Page  113 ï~~2005 THE MICHIGAN BOTANIST 113 THE DISTRIBUTION OF CHLORIS VERTICILLATA (TUMBLE WINDMILL GRASS, POACEAE) IN OHIO Michael A. Vincent W. S. Turrell Herbarium (MU), Department of Botany, Miami University, Oxford, Ohio 45056 USA, 513. 529. 2755, Chloris verticillata (Tumble windmill grass, Poaceae, Fig. 1) was first described by Thomas Nuttall from material collected in Arkansas (Nuttall 1835). Its original range appears to have been in the prairies from Louisiana west to Arizona, north to Colorado and South Dakota, and south through Iowa, Missouri and Arkansas (Barkworth 2003; Great Plains Flora Association 1986). The species has spread west into Nevada and California, and east to Illinois, Indiana, Michigan, Ohio, Kentucky, Tennessee, Pennsylvania, New York, Maryland, Rhode Island, Virginia, South Carolina, Connecticut, Massachusetts, and New Jersey (Barkworth 2003; Kartesz and Meacham 1999). It has also been reported from Delaware (McAvoy 2004) and Wyoming (Wyoming Native Plant Society 1982). Tumble windmill grass (also known as windmill grass and finger windmill grass) is a tufted perennial species growing from 10 to 40 cm tall. Its stems are erect to decumbent, often strongly jointed at the nodes, with flattened sheaths, and often rooted at the nodes. The leaves are often concentrated at the base of the stem, with flat blades ranging from 4 to 15 cm in length and 2 to 4 mm wide. Inflorescences of the species are purple to pinkish, when young consisting of upwardly pointed branches emerging from a folded leaf at the tip of the stem. As the inflorescence emerges, the branches flatten out to a verticillate arrangement of 2 to 4 or more whorls of branches, giving the species its characteristic appearance (Fig. la). The inflorescence measures from 15 to 28 cm in width at maturity, and up to 15 cm from the lowest whorl of branches to the upper tip. The spikelets (Fig. lb), composed of 2 glumes and 1 fertile (lower) floret and 1 sterile (upper) floret, measure about 3 mm in length, with an awn up to 10 mm in length at the tip of each lemma. As the fruits mature, the inflorescence fades to straw-colored or slightly pink, often breaking loose from the plant and being blown around by wind. The spread of Chloris verticillata from its historical range appears to have several causes. First, the species may have expanded its range naturally due to the wind-borne nature of its inflorescence, which can be blown around quite effectively by strong winds (personal observation). Secondly, the interstate highway system may play a role in its distribution, as may rail lines. The species may be weedy or invasive in at least some regions of the United States (e.g., Kansas, problem weed in lawns [Haddock 2004]; Michigan, considered a wide-spread non-native species [Michigan Association of Conservation Districts 2004]; Nebraska, considered invasive [Stubbendieck et al. 1994]; US Forest Service East

Page  114 ï~~114 THE MICHIGAN BOTANIST Vol. 44 a. I f FIGURE 1. Chloris verticillata (Vincent 7993, MU). a. Inflorescence (bar = 2cm). b. Closeup of spikelets (bar = 1.3cm).

Page  115 ï~~2005 THE MICHIGAN BOTANIST 115 ern Region, considered a wide-spread non-native species [USFS 2005]), and is listed in a compendium of world weeds (Randall 2002). In no-till agricultural fields in its native range, tumble windmill grass can become a significant problem after a few cycles of crop rotations (Anderson 2004; Hagny 2003). It is also available through horticultural sources (e.g., Bluestem Nursery 2005), and thus may spread from cultivation. While this species of grass may be included in some lawn grass seed mixtures, I was unable to find any documentation to verify it. The species may serve as a non-maize reservoir for Western Corn Rootworm (Wilson and Hibbard 2004). The grass is unpalatable to grazing cattle (Haddock 2004). The grass has been utilized by barn swallows as a component of nests (Whelan 2003). Confusion abounds in the literature on the flora of Ohio regarding the presence and distribution of Chloris verticillata in the state. The first published report of Chloris verticillata in Ohio was that of Weishaupt (1985), in her key to grasses in the vegetative