Creeping bentgrass (Agrostis stolonifera) has been utilized as a forage species for centuries, and as a turf species for many decades. It now occurs worldwide in temperate areas. However, basic information on Agrostis stolonifera from the non-agronomic biological and ecological literature had not previously been consolidated. Consequently, this white paper presents a general overview on Agrostis stolonifera biology and ecology as a native and naturalized species, with certain aspects focusing on the United States and Canada. This synthesis and summary provide perspective on Agrostis stolonifera and a pathway to the diverse literature 1 , to advance understanding, research and application.

 or by not realizing the population has more than one species. 1.5. For example, there is a need in USA for increased awareness to better distinguish A. capillaris and A. stolonifera from A. castellana (e.g.

Purpose

This white paper is intended to provide background scientific knowledge on the biology and ecology of Agrostis stolonifera. No overview of the basic biology and ecology of the species was available prior to this undertaking.

Context

Agrostis stolonifera (creeping bentgrass) is the species most utilized for golf courses in temperate areas worldwide. A strain has been genetically engineered to be tolerant to the herbicide glyphosate, in order to control especially the weedy species Poa annua (annual bluegrass) on golf courses. The USDA's Animal and Plant Health Inspection Service (APHIS) has been petitioned to grant non-regulated status to this glyphosate-tolerant strain.

Glyphosate is the most extensively used herbicide worldwide, and Agrostis stolonifera is an outcrossing perennial widespread in managed and natural environments. It hybridizes with many species of Agrostis (bentgrass) and Polypogon (rabbitsfoot grass) found in the United States. Creeping bentgrass and several of the hybridizing species can be weedy or invasive in some situations. The white paper is a summary document, which is heavily referenced, to provide a scientific overview on biology and ecology and a guide to the non-agronomic information about Agrostis stolonifera.

History

As part of the regulatory review process on the petition, a December 2003 version of the "Perspective" comprised much of the APHIS Biotechnology Regulatory Services (BRS) preliminary risk assessment, which APHIS-BRS has referenced in several Federal Register notices relating to the petition. The Perspective's scientific information played an important part in the APHIS-BRS decision to prepare an Environmental Impact Statement (EIS) under the National Environmental Policy Act to assess potential impacts from granting the petition and has been useful for some aspects of the EIS drafting.

Overview of the Genus Agrostis and its Species

1.1. Some field grasses were called Agrostis by Theophrastus (ca. 371-287 BC), the director of Aristotle's garden in Athens [START_REF] Greene | Landmarks of Botanical History, Parts 1 and 2[END_REF]. Bentgrasses (Agrostis) are in the convenient grouping called cool-season grasses, with a C 3 photosynthetic pathway (cf. [START_REF] Campbell | Climate profiles of temperate C 3 and subtropical C 4 species in New Zealand pastures[END_REF][START_REF] Goverde | Species-specific reactions to elevated CO 2 and nutrient availability in four species[END_REF]. Within the family Poaceae (Gramineae), the genus Agrostis is in the tribe Poeae (including Aveneae and Agrostideae), which also contains oats (Avena), and in the subtribe Agrostidinae (Soreng et al. 2003;cf. Clayton & Renvoize 1986;[START_REF] Watson | The Grass Genera of the World[END_REF], 1998, 1999;[START_REF] Mabberley | The Plant-Book: A Portable Dictionary of the Higher Plants[END_REF]. The genus might include 150-200 species, occurring primarily and probably originating evolutionarily in the Northern Hemisphere in western Eurasia (the European region), with some species having evolved in the Southern Hemisphere and temperate to coldtemperate areas of tropical mountains. The generic limits of Agrostis are not obvious (e.g. [START_REF] Phillips | Notes on grasses (Poaceae) for the Flora of China, I: Deyeuxia[END_REF][START_REF] García | The F 1 hybrids of A. stolonifera and A. capillaris[END_REF], and sometimes Podagrostis [START_REF] Björkman | Studies in Agrostis and Related Genera[END_REF], Bromidium (Rúgolo de [START_REF] Rúgolo De Agrasar | Revalidación del género Bromidium Nees et Meyen emend. Pilger (Gramineae)[END_REF] and Lachnagrostis (Edgar 1995;[START_REF] Jacobs | The genus Lachnagrostis (Gramineae) in Australia[END_REF]) are considered distinct (Soreng et al. 2003). The noun "bent" is a Middle English word for a grassy place and/or its grasses (from Old English "beonot-"); broad secondary meanings include a reedy grass or stiff stalk of coarse grass [START_REF] Mw | Merriam-Webster's Collegiate Dictionary, 11th Ed[END_REF]. As a common name, various species in more than a dozen genera of grasses have been called bents (e.g. reed bent, silky bent), and as well a sedge (Carex) and a bulrush (Scirpus), but bent has been associated especially with species of the genus Agrostis since at least 1677 [START_REF] Piper | Bent[END_REF][START_REF] Mw | Merriam-Webster's Collegiate Dictionary, 11th Ed[END_REF].

1.2. In the United States, 35 species of Agrostis in the broad sense (sensu lato) are native or naturalized, with 17-19 of them also in Canada (see Table 1). There are 26-29 native species of Agrostis in the USA, and 7-9 established introductions, mainly from Eurasia (7 or 8 of these species are entirely introduced, 1 or 2 mostly so). If the genus is instead considered in the narrow sense (sensu stricto) (Soreng & Peterson 2003), two of the native species would be placed in Podagrostis (Grisebach) Scribner & Merrill and one each of the introduced species in Bromidium Nees & Meyen and Lachnagrostis Trinius (Table 1). Overall taxonomy within the genus Agrostis is unsettled and difficult. There is no comprehensive worldwide or definitive U.S. taxonomic treatment (cf. Hitchcock & Chase 1951;[START_REF] Carlbom | A Biosystematic Study of Some North American Species of Agrostis L. and Podagrostis[END_REF][START_REF] Simpson | A Study of Species Complexes in Agrostis and Bromus[END_REF][START_REF] Harvey | Agrostis, Bent grass[END_REF][START_REF] Harvey | Agrostis L[END_REF][START_REF] Kartesz | Synthesis of the North American Flora[END_REF]), but there are some thorough floristic treatments and a few definitive regional taxonomic studies (e.g. [START_REF] Philipson | A revision of the British species of the genus Agrostis Linn[END_REF][START_REF] Björkman | Studies in Agrostis and Related Genera[END_REF][START_REF] Widén | The Genus Agrostis in Eastern Fennoscandia. Taxonomy and Distribution[END_REF]Romero García et al. 1988b;[START_REF] Tutin | Agrostis L[END_REF][START_REF] Rajbhandari | The genus Agrostis in Nepal, 1[END_REF][START_REF] Rajbhandari | The genus Agrostis in Nepal, 2[END_REF][START_REF] Koyama | Grasses of Japan and its Neighboring Regions: An Identification Manual[END_REF]Edgar & Connor 2000;[START_REF] Rúgolo De Agrasar | Las especies del género Agrostis (Gramineae: Agrosteae) de la Argentina[END_REF], 1997a). Consequently, the number of North American species stated above is a distillation reflecting different judgements. 1.3. Cytological study of chromosomes (karyology) along with experimental hybridization have provided central taxonomic insights, as polyploidy (multiplication of the basic set of paired chromosomes) plays a significant role in the evolution of the genus and within some species (e.g. [START_REF] Sokolovskaya | A caryo-geographical study of the genus Agrostis[END_REF][START_REF] Davies | The breeding affinities of some British species of Agrostis[END_REF][START_REF] Jones | The cytology of some British species of Agrostis and their hybrids[END_REF]Jones , 1956aJones -1956c;;[START_REF] Björkman | Chromosome studies in Agrostis. (A preliminary report[END_REF][START_REF] Björkman | Chromosome studies in Agrostis[END_REF][START_REF] Björkman | Studies in Agrostis and Related Genera[END_REF][START_REF] Carlbom | Premeiotic reduction in Agrostis L., Potentilla L., and Triticum L[END_REF][START_REF] Vovk | Khromosomni chisla vidiv ta form ukrainskikh mitlic (rid Agrostis L.) [Chromosome numbers of Ukrainian Agrostis L. species and forms][END_REF][START_REF] Widén | The Genus Agrostis in Eastern Fennoscandia. Taxonomy and Distribution[END_REF][START_REF] Sokolovskaya | Kariosistematicheskoye issledovanie Dal'nego Stochnykh vidov Agrostis L. [Karyosystematic investigation of the Far Eastern species of Agrostis L.][END_REF][START_REF] Tateoka | A contribution to the taxonomy of the Agrostis mertensii-flaccida complex (Poaceae) in Japan[END_REF][START_REF] Tateoka | Agrostis mertensii (Poaceae) in Hokkaido[END_REF][START_REF] Rumball | Plant introduction trials. Performance of Agrostis species at Palmerston North[END_REF]Romero García & Blanca 1988;Romero García et al. 1988a;Frey 1997;[START_REF] Tomić | Dry matter and seed production in some inbred lines of three Agrostis species. Herbage Seed as a Key Factor for Improving Production and Environmental Quality[END_REF][START_REF] Tomić | Genetic resource of five Agrostis species present in Serbia and their utilisation[END_REF][START_REF] Bonos | Ploidy determination in Agrostis using flow cytometry and morphological traits[END_REF][START_REF] Vergara | AFLP analyses of genetic diversity in bentgrass[END_REF]. Substantial taxonomic insight also has come from the study of floret anatomy and histology (e.g. [START_REF] Björkman | Studies in Agrostis and Related Genera[END_REF]Romero García et al. 1988b). Most species tend to be accepted as distinct, i.e. not controversial taxonomically, although many species have received little scientific attention. The general stability includes all five of the species of main agronomic interest (Tables 1 &2) [START_REF] Beard | Beard's Turfgrass Encyclopedia for Golf Courses, Grounds, Lawns, Sports Fields[END_REF][START_REF] Aldous | Sports Turf and Amenity Grasses: A Manual for Use and Identification[END_REF]):

• creeping bentgrass, Agrostis stolonifera [START_REF] Warnke | Creeping bentgrass (Agrostis stolonifera L.)[END_REF]); • colonial bentgrass, Agrostis capillaris [START_REF] Ruemmele | Agrostis capillaris (Agrostis tenuis Sibth.) colonial bentgrass[END_REF]); • velvet bentgrass, Agrostis canina [START_REF] Brilman | Velvet bentgrass (Agrostis canina L.)[END_REF][START_REF] Brilman | Velvet bentgrass: Rediscovering a misunderstood turfgrass[END_REF]; • dryland (including Highland) bentgrass, Agrostis castellana (Brede & Sellmann 2003); • redtop, Agrostis gigantea [START_REF] Brede | Three minor Agrostis species: Redtop, highland bentgrass, and Idaho bentgrass[END_REF][START_REF] Balasko | Bluegrasses, ryegrasses, and bentgrasses[END_REF].

1.4. Nevertheless, using the subtle biological distinctions frequently requires acumen and expertise for identification of the species of whole plants, whether in flower or not (e.g. [START_REF] Widén | The Genus Agrostis in Eastern Fennoscandia. Taxonomy and Distribution[END_REF][START_REF] Tutin | Agrostis L[END_REF][START_REF] Steiner | Zur Echtheitsbestimmung mittels morphologischer Merkmale der Spelzfrüchte bei Agrostis-Arten [Verification of species of Agrostis by means of morphological characters of the florets[END_REF][START_REF] Steiner | Zur Reproduzierbarkeit quantitativer Messungen an Spelzfrüchten bei der Echtheitsbestimmung in der Saatgutprüfung (Agrostis spp., Festuca sssp. [sic]) [Reproducibility of quantitative measurements of florets for verification in seed testing (Agrostis spp., Festuca sssp.)][END_REF]Romero García et al. 1988b;[START_REF] Chicouène | Note sur les Agrostis du Massif Armoricain: Identification, écologie, répartition[END_REF][START_REF] Hartvig | Agrostis L[END_REF]Sell & Murrell 1996;[START_REF] Copping | Identification of Agrostis species in Inverness-shire. Inverness-shire Surv[END_REF][START_REF] Cope | Problems with bents -identifying British Agrostis[END_REF][START_REF] Adams | Grass gripes[END_REF]Batson 1998a;Behrendt & Hanf 1979;[START_REF] Nittler | Seedling difference among Agrostis species and varieties[END_REF], or the seeds [START_REF] Hillman | The seeds of redtop and other bent grasses, Part II in C.V. Piper and F.H. Hillman, The Agricultural Species of Bent Grasses[END_REF][START_REF] Hillman | Identifying turf-grass seed[END_REF][START_REF] Musil | Diagnostic characters of seed of the commercial species of Agrostis and certain species occurring incidentally with them[END_REF][START_REF] Musil | Identification of Crop and Weed Seeds[END_REF][START_REF] Van Der Burg | Identification of Agrostis spp[END_REF]. Identification can be facilitated by lab analysis (e.g. [START_REF] Yamamoto | Electrophoretic identification of cross-pollinated bentgrass species and cultivars[END_REF][START_REF] Clark | Identification of cultivars of Agrostis species by polyacrylamide gel electrophoresis of seed proteins[END_REF][START_REF] Freeman | Esterase isoenzyme electrophoresis as a method of separating colonial and creeping bentgrass mixtures[END_REF][START_REF] Ohmura | Identification of Poa pratensis L. and Agrostis spp. cultivars by random amplified polymorphic DNA (RAPD) markers[END_REF][START_REF] Hollman | Identification of putative velvet bentgrass clones using RAPD markers[END_REF][START_REF] Scheef | Development of species-specific SCAR markers in bentgrass[END_REF][START_REF] Ridgway | Identification of roots from grass swards using PCR-RFLP and FFLP of the plastid trnL (UAA) intron[END_REF][START_REF] Bonos | Ploidy determination in Agrostis using flow cytometry and morphological traits[END_REF]. Routinely, diverse studies either lack full identification, 1.6. Agrostis is additionally complex in naturally forming F 1 (first-generation) hybrids, which are generally low in fertility or sterile but sometimes vegetatively vigorous, and occur perhaps not infrequently between various species. Such hybrids varyingly include all five of the main agronomic species (cf. [START_REF] Brede | Three minor Agrostis species: Redtop, highland bentgrass, and Idaho bentgrass[END_REF][START_REF] Warnke | Creeping bentgrass (Agrostis stolonifera L.)[END_REF], as well as Agrostis idahoensis, Idaho bentgrass (of agronomic interest - [START_REF] Brede | Three minor Agrostis species: Redtop, highland bentgrass, and Idaho bentgrass[END_REF][START_REF] Brede | GolfStar', a turf, ornamental, and reclamation cultivar of North American native Idaho bentgrass[END_REF]), A. vinealis, and several other germane species (see Tables 1 & 2, and Davies 1953;Jones 1956aJones -1956c;;[START_REF] Camus | Graminées hybrides de la flore française (genre Bromus excepté)[END_REF][START_REF] Björkman | Studies in Agrostis and Related Genera[END_REF][START_REF] Widén | The Genus Agrostis in Eastern Fennoscandia. Taxonomy and Distribution[END_REF]Bradshaw 1975aBradshaw , 1975b;;Sell & Murrell 1996;[START_REF] Lambinon | Agrostis ×fouilladeana Lambinon & Verloove, nothospec. nov., en Belgique et dans le nord de la France[END_REF][START_REF] Paunero | Las especies españolas del género Agrostis[END_REF]Romero García et al. 1988b;[START_REF] García | The F 1 hybrids of A. stolonifera and A. capillaris[END_REF]Edgar & Connor 2000;Batson 1998a;[START_REF] Wipff | Determining gene flow of transgenic creeping bentgrass and gene transfer to other bentgrass species[END_REF], 2001;Wipff & Fricker 2001;Belanger et al. 2003cBelanger et al. , 2003b;;Christoffer 2003). There are a few clear examples of F 2 hybrids (i.e. the second generation, F 1 hybrids crossed with each other) (Bradshaw 1975a;Sell & Murrell 1996), and of backcrosses of F 1 hybrids to a parental species (Forde 1991;Edgar & Connor 2000;Batson 1998a;Belanger et al. 2003b). In some areas where the Agrostis species are native or naturalized, the hybrids are recognized as sometimes being a significant component of vegetation [START_REF] Widén | The Genus Agrostis in Eastern Fennoscandia. Taxonomy and Distribution[END_REF]Bradshaw 1975a;Sell & Murrell 1996;Edgar & Connor 2000). This composition can occur even with sterile plants because of their vegetative reproduction. 1.7. A summary on the pertinent hybridizations is in Figure 1 and Table 1. In Table 2 are basic details (not all-encompassing) on these species in the USA and their crossing (which is inherently complicated but not atypical in grasses and some other wild plants), along with additional information as follows:

