The Biology of Canadian Weeds. 141. Setaria faberi Herrm.
Robert E. Nurse1, Stephen J. Darbyshire2,Ce´cile Bertin3, and Antonio DiTommaso4
1Agriculture and Agri-Food Canada, Greenhouse and Crop Processing Centre, 2585 County Road 20, Harrow, Ontario, Canada N0R 1G0 (e-mail: [email protected]); 2Agriculture and Agri-Food Canada, Central Experimental Farm, Saunders Building #49, Ottawa, Ontario, Canada K1A 0C6; 3Department of Horticulture, Cornell University, Ithaca, NY, USA 14853; and 4Department of Crop and Soil Sciences, Cornell University, Ithaca, NY, USA 14853. Received 17 March 2008, accepted 13 November 2008.
Nurse, R. E., Darbyshire, S. J., Bertin, C. and DiTommaso, A. 2009. The Biology of Canadian Weeds. 141. Setaria faberi Herrm. Can. J. Plant Sci. 89: 379 404. Setaria faberi, commonly known as giant foxtail, is an annual graminaceous weed that is native to eastern China, has colonized eastern North America and is expanding its range westward. This species is primarily self-pollinated and the only mechanism of reproduction is by seed. Adult plants may reach 2 m in height and produce over 2000 seeds per panicle. Seeds may possess non-deep physiological dormancy when freshly produced, and can form small persistent seed banks. If not controlled, S. faberi populations can cause severe yield reductions in corn and soybean crops. Several herbicides are available to provide chemical control; however, resistance to some modes of action, (ALS, ACCase, and Photosystem II) have been identified in Canada and the United States. Leaves and seeds of this species provide a food source to several species of mammals, birds, and insects.
Key words: Setaria faberi, giant foxtail, growth, development, seed germination, diseases, herbicide
Nurse, R. E., Darbyshire, S. J., Bertin, C. et DiTommaso, A. 2009. La biologie des mauvaises herbes au Canada. 141. Setaria faberi Herrm. Can. J. Plant Sci. 89: 379 404. La se´ taire ge´ ante (Setaria faberi) est une gramine´ e annuelle originaire de l’est de la Chine. Apre` s avoir colonise´ l’est de l’Ame´ rique du Nord, elle poursuit son expansion vers l’ouest. Essentiellement autogame, l’espe` ce n’a que les graines pour moyen de reproduction. Le plant adulte peut atteindre jusqu’a` 2me` tres de hauteur et chaque panicule produit au-dela` de 2 000 semences. Fraıˆ ches, les semences entrent en dormance a` faible profondeur et peuvent engendrer de petites re´ serves de graines persistantes. Si on ne lutte pas contre eux, les peuplements de S. faberi entraıˆ nent a` l’occasion de fortes diminutions de rendement chez le maı¨ s et le soja. Bien que plusieurs herbicides offrent un moyen de lutte chimique, au Canada et aux E´ tats-Unis, la plante re´ siste a` certains de leurs modes d’action (ALS, ACCase et Photosystem II). Plusieurs espe` ces de mammife` res, d’oiseaux et d’insectes vivent des feuilles et des graines de la se´ taire ge´ ante.
Mots cle´s: Setaria faberi,se´ taire ge´ ante, croissance, de´ veloppement, germination, maladies, herbicides
1. Name at the tip, usually finely pubescent with soft straight Setaria faberi R. A. W. Herrm. * Incorrect spelling: hairs on the upper surface or both surfaces (see also Setaria faberii * giant foxtail, Chinese foxtail, Chinese Section 2c). The inflorescence is a dense spike-like millet, giant bristle grass, Japanese bristlegrass, nodding panicle, usually nodding or drooping from near the foxtail; se´ taire ge´ ante; se´ taire de Faber (Rominger 1962; base and 6 20 cm long. The inflorescence rachis is Darbyshire 2003; USDA, NRCS 2007). Poaceae (Gra- densely villose with long whitish hairs and the spikelets mineae), grass family, Gramine´ es. European and Med- are densely grouped in a green, soft bristly, elongated iterranean Plant Protection Organization Code (Bayer head, which resembles a slender bottle brush. On the Code): SETFA. pedicels at the base of each spikelet are (1) 3 (6) bristles about 10 mm long, which remain on the plant after the 2. Description and Account of Variation spikelet has fallen from its minute cup-like base. The (a) Species Description * The following description is spikelets are 2.5 3 mm long; the lower glumes are about based on published information (Hitchcock 1950; Fair- 1 mm long and acute; the upper glumes about 2.2 mm brothers 1959; Pohl 1962; Rominger 1962), supplemen- long, 2/3 to 5/6 as long as the upper fertile lemma and ted with measurements from herbarium specimens made obtuse; the lower sterile lemmas are about 2.8 mm long; by the authors. Measurements are given as the typical the lower paleas are about 2/3 as long as the lower range with unusual extremes in parentheses. lemmas; the upper lemmas are pale greenish or yellow- Plants are annual. Stems single or branching near the ish, finely to distinctly transversely wrinkled (rugose); base, (8) 50 200 (250) cm high; leaf sheaths are smooth, and, the upper palea is similar to upper lemma. The with the upper margins hairy. The ligule is a fringe of exserted anthers are about 1 mm long. short hairs about 2 mm long. Leaf blades are flat, 15 30 Chromosome counts reported from Canadian popu- cm long and 10 20 mm wide, narrowing to a fine point lations are 2n 36 (Warwick et al. 1987, 1997). The 379 380 CANADIAN JOURNAL OF PLANT SCIENCE same tetraploid number has also been reported from the of S. faberi and S. viridis. For photographiccompar- United States (Fairbrothers 1959; Pohl 1962), Japan isons among S. pumila, S. viridis, S. faberi, and S. (Kishimoto 1938; Tateoka 1955) and Russia (Probatova verticillata see Bouchard and Ne´ ron (1991). and Sokolovskaya 1983). 3. Economic Importance (b) Similar Species and Distinguishing Features * Five (a) Detrimental * Considered a major weed worldwide, species of Setaria have been reported in Canada. A S. faberi is an annual grass found in a wide range of comparison of some morphological characteristics is cropping systems, but causes the most economic losses given in Table 1 and a key to the species is given in in field corn (Zea mays L.), and soybean [Glycine max Appendix 1. Setaria faberi can be distinguished from (L.) Merr.] and is responsible for crop yield reductions, other grasses by the presence of a hairy ligule and hairs increased cleaning costs, and the necessity for imple- on the upper side and margins of the leaf blade (see also mentation of expensive cultural and chemical control section 2a). The inflorescence is characteristic of the measures (Knake 1977; Zimdahl 1980). The detrimental foxtail family and often droops once mature (Fig. 1). effects of S. faberi also include production of secondary allelopathic chemicals, serving as a secondary host for (c) Intraspecific Variation * No intraspecific taxa have insect pests, and the ability to survive in a persistent seed been described. However, variation in the leaf blade bank (see also section 8c). pubescence has been reported. Forms with both leaf Setaria faberi is often found in mixed communities blade surfaces pubescent as well as non-hairy (glabrous) with other Setaria species such as S. pumila, and have been reported from within populations of typical S. viridis. Of the three species S. faberi is considered plants (Monachino 1959; Pohl 1962) (See also Section the most competitive species due to its adaptability to 7c). Fairbrothers (1959) reported the morphological diverse environments, delayed germination and high variation found in populations from New Jersey, mea- fecundity (Santelman et al. 1963; Schreiber 1965; Knake suring 150 plants (12 populations) grown in the field and 1977; Buhler et al. 1997; Moechnig et al. 2003). 20 plants (10 populations) grown in the greenhouse. Generally, comparisons among species for effects on Wang et al. (1995) reported that S. faberi is nearly crop yield are hard to distinguish, because accounts of monomorphic as a species, which is remarkable crop yield losses are often reported for mixed stands of given the geographically diverse sampling used for their two or more of the Setaria species. Although, in a 3-yr study, which included accessions from at least three study, Staniforth (1965) demonstrated that the vigorous different continents. Strong inbreeding processes (cleis- growth of S. faberi caused greater soybean yield togamy or self pollination) occur, because little hetero- reductions than did S. pumila or S. viridis. Moechnig zygosity was detected (Wang et al. 1995). Like S. pumila et al. (2003) developed empirical models describing the (Poir.) Roem. & Schult. ( S. glauca of authors) and competitiveness of S. faberi in relation to Chenopodium S. parviflora (Poir.) Kergue´ len ( S. geniculata album L. in corn. Competition coefficients that were P. Beauv.), S. faberi, populations are largely homoge- based on relative leaf area revealed that S. faberi was nous, but are differentiated from one another in the more competitive than C. album. In other mixed fixation of different alleles at one or more loci (Wang communities such as old-field successions, S. faberi is et al. 1995). When Benabdelmouna et al. (2001a) often ranked as the most competitive graminaceous performed an analysis of phylogeneticand genomic weed (Wakefield and Barrett 1979; Parrish and Bazzaz relationships among species in the genus Setaria they 1982; Facelli and Pickett 1991b). found that S. faberi rDNA contained an additional 450 bp that were not present in other species within the genus. Corn and Soybean * The presence of S. faberi at low (d) Illustrations * The habit of S. faberi is illustrated in population densities within a corn crop is sufficient to Fig. 1 and details of the spikelets are illustrated in Fig. 2. cause significant yield losses. For example, S. faberi Pohl (1962) provided illustrations to contrast the seeds caused a 24% reduction of corn yield when present in
Table 1. Comparison of Setaria species in Canada
Character S. italica S. faberi S. pumila S. viridis S. verticillata
Culms (cm) 100 250 50 200 30 130 20 250 30 100 Ligules (mm) 1 22 9 11 2to1 Blades (mm wide) 10 30 10 20 4 10 4 25 5 15 Panicle (cm) 8 30 6 20 3 15 3 20 5 15 Rachis hispid to villose densely villose hispid hispid to villose retrorsely hispid Bristles (mm) to 12 9 10 3 85 10 4 7 Spikelets (mm) 9 3 2.5 3 (2)3 3.4 1.8 2.2 2 2.3 Upper glume to lower lemma 9 3/4 9 2/3 9 1/2 9 9 NURSE ET AL. * SETARIA FABERI HERRM. 381
Fig. 1. Setaria faberi habit. Inset: upper (adaxial) surface of leaf blade. the interrow, and 37% reductions when between the Conley et al. (2003a, b) quantified the relative rows (Donald and Johnson 2003). In a 3-yr study in competitive ability of S. faberi in soybean and concluded Illinois, 17 S. faberi plants m 1 of crop row were that the critical period of S faberi removal was at the V2 responsible for a 25% reduction in corn yield (Knake leaf stage, and that if S. faberi was present at popula- and Slife 1961). Also in Illinois, four S. faberi plants tions of 64 plants m 2 then soybean yield may be m 2 reduced soybean yield 5%, and 200 plants m 2 reduced by up to 86%. Mulugeta and Boerboom (2000) reduced soybean yield by 30% (Knake 1990). Knake also demonstrated that the critical period of S. faberia (1987) generalized that each kilogram of S. faberi removal in soybeans was between the V2 V4 stages. aboveground biomass results in a 1 kg decrease in Harrison et al. (1985) described a soybean yield reduc- crop yields. Lindquist et al. (1999) developed a rectan- tion of 26% when S. faberi was present at 10 plants m 1 gular hyperbola-based model to predict the threshold of soybean row in Illinois. Even at these lower densities density of foxtail species (predominately S. faberi)in (B 16 plants m 2), S. faberi individuals may produce corn and concluded that while variable by year, the up to 28 000 seeds plant 1 (Conley et al. 2003a). economic threshold was 3 to 4 foxtail plants m 1 of Seeding soybeans in narrow rows (38 cm) versus wide row. In Michigan, a hyperbolic equation described up to rows (76 cm) may reduce foxtail biomass and maximize a 14% reduction in corn yield from the presence of 10 S. soybean yield; however, this strategy does not reduce the faberi plants m 1 of row (Fausey et al. 1997). total number of seeds produced (Schmidt and Johnson 382 CANADIAN JOURNAL OF PLANT SCIENCE
Fig. 2. Setaria faberi: A. Inflorescence branch; B. Spikelet, adaxial view showing second glume and fertile lemma; C. Fertile floret, abaxial view showing fertile lemma; D. Fertile floret, adaxial view showing fertile palea and lemma margins.
