern half of the state, north to Macon. Fulton, and McDonough counties. Records from Illinois represent the northern and northeastern limits for the species. The range extends southward to n. LA and westward through s. MO and AR to e. NE, s.e. KS. and e. OK. 182 Illivois Nati ral History Si rvey Bi lletin Vol. 34 An. 2 29. Aster umbellatus MiWeT Flat-top Aster Herbaceous, with a creeping horizontal rhizome system. New shoots initiated at intervals along or at the ends of stoloniform rhizome strands resulting in colonial stands: autumnal rosettes lacking. Stems mostly single at any point of emer- gence, erect, 5()-l 30 cm tall, usually unbranched below the capitulescence, glabrous in the lower portion, commonly puberulent with antrorsely hooked trichomes above, the branches somewhat winged or ridged from decurrent leaf bases. Leaves (except the lowermost) more or less uniform in contour, sessile or subpetiolate. those of the midstem ovate to elliptic-lanceolate, 4—12 cm long and 1-3(4) cm wide, with a prominent midrib that is abaxially keeled in the basal or subpeliolar portion and with 6-8 pairs of pinnate and anastomosing secondary nerves, as well as a conspicuous reticulum of tertiary veins with isodiametric areolae, the upper surface dark green, glabrous or minutely scabrous, the lower surface bluish green, scabrous-puberulent, at least along the midrib and major nerves, the apex acuminate, the margins entire, scabrous, the base cuneate and decurrent; lower cauline leaves smaller, often spatulate, those near ground level bracteiform; rameal leaves relatively few, much reduced in size but otherwise resembling the larger leaves except for the few small, subulate, puberulent bracts subtending the head. Capitu- lescence flat- or round-topped, each individual cluster corymbiform. Flowering heads 1-1. ."1(2) cm in diameter when the rays are fully extended, often crowded, subsessile, or on slender puber- ulent peduncles 0.2-2.5 cm in length. Involucre campanulate or turbinate, 3.5—4.5(5) mm high, the phyllaries imbricated in 4 or 5 series. Phyllaries strongly graduated, the outer ones 1-2 mm long, the inner 3^(6) times as long, more or less appressed at tlowering time but widely spreading in fruiting heads, puberulent or glabrous on the back; outer (louennost) phyllaries subulate, the median and inner ones oblong to linear-lanceolate, the green arcoles forming broad bands to the ba.se as an extension of the midrib, the apex acute to obtuse or sometimes rounded, the margins erose-hyaline and irregularly ciliolate. Receptacle alveolate with long sharp teeth. Ray florets (2)6-15 (or more), the corollas X-10 mm long, often with a few trichomes on the slender tube, the rays whitish, comparatively broad. 2-3 mm wide. Disk florets (8)12-20. the corollas funnelform. 4-7 mm long, thinly puberulent on the tube, the limb partway slenderly tubular, abruptly flared ca 1 mm above the point of insertion of the filaments, cream-colored turning purple after anthesis. the lobes relatively long and reflexed. the fraction of lobe/expanded limb ponion 0.6-0.7. the apparent tube (i.e.. including the tubular limb portion) distinctly longer than the expanded limb portion. Pappus "double" (according to literature references) but actually composed of 3 series of bristles. \v hitish or somew hat discolored; bristles of the inner series about as long as the disk corolla, firm, clavel- lately expanded toward an acute apex, those of the middle series slightly shorter, tapered toward an attenuate apex, and those of the outer series minute. 1 mm long or less, slender, and attenuate. Achenes obovate to oblanceolate in contour, compressed. 2.5-3.5 mm long and 1-1.2 mm across, light brown, sparsely puberulent (Illinois plants) or sometimes glabrous, w ith 5 or 6 prominent, glossy, golden-brown ribs. 2/i = 18. Including A. puhentior Cronquist ( 1947). IDiiellini^erici umhcUcihi (Miller) Nees; A. infirmus misapplied, not of Michau.x — Brendel 1887.] (Late July) August-September (early October). Low. damp, mostly open ground: swamps, seep areas, thickets near streams and rivers, and clearings in woods. Occasional in the northern half of the state, south to Menard and Cass counties. The range of the species extends from NF south to GA. n. FL. LA. s. AR. and e. TX. west to e. AB. SK. and ND. with a few stations in \.\. NE. and s. OK. Based solely on geographic considerations. plants of Illinois would be placed in var. puhens A. Gray (cf. Femald 1950) or. at the rank of species, in Aster puhentior (cf. Gleason 1952: Gleason and Cronquist 1963). Too much variability is evident in the specimens examined, however, to permit a clear separation of this variant from \ar. umhelhmis. Gray (1884) distinguished plants of var. puhens merely by May 1989 Asif.H AM) Bh\( Hi u Jis i\ Illinois 18? more copious pubescence on the lower leaf surface. Cronquisi (1947). in his argument for species rank, considered additional characteris- tics, e.g.. puberulent phyllaries and smaller heads ( 12-22 florets) compared with glabrous phyllar- ies and larger heads (23-54 florets) in M. umhellatus var. umhcllaliis. In applying these criteria, however, one may find thai both taxa occur side by side in northern Illinois. 