• Notable botanical synonyms (thus subordinate, not just alternative names); • Common names (cf. [START_REF] Monteith | Classification of redtop and the common bent grasses[END_REF][START_REF] Usga | When is a bent grass a creeping bent?[END_REF]); • Ranges (where native, and along with the Table 1 information, where naturalized);

• Ploidy (a polyploid has 3 or more sets of chromosomes; frequent levels are bolded) (cf. [START_REF] Levy | The impact of polyploidy on grass genome evolution[END_REF][START_REF] Gaut | Evolutionary dynamics of grass genomes[END_REF][START_REF] Soltis | Advances in the study of polyploidy since Plant Speciation[END_REF]: n = x = 7, the basic set of chromosomes in a male or female gamete, haploid 2n = 2x = 14, the paired parental sets of chromosomes usual in body cells, diploid 2n = 3x = 21, three sets of chromosomes usual in the plant's body cells, triploid 2n = 4x = 28, four sets of chromosomes usual in plant's somatic cells, tetraploid: allotetraploid (amphidiploid) = 2 unlike paired sets, AABB (e.g. hybrid's sets doubled) autotetraploid = 4 sets alike, AAAA (e.g. within sp., crossing plants' sets doubled) segmental allotetraploids = ~ alike, ± halfway, ~ unalike: e.g. AAĀĀ, AAÃÃ, AAẬẬ 2n = 5x = 35, five sets of chromosomes usual in plant's somatic cells, pentaploid 2n = 6x = 42, six sets of chromosomes usual in plant's somatic cells, hexaploid 2n = 7x = 49, seven sets of chromosomes usual in plant's somatic cells, heptaploid 2n = 8x = 56, eight sets of chromosomes usual in plant's somatic cells, octoploid aneuploidy = inexact multiple of chromosomes in all body cells, e.g. 2n -1, 2n + 2 aneusomaty = chromosome number varies ± as in aneuploidy within one plant's body cells • Genomic configuration (i.e., subgenomes or ancestral chromosome sets, e.g. AABBCC);

• Agrostis stolonifera hybrids with three species of the genus Polypogon, rabbitsfootgrass or beardgrass (e.g. Conley 1993;[START_REF] Barkworth | Polypogon Desf[END_REF][START_REF] Giraldo-Cañas | El género Polypogon (Poaceae: Pooideae) en Colombia[END_REF].

1.8. Natural intergeneric hybrids have been reported in Europe [START_REF] Stace | Agrostis L. × Calamagrostis Adans[END_REF] and Taiwan between a few species of Agrostis and Calamagrostis Adanson, which has one naturalized and 25 native species in the United States and Canada. The hybrid of Agrostis stolonifera and Calamagrostis tenella has been called ×Agrocalamagrostis stebleri. However, the species that was in Calamagrostis is now considered to be an Agrostis, called Agrostis schraderiana (or A. agrostiflora) (synonym C. tenella) [START_REF] Hess | Flora der Schweiz und Angrenzender Gebiete[END_REF][START_REF] Tutin | Agrostis L[END_REF], and it does not occur in the New World. Natural interspecific hybrids involving other species of Agrostis have also been found, but either one or both of the parental species do not occur in the United States and Canada.

Distribution and Conservation of Agrostis Species

2.1. Many species of Agrostis are natively widespread in temperate areas, whereas other indigenous species or taxa (i.e. subspecies or botanical varieties) are regional, local or rare (WCMC 1998). Using the Natural Heritage Network's assessment criteria [START_REF] Master | Vanishing assets: Conservation status of U.S. species[END_REF], several Agrostis in the United States are variously rare (Table 1). There is an endemic species in Hawaii, several endemic taxa in California and/or Oregon, an endemic species in Wyoming (Yellowstone National Park), and a rare species in Wyoming, California and Russia (Kamchatka Peninsula) (CNPS 2001(CNPS , 2005;;[START_REF] Onhic | Rare, Threatened and Endangered Species of Oregon. Oregon Natural Heritage Information Center [ONHIC[END_REF][START_REF] Tercek | Genetic and historical relationships among geothermally adapted Agrostis (bentgrass) of North America and Kamchatka: Evidence for a previously unrecognized, thermally adapted taxon[END_REF][START_REF] Tercek | Heat avoidance life history strategy controls the distribution of geothermal Agrostis in Yellowstone[END_REF][START_REF] Tercek | Ecology and evolution of geothermally adapted Agrostis of North America and Kamchatka[END_REF].

2.2. None of these rare Agrostis are now listed as endangered or threatened under Federal law; under State law, in California A. blasdalei var. marinensis (Marin bentgrass) was listed in 1978 as rare, but in 1998 no plants could be found at its single known site, and it is considered a synonym under the species by [START_REF] Harvey | Agrostis, Bent grass[END_REF][START_REF] Harvey | Agrostis L[END_REF], CNPS (2001CNPS ( , 2005) ) and [START_REF] Kartesz | Synthesis of the North American Flora[END_REF] rather than being a distinct biological entity. Creeping bentgrass (A. stolonifera) probably occurs in meadow or wetland habitats of some threatened or endangered species of animals and plants listed under Federal law, although confirmation may be needed on the identification of this grass in their habitats (e.g., for the threatened western orchid Ute ladies-tresses, Spiranthes diluvialis).

The Species Creeping Bentgrass, Agrostis stolonifera

3.1. Agrostis stolonifera L. is a robust, fast-growing perennial which is biologically and ecologically very variable, phenotypically plastic (adjusting locally) and evolutionarily adaptive (i.e. genetically), with vegetative spread and reproduction by stolons (horizontal aboveground stems or runners), wind-pollinated flowers, and tiny (0.07 mg) seeds (i.e. caryopses) dispersed by wind, water and animals (Sell & Murrell 1996;Romero García et al. 1988b;[START_REF] Grime | Comparative Plant Ecology: A Functional Approach to Common British Species[END_REF]Kik et al. 1990aKik et al. , 1990b[START_REF] Kik | Variation of relative growth rate and survival in ecologically contrasting populations of Agrostis stolonifera[END_REF][START_REF] Grime | Relative growth-rate: Its range and adaptive significance in a local flora[END_REF][START_REF] Kurchenko | Ontogenesis and developmental stages of bentgrass (Agrostis stolonifera L.) swards.] Biulleten' Moskovskogo Obshchestva Ispytatelei Prirody[END_REF][START_REF] Eriksson | Seedling dynamics and life histories in clonal plants[END_REF][START_REF] Shipley | Regeneration and establishment strategies of emergent macrophytes[END_REF][START_REF] Davies | Population differentiation in Festuca rubra L. and Agrostis stolonifera L. in response to soil waterlogging[END_REF]Misra & Tyler 2000b;[START_REF] Aston | which is a trait of agronomic interest (e.g[END_REF]Jónsdóttir 1991b;Ahmad & Wainwright 1976;[START_REF] Wu | Esterase isoenzymes in populations of Agrostis stolonifera L[END_REF]Wu , 1981;;[START_REF] Kik | Ecological genetics of salt resistance in the clonal perennial, Agrostis stolonifera L[END_REF][START_REF] Badiani | Foliar antioxidant status of plants from naturally high-CO 2 sites[END_REF][START_REF] Kaligarič | Vegetation patterns and responses to elevated CO 2 from natural CO 2 springs at Strmec (Radenci, Slovenia)[END_REF][START_REF] Teyssonneyre | Effects of elevated CO 2 and cutting frequency on plant community structure in a temperate grassland[END_REF][START_REF] Bradshaw | Evolution and stress -genotypic and phenotypic components[END_REF]cf. Donohue 2003;[START_REF] Grime | The role of plasticity in resource capture by plants[END_REF]. Generally the species is an allotetraploid (Jones 1956b(Jones , 1956c;;[START_REF] Warnke | Isozyme analysis supports allotetraploid inheritance in tetraploid creeping bentgrass (Agrostis palustris Huds.)[END_REF][START_REF] Chakraborty | Linkage map construction in allotetraploid creeping bentgrass (Agrostis stolonifera L.)[END_REF], but it also has cytotypes of higher ploidy and other such chromosomal complexity (aneuploidy, aneusomaty, and accessory or B-chromosomes), at least within its native range in the Old World -see Table 2 and [START_REF] Kik | The distribution of cytotypes in ecologically contrasting populations of the clonal perennial Agrostis stolonifera[END_REF][START_REF] Kik | Ploidy level and somatic chromosome number variation in Agrostis stolonifera[END_REF]. Correlations usually are not taxonomically consistent within its very broad natural range, but sometimes the main ecological and/or genetic tendencies in a region are featured either as informal groups or formally as botanical varieties (e.g. [START_REF] Philipson | A revision of the British species of the genus Agrostis Linn[END_REF][START_REF] Paunero | Las especies españolas del género Agrostis[END_REF][START_REF] Jansen | Flora Neerlandica[END_REF][START_REF] Widén | The Genus Agrostis in Eastern Fennoscandia. Taxonomy and Distribution[END_REF]Romero García et al. 1988b;Sell & Murrell 1996).

3.2. For example in Great Britain and Ireland, Sell & Murrell (1996) A polder population (on estuary land obtained over a decade earlier by holding back the sea) had all three biotypes, which were genetically distinct.

3.4. The related species Agrostis capillaris L. (synonym A. tenuis Sibth.) has similarly been observed to be quite variable and fine-tuned to a range of habitats (e.g. Bradshaw 1959aBradshaw , 1959d[START_REF] Bradshaw | Population differentiation in Agrostis tenuis Sibth. III. Populations in varied environments[END_REF][START_REF] Kershaw | An investigation of the structure of a grassland community. I. The pattern of Agrostis tenuis[END_REF][START_REF] Mcneilly | Evolution in closely adjacent plant populations. IV. Barriers to gene flow[END_REF][START_REF] Chadwick | Population differentiation within Agrostis tenuis Sibth. in response to colliery spoil substrate factors[END_REF][START_REF] Tasker | The microdistribution of Agrostis tenuis on colliery spoil in relation to spoil chemical variability[END_REF][START_REF] Acheroy | Étude sur des populations d'Agrostis tenuis Sibth.: Variation morphologique et écotypes édaphiques[END_REF][START_REF] Roxburgh | Dune slack vegetation in southern New Zealand[END_REF][START_REF] Funtova | Differentiation of populations of Agrostis tenuis Sibth. and Dactylis glomerata L. in the flood meadow habitats of the Luga River[END_REF][START_REF] Leyer | Predicting plant species' responses to river regulation: The role of water-level fluctuations[END_REF][START_REF] Helgadóttir | Competitive interactions between populations of Poa pratensis and Agrostis tenuis from ecologically contrasting environments[END_REF], 1986;[START_REF] Rothanzl | Struktura Genet Agrostis capillaris v Heterogenním Prostředí Vytvořeném Mraveništi Druhu Lasius flavus [Genet structure of Agrostis capillaris in heterogeneous environments created by anthills of Lasius flavus[END_REF][START_REF] Dostál | Ant-induced soil modification and its effect on plant below-ground biomass[END_REF][START_REF] Zhao | Genetic diversity in colonial bentgrass (Agrostis capillaris L.) revealed by EcoRI-MseI and PstI-MseI AFLP markers[END_REF][START_REF] Balocchi-L | Caracterización de sitios de crecimiento de seis especies pratenses nativas y naturalizadas del Dominio Húmedo de Chile[END_REF][START_REF] Nelson | Rol de las especies pratenses nativas y naturalizadas en las praderas permanentes del Sur de Chile[END_REF][START_REF] Wilson | The genetics of naturalization: A comparison of morphological variation within and between populations of Agrostis capillaris L. as an exotic in New Zealand and as a native in Britain[END_REF].

Creeping Bentgrass Distribution and Utilization

4.1. Creeping bentgrass has become naturalized in temperate to cold-temperate regions throughout the world (cf. [START_REF] Woodward | The dynamics of leaf extension in plants with diverse altitudinal ranges. I. Field observations on temperature responses at one altitude[END_REF][START_REF] White | Cold acclimation and deacclimation in coolseason grasses[END_REF][START_REF] Tompkins | Dehardening of annual bluegrass and creeping bentgrass during late winter and early spring[END_REF][START_REF] Tompkins | Effects of ice cover on annual bluegrass and creeping bentgrass putting greens[END_REF], for example in New Zealand, southern Australia including Tasmania, South Africa, South America including Tierra del Fuego, Patagonia and the Andes, North America, and on remote islands such as Hawaii, the Juan Fernández Islands, the Falkland Islands, Gough Island and Tristan da Cunha. Agrostis stolonifera is native in Eurasia principally (also Iceland and North Africa), but has an ambiguous status (perhaps native locally) in northern USA and/or in Canada at some salt marshes and freshwater lakes (Hitchcock & Chase 1951;[START_REF] Voss | Michigan Flora, Part 1, Gymnosperms and Monocots[END_REF][START_REF] Dore | Grasses of Ontario[END_REF][START_REF] Harvey | Agrostis L[END_REF]). However four close relatives, the other main agronomic species (Tables 1 &2), are clearly native only in Eurasia or Europe (e.g. [START_REF] Widén | The Genus Agrostis in Eastern Fennoscandia. Taxonomy and Distribution[END_REF]Romero García et al. 1988a;[START_REF] Warnke | Isozyme analysis supports allotetraploid inheritance in tetraploid creeping bentgrass (Agrostis palustris Huds.)[END_REF][START_REF] Vergara | AFLP analyses of genetic diversity in bentgrass[END_REF]. In the USA, creeping bentgrass is definitely mostly if not entirely naturalized, and probably arrived well before the 1750s (cf. [START_REF] Sauer | The settlement of the humid East[END_REF][START_REF] Sauer | The settlement of the humid East[END_REF][START_REF] Cronon | Changes in the Land: Indians, Colonists, and the Ecology of New England[END_REF][START_REF] Richardson | An Essay on Agriculture; …to which is added, A Memoir…on the Nature and Nutritive Qualities of Fiorin Grass, with Practical Remarks on its Abundant Properties, and the Best Mode of Cultivating that Extraordinary Vegetable[END_REF][START_REF] Odland | Bent grass seed production in Rhode Island[END_REF][START_REF] Monteith | Classification of redtop and the common bent grasses[END_REF] Diversity welcomed the "International Seed Treaty" (IT/PGRFA), anticipating it will be a key international agreement dealing with issues concerning plant genetic resources for food and agriculture (cf. Fowler 2004;PGGR 2003).