condition, where she lists the grass as found in three southwestern counties. The species was also included in an addendum (on page 293) in the back of the eighth printing of the third edition of Weishaupt's Vascular Plants of Ohio (Weishaupt 1987), and confusion about its date of publication makes it appear that the first report was for 1971, which is the date of the first printing of that edition; the actual printing date of the eighth printing of the third edition was April 1987 (Kendall-Hunt Publishing, pers. comm. 15 June 2004). While the range of the species includes Ohio in Barkworth (2003), the species is mapped for only one county in the central portion of the state. Chloris verticillata is also listed as present in Ohio by Kartesz and Meacham (1999). Cooperrider et al. (2001) do not list the species as part of the Ohio flora in their Seventh Catalogue. The species is not listed as occurring in Ohio in any of the following publications: Schaffner (1917, 1928), Hitchcock (1935), Hitchcock and Chase (1950), Fernald (1950), Pohl (1954), Weishaupt (1967), Anderson (1974), Gleason and Cronquist (1991). Indeed, in Anderson's 1974 monograph of the genus, he neither mentions nor maps any specimens east of central Iowa. It is also apparent that specimens from sites west of New Mexico and Colorado were unknown to him. Since several Ohio populations of tumble windmill grass are known to me, I decided to investigate further and see the extent to which it occurs in the state. Specimens were examined from the following herbaria: CM, F, ISC, KE, KNK, MICH, MO, MU, OS. A total of 31 sheets of Chloris verticillata were examined for the study, representing 17 separate collections (see Appendix 1). Populations of the species were documented from nine Ohio counties: Butler, Clark, Franklin, Greene, Hamilton, Miami, Montgomery, Pickaway, and Sandusky. The grass was found in lawns, especially along roads and railroad tracks. Chloris verticillata does not seem to be an invasive species in Ohio. Populations examined appeared to be relatively small and restricted. The lawns in which the species was found contained mixtures of several lawn grasses, forbs, and weeds, and were not particularly negatively impacted by the presence of the species. Indeed, the only time the grass appears anything out of the ordinary is at flowering, when the reddish inflorescences project above the leafy portions of

Page  116 ï~~116 THE MICHIGAN BOTANIST Vol. 44 the plants. Since several of the populations have persisted for decades, it appears that the grass will remain a part of Ohio's weedy flora, and will perhaps spread slowly in the state. ACKNOWLEDGMENTS I wish to thank the following herbaria for access to or information about specimens: CM, F, ISC, KE, KNK, MICH, MO, MU, OS. Thanks to Allison W. Cusick, who provided information pertinent to the study. LITERATURE CITED Anderson, D. E. 1974. Taxonomy of the genus Chloris (Gramineae). Brigham Young University Science Bulletin, Biological Series, 19(2): 1-133. Anderson, R. L. 2004. Impact of subsurface tillage on weed dynamics in the central Great Plains. Weed Technology 18: 186-192. Barkworth, M. E. 2003. Chloris. Pp. 204-218. In: Barkworth, M. E., K. M. Capels, S. Long, and M. B. Piep (eds.). Magnoliophyta: Commelinidae (in part): Poaceae, part 2. Flora of North America North of Mexico, volume 25. Oxford University Press, New York, NY. Bluestem Nursery. 2005. Full-sun to part-shade native grasses needing or tolerating irrigation. (accessed 26 Jan 2005). Cooperrider, T. S., A. W. Cusick, and J. T. Kartesz. 2001. Seventh Catalog of the Vascular Plants of Ohio. Ohio State University Press, Columbus, OH. Fernald, M. L. 1950. Gray's Manual of Botany (ed. 8). American Book Company, New York, NY. Gleason, H. A. and A. Cronquist. 1991. Manual of Vascular Plants of Northeastern United States and adjacent Canada. New York Botanical Garden, Bronx, NY. Great Plains Flora Association. 1986. Flora of the Great Plains. University Press of Kansas, Lawrence, KS. Haddock, M. 2004. Kansas wildflowers and grasses: windmill grass. wildflowers/windmillgrass.html (accessed 1 Feb 2005). Hagny, M. 2003. Leveraging biology: discovering efficiencies in cropping systems. Pp. 34-41. 