1 do not. therefore, recognize any varieties of this species for the fiora of Illinois. 30. Aster undulatus L. Wavy-leaved Aster Herbaceous, with branched caudiciform rhizomes that turn woody with age but also some- times with short stoloniform rhizome strands. New shoots arising at the base of old stems or from the rhizomes, the plants usually forming scattered individual clumps. Stems 1 -several, erect. 4()-l20cm tall, with ascending or divaricate branches above the middle, densely and uni- formly hirtellous or villous on the upper stem portion and in the branches, somewhat pubescent in lines or glabrescent in the lower stem portion. Leaves polymorphic, the basal and lower cauline ones petiolate. those higher up on the stem subsessile or sessile and strongly clasping: upper surface of the blade mostly scabrous to hirsute (rarely glabrous), the lower surface hirtellous or loosely villous; leaves of new shoots and of the vernal rosettes with ovate-oblong to suborbicu- late blades 1-6 cm long and 1—1 cm wide, purplish below, with two or more pairs of somewhat arching and anastomosing secondary nerves, the apex obtuse or rounded, the margins shallowly crenate-serrate to subentire. the base cordate or truncate to rounded, sometimes oblique, the petioles usually purplish, hirsute or pilose, winged, dilated, and sheathing at the base; principal cauhne leaves variable in shape and size, gradually reduced upward on the stem, the lower ones petiolate, those higher up sessile and often constricted near or below the middle, the blades ovate to lanceolate in outline, j(-12( 14) cm long and ( I )2-5 cm wide, the secondary venation mostly obscured by indument. the apex acute to attenuate or acuminate and callus-pointed, the margins crenate-serrate or entire, and scabrous, the petioles progressively shorter and more broadly winged upward on the stem, conspicu- ously dilated toward the auriculate or cordate, clasping base; rameal leaves more or less abruptly reduced in size, oblong or lanceolate to linear-lanceolate, acute to attenuate, niucronulate, subentire to entire, sessile, slightly clasping or sheathing; leaves of the peduncles often numer- ous, firm, bracteiform, 2-3 mm long, appressed or ascending, intergrading with the phyllaries. Capitulescence paniculiform. often notably overtopping the leafy stem portion but also sometimes with branches arising from nodes near the middle of the stem, the ultimate head-bearing branchlets ascending or divaricate, often racemifomi. Flowering heads 1.2-1.3(2) cm in diameter when the rays are fully extended, usually not crowded but sometimes secund, the peduncles 0.3-3(5) cm long, often densely bracteate, hirtellous. Involucre campanulate or hemispherical. 4-6(7) mm high, the phyllaries imbricated in 4 or 5(6) series. Phyllaries strongly graduated, appressed, or those of the outer series spreading, lanceolate, 1.5-2.5 mm long, the inner ones linear-oblanceolate, 4—5(6) mm long, puberulent on the abaxial surface and also thinly so on the adaxial surface, the green areoles rhombic or oblanceolate. the apex sharply acute. attenuate, or acuminate with somewhat inrolled margins, often mucronulate, the margins erose- hyaline, irregularly ciliolate, and with a scarious rim to near the tip, the basal 7'-7> portion scarious, or sometimes the midrib green to the base. Receptacle alveolate with mostly rounded teeth. Ray florets (12)15-25, the corollas 8-12 mm long, purple or blue, glabrous or nearly so. Disk florets 15-25. the corollas narrowly funnelform, (4)5-6 mm long, glabrous or with a few irichomes near the throat, the limb abruptly dilated at the throat, cream-colored or light yel- low turning purple after anthesis, the lobe/linih fraction 0.2-0.25. the tube shorter than the Imib. Pappus simple, the bristles about as long as the disk corolla or slightly shorter, cream-colored or slightly rose-tinged, soft, slender, and attenuate. Achenes oblong-obovoid. somewhat compressed. 2.5-3(3.5) mm long and 1.2-1.5 mm across, dull purple or light brown, sparsely puberulent at least 184 Illinois Natlral History Si rvey Bllletin Vol, 34 Art. 2 toward the top, with 4 or ."i straw-colored ribs, 2n - .^2; reported chromosome counts ofZ/i = 18 and 36 are probably in error (A.G, Jones 1977. 1980b). August-October, Dry or well-drained soils: loamy or rocky slopes at forest edges and open- wooded bluffs underlain by sandstone or limestone. Uncommon, restricted to the southern tip of the state: records from Jackson and Alexander counties mark the western limit for this extremely variable species. The range extends from NS and ME to c. FL. west to s. ONC?). OH. through s. IN. s. IL, and TN to c. MS and s.e, LA, Aster undulatus has been included in the Illinois list of threatened species but is one of the more common asters in the mountains and woodlands to the east and southeast of our state. Although several varieties have been described. I am unable to assign Illinois populations to any of them. Even though the plants seem to differ somewhat from the type specimen of A. umhda- liis. 1 am treating them at this time sensii hito or as belonging in var. umlulciiiis. There is evidence of occasional hybridization and intergradation with such closely related species as A. drum- mondii [e.g.. AG. Jones 4089 (ILL) from Gallatin Co.) and A. ootemangicnsis [e.g,. K. Wilson 3164 (ISM) from Pope Co,], Collections that form the basis for A. iindidatus sensu Mead (1846): Higlcy and Raddin ( 1X91 ); Pepoon (1927); Kibbe (1952); and Dobbs (196.