2004). This may be problematic, because after 35 yr of of lucerne (Medicago sativa L.) and/or triticale (Tritico- continuous crop rotation and tillage, S. faberi seed secale sp.) into a rotation with corn and soybeans banks were found to be highest (6890 seeds m 2)of effectively reduced field populations of S. faberi in several weed species in a no-till corn and soybean Iowa (Heggenstaller and Liebman 2005). rotation in Ohio (Cardina et al. 2002). Setaria faberi is also a problem in container-grown Differences in germination timing, growth and com- ornamentals. For example, the growth and development petitiveness of S. faberi cause the species to be less of a of the container-grown bush cinquefoil (Potentilla problem in a 3-yr corn soybean winter wheat rotation fruticosa L.) were negatively affected when grown with than in a corn soybean or a continuous corn cropping graminaceous weeds, such as S. faberi (Walker and system (Staniforth 1965; Schreiber 1992). The addition Williams 1989). Pohl (1951) also reported that it was a NURSE ET AL. * SETARIA FABERI HERRM. 383 common weed in Iowa gardens. Seeds of S. faberi are 4. Geographical Distribution also frequently a problem as contaminants of seed Although S. faberi is native to eastern Asia (Wang et al. commodities such as millet (Chase 1948) and clover 1995), its range has greatly expanded as a result of (Pohl 1951). See also Section 3c. global trade and human-mediated dispersal (Haflinger There is quantitative evidence that S. faberi may and Scholz 1980). It occurs as an introduced species in produce secondary alleopathic chemicals that may eastern Canada in Southern Ontario and Quebec(Fig. impact the growth of other species. For example, 3). Simultaneously, the species also colonized most of leachates from live and desiccated S. faberi tissue caused the eastern United States as well as California and up to a 36% reduction in loblolly pine (Pinus taeda L.) Arizona (USDA, NRCS. 2007). Elsewhere it is reported seedling growth (Gilmore 1980). Using similar metho- as a non-native weed in central Europe, Russia, the dology, Keoppe and Bell (1972) demonstrated that S. Middle East, and eastern Asia (Wang et al. 1995). The faberi exudates from mature plants impacted corn northern limit of its distribution in North America is growth by up to 40% during the first month after about 46.58N latitude in Canada (Warwick et al. 1987; crop emergence. herbarium specimen data). Setaria faberi may also serve as a potential host for several detrimental insects. In greenhouse studies, the 5. Habitat western corn rootworm [Diabrotica virgifera virgifera (a) Climatic Requirements * This species has a wide LeConte (Coleoptera: Chrysomelidae)] had good survi- distribution, suggesting that S. faberi has adapted to a vorship on S. faberi (Chege et al. 2005; Oyediran et al. broad range of environmental conditions. In the north- 2005). Clark and Hibbard (2004) tested 28 species of ern Canadian range of S. faberi, seeds experience grasses for non-corn host suitability for western corn January temperatures of 17.68C, and plants experi- rootworm and S. faberi ranked 13th out of the 28 species, ence July/August maximum temperatures of 27.98C but was shown to be a less suitable host than S. viridis.In (Table 2). In the same area precipitation ranges from a corn fields, S. faberi can also be a suitable host for the low of 42.6 mm to a high of 127.8 mm during the Northern Corn Rootworm [Diabrotica barberi Smith & growing season (Table 2). In the United States, the temperature extremes during the calendar year are Lawrence (Coleptera: Chrysomelidae)] (Oyediran et al. 15.48C (low) and 40.18C (high), and the precipitation 2004). extremes are 1.3 mm (low) and 204.2 mm (high) (Table 2). (b) Beneficial * Willson and Harmeson (1973) found that when presented with a choice of seeds from six (b) Substratum * A wide range of soils support the different weed species, cardinals and song sparrows germination and growth of this species. In Quebec, S. preferred to consume seeds of S. faberi. The seeds faberi seeds were found in soil seed banks of the clay of this species may also be a valuable food source for loam soil series: St. Urbain, Du Jour, Ste. Rosalie, and other granivorous birds and rodents and, when St. Blaise (fine, mixed, mesic, Typic Humaquept) found near wetland areas, S. faberi seeds may be (Vanasse and Leroux 2000). The occurrence of S. faberi consumed by various species of waterfowl (Anonymous has been documented on soils ranging from sandy loam 2003). (BrunisolicGray Brown Luvisol; 83% sand, 5% silt, As a pioneer species of open disturbed soils, S. faberi 13% clay, 2.6% organic matter and pH 6.4) to clay loam is sometimes useful for soil stabilization. In old-field (OrthicHumicGleysol, mixed, mesic,and poorly communities where sludge has been discarded by drained; 30% sand, 40% sand, 30% clay, 4.5% organic municipalities, S. faberi has been shown to readily matter, and pH 7.8) in southwestern Ontario (R.E. uptake and sequester Cd, Cu, Pb, and Zn in roots, Nurse, personal observations). In Ohio, S. faberi shoots, leaves and seeds (Levine et al. 1989; Peles et al. populations were established on a silt loam (fine, mixed, 1998). When grown in mixed communities with grass mesic, Typic Fragiaqualf) that was 11% sand, 75% silt, species that are used for prairie restoration, low densities 14% clay and 1.9% organic matter (Cardina et al. 2007). of S. faberi have been shown to be beneficial as a nurse Populations of S. faberi in Michigan were also found in species (Endress et al. 1999). fields with silt loam soils (43% sand, 40% silt, 17% clay, 1.1% organicmatter and pH 6.7) (Davis and Renner (c) Legislation * In Canada, S. faberi is not currently 2007). listed as a noxious or invasive weed in any province or territory (Agriculture Canada 1986; Darbyshire 2003), (c) Communities in Which the Species Occurs * Setaria although it is currently listed as a prohibited noxious faberi thrives in a wide range of communities through- weed, Class 1, in the Weed Seeds Order, 2005, under the out Ontario, Quebec, and the United States, including Canada Seeds Act. Presently in the United States S. cultivated fields, gardens, railway lines, roadsides, and faberi has been documented in at least thirty-five states disturbed areas (Darbyshire 2003). and is currently considered a noxious weed in AR, CA In old fields, S. faberi is an early successional species, and KY (USDA, NRCS 2007). typically occurring in fields that are less than 4 5 yr old. 384 CANADIAN JOURNAL OF PLANT SCIENCE
Fig. 3. Distribution of Setaria faberi in Canada. Locations based on 257 herbarium specimens from the following herbaria: CAN, DAO, HAM, MT, QFA, QUE, TRT, TRTE, and UWO [abbreviations according to Holmgren et al. (1990)].
In a 4-yr-old Ohio abandoned field undergoing second- carolinense (Hunt et al. 1979; Facelli and Pickett 1991a). ary succession, S. faberi was found growing with Festuca Raynal and Bazzaz (1975) found S. faberi in abandoned elatior L., Poa pratensis L., Solidago canadensis L., Illinois fields with Ambrosia artemisiifolia, Polygonum Ambrosia artemisiifolia L., Barbarea vulgaris R. Br., pensylvanicum L., Abutilon theophrasti,andChenopo- Oenothera biennis L., Phalaris arundinacea L., Melilotus dium album. Although rarely found in more advanced officinalis L. (Pall.), and Phleum pratense L. (Sedlacek et successional systems, Maly and Barrett (1984) reported al. 1988). In a 1-yr-old successional field in Illinois, S. S. faberi, Ambrosia artemisiifolia, and A. trifida L. to be faberi co-occurred with Abutilon theophrasti Medic., and present in a 7-yr-old old field that had been supple- Polygonum pensylvanicum L. (Wieland and Bazzaz mented with fertilizer or sludge. In Ohio, S. faberi 1975). Setaria faberi was also found in a first-year old dominated successional corridors. The successional field in Indiana with Abutilon theophrasti, Acalypha corridors were established by plowing and disking an virginica L., Ambrosia artemisiifolia, Asclepias syriaca area and then seeding it to Festuca spp., Poa pratensis, L., Chenopodium album, Cirsium arvense (L.) Scop., and Trifolium hybridum L. The corridors were then left Datura stramonium L., Daucus carota L., Digitaria undisturbed and allowed to enter secondary succession sanguinalis (L.) Scop., Hedeoma pulegioides (L.) Pers., for 10 yr. After this period S. faberi was found along Oxalis stricta L., Solidago canadensis, Solanum caroli- with Ambrosia artemisiifolia, A. trifida, Asclepias syr- nense L., Taraxacum officinale G.H. Weber ex Wiggers, iaca, Convolvulus arvensis L., and Polygonum persicaria Trifolium spp. and Triticum aestivum L. (Squiers 1989). L. (Kemp and Barrett 1989). In an Illinois successional old field grassland community In Ontario, S. faberi has been observed growing in S. faberi co-existed with Panicum anceps Michx., mixed stands within corn with Amaranthus retroflexus Cardamine hirsute L., Andropogon virginicus L., Sor- L., Amaranthus powellii S. Wats., Abutilon theophrasti, ghum halepense (L.) Pers., Festuca arundinacea Schreb., Ambrosia artemisiifolia, Setaria viridis (L.) Beauv., Tridens flavus (L.) A.S. Hitchc., Lespedeza cuneata Echinochloa crus-galli (L.) Beauv., Setaria pumila, Era- (Dum.-Cours.) G. Don., and the shrubs Elaeagnus grostis cilianensis (All.) E. Mosher, Panicum capillare L., umbellata Thunb., Rubus pensilvanicus Poir., and Rosa Panicum dichotomiflorum, Chenopodium album, Solanum multiflora Thunb. (Spyreas et al. 2001). In old-field ptychanthum Dun., and Polygonum spp. (Hamill et al. communities in New Jersey, S. faberi was the dominant 2004; R. Nurse, personal observations). Vanasse species and highly competitive with Ambrosia artemisii- and Leroux (2000) found S. faberi seeds in the soil folia, Calystegia sepium L., Digitaria sanguinalis, Oxalis seed bank of corn and soybean fields in Quebec stricta, Panicum dichotomiflorum Michx., and Solanum along with Abutilon theophrasti, Ambrosia artemisiifolia, Table 2. Maximum and minimum mean temperature and precipitation throughout the North American range of Setaria faberiz
Means (from monthly normals)
Data Daily max. Daily min. Max. total Min. total
Country City Prov/State Latitude Longitude Total years Temp. (8C) Month Temp. (8C) Month Precip. (mm) Month Precip. (mm) Month
CAN QuebecQC 46 848?N71823?W 25 25.0 Jul. 17.6 Jan. 127.8 Jul. 70.6 Feb. CAN Montreal QC 45828?N73845?W 30 26.2 Jul. 14.7 Jan. 92.7 Aug. 61.5 Feb. CAN Ottawa ON 45819?N75840?W 30 26.5 Jul. 15.3 Jan. 90.6 Jul. 58.9 Feb. CAN Kingston ON 44813?N76835?W 26 24.8 Jul. 12.2 Jan. 99.0 Dec. 58.8 Jul. CAN Toronto ON 43840?N79837?W 30 26.8 Jul. 10.5 Jan. 79.6 Aug. 42.6 Feb. CAN Niagara Falls ON 43808?N79805?W 25 27.2 Jul. 7.9 Jan. 95.2 Sep. 67.4 Feb. CAN Guelph ON 43833?N80813?W 21 25.9 Jul. 11.4 Jan. 95.9 Aug. 50.8 Feb. CAN London ON 43801?N81809?W 30 26.3 Jul. 10.1 Jan. 97.7 Sep. 60.0 Feb. CAN Windsor ON 42816?N82857?W 30 27.9 Jul. 8.1 Jan. 96.2 Sep. 57.3 Feb. US Little Rock AR 34845’N 92814?W 30 33.8 Jul. 0.7 Jan. 145.5 Nov. 74.4 Aug. US Phoenix AZ 33826?N 112800’W 30 40.1 Jul. 6.3 Jan. 27.2 Mar. 2.3 Jun. US Sacramento CA 38830?N 121830’W 30 33.6 Jul. 3.2 Dec. 97.5 Jan. 1.3 Jul. US Hartford CT 41856?N72841?W 30 29.4 Jul. 8.2 Jan. 111.5 May. 75.2 Feb. US Tallahassee FL 30824?N84821?W 30 33.3 Jul. 4.3 Jan. 204.2 Jul. 82.6 Oct. US Atlanta GA 33838?N84826?W 30 31.9 Jul. 0.8 Jan. 136.7 Mar. 79.0 Oct. US Des Moines IA 41832?N93840?W 30 30.0 Jul. 11.3 Jan. 116.1 Jun. 26.2 Jan. US Springfield IL 39851’N 89841?W 30 30.3 Jul. 8.3 Jan. 103.1 May. 41.1 Jan. US Indianapolis IN 39843?N86816??W 30 29.8 Jul. 7.5 Jan. 112.3 Jul. 61.2 Feb. US Topeka KS 39804?N95838?W 30 31.7 Jul. 8.2 Jan. 124.0 Jun. 24.1 Jan. US Frankfort KY 38814’N 84852??W 30 30.5 Jul. 6.2 Jan. 117.1 May. 78.2 Feb. US Baton Rouge LA 30832?N91809?W 30 32.7 Aug. 4.6 Jan. 157.2 Jan. 96.8 Oct. US Boston MA 42822?N71801?W 30 27.9 Jul. 5.5 Jan. 101.1 Nov. 77.7 Jul. US Baltimore MD/DC 39810?N76841?W 30 30.7 Jul. 4.7 Jan. 101.1 Sep. 76.2 Apr. US Lansing MI 42847?N84835?W 30 27.8 Jul. 10.1 Jan.. 91.4 Jun. 36.8 Feb. AL. ET NURSE US Minneapolis MN 44853?N93814’W 30 28.5 Jul. 15.4 Jan. 110.2 Jun. 20.1 Feb. US Jefferson City MO 38835?N92811?W 30 31.9 Jul. 7.9 Jan. 123.2 May. 41.9 Jan. US Jackson MS 32819?N90805?W 30 33.0 Jul. 1.7 Jan. 151.9 Apr. 82.0 Sep. US Raleigh NC 35852?N78847?W 30 31.7 Jul. 1.3 Jan. 109.0 Jul. 71.1 Apr. US Lincoln NE 40850?N96846?W 30 32.0 Jul. 11.4 Jan. 107.4 May. 16.8 Feb. US Concord NH 43812?N71830?W 30 28.3 Jul. 12.4 Jan. 90.7 Nov. 59.9 Feb.