2003 No-till Under Cover Proceedings. South Dakota No-till Association, Pierre, SD. http://www. (accessed 28 Jan 2005). Hitchcock, A. S. 1935. Manual of the Grasses of the United States. USDA Miscellaneous Publication 200, US Government Printing Office, Washington, DC. Hitchcock, A. S. and A. Chase. 1950. Manual of the Grasses of the United States (ed. 2). USDA Miscellaneous Publication 200, US Government Printing Office, Washington, DC. Kartesz, J. T., and C. A. Meacham. 1999. Synthesis of the North American Flora, Version 1.0. North Carolina Botanical Garden, Chapel Hill, NC. McAvoy, W. 2004. Non-native plants of Delaware. Unpublished report. Delaware Natural Heritage Program, Delaware Division of Fish and Wildlife, DNREC, Smyrna, DE. http://www. (accessed 28 Jan 2005). Michigan Association of Conservation Districts. 2004. Invasive plants. pdfs/AppC.pdf (accessed 28 Jan 2005). Nuttall, T. 1835. Transactions of the American Philosophical Society, new series, 5:150 Pohl, R. W. 1954. How to Know the Grasses. W. C. Brown, Dubuque, IA. Randall, R. P. 2002. A Global Compendium of Weeds. R. G. and F. J.Richardson, Meredith, Victoria, Australia. Schaffner, J. H. 1917. The Grasses of Ohio. Ohio Biological Survey Bulletin 9 Schaffner, J. H. 1928. Field Manual of the Flora of Ohio and adjacent Territory. R. G. Adams, Columbus, OH. Stubbendieck, J., G. Y. Friscoe, and M. R. Bolick. 1994. Weeds of Nebraska and the Great Plains. Nebraska Department of Agriculture, Bureau of Plant Industry. Lincoln, NE. USFS (United States Forest Service). 2005. Eastern Region invasive plants, ranked by degree of in

Page  117 ï~~2005 THE MICHIGAN BOTANIST 117 vasiveness as based on information from states. weed/Sec3B.htm (accessed 26 Jan 2005). Weishaupt, C. G. 1967. Gramineae. Pp. 61-174. In: Braun, E.L. The Monocotyledonae: Cat-tails to Orchids. The Vascular Flora of Ohio, Volume 1. Ohio State University Press, Columbus, OH. Weishaupt, C. G. 1985. A descriptive key to the grasses of Ohio based on vegetative characters. Ohio Biological Survey Bulletin New Series 7(1). Weishaupt, C. G. 1987. Vascular Plants of Ohio (ed. 3, 8th printing). Kendall/Hunt Publishing Co., Dubuque, IA. Whelan, E. 2003. Composition and structure of barn swallow nests. Transactions of the Oklahoma Junior Academy of Science 30: 1-5. (accessed 28 Jan 2005). Wilson, T. A. and B. E. Hibbard. 2004. Host suitability of nonmaize agroecosystem grasses for the western corn rootwom (Coleoptera: Chrysomelidae). Environmental Entomology 33: 1102-1108. Wyoming Native Plant Society. 1982. New state records (Chloris verticillata). Castilleja Newsletter. October 1982: 2. APPENDIX 1. SPECIMENS EXAMINED OHIO. Butler County: Oxford, tree lawn on south side of Spring Street just east of the railroad tracks, 23 Aug 1997, M. A. Vincent 7993 (F, ISC, KNK, MICH, MO, MU, OS); Oxford, vacant lot west of the railroad, Spring Street, 28 Jun 1982, K. W Dougherty 682-30 (MU); ibid, 6 Jul 1982, K. W Dougherty 782-01 (MU). Clark County: Springfield, grassy area, corner of Greenmont and Highland, Aug/Sep 1975, C.G. Weishaupt s.n. (OS). Franklin County: Columbus, lawns at Ohio Woolgrowers warehouse, 1 Sep 1986, A. W Cusick 25795 (KE, MICH, OS); ibid, 11 Sep 1986, A. WCusick 25866 (CM, MU); Columbus, Greenlawn Cemetery, A. W Cusick 34040 (CM). Greene County: Fairborn, yard on Central Ave., Sep 1974, C.G. Weishaupt s.n. (OS); Fairborn, along railroad, Kauffman Ave., 2 Sep 1975, C. G. Weishaupt s.n. (OS); Fairborn, along railroad, Central Ave., 15 Aug 1980, C. G. Weishaupt s.n. (OS). Hamilton County: Cincinnati, lawn, Eden Park, 24 Jul 1974, J. H. Carpenter 111 (KNK, OS); Cincinnati, Spring Grove Ave., 24 Aug 1991, J. W Thieret 57168 (KE); Wyoming, county fairgrounds, 11 Sep 1996 A. W Cusick 33408 (CM, MICH, OS). Miami County: Troy, grassy area along Great Miami River at Troy Community Park, Main Street, 5 Sep 2002, M. A. Vincent 10867 (MU). Montgomery County: Dayton, along railroad tracks, 20 Aug 1979, W R. Carr 2078 (KE, OS). Pickaway County: South Bloomfield, US Rt 23 & Main St., tree lawn, 25 Sep 2003, A. W Cusick 36147 (MU, OS). Sandusky County: Madison Twp., North Union Cemetery, 26 Sep 2000, A. W Cusick 35666 (MU).