^) non L, do not belong in this species. 31. Aster urophyllus Lindley in DC. Herbaceous, with a stout caudicifomi rhizome system lacking stoloni- fomi strands. New shoots arising at or near the base of old stems, the plants forming scattered individual clumps. Steins 1 -several, erect, brittle. 40-1 20 cm tall, with stiftly ascend- ing, somewhat ridged branches from above the middle, sparsely puberulent in the upper portion and m the branches, the indunicnl usually in decurrent lines (rarely unifonmly distributed around the stem), the lower stem portion glabrescent or glabrous. Leaves polymor- phic, the basal and larger cauline ones peliolate. those of the upper stem subsessile to sessile: leaves of new shoots and of the vernal rosettes with ovale or lanceolate blades 4—12 cm long and 2-5 cm wide, nearly glabrous, or often minutely scabrous above and thinly pubescent below, the secondary venation weakly expressed, the apex acute to acuminate, the margins shallowly crenate-serrate. the base cordate or truncate to rounded, the petioles 5-1 5 cm long, slender or narrowly winged, dilated, and sheathing at the base: principal cauline leaves gradually reduced in size upward on the stem, the blades ovate to lanceolate. 5-12 cm long and 2-5 cm wide. similar to the basal leaves in venation and vestiture. the apex acuminate to attenuate with a sharp, often purple callus point, the margins crenate-serrate to subentire. the base truncate or rounded, the petioles progressively shorter and more broadly winged upward on the stem, slightly sheathing: rameal leaves abruptly reduced in size, lanceolate to linear-lanceolate, glabrous or nearly so, attenuate and callus- pointed, entire, scabrous-margined, cuneate and slightly decurrent; leaves of the peduncles bracteifomi, linear or subulate. 2—4 mm long. mostly ascending (rarely spreading to recuned). mtergrading with the phyllaries. Capitulescence typically a dense, narrow, ovoid or pyramidal panicle in the upper 'A -'/{'/:) portion of the stem, commonly overtopping the leafy portion, the head-bearing branchlcts siiftls ascending, often racemifonn. Flowering heads 1-1. 2( 1.5) cm in diameter when the rays are fully extended. usually crowded and sometimes secund. sub- sessile or on often densel\ bracteate. puberulent peduncles 0,3-2 cm long. In>olucre cylindrical or turbinate. 4.5-6(7) mm high, the phyllaries imbricated in 4 or 5 series. Phyllaries appressed or somewhat recur\ed-spreading, graduated, the outer subulate, 2-3 mm long, the inner linear- lanceolate, .5-6(7) mm long, glabrous, searious over most of the surface area, the light green areoles very slender, linear to linear-oblanceolaie. the apev long-attenuate, lemiinaling in an often recurxed. spinulose niucro. the margins erose- h\ aline and irregularly ciliolate. Receptacle ahcolate with sharp teeth. Ra) florets (8)10-12(14). the corollas 6-8(10) mm long. t\picall\ wliiio (rarcU la\cnder). sparsely Mav \9m Asif-K AM) Bku iiyu II', IN li-i-isnis IS5 puberulent on the lube and throal. Disk florets 10-15. the corollas funnelform. abruptly dilated. (3.5)4—5 mm long, glabrous or with a few trichomes near the throat, the limb cream-colored turning pinkish after anthesis. the lobe/limb fraction 0.2-0.25. the tube much shorter than the limb. Pappus simple, the bristles shorter than the disk corolla, whitish or slightly discolored to rose-tinged, soft, slender, and attenuate. Achenes oblong-obovoid. somewhat compressed. l.X-2.5 mm long and ca I mm across, dull purple or brown, glabrous, with 4 or 5 ribs. 2n = 16. [A. saf>htif(iliiis sensu auct. (pro pane) non Wedem. ex Willd. (see A.G. Jones 1980b; Jones and Hiepko 1981 ). A. hiriellus Lindley in DC. A.sagiuifolius var. hiriellus (Lindley in DC.) Burgess in Britton & Brown f. hiriellus (Lindley in DC.) Shinners ( 1941 )—Steyemiark 196.1. A. saiiillifiiliiis var. urophyllus (Lindley in DC.) Burgess in Britton & Brown—Deam 1940. A. conlifdliiis var. moraliis sensu Fern. ( 1950) non (Shinners) Shinners.] August-October. Loamy or rocky soils in dry or mesic situations: woodland edges, open upland woods, and thickets, also in disturbed ground of pastures, roadsides, and railroad rights- of-way. Common throughout much of the state. The range extends in the East from s.e. Canada and ME south to GA, AL, and n. FL, and in the West from MN through lA, MO, and s.e. NE to s.e. KS, with scattered stations recorded from AR and MS. 186 iLLiNors Natural History Slrvey Bulletin Vol. 34 Art. 2 Description of Brachyactis Species Brachyactis ciliata (Ledeb.) Ledeb. Rayless Aster Taprooted and somewhat succulent annual. Stem erect, 10-70 cm tall, bluish or yellowish green, often red-tinged, glabrous or with a few bristly trichomes in the leaf axils and along the decurrent leaf bases; branches