US Newark NJ 40843?N74810?W 30 29.6 Jul. 4.2 Jan. 118.9 Jul. 75.2 Feb. * US Albany NY 42845?N73848?W 30 27.9 Jul. 10.4 Jan. 95.0 Jun. 57.7 Feb. US Wooster OH 40847?N81855?W 30 27.8 Jul. 8.4 Jan. 102.4 Aug. 50.0 Feb. 385 HERRM. FABERI SETARIA US Oklahoma City OK 35823?N97836?W 30 33.9 Jul. 3.2 Jan. 138.2 May. 32.5 Jan. US York PA 39855?N76845?W 30 30.3 Jul. 6.2 Jan. 109.5 Jun. 70.4 Feb. US Providence RI 41843?N71826?W 30 28.1 Jul. 6.5 Jan. 112.5 Mar. 80.5 Jul. US Columbia SC 33857’N 81807?W 30 33.4 Jul. 1.1 Jan. 140.7 Jul. 73.2 Nov. US Pierre SD 44823?N 100817?W 30 31.8 Jul. 13.5 Jan. 88.6 Jun. 12.2 Dec. US Nashville TN 36807?N86841?W 30 31.5 Jul. 2.3 Jan. 128.8 May. 72.9 Oct. US Richmond VA 37831?N77819?W 30 30.8 Jul. 2.4 Jan. 118.6 Jul. 75.7 Feb. US Montpelier VT 44812?N72835?W 30 25.6 Jul. 13.6 Jan. 101.9 Aug. 49.8 Feb. US Madison WI 43808?N89821?W 30 27.8 Jul. 12.6 Jan. 110.0 Aug. 31.8 Jan. US Charleston WV 38823?N81835?W 30 29.4 Jul. 4.3 Jan. 123.4 Jul. 67.8 Oct. zSource for Canadian data: Environment Canada (http://www.climate.weatheroffice.ec.gc.ca/climate_normals/index_e.html). Source for US data: US National Climatic Data Center (http://cdo.ncdc.noaa.gov). 386 CANADIAN JOURNAL OF PLANT SCIENCE
Echinochloa cus-galli, Oxalis stricta, Panicum dichotomi- to North America near New York city (Fairbrothers florum, Setaria pumila, Setaria viridis, Taraxacum offi- 1959) as a contaminant of millet imported from China in cinale, Plantago major L., and Chenopodium album.In the 1920s (Chase 1948). In 1936, S. faberi was reported corn fields throughout the United States, S. faberi is to be growing in northern Virginia (Allard 1941) where most commonly associated with Abutilon theophrasti, several varieties of millet were being cultivated between Amaranthus powellii, A. retroflexus, Ambrosia artemisii- 1909 and 1925. folia, A. trifida, Bromus japonicus L., Cannabis sativa L., After its introduction into North America, S. faberi Capsella bursa-pastoris (L.) Medik., Chenopodium al- rapidly spread throughout the eastern half of the United bum, Cirsium arvense, Conyza canadensis (L.) Cronquist, States (Fernald 1944; Evers 1949; Pohl 1951; Wood Cyperus esculentus L., Digitaria sanguinalis, Echinochloa 1956; Santelmann and Meade 1961). By 1950 it was crus-galli, Erigeron annuus (L.) Pers., E. strigosus Muhl. described as "becoming a weed in waste and cultivated ex Willd., Ipomoea hederacea Jacq., Muhlenbergia fron- ground, apparently spreading rapidly, New York to dosa (Poir.) Fern., Panicum dichotomiflorum, Polygonum Nebraska and Arkansas, North Carolina, Kentucky, pensylvanicum, Setaria pumilla, Sinapis arvensis L., and Tennessee" (Hitchcock 1950). It is now also present Taraxacum officinale (Abul-Fatih and Bazzaz 1979; throughout the eastern United States to South Dakota, Buhler 1998; Pavuk et al. 1997). In glyphosate resistant Kansas and Oklahoma as well as Arizona and Califor- soybean fields in Iowa, S. faberi was found in mixed nia (USDA, NRCS 2007). The extensive use of 2,4-D to populations with Abutilon theophrasti, Amaranthus spp., control dicotyledonous weeds has contributed to a Chenopodium album, Digitaria sanguinalis, Echinochloa population shift to grass weeds such as Setaria spp. crus-galli, Elytrigia repens, Polygonum convolvulus, Poly- (Rominger 1962; Dekker 2003a). Moreover, the devel- gonum pensylvanicum, Portulaca oleracea, Setaria opment and utilization of selective herbicides in corn pumila, S. viridis, and Xanthium strumarium L. (Buck- and soybean production has exacerbated this trend. In elew et al. 2000). Setaria faberi was one of the top three the 1970s, S. faberi was also reported to be present in the dominant species in oats prior to a burning of the field Red River Valley of North Dakota (Dekker 2003a). with fire. After the burn, S. faberi became the most Currently, the North American range of S. faberi dominant species (Crowner and Barrett 1979). In Ohio continues to expand. wheat fields, S. faberi was found along with Ambrosia artemisiifolia, Aster pilosus Willd., Chenopodium album, Cirsium arvense, Dactylis glomerata L., Daucus carota, 7. Growth and Development Erigeron annuus L., Melilotus officinalis, Phleum pra- (a) Morphology * The height and biomass of S. faberi tense, Polygonum pensylvanicum, Trifolium repens L., plants is largely dependent on the maternal environment and Triticum aestivum (Sedlacek et al. 1988). (Dekker 2003a). Setaria faberi can reach 2 m in height Setaria faberi has also been found among mixed with leaves between 8 and 17 mm wide. The leaves are populations in grassed waterways with Agropyron minutely pubescent beneath and strigose on the upper smithii (Rydb.) A. Lo¨ ve., Bromus inermis Leyss., Hor- surface. The panicles, cylindrical, densely flowered, and deum jubatum L., and Panicum virgatum L. in Iowa spicate are flexuous or slightly arching, 1 17 cm long (Bryan and Best 1991). and 2 3 cm thick. Based at the soil surface, the thread- like subcrown of S. faberi provides nutrients from the 6. History seed to the emerging plant until the crown roots are In Canada, S. faberi was first detected in 1975 in corn established. Before the crown root is established, the fields near Nicolet and Saint-Hyacinthe, Quebec (Do- seedlings are very sensitive to physical control measures yon et al. 1988; herbarium specimen data). Since then it such as rotary hoeing (Knake 1987). has rapidly expanded its range and abundance in the southern part of the province (Doyon et al. 1988). The (b) Perennation * No perennial growth of this annual first specimens of S. faberi from Ontario were collected grass has been reported. It overwinters only as seeds. in 1976 along railway tracks around Toronto, Fort Erie and Niagara Falls (Catling et. al. 1977). By 1981 it had (c) Physiological Data * The C4 photosyntheticpath- been collected in corn fields in eastern Ontario (War- way of S. faberi is of the NADP-malicenzyme type wick et al. 1987). By the time of its discovery in southern (Gutierrez et al. 1974). Like other Setaria species, S. Canada, S. faberi was clearly well established. Its arrival faberi competes efficiently for resources due to its highly seems to have coincided with the expansion of corn plasticgrowth (Dekker 2003a). However, some studies cultivation in the 1960s (Doyon et al. 1988) and to have have shown that in its early stages of growth, S. faberi is been established from several independent sources not competitive for light, moisture or nutrients (Schrei- (Warwick et al. 1987; Doyon et al. 1988). ber and Williams 1967). Many physiological mechan- In the United States, S. faberi was first collected at isms, mostly not yet characterized, are responsible for Philadelphia, PA, in 1923 (McCormick and Kay 1968) these plasticresponses. These in turn, influencethe and at Long Island, NY in 1925 (Fairbrothers 1959). It stature and fitness of the plants especially as they relate has been hypothesized that seeds were also introduced to the size and extent of tillering (Dekker 2003a). NURSE ET AL. * SETARIA FABERI HERRM. 387
Setaria spp. exhibited an exceedingly low degree of and waste areas where S. faberi is commonly the genetic variation when accessions from at least three dominant species (Schreiber 1977). Leachates extracted different continents were sampled (Wang et al. 1995). from live and dead tissue of S. faberi were shown to Setaria geniculata shows the highest geneticdiversity decrease loblolly pine seedling growth by up to 36% followed by S. viridis, S. pumila and S. faberi which is (Gilmore 1980). These leachates also reduced the ability nearly monomorphic. Heterozygosity is rarely detected of the pine seedlings to take up potassium, phosphorus in populations of S. faberi, suggesting that strong and magnesium, while enhancing their ability to take up inbreeding occurs through self-pollination. Populations nitrogen and manganese. Extracts from mature S. faberi of S. faberi are homogeneous but are differentiated from stem tissue resulted in a 50% reduction in the seed one another via fixed alleles at one or more loci. germination and seedling growth of Italian ryegrass Although differences were noted in the pattern of (Lolium perenne L.) and alfalfa (Medicago sativa L.) regional differentiation, S. faberi was nearly invariant. (Martin and Smith 1994). Mester and Buhler (1991) reported that soil tempera- There are concerns that increased competitiveness tures did not affect S. faberi seedling development; between crop and weeds might evolve as atmospheric however, when mulch was added to the soil, the soil CO2 concentration increases. Therefore, studies were temperatures decreased, which in turn increased S. conducted to evaluate the effect of atmospheric CO2 faberi shoot and root growth. The early root growth enrichment on C4 grasses including S. faberi in response (4 to 7 d after seed germination) of S. faberi was the to water stress. Sionit and Patterson (1984) reported that lowest when compared with three other Setaria species under water stress, the stomatal conductance under high (Orwick and Schreiber 1975). However, root diameter CO2 conditions is not significantly affected; whereas and root surface area and volume were highest amongst turgor pressure and net photosyntheticrates increased. the Setaria species when the root lengths were equated Garbutt et al. (1990) tested the growth response of S. by substituting the length of S. faberi roots with the root faberi to three ambient CO2 concentrations (350, 500, radii of each species. Due to these characteristics, and 700 mLL 1). Their study yielded the following control with soil applied herbicides should be improved results: (1) there was no detectable effect of elevated for S. faberi in a wider range of soil herbicide activation CO2 on timing of seedling emergence; (2) the water zones (Orwick and Schreiber 1975). The below-ground potential of shoots increases with increasing CO2 con- root biomass of S. faberi is shallow and fibrous, but is centrations; (3) stomatal conductance decreased with able to occupy a large volume and can overlap with increasing CO2 concentrations when light levels were several other successional species (i.e., Abutilon theo- high; (4) intercellular CO2 levels increased as ambient phrasti, Ambrosia artemisiifolia,andChenopodium al- CO2 levels increased; (5) flowering was delayed at the 1 bum), thus allowing S. faberi to compete for nutrients highest CO2 concentration (700 mLL ); and (6) such as nitrogen, potassium, calcium, and magnesium reproductive biomass did not change with increasing (Wieland and Bazzaz 1975; Parrish and Bazzaz 1976). CO2. Flowering was also delayed for greenhouse-grown Soil nitrogen level has been shown to influence the plants when nutrient levels were low and CO2 levels growth of S. faberi (Salas et al. 1997). Total dry biomass elevated (McConnaughay et al. 1993). However, the increases with increasing nitrogen rates, regardless of overall growth of the plants was not affected by elevated whether nitrogen is applied as NH4 or NO3 (Salas et al. CO2. Other studies have also reported photosynthetic 1997). However, the authors showed that seed produc- rate in S. faberi to be increased (Ziska and Bunce 1997) tion was reduced at higher nitrogen rates when nitrogen or unchanged (He et al. 2002) in response to elevated was applied as NH4. Schreiber and Orwick (1978) CO2, and no effect on above-ground biomass produc- measured the dry weight of S. faberi shoots from plants tion or seed production (Bazzaz and Garbutt 1988; grown in sand-soil nutrient solutions where nitrogen Ziska and Bunce 1997). levels varied 0.5, 1, or 3 times that of the nitrogen The amount of solar UV-B reaching the earth’s present in a Hoagland’s (Hoagland et al. 1938) solution. surface has dramatically increased in recent years due Dry weights of the shoots were significantly affected by to ozone depletion (Kane 1998). This increase in UV-B nitrogen level and plant biomass increased when nitro- radiation has been shown to influence the growth of gen level was increased by a factor of three. This result some grass species (Tosserams et al. 1996; 1997). demonstrated a direct relationship between S. faberi Cybulski and Peterjohn (1999) demonstrated