4.3. The IT/PGRFA could intend the inclusion of creeping bentgrass even though not considering it a major grass forage. With some differences in use (circumscription) of Agrostis names (section 5.9), the history of the A. stolonifera listing warrants checking for confirmation of the species intended by reviewing how the name was brought into consideration. [START_REF] Hammer | Crops of European origin[END_REF] recognize all five main agronomic species (A. canina, A. capillaris, A. castellana, A. gigantea, A. stolonifera) as minor crops of Central Europe with genetic resources of value because of their utilization for fodder (e.g. [START_REF] Balasko | Bluegrasses, ryegrasses, and bentgrasses[END_REF][START_REF] Defrance | Redtop and the bentgrasses[END_REF][START_REF] Tomić | Dry matter and seed production in some inbred lines of three Agrostis species. Herbage Seed as a Key Factor for Improving Production and Environmental Quality[END_REF][START_REF] Tomić | Genetic resource of five Agrostis species present in Serbia and their utilisation[END_REF][START_REF] Chorlton | A forage grass and legume plant collecting expedition in Czechoslovakia 1992[END_REF][START_REF] Chorlton | A forage grass and legume plant collecting expedition in Portugal[END_REF][START_REF] Karlsen | Agronomic value of some north Norwegian populations of Agrostis capillaris L[END_REF][START_REF] Laser | Effect of legume proportion and physiological age on forage quality and the suitability of Agrostis capillaris L. and Festuca rubra L. for silage making[END_REF][START_REF] Dumortier | Effects of harvesting dates and frequencies on above-and below-ground dynamics in Belgian wet grasslands[END_REF][START_REF] Kratochwil | Responses of flower phenology and seed production under cattle-grazing impact in sandy grasslands[END_REF][START_REF] Hill | Long-term effects of excluding sheep from hill pastures in North Wales[END_REF][START_REF] Balocchi | Caracterización fenológica y productiva de Agrostis capillaris y Holcus lanatus en el Dominio Húmedo de Chile[END_REF][START_REF] Flores-M | Efecto del período de rezago sobre la calidad de cinco gramíneas forrajeras y su predicción por NIRS[END_REF][START_REF] Faville | Allozyme variation within and between three populations of browntop (Agrostis capillaris)[END_REF][START_REF] Bañuelos | Effect of grazing history, experimental defoliation, and genotype on patterns of silicification in Agrostis tenuis Sibth[END_REF][START_REF] Pieper | Redtop seed production in Illinois[END_REF][START_REF] De Coninck | Fiorin Graes eller Agrostis stolonifera[END_REF][START_REF] Richardson | An Essay on Agriculture; …to which is added, A Memoir…on the Nature and Nutritive Qualities of Fiorin Grass, with Practical Remarks on its Abundant Properties, and the Best Mode of Cultivating that Extraordinary Vegetable[END_REF][START_REF] Haggar | The seasonal productivity, quality and response to nitrogen of four indigenous grasses compared to Lolium perenne[END_REF][START_REF] Frame | Herbage productivity of a range of grass species in association with white clover[END_REF][START_REF] Sheldrick | Dry-matter yield and response to nitrogen of an Agrostis stolonifera-dominant sward[END_REF], with both of the latter species of additional value for revegetation ("recultivation") and soil stabilization and improvement (cf. [START_REF] Smith | The use of metal-tolerant plant populations for the reclamation of metalliferous wastes[END_REF][START_REF] Wainscott | New gulley stopper … creeping bent[END_REF], and with all but A. gigantea of "ornamental" value (i.e. as lawn grasses or turf). An overview on this species (redtop) with its turf history is given by [START_REF] Brede | Three minor Agrostis species: Redtop, highland bentgrass, and Idaho bentgrass[END_REF].

Creeping Bentgrass -Establishing a Major Turfgrass

5.1. In the 1890s seeds of a few Agrostis species, sometimes including small amounts of A. stolonifera, were gathered in mixtures from semi-wild pasture populations in central Europe (present-day Austria and Hungary) and likely first imported to establish golf courses [START_REF] Duich | The bent grasses[END_REF][START_REF] Warnke | Creeping bentgrass (Agrostis stolonifera L.)[END_REF]. A large bentgrass seed trade (with ± 6,000,000 seeds per pound) continued for several decades, especially from a range of semi-wild locales across Germany [START_REF] Edler | Bent seed production in Germany[END_REF]) (annual imports 388,500 lbs to 294,000 lbs from 1927-1930 -USGA 1930b), and from an increasing production in New Zealand (USGA 1922b(USGA , 1922c(USGA , 1925b(USGA , 1930b;;[START_REF] Hillman | South German mixed bent seed described[END_REF][START_REF] Hillman | Identifying turf-grass seed[END_REF][START_REF] Henry | Changes in the bent grass seed market as viewed by the seed analyst[END_REF]cf. Levy 1924;[START_REF] Suckling | Genetic resources in high-rainfall hill pastures of New Zealand. I. Collection of ryegrass, browntop, and white clover[END_REF][START_REF] Nikolov | Investigations on the biology of seed germination of Agrostis capillaris L. and Festuca fallax Thuill. from grass coenoses on Vitosha Mountain[END_REF][START_REF] López | Características físicas y germinativas de la semilla de especies pratenses nativas y naturalizadas del Dominio Húmedo de Chile. I: Agrostis capillaris L., Arrhenatherum elatius[END_REF][START_REF] Balocchi | Caracterización fenológica y productiva de Agrostis capillaris y Holcus lanatus en el Dominio Húmedo de Chile[END_REF]. Moreover seed was gathered from free-living (naturalized) U.S. populations in coastal New England from about 1900, and chiefly 1924-1934 in the low coastal region of Oregon but also Washington and northwestern California, as well as farther inland in western Oregon and perhaps Washington [START_REF] Hillman | South German mixed bent seed described[END_REF][START_REF] Hillman | Identifying turf-grass seed[END_REF][START_REF] Henry | Changes in the bent grass seed market as viewed by the seed analyst[END_REF]USGA 1925a[START_REF] Usga | Kinds and quantities of turf seeds purchased by golf courses[END_REF][START_REF] Monteith | Classification of redtop and the common bent grasses[END_REF][START_REF] Carrier | Rhode Island bent seed situation[END_REF][START_REF] Odland | Bent grass seed production in Rhode Island[END_REF][START_REF] Schoth | Bent grass seed production in the Pacific Northwest[END_REF][START_REF] Schoth | Seed production of turf grasses on the Pacific Coast[END_REF][START_REF] Hyslop | Oregon bent grass seed certification[END_REF]. Additional seed was imported mostly from crop production in southeastern (maritime) Canada, as well as inland in Alberta [START_REF] Clark | Turf grasses in Canada[END_REF][START_REF] Tregillus | Rhode Island bent as a putting green turf[END_REF][START_REF] Malte | The commercial bent grasses (Agrostis) in Canada[END_REF][START_REF] Lelacheur | Canadian certified bent seed[END_REF][START_REF] Henry | Changes in the bent grass seed market as viewed by the seed analyst[END_REF][START_REF] Hillman | Identifying turf-grass seed[END_REF].

5.2. The many shipments of "South German mixed bent" seed were variously comprised of mostly Agrostis capillaris with some A. canina, about 10% or less A. gigantea, and sometimes a trace of A. stolonifera [START_REF] Hillman | South German mixed bent seed described[END_REF][START_REF] Hillman | Identifying turf-grass seed[END_REF][START_REF] Henry | Changes in the bent grass seed market as viewed by the seed analyst[END_REF]USGA 1922cUSGA , 1930b;;Piper & Oakley 1922b;Oakley 1923a;[START_REF] Monteith | Classification of redtop and the common bent grasses[END_REF]. Considering the relatively wild sources of seed from populations in Europe (cf. Hultén & Fries 1986), perhaps A. vinealis was also included (cf. Sell & Murrell 1996;[START_REF] Scholz | Agrostis vinealis[END_REF][START_REF] Vester | Experimentelle Untersuchungen zum vegetativen Kolonisation-und Konkurrenzverhalten bei Agrostis vinealis und A. capillaris[END_REF][START_REF] Ridgway | Identification of roots from grass swards using PCR-RFLP and FFLP of the plastid trnL (UAA) intron[END_REF]. [START_REF] Hillman | Identifying turf-grass seed[END_REF] reported that a distinct bentgrass seed which was being termed "hybrid seed" was coming from Germany under the names "creeping bent" and "German bent", in "recent years…practically all the seed of some lots", and that the real nature of these plants was under study.

5.3. The bentgrass seed imported from New Zealand (e.g. 162,600 lbs in 1929 and an impressive 508,900 lbs or 254 tons in 1930) was eventually found to be not solely Agrostis capillaris (cf. [START_REF] Suckling | Genetic resources in high-rainfall hill pastures of New Zealand. I. Collection of ryegrass, browntop, and white clover[END_REF]Rapson & Wilson 1992a, 1992b;Karataglis 1980b[START_REF] Karataglis | Gene flow in parapatric plant populations of Agrostis tenuis Sibth. and Festuca rubra L[END_REF]), but to include A. castellana (which earlier had been considered just a local ecological variation), and to additionally include its hybrids with A. capillaris [START_REF] Rumball | Plant introduction trials. Performance of Agrostis species at Palmerston North[END_REF][START_REF] García | The F 1 hybrids of A. stolonifera and A. capillaris[END_REF]Batson 1998a;[START_REF] Steiner | Zur Echtheitsbestimmung mittels morphologischer Merkmale der Spelzfrüchte bei Agrostis-Arten [Verification of species of Agrostis by means of morphological characters of the florets[END_REF][START_REF] Henry | Changes in the bent grass seed market as viewed by the seed analyst[END_REF]) -which have been called A. ×fouilladeana (rather than A. ×fouilladei P. Fourn. [1934], nomen nudum -see [START_REF] Lambinon | Agrostis ×fouilladeana Lambinon & Verloove, nothospec. nov., en Belgique et dans le nord de la France[END_REF]. Similarly, the inland bentgrass of western Oregon and Washington is increasingly being recognized not as a dryland or upland (i.e. non-coastal) variation of A. capillaris (marketed as 'Highland' bentgrass), but as a landrace of the species A. castellana [START_REF] Henry | Changes in the bent grass seed market as viewed by the seed analyst[END_REF][START_REF] Hillman | Identifying turf-grass seed[END_REF][START_REF] Schoth | Bent grass seed production in the Pacific Northwest[END_REF][START_REF] Schoth | Seed production of turf grasses on the Pacific Coast[END_REF][START_REF] Usda | Grass: The Yearbook of Agriculture[END_REF][START_REF] Usda | Seeds: The Yearbook of Agriculture[END_REF][START_REF] Madison | Turfgrass ecology: Effects of mowing, irrigation, and nitrogen treatments of Agrostis palustris Huds., 'Seaside' and Agrostis tenuis Sibth., 'Highland' on population, yield, rooting and cover[END_REF][START_REF] Griffin | Bentgrass for fairways[END_REF]Funk 998;[START_REF] Brede | Four more unconventional grasses to know and love[END_REF]Brilman 2001a;Ruemmele 2000a[START_REF] Ruemmele | Agrostis capillaris (Agrostis tenuis Sibth.) colonial bentgrass[END_REF][START_REF] Brede | Three minor Agrostis species: Redtop, highland bentgrass, and Idaho bentgrass[END_REF]cf. Vergara & Bughrara 2003).

5.4. The bentgrass seed from the imports and North America was widely distributed and sown on developing golf courses, and as well for municipal parks, playgrounds and fine lawns, with the intention that the most appropriate of the bentgrasses for a site would come through (cf. [START_REF] Wilson | The genetics of naturalization: A comparison of morphological variation within and between populations of Agrostis capillaris L. as an exotic in New Zealand and as a native in Britain[END_REF][START_REF] Rapson | Non-adaptation in Agrostis capillaris L. (Poaceae)[END_REF][START_REF] Lesica | Ecological genetics and the restoration of plant communities: Mix or match?[END_REF]. With golf's rapid geographic expansion, on most of the new courses Agrostis stolonifera emerged over the years to be dominant in the turf in many regions, except in the cool Northeast where A. canina sometimes out-competed the other bentgrasses [START_REF] Monteith | Classification of redtop and the common bent grasses[END_REF]USGA 1922cUSGA , 1930a;;Piper & Oakley 1922b, 1922a;Oakley 1923a;cf. Skogley 1973;[START_REF] Hurley | Velvet bentgrass -the putter's delight[END_REF][START_REF] Brilman | Velvet bentgrass: Rediscovering a misunderstood turfgrass[END_REF]Ruemmele 2000b;[START_REF] Rose-Fricker | Registration of 'Greenwich' velvet bentgrass[END_REF][START_REF] Hollman | Identification of putative velvet bentgrass clones using RAPD markers[END_REF]. Thus creeping bent's characteristics became familiar.

Experience for turf was also gained with A. capillaris (called Rhode Island bent, later colonial bent), which was introduced to New England early for pasture -it is mentioned in 1747 [START_REF] Odland | Bent grass seed production in Rhode Island[END_REF]) and likely was present a century earlier (cf. [START_REF] Cronon | Changes in the Land: Indians, Colonists, and the Ecology of New England[END_REF][START_REF] Sauer | The settlement of the humid East[END_REF]. Production to respond to the emerging desire (1900)(1901)(1902)(1903)(1904)(1905) for sports turf and lawns began by gathering seed from naturalized stands in Rhode Island and southern Connecticut, and subsequently Massachusetts. Imports from Germany and New Zealand as well as southeastern Canada later shifted the Rhode Island production to growing A. canina and A. stolonifera [START_REF] Odland | Bent grass seed production in Rhode Island[END_REF][START_REF] Hillman | Identifying turf-grass seed[END_REF]. Agrostis gigantea was also common in New England, and care needed to avoid gathering it, but there was deliberate adulteration too in this seed trade [START_REF] Carrier | Rhode Island bent seed situation[END_REF][START_REF] Tregillus | Rhode Island bent as a putting green turf[END_REF][START_REF] Hillman | Identifying turf-grass seed[END_REF]. [START_REF] García | The F 1 hybrids of A. stolonifera and A. capillaris[END_REF] reported finding Agrostis in Rhode Island with unusual morphology and aneuploidy (cf. [START_REF] Anderson | Hybridization of the habitat[END_REF]; these plants are now considered hybrids of A. capillaris and A. gigantea [START_REF] Björkman | Chromosome studies in Agrostis[END_REF][START_REF] Widén | The Genus Agrostis in Eastern Fennoscandia. Taxonomy and Distribution[END_REF]Bradshaw 1975a;[START_REF] García | The F 1 hybrids of A. stolonifera and A. capillaris[END_REF]cf. Sell & Murrell 1996), which are called A. ×bjoerkmanii.

5.5. So over several decades, creeping bentgrass transitioned from being considered a growth form or obscure minor kind of wild bent sometimes showing up in the "South German mixed bent" seed in small quantities, to a highly desirable foreign species in short supply (Piper & Oakley 1921, 1922a;[START_REF] Monteith | Classification of redtop and the common bent grasses[END_REF]). Through U.S. production efforts [START_REF] Monteith | Turf seed of higher quality[END_REF], including encouraging propagation by stolons rather than seeds, Agrostis stolonifera became the most successful and popular grass for putting greens [START_REF] Taylor | Turfgrass -its development and progress[END_REF][START_REF] Radko | Bentgrasses for putting greens[END_REF]), and now is grown extensively and managed intensively for greens as well as tees and fairways -and that is the chief focus in continuing to improve it agronomically (e.g. [START_REF] Hurley | Creeping bentgrass: The legacy and the promise[END_REF][START_REF] Morris | Bentgrasses and bermudagrasses for today's putting greens[END_REF][START_REF] Warnke | Creeping bentgrass (Agrostis stolonifera L.)[END_REF]). The species is additionally investigated as a forage [START_REF] Balasko | Bluegrasses, ryegrasses, and bentgrasses[END_REF][START_REF] Tomić | Dry matter and seed production in some inbred lines of three Agrostis species. Herbage Seed as a Key Factor for Improving Production and Environmental Quality[END_REF][START_REF] Tomić | Genetic resource of five Agrostis species present in Serbia and their utilisation[END_REF], and of interest for restoring some mine sites involving heavy metals [START_REF] Smith | The use of metal-tolerant plant populations for the reclamation of metalliferous wastes[END_REF]) and for water treatment [START_REF] Löser | Conditioning of heavy metalpolluted river sediments by helophytes[END_REF][START_REF] Tyrrel | Removal of ammoniacal nitrogen from landfill leachate by irrigation onto vegetated treatment planes[END_REF][START_REF] Mazer | Limitations to vegetation establishment and growth in biofiltration swales[END_REF]. Sell & Murrell (1996) state that A. stolonifera var. marina has been used in the formation of lawns. There are more than 1,000 holdings of genetic resources of Agrostis species in various countries [START_REF] Williams | Genetic resources of temperate native and low-input grasses in New Zealand and Australian collections[END_REF], and still exploration for germplasm of A. stolonifera for turf from populations in its native range (e.g. [START_REF] Romani | Collection and preliminary evaluation of native turfgrass accessions in Italy[END_REF]cf. Xie et al. 2003).