mostly ascending, often from near the base. Leaves bluish green, with a conspicuous midrib, the secondary venation faintly expressed; basal leaves spatulate, soon withered; principal cauline leaves linear, 3-10 cm long and 0.1-0.9 cm wide, gradually reduced in size upward on the stem, sessile, with clusters of smaller leaves produced in the axils, glabrous, the apex acute to short- attenuate, the margins entire and appressed ciliate, the base slightly dilated and sheathing; rameal leaves similar in contour, those of the jjeduncles few. Capitulescence a contracted panicle, or racemiform in poorly developed plants. Heads several to many, short-peduncled or subsessile. Involucre 6-10 mm high, the phyllaries imbricated in 3 or 4 series. Phyllaries subequal or somewhat graduated, occasionally those of the outer series longer than those of the middle and inner series, glabrous, linear to oblanceolate, (4)5-8 mm long, the outer and median ones herbaceous except for a narrow scarious margin, the apex acute or obtuse and mucronulate. Receptacle flat, shallowly alveolate with rounded teeth, or merely pitted in fruiting heads. Florets 40-80 (or more). Pistillate florets fertile, in several series, usually more numerous than the disk florets, the corollas slender, tubular, with a long-exserted pink style and no stamens; rays (ligules) absent or rudimentary. Disk florets perfect and fertile, the corollas tubular or narrowly funnelform, 3-4.5 mm long, the limb whitish aging to pink, with 5 very short lobes, the tube slender, yellowish green, longer than the limb. Style branches of the disk florets with attenuate, papillate, sterile appendages that are longer than the stigmatic lines. Pappus very prominent, considerably overtopping the corolla, the capillary bristles numerous, in 2 or 3 series, pure white or faintly pink, subequal in length, 4-6 mm long, soft and silky, attenuate, and minutely barbellate. Achenes oblong-obovoid. slender, slightly compressed, 1.5-2.5 mm long and 0.4-0.5 mm across, gray or whitish, often with purple streaks, pilose or strigillose, obscurely 2-A ribbed. ln= 14. Including B. angusta (Lindley in Hooker) Britton in Britton & Brown (see A.G. Jones 1984). [Asler brachyactis S.F. Blake—G.N. Jones 1945, 1950, 1963; Jones and Fuller 1955; Swink 1974; Mohlenbrock 1975, 1986; Swink and Wilhelm 1979; and others.] August-October. Disturbed ground: waste places, roadsides and railroad tracks. Occasional, recorded from only five counties of the Chicago region; adventive from the North and the West. The species range extends from NT and James Bay, ON, east to PQ, PE. and NB, and west through the northern half of the Great Plains to n. WA; the plants were also recorded from scattered stations in s.e. WI, n. LA, w. MO, w. NE, w. KS, w. OK, and s. and e. CO. The species was originally described (under the name Erigeron ciliatiis Ledeb. ) from collections made in Siberia. North American plants may be recognized as Brachyactis ciliata subsp. angusta (Lindley in Hooker) A.G. Jones [based on Tripolium anguslum Lindley in Hooker. Aster angustus (Lindley in Hooker) Torrey & Gray non Nees—Pepoon 1927. B. angusta—Hill 1902). This taxon. however, can probably not be distinguished from typical subsp. ciliata on any basis other than geograph> . Brachyactis is a small genus of two or three species with the center of distribution probably in North America. The genus can be distin- guished from Aster and other members of the Astereae by a combination of the following; pistillate florets that lack or have inconspicuous ray corollas, a prominent pappus that consider- ably overtops the disk corollas and is composed of numerous subequal bristles in several series, and the uncoinmon basic chromosome number of .V = 7. Most authors retain the taxon as a section of Aster (cf. Houle and Brouillet 1985; Semple and Brouillet 1980a, b). May \9H9 Asirn ANO Bhaciivu IIS IS Illjsois 187 Glossary of Descriptive Terms abaxial side ot an organ away from the axis achene dr\, indehiscent. one-seeded fruit aciculifurm needle-shaped acuminate tapering more or less abruptly lo a slender sharp point, the margins somewhat curved inward acute terminating in a sharp point at an angle ol between 45° and 90°, the margins straight (see also attenuate) adaxial side of an organ facing the axis adventive introduced from an adjacent or nearby region and spreading m the new region alveolate honeycombed anastomosing veins (or ner\es) connected by cross veins (or nerves) anthesis flowering time, i.e., period during which pollination takes place apex; apical tip; pertaining to the tip areole small space clearly marked out on a surface ascending directed upward after arising at an oblique angle or on a curve attenuate gradually tapering to a narrow and slender tip at an angle of less than 45', the margins more or less straight (see also acute) auriculate having auricles, i.e.. with ear-shaped appendages barbellate with minute barbs bracteate having bracts bracleiform bractlike bulliform appearing blistered, bubblelike callus a hard protuberance or thickening campanulate bell-shaped capitulescence cluster (often a large assem- blage) of flower