that the biomass and the amount of nitrogen available in the above-ground biomass of S. faberi was not influenced soil. by UV-B levels that were at least 90% of ambient UV-B Root residues collected from S. faberi have allelo- levels (8.1 kJ m 2 d 1) in West Virginia. pathic effects on corn root development (Schreiber and The daily water potential of S. faberi leaves is more Orwick 1978) and overall corn growth (Bell and Koeppe negative than that of Abutilon theophrasti and Poly- 1972). Additionally, other Setaria spp. except S. pumila gonum pennsylvanicum when the species co-exist in an do not develop in close proximity, possibly due to old-field successional community (Wieland and Bazzaz alleopathy. This allelopathic effect is enhanced by low 1975). Under optimal conditions, S. faberi can maintain 2 1 nitrogen levels such as those encountered in fencerows a photosyntheticrate of 38 mg CO 2 dm h at leaf 388 CANADIAN JOURNAL OF PLANT SCIENCE water potentials as low as 1.2 MPa and 30% of this produced 246 seeds per panicle. Overall seed production rate at leaf water potentials as low as 2.4 MPa. was shown to follow a curvilinear relationship with Drought stress and low temperatures have been panicle length with seed production in panicles 100 mm shown to increase epicuticular leaf wax levels in long averaging 323 seeds per panicle (Forcella et al. S. faberi (Hatterman-Valenti et al. 2006). Alternatively, 2000). Additionally, S. faberi fecundity (seeds per plant) growth under low light conditions decreases the amount increased linearly with increases in shoot biomass of leaf wax produced. This shift in leaf wax production (Conley et al. 2002). Seed production has also been may impact the absorption of some herbicides (see also linked to shoot biomass and this grass has been shown Section 11). to produce 88 and 154 seeds per gram of shoot biomass in each of two years (Moechnig et al. 2003). On a whole (d) Phenology * Setaria faberi is sensitive to shade, and plant basis, Schreiber (1965) showed production of more a decrease in incident radiation of 95% in soybean or than 10 000 seeds per plant. Seed production of about corn may cause cessation of growth (Knake 1987). 1000 seeds per plant was also found after additions of Generally, dry weight production decreases linearly swine manure compost in Iowa, but the manure addi- with increases in shade levels (Santelmann et al. 1963; tions did not affect total seed production (Liebman et al. Knake 1972). Peak flowering of greenhouse-grown S. 2004). faberi plants in Wisconsin occurred 63 to 67 d after Crop competition may significantly influence S. faberi planting of seeds (Volenberg and Stoltenberg 2002b). In seed production. Seed production is higher in corn than abandoned fields in Illinois, S. faberi populations soybean because of higher plant and panicle densities showed a bimodal pattern of seed germination where (Forcella et al. 2000). Estimates of 90 000 seeds m 1 of the densities of the populations first peaked in May and corn row have been reported (Fausey and Renner 1997). then in July/August (Raynal and Bazzaz 1975). A Bussan et al. (2000) recorded seed production of up to similar bimodal pattern of seedling emergence was also 4000 seeds per plant in competition with soybean. In a documented in the Northeastern United States (Myers soybean crop not treated with herbicides, the average et al. 2005). seed production was about 600 seeds per plant (11 400 seeds m 2) (DeFelice et al. 1989). In the same study, (e) Mycorrhiza * In greenhouse experiments, S. faberi sethoxydim did not reduce seed production per plant or was found to be only weakly colonized by arbuscular per panicle; however, it did reduce seed production to as mycorrhizal fungi with mean colonization rates up to low as 1020 seeds m 2. Early application of glyphosate 3% (Vatovecet al. 2005). The mycorrhizal responsive- reduced S. faberi seed production per plant by up to ness (effect on biomass production) was greater under 96% (3167 seeds in control vs. 114 seeds in glyphosate higher light and temperature conditions, although the treated plots) (Biniak and Aldrich 1986). colonization rate was not significantly different between Seeds of S. faberi are physiologically independent their two experiments. from the plant when mature, but do not disperse until the pedicel degrades (Dekker 2003b). Timing of dis- 8. Reproduction persal is variable and dispersal may occur by gravity, (a) Floral Biology * Setaria faberi is primarily self animal, wind, water, and humans (Dekker 2003b). pollinated (Dekker 2003a). Autogamy has been shown Additionally, birds, cattle and small mammals probably in the closely related S. viridis (Mulligan and Findlay assist in disseminating seeds (Rominger 1962; McCor- 1970) and agamospermy has been noted in the polyploid mick and Kay 1968). Its rapid and wide dispersal in S. macrostachya species complex (Emery 1957) and thus North America appears to have been primarily due to cannot be ruled out in S. faberi, although Brown and contamination of seed commodities such as millet, Emery (1958) found no evidence of apospory in the eight soybean, red clover and possibly garden flowers (Pohl species of Setaria studied. The pollen to ovule ratio of 1951; McCormick and Kay 1968). Other pathways likely S. faberi was determined to be 1212941.3, in compar- to have contributed to its spread include manure, hay, ison to a ratio of 754.7942.1 for the closely related livestock bedding, contaminated machinery and bird S. pumila (Cruden 1977). seed mixtures (McCormick and Kay 1968).
(b) Seed Production and Dispersal * Seed production in (c) Seed Banks, Viability of Seeds and Germination * S. faberi has been estimated to range from 165 to 2127 Germination occurs throughout the growing season seeds per panicle (Dekker 2003b). Variability in seed (King 1952) and S. faberi may form a small persistent production is directly related to the number of fertile seed bank (Buhler and Hartzler 2001; Leck and Leck spikelets per panicle and this characteristic is plastic 1998). Seeds within the seed bank are subject to (Dekker 2003b). Seed numbers were generally greater depletion and may decline rapidly if there is high annual for S. faberi plants that emerge early in the season seedling emergence (Forcella et al. 1997; Buhler 1999). (Schreiber 1965). Maximum seed rain occurs from July Zhang et al. (1998) showed that 80% of above-ground through December (Peters et al. 1963; Dekeer 2003b). S. faberi populations can be explained by the analysis Forcella et al. (2000) found that on average S. faberi of the soil seedbank. According to Buhler (1999), a NURSE ET AL. * SETARIA FABERI HERRM. 389 high-density weed seed infestation in Minnesota com- Seedling emergence from the seed bank is negatively prised 60 000 seeds m 2 of S. faberi, whereas a low affected by tillage (Buhler and Daniel 1988; Buhler and infestation was 1000 seeds m 2 (Buhler 1999). Leck and Oplinger 1990). Buhler et al. (1996) found that S. faberi Leck (1998) found an early seed bank of about 10 000 emergence was greater in no-till (100 to 1700 plants seeds m 2 in a successional old-field, which decreased to m 2) versus conventionally tilled plots (100 to 500 an average size of less than 2000 seeds m 2 after 4 yr. In plants m 2). Alternatively, Myers et al. (2005) found no a corn/soybean rotation in Wisconsin, the seedbank effect of tillage on emergence pattern and that tillage oscillated between 5000 and 10 000 seeds m 2 over a 20- had little effect on the total number of seedlings present yr period (Bussan and Boerboom 2001). Rothrock et al. in the field. When corn residue was added to the plots, (1993) found much smaller persistent seed banks in emergence was further reduced by 42 to 95%. Similarly, Indiana where size ranged from 152922 to 1994 seeds no-till plots yielded 500 800 plants m 2 versus about m 2. The authors measured a seed bank size of about 200 plants m 2 in conventional tillage 60 d after crop 164 seeds m 2 10 yr prior to the start of the experiment planting in Wisconsin (Buhler and Oplinger 1990). In suggesting that the persistent seed bank of S. faberi is Minnesota, S. faberi comprised 32% of the viable seed stable. The persistence of the S. faberi seedbank across bank and 5% of the total seed bank under long-term eight mid-western US states was found to be 60% of its conventional tillage (Reuss et al. 2001). In Indiana, original size after 3 yr (Davis et al. 2005). tillage may result in 55% of seeds becoming seedlings Tillage and crop type have substantial effects on seed (Rothrock et al. 1993). In Iowa, S. faberi seeds emerged bank size as well as the depth of seeds within the soil only in the first 3 yr of a 4-yr study in tilled soil. In the profile. The maximum depth for S. faberi seedling first year, 30% of the original seed bank emerged and emergence is estimated at 10 12 cm (King 1952; Mester the seed bank was reduced by 42% after 3 yr (Buhler and Buhler 1986), although this is undoubtedly depen- and Hartzler 2001). A similar study showed that while dant on soil compaction and moisture. In a 30-yr long 43% of seeds emerged in the first 2 yr of a burial study, tillage trial in Ohio, Cardina et al. (1998) found that seed only 1% of the total seeds emerged in the third year banks were larger in no-till soils (1140 seeds m 2) when (Hartzler et al. 1999). For seedlings that do emerge, at compared with soils that were chisel (210 seeds m 2)or least 10% of the emergence was shown to occur after moldboard (70 seeds m 2) plowed. Tillage reduced seed 180 degree days (DD) (from late April to early May in bank size in the top 5 cm of soil from 139 to 54 seeds Pennsylvania) with 75% emergence between 200 to 500 m 2 in Indiana (Rothrock et al. 1993). After 35 yr of DD (Myers et al. 2004). The density of seedling continuous crop rotation and tillage, S. faberi seed emergence in their study ranged from 148 plants m 2 banks were found to be highest (6890 seeds m 2)ina to 1040 plants m 2. Menalled et al. (2005) demonstrated no-till corn and soybean rotation at one site and in a that the addition of composted swine manure decreased continuous corn rotation (also 6890 seeds m 2) on no- S. faberi seedling emergence from 15 to 57%. till soils at a second site (Cardina et al. 2002). Reduced Herbicide control of S. faberi reduces the soil seed tillage or no-till has been shown to concentrate S. faberi bank. In Iowa, when herbicides were banded or omitted, seeds in the upper horizons of the soil profile. In Setaria spp. dominated the weed seed bank. Setaria spp. Wisconsin, S. faberi was the largest component of seeds only constituted 25% of the seed bank with broadcast in the soil seed banks of no-till soils with 40% of S. spraying (Hoffman et al. 1998). Buhler (1999) compared faberi seeds emerging from the top 1 cm of no-till soils different management practices and found that the seed and less than 5% at a depth of 4 cm (Buhler and Mester densities in weed-free (herbicide, cultivation, bi-weekly 1991). Several other studies have also documented that handweeding) plots were reduced from 57 000 to 9000 S. faberi seeds remain within the top 5 cm of the soil seeds m 2 in 4 yr. In the same study, herbicides alone profile in no-till fields and old field communities (Big- reduced seed banks from 61 000 to 36 000 seeds m 2, wood and Inouye 1988; Hoffman et al. 1998; Leck and while mechanical control increased seed banks from 59 Leck 1998; Reuss et al. 2001). In Iowa, a soybean/corn 000 to 76 000 seeds m 2. The greatest decreases were in rotation had the majority of seeds in the top 5 cm, while the first two years (78%) which then stabilized in a continuous corn rotation caused seeds to move deeper subsequent years at about 15% of the initial density within the profile (10 15 cm) (Hoffman et al. 1998). In (Buhler 1999). Glyphosate application at the first head- Maryland, dormant seeds of S. faberi were buried at ing stage reduced the weight of S. faberi seeds (1.026 g depths of 2.5, 7.6 and 15.2 cm and germination was 1000 1 control vs. 0.539 g 1000 1) and germination by monitored after burial lengths of between 3 and 6 mo. up to 95% (Biniak and Aldrich 1986). After 6 mo of burial, seeds entered into