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DISTRIBUTION AND TAXONOMY OF SYMPHYOTRICHUM SERICEUM AND S. PRATENSE (ASTERACEAE: ASTEREAE) PDF

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AND TAXONOMY SYMPHYOTRICHUM OF SERICEUM DISTRIBUTION AND PRATENSE (ASTERACEAE: ASTEREAE) S. L CTheoWitsell Ronald Jones of Dept. Biological Sciences 1500 Tower Building 323 Eastern Kentucky University Center Street :hmond Kentucky USA. Arkansas 7220 40475, Rock, U.S.A. Little 1, t Nesom Guy L h 2925 USA Fort Worth, Texas 76109, guynesoim@sbcglob( im.com. http://guynesc il.net; ABSTRACT Nesom Nesom and two Symphyotrichum sericeum (Vent.) pratense (Raf.) are closely related taxa of central S. & and eastern United In most treatments they have been maintained as distinct species Correll States. (e.g., When Nesom Johnston 1970, Cronquist 1980, Jones 1990, 1994, Brouillet et 2006). the taxa are considered al. & Wunderlin Hansen to be conspecific (see 2004), pratense treated as sericeum var. microphyllum (DC.) 5. is S. & & known many Wunderlin Hansen. Symphyotrichum was years Aster B.F. pratense for as phyllolepis Torr. from Texas North Dakota Symphyotrichum A. Gray. sericeum generally distributed in the central to states, is and southern Canada, western Michigan. Symphyotrichum pratense largely concentrated in eastern Texas to is — and western Louisiana. Populations of plants in this complex that are disjunct further eastward to Florida, — Virginia, and Kentucky are somewhat intermediate in phyllary morphology and have been variously treated number and widely as sericeum, pratense, or sericeum var microphyllus, a of states treat these often S. S. S. Chromosome need both scattered populations as counts also clarification, as ploidy levels (2n state-rarities. = 10 and In = 20) have been published for both taxa (Brouillet et 2006). The objective of this paper is to al. update information on complex, geographical point out distinguishing available this clarify distributions, chromosome and summarize and review information, features, identify typical habitats associates, available the state-rarity status of the taxa. 732 Journal of the Botanical Research Institute of Texas 2(1) The present study provides detailed view of the geographic distribution of Symphyotrichum sericeum a MO, BRIT-SMU, EKY, GA, LSU, from and based primarily on study of herbarium specimens pratense, a S. NCU, NLU, TENN, TEX-LL, UNC, VDB, and Maps augmented by records from various published VPI. are A chromosome numbers was The on and also reviewed. associates, sources. available literature habitats, new chromosome from Kentucky population based on mitotic counts from root tip squashes report a is from germinating achenes. The key ased on a detailed examination of morphological characteristics of is from two herbarium specimens across the ranges of the taxa. SYMPHYOTRICHUM SERICEUM from Arkansas and and Habitat.—The primary Symphyotrichum sericeum Distribution distribution of is Oklahoma southern Canada western Michigan and Indiana, with disjunct populations in central to to sandy dunes, Texas Over the larger part of range, sericeum occurs in a variety of habitats, e.g., (Fig. its S. 1). open outcrop— with woods, sandstone glades, over granite but typically associated seasonally dry, glades, on Edwards where semi-open Symphyotrichum sericeum in Texas mostly occurs the Plateau, habitat. it to An grows on limestone commonly on open, rocky slopes and roadbanks. early collection supposedly hills, Edwards made the (Harris Co.: Lindheimer BRIT) similar in every respect to Plateau plants far to east 78, is number and may be mislabeled. Swink and Wilhelm (1994) a of associates of this species in Illinois, list and including Amorpha canescens, Schizachyrium scoparium, and Bouteloua curtipendula in dry hill prairies, Lake Anemone and cernua in sand barrens near Spiranthes sites cylindrica, Liatris aspera, Liatris cylindracea, and sometimes under on Michigan. In Wisconsin occurs dry prairie hillsides, outcrops, bluffs, at sites it open dominated by Schizachyrium scoparium, with oaks, jack pine, or red cedar, but usually in the in habitats & Manitoba and (Cochrane 2000). In the Dalea purpurea, eupatorioides Asclepias Brickellia litis viridiflora, species occurs in remnant tall-grass prairie sites and in openings in Quercus macrocarpa/Populus tremuloides & Hamel woodlands (Foster 2006). The primarily associated with the Central Lowlands Physiographic Province in the U.S., but species is A and Symphyotrichum also occurs in the Ozark Plateau Province of Arkansas