heads (commonly syn- onymi/ed with inflorescence l caudex the usually condensed, thick, tough, persistent, largely underground base of an otherwise herbaceous stem caudiciform stein base shaped like a caudex. I.e.. condensed, thick, and lough cauline pertaining to or arising from the stem cespitose growing in tufts, e.g., several stems from a common base ciliate fringed v. iih slender trichomes (see also fimbriate) ciliolate minutely ciliate. i.e.. the trichomes short cinereous with ash-colored (light gray) indumenl clasping the base of a leaf or petiole nearly or completely surrounding the stem clavellate minutely club-shaped, i.e.. expanded toward the apex cordate heart-shaped, the base with rounded lobes and a sinus corolla perianth whorl composed of petals (united in the Asteraceae) corymbiform shaped like a corymb, i.e.. a flat- topped capitulescence with the lower (outer) branchlels and peduncles longer than the upper (inner) ones crenate toothed with rounded teeth cuneate wedge-shaped with the narrow end at the point of attachment cymiform shaped like a cyme, i.e., resembling an inflorescence in which the central or terminal flower opens first cymule a small few-flowered cyme; refers in the Asteraceae to an ultimate cluster of the cymiform capitulescence deciduous falling off at maturity or after withering decumbent reclining on the ground but with the tips ascending decurrent pertaining to leaf bases or veins that continue downward from the nodes in the form of wings, ribs, or lines of indument dichulomous forked with two nearly equal branches dilated expanded in width or widened in diameter disk floret flower of the central portion of a head, the corolla regular, 5-lobed divaricate spreading at a wide angle erose with the margin appearing eroded or gnawed falcate sickle-shaped fasligiale with the branches partway parallel and close together; broomlikc niament the stalk ponion of a stamen that supports the anther fimbriate fringed with coarse trichomes or processes (see also ciliate) foliaceous leaflike or leafy funnelform resembling or shaped somewhat like a funnel fusiform spindle-shaped, swollen near the middle and tapering at both ends glabrescent nearly glabrous or becoming glabrous 188 iLLiNors Natiral History Sirvey Bllletin Vol. 34 An. 2 glabrous smooth, i.e., without vestiture glandular invested with glands, i.e., with structures that produce a viscid sap glaucous covered with a waxy, bluish bloom that often rubs off easily habit general appearance head capitulum = unit of capitulescence (the individual florets sessile) hirsute having coarse or stiff, spreading or ascending trichomes hirtellous minutely hirsute, i.e., with short, spreading trichomes hispid having rigid spreading bristles or spinules hispidulous minutely hispid hyaline translucent or colorless imbricated overlapping, i.e., shinglelike in a spiral arrangement indument hairy covering indurate hardened inrolled rolled inward or downward at the edges involucre one or more series of bracts (phy Mar- ies) subtending the florets of a head (in the Asteraceae) isodiametric with all the diameters of an areole about equal m length keel; keeled a central abaxial ridge formed by the midrib of a compressed phyllary, bract, or leaf; forming a keel lanate woolly, with long, tangled, curly trichomes lanceolate lance-shaped; much longer than broad, widest near the base and tapering to the apex ligule the strap-shaped limb of a ray floret m the Asteraceae limb upper, expanded portion of a corolla ( m disk tlorets, the portion above the zone of attachment of the filaments) as distinct from the tube portion mucro a sharp, short, and abruptly narrowed point or protuberance mucronate tipped with a mucro mucronulate diminutive of mucronate node the place on the stem where leaves are attached and branches arise ob- Latin prefix meaning inverted, e.g., obovate = inverse of ovate oblique slanting, not straight up, or at an angle that IS not 90° oblong at least two times longer than broad \k ith nearly parallel sides obtuse blunt; if sharp-pointed, wjih an angle of more than 90" ovate; ovoid egg-shaped, with the broader part near the base panicle a variously branched compound inflorescence (in the Asteraceae a compound capitulescence with pedunculate flower heads) paniculiform resembling or shaped like a panicle papillate nipplelike pappus the specialized outer penanth whorl in the Asteraceae composed of bristles (in Aster), awns, or scales peduncle the stalk of a head perfect having both functional stamens and pistils petiole; petiolate the stalk of a leaf; having a stalk phyllary bract of the involucre pilose invested w ith long, soft, slender trichomes pinnate arising from both sides of the axis or midrib pistillate having pistils but no functional stamens polymorphic exhibiting several forms (morphs) of the same organ pro parte in part pro sp. described as a species puberulent minutely or finely hairy pubescence; pubescent hairiness; hairy puUinale svsollen or cushion-shaped puncticulale has ing minute colored or translucent dots pustulate w iih pimplelike or blisierlike raised areas racemiform resembling or shaped like a raceme, i.e.. an