secondary Viability and germination of freshly collected S. faberi dormancy. Percentage germination was greatest at the seeds is variable. Viabilty was about 80% in Illinois 2.5 cm depth (Taylorson 1986). The highest seedling (King 1952), 63 75% in western Minnesota and South emergence of S. faberi in Michigan was from a depth of Dakota (Forcella et al. 2000), 91% in Minnesota (Reuss 1 cm (74% after 21 d) and 1 to 2.5 cm (64% after 21 d). et al. 2001), 78 to 92% in Iowa (Leon et al. 2004), and Only about 3% of seeds emerged from a depth of 7.5 cm 90% in New York (DiTommaso and Nurse 2004). or greater (Fausey and Renner 1997). Germination of seeds collected from within a corn/ 390 CANADIAN JOURNAL OF PLANT SCIENCE wheat rotation ranged from 21 to 59% in Iowa (Davis 26, and 32oC (producing 25 different alternations) with and Liebman 2003). Seed collected in Minnesota from a a 16-h photoperiod in comparison to each at a constant field under long-term conventional tillage, and seeded to temperatures for 24 h (40 vs. 70%, respectively). annual crops had viability increased from 2 to 42% after Germination of S. faberi seeds occurs over a wide an over-wintering period, possibly due to after-ripening range of temperatures. Optimal germination was at or additional seed deposition. Highest seed viability was 248C for seeds collected in Iowa (Leon and Owen found in soil aggregates of 1 2 mm size (25%), but 2003). At constant temperatures, higher percentage viability of 21% was also found in aggregates 12 mm germination was obtained at 208C (77%) than at 308C in size (Reuss et al. 2001). When buried in soil at a depth (61%) (Fausey and Renner 1997). Seed germination of of 7.6 cm, S. faberi seeds lost viability most rapidly out 81% was obtained at an alternating temperature of 14/ of six weed species tested in Maryland (Taylorson 1972). 288C for 9 h day/15 h night, which simulated June and In the same study, approximately 35, 2, and 67% of July temperatures and photoperiods in Michigan (Fau- seeds remained viable at burial depths of 2.5, 7.6, and sey and Renner 1997). King (1952) found the highest 15.2 cm, respectively. In a burial study in Iowa, seed germination with pre-treatments at alternating tempera- viability of S. faberi decreased from 93% at the time tures of 21/378C for 8 h day/16 h night (48%) as of burial to B20% after only one year (Buhler and opposed to 21/308C (20%) or constant temperatures Hartzler 2001). (10%) of 21, 30 or 378C. Setaria faberi seeds collected in Two unidentified soil fungi were reported in caryopses Iowa germinated best at temperatures between 20 and of S. faberi by Pitty et al. (1987) which had a detrimental 248C, reaching a plateau at 288C. Temperatures above effect on germination. The extent of colonization by the 308C reduced the germinability of seeds (Leon and fungi was correlated with soil depth and tillage system Owen 2003). A similar study in Iowa showed that the (see also Section 13bi). minimum and optimal temperatures for germination Seeds of S. faberi possess non-deep physiological were 14 and 248C, respectively (Leon et al. 2004). In the dormancy (Baskin and Baskin 1998) that is affected by same study, increased amplitude of the diurnal tempera- temperature, aeration and moisture (King 1952). Ex- ture alternation increased germination to over 50% at periments by King (1952) suggest the presence of water- mean temperatures between 20 and 258C. Germination soluble germination inhibitors; however, the authors did at 10, 15 and 208C was 20, 77 and 72%, respectively. not make conclusions as to the source of the inhibitors. Moist stratification was not effective in breaking Environmental factors may also cause non-dormant or dormancy of seeds collected in a different Iowa study, seeds that have previously lost dormancy to enter into a but the viability of non-germinating seeds was low. secondary dormancy (Taylorson 1982). Temperatures Germination started when soil temperatures were at 10 greater than 308C were needed to induce secondary to 158C and peaked when soil temperatures reached dormancy, possibly due to changes in membrane func- 208C. Natural seed banks required about 100 GDD tion and metabolism (Taylorson 1982; Taylorson 1986; (base 108C) to reach 50% emergence (Leon and Owen Kegode and Pearce 1998; Leon et al. 2004). Forcella et 2004). Pre-chilling seeds at 58C for 1 mo followed by the al. (1997) suggested that once soil temperatures reach piercing of the seed coat was effective in relieving 178C within the top 10 cm, conditions are appropriate dormancy of seeds collected in Missouri (Stanway for S. faberi seeds to enter secondary dormancy. Levels 1971). Moreover, dormancy was reduced by dry storage of dormancy in freshly collected seeds are quite variable. at room temperature for 1 yr. In monoculture field pot An attempt to germinate freshly collected seeds from experiments in Michigan, seedling emergence was 22% 7 Missouri at alternating temperatures of 208C (8 h light) d after planting (DAP) at May temperatures of 7/208C; to 308C (16 h dark) for 21 d revealed that almost all and 33% 21 DAP. Overall, S. faberi had higher seeds were dormant (Stanway 1971). Similarly, Mulu- germination at lower versus higher temperatures (Fau- geta and Stoltenberg (1997) reported high levels sey and Renner 1997). ( 77%) of seed dormancy in freshly collected seeds in Dormancy in S. faberi may be partially regulated by Wisconsin. Low temperatures ( 20 to 508C) and phytochrome. In Iowa, light quality did not affect seed aerobic conditions have been shown to effectively slow germination; however, far-red light reduced germination after-ripening and retain seed viability (Dekker 2003a). in pre-chilled seeds (Leon and Owen 2003). Large Conversely, accelerated after-ripening occurs when changes in far-red light levels were found to only add seeds are exposed to temperatures greater than 508C small increments of germination to an already small for up to 14 d increasing percentage germination in population of dark germinating seeds (Taylorson 1972). comparison to a control at lower temperatures (Fausey Taylorson (1982) showed that 0.1 M ethanol and other and Renner 1997; Dekker 2003a). Taylorson and Brown chemicals such as methanol, and 1-propanol prevented (1977) found that dormancy was best overcome by after- secondary dormancy induction in S. faberi. Germination ripening seeds in the light at constant temperatures of was initially insensitive to light, but a response to red 40, 50, and 608C for between 3 and 14 d. Leon et al. irradiation was triggered by the addition of 0.5 M (2004) found that seed germination was increased by ethanol. Furthermore, the ethanol allowed germination combinations of alternating temperatures of 8, 14, 20, to occur after a treatment had been previously applied to NURSE ET AL. * SETARIA FABERI HERRM. 391 induce secondary dormancy (Taylorson 1982). Gallagher herbicide s-metolachlor were also incorporated into the and Cardina (1998) observed that field emergence of soil. S. faberi was increased by 30% when tillage operations were performed during the day rather than at night. (d) Vegetative Reproduction * Setaria faberi reproduces Maternal environment may influence dormancy levels only by seed, although tillering readily occurs. Tillering of seeds prior to dispersal from the maternal plant. in S. faberi is known to be strongly influenced by Studies in Iowa showed that seeds of S. faberi produced photoperiod and competition (Dekker 2003). For ex- by a greenhouse population were more dormant than ample, under long photoperiods and reduced competi- seeds collected from a field population. This was tion S. faberi may produce several primary, secondary possibly the result of development and maturation of and tertiary tillers, all with the ability to set seed. Under seeds at higher temperatures (Kegode and Pearce 1998). shorter photoperiods, there may be no tillering at all In the same study, moist stratification increased overall (Dekker 2003). Tillers may also root when in close seed germination and reduced differences in percentage proximity to moist soil (Santleman et al. 1963). germination between field and greenhouse produced seed lots. These authors suggest that geneticcontrols 9. Hybrids may also be regulating seed dormancy in S. faberi An extremely rare allo(tetra)polyploidization event may because when germination was tested for seeds from have caused a fertile hybridization product in southern different populations in a common environment, the China where an ancient cross of Setaria viridis (L.) Beauv. respective populations retained differences in germina- with S. adhaerans (Forssk.) Chiov., led to the formation tion (Kegode and Pearce 1998). of the tetraploid species, S. faberi (Li et al. 1942). Pickett and Bazzaz (1978) demonstrated that S. faberi Benabdelmouna et al. (2001b) using genomicin situ seeds would not germinate in waterlogged soil. In fact, hybridization showed that in S. faberi, one set of S. faberi seeds were very tolerant to moisture stress 18 chromosomes shared the genetic makeup of S. viridis, (Raynal and Bazzaz 1973) and, in some cases, germina- and the second set of 18 chromosomes with S. adhaerans. tion of S. faberi seeds increases under moisture-stressed Seeds were produced when S. viridis (n 9) and S. faberi conditions (Taylorson 1986). Seed germination was 55% (n 18; tetraploid) were cross pollinated (Willweber- in water, but increased to 90% after pre-treatment in Kishimoto 1962). Li et al. (1942) and Pohl (1962) reported polyethylene glycol (PEG) 8000 at 0.3 MPa or less producing the artificial hybrids S. faberi S. italica (L.) (Taylorson 1986). Continuous exposure to water poten- P. Beauv. as well as (S. viridis S. italica) S. faberi, tials of 0.5 MPa or less inhibited germination, but but the crosses were sterile. caused an increase in germination when seeds were later exposed to water (up to 96% vs. 52% in control). Leakage of electrolytes over the first 15 min of imbibi- 10. Population Dynamics tion was reduced by PEG due to its direct effect on Setaria faberi is very sensitive to shade and plant height water uptake (Taylorson 1986). Additionally, seed is reduced by more than 50% when plants are heavily germination was found to be positively affected by shaded (Peters et al. 1963). This annual grass is most increasing O2 levels above 20%, although germination commonly classed as a dominant early successional was not affected when O2 levels were lowered from 20 to species (Parrish and Bazzaz 1982). Being a drought- 10% (Dekker and Hargrove 2002). Carbon monoxide tolerant species with a high photosynthetic rate con- (CO) stimulated germination when seeds were exposed tributes to the competitiveness of S. faberi in early to low levels of the gas (Dekker and Hargrove 2002). successional communities (Perozzi and Bazzaz 1978). Carbon dioxide (CO2) increased the rate of germination Setaria faberi was the most dominant and most when applied exogenously at a concentration of 20 competitive species in early successional communities 1 mmol mol ;CO2 also accelerated the entry of seeds tested in Illinois and densities of this species were always into secondary dormancy when environmental condi- highest, except when Aster pilosus was present in the tions suitable for germination were not present system (Perozzi and Bazzaz 1978). Of the 45 species (Yoshioka et al. 1995). found in an old-field system in Ohio, S. faberi was Soil nutrient levels may also influence S. faberi seed dominant, even after the application of amitrole plus germination. Under nutrient levels of 0.03 g N, 0.06 g ammonium thiocynate (Wakefield and Barrett 1979). P, and 0.03 g K, the rate of germination of Setaria Endress et al. (1999) suggest that S. faberi is better suited seeds was reduced in comparison to seeds sown in soils for interspecific competition than intraspecific competi- that only had 1/4 or 1/8 the level of nutrients (Parrish tion due to giant foxtail’s ability to outcompete other and Bazzaz 1985). The emergence of S. faberi seedlings species under increasing atmospheric ozone concentra- was stimulated when at least 249 g m 2 hairy vetch tions and decreasing water availability. In a greenhouse (Vicia villosa Roth) residues were placed on the soil experiment where S. faberi was grown at different ratios surface, presumably due to an increase in available in mixture with the perennial grasses Andropogon nitrogen provided by the vetch (Teasdale et al. 2005). gerardii Vitman, and Sorghastrum nutans (L.) Nash; This effect was enhanced when reduced rates of the S. faberi always outcompeted these two species. 392 CANADIAN JOURNAL OF PLANT SCIENCE
