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The Foxtail (Setaria) Species-Group

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The Foxtail (Setaria) Species-Group Invited Review The Foxtail (Setaria) Species-Group Jack Dekker, Weed Biology Laboratory, Agronomy Department, Iowa State University, Ames, IA 50011 USA; [email protected] Abstract. The weedy Setaria species (giant, green, yellow, knotroot and bristly foxtail) compose one of the worst weed groups interfering with world agriculture, and in other disturbed and managed habitats. These five weed species, together with their crop counterparts (foxtail millet, korali), form the foxtail species-group. Five successive waves of Setaria spp. invasion from pre-agricultural times to the present have resulted in widespread infestation of the disturbed, arable, temperate regions of the earth. These invasions have resulted in considerable economic and environmental costs. The success of the Setaria species-group is due to their intimate evolutionary relationship with humans, disturbance, agriculture and land management. The ability to rapidly adapt to local conditions is the hallmark of this weedy group. Genotypic and phenotypic biodiversity provide this species-group with traits that allow them to invade, colonize, adapt to, and endure in a wide range of habitats around the world. The phenotypic life history traits important to weedy Setaria spp. success begin with the induction of dormancy in seed during embryogenesis. The formation of long-lived, heterogeneous seed pools in the soil is the inevitable consequence of the dormant seed rain. In soil seed pools, after-ripening, the occurance and timing of seedling emergence, and the induction of secondary (summer) dormancy are regulated by seasonally and diurnally varying soil oxygen, water and temperature signals. Precise and variable timing of seedling emergence ensures Setaria a dominant place in disturbed and managed communities during the growth and reproductive phases that follow. Once established in a community, phenotypic plasticity inherent in an individual weedy Setaria sp. plant allow it to maximize its growth, form and reproduction to the specific local conditions it encounters, including competitive interactions with neighbors. Traits controlling the plastic development of plant architecture include the ability to form one or more tillering shoots whose stature and number are precisely sized to local conditions. A complex pattern of branching, from plant to spikelet, provides diverse microenvironments within which different levels of dormancy are induced in individual seeds on a panicle, and among panicles on a common plant. Traits for adaptation to stress in weedy Setaria spp. include tolerance to many inhibitory chemicals (e.g. herbicides, salt), mechanical damage and drought. Genetic traits such as self-pollenation and small genome size contribute to a highly diverse collection of locally adapted genotypes and phenotypes ready to exploit any opportunities provided by a cropping system. Self-pollenating Setaria spp. exist in wild, weed and crop variants, an ideal genetic condition ensuring both long- term stability and novelty. Weedy Setaria spp. populations have low to exceedingly low amounts of total genetic variation, unusually low intra-population genetic diversity, and unusually high genetic diversity between populations compared to an "average" plant species. These traits result spatially in local populations that are unusually homogeneous, typically consisting of a single multilocus genotype. Either a generally- or specifically-adapted genotype of an individual Setaria species might predominate in that local population. Across the landscape different single-genotype populations dominate particular local sites, providing novel genetics to the region via dispersal and gene flow when conditions change. Across North America, populations of S. viridis and S. geniculata are genetically differentiated along a north- 1 Invited Review south gradient. The past history of invasion and colonization, the successful life histories of locally adapted weedy Setaria spp., and the evolutionary potential of this weed group emphasize the need for accurate predictions of its behavior. Weedy Setaria spp. management is the mangement of local selection pressure and the consequential adaptation. Farmers, land managers, policy-makers and regulators, homeowners and consumers need accurate information about weedy Setaria spp. to predict and guide management decisions based on economics, risk and environmental sustainability. Nomeclature: green foxtail , Setaria viridis, subspecies viridis (L.) Beauv. SETVI; foxtail millet, Setaria viridis, subspecies italica) SETIT; yellow foxtail, Setaria glauca (Weigel) Hubb. SETLU; giant foxtail, S. faberii Herrm. SETFA; bristly foxtail, Setaria verticillata (L.) Beauv. SETVE; knotroot