elongate capitulescence or branch with pedunculate heads rameal of or pertaining to the branches ray floret of the outer series of fiorets in a head, the corolla strap-shaped (= ligulatel receptacle the expanded end of the peduncle. I.e.. the end bearing the aggregate of lloreis reflexed abruptly bent or turned downward remote disiantl\ spaced reticulate; reticulum interconnected like a network rhizome an elongate, often branched under- ground stem, usuall\ with minute scale leaves and rooting at or producing new shixits from the nodes May 1W4 ,4s;/ K AM) Uk \( //) >i ;/s IS Illinois 1X4 rhombic shaped like a rhombus, i.e., like an equilateral, oblique-angled parallelogram; more or less diamond-shaped roselte a cluster of leaves with very short intemodes, arranged in a compact, spiral (near-circular) series rugose; rugulose wrinkled scabrellous minutely scabrous scabrous rough to the touch, the stiff irichomes pointing in one direction scarious thin, dry, membranous, not green secund directed to one side of the stem or branchlel sensu aucl. according to author(s) sensu lato in a broad sense sensu siricto in a narrow sense sericeous silky, with appressed. soft, glossy irichomes serrate with sharp teeth thai point forward serrulate finely serrate sessile lacking a stalk sheathing closely enveloping spatulale spatula-shaped or spoon-shaped spiciform shaped like a spike, i.e.. a simple. elongate capitulescence with sessile heads spinule; spinulose short spine; minutely spiny squarrose having the parts recurved at the tip (mostly applied to phyllaries) stamen pollen-bearing organ stigmatic pertaining to the portion of the style branches receptive to pollen stolonirorm shaped like stolons (applied to long-creeping horizontal rhizomes) striate marked with fine longitudinal lines or ridges strigillose minutely strigose. i.e.. the trichomes very short strigose with the trichomes appressed and lined up in one direction style the elongated part of the pistil above the ovary subulate awl-shaped; slender, tapering to a sharp point sufTruticose plants woody only at the base and herbaceous over most of the above-ground portion terete circular in cross section throat expanding zone in a corolla at the junction of tube and limb thyrsiform shaped like a thyrse. i.e.. like a compact or contracted panicle tomentose invested with short-haired, matted, woolly pubescence trichome an epidermal structure (hair, bristle. or prickle) truncate ending abruptly as if cut off turbinate lop-shaped, i.e., inversely conical umbelliform resembling or shaped like an umbel, i.e., a flat-topped capitulescence in which several peduncles more or less arise from a common point urceolate urn-shaped, e.g., a corolla that is abruptly contracted just below the mouth vestiture any covering of the surface, i.e., indument and/or glands villous woolly, the trichomes long, soft, and curly but not matted or tangled 190 Ii I isois Nati ral History Sl r\ fy Bi lletin Vol. 34 An. 2 Literature Cited Allen, G.A. 1985. The hybrid origin of Asler ascendens (Asteraceae). American Journal of Botany 72:268-277. Benke, H.C. 1928, Some Illinois asters and a new variety of A mulliflonis. Rhodora 30:77-79. Benke, H.C. 1930. Aster amelhystinus an obvious hybrid. Rhodora 32:1-3, pi. 192. Benke, H.C. 1932a. Some field notes: a new variety and some forms of plants from the Middle West; also two forms from Massachu- setts. Rhodora 34:4-12. Benke, H.C. 1932b. New asters from Illinois. The American Midland Naturalist 13:326-328. Blake, S.F. 1930. The names Aster encoides and A. mulliflonis. Rhodora 32:136-140. BoiviN, B. 1972. Flora of the prairie provinces. Part 3. Phytologia 23:1-140. Brendel, F. 1887. Flora Peoriana. J.W. Franks and Sons, Peoria, IL. Cronqi iST, A. 1947. Notes on the Compositae of the northeastern United States. V. Astereae. Bulletin of the Torrey Botanical Club 74:142-150. Cronquist. A. 1980. Vascular flora of the southeastern United States. Vol.1. Asteraceae. The University of North Carolina Press, Chapel Hill. Deam, C.C. 1940. Flora of Indiana. Indiana Department of Conservation, Indianapolis. DoBBS, R.J. 1963. Flora of Henry County, Illinois, an annotated list of the vascular plants. Natural Land Institute, Rockford, IL. Fell, E.W. 1955. Flora of Winnebago County, Illinois, an annotated list of the va.scular plants. The Nature Conservancy, Washington. DC. Fernald, M.L. 1950. Gray's manual of botany. 8th ed. American Book Company, New York. (Corrected printing, 1970). Gleason, H.A. 1952. The new Bntlon and Brown illustrated flora of the northeastern United Stales and adjacent Canada. The New York Botanical Garden, New York. (3rd printing, slightly revised, 1963). Gleason, H.A., and A. CRONyi ist. 1963. Manual of va.scular plants of northeastern United States and adjacent Canada. Van Nostrand-Reinhold Company, New York. Gray. A. 1884. Synoptical flora of North Amenca. Vol. 1, Pan 2. Iveson, Blakeman, Taylor and Company. New York. Hir.LEY, W.K., AND C.S. Raddis. 1891. The flora of Cook County. Illinois, and a part of Lake County. Indiana. Bulletin of the Chicago Academy of Sciences 2:i-xxiii, 1-168. Hill. E.J. 1902. Notes on migratory plants. Bulletin of the Torrey Botanical Club 29:564-570. Holmgren. P. K„ W. Kelkes, and E.K. ScHOFiELD. 