5.6. Ordinarily the modern turf cultivar (e.g. [START_REF] Robinson | Registration of 'SR 1020' creeping bentgrass[END_REF][START_REF] Brauen | Registration of 'Putter' creeping bentgrass[END_REF][START_REF] Hurley | Registration of 'Southshore' creeping bentgrass[END_REF][START_REF] Engelke | Registration of 'Crenshaw' creeping bentgrass[END_REF] is seeded, and is a clustering of A. stolonifera genotypes bred and selected for a set of distinctive traits (e.g. [START_REF] Holt | Variation in spreading rate and growth characteristics of creeping bentgrass seedlings[END_REF][START_REF] Lush | Turf growth and performance evaluation based on turf biomass and tiller density[END_REF][START_REF] Engelke | Bentgrass breeding -Texas style[END_REF][START_REF] Cattani | Relationship of shoot morphology between seedlings and established turf in creeping bentgrass[END_REF][START_REF] Cattani | Early plant development in 'Emerald' and 'UM67-10' creeping bentgrass[END_REF][START_REF] Cattani | Tillering, internode development, and dry matter partitioning in creeping bentgrass[END_REF][START_REF] Sweeney | Root weight, nonstructural carbohydrate content, and shoot density of high-density creeping bentgrass cultivars[END_REF][START_REF] Murphy | Assessing cool-season turfgrasses for performance under simulated traffic stress[END_REF][START_REF] Dacosta | Water use and requirements for bentgrasses under fairway conditions[END_REF][START_REF] Croce | Agrostis cultivar characterizations for closely mowed putting greens in a Mediterranean climate[END_REF][START_REF] Rogers | What's available in creeping bentgrasses[END_REF][START_REF] Huff | Comparing the new bents[END_REF][START_REF] Stier | Managing the new 'alpha'-bents[END_REF][START_REF] Wilkinson | Electrophoretic identification of Agrostis palustris and Poa pratensis cultivars[END_REF][START_REF] Warnke | Relationships among creeping bentgrass cultivars based on isozyme polymorphisms[END_REF]Golembiewski et al. 1997aGolembiewski et al. , 1997b;;[START_REF] Caceres | RFLP markers are an effective tool for the identification of creeping bentgrass (Agrostis stolonifera L.) cultivars[END_REF]Casler et al. 2003;[START_REF] Kubik | Creeping bentgrass (Agrostis stolonifera L.) cultivar identification using simple sequence repeats[END_REF], rather than being a single genetic line distributed by vegetative propagation as in earlier decades (e.g. [START_REF] Metcalf | Stolon-planting versus seeding for putting greens[END_REF]USGA 1924cUSGA , 1924bUSGA , 1924aUSGA , 1930a;;Piper & Oakley 1921, 1924;[START_REF] Carrier | Vegetative planting[END_REF][START_REF] Carrier | The vegetative method of planting creeping bent[END_REF]Oakley 1923b[START_REF] Oakley | Some suggestions for beginners on the vegetative planting of creeping bent[END_REF][START_REF] Radko | Bentgrasses for putting greens[END_REF][START_REF] Bengeyfield | Turf establishment -stolonizing[END_REF]cf. Abramashvili 1978).

5.7. From its 1923 beginnings in the Pacific Northwest Coast region [START_REF] Schoth | Bent grass seed production in the Pacific Northwest[END_REF][START_REF] Schoth | Seed production of turf grasses on the Pacific Coast[END_REF][START_REF] Hyslop | Oregon bent grass seed certification[END_REF]USGA 1930b;[START_REF] Monteith | Turf seed of higher quality[END_REF][START_REF] Ferguson | West Coast trip[END_REF]Pojar & MacKinnon 1994;[START_REF] Franklin | Natural Vegetation of Oregon and Washington[END_REF], the production of seed of bentgrasses for turf for the USA and internationally has become a modern industry concentrated in western Oregon in the Willamette Valley (125 mi × 25-30 mi) between the coast and Cascade Range of mountains. About 6,000 acres are in production, mostly for creeping bentgrass [START_REF] North | The relative seed yields in different species and varieties of bent grass[END_REF][START_REF] Cattani | Seed yield and yield components of creeping bentgrass cultivars[END_REF]cf. Wright 1980;[START_REF] Jonassen | Virkning av liten plantetetthet på frøavling av engsvingel, hundegras, rødsvingel og engkvein [Effect of low planting density on seed yield of meadow fescue, cocksfoot, red fescue and common bentgrass[END_REF][START_REF] Rumball | Grasslands Egmont' amenity browntop (Agrostis capillaris L. syn. Agrostis tenuis Sibth.)[END_REF][START_REF] Chastain | Vegetative plant development and seed production in cool-season perennial grasses[END_REF]. Background on this region is provided in several accounts [START_REF] Bowen | The Willamette Valley: Migration and Settlement on the Oregon Frontier[END_REF][START_REF] Johannessen | The vegetation of the Willamette Valley[END_REF][START_REF] Habeck | The original vegetation of the mid-Willamette Valley[END_REF][START_REF] Nelson | The grasses of Salem, Oregon, and vicinity[END_REF][START_REF] Mueller-Warrant | Geographic distribution of prominent weeds of grass seed production[END_REF][START_REF] Clark | Fire, mowing, and hand-removal of woody species in restoring a native wetland prairie in the Willamette Valley of Oregon[END_REF][START_REF] Hulse | Envisioning alternatives: Using citizen guidance to map future land and water use[END_REF].

5.8. The U.S. turfgrass industry frequently equates creeping bentgrass with Agrostis palustris or sometimes A. stolonifera var. palustris, but this usage does not agree with the detailed botanical concepts of [START_REF] Hubbard | Grasses: A Guide to Their Structure, Identification, Uses, and Distribution in the British Isles, 3rd Ed[END_REF] or Sell & Murrell (1996) where the plants are native, and are also utilized. This U.S. habit or convention may reflect the influence of the manual by Hitchcock & Chase (1951, and Misra & Tyler 2000b, 2000a;[START_REF] Kik | Population responses in Agrostis stolonifera to selective forces in inland and coastal habitats[END_REF]Kik et al. 1990aKik et al. , 1990b[START_REF] Kik | The distribution of cytotypes in ecologically contrasting populations of the clonal perennial Agrostis stolonifera[END_REF]. Another legacy problem lingering in the USA and Canada is an overly broad scope in use of the name A. stolonifera (or A. alba) (e.g. [START_REF] Malte | The commercial bent grasses (Agrostis) in Canada[END_REF][START_REF] Gleason | The New Britton and Brown Illustrated Flora of the Northeastern United States and Adjacent Canada[END_REF][START_REF] Gleason | Manual of Vascular Plants of Northeastern United States and Adjacent Canada[END_REF][START_REF] Steyermark | Flora of Missouri[END_REF][START_REF] Munz | Supplement to "A California Flora[END_REF][START_REF] Cronquist | Intermountain Flora: Vascular Plants of the Intermountain West[END_REF][START_REF] Stubbendieck | North American Range Plants[END_REF]but cf. Welsh et al. 1993), occasionally even using the name A. stolonifera var. stolonifera (rather than A. stolonifera var. major) for the plants usually called redtop and now well accepted as A. gigantea (e.g. in Europe, [START_REF] Tutin | Agrostis L[END_REF]Sell & Murrell 1996;and in North America, Fassett 1951;[START_REF] Voss | Michigan Flora, Part 1, Gymnosperms and Monocots[END_REF][START_REF] Bailey | Hortus Third: A Concise Dictionary of Plants Cultivated in the United States and Canada[END_REF][START_REF] Mcneill | Taxonomic and nomenclatural notes on Ontario, Canada grasses[END_REF][START_REF] Dore | Grasses of Ontario[END_REF][START_REF] Pohl | How to Know the Grasses, 3rd Ed. Agrostis, Bentgrass[END_REF][START_REF] Gleason | Manual of Vascular Plants of Northeastern United States and Adjacent Canada, 2nd Ed[END_REF][START_REF] Harvey | Agrostis, Bent grass[END_REF][START_REF] Harvey | Agrostis[END_REF][START_REF] Yatskievych | Steyermark's Flora of Missouri[END_REF].

5.9. Four themes (discovery, research, biological classification, scientific nomenclature) are interwoven in the 250-year historical to current usage of several Agrostis names, reflecting improved understanding but lags in communication (cf. [START_REF] Spooner | Plant nomenclature and taxonomy: An horticultural and agronomic perspective[END_REF].

(1) The natural variation of creeping bentgrasses in Europe led to early recognition of many formal distinctions, such as Agrostis stolonifera L. ( 1753 [START_REF] Kurchenko | Critical notes on bents of Agrostis stolonifera group: The new species A. diluta (Poaceae)[END_REF]. The usage of common names in North America (Tables 1 &2) has also become more stable (cf. [START_REF] Henry | Changes in the bent grass seed market as viewed by the seed analyst[END_REF][START_REF] Monteith | Classification of redtop and the common bent grasses[END_REF][START_REF] Usga | When is a bent grass a creeping bent?[END_REF], but can still be variable, and entangled or confused.

5.10. Consequently, when going beyond the extensive information on turf mainly for golf courses, in the North American literature in particular alertness is needed as to the species intended (see Table 2 synonymy). However, the identity of the plants studied frequently cannot be ascertained from the report, and the research is less useful, because several Agrostis species occur in the area of study and the plant characteristics were not distinguished nor a taxonomic reference stated (nor the actual originating source clear if research plants were supplied). As provisional pragmatic assumptions, A. alba tends to mean redtop (A. gigantea); A. stolonifera to mean creeping bentgrass but sometimes redtop, and occasionally both (i.e. the name used broadly or sensu lato, sometimes with A. stolonifera var. major for redtop and A. stolonifera var. palustris for creeping bentgrass, but possibly A. stolonifera var. stolonifera used for either); and A. palustris is A. stolonifera (creeping bentgrass). This confusion makes other literature especially useful in North America for gaining insights on the population dynamics and ecology of the species creeping bentgrass (A. stolonifera, synonym A. palustris) in natural to variously managed systems.

Creeping Bentgrass Habitats, Ecology, and Reproduction

6.1. In Europe where Agrostis stolonifera has been taxonomically clear for decades and much more studied in the general environment (although not as much as an amenity turfgrass), the habitats of this common species range from salt marshes and dunes to chalk cliffs, inland marshes, streamsides and ditches, grasslands and pastures, including locales periodically flooded, open woodlands, damp arable land, paths, urban parks and lawns, and rough ground such as roadsides (Sell & Murrell 1996;[START_REF] Hubbard | Grasses: A Guide to Their Structure, Identification, Uses, and Distribution in the British Isles, 3rd Ed[END_REF][START_REF] Jansen | Grassen om het IJsselmeer. V. Agrostis[END_REF][START_REF] Grime | Comparative Plant Ecology: A Functional Approach to Common British Species[END_REF][START_REF] Silvertown | Hydrologically defined niches reveal a basis for species richness in plant communities[END_REF][START_REF] Schröder | Rejecting the mean: Estimating the response of fen plant species to environmental factors by non-linear quantile regression[END_REF]Rozema & Blom 1977;[START_REF] Eertman | Restoration of the Sieperda tidal marsh in the Scheldt Estuary, The Netherlands[END_REF][START_REF] Burgess | Vegetation change on the Ouse Washes wetland, England, 1972-88 and effects on their conservation importance[END_REF][START_REF] Gipiškis | Effect of the interaction between biological properties of herbage species and factors of cultivation techniques on the formation of swards in the meadows of the Lower Nemunas at various levels of flooding[END_REF][START_REF] Raven | Vegetation changes within the flood relief stage of two-stage channels excavated along a small rural clay river[END_REF][START_REF] Panter | Rapid changes in the vegetation of a shallow pond in Epping Forest, related to recent droughts[END_REF][START_REF] Gilbert | Influence of seed mixture and hydrological regime on the establishment of a diverse grassland sward at a site with high phosphorus availability[END_REF][START_REF] Van Eck | Is tolerance to summer flooding correlated with distribution patterns in river floodplains? A comparative study of 20 terrestrial grassland species[END_REF][START_REF] Kennedy | Experimental growth responses to groundwater level variation and competition in five British wetland plant species[END_REF][START_REF] Kohler | Macrophyte distribution and habitat quality in the eutrophic Bjorka-Kavlinge River (Skåne, South Sweden) [Makrophyten-verbreitung und Standortqualitat im eutrophen Bjorka-Kavlinge-Fluss (Skåne, Sudschweden)[END_REF][START_REF] Hald | Impact of agricultural fields on vegetation of stream border ecotones in Denmark[END_REF][START_REF] Croxton | A comparison of green lanes and field margins as bumblebee habitat in an arable landscape[END_REF][START_REF] Roovers | Plant species variation across path ecotones in a variety of common vegetation types[END_REF][START_REF] Cornish | The origin and structure of the grassland types of the central North Downs[END_REF][START_REF] Thompson | Urban domestic gardens (III): Composition and diversity of lawn floras[END_REF][START_REF] Ross | Vegetation change on highway verges in south-east Scotland[END_REF]. Similarly in the USA, the species occurs in coastal habitats (e.g. [START_REF] Caldwell | A floristic and vegetation analysis of a freshwater tidal marsh on the Merrimack River, West Newbury, Massachusetts[END_REF]Wu 1981;USGA 1925a) and has become widely established inland, for example reported in Missouri habitats [START_REF] Yatskievych | Steyermark's Flora of Missouri[END_REF]) along the margins of ponds and lakes and on streambanks, in moist disturbed areas, various grasslands (including upland prairies), moderately moist (mesic) to rather dry upland forests, on roadsides and railroad embankments, and in sidewalk crevices.

6.2. Creeping bentgrass is grazed by cattle, sheep and horses, as well as rabbits and hares and migratory wildfowl in Europe such as the wigeon Anas penelope and various species of geese [START_REF] Cadwalladr | Wigeon (Anas penelope L.) conservation and salting pasture management at Bridgwater Bay National Nature Reserve, Somerset[END_REF][START_REF] Owen | The feeding ecology and conservation of wigeon wintering at the Ouse Washes, England[END_REF][START_REF] Loonen | The significance of water as a key trigger enabling wigeon Anas penelope to feed on grass leaves of Agrostis stolonifera in a wet meadow system in Lake Lauwersmeer[END_REF][START_REF] Groot Bruinderink | The impact of wild geese visiting improved grasslands in The Netherlands[END_REF][START_REF] Chang | Seed dispersal by small herbivores and tidal water: Are they important filters in the assembly of salt-marsh communities?[END_REF], and it is used for nest building by grebes [START_REF] Ridley | The Dispersal of Plants Throughout the World[END_REF]. Agrostis castellana may be grazed by Canada geese (Branta canadensis) (cf. [START_REF] Conover | Herbivory by Canada geese: Diet selection and effect on lawns[END_REF]. Throughout the year, creeping bentgrass at impoverished mine sites can be the main food of the Eurasian field vole, Microtus agrestis [START_REF] Stoddart | Does trap odour influence estimation of population size of the short-tailed vole, Microtus agrestis?[END_REF][START_REF] Hunter | Ecotoxicology of copper and cadmium in a contaminated grassland ecosystem, III. Small mammals[END_REF]. It is a larval food plant for the widespread small heath butterfly Coenonympha pamphilus [START_REF] Goverde | Effects of elevated CO 2 on development and larval food-plant preference in the butterfly Coenonympha pamphilus (Lepidoptera, Satyridae)[END_REF]. Agrostis stolonifera can host fungal endophytes (cf. [START_REF] Clay | Evolutionary origins and ecological consequences of endophyte symbiosis with grasses[END_REF][START_REF] Faeth | Are endophytic fungi defensive plant mutualists?[END_REF][START_REF] Malinowski | Adaptations of endophyte-infected coolseason grasses to environmental stresses: Mechanisms of drought and mineral stress tolerance[END_REF][START_REF] Schardl | Symbioses of grasses with seedborne fungal endophytes[END_REF], as can A. capillaris, A. castellana, A. gigantea and several native Agrostis species [START_REF] Spooner | Epichloë in Britain[END_REF][START_REF] Aldous | Evaluation of endophyte in creeping bentgrass (Agrostis stolonifera) and common bentgrass (A. capillaris)[END_REF][START_REF] Aldous | Endophytes in creeping bentgrass (Agrostis stolonifera and A. palustris)[END_REF][START_REF] Thrower | Uptake of sugars by Epichloë typhina (Pers. ex Fr.) Tul. in culture and from its host Agrostis stolonifera L[END_REF]Bradshaw 1959b;[START_REF] Saikkonen | Endophytic fungi in wild and cultivated grasses in Finland[END_REF][START_REF] Zabalgogeazcoa | Fungal endophytes in grasses from semi-arid permanent grasslands of western Spain[END_REF][START_REF] White | Endophyte-host associations in grasses. XVI. Patterns of endophyte distribution in species of the tribe Agrostideae[END_REF][START_REF] White | Endophyte-host associations in forage grasses. XV. Clustering of stromata-bearing individuals of Agrostis hyemalis infected by Epichloë typhina[END_REF][START_REF] Zenbayashi | Acremonium endophytes found in Agrostis clavata Trin., A. clavata Trin. subsp. matsumurae Tateoka and Poa trivialis L. in Japan[END_REF].