foxtail, S. geniculata SETGE INTRODUCTION The foxtails are members of the Setaria genus and are one of the worst weed groups interfering with North American and world agriculture and land management (Holm et al., 1977, 1979, 1997). The weedy Setaria spp. include S. viridis subspecies viridis (L.) Beauv. (green foxtail), S. glauca (Weigel) Hubb. (yellow foxtail), S. faberii Herrm. (giant foxtail), S. verticillata (L.) Beauv. (bristly foxtail), and S. geniculata (Lamarck) Beauv. (knotroot foxtail) (Pohl, 1951, 1966; Rominger, 1962). The genus Setaria also contains the crop foxtail millet (S. viridis subsp. italica) whose geographic distribution and evolutionary history is intimately connected with the weedy members of Setaria (de Wet, 1979, 1995). Therefore, a review of the agricultural Setaria appropriately begins at the level of the Setaria species-group (a group of closely related species, usually with partially overlapping ranges; Lincoln et al., 1998), a wild-crop-weed complex (de Wet, 1966). Herein the term Setaria species-group (spp.-gp.) will be used when referring to properties shared by the five foxtail weed species and foxtail millet. Reviews of Setaria spp.-gp. have been published previously (S. glauca and S. verticillata, Steel et al., 1983; S. viridis, Douglas et al., 1985; Setaria spp.-gp., Dekker, 2003). The success of the Setaria spp.-gp. is due to their intimate evolutionary relationship with humans, disturbance, agriculture, and land management. Genotypic and phenotypic biodiversity provide this species-group with traits that allow them to invade, colonize, adapt to and endure in a wide range of disturbed habitats in temperate, tropical, and sub-tropical regions. Why have the weedy Setaria spp. spread globally over the last 5-9000 years? How did they get to be such a major weedy pest? What are the specific traits that allow them to dominate agricultural fields? Why is Setaria still a dominant species-group in North American agriculture after continuous use of highly effective herbicides for almost 50 years? What will the Setaria species-group do in the future? What do we know, not know, and need to know to manage them? The answer to these questions begins with the history of global invasion of the Setaria spp.-gp. Subsequent success of the foxtails is revealed by their local adaptation. The consequences of this local adaptation and evolution are apparent in its life history. The conclusions implied by Setaria spp.-gp. invasion, local adaptation and life history could advise future management systems. 2 Invited Review HISTORY OF GLOBAL INVASION `There appears to be five major phases, or waves, of wild-weed-crop Setaria spp. invasion in earth's history. The origin of the genus and the original wild Eurasian Setaria sp. was probably Africa, based on the large number of Setaria spp. from there (74 of 125), as well as genomic evidence of tropical origins (Lakshmi and Ranjekar, 1984; Prasado Rao, 1987; Rominger, 1962; Simpson, 1990; Stapf and Hubbard, 1930). The tropical genus Setaria dispersed to Eurasia and adapted to a wider range of environmental conditions and habitats. In the second phase, a S. viridis weedy progenitor species spread over Eurasia as a wild colonizing species. The wild, progenitor Setaria spp. that invaded Eurasia in many locations after leaving its African home was most likely a diploid annual similar to S. viridis, subsp. viridis (Kihara and Kishimoto, 1942; Koernicke and Werner, 1885a, b; Li, 1934; Li et al., 1942, 1945; Prasado Rao et al., 1987; Rominger, 1962; Simpson, 1990; Werth, 1937; Willweber-Kishimoto, 1962). Speciation and adaptation followed this spread in geographic distribution to subtropical, and then temperate, regions. Parallel to this, with possibly more ancient antecedants, S. glauca arose and spread throughout China. With domestication, both S. viridis and S. glauca then spread as a weed and a crop (foxtail millet and korali). Foxtail millet was a crop in China 6000 years B.P. (Before Present) (Cheng, 1973; Li and Wu, 1996; Naeiri and Belliard, 1987), while its use as a crop in Europe dates to 3600 B.P. (Dembinska, 1976; Helbaek, 1960; Neuweiler, 1946; de Wet and Harlan, 1975). Setaria glauca seed were gathered from wild plants and later cultivated in India (de Wet et al., 1979; de Wet, 1992). Other weedy Setaria species arose from S. viridis-like ancestors (Khosla and Sharma, 1973), including polyploid specialists such as S. verticillata, S. verticillata and S. faberii. In the third wave, weedy Setaria spp. spread to the New World at two different times: pre-Columbian (before ca. 1500 A.D.) and post-Columbian invasions (after ca. 1500 A.D.). The pre-Columbian origins of S. geniculata, the only weedy Setaria native to the New World, suggest an ancient dispersal event eastward from the Eurasia to the Americas (Rominger,
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