Missouri. recent collection of it sericeum from open shale barrens on a southwest-facing slope in central Arkansas, is apparently the first record from the Ouachita Mountains Province. This disjunct from the nearest populations about 155 site is km on Ozark Voucher: Garland Co.: Ouachita Mountains, Ouachita National the northeast the Plateau. to and Walnut Creek on upper above Forest, steep S-facing shale barrens (Mazarn Formation) slope, east of 750 near point which flows into Lake Ouachita, Crystal Springs 7.5' quadrangle, steep glades, elev at it A (ANHC,BRIT, UARK). hundreds of plants in high quality shale barrens, 29 Sep 2006, Witsell 06-471 ft, km above northwest second population was subsequently found in similar habitat of the 1.9 site. Chromosome Numbers.— There are about a dozen published diploid counts for Symphyotrichum seri- & & Semple Semple ceum, these mostly from the northern portions of range (Jones 1980; Brouillet 1980; its Chmielewski Semple 1989; Semple 2002), with the most southerly count from Izard Co., Ar- 1987; et et al. al. kansas (Sherif 1983). There apparently only a single report of a tetraploid count (Love 1982) for S. seri- et is al. same comm.). from Semple, ceum, count from Canada; there are also diploid counts this vicinity pers. this (J. Nesom synonym Taxonomy.— Symphyotrichum lucayanum as a of Brouillet (2006) treated (Britt.) et al. more Nesom, based on sericeum, but lucayanum here considered to be closely related to S. concolor (L.) S. 5. is The and morphology and geographical proximity. in capitulescence phyllary their closer their similarities Nesom was by widespread the southeastern U.S.A. Symphyotrichum lucayanum treated (1994) as a in latter is based on very loosely racemiform capitulescence with interspersed leaves only slightly distinct species, its more reduced or not from the proximal cauline, different from the densely congested, bracteate capit- at all ulescence of Turner (1982) treated the taxon as a synonym of Aster concolor L., and the type of concolor. S. & was by 1974 lucayanum Uillspaugh 2448, NY!) annotated W.T. Gillis in as A. concolor. Aster (Britton Britt. State-Rarity.—Symphyotrichum sericeum currently ranked as S2 (Threatened or Rare) in Arkansas, is Canada Threatened Indiana, Michigan, and North Dakota (NatureServe 2008). listed as Nationally in is Symphyotrichum sericeum and pratense Jones et al„ Distribution of S. from Sym d are from vouchered observations; dotted circles are literature Distribution of circles Fig. 1 i . Journal of the Botanical Research Institute of Texas 2(1) by and NatureSer Brouillet (2006) et ir al. — Distribution and Habitat. The densest distribution of Symphyotrichum pratense in Texas, Louisiana, and is — southern Arkansas scattered, disjunct clusters of populations occur eastward over a wide geographic range and in a variety of physiographic provinces Symphyotrichumpratense in Texas characteristically occurs (Fig. 2). in sandy soil (sand, sandy loam, sandy clay, silty clay, silty clay-loam), being largely restricted to the follow- Oak Woods ing natural Piney Woods, and Gulf Coast and Marshes, and Blackland regions: Prairie, Prairie Within Texas. Dallas 12032 Prairie. these regions, less frequent in blackland prairie: Co.: Lundell is e.g., it & who name (LL, SMU). Shinners recognized the legitimacy of the Rafinesque ahead of the Torrey (1950), Gray name, and showy open oak woods and described this species as frequent plant of transition-belt "a and and prairies thoughout east Texas ." In Texas, pratense also occurs in longleaf loblolly pine areas in S. . . and been from and coastal prairie grassland, has collected roadsides, clearings, fencerows. fields, it and In Louisiana the plants occur in pine flatwood blackland prairie, chalk calcareous clays, sites, hills, A on as well as along dry roadsides. blackland prairie locality for Symphyotrichum pratense is the "Kieffer Winn Brown 8220 (SMU) and from same (LSU Herbarium Prairie:" Par.: five other collections the locality Online Database 2008). At the Kieffer Prairie pratense grows with associates such as Schizachyrium sco- 5. and parium, Andropogon gerardii, Sorghastrum nutans, Echinacea pallida, Oligoneuron rigidum, Vernoniamissurica (USDA Forest Service 2008). open In the Gulf Coastal Plain of Arkansas, typical habitat for Symphyotrichum pratense saline is — barrens treeless openings in Pinus taeda/Quercus flatwoods with clay soils containing high levels of similis who sodium and/or magnesium. Eric Sundell, discovered the species in Arkansas the Warren Prairie at first number (Sundell 1983), noted that the plants occurred with prickly pear and dwarf palmetto and that a of other rare species were associated with these including Geocarpon minimum and Schoenolirion wrightil sites, USDA NRCS The (USDA, and Newton Arkansas Plants website 2008) Izard counties in for pratense, lists 5. but these records from northern Arkansas are based on collections of sericeum. 