1981. Index Herbariorum. Ed. 7. Regnum vegetabile vol. 106. HoLLE, F., and L. Brol illet. 1985. Chromo- some number determinations in Aster section Conyzopsis (Asteraceae). Britionia 37:369-372. Jones, A.G. 1974. Aster ericoides L. east of the Mississippi River. The American Midland Naturalist 92:466-475. Jones, A.G. 1977. New data on chromosome numbers in Asler section Heterophylli (Asteraceae) and their phylogenetic implica- tions. Systematic Botany 2:334-347. Jones, A.G. 1978a. Observations on reproduction and phenology in some perennial asters. The American Midland Naturalist 99:184-197. Jones, A.G. 1978b. The taxonomy of Aster section Miiltifli>ri (Asteraceae).— 1. Nomen- clatural review and formal presentation of taxa. Rhodora 80:319-357. Jones, A.G. 1978c. The taxonomy of Aster section MM/f//7()n (Asteraceae).—II. Biosys- tematic investigations. Rhodora 80:453-490. Jones. A.G. 1980a. A classification of the New World species of Aster (Asteraceae). Brittonia 32:230-239. Jones, AG. 1980b. Data on chromosome numbers in Aster (Asteraceae), with com- ments on the status and relationships of cenain North .Amencan species. Brittonia 32:240-261. Jones, A.G. 1982. \ ir(;uliis Raf. vis-a-vis /t.s/cr L. (Asteraceae). Taxon 31:714. Jones, A.G. 1983. Nomenclatural changes in Aster (Asteraceae). Bulletin of the Torrey Botanical Club 110:39-12. Jones. A.G. 1984. Nomenclatural notes on Asicr (Asteraceae)—II. New combinations and some transfers. Phytologia 55:373-388. May msy ASTFK AUD Bn \CHrA(TIS IN ILLINOIS 191 Jones, A.G. 19H5. Chromosomal features as generic criteria in the Astereae. Taxon 34:44-54. Jones, A.G. 1987. New combinations and status changes in Asier ( Asteraceae). Phytologia 63:1.31-133. Jones, A.G.. and P. Hiepko. 1981. The genus Asler S.I. (Asteraceae) in the Willdenow Herbarium at Berlin. Willdenowia 1 1 :34.V360. Jones, A.G., AND P.P. Lowrv. 1986|I987|. Types and selected historic specimens of Aster S.I. (Asteraceae) in the Herbarium, Laboratoire de Phanerogamie, Museum National d'Histoire Naturelle. Paris (P). Bulletin de Museum National d'Histoire Naturelle, Paris, Series 4, Section B ( Adan- sonia)8(4):393^l2. Jones, A.G., and D.A. Young. 1983. Generic concepts of Asler (Asteraceae): a comparison of cladistic, phenetic, and cytological approaches. Systematic Botany 8:71-84. Jones. G.N. I94.'>. Flora of Illinois. The Ameri- can Midland Naturalist Monograph 2. The University Press. Notre Dame, IN. Jones, G.N. 1930. Flora of Illinois. 2nd ed. The American Midland Naturalist Monograph .3. The University Press. Notre Dame, IN. Jones, G.N. 1963. Flora of Illinois. 3rd ed. The American Midland Naturalist Monograph 7. The University Press, Notre Dame, IN. Jones, G.N., and G.D. Fuller. 1955. Vascular plants of Illinois. University of Illinois Press, Urbana, and Illinois State Museum, Spring- field. Jones, R.L. 1983. A systematic study of Asler section Potenles (Asteraceae). Sida 10:41-81. KiBBE, A.L. 1952. A botanical study and survey of a typical mid-western county (Hancock County, Illinois). Published by the author, Carthage, IL. Mead, S.B. 1846. Catalogue of plants growing spontaneously in the state of Illinois, the principal part near Augusta, Hancock County. Prairie Famier 6:35-36, 60, 93, 1 19-122. MoHLENBKocK. R.H. 1975. Guide to the vascular flora of Illinois. Southern Illinois University Press, Carbondale and Edwardsville. Mf)iiLENBRorK, R.H. 1986. Guide to the vascular flora of Illinois. Revised and enlarged |2nd| ed. Southern Illinois LInivcrsity Press, Carbondale and Edwardsville. MoHi.ENBRorK, R.H., AND D.M. Ladd. 1978. Distribution of Illinois vascular plants. Southern Illinois University Press, Carbon- dale and Edwardsville. MOHLENBROCK, R.H., AND J.W. VoiGT. 1959. A flora of southern Illinois. Southern Illinois University Press, Carbondale. Pepoon, H.S. 1927. An annotated tlora of the Chicago area. Chicago Academy of Sciences, Natural History Survey Bulletin 8:1-554. Reveal, J. K., andC.S. Keener. 1981. Virgidus Raf ( 1837). an earlier name (or Lasallcii Greene ( 1903) (Asteraceae). Taxon .30:648-651. Semple, J.C. 1979. The cytogeography of Asler lanceokiliis Willd. (synonyms: A. simplex and A. paniculaius) in Ontario with additional counts from populations in the United States. Canadian Journal of Botany 57:397^02. Semple. J.C, and R.A. Brammall. 1982. Wild Aster tanceolaliis x laleriflorus hybrids in Ontario and comments on the origin of A. onlarioiiis (Compositae—Astereae). Canadian Journal of Botany 60:1895-1906. Semple, J.C, and L. Brolullet. 1980a. A synopsis of North American asters: the subgenera, sections and subsections of Aster and Lasallea. American Journal of Botany 67:1010-1026. Semple, J.C, and L. Brouillet. 1980b. Chromosome numbers and satellite chromo- some morphology in Aster and Lasallea. American Journal of Botany 67:1027-1039. Semple, J.C. and J.G. Chmielewski. 1985. The cytogeography of Aster pilosus (Composi- tae—Astereae) II. Survey of the range, with notes on A. depaiiperatus. A. parvieeps and A. porteri. Rhodora 87:367-379. Semple, J.C, and J.G. Chmielewski. 