Introduced Forages

1. Crested wheatgrass seeding- Traditionally range has been defined so as to include grazing lands that were reseeded to non-native (= introduced) plants that are managed as if they are native (or naturalized) species and manipulated more by extensive ecological inputs, especially grazing management, than by intensive agronomic methods. The case exemplar for this inclusion of introduced species into the category of range plants and native grazinggrounds was probably the reseeeding of abandoned crop land in the Northern Great Plants and Intermountain Region of North America to the Eurasian bunchgrasses known as crested wheatgrass (Agropyron desertorum and A. cristatum).Crested wheatgrass was introduced from Russia and Siberia and used to reseed "go-back land" (abandoned farm land undergoing secondary succession) and marginal crop fields during the first third of the Twentieth Century. These are still, acreage- wise, the major introduced range plants in North America. This is the Berger Tract of crested wheatgrass south of Twin Falls Idaho. It was primarily desert crested wheatgrass (A. desertorum). October.

No FRES or Kuchler designation because this is not native or potential natural vegetation. (Refer to Literature Review for the limitations of limiting the designation of range strictly to indigenous species.) SRM 614 (Crested Wheatgrass). It should be remembered that some of native range plants do return by secondary succession to lands reseeded to crested wheatgrass. This is especially the case for shrubs like big sagebrush (shown here invading what was once likely a bluebunch wheatgrass-big sagebrush shrub steppe). Rabbitbrush, greasewood, and horsebrush are other examples of the inevitable invasion of woody plants into grasslands in absence of recurrent fire. These shrubs make important contributions to animal diets and they should be recognized as part of the potential natural vegetation (that is, of course, at population levels below those on ecologically deteriorated ranges caused by past cultivation, overgrazing, underburning, etc.).

3. The way it looks- Stand of crested wheatgrass (A. cristatum) on previously farmed land in the Central Great Plains showing the physiogonomy of a typical population of this species as well as the extent of bare ground in interspaces among plants of this highly cespitose species. Even with this relative amount of bare soil surface there is adequate cover (probably both basal and foliar are important) by crested wheatgrass to protect the soil from what would otherwise be the ravishes of wind- and water-caused erosion.

Platte County, Wyoming Late June: peak standing crop (or nearly so) and height of anthesis (see next slide).

4. Exchanging gametes in the Great Plains breeze- Peak anthesis in crested wheatgrass (A. cristatum) in a seeding on an abandoned field in the Central Great Plains. These spikes were on shoots of plants introduced in the preceding slide. Whoever thought that flowers of grasses were "dull", "drab", "boring" or in some way inferior to inflorescences of "real flowers" never took the time to enjoy the grandure of blooming grass such as in this introduced "salvation species".

Platte County, Wyoming Late June: peak standing crop (or nearly so) and height of anthesis.

5. For those who missed it the first- Closer-in photographs of spikes of Agropyron ciristatum in full anthesis. More detail for students in Agrostology and, coincidentially, for those who think that it "ain't" a flower unless its the size of a tea mug with brilliant, dazzling colors. These shoots were of plants in the stand fetured in the two preceding photographs-and-captions.

Platte County, Wyoming Late June: peak standing crop (or nearly so) and height of anthesis.

8.   Stand of immature crested wheatgrass ( mostly A. cristatum)- Crested wheatgrass at the phonological stage known as the boot stage. This is the stage of growth in grasses and grasslike plants when the inflorescence is enclosed in the sheath of the uppermost leaf (all editions of Forages as for eg. Barnes et al., 1995, p. 488). This is often the phenological stage at which herbage or forage (biomass) yield and nutritive value or content (ie. nutrient yield) are at optimum levels, and with respect to each other. Crested wheatgrass is clearly not up to the standard of wheat pasture, but it is good quality by most range (and tame pasture) standards. This example demonstrates why this extraordinary introduced range grass has been so widely planted and how it can complement native range and increase ranch productivity and profitability. Flying D Ranch, Gallatin County, Montana. June. SRM 614 (Crested Wheatgrass).

9.  Crested wheatgrass (mostly A. cristatum) pasture at peak standing crop- This photograph was taken on the same day as the preceding slide, but here on this shallower, more upland land (a bench) crested wheatgrass has advanced to maturity at an earlier date and in this drier than typical spring. (Note the dried, mature seed stalks.) On this less fertile and mesic site this introduced Eurasian grass will continue to survive (including, of course, to reproduce) as naturalized range and serve as what local ranchers call “dryland pasture”. (Compare to irrigated pasture on flood meadow three slides below.)

As mentioned for the previous slide, crested wheatgrass does not produce the highest quality grass forage imaginable. Yet, as evidenced by these registered Line One Herefords, diets from crested wheatgrass range are adequate for phenotypic expression of the genetic potential of high performing grazing animals. 

No FRES or Kuchler designation. SRM 614 (Crested Wheatgrass). Cooper Hereford Ranch, Gallatin County, Montana. June.

Alfalfa is one of man’s most important agronomic forage legumes, especially as a hay crop. Most cultivars of  common alfalfa (M. sativa) can be grown without irrigation only in sub-humid or wetter regions. There are a few alfalfa cultivars that have been selected for dryland agriculture in semi-arid regions. These are generally either M. sativa X  M. falcata  hybrids or M.  falcata selections. Ladak is one of the latter which holds some promise for seeded range. Gallatin, County, Montana. June.

Alfalfa, like the other major agronomic legumes, is a member of the Papilionoideae (= Faboideae) subfamily). Papilionaceous legumes have a corolla consisting of five petals: the largest is a single petal known as the standard or banner, two central petals fused as the keel, and two “side” (lateral) petals known as wings. The major agronomically important nodulated legumes— those entering into mutualistic symbiosis with nitrogen-fixing bacteria like Rhizobia species which form nodules on the host roots— are all in this subfamily, the Papilionoideae.

14. Desert crested wheatgrass (Agropyron desertorum) range- Desert crested wheatgrass instead of "regular" crested wheatgrass is grown in drier areas to which this group of Eurasian perennial bunchgrasses is adapted. This desert crested wheatgrass stand was planted in the High Lava Plains province (Franklin and Dyrness, 1973, p.6) of southcentral Oregon. The native Wyoming big sagebrush had re-established from the soil seedbank in this seeded single-species stand of naturalized bunchgrass to provide an outstanding example of the "right blend" of crested wheatgrass and big sagebrush. Physiognomy, structure, and species composition of this naturalized range was "danged near perfect".

Harney County, Oregon. June. Estival aspect; peak standing crop. No FRES number or Kuchler unit for this non-native (but naturalized) range cover type. SRM 614 (Crested Wheatgrass).

15. Naturalized range of desert crested wheatgrass- These two views of the same range (a different range from that presented in the preceding photogarph) that developed from a seeding of desert crested wheatgrass provided the student with an all-inclusive picture of "the way a crested wheatgrass range should look". The single-species stand of this Eurasian bunchgrass had been invaded by scattered plants of Wyoming big sagebrush resulting in the species composition, community structure, and, consequently, proper ecosystem function of one of the single most important and valuable cover types of rangeland in parts of the Northern Great Plains and Intermountain Region of North America.

At various points in this publication beginning students in Range Management have been advised to familarize themselves with the classification of grazing lands based on longevity or duration of pasture developed by Vallentine (2001, ps. 7-13). Vallentine's treatment was comprehensive and covered all grazing lands, both natural or native and introduced or agronomic pasture (a dichotomy that Vallentine [2001, ps.7, related 15-16] showed to be unreliable for certain uses). The Vallentine classification of grazing lands is, in the current author's experience, the most reliable, comprehensive, and useful of any other such treatment of grazing lands (= grazinglands as one word, if the reader prefers).

In the comprehensive, pasture longevity-based system of Vallentine (2001, ps. 7-13) crested wheatgrass grazing land falls under the categories of "long-term grazing land (synonym range), seeded range (introduced species)". Simply put, crested wheatgrass is introduced range. In fact, crested wheatgrass was probably the single most common example of why the term range was first officially defined by the American Society of Range Management (Huss, 1964) to include "... lands that are revegetated naturally or artificially to provide forage cover that is managed like native vegetation".

Technical note on syntax of the language: It is redundant and affected usage to use the words "native range" except in context of drawing the clear distinction between native range vs. introduced range where the latter is the shortened version of the Vallentine (2001, p. 8) term of "seeded range (introduced species)". If one is comparing or distinguishing between range made up of seded introduced species and range comprised of seeded native species or contrasting seeded introduced range to native (non-seeded) range, the use of native range is quite appropariate. Otherwise it is inappropriate and marks one as a novice, pedant or greenhorn. For example, to speak of sagebrush steppe or tallgrass prairie as "native range" without contrast or comparison to, say, crested wheatgrass range or weeping lovegrass (Eragroistic curvula) range is stilted, redundant, and unnecessary verbage. In context of the latter examples, the foolish use of "native range" is the equivalent of describing livestock as "domestic livestock" (without contrast to feral livetock). Use the language of our discipline and profession properly so as to foster respect for them, or do not use our "lingo" at all.

Crested wheatgrass (both A. desertorum and A. cristatum) have proven to be some of the useful non-domesticated plants introduced into North America. These two species have been shown time and again to be invaluable for soil and water conservation and wildlife habitat (feed and cover) as well as in provision of forage for livestock.

Harney County, Oregon. June. Estival aspect;

16. Crested whetgrass range in the Great Plains- Former cropland reseeeded to crested wheatgrass (Agropyron cristatum) in the Central Great Plains. This pasture of naturalized range had been recently grazed by cattle that had been moved to other pasture. Photograph taken under overcast sky such that grass could not be shown to best clarity.

Laramie County, Wyoming. Late June (early summer).

17. Naturalized amid the pines- Crested wheatgrass (A. cristatum) range in the ponderosa pine zone at western edge of Central Great Plains (foothills of the eastern side of the Rocky Mountains). Much of this crested wheatgrass appeared to have been from natural seeding of this species rather than by human planting. Crested wheatgrass had literally taken over much of the range in this area. Some of this had been abandoned fields tht had been reseeded. This resulted in such a seed source that abused ranges became naturally revegetated by crested wheatgrass. This phenomenon had likely reduced the species diversity, but at least the soil wass stabilizeed and the land producing forage for livestock and wildlife. It was certainly a testament to the adaptation of this species. toAbout the only native grasses of consequence were Junegrass (Koeleria cristata= K. pyramidata) and several bluegrass species. The small, whitish or "silvery" colored shrubs were plains silver sagebrush (Artemisia cana).

Platte County, Wyoming. Late June (early summer). Soft-dough stage.

18. Individual plants of desert crested wheatgrass (Agropyron desertorum)- These two robust and ungrazed specimens of desert crested wheatgrass provided examples their species and illustrated cespitose morphology of bunchgrasses in general.

Once individual plants have developed to this stage of maturity and have aquired this extreme degree of rank growth they are no longer palatable to grazing animals (unless animals are feeling severe "hunger pangs" if not suffering from "holler belly"). These two grass plants were chosen from ungrazed areas to give viewers "purple ribbon" textbook examples of this species. In a grazed pasture an individual such as either of these "grand champions" is known as a wolf plant: "1) an individual plant that is generally considered palatable, but is not grazed by livestock, 2) an isolated plant growing to extraordinary size, usually from lack of competition or utilization" (Bedell, 1989).

Harney County, Oregon. June. Peak standing crop; soft-seed phenological stage.

19. Spikes of desert crested wheatgrass- Inflorescences of grasses in the Tritaceae or, also, Hordeae (wheat or barley tribe) have inflorescences known as spikes which are distinguished by having spikelets sessile (without a pedicel or short "stalk") on the rachis (central axis of a grass inflorescence).

Harney County, Oregon. June. Soft-seed phenological stage.

20. Desert crested wheatgrass in Intermountain Region- Seeding of desert crested wheatgrass on an old field (abandoned cropland) in the Great Basin. Exact agricultural, especially agronomic, history of this land was largely lost to time except for local oral tradtion which was rapidly diminishing. A would-be grain grower undoubtedly broke out this land on which virgin vegetation was probably some form of sagebrush shrub steppe with Wyoming big sagebrush (Artemisia tridentata ssp. wyomingensis), wheatgrasses, including western wheatgrass (Agropyron smithii), galleta (Hilaria jamesii), needle-and-thread- (Stipa comata), and Indian ricegrass (Oryzopsis hymenoides). Following abandonment or reverting to Public Domain (probably failure to "prove up" on the homestead) the devasted land was reseeded to the Eurasian desert crested wheatgrass.

This reseeded range was used as an example of countless thousands of acres of land that went from virgin range to vanished dreams of the family farm (on land hardly more than desert) back to introduced range.

Juab County, Utah. Bureau of Land Management, Fillmore Field Office. June.

21. Mixed seeding of introduced range- A mixed seeding of desert crested wheatgrass and intermediate wheatgrass (Agropyron intermedium) on depleted range. Range vegetation on this Great Basin land had been extremely degraded by past improper mamagement that most likely was a combination of uses including attempted production of field crops such as small grains. By reseeding and judicious grazing management this naturalized range vegetation was what Vallentine (2001, p. 8) classified as Seeded Range (Introduced Species), one kind or category of Long-term Grazing Lands. With proper management this crested wheatgrass-intermediate wheatgrass introduced range will produce plentiful forage for livestock and wildlife indefinitely as well as protecting previously abused land.

Juab County, Utah. Bureau of Land Management, Fillmore Field Office. June.

22. Outstanding habitat on an old seeding- Two views of an extremely old seeding (perhaps past the half century mark) of desert crested wheatgrass on which Wyoming big sagebrush invaded. This is the stage of vegetation development on introduced range like crested wheatgrass that is ideal for multi-animal species use (ie. common grazing by several species of range animal). Re-establishment (invasion) of big sagebrush by secondary succession added the proper cover and density of a desirable native range shrub that can now provide browse for big games species like pronghorn and mule deer and cover for shade and nesting space by various bird species.

Big sagebrush at this relative proportion will have minimal influence on crested wheatgrass yield and cattle or sheep gains. It does provides valuable browse for wildlife that consume proportionately more biomass from woody plants than do grass-preferring cattle. The scattered plants of big sagebrush trap blowing snow which upon melting increases soil moisture that benefits grass as well as sagebrush. Big sagebrush retains most of its leaves during winter which serve as browse for various species including native ruminants and even sheep. Under conditions of heavy snow accumulation sagebrush is a source of emergency feed.

If and when big sagebrush cover becomes excessive, prescribed fire of the proper regime (season, intensity, etc.) or simple application of herbicide like 2,4-D can be used to reduce sagebrush to a more desirable density, cover, age of plants, etc. Care must be taken when using prescribed fire or too much sagebrush can be killed and cheatgrass increased to excessive quantities. Management is key and management means proper action.

Juab County, Utah. June.

23. Mixed seeding of intermediate wheatgrass and smooth brome (Bromus inermis)- Former cropland retired under Copnsrvation Reserve program and reseeded to a half-and-half mix of intermediate wheatgrass and smooth bromegrass. Year of average precipitation (though coming out of a four-year drought) so this pasture was not being grazed by livestock.

Intermediate wheatgrass in foreground and smooth bromegrass in background of both photographs.Details of pasture sward presented in second slide.

Dawes County, Nebraska. Late June (estival aspect); peak standing crop and early grain-ripe stage of phenology.

24. Intermediate wheatgrass (Agropyron intermedium)- This is another Eurasian species that was introduced and subsequently naturalized, at least locally, across much of the northern portion of the Western Range. At phenological stages below maturity intrmediate wheatgrass produces high yields of good to excellent quality forage. Intermediate wheatgrass has been seeded as a single species, much like crested wheatgrass, as well as a component species of range reseeding mixtures. Like crested wheatgrass, intermediate wheatgrass is also valuable for native ruminants. This demonstrated fact has sometimes been often overlooked by detractors who decry seeding of non-native species and establishment of monocultures (= single-species stands). In the author's observation intermediate wheatgrass has been nowhere as "successful" on the Western Range as have the crested wheatgrass species.

Lincoln County, Colorado. June.

26. Plants of intermediate wheatgrass- Four plants (genetic individuals) of the cespitose intermediate wheatgrass were growing on this reseeded landing of a ponderosa pine (Pinus ponderosa) forest. Following selective logging of the uneven-age forest intermediate wheatgrass volunteered on local parts of the understorey. The landing and logging trails were apparently reseeded to this introduced species.

Ochoco National Forest, Wheeler County, Oregon. June.

28. Intermediate wheatgrass in the Palouse Prairie Region- A former native bunchgrass steppe in heart of the Palouse Hills was converted into this dry-land pasture of intermediate wheatgrass with substantially greater grazing capacity. Another melanchology instance of what Charlie Russell dubbed "Trails Plowed Under" as the native grassland gave way to more agriculturally productive --and profitable-- tame (ie. agronomic) pasture.At least it was still grazing land. Furthermore, in this instance the introduced forage grass provided complementary pasture for adjoining cropland also used for grazing (eg. small grains pasture, crop aftermath) or harvested forage (primarily hay) along with some scattered remnants of Palouse Prairie range.

This was also an example of wise and efficient use of water. This tame pasture was being managed strictly as a rain-fed forage crop even though this field was bottomland with fertile soil such that domesticated, high-yielding crops would respond faovrably (physiologically if not profitably) to irrigation. In such a grassland agriculture management system limited water and expensive fossil fuels could be used to greater advantage on other fields for other crop species. Utilization of fields like this for higher-producing agronomic forages also enabled puirebred beef cattle to express their genetic potential which could then be evaluated. Such evaluation for genetic potential (eg. progeny testing for economically important traits) would not be possible (at least not to such extent) on natural pastures like the very limited remaining Palouse Prairie range that in such a fertile area was limited to steeper or rougher land which was marginal for field crops like small crops and oil seeds. An example of wise use management of financial and cultural resources as well as natural resources.

Whitman County, Washington. June, early hard dough stage of grain.

31. Pasture of Jose tall wheatgrass (Agropyron elongatum c. Jose)- A seeding of Jose tall wheatgrass grown as tame pasture and used as complementary pasture to a range-based cattle operation. This single-species stand was growing on land that was naturally subirrigated (the soil profile is adjacent or close to a very shallow surface aquifer; often accompanied by capillary flow from streams flowing in close proximity, as in the case shown here where eastern cottonwood trees indicte presence of such stream). This photograph was taken in 2002 during the most extreme drought in Colorado history. No precipitation fell during this growing season. Instead all soil moisture was from that stored in soil and attributable to subirrigation.

Tall wheatgrass is another introduced (from Eurasia) domestic grass species. It does particularily well in the Great Plains (from New Mexico to Canada) where it is often grown under irrigation and produces high herbage yields. This is probably the largest and latest-maturing of the introduced wheatgrasses. It tends to become rank and coarse at which stage it becomes unpalatable, but as a function of it's late-maturity tall wheatgrass has a longer green-feed period. For these reasons tall wheatgrass is more commonly planted and managed as a single-species stands (monocultures) rather than in mixtured seedings (polycultures). Consistent with it's height and general large size, tall wheatgrass has vast, deeply penetrating root systems tht contribute to its' general drought tolerance (see slide again). It also does well or alkaline and/or saline sites, a feature which makes it adapted to flood plains in semiarid climates.