5. Eastward Alabama, Florida, Georgia, Kentucky, Mississippi, Tennessee, and Virginia), Symphyotrichum (to and pratense occurs mostly in calcareous habitats, chalk barrens, cedar glades, limestone outcrops. Krai e.g., (1981) reported on the species Coastal Plain in Alabama (Sumter Co.) and Florida (Gadsden Co.) and at sites noted that the plants in Alabama were found in blackland prairie patches and that those in Florida were The was from the coast. species subsequently discovered in Decatur Co., Georgia, just across the state line & Gadsden on Decatur 80099 Co., in calcareous glades a Co.: Godfrey Gholson (GA). In Mississippi bluff: McCook, The the species occurs in association with calcareous chalky outcrops pers. comm.). central (L. Alabama and Barone The most Mississippi populations are located in the Black Belt prairie lands (see 2005). Rim Alabama Highland northerly populations in occur in limestone prairie barrens within the province, many with along other rare species, including Dalea Leavenworthia alabamica, Cypripedium can- gattingeri, and magnicamporum (Webb didum, Eriogonum longifolium var. harperi, Schoenolirion croceum, Spiranthes et al. 1997). At most northerly range limits in Kentucky, Symphyotrichum pratense found in glades or barrens its is Low Rim habitats of the Interior Plateaus (chiefly Highland sites) with Juniperus virginiana and Quercus woody marilandica the dominants and with Andropogon Schizachyrium scoparium, Sorghastrum as gerardii, and These nutans, Sporobolus compositus, as the conspicuous grasses. habitats often contain S. vaginiflorus a variety of other rare or infrequent plants, such as Cypripedium candidum, Echinacea pallida, Liatris aspera, and Lithospermum Dalea Dalea Liatris cylindracea, canescens, Silphium pinnatifidum, Candida, purpurea, Phys- ostegia virginiana (Cranfill 1991; Kentucky State Nature Preserves Commission 2008). The barrens sites in Rim Tennessee (Highland and Appalachian Plateau and in southwestern Virginia (Valley and Ridge) sites) Jones et Distribution of Symphyotrichum sericeum and S. pratense al., dotted from Distribution of Symphyotrichum pratense. Solid circles are from vouchered observations; circles are literature records. 2. Fig. number (DeSelm Ludwig are very similar to the Kentucky sites and also contain a of very rare species 1990; 1999; Tennessee Natural Heritage Program 2008). In northern Georgia the plants occur at both Appalachian and and Ridge comm.). In the Coosa watershed of Georgia, Plateau Valley Patrick, pers. rive at sites (T. : with habi include other iated this at Asclepias hirtella, pra, S. Hypericum Buchneraamericana, Dalea Echinacea urn dolabrijorme, gattingeri, pallida, \tra\is, I Depa and magn: (Georgia sphaerocarpum, squarrosa, Spiranthes Liatris 7i 2007). Recent investigations of dolomite and limestor n southwestern Virginia have disc - by Ludwig subsequently Symphyotrichum reported treated several populations of pratense initially (1999), We by Weakley and Virginia Botanical Associates (2008). have studied the Brouillet (2006), (2007), et al. W km following vouchers: Russell Co.: E of Carterton, one-acre barren at top of SW-facing slope on end 1 May 20 1996 Wie- of a short ridge, dense grass cover with dolomitic limestone cobbles, [not yet in flower], km hardwood with boldt 9415 (VPI); river knob 1 E of Carterton, top of SW-facing slope, substrate cobbly and 2100 dominant with aspera Andropogon scattered over dolomitic limestone, species Liatris gerardii, (?) ft, most of one-acre barren, 17 Sep 1996, Wieboldt 9605 (VDB, VPI); E side of Mill Creek where cuts through it km N between Clinch River and Reeds of Jessees Mill, SW-facing limestone hillside in ridge Valley, ca. 1.9 3082 2040 20 Sep Ludwig open grassland dominated by Schizachyrium scoparium, ca. plants, 11 1996, ft, Wise Fleming 14122 (GMUF). Ludwig (1999) noted that this species has been found at six Russell (VPI). Co.: , 736 Journal of the Botanical Research of Texas Institute 2(1) Chromosome Numbers.