In an old-field community in New Jersey, S. faberi flufenacet was tank-mixed with atrazine, dicamba, or always out-competed other grasses such as Panicum dicamba/atrazine there was no improvement in percen- dichotomiflorum (Facelli and Pickett 1991b). However, tage control. However, tank mixtures with metribuzin or when growing in stands with common dicotyledonous linuron did improve percentage control to 98% (PMRA annuals such as Abutilon theophrasti, Ambrosia artemi- 2000). siifolia, and Amaranthus retroflexus, S. faberi was the A model developed by Bussan and Boerboom (2001) least competitive species (Bazzaz and Garbutt 1988). for control of S. faberi in a corn/soybean rotation The population density of S. faberi was reduced from showed that over the long term S. faberi populations 100 plants m 2 to 10 plants m 2 by heavy leaf litter could be managed more economically when herbicide from Quercus spp. Facelli and Pickett (1991b) specu- rates are reduced unless the initial seedbank populations lated that this effect was regulated primarily by intras- were high. Broadcast applications of alachlor (3.3 kg a.i. pecific competitive interactions among S. faberi ha 1) plus glyphosate (0.4 kg a.e. ha 1) and bromox- individuals. The net primary production (NPP) con- ynil (0.4 kg a.i. ha 1) provided 100% control of S. tribution of S. faberi was estimated to be B5% of the faberi populations in conventionally grown corn in total NPP in an old-field successional system in Ohio Minnesota (Buhler 1998). Mesotrione (240 g a.i. ha 1) (Sedlacek et al. 1988). applied alone (PRE or POST) provided less than 33% Densities of S. faberi range from 98 to 1040 plants control of S. faberi in conventional corn grown in m 2 in corn fields throughout the Northeastern United Virginia (Armel et al. 2003b, c). Percentage control was States (Myers et al. 2005). In a soybean field in increased to greater than 84% when mesotrione (240 g Wisconsin comprised of 27 different weed species, S. a.i. ha 1) was tank-mixed with acetochlor (1800 g a.i. faberi accounted for 62 75% of the weed density (147 to ha 1), or imazethapyr (70 g a.i. ha 1). When applied to 452 plants m 2) (Mulugeta and Boerboom 2000). In glyphosate-resistant corn, a tank mixture of mesotrione soybean fields in Wisconsin, S. faberi was more compe- (70, 105, or 140 g a.i. ha 1) plus glyphosate (560 g a.i. titive than Chenopodium album, and a S. faberi popula- ha 1) provided up to 75% control compared with B tion density of 64 plants m 2 resulted in more than an 25% with using mesotrione (70, 105, and 140 g a.i. 86% soybean yield loss. In this same study, the critical ha 1) alone (Armel et al. 2003d). All PRE and mid- period of S. faberi competition in soybeans was deter- POST applications of glyphosate (1100 g a.i. ha 1), mined to be up to the V2 leaf stage of the crop (Conley acetochlor (2000 g a.i. ha 1), atrazine (1100 g a.i. ha 1), et al. 2003b). In a similar study, Conley et al. (2003a) metolachlor (2200 g a.i. ha 1), dimethenamid (1500 g demonstrated that S. faberi could cause up to 75% yield a.i. ha 1) alone and in tank-mix combinations provided loss in soybeans if allowed to germinate prior to the V2 greater than 95% control of S. faberi in Kentucky stage of the crop. Weed shoot biomasses of 2000 and (Ferrell and Witt 2002). At least 90% control was 3500 kg ha 1 reduced soybean yield by 21 and 32%, obtained for plants 23 to 30 cm in height when respectively. When S. faberi emerged in the field at the sequential (2 or more) applications of glyphosate (800 V3 or later stage of soybeans there was no yield loss. At g a.i. ha 1) were applied to glyphosate-resistant corn in low densities (B16 plants m 2), soybean yield was Ohio (Gower et al. 2002). In glufosinate-resistant corn reduced by only 5.1%; however, S. faberi produced up in Missouri, control levels of 90% were obtained when to 28 000 seeds per plant (Conley et al. 2003a). acetochlor (1120 g a.i. ha 1) PRE followed by atrazine The relative fitness of acetyl-coenzyme A carboxylase (1120 g a.i. ha 1) plus glufosinate (290 g a.i. ha 1) (ACCase)-resistant S. faberi individuals was not reduced POST was applied. Glufosinate alone provided less than relative to ACCase-susceptible individuals when sub- 80% control (Bradley et al. 2000). Trifluralin (1.12 to jected to inter- and intra-specific competitive environ- 1.40 kg a.i. ha 1) applied PRE in a conservation tillage ments (Wiederholt and Stoltenberg 1996). (fall chisel plowed) rotation of corn and wheat in Indiana and Kansas provided greater than 90% control 11. Response to Herbicides and Other Chemicals of S. faberi (Regan 1995). Applications of halosulfuron- The increased occurrence of S. faberi in corn and methyl (36 g a.i. ha 1), nicosulfuron (18 g a.i. ha 1), or soybean fields has provided valuable information with dicamba (280 g a.i. ha 1) all provided at least 89% regards to herbicide efficacy and resistance. This review control in Delaware (Isaacs et al. 2002). Sethoxydim is by no means exhaustive, but does provide a summary (213 g a.i. ha 1) alone provided 96% control and tank of major herbicides efficacies and known resistance. It mixtures of sethoxydim (213 g a.i. ha 1) plus bentazon should also be noted that in the scientific literature, (1120 g a.i. ha 1), dicamba (280 g a.i. ha 1), dicamba there are sometimes no clear distinctions made between (280 g a.i. ha 1) plus atrazine (588 g a.i. ha 1), and the various Setaria spp. in relation to chemical control sethoxydim (213 g a.i. ha 1) plus bentazon (1120 g a.i. efficacy (Steel et al. 1983). ha 1) plus atrazine (1166 g a.i. ha 1) reduced the levels Flufenacet was applied to conventionally grown corn of control to between 60 and 89% in sethoxydim- and soybeans in Ontario and Quebec. When applied resistant corn in Delaware (Isaacs et al. 2003). alone pre-emergence (PRE) at a rate of 570 g a.i. ha 1, In no-till corn in Illinois, herbicides applied PRE flufenacet provided 95.7% control of S. faberi. When provided more consistent control (90 to 98%) than the NURSE ET AL. * SETARIA FABERI HERRM. 393 same herbicides applied early pre-plant (0 to 92%) glyphosate (840 g a.i. ha 1) POST resulted in 80 to (Krausz et al. 2000). Ninety-five percent control was 100% control of S. faberi in no-till narrow-row glypho- obtained after an application of metolachlor (3400 g a.i. sate-tolerant soybean in Missouri (Dirks et al. 2000a). If ha 1) and flufenacet (920 g a.i. ha 1) plus metribuzin glyphosate was applied at planting at a rate of 430 g a.i. (230 g a.i. ha 1) and 87% control was obtained with ha 1 sulfentrazone (220 g a.i. ha 1), S. faberi popula- acetochlor (2700 g a.i. ha 1) or dimethenamid (1600 g tions were reduced more than if glyphosate was applied a.i. ha 1) in Illinois (Bunting et al. 2003). A burndown alone at a rate of 430 g a.i. ha 1 in no-till narrow-row application of glyphosate (630 g a.e. ha 1) plus a full glyphosate-tolerant soybeans in Missouri (Dirks et al. label rate of the residual herbicides acetochlor (2240 g 2000b). Flumioxazin applied at a rate of 70 g a.i. ha 1 a.i. ha 1) and atrazine (2240 g a.i. ha 1) applied early in no-till soybeans in Missouri resulted in variable preplant provided less than 80% control in Missouri control of S. faberi ranging from 18 to 81% control (Hellwig et al. 2003). An imazethapyr (470 g a.i. ha 1) (Niekamp and Johnson 2001). In the same study, application provided better residual control ( 96% percentage control was also variable and ranged from control) (Hellwig et al. 2003). Percentage control 110 d 65 to 93% if sulfentrazone was applied at a rate of 280 g after no-till corn planting was only 73% after an early a.i. ha 1. The addition of clomazone (840 g a.i. ha 1) preplant treatment of atrazine (2.2 kg a.i. ha 1) alone, clomazone (840 g a.i. ha 1) chlorimuron (60 g compared with 99% control when the same amount of a.i. ha 1), or pendimethalin (1350 g a.i. ha 1) herbicide was evenly divided between early preplant and chlorimuron (60 g a.i. ha 1) into a tank-mix with either preemergence applications in Wisconsin (Buhler 1991). herbicide provided substantially more effective S. faberi In Virginia, PRE or POST applications of mesotrione control. (70, 105, and 140 g a.i. ha 1) did not adequately control When metolachlor (2240 g a.i. ha 1) was applied with S. faberi (Armel et al. 2003a; Armel et al. 2003b). The metribuzin (425 g a.i. ha 1) clomazone (280 g a.i. best season long control of S. faberi in Pennsylvania was ha 1) PRE, flumiclorac (45 g a.i. ha 1) POST, or with metolachlor (2.2 kg a.i. ha 1), microencapsulated metribuzin (425 g a.i. ha 1) PRE and flumiclorac (45 g alachlor (2.8 kg a.i. ha 1), or EPTC (4.5 kg a.i. ha 1) a.i. ha 1) POST, S. faberi populations were reduced by (Ritter et al. 1989). at least 85% in soybean fields in Michigan (Fausey and In glyphosate-tolerant soybeans in Wisconsin, gly- Renner 2001). In Illinois, the addition of lactofen (146 g phosate applied at rates as low as 210 g a.i. ha 1 to S. a.i. ha 1) to a tank-mix with imazethapyr (67 g a.i. faberi plants up to 28 cm in height resulted in 90% ha 1) decreased S. faberi control in a soybean crop control (Ateh and Harvey 1999). Glyphosate (420, 630, relative to an application of imazethapyr alone (Hart et and 840 g a.i. ha 1) applied at the V2, V4, and R1 al. 1997). In the same study, S. faberi control with stages of glyphosate-tolerant soybean gave excellent sethoxydim (213 g a.i. ha 1) and fluazifop (190 g a.i. control of S. faberi in both 18 cm and 76 cm row ha 1) fenoxaprop (45 g a.i. ha 1) was equal to control soybean (Mulugeta and Boerboom 2000). When gly- provided by application of imazethapyr alone. The phosate was preceded by a residual herbicide in no-till application of sulfentrazone at a rate of 280 g a.i. glyphosate-tolerant soybeans, populations of S. faberi ha 1 resulted in 97 to 100% control of S. faberi in were reduced by 90% in Wisconsin (Corrigan and soybean in Illinois (Krausz and Young 2003). Harvey 2000). In glufosinate-tolerant soybeans in Mis- Setaria faberi is also an important economic weed in souri, glufosinate applied at 290 and 400 g a.i. ha 1 several other crops. In a Virginia potato crop, rimsul- provided greater than 85% S. faberi control. However, furon at a rate of 35 g a.i. ha 1 PRE or POST two applications of glufosinate (290 g a.i. ha 1 followed controlled 90% of S. faberi (Ackley et al. 1996). by 290 g a.i. ha 1) during the cropping season provided Oxyfluorfen at rates of 280, 414, 560, and 1120 g a.i. followed by 90% control (Beyers et al. 2002). A 28-g a.i. ha 1 resulted in 71, 85, 90, and 100% control of S. ha 1 rate of quizalofop with 1.5% petroleum oil faberi, respectively, in comparison to a cultivated concentrate (POC) resulted in up to 99% control of S. control in transplanted broccoli in Tennessee (Eaton et faberi, while a 56 g a.i. ha 1 rate of sethoxydim with al. 1990). Isoxaben (840 g a.i. ha 1) provided less than 1.5% POC resulted in up to 79% control in soybean 70% control of S. faberi 12 wk after treatment (WAT) fields in Illinois (Beckett et al. 1992). and pronamide (2200 g a.i. ha 1) did not provide A PRE application of (metolachlor at 2220 g a.i. control of S. faberi in landscape crops in Kentucky ha 1 metribuzin at 314 g a.i. ha 1 chlorimuron at (Setyowati et al. 1995). In the same study, oxyfluorfen 54 g a.i. ha 1) and POST application of (quizalofop at (1100 g a.i. ha 1) and dithiopyr (2200 g a.i. ha 1) 62 g a.i. ha 1 thifensulfuron at 4 g a.i. ha 1 provided adequate control. In Tennessee snap bean chlorimuron at 4 g a.i. ha 1) resulted in 95% control fields, sethoxydim (280 g a.i. ha 1) provided 96 100% of S. faberi in no-till soybeans in Illinois (Bradley et al. control of S. faberi (Mullins et al. 1993). 