Beef producers have to "stay on top of" tall wheatgrass. This species is less tolerant of oversue than most wheatgrasses , but if it is not kept grazed to a height and state of maturity that is palatable to cattle tall wheatgrass "gets ahead of" the stock and their performance will suffer as forage intakes drop precipituously.

One of the amusing (at least to this author) academic aspects of tall wheatgrass is the "muscial chairs" gyrations performed by agrostologists in regards the scientific name of tall wheatgrass. In each of the last editions of the Iowa State classic text, Forages, the simple common name of tall wheatgrass was shown as Agypyron elongatum (1973), Elytrigia pontica (1985), and Thinopyrum ponticum (1995). Maybe such name changes are beneficial, sort of like fashion changes in neck ties and hemlines. They keep professors from becoming "stale". Is this an example of "continuing education"? Rangemen have to re-learn scientific names so they will know the correct answer in case there becomes a game, The Range of the Trivial. Then knowing the latest species name for tall wheatgrass would be about as important as knowing the name of Tonto's or Dale Evans' horse. Scout and Buttermilk (I can handle those).

34. Quackgrass, quichgrass, or couchgrass (Agropyron repens= Elytrigia repens)- Quackgrass is another of the introduced Eurasian wheatgrasses. According to various authors in Iowa State's Forages text the historical details regarding introduction of quackgrass remain obscure. It was introduced into North America prior to the Revolutionary War. Quackgrass is now one of the most widely distributed species of the wheatgrass group being found on every continent.

The situation regarding quackgrass is similar to that of other introduced grasses which in numerous locations became more important as weeds than beneficial crops. Quackgrass is in this respect the "Johnsongrass of the north". Other introduced grasses that have these ruderal (naturalized) and noxious (plant pest; weed) aspects include (though generally to a lesser degree of noxiousness) Kentucky bluegrass, bahiagrass, bermudagrass, tall fescue, and King Ranch bluestem. Quackgrass has naturalized in North America from parts of Oklahoma and Arkansas far into the Canadian provinces. Quackgrass is especially noxious in the northern United States and Canada where it is particularily aggressive and persistent. In fact, quackgrass is probably the worst (or next to the worst) weed in much of this vast region.

While understanding the weediness of quakegrass and farmers antipathy toward it, many forage specialists have consistently pointed to the highly desirable features of quakegrass as a forage crop, especially under minimal input management. For instance, authors in Forages (beginning in 1951) consistently acknowledged the benefits of quackgrass while pointing out that control of it as a weed was not as serious a problem as it once was. Authors in recent editions of Forages pointed out that quackgrass, though a weed, was so well-adapted to habitats in the far north that it was now one of the major grazed and harvested forages in Alaska. Pieper (1939, ps. 249-250) remarked that much of the hay in New England that was alleged to be timothy was "in reality quack grass". He concluded: "While quack grass is one of the worst weeds with which a farmer has to contend, this fact should not obsucre the merits which the plant really possesses".

One of the most recent and promishing practices in wheatgrass breeding was development of interspecific hybrids. Quackgrass was one of the major parental species in this effort, in particular for hybridization with bluebunch wheatgrass and crested wheatgrass. One such cross was the quackgrass x tetraploid bluebunch wheatgrass hybrid. In 1981 two of these synthetic hybrids were released jointly by the Agricultural Research Service and the Utah Agricultural Experiment Station (Asay and Dewey, 1981). In 1989, the cultivar NewHy wheatgrass (Agropyron spicatum x repens= Pseudoroegneria spicata x Elytrigia repens) that was developed from this breeding program was released. Like the hybrid equine, the mule, this Agropyron hybrid combined desirable traits of both species in a single new organism with the increased performance due to heterosis ("hybrid vigor"). With the drought tolerance and high palatability of the native bluebunch wheatgrass combined with the grazing tolerance of quackgrass-- and exhibiting heterosis-- NewHy excels in all three traits. NewHy has proven to especially well-adapted to harsh habitats such as saline environments. It holds potential for use in reclamation. Such developments have proven over and over the wisdom of the tongue-in-cheek remark of Ralph Waldo Emerson, "A weed is a plant whose virtues have not yet been discovered".

The more common importance of quackgrass is as a naturalized range plant in the northern portions of North America. For example, in northeastern North America much of the original forest land is now used primarily as naturalized range. These were cut-over forests that are undergoing secondary succession, but on which introduced pasture species like quackgrass, timothy, orchardgrass, redtop and bentgrasses (Agrostis species) naturalized and, for the forseeable future, provide a grazable resource. In it's National Range and Pasture Handbook the Natural Resources Conservation Service (1997, ps. 2-3, 5.1-21) described such grazing lands as native and naturalized pasture. It was not completely clear how some of these grazing lands differed from permanent forest range, but cut-over forestland not being managed (at least not intensely) by forest management practices and used primarily for grazable/ browsable understorey appeared to this author to pretty much catch the spirit of this category of grazing land.

"Naturalized pasture is land that was forest land in historic climax, but is being managed primarily for the production of forage rather than the production of wood products. It is managed for forage production with only the application of grazing management principles. The absence of the application of fertilizer, lime, and other agronomic type practices distinguish this land use from pasture." (Natural Resources Conservation Service, 1997, p. 5.1-21).

In other words, this form or category of pasture or grazingland is essentially naturalized forest range (ie. the range portion-- the grazable understorey-- is dominated by introduced and now naturalized species). On such naturalized range (naturalized grazingland) in more northern portions, especially in the eastern half of North America, quackgrass is a dominant species. Quackgrass was included here from that perspective as well as it's recent contributions to wheatgrass breeding.

The quackgrass shown in these two slides was in the soft-dough stage. It formed exclusive stands in bar ditchs and fencerows in the fashion typical over much of the geographic regions to which it has naturalized. This quackgrass was too mature to make good-quality hay, but when quackgrass is cut in the boot stage it will produce high yields of high-quality grass hay.

Agriculture Canada Research Centre, Lethbridge, Alberta. July.

36. Rootcrown (proaxis) and rhizomes of quackgrass- The rootcrown as used in Agrsotology is that general part of the grass plant where the shoot and root merge. It is not necessarily a union in the sense that the ligule-collar is the union of leaf blade and sheath, but it is a distinctive morphological part and the one from which much of grass growth is initiated. Tillers (known also, and in less technical terms, as suckers or stools) emanate from the proaxis. This is more true for cespitose species (bunchgrasses) than for sod-forming grasses, but rootcrowns are central growth regions in all grasses and grasslike plants. Some grasses reproduce asexually from (by) both tillers and rhizomes, by stolons, or by all threee main forms of secondary shoots). Quackgrass has rhizomes as well as a conspicuous rootcrown. Both of these organs were shown in this slide.

Asexual reproduction by rhizomes traditionally was the explanation given for the aggressiveness and persistence of quackgrass, and thus the main factor in it's noxious (extremely weedy) characteristic. A rangeman or grazier who cherishes quackgrass for it's high yield of nutritious forage can turn that fact around and point out that rhizomes are the main reason why quackgrass is so valuable as a naturalized forage species. Numerous authors such as Pieper (1939, p. 250) pointed out that quackgrass rhizomes have been harvested and fed as if hay which allegedly rivaled in nutritive value the better legume hays. Rhizomatous grasses like quackgrass and Johnsongrass usually benefit (improved stand maintenance as well as increased herbage yields) from some form of soil tillage. In other words, cultivation is not a control method for rhizomatous weeds. Excessively heavy grazing is a more effective method of control. The lesson for grassmen is to be certain to practice sound grazing mangement of rhizomatous grasses like quackgrass or they will be grazed out.

Provencial Prison grounds, Lethbridge, Alberta. July.

This smooth bromegrass seeding in Harvey Valley, Lassen National Forest, California was used to restore depleted range more efficiently and with less risk of crop failure than if attempting to reestablish native grass species such as bluebunch wheatgrass or Idaho fescue. In such situations smooth brome provides vital vegetative cover faster and more reliably than native species and is superior in reduction of accelerated erosion and protection of critical watersheds. It not only provides nutritious forage for both livestock and wildlife but it can actually increase forage (and total nutrient) yield and, ultimately, animal turn-off. Note that as with crested wheatgrass, native shrubs (eg. the sagebrush growing here) come in on range reseeded to introduced grasses and help form a range plant community in contrast to the monocultures or single-species stands of agronomic pastures. Again, as this is not native (potential natural or climax) vegetation it has neither FRES nor Kuchler unit recognition. No SRM designation either.

39. Smooth brome pasture in the Great Plains- Pasture of smooth brome at peak standing crop being grazed by Angus cattle.First of these two photographs taken at full-sun; second photograph taken a few minutes later under an overcast sky. These paired photographs illustrated the effects of shading on color contrast. Panicles of smooth brome showed more distinctly (with more clarity) under overcast atmospheric conditions, but the yellowish-tinged green of smooth brome at an early stage of grain ripening showed more clearly in absence of shading. Also cumulus with their white/gray coloration showed up better in full sun (absence of cloud cover between sun and photographed subject). This example demonstrated that appropriateness of light intensity or, for that matter, quality or kind of light ("light conditions") depends on what is being featured in the photograph and, in context of instructional use of photographs, the lesson trying to be taught.

The herbage in this pasture was beyond best stage of plant maturity for animal performance. As a general rule of thumb, the lower the stage of plant maaturity the higher the nutritive value of forage. Forage quality must, however, be balanced, with herbage yield when optimum quantities of specific nutrients and feed properties (eg. voluntary intake of dry matter and usable energy, digestability, palatability) are being determined. These in turn vary with kind and class of pasture animal. Furthermore, it is neigh on to impossible to keep forage plants in immature phenological stages and maintain the forage crop, the pasture. Compromise between grazing animal and grazed crop is an unavoidable part of pasture management. Such tradeoffs are more of greater management concern when there are options as, for instance, in tame or agronomic pasture (or, eaven, agronomic species managed as range) such as that shown here.

Sheridan County, Nebraska. Late June (estival aspect); peak standing crop and early grain-ripe stage of phenology.

40. Sward of smooth brome at maximum biomass- Stand of Bromus inermis at peak standing crop with caryopses in early ripening stage of phenology. This stand was in the Conservation Reserve Program in a year of average moisture conditons so that it was not being grazed by livestock.

Dawnes County, Nebraska. Late June (estival aspect); peak standing crop and early grain-ripe stage of phenology.

Location note: an example of a mixed pasture seeding of smooth brome and intermediate wheatgrass (Agropyron intermedium) was included above under intermediate wheatgrass (Hordeae or Tritaceae).

41. Grazing land type conversion: from small grains to smooth brome- On this steep, rolling, highly eroidable land in the Palouse Hills a native bluebunch wheatgrass (Agropyron spicatum)-Idaho fescue (Festuca idahoensis) bunchgrass prairie was converted into a crop field that was used for production of various crops like wheat (Triticum aestivum) and lentils (Lentilla lens) before it was restored to the soil-protecting cover of perennial grass produced by smooth brome. This monoculture stand had remarkably little cover of cheatgrass or other introduced Eurasian forage grasses that have been widely planted in this area (some of which were covered below). It provided a good example of smooth bromegrass forage at its best. This stand was being grazed when the photograph was taken. It had been used alterntively for hay or pasture.

Latah County, Idaho, June (early summer), soft dough phenological stage.

44. Rescue grass or rescuegrass (Bromus unioloides= B. catharaticus)- Two views of rescuegrass, an introduced (from its native Argentina), short-lived perennial (that frequently responds to cultivation, grazing, or winterkill as an annual) bromegrass that has naturalized widely over the southern United States and parts of Mexico. Hitchcock and Chase (1950, p. 34), Gould (1975, p. 84), Hatch and Pluhar (1993, p. 161), and Alderson and Sharp (1995, p. 26) described rescuegrass as an annual or biennial, but other authors (including those for all editions except the sixth of Forages) interpreted this species as a short-lived perennial. Barkworth et al. (2007, ps. 199-201) distinguished between Bromus catharticus var catharticus, an annual or biennial, and B. catharticus var. elatus, a perennial. (Incidentially, readers will note that the experts still cannot agree on the proper specific epithet for this species: they have gone back and forth between unioloides and catharaticus for decades.)

Rescuegrass was one of the first Bromus species that was intentionally introduced into North America. It is also one of the few introduced bromes that is not Eurasian or Old World in origin. Based on official or documentable records rescuegrass was introduced into North America in 1853 and widely advertised by B.V. Iverson of Columbus, Georgia who is also credited with giving this species the American English name of rescuegrass (Piper, 1939, p. 279). However, according to the first edition of Forages (Bennett in Hughes et al. 1951, p. 374), and apparently later quoted from this source by Phillips Petroleum Company (1963, p. 22) and Nicholson (2006, p. 22), this species was first identified in the United States in 1806. Regardless, rescuegrass has 1) been on North American soil for over two and a half centuries and 2) naturalized over a large geographic area much like the various species of annual Eurasian bromes. In fact, B. unioloides often grows from autumn through early to mid spring in association with naturalized B. japonicus and B. tectorum throughout much of the southeastern and southcentral regions of North America. (This was described in more detail three paragraphs below.)

The agronomic "state of affairs" of rescuegrass has been similar to that of many introduced grasses and legumes: it did not become the "wonder forage" that it was purported and hoped to be but instead naturalized so as 1) to not require management by farming practices (intensive management) and 2) persist as either a weed or less productive crop plant that is best used as an opportunistic forage. This is, of course, the common situation with such introduced species as Johnsongrass (Sorghum halepense), quackgrass (Agropyron repens) , Vaseygrass (Paspalum urvillei), bahiagrass (Paspalum notatum), white and yellow sweet clovers (Melilotus alba and M. officinalis), and sericea lespedeza (L. sericea= L. cuneata).

Nonetheless, rescuegrass does commonly live up to its name by furnishing advantageous (often fortuitous) feed, especially on overgrazed southern ranges, small subsistence and "hobby farm" pastures, and "go-back land" that might otherwise have little but warm-season invader species. Also, rescuegrass provides a relatively dependable source of forage during the cool-season (including the dead of winter in the South) when both livestock and wildlife would otherwise be forced to subsist on either harvested roughage, mature, weathered, standing, warm-season forage, or browse. It should be underscored, however, that most such producers (and their unfortunate livestock) so "rescued" were at best marginal or, more likely, poor managers and poor husbandmen of crop plants and animals.

Rescuegrass does sometimes occur as a more-or-less single species stand (= monoculture), especially on seeded pastures, but more commonly this self-seeding annual, biennial, or short-lived, perennial is one--albeit it often the dominant one--species among several associated species. Plants growing in association with rescuegrass include other naturalized, as well as adventive and native species (usually in that relative order). Plants growing on rescuegrass-dominated ranges and pastures include exotic annual or perennial grasses such as Eurasian annual bromes (noted above), wild oats (Avena fatua), ryegrass (Lolium perenne, L. multiflorum), and goatgrass (Aegilops cylindrica); native annual grasses like little barley (Hordeum pusillum); plus exotic forbs, especially annuals including crucifers like shepherd's purse (Capsella bursa-pastoris) and Eurasian mustards (Brassica spp.); composites such as prickly sow-thistle (Sonchus asper), prickly lettuce (Lactuca serriola), and common groundsel (Senecio vulgaris); and the geranium family member, redstem filaree (Erodium cicutarium). Grazinglands composed of these kinds of introduced species (many or most of them weeds in both agronomic and ecological usages) in association with rescuegrass constitute medium-term transitory pasture/range at best or perhaps a form of short-term grazingland (Vallentine, 2001, ps. 11-13). Rescuegrass also grows on local habitats (microsites) characterized by disturbance within native grasslands and savannahs as well as revegetating sacrifice areas and old barnyards and in patches or more open spaces on permanent pastures of warm-season grasses as, for instance, in the cool growing season on dormant bermudagrass (Cynodon dactylon). In these latter instances rescuegrass functions like an overseeded small grains crop (except that the naturalized rescuegrass is self-perpetuating).

There have been numerous cultivars and accessions of rescuegrass developed and released over the years, including some from New Zealand (Alserson and Sharp, 1995, ps. 26-28).

Overall, rescuegrass is generally more welcome than not. This species definitely does not produce toxicity problems like those of Johnsongrass and the sweet clovers or yield forage that at maturity is as low in nutritive value as that of introduced warm-season forage species or even some of the more productive, introduced cool-season grasses like tall fescue (Festuca arundinaceae).

Erath County, Texas. April, peak standing crop.

45. Shoot apices of rescuegrass- Top of shoot (first photograph) and panicle (second photograph) of rescuegrass. The inflorescence of the Bromus species is a conspicuous and usually relatively large panicle.

Erath County, Texas. April, hard-dough stage in grain.

46. Spikelets of rescuegrass- Rescuegrass is usually a prolific grain-producer. This is the necessary condition for most short-lived perennials. It also one of the likely reasons why rescuegrass can persist on grazinglands as an annual. It seemed likely that the common high yields of grain with concentrations of energy and nutrients contributed to nutritional value of animal diets on rescuegrass pasture.

Erath County, Texas. April, hard-dough stage in grain.

Tall fescue is only fair in forage nutritive value as a result of the endophytic toxins which cause detrimental effects on grazing animals (ps. 110, 348-349 in Barnes et al., 1995), but the growers who rely on it obviously accept the tradeoffs and tall fescue continues to be planted and maintained as the major cool-season permanent pasture grass in the USA.

 From the view of stockraising on ranching operations tall fescue is tame pasture that is complementary or supplementary to the range base. Naturalized or introduced range such as crested wheatgrass is also typically complementary pasture (enhances the usefulness of native pastures which provide most of the ranch’s forage needs) but this category of introduced forage is managed more extensively, more ecologically than agronomically, (ie. it is range and managed as range). By contrast tall fescue is managed more agronomically (ie. it is not range). As discussed in the introduction, however, the distinction between tame and native pasture is not always clear and can be downright arbitrary. Tall fescue under minimal management is apparently a case in point. Shiflet (1994) designated and described tall fescue as a rangeland cover type (SRM 804). It was included here in that context. Tall fescue appears to have naturalized in some areas. It is sometimes an invader of abused tallgrass prairie especially along the boundaries between fescue pastures and range or even prairie hay meadows where the cool-season bunchgrass can be particularly noxious. No FRES or Kuchler unit because it is not natural vegetation. Adair County, Oklahoma. May.

50. Naturalized in the central southland- Example of tall fescue naturalization and successful invasion in the tallgrass parairie and oak-hickory forest types in the Ozark Plateau. If this land could talk it would tell of how a former tallgrass prairie within a larger area of surrounding oak-hickory forest was overgrazed about 90 to 100 years ago in subsistence-level diary farming that resulted in invasion (expansion) of hardwood and shrub-dominated old field vegetation which was subsequently invaded by naturalized tall fescue and converted into a tall fescue grassland with broomsedge bluestem (Andropogon virginicus) as the major associate. Along the perimeter of this naturalized grassland common bermudagrass was common along with weedy and ruderal grasses such as hairy crabgrass (Digitaria sanguinalis) and goosegrass (Eleuisine indica).