— The 2n=20 first report of the tetraploid level of for Symphyotrichum pratense who was from Gonzales Texas, by Semple and noted count had been Co., Brouillet (1980), that a diploid made earlier for the species, but that report was unvouchered and part of a 1945 embryological study by German anatomists. Semple and Chmielewski published counts from Grimes and (1987) diploid plants for Walker counties Texas and count Milam Two Semple in a tetraploid for plants in Co., Texas. years later et. we (1989), reported a diploid count of 2n=10 for plants from Gadsden Co., Florida. In addition, report al. KY a previously unpublished 2n=20 count for pratense: Kentucky. Hardin Co.: cedar glade off 2762, 6 S. Nov 5457 1987, R.L.Jones (EKY). — Taxonomy. main In the part of the range of Symphyotrichum the plants have phyl- pratense, typically and lary surfaces mostly glabrous strongly contrasting with the spreading- margins. Phyllaries of the ciliate Virginia plants are unusual, having densely sericeous surfaces, with or without strongly margins, but ciliate resemble typical pratense of Texas and Louisiana in the proximal indurate portion of the phyllary being S. relatively short, the distal leafy portion being relatively broad and long, and the overall phyllary length about Some 1.5 cm. Their geographic position also obviously suggests that they are part of pratense. plants from S. Roane Tennessee are similar in their phyllary features Co.: Morton 1307, SMU). (e.g., The description of Aster montanus Nutt. seems to refer to Symphyotrichum pratense; Nuttall noted that it was "nearly allied to the preceding [Aster sericeus], but distinct." Torrey and Gray (in N. Amer.) treated Fl. known They noted was from and North as A. sericeus also that the species "Tennessee Carolina var. it p. On near the mountains." 1816 he crossed from Tennessee and went on Nuttall's Asheville, N.C., trip, to name to Rutherfordton and Morganton (Graustein 1967). Gray Synop. N. Amer. formalized the (in Fl. at ) varietal rank, as Aster sericeus var. montanus (Nutt.) Gray, noting that the involucre "sometimes glabrate is An and villose-ciliate; approaching the next species [A. phyllolepis]" image of the holotype of A. montanus cm much was obtained from PH, and an examination indicates that the involucral widths are about 2 wide, we than and name synonomy larger typical sericeum, therefore place this in the of pratense. 5. S. known Today nor North Gray neither sericeum pratense occur in Carolina. N. Amer.) S. S. is to (in Fl. North and listed Carolina as included in the range of A. Synop. N. Amer.), he sericeus var. later (in Fl. p, specified Buncombe County as the of the collection. The specimen upon which Gray apparently based site comment was NY, and an image was The from two his located at obtained. sheet bears three plants dif- areas— two from Minnesota and was from ferent are identified as Aster while the other collected sericeus, "Buncombe, North and specimen Carolina, Gibbes" identified as "A. sericeus (montanus Nutt.)." This latter p. known heads and exhibits the large of pratense, here assigned to that taxon; represents the only record 5. is it from North been and of the species Carolina. pratense has not subsequently rediscovered in the can state S. be assumed be extirpated to there. monk anus Nutt., Gen. N. A]ner. Pi. 2:156. 1818 (non A. inus All. 1785). Aster sericeus var. No n nnessee. other collection c Nuttall September to early Octob< 1816, fide Graustei 1967], annotate Lata, [T. s.n., >r When these disjunct populations of Symphyotrichum pratense were discovered in eastern especially first U.S., more at the northerly sites in Alabama, Tennessee, and Kentucky, they were often identified as sericeum. 5. We They now such confirm examined are listed as in several state natural heritage databases. speci- still all mens from Alabama, Florida, Georgia, Kentucky, Mississippi, North Carolina, Tennessee, and Virginia as — main State-Rarity. Symphyotrichum pratense generally rare outside range in Texas and Louisiana is its — it is currently considered Threatened (S2) in Arkansas, Special Concern (S3) in Kentucky, and SI (critically imperiled in Alabama, Georgia, Mississippi, Tennessee, and Virginia (NatureServe 2008). Although there ) is known only one county occurrence only where of in Florida (the state it still is treated as a variety of S. seri- among ceum, pratense currently not listed that rare species (Florida Natural Areas Inventory 2008). S. is state's Jones et Distribution of Symphyotrichum sericeum and pratense al., S. Taxa.