6.3. The numerous tiny seeds of creeping bentgrass are dispersed by the wind and on water [START_REF] Wolters | Plant colonization after managed realignment: The relative importance of diaspore dispersal[END_REF], sometimes to lake islands [START_REF] Nilsson | Species richness and dispersal of vascular plants to islands in Lake Möckeln, southern Sweden[END_REF][START_REF] Roden | Persistence, extinction and different species pools within the flora of lake islands in western Ireland[END_REF], and consumed and sometimes dispersed (still viable) by cattle, sheep, horses, white-tailed deer (Odocoileus virginianus), fallow deer (Dama dama), and brent geese or brants (Branta bernicla) [START_REF] Welch | Studies in the grazing of heather moorland in north-east Scotland. IV. Seed dispersal and plant establishment in dung[END_REF]Cosyns et al. 2005a;[START_REF] Myers | Seed dispersal by whitetailed deer: Implications for long-distance dispersal, invasion, and migration of plants in eastern North America[END_REF][START_REF] Gill | The impact of deer on woodlands: The effects of browsing and seed dispersal on vegetation structure and composition[END_REF][START_REF] Chang | Seed dispersal by small herbivores and tidal water: Are they important filters in the assembly of salt-marsh communities?[END_REF], and the seeds are also dispersed externally on cattle hair [START_REF] Couvreur | Large herbivores as mobile links between isolated nature reserves through adhesive seed dispersal[END_REF]). In addition to the seeds, jointed pieces of the stolons (i.e. with nodes) can be carried downstream by the water [START_REF] Mitlacher | Restoration of wooded meadows -a comparative analysis along a chronosequence on Öland (Sweden)[END_REF][START_REF] Widén | The Genus Agrostis in Eastern Fennoscandia. Taxonomy and Distribution[END_REF][START_REF] Nilsson | A comparison of species richness and traits of riparian plants between a main river channel and its tributaries[END_REF][START_REF] Andersson | Plant dispersal in boreal rivers and its relation to the diversity of riparian flora[END_REF][START_REF] Vogt | Water-borne seed transport and seed deposition during flooding in a small river-valley in Northern Germany[END_REF][START_REF] Boedeltje | Plant dispersal in a lowland stream in relation to occurrence and three specific life-history traits of the species in the species pool[END_REF][START_REF] Boedeltje | Dispersal phenology of hydrochorous plants in relation to discharge, seed release time and buoyancy of seeds: The flood pulse concept supported[END_REF][START_REF] Goodson | Evidence for hydrochory and the deposition of viable seeds within winter flow-deposited sediments: The River Dove, Derbyshire[END_REF]Levine 2000Levine , 2001Levine , 2003;;[START_REF] Wolters | [END_REF]. Seeds of A. canina and A. capillaris also are known to be viably dispersed by passing through livestock [START_REF] Bruun | The past impact of livestock husbandry on dispersal of plant seeds in the landscape of Denmark[END_REF]Pakeman et al. 2002;Cosyns et al. 2005aCosyns et al. , 2005b)), A. capillaris through rabbits (Oryctolagus cuniculus) (Cosyns et al. 2005b), and A. gigantea through white-tailed deer [START_REF] Myers | Seed dispersal by whitetailed deer: Implications for long-distance dispersal, invasion, and migration of plants in eastern North America[END_REF]) and as well by water [START_REF] Merritt | Plant dispersal along rivers fragmented by dams[END_REF]. Seeds of A. capillaris are also dispersed externally by domesticated and wild ungulates (via hair and hooves) [START_REF] Couvreur | Large herbivores as mobile links between isolated nature reserves through adhesive seed dispersal[END_REF][START_REF] Mouissie | Estimating adhesive seeddispersal distances: Field experiments and correlated random walks[END_REF][START_REF] Heinken | Trägt Schalenwild durch Epizoochorie zur Ausbreitung von Gefäßpflanzen in mitteleuropäischen Wäldern bei? Eine Fallstudie aus Nordostdeutschland[END_REF] and are eaten and apparently dispersed by ants, and the plants are sometimes associated with ant-mounds [START_REF] Rothanzl | Struktura Genet Agrostis capillaris v Heterogenním Prostředí Vytvořeném Mraveništi Druhu Lasius flavus [Genet structure of Agrostis capillaris in heterogeneous environments created by anthills of Lasius flavus[END_REF][START_REF] Kovář | Vegetation of ant-hills in a mountain grassland: Effects of mound history and of dominant ant species[END_REF][START_REF] Kovář | Ant herbivory -a significant factor in population dynamics of Veronica and other temperate plant species?[END_REF][START_REF] Dostál | Effect of three mound-building ant species on the formation of soil seed bank in mountain grassland[END_REF]).

6.4. Agrostis stolonifera is documented [START_REF] Grime | Comparative Plant Ecology: A Functional Approach to Common British Species[END_REF] to have a "competitiveruderal" ecological strategy in the well-known C-S-R (competition-stress-ruderality) system of plant strategies or functional types [START_REF] Grime | Evidence for the existence of three primary strategies in plants and its relevance to ecological and evolutionary theory[END_REF](Grime , 1988[START_REF] Grime | Plant Strategies, Vegetation Processes, and Ecosystem Properties[END_REF], which thus includes weedy characteristics [START_REF] Booth | Weed Ecology in Natural and Agricultural Systems[END_REF][START_REF] Baker | Characteristics and modes of origin of weeds[END_REF][START_REF] Baker | The migration of weeds[END_REF][START_REF] Baker | The evolution of weeds[END_REF][START_REF] Keeler | Implications of weed genetics and ecology for the deliberate release of genetically engineered crop plants[END_REF][START_REF] Keeler | Can genetically engineered crops become weeds?[END_REF] in relation to competitors and disturbance (e.g. [START_REF] Schippers | Herbaceous plant strategies in disturbed habitats[END_REF][START_REF] Hill | Hemeroby, urbanity and ruderality: Bioindicators of disturbance and human impact[END_REF][START_REF] Wilcox | Early plant succession on former arable land[END_REF][START_REF] Marshall | Interference between sown grasses and the growth of rhizome of Elymus repens (couch grass)[END_REF][START_REF] Goldsmith | Interaction (competition) studies as a step towards the synthesis of sea-cliff vegetation[END_REF][START_REF] Owen | Plant species and community responses to sand burial on the machair of the Outer Hebrides, Scotland[END_REF][START_REF] Lenssen | Does disturbance favour weak competitors? Mechanisms of changing plant abundance after flooding[END_REF]. The plant's roots (cf. Fitts 1925a;[START_REF] Murphy | Turfgrass root systems evaluated using the minirhizotron and video recording methods[END_REF][START_REF] Boeker | Root development of selected turfgrass species and cultivars[END_REF]; [START_REF] Lehman | Heritability estimates of creeping bentgrass root systems grown in flexible tubes[END_REF][START_REF] Steer | Shifts in the microbial community in rhizosphere and non-rhizosphere soils during the growth of Agrostis stolonifera[END_REF][START_REF] Beard | Relationship of creeping bentgrass (Agrostis palustris Huds.) root growth to environmental factors in the field[END_REF][START_REF] Ralston | Effect of temperature and water-table depth on the growth of creeping bentgrass roots[END_REF][START_REF] Krans | Some effects of subirrigation on bentgrass during heat stress in the field[END_REF][START_REF] Bowman | Root architecture affects nitrate leaching from bentgrass turf[END_REF][START_REF] Christians | Root activity in creeping bentgrass thatch[END_REF]) and shoots (stolons) actively forage in space, exploiting pockets of nutrient enrichment and vegetation gaps [START_REF] Crick | Morphological plasticity and mineral nutrient capture in two herbaceous species of contrasted ecology[END_REF][START_REF] Hunt | Growth and root-shoot partitioning in eighteen British grasses[END_REF][START_REF] Grime | Comparative Plant Ecology: A Functional Approach to Common British Species[END_REF]cf. Glimskär & Ericsson 1999;[START_REF] Glimskär | Estimates of root system topology of five plant species grown at steady-state nutrition[END_REF]. Being a clonal perennial organism, the plant can function in a modular way (e.g. Jónsdóttir 1991bJónsdóttir , 1991a;;[START_REF] Tamm | Classifying clonal growth forms based on vegetative mobility and ramet longevity: A whole community analysis[END_REF], with the leafy plantlets (rooted tillers) along a stolon able to become somewhat independent [START_REF] Agha | Effect of local shading and drought both singly and combined on tiller ramets of Agrostis stolonifera L[END_REF][START_REF] Marshall | Mineral nutritional inter-relations amongst stolons and tiller ramets in Agrostis stolonifera L[END_REF]. Consequently, severed stolons or dispersed pieces of jointed stolon are more readily able to establish new plants [START_REF] Boedeltje | Plant dispersal in a lowland stream in relation to occurrence and three specific life-history traits of the species in the species pool[END_REF][START_REF] Widén | The Genus Agrostis in Eastern Fennoscandia. Taxonomy and Distribution[END_REF]cf. Fitts 1925b;[START_REF] Carrier | Vegetative planting[END_REF][START_REF] Carrier | The vegetative method of planting creeping bent[END_REF].

6.5. The species often reproduces primarily vegetatively, spreading or "creeping" by the stolons (cf. [START_REF] Hoeltzener | Tillering patterns in turfgrass species with distinct clonal tactics[END_REF], which may become divided (e.g. by an animal or machine) and continue growing from the stolon nodes as separate plants, ramets with the same genotype (thus clones). In the intensely managed golf course, the turf is mowed very short often (cf. [START_REF] Mccartney | Auditing non-hazardous wastes from golf course operations: Moving from a waste to a sustainability framework[END_REF] and sometimes cut vertically to reduce thatch; consequently flowering is curtailed but fragmentation of plants is likely. In unstable natural habitats such as dunes, more seeds are produced than in grassier habitats such as a meadow. A single panicle may average from 100-200 to as many as 1,480 florets in semi-natural habitats [START_REF] Prieto-Baena | Pollen production in the Poaceae family[END_REF], and the plant may have panicles at somewhat different stages of maturity. Separate plants of the same genotype (i.e. ramets of the same genet) adjust to their immediate surroundings, so do not necessarily flower simultaneously. For details on these aspects, see [START_REF] Smith | The pattern of distribution of Agrostis and Festuca plants of various genotypes in a sward[END_REF][START_REF] Grime | Comparative Plant Ecology: A Functional Approach to Common British Species[END_REF][START_REF] Boedeltje | Plant dispersal in a lowland stream in relation to occurrence and three specific life-history traits of the species in the species pool[END_REF][START_REF] Kik | Population responses in Agrostis stolonifera to selective forces in inland and coastal habitats[END_REF]Kik et al. 1990aKik et al. , 1990b[START_REF] Kik | Ploidy level and somatic chromosome number variation in Agrostis stolonifera[END_REF]Wilson & Thompson 1989;[START_REF] Purves | Fine-scale spatial structure in a grassland community: Quantifying the plant's-eye view[END_REF][START_REF] Lenssen | Experimental ramet aggregation in the clonal plant Agrostis stolonifera reduces its competitive ability[END_REF][START_REF] Tamm | Classifying clonal growth forms based on vegetative mobility and ramet longevity: A whole community analysis[END_REF]Jónsdóttir 1991aJónsdóttir , 1991b;;[START_REF] Olff | Species dynamics and nutrient accumulation during early primary succession in coastal sand dunes[END_REF]Pakeman & Marshall 1997. 6.6. The flowering period, the seasonal duration (months, weeks or days) of the species bringing forth flowers, varies with the region where the plants occur, for example with a shorter season in the North (e.g. Widén 1971;cf. Heide 1994;[START_REF] Cooper | The inductive requirements for flowering of some temperate grasses[END_REF].

The daily duration (1-4 hrs) of a plant's flowering perhaps differs somewhat among the Agrostis species, but also depends on the day's weather. The anthesis period, when the flower begins and ends shedding its pollen during the 24-hr day, is characteristic of the Agrostis species and fine-tuned to the local conditions. In field-plot or greenhouse and lab studies (e.g. [START_REF] Philipson | A revision of the British species of the genus Agrostis Linn[END_REF][START_REF] Davies | The breeding affinities of some British species of Agrostis[END_REF][START_REF] Ponomarev | The daily rhythm of pollination and its role in the formation of grass species[END_REF][START_REF] Fei | Estimation of pollen viability, shedding pattern, and longevity of creeping bentgrass on artificial media[END_REF], the pollen of creeping bentgrass shed for a few mid-morning hours (exact times differing with locale) or again (but perhaps not usually) in early-mid afternoon, and the pollen grains were viable for no more than 3 hours. Agrostis stolonifera in seminatural habitats may average 545 pollen grains per anther, thus 1,635 per floret, and up to some 2,420,000 per panicle [START_REF] Prieto-Baena | Pollen production in the Poaceae family[END_REF]. The species is mainly sexually outcrossing [START_REF] Davies | The breeding affinities of some British species of Agrostis[END_REF], with some turf strains perhaps obligately so, but it can also self-pollinate (e.g. Belanger et al. 2003b;[START_REF] Tomić | Dry matter and seed production in some inbred lines of three Agrostis species. Herbage Seed as a Key Factor for Improving Production and Environmental Quality[END_REF]) (a highly selfing turf clone was recently detected - [START_REF] Warnke | Isozyme analysis supports allotetraploid inheritance in tetraploid creeping bentgrass (Agrostis palustris Huds.)[END_REF]. Agrostis species in a locality might be reproductively somewhat isolated temporally by the sequence of their daily anthesis, for example with A. canina releasing pollen pre-dawn -dawn, A. stolonifera ± mid-morning, and A. capillaris variously early-mid-late afternoon. Reports for A. gigantea seem to be inconsistent; A. castellana might be earlier than A. stolonifera; and A. vinealis is bimodal with ± pre-dawn and evening shedding (cf. Christoffer 2003).

6.7. In experimental field-plot studies on pollen dispersal and intraspecific (withinspecies) crossing and potential hybridization with other species of Agrostis and with Polypogon (Wipff & Fricker 2001;Belanger et al. 2003b;Christoffer 2003), viable pollen of creeping bentgrass went as far as 1,400 (or 1,417) feet (determined by testing progeny). Calculating from their data on the intraspecific progeny from an experimental array of linearly placed plants, Wipff & Fricker (2001) estimated that plants would be receiving donor pollen and crossing at a 0.02% level up to 4,296 ft (about 0.8 mile) away from the pollen-dispersing (i.e. source) plants. Based on independent data from a fairly similar experiment (Belanger et al. 2003b), pollen dispersal and as well variation in fitness of the pollen donor were considered in a computer model [START_REF] Meagher | Using empirical data to model transgene dispersal[END_REF].