This overgrazed pasture (in effect an old field or go-back land) became dominated by--in many portions almost exclusively made up of--Kentucky 31 tall fescue that volunteered from seed from an adjacent hay field and by feeding of tall fescue hay from that neighboring land. This was an example of a typical and widespread development with the result that tall fescue, especially endophyte- infected cultivars like Kentucky 31, became a major weed throughout much of the southcentral and southeastern regions of the United States such as shown here in the Ozark Plateau. When tall fescue was first being seeded throughout the Ozarks farmers and stockmen argued over whether this species would self-seed so as to "fill in the gaps" between widely spaced plants of this bunchgrass on newly planted fields. Initially, establishment of new plants from older ones established by drill seeding was not apparent and there was general agreement among producers that second, third, etc. generation plants of tall fescue was not likely. If you had a "skimpy" stand it would stay that way and reseeding was advised.

Eventually it became obvious that over course of, say, five to ten or so years tall fescue had not only "filled in the gaps" where this was desired, but that it was also spreading by seed slowly but inexorably into adjoining fields, tame pastures, and even into tallgrass prairie (especially those that had been routinely hayed late in the summer or overgrazed).Once established--by planting or natural seeding--tall fescue is--for purposes of comparison--even more persistent (considerably more persistent) than naturalized Johnsongrass. This is due to the extreme tolerance to severe overuse and general grazing mismanagement that is the "order of the day" throughout much of the vast region over which tall fescue became a defacto "king grass". This is because endophyte-infected tall fescue, unlike grazing-susceptible Johnsongrass, cannot be grazed out readily (if at all).

Tall fescue is now an established plant across landscapes in the South, for better or worse. Under conditions like that shown here the natualization of tall fescue has clearly been for the better. Tall fescue has prevented soil erosion in the southern humid zone where heavy rains possible at any time of the year can have devastating impacts on abused land. Perhaps most consistently tall fescue has provided forage--along with soil protection--where there would otherwise be little except that provided by annual grasses and unpalatable perennials like broomsedge bluestem. Across the region over which tall fescue has naturalized is probably one of the most effective means of preventing or reducing infestations of annual and biennial weedy forbs that were previously much more abundant on the "grubbed-out" land such as much of that in the Ozark Plateau . On the other hand, this latter feature was undoubtedly a big factor in reduction in populations of northern bobwhite (Colinus virginianus) in areas once famous for this extradinary upland game species.

Along with tolerance to abuse from grazing and trampling, tall fescue is drought-tolerant due largely to drough avoidance that is inherent with a cool-season species in the South. Tall fescue goes dormant by mid-winter and late summer so tht it naturally avoids the the two major periods of dry edaphic conditions.

Ottawa County, Oklahoma. Late December: early winter and still this much green feed.

51. Stand of orchardgrass (Dactylis golmerata)- Orchardgrass is another Eurasian perennial grass that was introduced into and widely planted throughout North America. Single species stands of orchardgrass such as this one are common in the central and upper Midwest. In areas of northern Missouri, Iowa, IllinoisMichigan, etc. orchardgrass (or the British, cocksfoot) is a Eurasian cool-season, perennial grass species for both pasture and hay. Orchardgrass is a important species with wide application in grassland agriculture. In his landmark book, Theory and Dynamics of Grassland Ageiculture (Harlan, 1956) described grassland agriculture: "A system of agriculture in which a major emphasis is placed on grasses, legumes, and other fodder or soil-building crops. The system may be extensive as in many natural grassland or desert shrub areas, or it may be extremely intensive. The sod crops may be permanent or simply part of a rotation. The principal feature of grassland agriculture is the dependence of the system on grass crops for soil building and animal nutrition. Grassland agriculture, is therefore, a dynamic system in which the soil, the plant , and the animal are intimately and inseparably interlocked." In Forages-The Science of Grassland Agriculture the editors defined grassland agriculture as: "Farming system that emphasizes the importance of grasses and legumes in livestock and land management" (Hughes and numerous other editors down through the years, 1962, 1973, 1985, 1995, 2003). Vallentine (1990) defined grassland agriculture as "A land management system emphasizing cultuvbated forage crops, pasture, and range for livestock [and wildlife] production and soil stability." (This definition was more inclusive than his definition in the second edition [Vallentine, 2001]).

More details of orchardgrass as a pasture and naturalized range species were preented in the immediately following caption.

Ottawa County, Oklahoma. May.

53. Feet on top- The panicle of orchardgrass, or as the Limies know it, cocksfoot. It would seem that one needs a vivid imagination to see a chicken foot (where's the cock's spur?) in the inflorescences of this species, but several panicles were provided that might facilitate such imaging. For less poetic and more practical folks this was just a view of flower clusters of an agronomic forage grass that naturalized to occupy an apparently empty niche, provide feed for livestock and wildlife, and add biodiversity to North American range.

Ottawa County, Oklahoma. May; peak anthesis.

55. Subirrigated meadow seeded to Kentucky bluegrass- Poa pratensis, another member of the Festuceae or Poeae tribe of Gramineae, is one of the more valuable introduced grasses in the tame pastures of eastern North America. It makes outstanding pasture for horses and was once used to raise grass-fat cattle for eastern urban markets. Today higher-yielding introduced species like tall fescue and orchardgrass are grown instead of Kentucky bluegrass over much of its former area. On moister sites in semi-arid regions Kentucky bluegrass is still one of the most persistent introduced pasture grasses in North America. This is due primarily to its grazing tolerance and fairly aggresive spread by abundant rhizomes. In fact, with exception of tall fescue Kentucky bluegrass is the (at least one of the) most grazing-tolerant of all major cool-season pasture grasses in North America.

The bottomland mountain pasture shown here included some sedge and native grass species (eg. blue grama and western wheatgrasss), but it is overwhelmingly populated by Kentucky bluegrass. Abundant subsurface moisture is present over much of the growing season and the introduced bluegrass has persisted. Smooth bromegrass is another introduced grass that does well on moist, fertile soils, but it is not as tolerant of heavy grazing as Kentucky bluegrass. Kentucky bluegrass is a weed on overgrazed tallgrass and even mixed prairie ranges, but on sites such as the one seen here and with proper grazing Kentucky bluegrass makes a fine complement to other kinds of pasture. Teton County, Wyoming. July. No FRES, Kuchler, or SRM units.

56. Sandhills subirrigated pasture- Pasture of a mixture of Kentucky bluegrass and creeping foxtail (Alopecurus arundinaceus, Garrison, cultuvar) on a subirrigated meadow in Nebraska Sandhills. Some other incidental plant species (including annual Mediterranean Bromus spp.) were present. Species composition of this pasture was not determined but an approximatae "half-and-half" would not be far off.

Details of creeping foxtail presented below under Aveneae.

Cherry County, Nebraska. late June (early summer). Predominately hard-grain for foxtail and seed-shatter stage for bluegrass.

57. Kentucky bluegrass- Exact origin of Poa pratensis has long been a matter of conjecture and confusion. The species is clearly a Eurasian species, but there is some evidence that it was also indigenous to northernmost North America. It most ceretainly was not native to southeastern North America, folklore and poetical names like Bluegrass State (hence Bill Monroe and his Bluegrass Boys) notwithstanding. Rather, and like so many other introduced forage species, Kentucky bluegrass "hitched a ride" with pioneering western Europeans and became one more plant species frequently called "the white man's footprint". Long before the "dark and bloody ground" of Kentucky got around to the settlement state of civilization hunting shirt Kentuckians Poa pratensis had naturalized over much of the land that later became a commonwealth and star in the Union. This naturalization phenomenon was responsible for the widely and variously used designations of Bluegrass this or that in the Bluegrass State, including the Bluegrass section or Bluegrass region of the interior low plateau physiographic province (Fenneman, 1938, ps. 419, 427-431, 441).

(Incidentally, the lack of native status of Kentucky bluegrass is most appropriate for the wonderful genre of Country-Western music that bears its name: Bluegrass Music is not indigenous among Kentucky and Tennessee crooners either. It too is an adaptation [and a skillfully market one] of an import from the "old country" of the British Isles, especially Scots Irish, with colorful additions from Africa via slaves and the slave trade (eg. the banjo), Germany, and God alone knows where else.)

For a century or more Kentucky bluegrass has been one of the most widely distributed grasses in North America. Authors in the Iowa State text, Forages (Barnes et al. 1995, p. 358) indicated that this introduced perennial occurred in every state of the Union. It is likely found in all of the Canadian provinces as well. Wedin and Huff in Moser et al. (1996, ps. 668-669) concluded that Kentucky bluegrass occurred in most, if not all, regions of the Americas, Eurasia, and North Africa as well as being progagated in Australisia. Given this almost universal distribution Poa pratensis shows tremedous ecotypic variation in morphology.

The ungrazed individual see here grew on a bluff on a north slope deep in an Ozark oak-hickory forest where it made itself "to home" every bit as much as the native plants in the forest community. Ottawa County, Oklahoma. April.

59. A rank, dense stand- This stand of rank-growing Kentucky bluegrass was in the Palouse Hills where, like so many areas of North America, it has naturalized to become as much a part of the range plant community as the native bluegrasses, fescues, bromes, and wheatgrasses. In fact, under human-induced disturbances such as improper grazing (eg. improper degree and season of use), heavy traffic, or disruption of land surface Kentucky bluegrass is able to outcompete, survive longer, and, in general, thrive when and where natives "give up the grass ghosts". This phenomenon is commonplace from the Flint-Osage Hills Region through the Rocky Mountains across to the Pacific Slope and Coast Range, and on grassland, savanna,and forest.

The coarse-stemmed plants in this and the following slide indicated the size and arrangement of tillers in an ungrazed plant in contrast to the closely cropped plant presented in the preceding slide. Obviously, proper grazing management should whenever possible strive to maintain Kentucky bluegrass in the leafier and less stemmy state that has more palatable and nutritions forage than that of the coarse or rank state like the plant shown in this photograph. Conversely, if the objective is seed production (either as a commodity for the grass seed market or for reproduction to restore grass on a degraded range or pasture) the plant shown here was representative of the form or degree of plant development that is very effective for that management goal.

Latah County, Idaho. June (early summer), anthesis.

60. Shoots, vertical and horizontal- The strongly rhizomatous and abundant tillering features of Kentucky bluegrass was evident in this photograph of what was probably a single plant of this species. In addition to these vegetative means of reproduction Kentucky bluegrass is a strong seed-producer, a characteristic also evident in this plant. The combination of both effective asexual and sexual reproduction along with tolerance of close grazing makes this species a fierce competitor with native cool-season grasses, especially under conditions of range abuse like overgrazing.

This photograph also demonstrated that a single plant of this species is capable of producing considerable herbage available to grazing animals. It was explained in the preceding caption that bluegrass plants like these which have relatively high proportions of culm compared to leaf material do not provide high-quality forage like those plants with more leaves and less culm material. There is a tradeoff between herbage yield and forage quality. The objective from the perspective of range animal nutrition is to optimize yield of nutrients (not gross herbage production or maximum phytomass).

Latah County, Idaho. June (early summer), anthesis.

61. Panicles of Kentucky keep on shining- Bill Monroe and his Bluegrass Boys should be favorably impressed with the simple beauty and prolific performance of these sexual shoots of Kentucky bluegrass at peak anthesis. These panicles were on the plants introduced in the last two slides. They were more elongated with more whorls of panicle branches than is the typical case. Panicles of Kentucky bluegrass were described by Hitchcock and Chase (1950, p. 115) as being "pyramidal or oblong-pyramidal". Most of the ones seen here were of the latter shape.

Latah County, Idaho. June (early summer), anthesis.

63. Details of Kentucky bluegrass panicle- This typical pyramid-shaped panicle of Kentucky bluegrass featured spikelets at anthesis. Hitchcock and Chase (1950, p. 115) described the lower whorl of panicle branches as being composed of five with a central long one, two short, and two intermediate length branches. There are usually three to five florets per spikelet (and a lot of spikelets per panicle branch) in this species. Poa pratensis produces abundant seed crops. It is a "survivor".

Latah County, Idaho. June (early summer), anthesis.

In California and Oregon ryegrass and tall fescue are grown as agronomic pasture species and here, too, these species “escaped cultivation” and naturalized to the extent that they are now members of the once-exclusively native range plant community on a “permanent” basis in an ecological as well as an agronomic sense. So far this is less the case in the Pacific Northwest than in the Southeastern Region.

Ryegrass pasture was included with range cover types following tall fescue because it is the species most similar to it in adaptation and extent of naturalization in what the SRM defined as the Southeastern Region (Shiflet, 1994, p. 111). No SRM designation for this one and no FRES or Kuchler units because ryegrass is not native.

 Lamar County, Texas. May. (In Texas, southern Oklahoma, and Arkansas ryegrass is perennial, or weakly perennial, so L. perenne seems the most precise species designation, but as seen in this and the next three slides Texas-grown ryegrass also produces seed-laden spikes perhaps qualifying it for L. multiflorum. Nomenclature controversy continues.)      

 The main cultural controversy for ryegrass was in California where for years it was standard practice to reseed burnt chaparral to ryegrass thereby often effecting a type conversion from shrubland to grassland, for some years anyway. This was controversial because on most of these sites California chaparral was climax or natural vegetation and the deeper-rooted shrubs were superior to grasses for watershed protection and as safeguards against massive mudslides that were apt to occur in the following rainy seasons. (Ironically, prevention of mudslides was presumedly the reason for seeding ryegrass to begin with.) The seeded grass would suppress development of seedlings and, to some extent, resprouts of chaparral species. Type conversions of brushfields or brushlands, as chaparral was often called, was an accepted practice for suitable sites, but wholesale conversion to ryegrass (or any other herbaceous species) was justly criticized until the practice was at least partly curtailed. Sampson and Jespersen (1963, ps. 21-32) discussed conversion of chaparral to grassland.

65. Perennial ryegrass (L. perenne)- Bottomland pasture of perennial ryegrass at peak standing crop (hard seed stage just prior to seed-shatter). Ryegrass at this phenological stage is long past it’s nutritional prime, but this photograph shows the high yield of herbage biomass possible from this introduced agronomic grass when after seeding and establishment it is managed solely through grazing management and as if it was native. From the Southeast, including east Texas, to the Cross Timbers and better sites of the Edwards Plateau ryegrass is outstanding complementary pasture to both natural and agronomic pastures, and it is permanent pasture or medium-term grazing lands. Ryegrass pasture like this can increase ranch cash flow and profitability, allow greater flexibility in overall ranch management, and reduce grazing on or allow winter deferment of “go-back land” range supporting cool-season natives like Texas wintergrass and Canada wildrye.

Hamilton County, Texas. May; peak standing crop.

66. One farmer's weed is another grazier's crop- Stand of Lolium perenne that invaded a field of winter wheat (Triticum aestivum). Ryegrass has naturalized in many areas of North America. Under certain conditions L. perenne has become a weed just like other introduced forage species including Johnsongrass (Sorghum halapense), Kentucky bluegrass (Poa pratensis), quackgrass (Agropyron repens), rescuegrass (Bromus catharticus= B. uniodes),, bermudagrass (Cynodon dactylodon), and tall fescue (Festuca arundinacea) as explained in Whitson et al. (1992).

(Incidentially ryegrass and tall fescue produce hybrids which fact was used as basis for renaming the latter by Barkworth et al. [2007], another bit of stretched logic thst was rejected by the author of Range Types.)

To return to the example provided here, this situation presented a major weed problem to the wheatgrowers. Ryegrass was so aggressive and well-adapted to cold conditions that it "choked out " the wheat crop in spots (like that shown here). Cultural and chemical treatments proved ineffective in controlling spread of ryegrass in this wheat field. This case of weed control was secondary to it serving as an example of the value and adaptation of ryegrass as an introduced forage species that naturalized, and now--"from now on out"--constitutes a pasture and range grass in North America. Or more simply put:"like or lump it" and "use it or lose it". This was outstanding winter pasture for cattle and deer. This stand showed just how naturalized Lolium perenne has become in parts of North American range.

Ottawa County, May; peak standing crop, soft-dough stage.

70. A study in spikelets; a lesson in detail- Proper nomenclature of perennial ryegrass (Lolium perenne) versus Italian ryegrass (L. multiflorum) has been a long-running debate among agrostologists and a source of confusion among practical pasturemen. For Missouri (and, hence, neighboring states) Kucera (1998, p. 46) recognized ryegrass as L. perenne and pointed out that "this species intergrades with closely related L. multiflorum", this latter being traditionally interpreted as Italian ryegrass. Kucera (1998, p. 46) joined these two legitimate taxa, which constitute "a continuum of forms", so that annual or biennial palnts with generally larger, more robust habit and that have "mostly awned" lemmas were interpreted as L. perenne var. multiflorum in contrast to L. perrenne var. perenne. Gould (1975, ps. 106-107) had earlier concluded that ryegrass in Texas was either annual or short-lived perennial and that Texas plants had considerable variation in both size and features of the whole plant as well as spikelets. The latter included the apparently pivotal characteristic of awned or awnless lemmas. Gould (1998, p. 107) concluded that splitting L. perenne into more than one species was not justified.

Then up and comes the what-should-be latest version of the "bible" of North American grasses in the series Flora of North American in which Barkworth et al. (2007, ps. 455-456) resort back to the two separate species of L. perenne and L. multiflorum based on awned versus mostly awnless lemmas and numbers of florets per spikelets with the range in numbers of florets overlapping between the two species! "The more things change, the more they stay the same". Clearly, Barkworth et al. (2007, 445-446) only increased the already existing state of confusion.

Even from the standpoint of keying out Lolium taxa a critical problem in dichotomy was that of "mostly awned" (Kucera, 1998, p. 46). Are lemmas "mostly awned" in the examples presented in these two photographs, or are they not? Obviously, the spikelets in these two photographs resemble most closely those of L. multiflorum in Figure 14.05 in Barkworth et al. (2007, p. 457).

Erath County, Texas. May; anthesis.

75. Timothy- Adult plant of common timothy at full-flower (anthesis). Drought-stunted plant that was only a fourth (or less) of normal adult height therby allowing a photograph that showed entire shoots in focus. In focus, that is in original slide. Even the best of photographers cannot be responsible for the condition of "picture perfect" after going through the "cap-and-ball" equipment and procedure of scanning as J-pegs.

Jackson County, Colorado. June.

76. Portion of timothy panicle- A close-up view of a timothy inflorescence showing spikelets. Threadlike tissues are the filaments of anthers that were beaten flat onto the panicle by recent heavy rains. Ottawa County, Oklahoma. June

The major use, and a most beneficial use, of redtop remains as a major hay crop in certain locales (eg. northwestern Montana). Redtop was first used for this purpose in the Hay and Dairy Region from New England (where it naturalized first) across to the Lake States until eventually it was displaced by timothy and smooth bromegrass. Redtop was also a pasture crop, second at one time only to Kentucky bluegrass (De France and Burger in Iowa State University, 1962, p. 239) but it was replaced as a pasture grass by species like tall fescue and ryegrass. Today, redtop barely rates one page in standard forage textbooks (eg. Iowa State’s Forages).