1987. Revision of the Aster lanceolatus complex, including A. simplex and A. hesperius (Compositae: Astereae): a multivariate morphometric study. Canadian Journal of Botany 65:1047-1062. Shinners, L.H. 1941. The ^enus Asler in Wisconsin. The American Midland Naturalist 26:398-420. Shinners, L.H. 1949. Aster ontarionis the same as A. pantotrichus I.A. missimricnsis). Rhodora 51:89-91. Steyermark, J.A. 1963. Flora of Missouri. Iowa State University Press, Ames. 192 Illinojs Nau ral History Survey Bllletin Vol. 34 An. 2 SwiNK, F. 1974. Plants of the Chicago region. 2nd ed. The Morton Arboretum. Lisle, IL. SwiNK, F.. AND G. WiLHEi.M. 1979. Plants of the Chicago region. Revised and expanded |3rd| ed. The Morton Arboretum, Lisle, IL. ToRREY, J., AND A. Gray. 1841. Flora of North America, Vol. 2, Part 1. Wiley and Putnam. New York. Wetmore, R.H., AND A.L. Delisle. 1939. Studies in the genetics and cytology of two species in the genus Aster and their polymor- phy in nature. American Journal of Botany 26:1-12. WiEciAND. K.M. \92^. Aster laieriflorus and some of its relatives. Rhodora 30:161-179. WiEGAND, K.M. \9?i?i. Aster paniculatus ind some of its relatives. Rhodora 35:16-38. I Mav 1^89 ASIF.R AND SR U7/1 « T/S 1\ III INOIS 193 Index to Scientific Species Names, Including Synonyms The names of accepted Illinois species are given in roman type. Bold-faced page numbers indicate species descriptions. Aster X amethystinus 141. I4fi. 156. 166. 166 ani>usnis 1 S6 anomalus 144. 145.149. ISO azureiis 139. 169 borealis 139. 141. 144. 148, 150-151. 161 hrachyculis 139. 143. IS6 carneiis 1 74 chasei 139. 141. 178 ciliolatus 141,144.146.152,152 commulatus 1 56 cordifolius 141, 144, 146, 151-152, 152, 185 depiiuperatus 170 diffiisus 162 (livarlcalus 1 59 drummondii 144. 145, 152, 153-154, 169. 184 dumosus 141, 144, 148, 151, 154-155, 158, 162 eatonii 1 5 1 ericoides 139, 141, 144. 147, 155-156. 156. 166. 170, 172 e.xiiiiiiis 1 56 falcatus 141.144,147,156 finkli 154 firmus 1 76 fragilis 140, 144. 148, 155, 157-158, 161, 162. 163, 168 furcatus 141.144,145,158-159 hirsulicaulis 1 62 hirlellus 185 honzi'inaUs 1 62 infirmiis 1 82 interior 1 6 1 junceus 1 50 junciformis 139, 150 Kunileini 1 67 laevis 144, 145, 147, 152,159-160, 161. 169, 177 lanceolatus 139, 144, 148, 151, 155, 158, 160, 160-161, 163, 168, 174, 177 laterillorus 144, 148, 158, 160, 161, 162-163, 168. 177 linariifolius 144, 146, 163 limllcyanus 152 lonnifoliu.s 151, 176, 177 lucidulus 176 lulescens 140 macrophyllus 141, 144, 145, 164-165 miser 1 62 missDiiriensis 1 68 nmllijlonis 1 56 novae-angliae 141, 144, 146, 156, 165-166, 166 n(ivi-hfli;ii 1 76 iH( iJcnliilis 151 oblongifolius 144, 146, 166-167 ontarionis 139, 144, 148, 158, 163, 167-168 oolentangiensis 139, 144, 145, 148, 160, 168-169, 184 punuiilalus 161 panidlrichus 139, 168 parviceps 139, 141, 144, 147, 169-170 patens 144, 146, 170-171. 176, 177 pawnlissimus 1 7 1 pcndulus 1 62 pilosus 139, 144, 147, 156, 170, 171-173 polxphyUii.s 1 72 praealtus 144, 147, 173-174, 177 prenanthoides 141, 144, 147, 174-175 pringlei 172 piurmicoides 139, 140, 143 pnhenliiir 1 82 puniceus 144, 147, 160, 161, 174, 175-177 sui;itlifoliiis 152, 153, 185 salicifoliiis 1 74 schreberi 141,144,145,177-178 sericeus 144, 146, 178-179 shortii 144, 145, 149, 154, 179-180 simplex 139, 161 suhasper 1 74 tataricus 139, 141, 144, 147, 180-181 iciniifoliiis 1 70 tradcscantii 161 turbinellus 144, 147, 181 umbellatus 144, 146, 182-183 undulatus 141. 144, 145, 153, 154, 183-184 urophyllus 144, 146, 152, 154, 180, 184-185 villosus 172 vimineus 157, 161, 162 Brachyactis (;/i,i;«.v/<; 186 ciliaia 140,143,145,186 Diplopappus linariifolius 1 63 lulescens 140 Doellingeria umhellalii 1 82 Erif;eron ciliatus 1 86 Solidago X lulescens 140 ptamiicoides 143 riddellii 140 Tripolium anf>iiMum 1 86 194 Illinois Natlrai History Sirvey Bllletin Vol. 34 Art. 2 Index to Vernacular (Common) Names Aromatic Aster 166 Azure Aster 168 Big-leaved Aster 164 Blue Aster 149 Blue Wood Aster 151 Brittle Aster 140, 157 Bushy Aster 154 Calico Aster 162 Crooked Aster 1 74 Crooked-stemmed Aster 1 74 Drummond's Aster 153 Flat-top Aster 182 Flax-leaved Aster 163 Forked Aster 158 Frost-flower 143 Frost-weed Aster 171 Hairy Aster 171 Heart-leaved Aster 151 Heath Aster 155 Large-leaved Aster 164 Late Purple Aster 170 Lindley's Aster 152 Many-rayed Aster 149 Michaelmas Daisy 143 New England Aster 165 Ontario Aster 167 Panicled Aster 160 Purple-stemmed Aster 175 Rayless Aster 1 86 Red-stemmed Aster 1 75 Rice-button Aster 154 Rush Aster 150 Savory-leaved Aster 1 63 Schreber's Aster 1 77 Shorts Aster 179 Side-tlowered Aster 162 Silky Aster 178 Sky-blue Aster 168 Small-headed Aster 169 Small White Aster 157 Smooth Aster 159 Smooth Blue Aster 159 Spreading Aster 170 Starved Aster 162 Starwort 143 Stiff Aster 139 Swamp Aster 175 Tartarian Aster 180 Wavy-leaved Aster 183 Western Heath Aster 156 White Prairie Aster 1 55 White Woodland Aster 1 62 Wild Aster 143 Willow Aster 173 Willow-leaved Aster 173 Wreath Aster 155 I I i i Illinois Natural History Survey 607 East Peabody Drive Champaign, Illinois 61820 A Division of the Illinois Department of Energy and Natural Resources