The model found that pollen dispersal and gene introgression (crossing and subsequent diffusion into a different population -cf. [START_REF] Anderson | Introgressive Hybridization[END_REF][START_REF] Anderson | Introgressive hybridization[END_REF][START_REF] Arnold | Natural Hybridization and Evolution[END_REF][START_REF] Arnold | Transfer and origin of adaptations through natural hybridization: Were Anderson and Stebbins right?[END_REF][START_REF] Lanchier | Voter model and biased voter model in heterogeneous environments[END_REF] would be limited at some sites and extensive at others, depending on local conditions such as exposure to the wind. In a large-scale experimental study [START_REF] Watrud | Evidence for landscape level, pollenmediated gene flow from genetically modified creeping bentgrass with CP4 EPSPS as a marker[END_REF][START_REF] Watrud | Long distance pollen-mediated gene flow from creeping bentgrass[END_REF], pollen dispersal and gene flow were found to occur mostly within about 1.25 miles and extend up to 13 miles. All four of the recent studies concluded that hybridization can occur in the field between A. stolonifera and some naturalized or native species (Tables 1 &2). This recent work using molecular techniques thus confirms and enriches earlier reports, from experimental hybridization and the study of chromosomes (e.g. [START_REF] Davies | The breeding affinities of some British species of Agrostis[END_REF]Jones 1956aJones -1956c;;[START_REF] Björkman | Studies in Agrostis and Related Genera[END_REF]Romero García & Blanca 1988;Forde 1991), and from fieldwork and the taxonomic analysis of plants' morphology (e.g. [START_REF] Murbeck | Studier öfver kritiska kärlväxtformer. II De nordeuropeiska formerna af slägtet Agrostis[END_REF][START_REF] Weber | Die mesophilen Straussgraswiesen der Marschen im Mittellaufe der Weser. Mit vergleichenden Ausblicken auf andere Pflanzenvereinen und Lebensgemeinshcaften. Ein Beitrag zur Kenntnis der Grasflur der Wesermarschen[END_REF][START_REF] Malte | The commercial bent grasses (Agrostis) in Canada[END_REF][START_REF] Fouillade | Sur les Agrostis alba, vulgaris, castellana et leurs hybrides[END_REF][START_REF] Juhl | An-euploidie und Systematik bei Agrostis stolonifera L. und Festuca rubra L. aus Schleswig-Holstein[END_REF]Bradshaw 1958;[START_REF] Widén | The Genus Agrostis in Eastern Fennoscandia. Taxonomy and Distribution[END_REF]Romero García et al. 1988b;[START_REF] Meerts | [END_REF]).

6.8. Of the 10 (or 11) species of Agrostis with which it is known that A. stolonifera could directly hybridize in USA (Figure 1, Tables 1 &2), the most likely crossing is with A. capillaris, forming A. ×murbeckii; and to a lesser extent crossing with A. gigantea. Agrostis capillaris is most likely to cross with A. castellana, forming A. ×fouilladeana, which can backcross into A. capillaris, and for some years these species and their hybrids were imported unknowingly from New Zealand all misnamed as "colonial bentgrass" (or "Agrostis capillaris") and were widely distributed (section 5.3). The species A. capillaris is also likely to cross with A. gigantea, forming A. ×bjoerkmanii, as found in Rhode Island (section 5.4). The various hybrids are for the most part sterile or with very low fertility, but can be vegetatively vigorous (Table 2). Hybridization and introgression have always been aspects of domestication and improvement of crops and ornamentals (cf. [START_REF] Gepts | A comparison between crop domestication, classical plant breeding, and genetic engineering[END_REF][START_REF] Anderson | The analysis of variation in cultivated plants with special reference to introgression[END_REF]. Various new lab techniques facilitate working with hybrid turfgrasses (e.g. Brilman 2001b;[START_REF] Ovesná | DNA analyses and their applications in plant breeding[END_REF], and efforts are underway to hybridize Agrostis species for traditional reasons such as bringing in disease resistance (e.g. Belanger et al. 2003aBelanger et al. , 2003b[START_REF] Belanger | Dollar spot resistant hybrids between creeping bentgrass and colonial bentgrass[END_REF][START_REF] Li | Identification of a gene in the process of being lost from the genus Agrostis[END_REF].

7.5. Of those four species, Agrostis gigantea (redtop), A. capillaris (colonial bentgrass) and A. castellana (dryland bentgrass) are more closely related to A. stolonifera (Table 2).

Redtop and colonial bentgrass are in general considered more weedy than creeping bentgrass (section 7.4). In the C-S-R system of plant functional types (Grime 1988[START_REF] Grime | Plant Strategies, Vegetation Processes, and Ecosystem Properties[END_REF], A. gigantea also is a competitive-ruderal strategist, whereas A. capillaris is a C-S-R generalist [START_REF] Grime | Comparative Plant Ecology: A Functional Approach to Common British Species[END_REF].

7.6. Some species of Agrostis have a robust capacity to adapt genetically and establish populations tolerant to heavy metals and metalloids, for example on soil at contaminated mine sites [START_REF] Bradshaw | Populations of Agrostis tenuis resistant to lead and zinc poisoning[END_REF][START_REF] Bradshaw | Industrialisation, evolution, and the development of heavy metal tolerance in plants[END_REF][START_REF] Bradshaw | Evolution and stress -genotypic and phenotypic components[END_REF]. All five of the main agronomic species as well as A. vinealis [START_REF] Farrow | The dynamics of natural selection for copper tolerance in Agrostis canina L. subsp. montana Hartm[END_REF]) and the common native A. scabra [START_REF] Archambault | Metal tolerance in Agrostis scabra Willd. from the Sudbury, Ontario area[END_REF][START_REF] Koch | The predominance of inorganic arsenic species in plants from Yellowknife, Northwest Territories, Canada[END_REF] are among these facultative metallophytes. Arsenic, cadmium, copper, lead, manganese, nickel and zinc can be prominently involved, and the plants on a site able to tolerate one or more of these ordinarily toxic substances. The zone between tolerant and nontolerant biotypes can be narrow, and without hybrids. Some of the research (cf. Bone & Agnes 2001;[START_REF] Levin | The recurrent origin of plant races and species[END_REF]) has led to classic evolutionary studies used in teaching (e.g. [START_REF] Briggs | Plant Variation and Evolution, 3rd Ed[END_REF], and to efforts for mine-site reclamation and ecological restoration (e.g. [START_REF] Cotter-Howells | Remediation of contaminated land by formation of heavy metal phosphates[END_REF][START_REF] Lepp | Patterns of soil copper contamination and temporal changes in vegetation in the vicinity of a copper rod rolling factory[END_REF][START_REF] Koe | Agrostis castellana and Agrostis delicatula on heavy metal and arsenic enriched sites in NE Portugal[END_REF]cf. Lesica & Allendorf 1999) and water-quality improvement by biofiltration [START_REF] Hares | Comparison of the heavy-metal content of motorway stormwater following discharge into wet biofiltration and dry detention ponds along the London Orbital (M25) Motorway[END_REF].

Polypogon monspeliensis also has good potential for management of wastewater contaminated with selenium or arsenic [START_REF] Ye | Possible use of constructed wetland to remove selenocyanate, arsenic, and boron from electric utility wastewater[END_REF].

7.7. The repertoire of Agrostis stolonifera [START_REF] Duwensee | Morphologie von Agrostis stolonifera (Poaceae) auf Schwermetallboden[END_REF] includes biotypes tolerant to copper [START_REF] Wu | Aerial pollution and the rapid evolution of copper tolerance[END_REF]Wu et al. 1975;[START_REF] Masarovičová | Effect of copper on growth and chlorophyll content of some herbs[END_REF][START_REF] Briggs | Plant Variation and Evolution, 3rd Ed[END_REF], copper and nickel [START_REF] Jowett | Populations of Agrostis spp. tolerant of heavy metals[END_REF], copper and zinc [START_REF] Wu | Zinc and copper uptake by Agrostis stolonifera, tolerant to both zinc and copper[END_REF], and lead and zinc (Gregory & Bradshaw 1965;cf. Harkot & Czarnecki 1999). A biotype of its hybrid Agrostis ×murbeckii tolerant to zinc has been detected (Gregory & Bradshaw 1965). Creeping bentgrass has also adapted to serpentine [START_REF] Marrs | The response of serpentine and non-serpentine Agrostis stolonifera to magnesium and calcium[END_REF], natural rock outcroppings high in magnesium and low in available calcium and water-holding capacity (Baker et al. 1993), thus an exclusive habitat where some species are unable to survive but some species have evolved and only occur (i.e. endemics).

7.8. Colonial bentgrass (the zinc-tolerant hybrid's other parent) has been studied most, and there are metal-tolerant cultivars for reclamation available commercially [START_REF] Jowett | Population studies on lead-tolerant Agrostis tenuis[END_REF]Gregory & Bradshaw 1965;[START_REF] Jain | Evolutionary divergence among adjacent plant populations. I. The evidence and its theoretical analysis[END_REF][START_REF] Mcneilly | Evolution in closely adjacent plant populations. III. Agrostis tenuis on a small copper mine[END_REF][START_REF] Mcneilly | Evolution in closely adjacent plant populations. IV. Barriers to gene flow[END_REF][START_REF] Antonovics | Evolution in closely adjacent populations. V. Evolution of selffertility[END_REF][START_REF] Jones | Physiological and biochemical basis for heavy metal tolerance in clones of Agrostis tenuis. 6th Colloquium on Plant Analysis and Fertilizer Problems[END_REF][START_REF] Mathys | Vergleichende Untersuchungen der Zinkaufnahme von resistenten und sensitiven Populationen von Agrostis tenuis Sibth[END_REF][START_REF] Gartside | Genetic studies in heavy metal tolerant plants. II. Zinc tolerance in Agrostis tenuis[END_REF][START_REF] Whalley | The potential for evolution of heavy metal tolerance in plants. I: Copper and zinc tolerance in Agrostis tenuis[END_REF][START_REF] Simon | Cadmium tolerance in populations of Agrostis tenuis and Festuca ovina[END_REF][START_REF] Simon | Aspects de la tolerance aux metaux lourds chez Agrostis tenuis Sibth., Festuca ovina L. et Armeria maritima (Mill.) Willd[END_REF][START_REF] Nicholls | Sensitivity of rooting and tolerance to copper in Agrostis tenuis Sibth[END_REF], 1985;[START_REF] Smith | The use of metal-tolerant plant populations for the reclamation of metalliferous wastes[END_REF]Karataglis 1980c;[START_REF] Karataglis | Lead and zinc tolerance of Agrostis capillaris L. and Festuca rubra L. across a mine -pasture boundary at Minera, North Wales[END_REF]Symeonidis et al. 1985a;[START_REF] Meharg | The mechanisms of arsenate tolerance in Deschampsia cespitosa (L.) Beauv. and Agrostis capillaris L[END_REF][START_REF] Watkins | Genetics of arsenic tolerance in Agrostis capillaris L[END_REF][START_REF] De Koe | Measuring maximum root growth instead of longest root elongation in metal-tolerance tests for grasses (Agrostis capillaris, Agrostis delicatula and Agrostis castellana)[END_REF][START_REF] Boon | Effects of a coppertolerant grass (Agrostis capillaris) on an ecosystem of copper-contaminated arable soil[END_REF][START_REF] Meerts | Soil seed banks in a heavy-metal polluted grassland at Prayon (Belgium)[END_REF][START_REF] Griffioen | The role of VA mycorrhiza in the heavy metal tolerance of Agrostis capillaris L. Agriculture[END_REF][START_REF] Griffioen | Mycorrhizal infection of an Agrostis capillaris population on a copper-contaminated soil[END_REF][START_REF] Malcová | Effects of inoculation with Glomus intraradices on lead uptake by Zea mays L. and Agrostis capillaris L[END_REF][START_REF] Rydlová | Effect of Glomus intraradices isolated from Pbcontaminated soil on Pb uptake by Agrostis capillaris is changed by its cultivation in a metal-free substrate[END_REF]. Agrostis capillaris also has biotypes with multiple-metal tolerance, to variously two, three or four metals, and that can have large within-population variability [START_REF] Hertstein | Tolerances of different populations of three grass species to cadmium and other metals[END_REF]Symeonidis et al. 1985b;[START_REF] Karataglis | Combined tolerance to copper, zinc and lead by populations of Agrostis tenuis[END_REF]Karataglis , 1980e, 1980aKarataglis , 1980d;;[START_REF] Humphreys | Relationships between tolerance to heavy metals in Agrostis capillaris L. (Agrostis tenuis Sibth.)[END_REF][START_REF] Turner | The sub-cellular distribution of zinc and copper within roots of metaltolerant clones of Agrostis tenuis Sibth[END_REF][START_REF] Mcneilly | Evolutionary processes in populations of copper-tolerant Agrostis tenuis Sibth[END_REF]). Separate evolutionary adaptations apparently have resulted in the similar metal-tolerance traits of some biotypes [START_REF] Nicholls | The possible polyphyletic origin of copper tolerance in Agrostis tenuis (Gramineae)[END_REF][START_REF] Al-Hiyaly | The effect of zinc contamination from electricity pylons -evolution in a replicated situation[END_REF], 1990, 1993). The other agronomic species and some metal-tolerance studies include A. canina (Gregory & Bradshaw 1965), A. castellana (De Koe et al. 1992;[START_REF] De Koe | Arsenate tolerance in Agrostis castellana and Agrostis delicatula[END_REF][START_REF] Schat | The role of phytochelatins in constitutive and adaptive heavy metal tolerances in hyperaccumulator and non-hyperaccumulator metallophytes[END_REF] and A. gigantea (Hogan et al. 1977a(Hogan et al. , 1977b;;[START_REF] Hogan | Tolerance and toxicity of cobalt, copper, nickel and zinc in clones of Agrostis gigantea Roth[END_REF], 1981;Rauser 1984aRauser , 1984b;;[START_REF] Mclaughlin | The distribution of Agrostis gigantea and Poa pratensis in relation to some environmental factors on a mine-tailings area at Copper Cliff, Ontario[END_REF].