Redtop's story is the same as that of Johnsongrass and, to a lesser degree, species like bahiagrass (Paspalum notatum), the sweet clovers (Melilotus spp.), and the vetches (Vicia spp.). These were the best agronomic forage crops for certain areas at the time of their introduction, but gradually superior forage species were found and the earlier introductions were more or less replaced by later, more productive, introduced species. The "prototype" agronomic forages either died out or, worse, became major weeds (eg. Johnsongrass). Even bahiagrass is a weed where Coastal bermudagrass can be grown because the former has pretty much naturalized while the latter requires intensive farming, including control of bahiagrass, in order to persist.  Like the rhizomatous Johnsongrass and quakgrass (Agropyron repens), redtop can become a weed on cropland. It can also be used as a hay crop that requires little management beyond proper haying practices. Redtop hay production is still common in restricted locations especially where redtop is one component of meadow hay.
 
Redtop can also be very important in riparian vegetation where its rhizomes are outstanding for stablization of creek banks. The Forest Service (1940, ps. g8-g12) Range Plant Handbook a time-honored, always relevant reference that should be a part of every rangeman's library stated that while cattle preferred almost any other cultivated grass to redtop it was "highly regarded as range forage plant" that was "extensively raised as a meadow hay and pasture grass in the valleys of the West" (p. g9). In certain areas of the Western Range nothing has changed since those words were written. As water becomes an ever-scarcer resource and more expensive commodity, crops like redtop that require minimal management will likely become more commonplace perhaps replacing premium hay crops like alfalfa (Medicago sativa) that require more intensive cultural practices and costly inputs of water and fertilizer. The old standbys like the old virtues seem never to be completely replaced.

Redtop at peak standing crop and anthesis along a stream bank. Rocky Mountain National Park, Colorado. Early August.

80. Hay field dominated by redtop- This agronomic hay field and the one shown in the slide immediately below were on a river flood plain and consisted of naturalized communities of several introduced (Eurasian), cool-season, perennial grasses (list of species given in description below). Redtop was the dominant species on both hay fields. Photographs of these two fields were included as examples of tame (domesticated and, usually, introduced) grasses raised for hay. These slides exemplified the role of agronomic hay grasses in ranching and grazing land (mostly range) management across a large part of the North American Western Range.

Throughout much of the Intermountain West flood plains of rivers have been converted into crop fields. Field crops include "spuds" (potato [Solanum tuberosum]), sugar beets (Beta vulgaris), dent corn (Zea mays indentata), and alfalfa (Medicago sativa) just for starters. In other areas of this North American agricultural region, known by various titles such as the Grazing and Irrigated Crops Region or Range Livestock Region (a large portion of the Western Rage Region), this flood plain crop land is devoted to hay production from various introduced grasses (mostly cool-season, Eurasian perennials).

This latter situation obtains in areas devoted to range livestock (primarily beef cattle) where hay feeding is necessitated by relatively long winters with snow and/or severe cold. Alfalfa is frequently too valuable or expensive for dry beef cows and replacement heifers so if it is raised it is usually sold as a cash crop. More commonly grass hay is grown because it is less costly and risky to produce and produces higher yields of dry matter with adequate nutrients and energy for beef cattle, most of which graze summer range and come into winter in body condition. Such grass hay produced from commensurate base property (land and/or water-based) determines the Animal Unit Months of feed that a livestock producer has the privilege to buy as a lessee or permittee from public lands range, most of which is administered by the Bureau of Land Management or U.S. Forest Service.

In this situation hay production is the key crop to the range livestock operation. Often this hay (it is either baled and stacked as loose hay) is produced from native flood meadows (flood plains still in native herbaceous vegetation). Such flood meadows were treated herein under Grasslands, Meadows. In many other instances, however, the native hay meadows were converted into hay fields of introduced species in order to get higher yields and/or higher quality hay. (Many of the native flood meadows were [are] dominated by plants like cattails [Typha spp.] or bulrushes [Scirpus spp.] that are not highly palatable to livestock.) Type conversions from such native wetlands to wetlands of introduced cool-season grasses result in improved forage production from a stockman's perspective. Some of the introduced agronomic species appear to have about the same benefit for most wildlife species as the native wetland vegetation. Maintenance of natural wetland communities along with hay production from introduced grasses on modified wetlands has increased plant species richness in the region and appears to have been a net benefit. This seems to have been especially the case when there is a mixture of private and public wetlands with the latter being mostly natural wetland ecosystems.

There are numerous introduced, cool-seasson, perennial grass species that have become established on these improved-for-livestock-production, type-conversion, flood meadows. Most of these Eurasian agronomic grasses became more-or less naturalized and typically continued to propagate themselves (often rapidly invading new fields) once introduced. These introduced grassses do not appear to have become weedy or to threaten native wetland plants. Most of these flood plain hay fields consist of diverse communities of several agronomic grasses that continued to change as new species were introduced and naturalized at various times.

Common introducted grass species used for domestic hay production that can be found within the Grazing and Irrigated Crops Region include redtop, timothy, smooth brome, tall fescue (Festuca arundinacea), reed canarygrass (Phlaris arundinacea), creeping foxtail (Alopecurus arundinaceus), and meadow foxtail (A. pratensis).

The two hay field grass communities presented here and in the immediately succeeding slide were dominated by redtop. Other introduced grasses were timothy (second most abundant, the associate, species), tall fescue, creeping foxtail, and meadow foxtail (in smallest, trace, amounts). Each of these species grew in patches of pure stands of their own species more often than as a uniform mixture of evenly distributed species (ie. a patch of creeping foxtail here, a spot of tall fescue over there, and redtop about everywhere). Some of these grasses like redtop and creeping foxtail are strongly rhizomatious whereas others like timothy and tall fescue are bunchgrasses whose shoots consist solely of tillers. Method of asexual reproduction may account for some of the species dispersion and population patterns within hay fields. These different grass species were introduced at various times over the last century or so. Time of (and since) introduction may also account for dominance rank.

The hay fields shown in this and the next slide were on the flood plain of Silvies River in the Harney Basin of the southern Columbia Plateau. These fields were being flood irrigated by a combination of natural river overflow and man-made modifications. Local ranch owners have water rights and they irrigate their hay fields and native meadows in the spring by flood irrigation out of Silvies River which is (hopefully) filled with runoff from snow melt water. Natural spring flooding has been modified by series of cannals, ditches, etc. Water usage is based on water rights and is carefully regulated and policed by hired "water masters" who manage (usually) to keep water theft at reasonable minimums. (Certain property owners who shall remain nameless were notarious water theives.)

Traditionally these hay fields were not routinely treated with commercial fertilizer. Rather, beef cattle were fed and wintered on these same bottomlands from which the hay had been harvested. After cattle were moved off to spring and summer range these fields were hayed and the annual cycle begain again when cattle were brought back to the fields in the fall to utilize dormant grass regrowth and haying aftermath. Drags were sometimes used to break up the dung. Most producers saw no benefit to application of commercial fertilizer on this extensive cropping system with its attendant recycling of soil nutrients. No manure spreader needed, thank you. Some cattlemen, however, did see benefit to soil fertilization and used the practice-- if and when their banker went along with the idea.

Silvies River. Harney Basin, Harney County, Oregon. June. Peak standing crop. Harvesting hay when grasses were at optimal stage of maturity was impossible on these mixed-species stands (naturalized plant communities). Most plants of timothy were at soft-seed stage, tall fescue was mostly at seed-shatter stage, and creeping foxtail was beyond seed-shatter. The dominant redtop was in early bloom which, as countless studies with forage grasses have shown, is past optimal-- not to mention--peak forage quality. But hey, delayed haying made for great teaching photographs and illustrated physiognomy and dominance of these naturalized hay grounds.

81. Redtop growing on flood hay field- Redtop was typically the dominant (certainly aspect dominant) of hay fields in much of the Intermountain and Northern Rocky Mountain Regions. That phenomenon was shown here on a field that was flood- irrigated out of the Silvies River in the Harney Basin of southeastern Oregon.

The cropping practice and culture of haying on flood plains in this immediate vcinity and the general Intermountain Region was described in the immediately preceding caption. That treatment dealt with hay fields of domesticated and introduced (from Eurasia), cool-season, perennial grasses like the field shown here. Further discusssion of haying native flood meadows (in other words, the natural wetland vegetation), including in the Harney and Blitzen Basins, was treated in the section: Grasslands, Meadows.

The agronomic crop shown in this photograph was obviously dominated by redtop. In fact, as was frequently the case in this vicinity, the hay field was mostly redtop. Typically timothy was the second most abundant species in these fields, but the aggressive creeping foxtail with its stout rhizomes was frequently the most common of these introduced but now naturalized perennial grasses. Tall fescue, a strict bunchgrass (no stolons or rhizomes), was able to hold its own in large patches.

In these hay field communities timothy and creeping foxtail bear a close resemblance to each other so that mis-identification is possible for the unitiated. Creeping foxtail has a spikelike panicle that resembles the contracted panicle of timothy, but panicles of creeping foxtail turn dark at maturity at which stage they can be readily distinguished from the panicles of timothy. Also creeping foxtail sheds its grains earlier in the growing season which aids in distinguishing between these two species of the Aveneae (timothy tribe).

Redtop was typically the most aggressive-- hence most abundant (greatest cover, highest density of shoots)-- of the introduced perennial grassses. Redtop has the largest adult size among these specific agronomic species in addition to spreading and vegetative growth by creeping rhizomes. Redtop has stout shoots and large broad leaves, features that give it a competitive advantage-- particularily where resources like water are not limiting-- over species that have smaller plants. These features would seem to be even more important when on hay fields where grazing selectivity is not a factor and mechanical harvest is delayed until shoot growth is mature and adult size is reached. Production of long, broad leaves on rank culms suggested that redtop would be a superior species for higher-quality, high-yield hay production. However it is not a highly palatable species and often yields rank, chewy hay. (So what else does a dry cow on a winter feed ground have to do but chew hay?)

Silvies River flood plain, Harney Basin, Harney County, Oregon. June. Peak standing crop.

82. Stand of reed canarygrass (Phlaris arundinacea)- This rank-growing, rhizomatous perennial of the timothy tribe produces high yields of coarse forage that is highly variable in nutritive value, but it produces sod which is unexcelled for erosion control and in rate of sod-formation in the northern latitudes of North America. Reed canarygrass was one of the first introduced grasses to be cultivated for hay and pasture in North America. It is less cold tolerant than such Eurasian species as timothy, but it is better adapted to poorly drained sites and has long been widely planted in the Pacific and Interior Northwest Regions for both permanent pasture and hay as well as for soil conservation. While less important than some other festucoid grasses (as for eg. tall fescue, orchardgrass, timothy, and smooth bromegrass) reed canarygrass remains one of the major agronomic grasses in parts of the northern Western Range Region. This is evidenced by the fact that all five editions of the indispensable text, Forages (Iowa State University collaborators), devoted a chapter to reed canarygrass. Those desiring detailed information are, as always, referred to that set which forms an “encyclopedia” of forage crops.

Reed canarygrass was included here because it so often complements diets of range livestock and wildlife and is integrated into ranching and forestry operations in areas where it has proved adapted. Idaho County, Idaho. June.

84. Field of reed canarygrass in the Palouse Prairie- Two views of a field of reed canarygrass that remained on former Palouse Prairie. This small tame pasture on fertile, arable land was situated among fields of greater income-generating crops like Canola and had somehow avoided the plow. No, not native but it was at least an island of perennial forage grasss in an otherwise sea of intensively managed cash crops.

Whitman County, Washington. June, early summer.

85. Reed canarygrass panicles- Inflorescences of reed canarygrass growing on the pasture shown immediately above. Panicles of this species can have a waxen color like those presented previously or they can have the purplish-red coloration of these which were in late anthesis and early post-anthesis.

Whitman County, Washington. June, early summer.

87.    Hardinggrass (Phlaris tuberosa var. stenoptera)- This perennial grass is regarded as a native of the Mediterranean Region. It was introduced as an agronomic forage crop into Australia where adaptation to this continent’s Mediterranean and similar climates seemed likely. From Australia Hardinggrass was brought to the Mediterranean climatic region of California’s Central Valley and Coast Range where it was seeded on annual grasslands and brushfields (chaparral) which were converted to grasslands (type conversions) as well as on cropland for tame pasture. Hardinggrass has been the major— in fact, about the only —perennial grass to be well-enough adapted to the combination of California’s wet, cool winters and hot, dry summers to produce forage of the quantity and quality of  the naturalized Medterranean annual grasses and forbs, or even to persist in the severe, often harsh, Mediterranean climatic zone of California.

Unlike the highly rhizomatous reed canarygrass this Phlaris species is pronouncedly cespitose which may account for its relatively slow rate of establishment. Slower rate of spread notwithstanding, Hardinggrass has joined the ranks of other Mediterranean grasses which were introduced— purposely or accidentally —into California to become naturalized range species. The perennial, cespitose Hardinggrass has reached naturalization to substantially less extent than the annuals (to be expected given the example of extirpation of the native cool-season, perennial bunchgrass species), but it is apparently “here for the duration” and offers obvious advantages as a range forage species over the annual grasses.

Crampton (1974, p. 112) offered this assessment: "Hardinggrass is one of the most valuable forage grasses seeded on the valley and foothill rangelands in California. It is useful in seeding prepared land in the oak woodland of the foothills and cleared brushlands. Hardinggrass is slow in developing but after several years forms sizeable bunches and reseeds itself fairly well but not uniformly over any given range. The value of seeding this perennial on typically annual forage rangeland lies in its prolonged period of green forage beyond the time annuals have completely dried. Established plants green up before the fall rains and provide much succulent forage during the winter when the annuals are yet small”.

Butte County, California. June.

88. Hardinggrass seeded on chaparral (“brushfield”) converted to grassland- Bunches of Hardinggrass seeded on annual grassland. These clumps of Hardinggrass at the edge of a clearing in California chaparral of common manzanita and chamise were heavily grazed by sheep and black-tail deer. This slide illustrates two fundamental points. First, it is “proof-positive” that this introduced perennial bunchgrass is both palatable and capable of survival (sexual reproduction by abundant seed-set and asexual reproduction by tillering are obvious). Secondly, it demonstrates the value of the edge effect along with patches and corridors in providing prime wildlife habitat. (Notice how basic concepts from Landscape Ecology worked their way into the “clearing”.)

These defoliated grasses are a textbook example of the phenomenon of grazing selectivity and clearly show that animals do not graze plants in shapes like those clipped by human-held shears. The combination of native shrubland and naturalized grassland supporting both annual and perennial grasses is an example of the thorough planning and attentive management that are indicative of “wise use” husbandry of natural resources. Range and pasture management practices involved in the vegetation shown here varied from plant exploration and introduction to brush control to grazing management to livestock and wildlife husbandry.

Hopland Field Station, Mendocino County, California. June.

89. Another member of Aveneae, the oat tribe, that was introduced to North America as a pasture and hay species about two centuries ago. Tall oatgrass is best adapted to cool, moist climates like the Pacific Northwest, both Coastal and Interior. It does best on well-drained soils and does tolerate wet soils such as those of flood meadows. Tall oatgrass is fairly palatable (compared to other high-yielding, introduced grass species), but it is not well-adapted to heavy grazing, drought, or high temperatures, standard conditions sooner or later on most grazing lands. As such tall oatgrass is much less common and naturalized to considerably less extent than similar perennial, cool-season European or Eurasian species of the Aveneae (eg. redtop, reed canarygrass) not to mention Eurasian members of the Festuceae like tall fescue (Festuca arundinaceae), Kentucky bluegrass (Poa pratensis), smooth bromegrass, and orchardgrass or of the Hordeae such as perennial ryegrass (Lolium perenne).

The good part about these limitations is that tall oatgrass is less likely to become a weed on range and pasture. In the Palouse Prairie Region the author observed thick stands of rank-growing tall oatgrass persisting (frequently dominating) herbaceous plant communities adjacent to pastures of smooth bromegrass and reed canarygrass that essentially had no tall oatgrass. It was apparent that the latter could not compete with the former species under moderate grazing by cattle. Tuber oatgrass is a variety of tall oatgrass (A. elatius var bulbosum) that produces and spreads asexually via clusters of corms on or just below the soil surface (Hitchcock and Chase, 1950, p. 305). Tuber oatgrass was treated as a weed of northwestern North America by the Western Society of Weed Science (Whitson et al., 1992, ps. 416-417) it being similar in vegetative reproduction to Johnsongrass (Sorghum halepense) and quackgrass. Plants in the examples presented here were not corm-producing (at least the author's search for these modified shoots in this stand was not fruitful).

Tall oatgrass has long held a recognized though restricted place as an important, introduced forage grass. It has received treatment in all editions of the Iowa State University text, Forages, and in both editions of the Phillips 66-Fort Hays State University Pasture & Range Plants. The latter provided a pertinent, to-the-point summary of this forage species.Tall oatgrass appears to have become a naturalized species in specific locales (on local mesic, moderate habitats) on North America grazing lands. For this reason (and for as comprehensive coverage as practical in this publication) tall oatgrass was included herein.

Latah County, Idaho. June (early summer), hard dough stage of caryopsis.

91. Spikelets of tall oatgrass- Spikelets of tall oatgrass are similar to those of the Phalaris species in having spikelets of two florets in which the terminal or upper floret is perfect and the lower floret is staminate or neuter. Features of spikelets, including the relatively large, geniculate awn on the lower floret, and general morphology of the panicle were shown here.

Latah County, Idaho. June (early summer), hard dough stage of caryopsis.

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92. Stand of meadow foxtail (Alopecurus pratensis)- Alopecurus is another genus of the Aveneae that provided useful introduced forage grasses in North America, especially in portions of the Northwestern Range Region. The two most important of these are meadow foxtail and creeping foxtail (A. arundinaceus) which are interpreted as being closely related taxonomically (they certainly have confusingly similar morphological features). Traditionally, these Eurasian perennial grasses have been treated with other introduced grass species such as tall oatgrass that are best adapted to cool, moist habitats. The perennial Alopecurus species are much more tolerant of flooding or waterlogged soils than is tall oatgrass. For this rea

son meadow and creeping foxtail are much more common on wet ground such as flood meadows where they appear to have naturalized along with other perennial Eurasian introductions including redtop, reed canarygrass, smooth brome, and tall fescue.

Both tall oatgrass and the foxtails are examples of introduced forage grasses that initially showed much "promish" (at least "potential"), but that have become less valuable agronomically over time (see for example coverage in the various editions of Forages). These species do seem to have naturalized and become lasting members of ranges and pastures thoughout their range of adaptation, but this also appears to be of restricted local abundance as, perhaps, pests (including arthropods and fungi) continue to evolve adaptations to better utilize these introduced food sources.