2001). Imazethapyr (55 g a.i. ha 1) applied early pre- When S. faberi was grown in a greenhouse or a corn plant reduced S. faberi population by 90% in no-till field, foramsulfuron alone (37 g a.i. ha 1), or foram- soybean in Wisconsin (Buhler and Proost 1992). Gly- sulfuron (37 g a.i. ha 1) a methylated seed oil (MSO) phosate (840 g a.i. ha 1) early preplant followed by provided higher control of S. faberi in comparison with 394 CANADIAN JOURNAL OF PLANT SCIENCE when a crop oil concentrate (COC) was added instead of shown to have phytotoxiceffectson S. faberi (Coleman the MSO (Bunting et al. 2004a, b, 2005). When and Penner 2006). Pyridazocidin, a microbial phyto- glufosinate (291 and 409 g a.i. ha 1), glyphosate toxin, showed rapid (within 1 to 2 d) chlorosis and (1120 g a.i. ha 1) or sulfosate (1120 g a.i. ha 1) were necrosis of S. faberi leaf tissue at a rate of 1400 g a.i. applied in fallow field areas, S. faberi populations were ha 1 (Gerwick et al. 1997). Ferulic acid, catechol, reduced by at least 90% in North Carolina (Corbett et salicylic acid and caffeic acid all showed some allelo- al. 2004). Antagonism occurred when a tank-mix of pathicpotential in preventing the germination of S. imazethapyr and glyphosate at a rate of 210 g a.i. ha 1 faberi seeds; however, in field situations adsorption to was applied on S. faberi grown in monoculture (Li et al. soil particles may reduce their activity (Vidal et al. 2002). Corn gluten meal applied at a rate of 490 g m 2 1998). Water-soluble substances found in alfalfa inhib- reduced weed cover in greenhouse-grown strawberry ited seed germination, but stimulated plant height, leaf plots in Iowa from 80% (of which S. faberi was area, and dry weight of S. faberi plants (Chung and included) to as low as 8% of a control. The efficacy of Miller 1995). corn gluten meal to control S. faberi was not reported In Saskatoon, a potential bioherbicide, Pyricularia (Nonnecke and Christians 1993). Diclofop applied at setariae Niskada was tested and found to have low 560 g a.i. ha 1, 840 g a.i. ha 1, and 1120 g a.i. ha 1 pathogenicity on S. faberi. When the fungus was paired controlled 75, 25.4, and 16.4% S. faberi, respectively, in with 0.25 the labelled rate of sethoxydim (label rate cucumber, soybean, sorghum and wheat (Schreiber et al. 200 g a.i. ha 1) there was a 10-fold increase in the 1979). Thiazopyr, a pre-emergence grass herbicide, pathogenicinfectionof S. faberi, and a 100% increase in inhibited the growth of S. faberi in a greenhouse study weed control in comparison to the pathogen alone (Peng by 50% when the herbicide was sprayed at a rate of 4 g and Byer 2004). a.i. ha 1. The addition of piperonyl butoxide (PBO) increased the efficacy of thiazopyr by a factor of 1.5 (Rao et al. 1995). Alachlor (2.24 kg a.i. ha 1), atrazine Herbicide Resistance (2.24 kg a.i. ha 1), butylate (3.36 kg a.i. ha 1), and Setaria faberi resistance has been reported in Ontario cyanazine (2.24 kg a.i. ha 1) all provided acceptable [ALS inhibitors (B/2)], Spain [Photosystem II inhibitors control ( 95%) of S. faberi in Wisconsin (Harvey 1974) (C1/5)], Iowa [Photosystem II inhibitors (C1/5) and and haloxyfop-methyl gave better than 95% control at ACCase inhibitors (A/1)], Maryland [(Photosystem II rates of 101 g a.i. ha 1 and 134 g a.i. ha 1 and showed inhibitors (C1/5)], Wisconsin [ACCase inhibitors (A/1) no antagonism when tank mixed with broadleaf herbi- and (ALS inhibitors (B/2))], and Minnesota [ALS cides in several northern, and mid-western states (Han- inhibitors (B/2)] (Heap 2007). son and Velovitch 1986). Triazine resistance in S. faberi was also identified in When S. faberi was planted in non-crop stands the Pennsylvania (Ritter et al. 1989) and Iowa (Thornhill herbicides simazine (1.1 kg a.i. ha 1), atrazine (1.1 kg and Dekker 1993). A resistant biotype tolerated atrazine 1 a.i. ha 1), and propazine (1.1 kg a.i. ha 1) all yielded at a rate of 9.0 kg a.i. ha , while a susceptible biotype 1 better than 90% control (Oliver and Schreiber 1971). In was injured by a rate of 2.2 kg a.i. ha (Ritter et al. the same study, the herbicide butylate only provided 1989). The inadequate control of S. faberi by atrazine 79% control at a 1.1 kg a.i. ha 1 rate and 94% control and other selective herbicides may be due to detoxifica- at a 3.4 kg a.i. ha 1 rate. Normally about 30% of tion by glutathione-S-transferases (GSTs) (Hatton et al. atrazine applied penetrates into the leaves and is 1995). These GSTs may detoxify atrazine, metolachlor, metabolized (McCall 1989). alachlor, and fluorodifen. Atrazine may only be phyto- The control of S. faberi was antagonized when toxicto young S. faberi plants (B35 d old). Up to 80% lactofen (70 g a.i. ha 1) was tank-mixed with either of S. faberi plants were affected by atrazine application imazamox (35 g a.i. ha 1) or imazethapyr (70 g a.i. (400 g a.i. ha 1) at ages below 35 d old. Glutathione ha 1), and when imazethapyr (70 g a.i. ha 1) was tank- (GSH) content was higher (700 nmols g 1 FW) in mixed with clethodim (140 g a.i. ha 1) in soybeans younger tissue than in older tissue (300 nmols g 1 FW). grown in Michigan (Nelson et al. 1998). In the same The GST activities of the whole plant remain un- study, S. faberi control with imazamox (35 g a.i. ha 1) changed. Setaria faberi may detoxify atrazine through or imazethapyr (70 g a.i. ha 1) plus a non-ionic GSH conjugation (Hatton et al. 1996). The levels of surfactant (0.25% vol/vol) was improved when 28% GST in S. faberi foliage are 20-fold lower than levels urea ammonium nitrate (2.6% vol/vol) was added into found in corn foliage. Low levels of GST may increase the tank mixture. the effectiveness of chemicals such as atrazine when The sesquiterpene lactone, artemisinin that is pro- applied at the seedling stage (Hatton et al. 1999). Two duced by the leaf tissue of Artemisia annua L. has been mechanisms of atrazine resistance were identified in shown to inhibit seed germination and seedling growth S. faberi growing in corn fields in Spain (De Prado et al. of S. faberi when applied at a concentration of 200 mg 2000). The resistant biotype of S. faberi was 10 times per 9-cm-diameter Petri dish (Chen and Leather 1990). more resistant to atrazine than the susceptible biotype. The short chain fatty acid, caprylic acid has also been Furthermore, the resistant biotype metabolized the NURSE ET AL. * SETARIA FABERI HERRM. 395 atrazine into the conjugate-atrazine faster than the mammals consume seeds of S. faberi (McCormick and susceptible biotype. The authors concluded that Kay 1968). S. faberi may have two resistance mechanisms (1) a Harvest mouse [Reithrodontomys megalotis (Baird)] mutation in the D1 protein leading to a lower affinity populations in Indiana were higher in grassy areas (that for atrazine and (2) an enhanced detoxification of included S. faberi) compared with areas that were atrazine to conjugate-atrazine. dominated by dicot species (Ford 1977). Studies by In Wisconsin, S. faberi was shown to have resistance Birkenholz (1967) and Pinkham and Mead (1970) to sethoxydim, an acetyl-coA carboxylase-inhibitor, suggested that the optimal habitat of the harvest mouse after repeated applications. A bioassay of 10 mg L 1 was in dense stands of Setaria spp.; however, Ford sethoxydim easily discriminated between susceptible and (1977) showed no direct correlation between Setaria spp. resistant biotypes after a 6-d exposure (Retrum and and harvest mouse population and that Setaria seeds Forcella 2002). High-level resistance to sethoxydim in S. were not used as a food source. The most important faberi may be due to an alteration in the target enzyme food source for the house mouse (Mus musculus L.) in (ACCase) (Shukla et al. 1997). In the same study, the Indiana was a mixture of Setaria seeds (Whitaker 1966). resistant biotype was also less sensitive to clethodim, In the same study, the authors studied foraging habits of fenoxaprop, and quizalofop. Resistance to fluazifop-P the prairie deer mouse (Peromyscus maniculatus bairdi- and sethoxydim in a carrot, onion, and corn rotation Wagner), which occupies the same habitat of the house was reported in Wisconsin in 1991. When sprayed at the mouse; however, Setaria seeds did not make up an labelled rates, fluazifop-P provided 42% control and important component of the diet of the prairie deer sethoxydim provided 47% control of S. faberi. When mouse. the same population was sprayed with imazethapyr, Spencer and Barrett (1980) found that owing to low nicosulfuron or clethodim few to no plants survived protein content, S. faberi seeds failed to sustain body (Stoltenberg and Wiederholt 1995). In a similar study, S. weight in meadow voles [Microtus pennsylvanicus (Ord.)] faberi cross resistance to fluazifop-P-butyl was identified that were force-fed the seeds. In a study focusing on the in a vegetable cropping system in Wisconsin where the feeding preferences of the prairie vole (Microtus ochro- resistant biotype was 10.6- and 319- fold more resistant gaster Wagner), S. faberi seed were preferentially chosen than the susceptible biotype (Volenberg and Stoltenberg by the voles even though this foxtail species was not 2002a). Greenhouse experiments have shown that the common in the study area (Pascarella and Gaines 1991). outcrossing rates between ACCase susceptible and This finding was supported by Zimmerman (1965), who ACCase resistant parent plants are low and range showed that prairie vole populations in Indiana prefer- from 0.24 to 0.73% (Volenberg and Stoltenberg 2002b). entially chose S. faberi seeds as a food source even though the seeds only accounted for 1.4% of their diet. 12. Response to Other Human Manipulations Haken and Batzli (1996) found that S. faberi seeds were When mowed close to the soil surface two or more the lowest ranking food choice for the prairie vole, while times, populations of S. faberi were eliminated or the S. faberi seeds were not chosen at all by the meadow suppressed in Missouri soybean and corn fields (Donald vole in Illinois. 2000a, b, 2006; Donald et al. 2001). Shading by the crop canopy also reduced weed growth in these studies. In the (ii) Birds * McCormick and Kay (1968) reported that absence of shade, S. faberi re-grew from tillers and was birds consumed seeds of S. faberi and Sylwester (1970) still able to set seed. Mowing of S. faberi in late August, reported that geese consumed foliage of Setaria species. during its reproductive phase reduced populations to In Illinois, Kendeigh and West (1965) listed S. faberi such an extent that it was replaced by Digitaria ciliaris seeds as a food source for the tree sparrow (Spizella (Retz.) Koeler as the most dominant species in a white arborea Wilson). West (1967) further documented that a clover field in Osaka, Japan (Asai et al. 1995). mixture of S. faberi and S. viridis seeds made up 70% of The addition of sludge (6-2-0, N-P-K) or fertilizer (34- the tree sparrow’s diet even though grass seeds only 11-0, N-P-K) to old-field plant communities where represented 39% of the total seeds available to the birds. S. faberi was present caused the populations to increase Also in Illinois, cardinals (Richmondena cardinalis L.) in number and biomass, especially when moisture was a and song sparrows (Melospiza melodia Wilson) were limiting factor (Bollinger et al. 1991). both shown to prefer S. faberi seeds as a food source, especially during the spring and summer when outdoor abient temperatures were approximately 258C (Willson 13. Response to Herbivory, Disease and Higher and Harmeson 1973). Plant Parasites Although S. faberi seeds accounted for up to 12% of (a) Herbivory the total seeds available in feeding areas for Bobwhite (i) Mammals * In Minnesota, goats were given the quails (Colinus virginianus L.) in southern Illinois, only choice of 12 weed species versus alfalfa or oats for seeds of S. pumila were found in the crops of captured consumption and S. faberi was deemed unpalatable quail (Bookhout 1958). This was supported by findings (Marten and Andersen 1975). Cattle and several small in Eastern Kansas where S. pumila and S. viridis, but not 396 CANADIAN JOURNAL OF PLANT SCIENCE