7.9. "What is it about the genome of Agrostis species that they can evolve tolerance to almost any metal anywhere…?" [START_REF] Macnair | Heavy metal tolerance in plants: A model evolutionary system[END_REF]). These species are considered metal excluders rather than hyperaccumulators [START_REF] Memon | Heavy metal accumulation and detoxification mechanisms in plants[END_REF][START_REF] Lasat | Phytoextraction of toxic metals: A review of biological mechanisms[END_REF][START_REF] Pollard | The genetic basis of metal hyperaccumulation in plants[END_REF]). Agrostis capillaris for example avoids cadmium, copper, lead, and zinc by precipitating them in the rhizosphere [START_REF] Dahmani-Muller | Strategies of heavy metal uptake by three plant species growing near a metal smelter[END_REF]. The genetics and physiology of such metal-tolerant plants are now better delineated and precise understanding is becoming possible (e.g. [START_REF] Macnair | The genetics of tolerance in natural populations[END_REF][START_REF] Macnair | The genetics of metal tolerance in vascular plants[END_REF][START_REF] Macnair | The genetics of metal tolerance and accumulation in higher plants[END_REF][START_REF] Schat | Metal-specific patterns of tolerance, uptake and transport of heavy metals in hyperaccumulating and nonhyperaccumulating metallophytes[END_REF][START_REF] Hall | Cellular mechanisms for heavy metal detoxification and tolerance[END_REF][START_REF] Memon | Heavy metal accumulation and detoxification mechanisms in plants[END_REF]. [START_REF] Gepts | A comparison between crop domestication, classical plant breeding, and genetic engineering[END_REF]. There are overviews on genetic engineering for turf and forage grasses (e.g. [START_REF] Wang | Recent advances in genetic transformation of forage and turf grasses[END_REF][START_REF] Wang | Forage and turf grass biotechnology[END_REF][START_REF] Maqbool | Past, present, and future of turfgrass biotechnology[END_REF]Chai & Sticklen 1998;[START_REF] Lee | Turfgrass biotechnology[END_REF], and on transgenic turfgrasses and risk assessment in considering the release of a particular modified plant into the environment (e.g. [START_REF] Johnson | A review of issues pertaining to transgenic turfgrasses[END_REF]Day & Lee 1997). Consequent experiments to better understand aspects of the biology of these species are the pollination and hybrid studies noted above (sections 6.6-6.7). The purpose of other experiments (in the greenhouse or a field trial) (e.g. [START_REF] Lee | Field evaluation of herbicide resistance in transgenic Agrostis stolonifera L. and inheritance in the progeny[END_REF][START_REF] Gardner | Relative fitness of glyphosate-resistant creeping bentgrass lines in Kentucky bluegrass[END_REF][START_REF] Gardner | Lateral spread of glyphosateresistant transgenic creeping bentgrass (Agrostis stolonifera) lines in established turfgrass swards[END_REF][START_REF] Fei | Greenhouse evaluation of fitness-related reproductive traits in Roundup®-tolerant transgenic creeping bentgrass (Agrostis stolonifera L.)[END_REF][START_REF] Reichman | Establishment of transgenic herbicide-resistant creeping bentgrass (Agrostis stolonifera L.) in non-agronomic habitats[END_REF][START_REF] Hart | Response of glyphosate-resistant and glyphosate-susceptible bentgrass (Agrostis spp.) to postemergence herbicides[END_REF]Wang et al. 2003) is to evaluate the transgenic plant's biology for any unexpected effects, or to study the trait put into the plant to see if the result is as desired. Semi-popular and professional trade articles on methods and potential benefits perceived in the genetic engineering of turfgrasses are also available (e.g. [START_REF] Lee | Herbicide-resistant creeping bentgrass[END_REF]Sticklen 2001;[START_REF] Harriman | Turf benefits of biotechnology on the horizon[END_REF][START_REF] Litrenta | Transgenics: The bright future of turfgrass[END_REF]Ostmeyer 2004aOstmeyer , 2004bOstmeyer , 2004c;;[START_REF] Jones | Approval for genetically engineered bentgrass creeps through agency turfs[END_REF]).

Biotechnological Developments in

8.2. In Agrostis, genetic engineering (on an experimental basis in several countries) has succeeded, and next steps are proceeding with at least four species: especially Agrostis stolonifera [START_REF] Krans | Callus induction, maintenance and plantlet regeneration in creeping bentgrass[END_REF][START_REF] Blanche | Improvement in callus growth and plantlets formation in creeping bentgrass[END_REF][START_REF] Tanpo | Callus induction and plant regeneration in creeping bentgrass (Agrostis palustris)[END_REF][START_REF] Terakawa | Plant regeneration from protoplasts isolated from embryogenic suspension cultures of creeping bentgrass (Agrostis palustris Huds.)[END_REF][START_REF] Zhong | Plant regeneration via somatic enbryogenesis in creeping bentgrass (Agrostis palustris Huds.)[END_REF][START_REF] Zhong | Transgenic plants of turfgrass (Agrostis palustris Huds.) from microprojectile bombardment of embryogenic callus[END_REF][START_REF] Sticklen | Genetic transformation in Agrostis palustris Huds. (creeping bentgrass)[END_REF][START_REF] Xiao | Efficient selection and regeneration of creeping bentgrass transformants following particle bombardment[END_REF][START_REF] Basu | Promoter analysis in transient assays using a GUS reporter gene construct in creeping bentgrass (Agrostis palustris)[END_REF][START_REF] Luo | Agrobacterium tumefaciens-mediated creeping bentgrass (Agrostis stolonifera L.) transformation using phosphinothricin selection results in a high frequency of single-copy transgene integration[END_REF][START_REF] Luo | Controlling transgene escape in GM creeping bentgrass[END_REF]Fu et al. 2005a;[START_REF] Wang | Rapid and efficient production of transgenic bermudagrass and creeping bentgrass bypassing the callus formation phase[END_REF], as well as A. canina [START_REF] Pitcher | Transformation of tall fescue and velvet bentgrass using Agrobacterium tumefaciens[END_REF][START_REF] Pitcher | Agrobacterium tumefaciens-mediated transformation of turfgrasses using the selectable marker phosphomannose isomerase[END_REF], A. capillaris [START_REF] Chai | Agrobacteriummediated transformation of herbicide resistance in creeping bentgrass and colonial bentgrass[END_REF][START_REF] Chai | Transgenic plants of colonial bentgrass from embryogenic callus via Agrobacterium-mediated transformation[END_REF] and A. gigantea [START_REF] Asano | Plant regeneration from suspension culture-derived protoplasts of Agrostis alba L. (redtop)[END_REF][START_REF] Asano | Electroporation-mediated and silicon carbide fiber-mediated DNA delivery in Agrostis alba L. (redtop)[END_REF][START_REF] Asano | Transgenic plants of Agrostis alba obtained by electroporation-mediated direct gene transfer into protoplasts[END_REF]. Various lab methods now exist (e.g. van den [START_REF] Van Den Eede | The relevance of gene transfer to the safety of food and feed derived from genetically modified (GM) plants[END_REF][START_REF] Rakoczy-Trojanowska | Alternative methods of plant transformation -a short review[END_REF] to deliver gene(s) (DNA) of interest into a plant's genome. At least five methods can transform material of Agrostis species: (1) microprojectile bombardment or biolistic transformation [START_REF] Hartman | Herbicide-resistant turfgrass (Agrostis palustris Huds.) by biolistic transformation[END_REF][START_REF] Lee | Herbicide-resistant creeping bentgrass[END_REF][START_REF] Lee | Herbicide-resistant transgenic creeping bentgrass[END_REF];

(2) silicon carbide fibers or whiskers [START_REF] Asano | Electroporation-mediated and silicon carbide fiber-mediated DNA delivery in Agrostis alba L. (redtop)[END_REF][START_REF] Dalton | Transgenic plants of Lolium multiflorum, Lolium perenne, Festuca arundinacea and Agrostis stolonifera by silicon carbide fibre-mediated transformation of cell suspension cultures[END_REF]);

(3) electroporation [START_REF] Asano | Electroporation-mediated and silicon carbide fiber-mediated DNA delivery in Agrostis alba L. (redtop)[END_REF][START_REF] Asano | Transgenic plants of Agrostis alba obtained by electroporation-mediated direct gene transfer into protoplasts[END_REF][START_REF] Asano | Herbicide-resistant transgenic creeping bentgrass plants obtained by electroporation using an altered buffer[END_REF][START_REF] Sugiura | Generation of herbicideresistant creeping bentgrass (Agrostis palustris Huds.) plants by electroporationmediated direct gene transfer into protoplasts[END_REF]); (4) polyethylene glycol [START_REF] Lee | Transformation and regeneration of creeping bentgrass (Agrostis palustris Huds.) protoplasts[END_REF][START_REF] Lee | Herbicide-resistant transgenic creeping bentgrass[END_REF]; and (5) Agrobacterium tumefaciens [START_REF] Pitcher | Transformation of tall fescue and velvet bentgrass using Agrobacterium tumefaciens[END_REF][START_REF] Yu | Agrobacterium-mediated transformation of creeping bentgrass using GFP as a reporter gene[END_REF][START_REF] Chai | Embryogenic callus induction and Agrobacterium-mediated transformation in bentgrass (Agrostis spp.)[END_REF][START_REF] Kim | Factors affecting the transformation of bentgrass (Agrostis spp.) based on Agrobacterium tumefaciens[END_REF][START_REF] Pitcher | Agrobacterium tumefaciens-mediated transformation of turfgrasses using the selectable marker phosphomannose isomerase[END_REF][START_REF] Chai | Agrobacteriummediated transformation of herbicide resistance in creeping bentgrass and colonial bentgrass[END_REF][START_REF] Chai | Transgenic plants of colonial bentgrass from embryogenic callus via Agrobacterium-mediated transformation[END_REF][START_REF] Luo | Agrobacterium tumefaciens-mediated creeping bentgrass (Agrostis stolonifera L.) transformation using phosphinothricin selection results in a high frequency of single-copy transgene integration[END_REF][START_REF] Han | Production of transgenic creeping bentgrass Agrostis stolonifera var. palustris plants by Agrobacterium tumefaciens-mediated transformation using hygromycin selection[END_REF][START_REF] Hu | Obtaining insect-resistant transgenic plants of creeping bentgrass (Agrostis stolonifera L.) mediated by Agrobacterium tumefaciens[END_REF]). The traits of interest (cf. Chandler The species are numbered in alphabetical order, so for example the number 30 = Agrostis stolonifera. See also Figure 1,and [START_REF] Master | Vanishing assets: Conservation status of U.S. species[END_REF]; in taxon's native range: G1 = critically rare; G2 = rare; G3 = vulnerable; G4 = apparently secure; G5 = widespread, abundant and secure; T (trinomial) = ranking for a botanical variety or subspecies; Q = a question (problem) in taxonomy (TNC/NatureServe 2002, ranks in brackets by APHIS/BRS). WEED CONCERN IN COUNTRY: S = Serious, P = Principal, C = Common, X = Behaves as weed, importance unknown [START_REF] Holm | A Geographical Atlas of World Weeds[END_REF] [START_REF] Dudley | Exotic plant invasions in California riparian areas and wetlands[END_REF]; and for Polypogon monspeliensis: Troublesome -California (Zedler & Kercher 2004). HYBRIDS: F1 = F 1 with sterility/fertility not reported; f = fertile (without details), s = sterile, v = variable in crossing or in reports (perhaps sterile and low or higher fertilities).

Species (Taxa)

Common Names 2006[START_REF] Fna | Grass Manual and Flora of North America [FNA] Grass Volumes: Publication Plans: Names and Mapping Synonyms[END_REF], but not Tzvelev 1983[1976], [START_REF] Probatova | Novye taksony sem. Poaceae s Dalnego Vostoka SSSR [New taxa of the Poaceae from the Far East of the USSR][END_REF]etc., Kharkevich et al. 2003[1985]

  Agrostis alba L." had to become a technical synonym for that species (and thus no longer available for use).(3) On a different theme, the determination and classification of biological entities, it was not obvious that the plants called A. stolonifera and the plants now called A. gigantea represented two species rather than variations of one biological species. If considered to be one (e.g.[START_REF] Malte | The commercial bent grasses (Agrostis) in Canada[END_REF]), the species' name had to be the earlier A. stolonifera, and the redtop plants were then called A. stolonifera

	), A. palustris Huds.
	(1762), and A. stolonifera var. compacta Hartm. (1832). (2) The name Agrostis gigantea
	Roth (1788) for redtop was its second name, regarded for a while as superfluous.
	However, redtop's earlier name of Agrostis alba L. (1753) unfortunately was based
	nomenclaturally (i.e. standardized) on a specimen found to be a Poa nemoralis L. (wood
	bluegrass), and so "

var. major (Gaudin) Farw. Within this broadly circumscribed single species, all of the more "creeping" bentgrasses would be called A. stolonifera var. stolonifera, or these plants could still be regarded to have additional inherently distinct populations such as A. stolonifera var. palustris (Huds.) Farw. or A. stolonifera var. compacta. (4) With increasing familiarity and scientific study, Agrostis gigantea has become understood to be a biologically distinct species (as Linnaeus had discerned; Table

2

), a hexaploid with rhizomes (underground stems) and panicles that stay open after flowering, whereas A. stolonifera is a predominant tetraploid with stolons and panicles that close after flowering. Similarly, Agrostis castellana has been recognized to be biologically distinct from A. capillaris, and A. vinealis distinct from A. canina. (5) On the other hand, from thorough study of many native European populations of Agrostis stolonifera (sensu stricto, i.e. creeping bentgrass in the narrow sense excluding "var. major"), broad ecological amplitude and extensive variability have been found without substantial inherent discontinuites (sections 3.1-3.2) (e.g.

Kik et al. 1990a;[START_REF] Vergara | AFLP analyses of genetic diversity in bentgrass[END_REF] cf.

  Agrostis 8.1. Many publications are providing substantial current overviews on aspects of the improvement of plants for various purposes and genetic engineering. Recent summaries include technological developments over 100 years and prospects (e.g. McCown 2003; Chandler 2003), and comparing crop domestication, conventional plant breeding, and genetic engineering

Table 1 . Summary on Agrostis (31-35 spp.) and Polypogon (8 spp.) in USA and Canada

 1 (5/31/2006) (see "White Paper: Perspective on Creeping Bentgrass, Agrostis stolonifera L." -USDA/APHIS/BRS, B. MacBryde)

  Table 2 for further information.

ORIGIN or INTEGRATION IN FLORA, and USE (O/I, U): Nv = Native, Nz = Naturalized, I = Introduced, extent of naturalization unknown; A = Agronomic. NATIVE RANK

  , A Geographical Atlas of World Weeds); H&S:Häfliger & Scholz 1981, Grass Weeds 2; and these additional references for Agrostis stolonifera: Serious -Australia pastures and/or crops (e.g.[START_REF] Brown | Biodiversity and potential utilization of blowngrasses (Agrostis spp.) in lowland Victoria[END_REF], Moderate invasive threat -California coast in wetlands

, U Distribution in USA & Canada Native Rank Weed Concern Hybrids With Other Species / Comments on Taxonomy &/or Range

  Podagrostis humilis perSoreng et al. 2003, FNA 2005. Incl. A. thurberiana (W US, Can.) per[START_REF] Harvey | Agrostis[END_REF][START_REF] Fna | Grass Manual and Flora of North America [FNA] Grass Volumes: Publication Plans: Names and Mapping Synonyms[END_REF][START_REF] Harvey | Agrostis, Bent grass[END_REF], Welsh et al. 1993 q.v., or [as P. thurberiana] Soreng et al. 2003; cf. Biek 2000). Intergrades with #1[START_REF] Harvey | Agrostis L[END_REF]).