As hay and livestock producers planted first one and then the next new "wonder grass" that "promished" to "revolution" forage production, their pastures, hay meadows, and adjoining ranges came to be mixtures of native and naturalized plant species. An example of this in the Northern Rocky Mountains was shown and described below.

The stands of meadow foxtail presented here were growing side-by-side (= shoot-by-shoot,-root-by-root) with the tall oatgrass shown immediately above on a hillside in the Palouse Hills (Latah County, Idaho; June (early summer).

93. Individual plants of meadow foxtail - Two examples of large, individual plants (genotypes or genetic individuals) of meadow foxtail growing (apparently naturalized and self-seeded) on a pasture and a waste area on former Palouse Prairie steppe. Meadow foxtail as an introduced, perennial forage grass could be summarized similarly to tall oatgrass, except that this current member of the Aveneae is better adapted to disturbance and stress, including heavy grazing, wet soils, and, being rhizomatous, is more readily or efficiently dispersed vegetatively (other than the corm-producing tall oatgrass).

Whitman County, Washington and Latah County, Idaho, upper and lower photographs, respectively. June (early summer).

94. Sexual shoots of meadow foxtail- .Against the dark background of early morning light grain-bearing shoots of meadow foxtail shoed up to good advantage in the pastoral Palouse Hills. These shoots were produced by plants that were growing were they had been established (self-seeded) from apparently naturalized plants. These plants were not waifs or adventive individuals in the fashion of roadside volunteer corn or wheat. Rather they were more of the nature of now-thoroughly adapted weeds that "ain't native, but we got here as soon as we could and we're staying".

Latah County, Idaho. June (early summer); grain-ripe phenological stage.

95. Panicles of meadow foxtail- Members of the Aveneae have a panicle type inflorescence. Some of these like timothy (shown and esscribed aobove) and the foxtails have a condensed panicle with secondary branches of the flower cluster being extremely short but nonetheless attached to the central axis. This central axis of the inflorescence is not, strictly speaking and contrary to common usage, a rachis By whatever name, the spikelets readily shed from the "core" or "main line" of the panicle leaving a bare stalk or core when dissemination is complete.

Latah County, Idaho. June (early summer); grain-ripe phenological stage.

96. Sandhills subirrigated pasture- Pasture of a mixture of Kentucky bluegrass and creeping foxtail (Alopecurus arundinaceus, Garrison, cultuvar) on a subirrigated meadow in Nebraska Sandhills. Some other incidental plant species (including annual Mediterranean Bromus spp.) were present. Species composition of this pasture was not determined but an approximatae "half-and-half" would not be far off.

Details of creeping foxtail presented below under Aveneae.

Cherry County, Nebraska. late June (early summer). Predominately hard-grain for foxtail and seed-shatter stage for bluegrass.

97. Just creeping along- Shoot base and top of roots (= rootcrown or proaxis) of Garrison creeping foxtail, complete with nice example of a rhizome. This plant was growing in the subirrigated Nebraska Sandhills pasture presented immediately above.

Cherry County, Nebraska. late June (early summer).

Grasses in the flood meadow pasture shown here included smooth and meadow brome, creeping foxtail, reed canarygrass, and timothy as the more common species and with scattered but conspicuous individuals of tall fescue, orchardgrass, and Great Basin wildrye. Idaho fescue and sedges were common along field edges and the outside of fencerows. No legumes were present.  No FRES, Kuchler, or SRM designations. Gallatin County, Montana. June.

99. Common bermudagrass (Cynodon dactylon)- Bermudagrass in all it's taxonomic forms ranging from common to the numerous cultivars, including F1 hybrids (both fertile and sterile), is the single most important permanent pasture grass in the southern United States. Bermudagrass is most likely native to southeast Africa. It was probably introduced into the North American Colonies before the Revolutionary War. By the late Nineteenth Century bermudagrass had become the most important tame pasture grass in the South. Research and development on bermudagrass has continued unabated even during World Wars. Cultivars like Coastal, Midland, Suwanee, Callie, Hardie, Brazos, Grazer and several Tiftons have probably been the single greatest factor in revolutionizing (and that is not an overstated verb) beef production in the South. Only crossbreeding of beef cattle would approach the profound impact of bermudagrass breeding in a vast region extending from Virginia across the Upland South of Tennessee and Kentucky through Arkansas and Oklahoma and "parts south" thereof. Plus, these higher-yielding cultivars are grown under irrigation across the Southwest from west Texas clear to and including California.

This form of grassland agriculture is some of the most intensive tame pasture management and forage production systems in the world. It is most certainly not a form of range management nor part of the discipline of Range Management. The cultivars of bermudagrass are cultigens (organisms, generally thought of as being at species or lower levels or taxa, that can exist only under domestication and not known in a wild form and presumably originating under domestication; the term is used more with plants than animals). These bermudagrass cultivars are as much dependent on continuing intensive human inputs (soil tillage and ferttilization practices, plant breeding, precise harvest) as is Zea mays. The subject of bermudagrass management and bermudagrass-based livestock production was deemed inappropriate for a publication devoted to natural pasture vegetation types. Readers desiring coverage of the agronomic management and production of bermudagrass are referred to the seemingly limitless literature available. This ranges from basic texts like Iowa State's Forages to "jillions" of publications cranked out by the Agricultural Experiment Stations and Agricultural Cooperative Extension Services of the southern states, the Agricultural Research Service, various seed companies, and private foundations (eg. the notable George Noble Foundation headquartered in southern Oklahoma).

In contrast to the cultigen nature of the bermudagrass cultivars, common bermudagrass has naturalized (at least under anthropogenic influences such as introduction and continuing disturbances aside from agronomic management) over a vast portion of North America. This area of naturalization ranges from Wet to Semiarid precipitation zones and from the general tropical zone northward nearly to southern Canada, literally from Atlanic to Pacific. In fact, bermudagrass is one of the more widespread forage species in the world. Bailey (1949, p.149) described the distribution of C. dactylon as "now widely spread over the globe".

Over much of this range of naturalization bermudagrass is used or serves as a naturalized range species, including several parts of North America. Bermudagrass has an ecological role on southern ranges similar to that of the naturalized festucoid species like Kentucky bluegrass, timothy, orchardgrass, and the various introduced wheatgrasses. In the classic Western Grazing Grounds and Forest Ranges Barnes (1913) included bermudagrass (complete with line drawing) along with buffalograss, curly mesquite, galleta, and threeawns as one of the "southwestern grasses". (Bermudagrass has been regarded as being less productive than the natives on semiarid range, but it does grow on plains grasslands.) Bermudagrass was included as one of the 200 North American range plant species on the Master Plant List for the International Range Plant Identification Contest under leadership of the Society for Range Management (Stubbendieck et al., 1981, 1982, 1986, 1992, etc.). In that context C. dactylon was included in the present publication.

Bermudagrass is sometimes a major naturalized forage species on ranges in the coastal prairies and marshes from Texas eastward along the Gulf and Atlanic Coasts. Common bermudagrass has been viewed as quite tolerant of close grazing, but it may be less tolerant of heavy defoliation than are such introduced bunchgrasses as King Ranch bluestem or even tall fescue. Common bermudagrass reproduces-- and can be propagated --by both seed (grain) and "sprigs" (the term used by growers when referring to stolons and rhizomes collectively). Bermudagrass is cross-pollinated, a factor that has contributed to it's rapid adaptation to a wide arrary of climatic, edaphic, and other environmental factors. Asexual reproduction by sprigs accounts for both the rapid spread and grazing tolerance of this sod-forming grass (the same as in cases of such native sod-formers as buffalograss or curly mesquite). However on rocky, shallow soils typical of those on which common bermuda is naturalized in the Midsouth and central highlands like the Ozark Mountains, bermudagrass stolons can be readily ripped up by grazing cattle and horses thereby greatly increasing degree of use and often wasting as much or more forage than was eater. This phenomenon coupled with the drought-avoidance inherent in cool-season perennials like tall fescue may explain why farmers and cattlemen commonly observe that bermudagrass, even the more adapted common bermuda, is not as tolerant of overuse (or even just heavy grazing) as it was originally touted to be. As with all forage and browse species there is no substitute for proper grazing management, beginning with the First Cardinal Principle of Range Management: Proper Degree of Use.

Bermudagrass is automatically classified as an ecological invader because it is an introduced or alien species on North American range. On range where common bermudagrass has naturalized, and is manged as if it was native, it has an ecological response on some of the more xeric range sites more like that of an increaser (similar to such stoloniferous species as buffalograss or curly mesquite, which are alternatively decreasers, increasers, or invaders depending on range site).

Erath County, Texas. July.

100. Bermudagrass sod- A stolon of bermudagrass was extended above the sward level to expose the stolon and modular unit of a "daughter or sister plant" produced along this extravaginal shoot of a clonal organism. Each "sister plant" is a ramet or module of the overall genetic individual (the genotypic plant) which is the genet or clone. A clone is a metapopulation of modules or ramets. Concept of clonal organism as applied to range plants, especially grasses, was covered quite well by Dahl in the reference edited by Bedunah and Sosebee (1995, ps. 27-30).

The upright position of the new module suggest that it is a tiller (an intravaginated shoot), but it is another stolon that grow horizonally and that will produce other stolons, each of which pierces the leaf sheath rather than emerging from within it. The latter form of shoot (with a maintained vertical orientation) is a tiller (also called "sucker" or "stool"). "Tiller" has sometimes been used in the generic sense for shoot. This is very poor form as it is extremely confusing to beginning students trying to master the concept of intra- (ie. vertical) vs. extravaginal (ie. horizonal) shoot.

Erath County, Texas. September.

101. Bermudagrass stolons- Shoots (the culm or stem and leaves) of grasses are, as was explained immediately above, either intravaginal or extravaginal (elongating up through the enveloping leaf of the phytomer or piercing through the leaf sheath of the phytomer, respectively). Intravaginal shoots are tillers and extravaginal shoots are either stolons (aboveground horizonal shoots) or rhizomes (belowground shoots). Stolons are also known as "runners"; rhizomes as "rootcrowns". Both are also called and known by commercial growers as "sprigs". (ie. stolons and rhizomes are "sprigs"). "Sprigs" more commonly refers to rhizomes when hybrid bermudagrasses like Coastal are "sprigged". Such "sprigs" are often baled and sold in that unit. Common bermudagrass is "sprigged" by planting the "runners" or "runners" complete with rhizomes. Adventituous roots develop at nodes along these horizonal shoots (again, clum + leaves and including flower cluster).

Grass species that develop exclusively or primarily by tillers when they undergo vegetative (asexual) reproduction have a cespitose habit or morphological form and are labeled "bunchgrasses". Grasses reproducing vegetatively by stolons and/or rhizomes are, by contrast, "sod-forming grasses". Some grasses produce both intra- and extravaginal shoots (eg.some varieties of big bluestem, Indiangrass, and sideoats grama produce short rhizomes as well as tillers).

Several stolons of common bermudagrass were shown here pioneering or invading a bare patch of soil on the famous Oxford Tract of the University of California, Berkeley. Alameda County, California. October.

102. Modules of of bermudagrass- This is the fundamental unit of the stoloniferous bermudagrass: a portion of the "runner" had two sister plants (ramets or modules) developing from buds of intercalary meristem along this stolon. One of these modules developed an inflorescence such the reader views both sexual and asexual reproduction of this "versatile species". One should note also roots forming at the node just below the newly forming clonal shoot. This is a textbook example of a clonal organism.

Erath County, Texas. September.

103. Flowering shoot apices of bermudagrass- Common bermudagrass in anthesis. Bermudagrass, like other perennial sod-forming and even tillering or bunchgrasses reproduce primarily by vegetative means (asexually), but most also set viable seed and are capable of sexual reproduction. Sterile hybrid bermudagrass cultivars like Coastal can only be propagated by "sprigging" (planting rhizomes and.or stolons), but common bermudagrass can be seeded. Results of seeding have been extremely variable and sprigging has been the more accepted practice. At one time seeding was more common. In fact, some cattlemen established reasonable good stands of common bermujdagrass by "feed seeding", the practice of mixing bermudagrass seed in with concentrate feeds or meal mixes so that bermudagrass grains were "planted" in dung across pastures. The author witnessed an example of where this method established common bermudagrass on formerly overgrazed pastures in the Ozark Plateau of eastern Oklahoma.

Erath County, Texas. September.

104. Inflorescences of bermudagrass (lateral-view)- The inflorescence type of bermudagrass and other members of the Chlorideae tribe has traditionally been regarded as consisting of one or more one-sided spikes that are solitary and digitate or racemose. Sometimes the Chlorideae inflorescence was described as a spikelike raceme with these solitary, digitate, or racemose along the rachis or main axis. More recently, authors have taken to using rather elaborate and more descriptive terms for inflorescences of the Gramineae, in particular for the panicle inflorescence type. In this regard there are several forms of a panicle having spicate primary unilateral branches. Bermudagrass could be interpreted as having a specialized panicle inflorescence described as a panicle of digitate spicate primary unilteral branches (Hignight et al. 1988, p.7).

Erath County, Texas. September.

106. Rhodesgrass (Chloris gayana)- This species is native to most of sub-Sharaian Africa. It was one of numerous other species including weeping lovegrass, Lehmann's lovegrass, bermudagrass, buffelgrass, and Johnsongrass that was introduced into North America from one part or another of Africa. Rhodesgrass was a relatively early introduction but, as as was proven to be the case with several of the earlier introductions, some of the other introduced species proved to be more productive, easier to establish, more readily maintained, more palatable or nutritious, more profitable, and generally more desirable forages. It was seen as revealing that Rhodesgrass received more coverage in early forage crop texts such as Forage Plants and Their Culture (Piper, 1939) than in recent ones (eg. later editions of the forage "bible" Forages out of Iowa State University Press, 1985, 1995).

According to Piper (1939, p. 282) the first recorded cultivation of Rhodesgrass was by Cecil Rhodes in 1895 and the species was introduced into the United States in 1902. For a period, Rhodesgrass was cultivated along the Gulf Coast from Florida to Texas and westward to southern Arizona and California. While Rhodesgrass is still commonly found over much of this coastal strip such occurrence is now more of occasional stands or isolated plants. Like many, if not most, introduced forage species Rhodesgrass apparently naturalized enough to maintain itself but usually only as a major or dominant species in periodically or formerly severly disturbed environments like roadways and abandoned areas (old fields, city lots, closed school yards) that are not subjected to continued heavy grazing or mowing.

Currently the center of importance of Rhodesgrass as an important agronomic forage in North America is south Florida. It is generally assumed that the main factor responsible for "demise" of Rhodesgrass in Texas (once its main production location in the United States) was sensitivity to Rhodesgrass scale or Rhodesgrass mealybug (Antonina graminis). This pest is also an alien species (from Asia). Rhodesgrass mealybug has been controlled to large degree in Texas through a classic biological control campaign using a parasitoid (Anagyrus antoninae) imported from India (Chada and Wood, 1960). In fact this was an early cooperative control program between United States Deparatment of Agriculture and Texas Agricultural Eaxperiment Station. The Texas cultivar, Bell (a diploid), has considerable resistance to Rhodesgrass mealybug. Notwithstanding, Rhodesgrass scale was a major factor in reduced populations and use of Rhodesgrass as a forage in south Texas.

Other limiting factors included limited winterhardiness and availability of superior forages such as buffelgrass (Kretschmer and Pitman in Forages, 1995, p. 298). Likewise, Rhodesgrass often has reduced palatability especially at advanced maturity and when lighter grazing results in rank forage (large, overly mature stalks). Coincident with this is the limited ability of Rhodesgrass to tolerant close grazing as compared to lower-growing grasses like bermuda and even more upright bunchgrasses like buffelgrass. Rhodesgrass does withstand trampling and frequently recovers quickly from abusive grazing when soil is fairly moist and fertile.

Rhodesgrass is one of the perennial woarm-season grasses most tolerant of salinity. Rhodesgrass also tolerates alkalinity better than many agronomic species. Rhodesgrass is drought-tolerant, but less so than buffelgrass. The important cultivars of Rhodesgrass are diploids or tetraploids (Kretschmer and Pittman in Iowa State University, 1995, p. 297). Rhodesgrass remains on of the most important domestic forage species in its native Africa and Australia, but in most of the southern range region it is mostly important as an infrequent though persistent naturalized grass that added biodiversity, erosion control, and opportunistic forage where forage is all too often severely limited.

Rhodesgrass is a stoloniferous species with clones developing along stolons and subsequently forming cespitose units as tillers proliferate from these daughter plants. This feature was visible in the example shown here where the smaller "plant" at the left was an offshoot from the older and larger bunch on the right.

Kenedy County, Texas. February, peak standing crop and soft grain stage of phenology.

107. Inflorescence of Rhodesgrass- An infloresecence from the specimen in the preceding photograph displays the arrangement of racemes in the flower cluster. Recent treatments of the grass inflorescence have deviated from the standard panicle, raceme, or spike type, kind, or pattern of earlier authors. Modern treatments of Gramineae taxonomy described the inflorescences of Chloris species as panicles of spicate primary unilateral branches that are verticillate, digitate, or aggregated on upper portions of the rachis (Highnight et al., 1988, ps. 7, 22; Hatch et al., 1999, ps. 15, 50-51, 101). The main, major, or largest unit arrangements of spikelets in several of the Chloris species, including C. gayana, are joined at base of the rachis (peduncle) such that these primary unilateral branches are close together forming a "tight" rather than "loose" or "open" infloresecence that is currently designated a panicle.

Kenedy County, Texas. February, soft grain phenological stage.

108. Rhodesgrass stand- Rhodesgrass growing on a saline disturbed habitat. Development of Rhodesgrass clones at intervals (usually on or near nodes) along short, stout stolons was shown in this photograph. Two other morphological features of Rhodesgrass were displayed in this example: 1) stemy or stalk-like feature (relatively large tillers, especially diameter-wise dimension of culms) at maturity and 2) abundant floral (hence, grain) production. Large culms are largely responsible for fairly low palatability and nutritive value of mature Rhodesgrass forage. This feature has often been described variously as "rank", "big-stemmed", or "a lot of straw". Such large stems (relative to overall size of plants) make for high yields of low-quality herbage due to high quantities of cellulose of low digestability that is responsible for low voluntary forage intake.

High levels of seed (grain) production explain why Rhodesgrass is easily established from seedings and readily self-seeds in bare spaces. High levels of seed production combined with asexual (vegetative) reproduction from stolons enaables Rhodesgrass to recover quickly from damage to stands resulting from drought, overgrazing, freezing, and Rhodesgrass scale (mealybugs). On the other hand, these recovery adaptations come at relatively high costs of eventual favorable soil moisture (ie. rain, sooner-or-later, and soils of high moisture-holding capacity) along with farily high soil fertility. Rhodesgrass is like other highly productive grasses in requiring high levels of soil nitrogen and water to achieve its potential for forage yields.

Boca Chica subdelta of greater Rio Grande Delta. Cameron County, Texas. February, full-maturity and early dormancy; seed-ripe to seed-shatter stage.