S. faberi were found in the crops and stomachs of M.E. Barr], and Tilletia zundelii Hirschh. on S. faberi bobwhite quail (Jennings 1941). in the United States, as well as Ustilaginoidea virens (iii) Insects * Menalled et al. (2000) documented up to (Cooke) Takah., Ustilago crameri Ko¨ rn. and Ustilago 12% removal of S. faberi seeds in Michigan as a result of neglecta Niessl [ Macalpinomyces neglectus (Niessl) non-selective post-dispersal seed predation by birds and Va´ nky] in China. ground beetles. In Iowa, invertebrate seed predators In Iowa, seed-borne fungi were studied in S. faberi [mainly Gryllus pennsylvanicus Burmiester (Orthoptera:- and S. viridis from fields in corn-soybean rotation by Gryllidae)] removed 30 to 90% of seeds in corn and Pitty et al. (1987). Fungal colonization of seeds was soybean crops between the months of July and Septem- investigated under conventional tillage and reduced ber (O’Rourke et al. 2006). Westerman et al. (2006) tillage systems when harvested by hand from the found that up to 67 and 80% of seeds were removed by inflorescences compared with seeds recovered from post-dispersal soil predators when S. faberi grew in a various soil depths. Extent of colonization under all corn or soybean field, respectively. Setaria faberi was treatments was greater in S. faberi than S. viridis, highly preferred as an oviposition host and food source although it was not determined whether this was for the moths Hydraecia immanis Guene´ e (Lepidoptera: attributable to differential susceptibility or correlation Noctuidae) and Papaipema nebris Guene´ e (Lepidoptera: with seed size. More than nine species of fungi were Noctuidae) in Illinois (Levine 1985, 1986) and as a host isolated from colonized seeds (22%) on the inflores- for P. nebris in Wisconsin (Alvarado et al. 1989). cences, the most common of which were Cochliobolus In greenhouse studies, the western corn rootworm carbonum R.R. Nelson, Epicoccum nigrum Link ( E. [Diabrotica virgifera virgifera LeConte (Coleoptera: purpurascens Ehrenb.) and Fusarium arthrosporioides Chrysomelidae)] had good survivorship on S. faberi Sherb. More than 16 species of fungi were isolated and may have chosen this grass as an alternate host to from buried seeds, the most common of which were corn (Chege et al. 2005; Oyediran et al. 2005). Clark and Alternaria alternata (Fr.) Keissl., E. nigrum and several Hibbard (2004) tested 28 species of grasses for non-corn unidentified sterile strains. At shallow soil depths (0 7.5 host suitability for western corn rootworm and S. faberi cm) S. faberi seeds showed greater colonization in the ranked 13th out of the 28 species, but was shown to be a reduced tillage system (60% vs. 32%), while in the less suitable host than S. viridis. In corn fields, S. faberi conventional tillage system colonization increased with may also be a suitable host for the northern corn soil depth (63% at 7.5 15 cm, 70.8% at 15 22.5 cm). Rootworm [Diabrotica barberi Smith & Lawrence (Co- The authors speculated that these colonization patterns leptera: Chrysomelidae)] (Oyediran et al. 2004). are related to different moisture and organicmatter Potato leafhopper [Empoasca fabae (Harris) (Homo- profiles in the two systems. An Endogone spp. found in ptera: Cicadellidae)] nymphs did not survive on S. faberi Indiana cultivated fields was found to colonize on and in in Illinois (Lamp et al. 1984). Similarily, the Hessian fly the seeds of several plant species; however, the fungus [Mayetiola destructor Say (Diptera: Cecidomyiidae)] was was not found on any S. faberi seeds that were examined unable to reproduce when S. faberi was presented as a (Houtcooper 1972). Davis and Renner (2007) documen- possible host (Zeiss et al. 1993). ted that fatal germination caused by soil-borne patho- genicfungi was non-existent for S. faberi seeds (iv) Nematodes and/or Other Non-vertebrates * Wong in Michigan soil that was inoculated with Pythium and Tylka (1994) showed that S. faberi was not a host ultimum. for the soybean cyst nematode (Heterodera glycines Cropping system was shown to influence S. faberi Ichinohe) in Iowa. seed mortality due to fungal community changes (Davis et al. 2006). The authors showed that seed mortality was highest in crop rotations that were conventionally (b) Diseases managed versus those where a reduced input approach (i) Fungi * Farr et al. (undated) give references to weed management was adopted. reporting Bipolaris setariae (Sawada) Shoemaker [ In Saskatoon, a potential bioherbicide, Pyricularia Cochliobolus setariae (S. Ito & Kurib.) Dastur], Colleto- setariae Niskada was tested and found to have low trichum graminicola (Ces.) G.W. Wilson ( Glomerella pathogenicity on S. faberi. When the fungus was paired graminicola D.J. Politis), Curvularia lunata (Wakker) with 0.25 the labelled rate of the herbicide sethoxydim Boedijn ( Cochliobolus lunatus R.R. Nelson & Haa- (labelled rate 200 g a.i. ha 1) there was a 10-fold sis), Fusarium equiseti (Corda) Sacc. ( Gibberella increase in the pathogenic infection of S. faberi, and a intricans Wollenw.), Nigrospora oryzae (Berk. & 100% increase in weed control in comparison to the Broome) Petch ( Khuskia oryzae H.J. Huds.), Nigros- pathogen alone (Peng and Byer 2004). pora sphaerica (Sacc.) E.W. Mason ( K. oryzae), Phaeosphaeria luctuosa (Sacc.) Otani & Mikawa, Phoma (ii) Bacteria *The deleterious rhizobacteria strains sorghina (Sacc.) Boerema, Dorenb. & Kesteren ( Aeromonas hydrophila, Aeromonas caviae, Agrobacter- Leptosphaeria sacchari Breda de Haan), Pyricularia ium radiobacter, Chromobacterium violaceum, Chryseo- grisea Sacc. [ Magnaporthe grisea (T.T. Hebert) monas luteola, Pseudomonas aureofaciens, Pseudomonas NURSE ET AL. * SETARIA FABERI HERRM. 397 putida, Pseudomonas fluorescens, and Xanthomonus Armel, G. R., Wilson, H. P., Richardson, R. J. and Hines, T. E. maltophilia were isolated from S. faberi roots in 2003d. Mesotrione alone and in mixtures with glyphosate in Missouri (Li and Kremer 2000, 2006). Li and Kremer glyphosate-resistant corn (Zea mays). Weed Technol. 17: 680 (2000) reported that the deleterious effects of the 685. aforementioned isolates were similar for both S. faberi Asai, M., Ito, M. and Kusanagi, T. 1995. Effects of mowing regimes on the growth and vegetation dynamics of established and S. viridis and Li and Kremer (2006) showed that the white clover (Trifolium repens L.) cover for weed suppression. rhizobacteria caused reductions of up to 77.2 and 79.1% Weed Res. (Japan) 40: 194 202. for root and shoot mass, respectively. Ateh, C. M. and Harvey, R. G. 1999. Annual weed control by glyphoste in glyphosate-resistant soybean (Glycine max). Weed (iii) Viruses * Brunt et al. (1996) refer to studies Technol. 13: 394 398. reporting susceptibility of S. faberi to cereal northern Baskin, C. C. and Baskin, J. M. 1998. Seeds: Ecology, mosaic cytorhabdovirus, maize chlorotic dwarf waika- biogeography, and evolution of dormancy and germination. virus, maize streak monogeminivirus and maize white Academic Press, New York, NY. 666 pp. line mosaic virus, but not susceptible to cereal northern Bazzaz, F. A. and Garbutt, K. 1988. The response of annuals in mosaiccytorhabdovirus. Setaria faberi was reported to competitive neighborhoods: effects of elevated CO2. Ecology be a suitable host for wheat streak mosaicvirus in 69: 937 946. Kansas (Christian and Willis 1993). Beckett, T. H., Stoller, E. W. and Bode, L. E. 1992. Quizalofop and sethoxydim activity as affected by adjuvants and ammo- nium fertilizers. Weed Sci. 40:12 19. (c) Higher Plant Parasites * No information was found. Bell, D. T. and Koeppe, D. E. 1972. Non competitive effects of giant foxtail on the growth of corn. Agron. J. 64: 321 325. Benabdelmouna, A., Abirached-Darmency, M. and Darmency, ACKNOWLEDGEMENTS H. 2001a. Phylogeneticand genomicrelationships in Setaria The authors thank Lorraine Smith and Lisa Boughner italica and its close relatives based on the molecular diversity at AAFC (Harrow) for their help locating articles for and chromosomal organization of 5S and 18S-5.8S rDNA the literature review. Barry Flahey of Agriculture and genes. Theor. Appl. Genet. 103: 668 677. Agri-Food Canada produced the drawings in Fig. 1 Benabdelmouna, A., Shi, Y., Abirached-Darmency, M. and and 2. We acknowledge the generous assistance of the Darmency, H. 2001b. Genomicin situ hybridization (GISH) herbarium curators who provided loans of specimens for discriminates between the A and the B genomes in diploid and study and distribution mapping (see caption to tetraploid Setaria species. Genome 44: 685 690. Fig. 3). Beyers, J. T., Smeda, R. J. and Johnson, W. G. 2002. Weed management programs in glufosinate-resistant soybean (Gly- Abul-Fatih, H. A. and Bazzaz, F. A. 1979. The biology of cine max). Weed Technol. 16: 267 273. Ambrosia trifida L. I. Influence of species removal on the Bigwood, D. W. and Inouye, D. W. 1988. Spatial pattern organization of the plant community. New Phytol. 83: 813 analysis of seed banks: an improved method and optimized 816. sampling. Ecology 69: 497 507. Ackley, J. A., Wilson, H. P. and Hines, T. E. 1996. Efficacy of Biniak, B. M. and Aldrich, R. J. 1986. Reducing velvetleaf rimsulfuron and metribuzin in potato (Solanum tuberosum). 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Yoshioka, T., Ota, H., Segawa, K., Takeda, Y. and Esashi, Y. out, the hairs without swollen bases; leaf sheath 1995. Contrasted effects of CO2 on the regulation of dormancy margins fringed with hairs, not keeled; spikelets and germination in Xanthium pennsylvanicum and Setaria 1.9 3 mm long ...... 2 faberi seeds. Ann. Bot. (Lond.) 76: 625 630. Zeiss, M. R., Brandenburg, R. L. and van Duyn, J. W. 1993. Suitability of seven grass weeds as Hessian fly (Diptera 2. Spikelets toward the base of panicle in spaced clusters Cecidomyiidae) hosts. J. Agric. Entomol. 10: 107 119. or verticils, with the rachis visible in between; leaf Zhang, J., Hamill, A. S., Gardiner, I. O. and Weaver, S. E. blades with a few hairs on upper surface; bristles 1998. Dependence of weed flora on the active soil seedbank. 4 7 mm long; bristles, rachis and upper culm Weed Res. 38: 143 152. with downwardly (or rarely upwardly) pointing Zimdahl, R. L. 1980. Weed crop competition: a review. Oregon barbs ...... S. verticillata State University, Corvallis, OR. 220 pp. Zimmerman, E. G. 1965. A comparison of habitat and food of 2. Spikelets not (or rarely) in spaced clusters on rachis; two species of Microtus. J. Mammal. : 605 612. 46 leaf blades hairy or hairless on upper surface; bristles Ziska, L. H. and Bunce, J. A. 1997. Influence of increasing carbon dioxide concentrations on the photosynthetic and 5 12 mm long; bristles, rachis and upper culm with growth stimulation of selected C4 crops and weeds. Photo- upwardly pointing barbs ...... 3 synth. Res. 54: 199 208. 3. Fertile floret separating from glumes and sterile lemma at maturity; panicle (excluding bristles) 7 15 Appendix A mm diam., cylindrical but often lobed or interrupted; fertile lemmas smooth and shiny, occasionally ob- Key to the species of Setaria in Canada: scurely transversely wrinkled ...... S. italica
1. Bristles below each spikelet 5 or more, orange 3. Entire spikelet falling at maturity leaving a cup-like or tawny; upper glume covering about half the scar on the pedicel; panicle (excluding bristles) 4 coarsely transversely wrinkled fertile lemma; upper 8(10) mm diam., narrowly cylindrical; fertile lemmas surface of leaf blade with a few long (greater than 4 finely or coarsely transversely wrinkled, not shiny 4 mm) hairs near the base, the hairs with swollen bases; leaf sheaths smooth on margins, keeled; 4. Spikelets 1.9 2.2(2.4) mm long; upper glume almost spikelets (2)3 3.4 mm long ...... S. pumila completely covering the fertile lemma; leaf blades smooth or scabrous ...... S. viridis 1. Bristles below each spikelet 1 3 (sometimes to 6 in S. faberi), green or purplish; upper glume covering 4. Spikelets 2.4 3 mm long; upper glume covering about 3/4 or more of the minutely transversely wrinkled or three-quarters of the fertile lemma; leaf blades usually smooth fertile lemma; upper surface of leaf blade evenly covered with soft hairs at least on the upper smooth or with shorter hairs (0.5 3.5 mm) through- surface ...... S. faberi