— Morphological Criteria for Distinguishing the Our examination specimens from th of much two out of the ranges of the taxa indicates that they can be reliably distinguished as follows: 1/4-1/3(-1/2) the total phyllary length, jsually distinctly shorter i < middle portion; phyllaries mostly 7 te mm Omm mm) < ' wide; involucral height mostly (7-11 high 3) I Symphyotrich ui sericeous itrigose (rarely he phyllary length, total mm > > llabrate portion; middle phyllaries mostly 8 long and 2.5 Symphyotrichum and and They pratense sericeum are distinct species of central eastern U.S. are largely S. al- common and lopatric in distribution, occur sympatrically only in Texas, where pratense and sericeum S. is S. Symphyotrichum with provinces— pratense associated range physiographic rare. a of chiefly Coastal is is it Low Plain in distribution, but its range extends to the Interior Plateaus, the Appalachian Plateau, and the and Valley Ridge. There an historical record from the Blue Ridge of North Carolina. Symphyotrichum is seri- ceum with Lowlands and Ozark and is largely associated the Central Province the Plateau, only sporadically into adjacent provinces to the south, north, and west. Disjunct populations and population systems are a prominent some feature of the biology of both taxa. There intermediacy in phyllary pubescence in the is easternmost disjunct populations, but these features could be interpreted as resulting from local genetic drift in populations far removed from the main range of the species, and there no evidence of any significant is gene interchange between the taxa. The available evidence indicates that sericeum consists of diploid 5. populations (with the exception of one anomalous report) and that pratense contains populations both S. at common much ploidy levels. Symphyotrichum sericeum is generally throughout of range, considered rare its only in the states at the eastern and southern periphery of range (Michigan, Indiana, and Arkansas), and its Nationally Threatened Canada. Outside Texas and and is in of Louisiana, pratense a rare plant con- 5. is is sidered critically imperiled in several states (Alabama, Georgia, Mississippi, Tennessee, and Virginia). The — been some identity of these disjunct eastern populations has the source of confusion in the past they have been variously treated by the different states as S. pratense or S. sericeum, and this latter name continues to be used by some Our eastern states for their rare species study concludes that specimens examined lists. all from populations in Alabama, Florida, Georgia, Kentucky, Mississippi, North Carolina, Tennessee, and We MO, TENN, TEX-LL and are grateful to staff at for help during visits there and to the curators and staff Tom we of all herbaria that provided loans. In particular, thank Wieboldt for a loan of specimens from VPI, Townsend Johnny (Natural Heritage Program, Virginia Dept. of Conservation and Recreation) information for GMUF on Virginia Andrea Weeks an image Wise County collections, at for of the (Virginia) collection of PH Symphyotrichum Alina an image pratense, Freire-Fierro of for of the holotype of Aster montanus, Jacquelyn NY Thomas Kallunki, Zanoni, and Lucy Klebieko of for providing a virtual herbarium loan of the speci- men from North Carolina, and John Pruski (MO) help with obtaining Brent Baker (UARK) for literature. provided distribution data for specimens compiled by the Arkansas Vascular Flora Deborah White Project, McCook provided from Kentucky Commission data the State Nature Preserves database, Lucile provided Wood information on Mississippi populations, Emily and Walter Kittredge specimen searches GH, for at Wendy and Zomlefer and David Giannasi information on Georgia we thank John for records. In addition, Semple chromosome on for his assistance in the gathering of information counts. Journal of the Botanical Research Institute of Texas 2(1) REFERENCES and Alabama. JA presence of the Black Belt of Mississippi 2005. ant distribution prairies in Barone, istorical Castanea 70:170-183. GA America North and 2006. Symphyotrichum. Flora of Allen, K.L Chambers, S.D. Sundberg. In: L, Semple, Brouillet, J.C. New 2+ Oxford North America North of Mexico. Univ. Press, Committee, 993+. of vols. eds. Flora Editorial 1 1 and 20:465-539. York Oxford. Vol. .Wisconsin and savanna Tech. No. 191 and 2000. of the Wisconsin prairie flora. Bull. Cochrane, H.H. Atlas T.S. Iltis. Madison. Wisconsin-Madison Herbarium, and Department Resources the University of of Natural and M.C Manual of the vascular plants ofTexas. Texas Research Foundation, Renner, Johnston. 970. Correll, D.S. 1 Castanea 6:228-267. Hardin County, Kentucky. 991 of Flora Cranfill, R. 1 . North Carolina Asteraceae. 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