109. Detail of Rhodesgrass inflorescence- Inflorescence from Rhodesgrass stand in preceding photograph. Descriptions of Gramineae inflorescences have traditionally been confusing with reasonably nonconfusing ones "hard to come by" at best. Describing flower clusters of the Eragrostoideae has been especially problematic. At present, inflorescences of Chloris species have probably been most precisely described as panicles of spicate primary unilateral branches in verticillate, digitate, or aggregated arrangements along the rachis (Hignight et al., 1988, ps.7, 22; Hatch et al., 1999, ps. 15, 50-51, 101). The primary branches of the infloresecnce in C. gayana originate from a common location on the peduncle (base of rachis) and extend upward resulting is a rather tight cluster (tightly clustered panicle).

Many full spikelets were clearly visible on the unilateral primary branches of this specimen. This was an example of abundant grain production in this warm-season, perennial forage grass. Basis of the characteristic self-seeding (reseeding) of Rhodes-grass was evident. Such plentiful sexual reproduction combined with effective asexual reproduction by stolons and tillers (preceding and succeeding slides) is responsible for ease of establishment and fairly effective maintenance of Rhodesgrass stands in the Guld coastal south of North America.

Berm of road, Boca Chica subdelta of greater Rio Grande Delta. Cameron County, Texas. February, seed-ripe or early seed-shatter phenological stage.

110. Individual plant of Rhodesgrass- Typical habit of a genetic individual of Rhodesgrass. This plant had produced abundant numbers of tillers resulting in a cespitose or bunchgrass growth form, but visible in right-of-center foreground was a new "clump" that had risen from a short, thick stolon off of the full-sized bunch. This vegetatively reproduced unit (a daughter plant) was producing new tillers and on its course toward development of another bunchgrass of the same genotype.

An extremely saline microsite on Boca Chica subdelta of greater Rio Grande Delta. Cameron County, Texas. February, full-maturity and early dormancy; seed-ripe to seed-shatter stage.

112. Pasture of weeping lovegrass- This field of weeping lovegrass was also a scrub-to-grassland type conversion. This lovegrass pasture was previously a sand shinnery oak (Quercus havardii) scrub type, a climax shrubland. (Slides of the shin oak ecosystem were presented under Shrublands, Shinnery Oak: view slides from Wheeler County, Texas for the climax scrub in the immediate area of this weeping lovegrass type conversion.) The weeping lovegrass in the photograph on the left (with cattle) was at the phenological stage of pre-anthesis but fully expanded panicle; the same stand in the photograph on the right was 10 days later when weeping lovegrass was post-bloom with grain at milk stage.

This stand of weeping lovegrass was "pretty" and presented a striking pastoral scene with big crossbred Charolais cattle in grass over "belly deep", but this is not proper management for weeping lovegrass. This grass "got ahead of the cattle" and became "rank" (ie. over-mature so as to be lower in general nutritive value and digestibility due to greater concentrations of low-quality fiber components, cellulose in particular, and lesser contents of digestible energy, protein, etc.). Most importantly, the forage reached the stage of maturity were it was extremely unpalatable which, in conjuction with lower digestibility, resulted in lower voluntary forage intake than if the lovegrass had been grazed heavier and consequently kept at earlier maturity stage(s). Conservative stocking and/or light degree of use like that seen here would be acceptable or perhaps even ideal management (depending on many factors) of native grass species like giant dropseed (Sporobolus giganteus), sand bluestem (Andropogon hallii), and sand lovegrass (Eragrostis trichodes). Proper grazing management of the introduced weeping lovegrass, however, is a heavier degree of use (to which this species is adapted) to maintain palatability and nutritive value. As is, it did make for a "purty picture" (but "pretty is as pretty does").

Wheeler County, Texas. June.

114. Portions of panicles of weeping lovegrass- Spikeles of weeping lovegrass were shown at pre-anthesis stage. Erath County, Texas. April.

115. Lehmann lovegrass (Eragrostis lehmanniana)- This native of southern Africa was introduced into North America in 1932 by the Soil Conservation Service to halt severe soil erosion and aid revegetation of land in arid and semiarid regions of the Southwest. Lehmann lovegrass had the essential features required for these purposes. It was readily established by seeding and produced abundant seed crops that allowed for readily available, economical planting material. Lehmann's lovegrass also reproduces asexually by adventituous rooting at nodes to increase cover and density within stands and increase stand persistance.

Given the then-desirable characteristics of aggressive, efficient reproduction and high herbage yields for semidesert habitats (and all this was economically feasible), Lehmann's lovegrass was viewed as a "wonder grass" by cattlemen, Extension agents, and soil conservationists. The Tucson (Arizona) Plant Materials Center released three cultivars of Lehmann's lovegrass: A-68 in 1950 and Kuivato and Puhuima in 1976. Lehmann's lovegrass was widely planted.

Unfortunately-- at least from one major respect-- Lehmann's lovegrass was too "successful", too "wonderful" for it's good. As has been the case for other aggressive introduced forage species like Johnsongrass, quackgrass, or kudzu, Lehmann's lovegrass became a major range weed (to some extent). It spread from severely depleted ranges where it functioned as needed into native range communities (many of which had experienced some deterioration from mismanagement and droughts) where it was not needed and where in time it replaced, to varying degrees, native range plants. Many of these were desirable climax grasses like sideoats grama, black grama, threeawns, tanglehead (Heteropogon contortus) and cane bluestem (Andropoogon barbinodis= Bothriochloa barbinodis var. barbinodis) that were superior forage species. Selective grazing by livestock, cattle in particular, of the more palatable native grasses and forbs provided a competitive advantage to the reproductively superior Lehmann's lovegrass. The predictable result was displacement of the native range plants by the now-naturalized introduced species.

Superimposed over all this was the sociopoliticoeconomic shift that accompanied the transformation of much of the Southwest from a rural, primary industry culture to an urban, service-based one. Emphasis (ie. "politics") shifted from ranching and efficient food-fiber production to "environmental concerns" ranging from such valid issues as biological diversity and preservation of natural plant and animal communities to jealous resentment of agricultural producers by city slickers who drive recreational vehicles, join the Sierra Club, and solve the problem of Lehmann's lovegrass invasion by converting the natural vegetation into crabgrass monocultures sandwiched between paved streets and cul-de-sacs. In essence, Lehmann's lovegrass is not "politically correct". It went from Lone Ranger status to the noxious plant list, the equivalent of the FBI "ten most wanted".

Whatever the complex of factors (some legitimate; some not, depending on assumptions and value judgments), the fact is that Lehmann's lovegrass is now an introduced species "here to stay" as both a naturalized range plant that is a component of native plant communities and as reseeded range to conserve soil and increase commodity turnoff (eg. beef production).

Yavapai County, Arizona. June.

118. Lehmann's lovegrass range in Texas Trans-Pecos Basin and Range- Lehmann's lovegrass has been seeded over a large portion of the greater Chihuhuan-Sonoran Deserts Region from Trans-Pecos Texas to central Arizona and south into the western states of Mexcio. Lehmann's lovegrass has naturalized over much of that "super-region", and beyond into adjacent regions such as the Rio Grande Plains.

The reseeded range presented here was typical of numerous old fields and overgrazed ranges that had been reseeded to this introduced species. Net results were stoppage of accelerated soil erosion (and general improved soil and water conservation) as well as provision of dependable perennial forage sources even in prolongued, destructive drought that is a key feature of arid regions. Some native shrubs had re-invaded this medium term grazing land (Vallentine) which a desirable successional development, as long as such invasion is not excessive. In fact, some native shrub invasion should be an anticipated as a "fringe benefit" especially for production of certain wildlife species including upland birds along with ungulate browsers.

To some extent (though to lesser degree) Lehmann's lovegrass has been to southwestern North America what the crested whetgrass species were to the Great Basin, Rorthern Rocky Mountain, and Northern Great Plains regions. That Lehmann's lovegrass has naturalized attest to the fact that this intrtoduced species does constitute a rangeland cover type. Interestingly, Lehmann's lovegrass was not included in the Society for Range Management publication, Rangeland Cover Types of the United States (Shiflet, 1994), even though crested wheatgrass and even tall fescue were described as rangeland cover types. "Go figure". This author concluded that the situation regarding SRM rangeland cover types is similar to that of SRM awards. What and who gets such recognition depends on willingness of someone to "write them up".

Jeff Davis County, Texas. June, immediate pre-bloom stage.

119. Lehmann's lovegrass and Longhorns- Longhorns may not love Lehmann's lovegrass (few, if any, of the introduced range and tame pasture species are as palabtable as the original native species), but this drought-tolerant, persistent, perennial grass is very productive of fairly nutritious forage across much of the vast region of southwestern North America. "Beats hell outa holler belly". Furthermore, where romance, recreation, wilderness, wildness, and trophy horns and hides count for more than profitable red meat production Lehmann's lovegrass is frrequently "just what the doctor ordered".

Creosotebush (Larrea tirdentata), zonal dominant of the Chihuhuan and Sonoran Deserts, had reinvaded this seeded field of Lehmann's lovegrass. At density and cover shown here such woody reinvasion was desirable and to be expected. Beginning range management students should remember that one of the advantages of reseeding range is to facilitate plant succession, and that includes climax shrubs (at proportions characteristic of climax vegetation or the desired seral state). Vegetation like that shown here is range not tame or agronomic pasture. This is naturalized rangeland not a farmed field crop where any species not seeded is a noxious plant.

Presidio County, Texas. June, full-bloom stage and peak standing crop.

120. This robust plant of Lehmann's lovegrass was growing in the Chihuhuan Desert-semidesert (Chihuhuan) grassland ecotone in the eastern part of the arid zone. This massive specimen was growing of the Longhorn range shown above. Such "purple-ribbon" plants illustrated how effectively Lehmann's lovegrass can survive and reproduce (both sexuallly ans asexually) in habitats that are less than hospitable if not downright hostile to most living things.If nothing else, Lehmann's lovegrass is an adaptation success story. This species should not be used as the poster plant for invasinve plant species.

Presidio County, Texas. June, full-bloom state and peak standing crop.

121. Lehmann's lovegrass at full-bloom stage. This plant was also growing on the Longhorn range presented above. Presidio County, Texas. June.

124. Kleingrass pasture- Field of kleingrass was in the northeastern portion of the Texas Edwards Plateau in early autumn following a number of years of prolonged drought. Peak standing crop at seed-ripe to seed-shatter phenological stage. This was an example of outstanding pasture and livestock management. This rancher had stocked properly for the drought which meant as low a stocking rate as possible to insure adequate forage for the indefinite duration of short feed conditions. Also, cattle husbandry was outstanding. Brood cows with husky calves were gentle even around the stranger who was "taking their picture" along with that of their naturalized range. A landscape photographer knows immediately which cattle have been treated with respect and which ones were "choused" and treated like rodeo stock.

Kleingrass in this advanced stage of maturity is much lower in nutritive value (higher in fiber while lower in digestability, energy, protein, etc.and therefore less voluntary forage intake). Heavier grazing would have kept kleingrass plants at less mature stage(s) and been superior grazing management-- in a typical growing season. In drought, especially a prolonged drought of several years duration, range managers must stock lighter to hedge against forages shortages. The result is more herbage or lower nutritional quality but plenty of feed left as "insurance".

Most importantly over the long run is that such conservative stocking rates and lighter degrees of utilization maximize the likelihood that the "grass" or the "range" (ie. the forage crop) will survive. It is very sad and discouraging to have to sell off genetically superior livestock and loose the financial investment in such animals, but it is even sadder and far more futile to try vainly to hold on to them until it is too late and the range has been overgrazed and the stockmen still had to sell livestock at a loss. Now the stockman's basic crop--"grass"--is lost (or will take many years to recover) along with his beloved livestock. Now investments in both livestock and land have been lost or greatly reduced in value.

The rancher who owned (or rented) this kleingrass pasture and these fine cattle knew what business he was in: the "grass" business, first; the cattle business, second.Outstanding stewardship.

San Saba County, Texas.October.

128. Field of guinea grass (Panicum maximum)- An old-field in floodplain of Rio Grande was naturally and densely populated by naturalized guinea grass. Cropping history of this go-back land was incompletely known as, for that matter, is history of naturalization of guinea grass. Naturalization of guinea grass was probably one of the most rapid and successful cases of the plant naturalization phenomenon is recent history. The prevailing account of naturalization of guinea grass is that grain and/or vegetative material of this exotic panicoid was spread from a branch station of the Texas Agriculturaal Experiment Station due to flooding caused by Hurricane Beluah in 1968. In only three or four decades (and perhaps as soon as a quarter century) guinea grass had naturalized throughout much of the southern portions of Rio Grande Plains and Coastal Marshes and Prairies vgetational areas of Texas.

Whether dominance by guinea grass over hundreds of thousands of acres will remain over time or whether this remarkable rate of spread will be halted or reversed by some heretofore unknown disease, parasite, insect, etc. pest remains to be seen. For now range invaded and dominated by guinea grass remains successfully vegetated by a reasonably palatable and highly productive introduced species.

Audubon Sanctuary, Cameron County, Texas. October, seed-set to seed-shatter stage.

129. Individual plant of guinea grass- One of the common characteristics of high-yielding (heavy production of biomass) forage species is large mature size of adult plants (think silage cultivars of dent corn for example or the more upright-growing cultivars of bermudagrass). This large specimen of Panicum maximum with its "countless" shoots exhibited the "big plant" feature. Guinea grass grows in large clumps and has a bunchgrass habit, but it is not a strictly cespitose species. Instead guinea grass spreads gradually by vegetative (asexual) reproduction from a rhizomatous rootcrown. so that clumps of genetic individuals continue to enlarge (more shoots of the same genotype are produced) and increase basal and aerial cover by the large species that was most likely originally a native of Africa. Guinea grass is now a major pasture grass in grazinglands throughout tropical and subtropical regions around Earth.

Audubon Sanctuary, Cameron County, Texas. October, seed-ripe phenological stage.

130. Guinea grass shoots- It has been remarked at various points in this publication that grasses are far from photogenic. Absence of large flowers with showy petals and instead delicate spikelets on dainty panicle branches atop jackstraw arrangements of half-lodged shoots as in this guinea grass does not make for a clear presentation of a plant species. Nonetheless, presented here was a view of guinea grass that showed lower portions of culms, leaves, and flowering shoot apices. Potential for high yields of forage biomass from this species was obvious.

Sabal Palm Audubon Sanctuary, Cameron County, Texas. October, full-bloom/ peak standing crop phenological stage.

131. Dallisgrass (Paspalum dilatum)- Dallisgrass was another one of the first warm-season perennial grasses introduced into the United States. Like Johnsongrass and King Ranch bluestem this species was introduced, at least initially, for a tame (= domestic) pasture grass. (It may have been introduced inadvertently.) Dallisgrass has long-since naturalized but to a lesser degree than the other two species. Dallisgrass is therefore much less of weed problem than the latter two aggressive ruderals. Like them, however, it tolerates heavy grazing and often persist under overgrazing and other forms of mismanagement. This undoubtedly accounts for much of it's popularity in certain locales.

The geographic range of Dallisgrass (a commemorative name after one of it's early advocates) is limited to the humid southeastern part of the continent barely extending north of the Deep South into Oklahoma and Arkansas. Over most of this region Dallisgrass is an occasional or, at most, a locally dominant grass. This is primarily on rangeland and forest range at lower successional states. Dallisgrass is generally a minor range species. In certain areas it is one of the more important permanent or tame perennial pasture grasses, but it is not as economically important as it once was. Dallisgrass was presented here to familarize students with a common roadside grass and one of the more obvious species of Paspalum.

This and the next three slides of dallisgrass were taken in July at peak standing crop and anthesis in Erath County, Texas.

135. Bahiagrass (Paspalum notatum)- Bahiagrass is the most important introduced (from it's native geographic range in northern through central South America) Paspalum in North America. As is the case for many of the other introduced grasses (eg. Dallisgrass, King Ranch bluestem, tall fescue) bahiagrass is tolerant of heavy defoliation. It is adapted to a wider range of soil conditions than most of the other introduced pasture grasses of the Southeast, but it's limits of adaptation are somewhat south of Dallisgrass the other major introduced Paspalum species. In addition to common bahiagrass (P. notatum var. notatum) there are several cultivars of this species all of which are established by seeding. As is the case for many, if not most, introduced grasses bahiagrass can be an economically important weed. This is often due to its greater persistence on pasture lands reseeded to the less persistent but economically superior improved cultivars of other domestic pasture species, especially of bermudagrass (Cynodon dactylon).

The importance of bahiagrass on range is that of a naturalized range species. It is especially common, and often locally important, as a forage source (though minor species overall) on forest ranges of the Southeast, both those with understories dominated by wiregrasses (Aristida spp.) and by bluestems (Andropogon or Andropogon, Bothriochloa, Schizachyrium, and Dichanthium spp.)

The example shown here is Pensacola bahiagrass (P. notatum var. saurae) which is the variety that is more frequent than common bahiagrass (P. notatum var. notatum) on disturbed areas in the western parts of the Southeast. This specimen was growing in the understorey of a longleaf pine (Pinus palustrus)- pinehill bluestem (Andropogon divergens) forest range subjected to routine prescribed burning. Fort Polk Army Post, Vernon Parish, Louisiana. July.

Common bahiagrass and the cultivars Argentine and Paragauyan are all of the taxonomic variety, notatum, that has 1) the more characteristic prostrate habit and 2) a high occurrence of apomixis. This is in contrast to Pensacola of the saurae variety and with 1) a pright habit and sexual (grain) reproduction.

Henderson County, Texas. September; anthesis stage of phenology.

139. Inflorescence of bahiagrass- The inflorescence type of the Paspalum species has traditionally been regarded as spikelike racemes (Hitchcock and Chase, 1950, p. 599) or simply as racemes (Chase, 1964, p.71).Gould and Shaw (1983, p. 223) described the Paspalum inflorescence as consisting of "one to many unilateral spikelike branches". By more recent and elaborate descriptions or classifications of the grass inflorescence the bahiagrass flower cluster would be a form of "spicate primary unilateral branches" with many or most species having "digitate spicate primary unilateral branches (Hignight, 1988, p.7).

Henderson County, Texas. September; anthesis.

141. Coarse bases- Rootcrowns and rhizomes of bahiagrass showing the coarse or rank size and structure of basal shoots of this introduced panicoid grass. Over much of the area where this exotic species naturalized it became a rank, aggressive weed that interferes with establsihment of more productive, more recently developed forage crops such as Coastal bermudagrass. Much of this weediness is due to successful propagation and invasion attributable to aggressiveness of the rhizomomes of this species.

Henderson County, Texas. October.

142. Stand of Vaseygrass (Paspalum urvillei)- This grass was introduced into the United States in the last quarter of the Nineteenth Century. Vaseygrass is closely related to Dallisgrass, but due to its more upright habit is less tolerant of heavy grazing. Guess which one of these species is more widely planted and has naturalized over a large portion of the southern states. Vaseygrass remains one of those perennial forage grass species that has "potential" (and like so many of these "potential" species most of this "potential" remains unrealized).