		Henderson's Bentgrass	Nv	CA-OR	[G2?]		Incl. A. aristiglumis (endemic CA);
	A.S. Hitchcock						perhaps 15 & 22 related or one sp.
							(Harvey 2004; Beetle 1945).
	16 Agrostis hooveri	Hoover's Bentgrass	Nv	CA endemic	[G2]
	Swallen
	17 Agrostis howellii	Howell's Bentgrass	Nv	OR endemic	G2		Also WA? (FNA 2006 draft map).
	Scribner
	Hybrids With A. stolonifera F1 F1 with 34; sF1 with 5, and 13 (rare?). F1; rare Syn. of Podagrostis aequivalvis Syn. of 19 Agrostis hyemalis 1 Agrostis aequivalvis (Trinius) Trinius Arctic or Northern Bentgrass Nv AK to OR; Canada (BC) G5? 18 Agrostis humilis Vasey Mountain or Alpine Bentgrass Nv W US (AK to NM); W Canada [G5] Winter Bentgrass Nv E US; PR; E Canada G5 per Soreng et al. 2003, FNA 2005. (Walter) Britton, (Ontario) Intergrades with 18 (Harvey 2004). 2 Agrostis anadyrensis Anadyr Bentgrass Nv AK? Sterns & Poggenburg [G4?] 20 Agrostis idahoensis Idaho Redtop or Bentgrass Nv, (A) W US; W Canada [G5?] Hybrid origin? (Welsh et al. 1993). Soczava 3 Agrostis avenacea J.F. Gmelin Pacific Bentgrass, Fairy Blowngrass Nv?, Nz Nv? HI, Nz CA, TX, SC, OH [G4G5] Invasive in California vernal pools (Zedler & Black 2004) Nash Syn. of Lachnagrostis filiformis Nv Australasia, New Guinea, Trinius WV & TN to SC; Canada per Soreng et al. 2003, FNA 2005. 21 Agrostis mertensii Northern or Arctic Bentgrass Nv AK-CO, ME-NY, G5
	22 Agrostis microphylla	Small-leaved Bentgrass	Nv	WA to CA & NV;	G4		perhaps Easter Island, HI Perhaps 15 conspecific (Harvey 2004).
	4 Agrostis blasdalei Steudel	Blasdale's or Cliff Bentgrass	Nv	CA endemic Canada (BC)	G2		F1 with 10 (Harvey 2004).
	A.S. Hitchcock 23 Agrostis nebulosa	Cloudgrass	I, (Nz) Nz in CA, OH,	[G3G4]		Incl. A. blasdalei var. marinensis Nv Iberian Peninsula
	Boissier & Reuter			perhaps elsewhere			per Harvey 1993, 2004 & CNPS
	24 Agrostis oregonensis	Oregon Redtop or Bentgrass	Nv	W US (CA to WY & AK);	G4		2001, 2005' but not Gov. California.
	5 Agrostis canina Vasey	Velvet Bentgrass	Nz, A	HI, OR, MN, MI, ME to TN; W Canada	G5	C (Finland, Russia); in H&S	sF1 with 21, and with 34	vF1; rare?
	Linnaeus 25 Agrostis pallens	Leafy or Dune Bentgrass,	Nv	Canada (BC & E) CA to WA & MT, MA;	[G4G5]		fF1 with 14. Incl. A. diegoensis	F1
	6 Agrostis capillaris Trinius	Colonial (or Rhode Island) Seashore Bentgrass	Nz, A	Most E & W states but few W Canada	[G5]	P (Chile, NZ, Russia), C, X;	vF1 & vF2 with 7, and with 13; F1 (cf. Harvey 1993, 2001; Biek 2000).	vF1; frequent?
	Linnaeus 25A Agrostis pauzhetica	Bentgrass, Browntop Thermal Bentgrass	Nv	in midcontinent; Canada CA & WY	[G1G2]	in H&S	with 8 and probably 14; F1 s? with 34
	7 Agrostis castellana Probatova	Dryland (incl. Highland)	(Nz), A WA, OR, CA, AL,	[G5]	X (Australia, NZ, Portugal)	Confounded with 6, q.v. (& Table 2) vF1; infreq.?
	Boissier & Reuter	Bentgrass		many states?; Canada?
	8 Agrostis clavata	Clubbed Bentgrass	Nv	AK; Canada (Yukon)	G4G5	C (Japan)	F1 with 6 and 29
	Trinius
	9a Agrostis clivicola	Coastal Bluff Bentgrass	Nv	CA endemic	G3?T3?Q		Syn. of 10 per Harvey 1993, 2004,
	Crampton var. clivicola						but not Kartesz 2006
	9b Agrostis clivicola	Point Reyes Bentgrass	Nv	CA endemic	G3?T1Q		Syn. of 10 per Harvey 1993, 2004,
	var. punta-reyesensis						FNA 2005, but not Kartesz 2006
	Crampton
	10 Agrostis densiflora	Dense-flowered Bentgrass	Nv	CA-OR	G3G4		F1 with 4 (Harvey 2004); F1?
	Vasey						with 12 (Carlbom 1967, p. 88).
	11 Agrostis elliottiana	Elliott's Bentgrass	Nv	Mainly S & mid-latitude US	G5
	J.A. Schultes			(FL-PA-KS to CA, & MA, ME)
	12 Agrostis exarata	Spike Bentgrass	Nv	Mostly W US (including	G5	C (Japan)	F1? with 10; F1 with 13?	sF1 with 30?
	Trinius			HI), also KY, VT; W Canada			(Welsh et al. 1993) & 29
	13 Agrostis gigantea	Redtop, Black Bentgrass	Nz, A	All states (including HI);	[G5]	C, X (Australia, Canada, NZ);		vF1;
	Roth			Canada		in H&S		occasional?
	14 Agrostis hallii Vasey	Hall's Bentgrass	Nv	CA-OR, & WA?	G4G5		fF1 with 25; and F1 with probably 6

O/I

vF1 & vF2 with 6 (occasional?); sF1 with 21 (rare?); 13 or perhaps 30 (i.e., A.s. s.l.) F1 with 12

[START_REF] Welsh | A Utah Flora[END_REF] cf. Harvey 2004

).

United States Department of Agriculture Animal and Plant Health Inspection Service Biotechnology Regulatory Services Riverdale, Maryland May 31, 2006

							5 5
							Agrostis
						34	vinealis Agrostis	canina	4	Agrostis blasdalei
				33		6	Agrostis
					Agrostis	capillaris
					variabilis
							Polypogon 7P	Polypogon 8P
			32				monspeliensis	viridis
			Agrostis trinii	?		Polypogon 3P	7	Agrostis castellana
							fugax
	29	Agrostis				8	Agrostis
		scabra				clavata
							30
							Agrostis
		25	Agrostis pallens				stolonifera	?	10	Agrostis
							densiflora
							?
			21	Agrostis			?
				mertensii
							12	Agrostis
							exarata
					20	Agrostis
					idahoensis
							14	Agrostis hallii	13	gigantea Agrostis	?

, Koyama 1987 F1 33 Agrostis variabilis Rydberg Alpine or Mountain Bentgrass Nv W US; W Canada (BC, AB) G5 F1? (≈ 20) with 29 (Welsh et al. 1993) Figure 1. Known natural and/or potentially natural (spontaneous) interspecific crosses among species of Agrostis and/or Polypogon in the United States and Canada (details in Tables 1 & 2 of "White Paper: Perspective on Creeping Bentgrass, Agrostis stolonifera L." -USDA/APHIS/BRS, B. MacBryde, ver. 5/31/2006).

Much of the information in this review is obtainable only by continually noting the references as they are cited and reading the titles.

-Wipff & Fricker 2001, but not Christoffer 2003.6 × 7(some 2n = 35): Partially fertile(Sell & Murrell 1996). Backcrossing into #6 in New Zealand[START_REF] García | The F 1 hybrids of A. stolonifera and A. capillaris[END_REF], Edgar & Connor 2000). Formerly Agrostis ×fouilladei, a nomen nudum[START_REF] Lambinon | Agrostis ×fouilladeana Lambinon & Verloove, nothospec. nov., en Belgique et dans le nord de la France[END_REF]).

USDA/APHIS/BRS (B. MacBryde, ver. 5/31/2006

Schreber Brown Bentgrass

Nz, Nv Nz several states?, Nv AK; Canada? G5?

F1 with 21; F1 s? with 6; sF1 with 5 vF1; rare? 1956b, 1956c;[START_REF] Warnke | Isozyme analysis supports allotetraploid inheritance in tetraploid creeping bentgrass (Agrostis palustris Huds.)[END_REF]; also, at least in Europe [START_REF] Harvey | Agrostis L[END_REF]), 2n = 5x = 35, A 2 A 2 A 3 A 3 A 3 or A 2 A 2 A 2 A 3 A 3 and 2n = 6x = 42, A 2 A 2 A 2 A 3 A 3 A 3 Also aneuploidy (frequent), B-chromosomes, aneusomaty [START_REF] Björkman | Chromosome studies in Agrostis[END_REF][START_REF] Kik | Ploidy level and somatic chromosome number variation in Agrostis stolonifera[END_REF], Frey 1997).

Reproduction of Agrostis stolonifera often mainly vegetative, by stolons (e.g. Kik et al. 1990b[START_REF] Kik | The distribution of cytotypes in ecologically contrasting populations of the clonal perennial Agrostis stolonifera[END_REF]. Sexual reproduction predominately outcrossing [START_REF] Davies | The breeding affinities of some British species of Agrostis[END_REF]Belanger et al. 2003b), with some cultivars perhaps obligately so [START_REF] Warnke | Isozyme analysis supports allotetraploid inheritance in tetraploid creeping bentgrass (Agrostis palustris Huds.)[END_REF]cf. Belanger et al. 2003b). Also highly selffertile clone, and selfing has been utilized [START_REF] Warnke | Isozyme analysis supports allotetraploid inheritance in tetraploid creeping bentgrass (Agrostis palustris Huds.)[END_REF][START_REF] Tomić | Dry matter and seed production in some inbred lines of three Agrostis species. Herbage Seed as a Key Factor for Improving Production and Environmental Quality[END_REF]).

2n = 2x = 14 reported by [START_REF] Tomić | Genetic resource of five Agrostis species present in Serbia and their utilisation[END_REF]; further taxonomic study may be warranted.

A. blasdalei Blasdale's or Cliff Bentgrass

Native California (endemic).

2n = 6x = 42 [START_REF] Harvey | Agrostis L[END_REF]). 4 × 10 4 × 10: USA: west-central CA [START_REF] Harvey | Agrostis L[END_REF]).

5 2n = 4x = 28, 6x = 42, and 8x = 56 [START_REF] Harvey | Agrostis L[END_REF], Frey 1997[START_REF] Taylor | Flora of the Queen Charlotte Islands, Part 2, Cytological Aspects of the Vascular Plants[END_REF]). 30 × 12 (apparently); (30? or) 13 × 12; 10 × 12 (perhaps); 12 × 29 30 × 12: NW USA? (Carlbom 1967).

(30? or) 13 × 12: USA: UT? [START_REF] Welsh | A Utah Flora[END_REF]. 12 × 29: USA: UT? [START_REF] Welsh | A Utah Flora[END_REF].

10 × 12 (perhaps, as F 1 seeds not grown out).

× 12:

Suspected in WA and OR, sterile (Carlbom 1967, pp. 109-110, 112). 2n = 6x = 42 [START_REF] Harvey | Agrostis, Bent grass[END_REF]). 6 × 14 (probable); 14 × 25 (apparently).

6 × 14: NW USA? (Pendergrass 2001). 14 × 25: NW USA? [START_REF] Carlbom | A Biosystematic Study of Some North American Species of Agrostis L. and Podagrostis[END_REF]). 2n = 2x = 14 [START_REF] Tinney | Chromosome behavior in Agrostis nebulosa[END_REF], Romero García & Blanca 1988, Frey 1997). 30 × 23 (no transgenic flow found in field test - Christoffer 2003).

No transgenic flow found (Christoffer 2003). 2n = 6x = 42, 8x = 56 (Harvey 1993, 2004, Frey 1997).

× 25

NW USA? [START_REF] Carlbom | A Biosystematic Study of Some North American Species of Agrostis L. and Podagrostis[END_REF]. 30 × 25 (transgenic test -Wipff & Fricker 2001, Christoffer 2003).

× 25:

Transgenic test: field - Wipff & Fricker 2001, Christoffer 2003. 14 × 25: Probable (and fertile) in OR (Carlbom 1967, pp. 98, 126) 2n = 4x = 28, A 1 A 1 A 2 A 2 or A 1 A 1 A 1 A 1 (somewhat as autotetraploid with 2x A. canina-like ancestry, or perhaps from cross of 4x A. canina [if such] and A. capillaris -Jones 1956b, cf. Romero García et al. 1988b, but the "A. canina" ploidy over 2x may not be A. canina -Romero García & Blanca 1988;cf. Vergara & Bughrara 2003). 30 × 34; 6 × 34 (A. ×sanionis); 21 × 34 All three: Fennoscandia, with 30 × 34 rare (Widén 1971); both 30 × 34 & 6 × 34: UK [START_REF] Hubbard | Grasses: A Guide to Their Structure, Identification, Uses, and Distribution in the British Isles, 3rd Ed[END_REF], Sell & Murrell 1996). 30 × 34;5 × 34 (Davies 1953[START_REF] Björkman | Chromosome studies in Agrostis[END_REF]); 6 × 34 (Davies 1953, but probably most were not hybrids per Widén 1971); × 34 [START_REF] Björkman | Chromosome studies in Agrostis[END_REF].

30 × 34 (2n = 28): Readily crossing per Harvey 2004; experimental hybrids completely sterile (Bradshaw 1975a); a few viable seeds [START_REF] Davies | The breeding affinities of some British species of Agrostis[END_REF]. Björkman 1954 also made a cross of 5x #30 and #34. 5 × 34 (2n = 21): Quite sterile [START_REF] Widén | The Genus Agrostis in Eastern Fennoscandia. Taxonomy and Distribution[END_REF]). 6 × 34 (2n = 28): Unclear facility of crossing in nature and experimentally (readily crossing per Harvey 2004); most experimental "hybrids" [START_REF] Davies | The breeding affinities of some British species of Agrostis[END_REF], Jones 1956b) likely instead selfs of #34 or #6 [START_REF] Widén | The Genus Agrostis in Eastern Fennoscandia. Taxonomy and Distribution[END_REF], Bradshaw 1975a). Probably natural hybrids sterile [START_REF] Widén | The Genus Agrostis in Eastern Fennoscandia. Taxonomy and Distribution[END_REF], Sell & Murrell 1996). 21 × 34 (2n = 42) [START_REF] Widén | The Genus Agrostis in Eastern Fennoscandia. Taxonomy and Distribution[END_REF]. [START_REF] Herbst | Notes on the grasses of Hawai ׳i: New records, name changes, and corrections[END_REF][START_REF] Barkworth | Polypogon Desf[END_REF], also Wipff & Fricker 2001).

Polypogon fugax

Sp.: 2n = 42 [START_REF] Björkman | Chromosome studies in Agrostis[END_REF]; native Iraq to Burma; not found in Hawaii since 1916 [START_REF] Herbst | Notes on the grasses of Hawai ׳i: New records, name changes, and corrections[END_REF], nor in CA or OR for many decades [START_REF] Barkworth | Polypogon Desf[END_REF]).

Intergeneric hybrid [START_REF] Björkman | Studies in Agrostis and Related Genera[END_REF] 2n = 4x = 28 [START_REF] Tutin | Agrostis L[END_REF], Sell & Murrell 1996). Sp.: 2n = 14, 28, 35 & 42 (Barkworth 2004[START_REF] Giraldo-Cañas | El género Polypogon (Poaceae: Pooideae) en Colombia[END_REF][START_REF] Harvey | Agrostis, Bent grass[END_REF]; native S Europe to Turkey.

Polypogon monspeliensis

UK, France, Spain (Sell & Murrell 1996[START_REF] Hubbard | Grasses: A Guide to Their Structure, Identification, Uses, and Distribution in the British Isles, 3rd Ed[END_REF][START_REF] Paunero | Las agrostídeas españolas[END_REF]; Canada (BC), W & SE USA [START_REF] Kartesz | Synthesis of the North American Flora[END_REF]; Chile (Rúgolo de Agrasar & Molina 1997b); New Zealand (Esler 1988, Edgar & Connor 2000); Australia [START_REF] Weiller | Rúgolo de Agrasar & Molina[END_REF](Weiller et al. -[2003]]); China (Qian & Sun 1998). 30 × Pm (transgenic test -Christoffer 2003).

× Pm:

Almost complete pollen and seed sterilities. Vigor varies, perhaps because of hybridization with different ecotypes of #30, which is more vigorous (Bradshaw 1975b;cf. Welsh et al. 1993) [START_REF] Barkworth | Polypogon Desf[END_REF][START_REF] Giraldo-Cañas | El género Polypogon (Poaceae: Pooideae) en Colombia[END_REF][START_REF] Harvey | Agrostis, Bent grass[END_REF]; native Eurasia.

UK, rare [START_REF] Hubbard | Grasses: A Guide to Their Structure, Identification, Uses, and Distribution in the British Isles, 3rd Ed[END_REF], Sell & Murrell 1996); USA: UT? [START_REF] Welsh | A Utah Flora[END_REF]. 30 × Pv (transgenic test -Christoffer 2003).

× Pv:

Experimental crossing facility uncertain. Parents highly self-incompatible; hybrid has complete pollen sterility (Bradshaw 1975b[START_REF] Hubbard | Grasses: A Guide to Their Structure, Identification, Uses, and Distribution in the British Isles, 3rd Ed[END_REF]). Short-lived perennial (Sell & Murrell 1996) [START_REF] Barkworth | Polypogon Desf[END_REF]).