At present the most that can be said of Vaseygrass is that it is one of many Paspalum species in southern states (especially Texas), and it is one of the minority of these Paspalum species that was introduced (and, even more in the minority, one that did not naturalize to the extent that some like bahiagrass and Dallisgrass did). Large individual plants of Vaseygrass are sometimes encountered in the southern pine range types where they may make valuable contributions of animal diets. Vaseygrass has higher seed yields than bahiagrass and Dallisgrass, but the cespitose growth habit of Vaseygrass versus the stoloniferous bahiagrass and more prostrate growth feature of Dallisgrass insure Vaseygrass a minority status on southern grazinglands. The rank-growing, often immense clumps of Vaseygrass make it an impressive high-yielding grass, but it has never been an important forage species. For instance, none of the five complete editions of the classic text, Forages, has devoted more than incidental coverage of Vaseygrass. Like most rank species, Vaseygrass becomes unpalatable at advanced maturity (like that shown here) and especially when dormant.

This small stand of Vaseygrass was growing on the bottom side of a pond dam-- protected from grazing. Erath County, Texas. September, phenological stage ranged from early anthesis to seed-ripe but plants were at peak standing crop.

143. Shoots of Vaseygrass- These mature shoots of Vaseygrass were in full-flower. At this stage of annual plant development forage is generally of low palatability and forage quality (both nutrient content and digestibility/voluntary intake) is much less than at earlier stages of maturity. Like a fat cow (and a thin woman) it "shore is purty" at this stage.

Erath County, Texas. September, peak standing crop

144. Inflorescence of Vaseygrass- General inflorescence type of Vaseygrass is a panicle; specifically it is a panicle with alternate spicate primary unilateral branches. That was a mouthful, but it was also very precise, straight-forward, and readily interpreted. Main units off of the rachis (ie. primary branches along the stem of the panicle) are alternate (vs. opposite) and have more-or-less sessile spikelets on only one side of these individual branches so that the same resemble spikes. Or, said another way, individual spikelike inflorescences are arranged in an alternate placement in an overall flower cluster that is a panicle.

Beginning students should learn quickly that there is a rational reason why specialized crafts and their supporting bodies of knowledge are rich in terminology. The technical terms facilitate rapid learning and ready communication among practitioners. This is the case whether the skill is saddle-making, welding, ranching. or open-heart surgery. The terminology regarding the inflorescence of Vaseygrass is no more technical or formitable than those dealing with carborators, saddles, or chain saws.

Erath County, Texas. September, spikelets in anthesis.

145. Details of Vaseygrass inflorescence- Numerous spikelets were "standing in line" along one side of primary unilateral branches resembling individual spike inflorescences were supported directly on the rachis. Thus the whole or overall flower cluster was a panicle of spikelike unbranched branches each of which supported numerous spikelets each of which was borne on the branch without a pedicel (or at least not much of one).

Erath County, Texas. September, anthesis to seed-ripe stage.

146. Details of mature spikelets- Inflorescence of Vaseygrass at phenological stages approaching grain-maturity. This naturalized, agronomic, perennial grass was the associate herbaceous species on an oak woodland or savanna range that had an an understorey dominated by naturalized domestic grasses at western perimeter of Pineywoods in east Texas. Panicle and flag leaf were shown in first photograph while the second slide featured ripening spikelets. (Coverage olf this forest range vegetation was presented in Woodlands and Forests chapters, Texas Pineywoods-II and Miscellaneous.

The Paspalum inflorescence type has traditionally been regarded as a panicle. Contemporary terminology described the major units of this panicle as "unilateral spicate branches" (Gould, 1975, p. 500). These branches in P. urvillei are ascending or erect and closely spaced among themselves and arising off both sides of the central axis (or rachis) of the panicle.

.Freestone County, Texas. October; hard-dough to grain-shatter stages of phenology (and all on the same branch of a panicle).

In fact, the introduction of buffelgrass has been one of the major advances in ranching in the area to which it is adapted. It is typically managed as range with no inputs except seeding and proper grazing management. It will respond favorable— though likely not economically— to fertilization and irrigation. This pasture was an abused South Texas Plains range deteriorated to the mixed brush state of mesquite, huisache, pricklypear, and the full array of the brush community. The depleted range was root-plowed in alternate strips 200-400 yards wide between strips of untreated brush (left for white-tail deer habitat) of the same width. Then the plowed lands were reseeded to common buffelgrass in February. Cattle were turned in on the reseeded range in July (five months later!) and grazed until the next February (grazed for seven months) and then rested until May (three months later) when this picture was taken. Chaparrosa Ranch, Zavala County, Texas. No FRES or Kuchler (not native). Mini-editorial: this kind of grazing land and that of stable Kentucky bluegrass, weeping lovegrass, Lehmann lovegrass, etc. is as deserving (or undeserving) of SRM cover type designation as is tall fescue!

151. Panicles of buffelgrass- These buffelgrass panicles were in anthesis with individual spikelets, as well as anthers, clearly visible. Buffelgrass spikelets bear bristles that are united at their bases. This arrangement of partially united bristles at base of a spikelet is an involucre. The Pennisetum involucre somewhat resembles the more completely fused or coalescing bristles that form the spiny bur of the Cenchrus species. This situation prompted some agrostologists to transfer buffelgrass to the the latter genus (ie. Cenchrus cilaris). The author of the current publication rejected this imaginative move as did the latest definitive, comprehensive treatment of Pennisetum and Cenchrus species (Wipff in Barkworth et al, 2003, ps. 525).

Note the flag leaf, the leaf immediately subtending (below) the inflorescence, in the photograph of the solitary panicle. Pima County, Arizona. June.

152. Range reseeded to King Ranch bluestem- A brush-infested old field was treated by mechanical brush control methods and reseeded to King Ranch bluestem (Andropogon ischaemum= Bothriochloa ischaemum, often B. ischaemum var. songarica) during the hey-day of the Great Plains Conservation Program. Following mechanical brush control (primarily root plowing to kill mesquite) this severely depleted land was drilled to King Ranch or yellow bluestem. KR bluestem was one of the first of several species of Old World bluestems introduced into North America. The common name, King Ranch bluestem, originated because this species was first introduced to the famous King Ranch from Asia.

The Old World bluestems (so called because they are native species in Africa, the Middle East such as Russia, and southern Asia like India) form a complex of such genera as Bothriochloa, Capillipedium, and Dichanthium (Voigt and Sharp in Barnes et al., 1995, p. 398). KR bluestem has been the most important of the several introductions from this complex owing perhaps to its early use-- and promotion-- by the famous King Ranch. "KR" has also proved to be extremely prolific due to its ease of establishment, winterhardiness, and drought tolerance. In the latter two characteristics B. ischaemum is clearly inferior to the bluestems native to locales where KR was introduced, but KR's capacity for rapid and sustained reproduction (both prolific seed production and abundant tiller formation from a compressed, grazing tolerant, basal portion) give this introduced grass a clear advantage over any of the native decreasers (and most of the increasers).

This is so much the case that KR bluestem which thoroughly naturalized from the early introductions of this species become a serious weed in various locations. This is a major problem, some allege the major problem, with many of the introduced forage species like Johnsongrass, bahiagrass, and common bermudagrass. The first releases of such species proved to be much less productive and nutritious than later cultivars of these species, but the original releases were superior competitors for any number of reasons and became noxious plants. For example, bahiagrass became a weed after it proved to be more competitive and persistent than superior cultivars of bermudagrass like Coastal and the various Tiftons. Johnsongrass requires no explanation beyond it's name though it remains a superior forage species under certain livestock production systems. KR bluestem was so widely seeded along highway rights-of-way and on severely eroding old fields (like that shown above) that it's prolific seed production resulted in it becoming widely established where it was not wanted (eg. bluestem ranges in the Texas Prairies and and Cross Timbers area, especially with overgrazing).

By the 1980s, however, improved cultivars of B. ischaemum (now designated either yellow bluestem or the generic Old World bluestem to distinguish it from the earlier strains called King Ranch) became available that were superior for such features as nutritive value, disease resistance, tolerance to iron-deficient soils, and winter-hardiness. These newer releases included Plains, WW Iron Master, and WW Spar. During the severe drought of the late 1990s and early 2000s the newer cultivars of B. ischaemum were some of the most dependable forage sources in the Texas and Oklahoma Panhandles area. Substantial acreages of highly erodible farmlands in the Southern High Plains region were reseeded to the improved strains of King Ranch bluestem under the Conservation Reserve Program.This was one of the "success stories" in which conservation increased chances of economic survival in times of rough weather and tough financial times, the very combination of conditions that led to the second wave of conservation in the 1930s during which federal agencies like the Agricultural Conservation and Stabilization Service, Soil Conservation Service, and Bureau of Land Management were born.

There have been several other Old World bluestem species introduced into the southern states, especially Texas, and the lower Midwest. Some of the other species to receive the early attention of grass breeders included Angleton bluestem (Andropogon aristatus= Dichanthium aristatum), Australian bluestem (A. intermedius= Bothriochloa intermedia), and Kleberg bluestem (A. annulatus= D. annulatum). Readers will note from the latter species that the King Ranch heritage has figured prominently in the breeding of Old World bluestems. Caucasian bluestem (B. caucasia) was found to be the most winter-hardy of the Old World species, but to this author's knowledge there have been no selected cultivars of this species released.

B. ischaemum apparently remains the only species of Old World bluestems for which there are on-going selection programs. Only time will tell if the superiority of new B. ischaemum cultivars for desirable traits overcomes the weedy tendency of this species, or if the new cultivars are less weedy and therefore less persistent under the harsh environments into which they were seeded. Voigt and Sharp in Barnes et al.(1995, p. 399) remarked that "[t]he weedy nature of the Old World bluestems attests to their aggressive establishment ability".

153. King Ranch bluestem on a reseeded field- Former cropland (cotton, sorghum, etc.) that had been abandoned and then "went-back" to brush (especially mesquite), annual bromegrass species, and Texas wintergrass was improved for forage production and soil conservation by reseeding to King Ranch bluestem, sideoats grama, and little bluestem following mechanical brush control. Following this combination of improvement practices the seeded grassland became (for all practical purposes) single species stands of King Ranch bluestem on lower portions (shown here) and sideoats grama on the higher, better drained parts of the former field.

Only occasional clumps of little bluestem had established from the reseeding operation. More plants of silver bluestem and white tridens had "volunteered" (become established through limited secondary plant succession). A few mesquites had reinvaded the cleared, seeded land.

Commanche County, Texas. October.

154. Dense stand of King Ranch bluestem- Density of shoots (almost every one of them sexually reproductive) typical of planted and naturalized stands of King Ranch bluestem. Such examples of plant life are technically local populations (ie. single species stands) and not communities, not vegetation, per se. This photograph illustrated why naturalized KR bluestem has long been a range weed under certain conditions. Such instances also showed, however, why this species was chosen at a time when getting any cover on bare soil was the "first and highest priority" of conservation. Even today this species is just as important (and for just the same reason) in certain areas.

A current example of this is thousands of acres of range in Texas' Edwards Plateau that continues to be overgrazed-- often to the degree of bare soil ("Believe It or Not")-- after decades of research and demonstations proved beyond doubt that such is mismanagement of both financial as well as natural resources. Such stocking rates, and consequent degrees of use, extend input of resources (of all kinds) and animal turnoff far beyond stage two of the economic production function (the rational, relevant stage of production that maximizes profit). Such rates of resource use result in temporary harvest in excess of sustained yield. These are market failures that deplete resources with negative spillover impacts on society. Such excessive rates (those that exceed sustained yield) can be achieved (but never sustained) only by subsidies such as supplemental feeding, higher death rates, and lower animal performance in actual terms. Under such instances of mismanagement due to forces like greed, vanity (eg. high numbers of animals for purposes of "bragging rights"), and stupidity (given today's level of education such hubris no longer qualifies as ignorance) King Ranch bluestem remains a Godsend and not a noxious plant.

Mills County, Texas. October.

155. Pasture of "Plains Bluestem"- A cow-calf herd was grazing this pasture of "Plains Bluestem". Old World "Plains Bluestem" is one of 600 accessions or cultivars tested in the southern Great Plains. Plains was released by the Oklahoma Agricultural Experiment Station in cooperation with the Agricultural Research Service. Bell and Caudle (www.rw.ttu.edu/newsletter/) summarized the "Plains" cultivar: "Plains bluestem is a blend of about 30 similar looking bluestem accessions collected from six different countries [Pakistan, Iran, Iraq, India, Turkey, and Afghanistan]. It was released commercially in 1972 and has been readily accepted and widely used from Missouri to Texas. It has become the standard by which all other Old World bluestems are evaluated". The white flowers in the field are those of field bindweed (Convolvulus arevensis). This Eurasian species is one of the world's worst weeds having a nearly global distribution and being extremely difficult and expensive to control. So far "Plains Bluestem" in this pasture has outcompeted the aggressive bindweed, even under obvious close grazing.

Carson County, Texas. June.

156. "Plains Bluestem" (Andropogon ischaemum= Bothriochloa ischaemum var. ischaemum)- A plant of "Plains Bluestem" in the pasture shown in the preceding photograph. The cultivars "Plains", "El Kan", "Ganada", "WW-Iron Master", and "WW-Spar" are of the taxonomic variety B. ischaemum var. ischaemum. whereas the actual or real "King Ranch Bluestem" is B. ischaemum var. songarica (Alderson and Sharp, 1993; www.forages.css.orst.edu/).

Carson County, Texas. June.

157. Sward of King Ranch bluestem- With exception of the large and/or ornamental grasses none of the Gramineae members are "photogenic". The absolute best way to examine grass species is to view live or preserved specimens. The best way to artificially portray grass features for overall clarity and everything-in-focus views for purposes of identification, morphological examination, etc. is line drawings. Photographs--even color (perhaps expecially color) photographs-- are no surrogate and a poor substitute for specimens or professional drawings, but the medium does dictate it's own terms even for those who did their best to keep the medium from hijacking the message.

One of the least "photogenic" of the grasses is the aesthetically ugly King Ranch bluestem. This rhizomatous, stoloniferous, generally tufted grass has it's basal leaves in a mat-like morphological form that hardly exceeds, say, three inches in height, but it quickly bolts it's, flowering shoots to heights of two feet (or more) from which it dispenses many small grains.This habit renders KR bluestem extremely tolerant of close defoliation and ideally adapted for survival under abusive defoliation. This same set of characteristics make it one of the most difficult of all range plants to portray in photographs. It is neigh on to impossible to get adequte depth of field so that inflorescences and basal portions of shoots are simultaneously in focus. This is true "in spades" when using slow film to accurately reflect color and get quality photographs that are relatively free of graininess.

This and the next six slides were presented in an attempt to provide the viewer with a reasonable representation of King Ranch bluestem. Erath County, Texas. September.

158. King Ranch bluestem- One plant of King Ranch bluestem was in the center of this photograph (edge of another can be seen in upper right hand corner). Individual plants of A. ischaemum (= B. ischaemum) have rhizomes and, frequently, stolons depending on such factors as genotype and intensity of defoliation. Emperical evidence (common observations and general descriptions in some of the manuals) indicated that these two forms of extravaginated shoots are more common under close grazing or mowing. There would seem to be considerable phenotypic variation in this species.

Erath County, Texas. September.

159. Individual plant of King Ranch bluestem- A solitary genetic individual of A. ischaemum at peak of anthesis was shown here. It was just explained (immediately above) that King Ranch bluestem reproduces asexually by stolons and.or rhizomes depending on genetics and phenotypic factors. Under repeated heavy mowing, grazing, traffic, and so on this tufted species takes on a mat-like appearance as shown here. This photograph illustrated (as best habit can) why King Ranch bluestem is often a aggressive weed much like Johnsongrass or quackgrass. Numerous flowering shoots had been produced by this individual and it was spreading by both stolons and rhizomes. This specimen grew on university grounds subjected to repeated, long-term, too-close mowing (ie. mechanical overgrazing) such that even common bermudagrass had been eliminated from the former lawn.

In many areas of Texas and southern Oklahoma such a regimen of overmowing produces a depletion grassland community composed of crabgrass (Digitaria spp.) and goosegrass (Eleusine indica)-- often accompanied by common grassbur (Cenchrus incertus) -- that takes over the former perennial sod. Under this combination of abiotic and biotic variables King Ranch bluestem is one of the few perennial grasses that can survive (ie. outcompete the therophytes). This is when the aggressive weediness of KR bluestem is a blessing. At very least, "KR" provides more protection and conservation of the basic natural resources of soil and water than the weedy annuals.

Erath County, Texas. September.

160. Shoot of King Ranch blusetem- An individual shoot of King Ranch bluestem at full-flower showed ("explained") the habit or basic morphological form of this tufted species. This shoot was from a plant that had both rhizomes and stolons and that had been subjected to overmowing for years (at least, this perennial grew on a yard that had been mowed too closely and too frequently for decades).

The first tuft of small leaves on the shoot (on right-side) were the first node-internode unit (the phytomer) above ground level. That part of the shoot below this phytomer with adventituous roots was the most basal part growing aboveground and it was prostrate on land surface. The shoot region of the next four phytomers had first a decumbent and then an ascending form such that most of the bluestem plant was below level of a rotary mower (or a cow's mouth) until the proverbial last moment when the terminal phytomer with the inflorescence "shot up overnight" (literally within four days of heavy rainfall). Those viewers who take a close look can see that almost two-thirds of total shoot length was in the terminal, flower-bearing phytomer.Furthermore, the internodes of the three phytomers below the terminal one did not elongate much until onset of the flowering phenological stage.

Habit of this species furnished students with an outstanding illustration of grazing adaptation by morphological modification, the net result of natural selection for heavy defoliation.

161. Shoot apices of King Ranch bluestem- Flowering shoots of a "culmless", "non-jointing", or "short-shoot" grass species. King Ranch bluestem is one of the species that fits readily into the cul group of grasses that have more fertile than infertile apices and with apical meristem kept unelevated until late in annual life cycle. Readers were referred to Heady and Child (1994, ps. 23-25). Concepts regarding shoot morphology of grasses was discussed very well in the SRM reference edited by Bedunah Sosebee (1995, ps. 30-39 passim).

Erath County, Texas. September.

162. Flower cluster of King Ranch bluestem- The inflorescence of the bluestem and panicgrass groups (the panicoid grasses) has traditionally been interpreted as a panicle of one to several racemes or, same thing essentially, a panicle of racemose branches. More precisely, the inflorescence type seen here was a panicle of subdigitate racemose branches. These racemose branches were opposite (more clear in succeeding slide).

Erath County, Texas. September

163. King Ranch bluestem at anthesis- The opposite racemose branches of the panicle of A. ischaemum. Anthers (filaments and anthers) and stigmas were fully exerted at "full-flower". King Ranch bluestem produces large numbers of caryopses. This is in addition to asexual reproduction by stolons and rhizomes. KR bluestem is a "survivor".

Erath County, Texas. September.

164. Johnsongrass (Sorghum halepense)- This introduced member of the bluestem tribe (Andropogoneae) is probably the most widely distributed naturalized warm-season, perennial grass in North America. Johnsongrass is a native of the Mediterranean Region of North Africa and the adjoining areas of Eurasia. It was introduced into the United States early in the Nineteenth Century and quickly naturalized. Johnsongrass was introduced into many other parts of the world-- both purposely and accidentially-- where it also naturalized to the point of weediness until today it generally listed as one of the "Worlds Ten Worst Weeds" by the Food and Agricultural Organization of the United Nations. Sorghum halepense is clearly one of the Dr. Jekyll and Mr. Hyde agricultural plants. It is one of the most productive of the temperate forage grasses with the ability to produce comparatively high yields under both extensive and intensive management (ie. it responds well to fertilization and careful cropping management yet is amazingly adapted to survival and production of high herbage yields under harsh environments). Johnsongrass is widely distributed across much of the Western Range Region and adjacent areas. It has been found on the Northern Great Plains of North Dakota within one county of the Canadian-US border and south into favorable sites on the Sonoran Desert of Mexico and from California to parts of New England. From the perspective of agronomic and horticultural practice Johnsongrass is a weed throughout this geographic range, but it also produces nutritious feed as forage for grazers and grains for native birds.

Like all Sorghum species (including S. bicolor or S. vulgare, the grain and silage sorghum), Johnsongrass is a poisonous plant. Readers should consult all the standard works cited earlier such as Kingsbury (1964), Cheeke and Shull (1985), and Burrows and Tyrl (2001). The latter discussed in encyclopedic detail how the Sorghum species caused animal losses due to any of four general poisonous principles: hydrocyanic (= prussic) acid poisoning, nitrate toxicity, primary photosensitization, and ataxia/cystitis/ tetratogensis (Burrows and Tyrl, 2001, ps. 929-945).

Although Johnsongrass was one of the first warm-season perennial grasses introduced into North America (and remains one of the best adapted and productive of these introductions) the aggressiveness, weediness, and toxic characteristics of this species greatly limits its use as a domestic forage crop. Instead the major beneficial aspect of Johnsongrass is as a naturalized species on range where under proper management it is a minor species among numerous native range plants. It is a relatively highly productive range species on the localized microsites of the "empty niche" it found in the New World. Under these conditions-- including proper grazing management-- Johnsongrass cannot out-compete the native panioid and eragrostoid grasses nor grow in such populations as to be a major poisonous plant. Instead it contributes to biological diversity as well as furnishing protection for soil and providing feed and cover for range animals.

The example shown here was growing in a pecan (Carya illinoisis) orchard as a companion, cover, and forage crop illustrating the various agronomic and horticultural uses of this crop (and the weed feature in the absence of proper management). Erath County, Texas. July.

167. More spikelets of Johnsongrass in bloom- Some more views of anthesis in Sorghum halepense were presented as examples of sexual reproduction in grass. It is important to bear in mind that applied practice such as management of pastures and ranges is grounded in sciences as fundamental as Botany or, even more fundmental, general Biology.

Ottawa County, Oklahoma. July.

168. Johnsongrass grains- Spikelets filled with mature caryopses in Johnsongrass. The persistence of this introduced panicoid grass (as both a range or pasture forage species and as the worst kind of a weed) is due in large part to the fact that it progagates readily both asexually by rhizomes and sexually by abundant grain production. The short rhizomes allow aggregation or "thickening of clumps" and invasion of smaller spaces among adult plants by new sister plants (clones) while shattered grains permit wider dispersal of new daughter plants (ie. new genotypes). This combination of asexual-sexual reproduction enables on-going evolution at the same time that immediate colonization takes place through phenotypic plasticity and growth of established mature individual genotypes. Johnsongrass does this -- and under the severe disturbance of land tillage-- better than almost any perennial grass native to the geographic range of the naturalized Johnsongrass. No wonder this ruderal species is such a survivor.

Erath County, Texas. October.

169. Rhizomes of Johnsongrass- Rhizomes are belowground extravaginated shoots. "Rootstock" is a confusing synonym for rhizome. (Rhizomes are modified shoots or stems and not roots at all, but the subterranean location and superficial resemblance of these shoots to roots accounts for this old and traditional synonymy.). Daughter shoots come off of the rhizomes by piercing the enveloping sheath such that each rhizome is first a new daughter shoot and subsequently a mother or parental shoot. This is an example of a clonal organism in which each new daughter plant is a clonal unit (= a module) of the same genetic individual. Modular organism is synonymous with clonal organism. The repeating units, the modules, are ramets while the entire genetic individual (the entire plant having the same genotype) is the genet.

Asexual (= vegetative) reproduction by rhizomes that are protected from many forms of defoliation (eg. grazing, fire, hail) is one of the factors responsible for the aggressive growth, persistence, and general weedy nature of Johnsongrass. As propaguules or disseeminules, rhizomes are dispersed less readily than some forms of diaspores (eg. many seeds and fruits), but they are very effective in insuring survival of S. halepense. Like other modified stems (tubers, corms, bulbs, etc.) rhizomes also have a food storage function. Various proportions of sucrose and glucose are stored in these plump rootstocks. The rhizomes in the pile shown in the first slide were unearthed in mid-spring (May) and have a "drawn-down" appearance following use of much of the storedd food over the winter dormant season. The fat rhizomes shown in the second slide were dug druing mid-summer during the replinishment or storage phase in preparation for dormancy and emergence at onset of the next warm, growing season. Erath County, Texas (May) and Ottawa County, Oklahoma (July), respectively.

170. Field of Johnsongrass- Johnsongrass' potential for high production was captured in this photograph taken at boot- to early inflorescence-emergence stage in north Texas (May). This is typically the phenological stage of most grasses at which to harvest forage standing crop for maximum yield of nutrients (energy as Total Digestible Nutrients or Digestible/Metabolizable Energy, crude protein, most minerals) on a per plant or per acre basis. This is not the most palatable or the most nutritious or nutrient-dense stage. That occurs at less mature (less phenologically advanced) stages of plant growth, but harvest at boot stage generally results in highest yield of most nutrients and nutrient properties with forage still having acceptable palatability.

This is most obvious in harvesting as hay, silage, or greenchop, but the principle also obtains for harvest by grazing. Johnsongrass is extremely sensitive to defoliation, especially close and/or repeated cutting or grazing. This has traditionally been explained on the basis that frequent and close cropping prevents both food storage in and total production of numbers and size of rhizomes. (See for eg. the first edition of Forages, 1951, ps. 365-366). Thus Johnsongrass plants arising from new seedings should be allowed to reach full maturity (seed-set stage) before being grazed the first season of growth. Even with established stands plants should be allowed to develop to the seed-set stage prior to end of growing season. This grazing or mowing safeguard assures adequate food storage in rhizomes and roots which is essential for plant survival during dormancy and for stand persistence. Practical experience and field research have shown that the best way to kill Johnsongrass on land where it is a weed is to graze it out by continuous, or at least frequent, close grazing or by repeated low mowing.

As a field crop Johnsongrass requires careful grazing or mowing management, a fact belied by the aggressive, rank growth and persistence of Johnsongrass under conditions where it is a weed. This is less of a problem on range where Johnsongrass is often less palatable than other associated and closely related grasses (eg. other panicoid grasses like bluestem species or Indiangrass on tallgrass prairie) and where proper use is usually lighter than on introduced pastures, including single-species stands of Johnsongrass raised as an agronomic crop.

Erath County, Texas. May.

172. A study in Johnsongrass (and of color)- This field of Johnsongrass was at peak standing crop at mid-seedset stage. The two slides (Kodachrome 64) were taken just seconds apart at approximately 1530 hours in mid-July. The first of these two photographs (upper one) was taken at full sun while the second (lower) photograph was taken after a large cumulus cloud passed between the sun and the subject resulting in loss of one f-stop. Exact f-stops and shutter speeds are irrelevant, but the difference between a full sun shot and a cloudy or overcast one made this readily observable difference. While the difference was not "day and night" it was real and important. The upper slide captured true color of Johnsongrass shoots as seen and interpreted by the human eye. Like sorghum (Sorghum bicolor= S. vulgare) and dent corn (Zea mays indentata) at seedset stage of maturity (phenology) Johnsongrass culms and, especially, leaves have a slight yellowish cast or hue (a yellowish-green). By contrast, presence of clouds (a photograph taken under overcast sky) presented the color of Johnsongrass shoots as more of a bluish-green hue that was more the color of bluestems than of sorghums.

Some plant photographers prefer a shaded, overcast, early morning or evening atmosphere so as to "soften" tones of colors, especially of petals having "lighter" or "paler" tones of blues, pinks, and pastels in general. Under some light conditions and with certain plant species this is desirable or even necessary to prevent "washing out" of pastels. For example, bright or full light will "blanch" or "bleach out" light pink veins or other pastel-colored pigments in the corolla of some species. This results in incorrect color portrayal and presents petal pigmentation as white rather than a pinkish tinge perceived by the eye under paler light or under indirect light. In this photographers experience, however, with these exceptions and under most situations full-un, direct, natural light provides the closest color on film to what a non-colorblind human eye has seen.

Furthermore, Kodachrome captures most--not all--coloration of rangeland and forest plants much more accurately (based on human eye perception of color) than do faster films like Ektachrome or Fujichrome. This is most true for yellows and reds. Plants whose corollas reflect pale pastels only in indirect light or overcast atmospheres should be photographed at those times (ie. when light is "less harsh" as under overcast sky or in early morning, late afternoon or evening; sometimes just before sundown or at sunup for example).

One problem that requires the photographer's careful attention when shooting in full sun or direct mid-day light is presence of shadows. This is probably the major problem when trying to get "half-way decent" photographs with good depth-of-field in dense forests. Under these situations overcast skies are sometimes superior to bright, direct light because light is diffused and does not result in shadows as on tree trunks or in as much shaded ground. Such light conditions are drastically different from those on grasslands, tundra, alpine, and most savannahs where full sun is usually superior for most landscape and plant pictures. Again though, photographers should be aware of shadows being formed. For instance, in the Johnsongrass stand shown here there were more shadows under full afternoon sunshine. The photographer got around this potential problem by 1) positioning his body and the camera at angles that eliminated or, at least, reduced shaded areas or patches and 2) taking an extra photograph or two to increase likelihood of getting a closer-to-perfect shot.

Ottawa County, Oklahoma. July; early seedset (soft-dough) stage.

Various nodulated legumes, the papilionaceous legumes (Papilionoideae subfamily of Leguminosae), that are primarily agronomic species have been introduced to be managed either 1) extensively as range plants or 2) intensively as field crops for tame pasture and/or hay production.The number of publications by the various Agricultural Experiment Stations and Extension Services, Forest Service, text and reference books, private organizations such as seed companies, etc. is staggering and beyond any comprehensive cataloguing. A good introductory source for beginners and oldsters alike is the Legumes section of Pasture and Range Plants by Phillips Petroleum Company (1963).This delightful series (now bound as one book) has proved invaluable to several generations of pasture-and rangemen. Perhaps the most longstanding authoritative text with the clearest summaries of major forage legumes between one cover is the classic text, Forages, by Iowa State Press and under numerous authorships. Another standard text of equal persistence, but of less consistency in revision, is Principles of Field Crop Production (Martin et al., 1976). Though woefully "out-of-date", as of this writing Martin et al. (1976) remains the only comprehensive text that covers actual culture or husbandry (ie. a farming manual) for all major field crops in North America, including those used as forage crops. Forage Plants and Their Culture (Piper, 1939) is "ancient" by academic dating standards, but as a fundamental text-reference for many of the introduced species that have naturalized (eg. annual lespedezas, sweet clovers, Johnsongrass, vetches) it is still one of the standards. (Some of the "antiquated" works are more useful for the extensive-input approach of Range Managaement than are the more current texts.) The American Society of Agronomy, Crop Science Society of America, and Soil Science Society of America have published jointly a number of monographs on individual species and groups of species of legumes over the years. Clover Science and Technology edited by Taylor (1985) is an example.

Probably the best general source and detailed botanical descriptions of all domesticated forage species is the dated, but still only encyclopedic coverage, Manual of Cultivated Plants (Bailey, 1949). Descriptions, nomenclature, and range of naturalized species have been shown by most of the various flora or manuals.

The major management "challenge" associated with legumes, especially the introduced species that have been selected for high yields and high nutritive value (a major component of which is palatability), is their persistence or maintenance in either diverse range plant communities, mixed-species tame pastures, or single-species stands. Simply stated, legumes (especially domestic species), are so much more palatable than their "neighbors" that grazers tend to preferentially select and graze them out. Legumes are more nutritious (eg. higher in protein, energy, and minerals like calcium), hence more palatable, and they become ice cream species. Legumes are even more attractive to certain species of phytagophus insects. Most legume species are less highly adapted to defoliation than the grasses are. Under roughly equal degrees of use, competitive advantage shifts to the more grazing-adapted grasses which have already been the beneficiaries of nitrogen-fixation by the host legumes and their symbiotic bacteria. Most of the forage legumes have more upright morphological forms with their apical meristems more exposed to agents of defoliation (fire, hail, and wind as well as herbivores). Some species like white clover (Trifolium repens) are low-growing and reproduce well vegetatively, but most legumes are nowhere in the same league as grasses and grass-like plants when it comes to grazing adaptations and tolerance of defoliation.

Discovery of which Rhizobium species are specific to the various legume species and development of technology for innoculation of legume seed were other factors limiting introduction and establishment of forage legumes.

Even when legumes can be established on range and mixed pastures their maintenance remains a big problem. Legumes are much easier to manage and maintain as harvested forages, usually hay. This fact explains the presence of alfalfa (Medicago sativa), one of the most water-demanding forages, as a major field crop in many areas of the Western Range, even in the Intermountain West.

"Clover" is a word of Middle English origin as are numerous terms used in Agriculture such as "corn". Several Middle English words such as "clover", "corn", or "cattle" are generic terms that have both 1) retained their original general meaning and 2) taken on newer and more specific meanings. "Clover" has been applied as a common name to numerous plant taxa, usually, but not always exclusively, to members of the legume family. Within the Leguminosae, "clover" traditionally was ascribed to several genera. Such use of "clover" was then modified with adjectives preceding the main noun of "clover". Thus there became "bur-clover", "sweet clover", "Korean clover", "bush clover", etc. The other use of "clover" in reference to legumes is to species within the genus Trifolium in which case adjectives in the common name refer to species rather than genera as in previous cases like "sweet clover" or "bur-clover". To avoid any more confusion than already exist, "clovers" in Trifolium have often been designated the "true clovers" in which case adjectives describing "clover" indicate species (eg. white clover, crimson clover, red clover, alsike clover, rose clover, rabbitfoot clover). Trifolium is from the Latin for three leaves in reference to the compound leaf that typically consist of three leaflets. (Incidentially, it is the trifoliate clover leaf and not the "four-leaf clover" of Landy Luck or the 4-H Club that is the shamrock, national emblem of Ireland. The shamrock is, of course, one of the Christian symbols for the Trinity. The traditional account is that the King of Ireland could not grasp the concept of the triune God whereupon Saint Patrick plucked the tripartite clover leaf to show how God could exist as three persons in one.)

The following "clovers"-- at both generic (taxonomic adjective for genus) and specific (taxonomic adjective for species) levels-- are some of the more important ones as field (ie.agronomic) and/or range legumes.

176. White clover (Trifolium repens)- This may well be the most widespread of the true clovers (Trifolium species), at least as a local dominant. White clover is native to Eurasia but has naturalized in the wake of human migration, including to the New World. The American Indian referred to this species as the "white man's footprint" because with it's spreading habit and bright white inflorescences it was one of the most conspicuous alien species brought with the mass human movement that began with the voyages of Christopher Columbus. Unlike most of the naturalized alien species which became plant or animal pests white clover had what is generally regarded as a beneficial impact as result of it's invasion.

The specific epithet repens is Latin for "prostrate and rooting" and refers to the stoloniferous and adventituous rooting habit of this perennial species. (Observe the carpet-like sward of the stand shown above.) T. repens has remarkable phenotypic plasticity (often a characteristic of domesticated species as in domestic swine for instance) and genetic polymorphism that enables this species to adapt through natural selection to new environments and micro-habitats very quickly (see for eg. Begon et al., 1990, ps. 38-39). White clover is, of course, one of the classic examples of a clonal organism. Plus it is a prolific seed-producer. This combination of features, along with the mutualism of legume and bacterial nitrogen fixation, has allowed white clover to naturalize over much of North America, including more mesic range areas.

Forage quality of white clover is exceptional, but herbage yield is usually lower than that of associated grass and the weedier forb species. It is preferred feed by about all herbivores. Ladino white clover is a cultivar of the taxonomic variety T. repens var. giganteum. It's use is probbly best reserved for tame pastures. Viewers are reminded of the texts and references suggested above and of the numerous publications by their state Agricultural Experiment Stations and Extension Services.

Ottawa County, Oklahoma. June.

178. Shoots of white clover- Detail of stem, leaf, and inflorescence of common white clover. Note leaflets of the trifoliate, compound leaf. Erath County, Texas. April.

179. Stolon of white clover- The"runner" of common white clover is a ramet or clone of this highly stoloniferous species. Another example of a modular plant. Ottawa County, Oklahoma. June.

181 Crimson clover (Trifolium incarnatum)- Most of the introduced true clovers (domesticated Trifolium species) are better adapted to northern parts of North America because they tend to be the more mesophytic of the mesophytes and to have lower yields under regimes of higher temperatures and drier soils. Crimson clover is less extreme in this regard and is one of the Trifoilum species better able to tolerate conditions more characteristic of the American South. Crimson clover is a winter annual and it can utilize the generally more moist soil conditions that occurr in winter and spring months when evapotranspiration is lower and, in the spring, when rainfall is greater. Crimson clover is not common in southeast or southcentral portions of the continent, but it is less uncommon there than are many species. Crimson clover does relatively well in Arkansas, Oklahoma, and Texas if carefully managed. It is one of the best introduced legumes for winter forage production in southeastern North America. Crimson clover is one of the most strikingly colorful of the cultivated legumes and is sometimes seeded along highway rights-of-way as part of beautification programs.

Erath County, Texas. April.

184. Rose clover (Trifolium hirtum)- This attractive member of the true clover genus is another winter annual that is well- adapted to the South for the same reasons as explained for crimson clover immediately above. Rose clover has generally been somewhat more productive than crimson clover under growing conditions and habitats of more limited soil moisture. Rose clover was one of several clovers introduced into California to take advantage of the generally high winter moisture regime in the Mediterranean climate of the Coast Range and Central Valley portions. Growers shifted from rose clover to subterranean clover as the latter proved to be more productive and persistent as a component of improved California annual grassland.

Erath County, Texas. April.

185. Hop clover, yellow hop, or field hop (Triflolium agrarium)- Here was presented yet another cool-season annual true clover introduced from Eurasia. Yellow hop has so thoroughly naturalized over much of the southern part of the continent and along the Pacific Coast that many farmers and ranchers assume it is a native species. Yield of this and the species often called little hop (T. dubium) are low in comparison to most other true clovers, but these self-seeding annuals are sure yielders because they complete their life cycle during spring to early summer when seasonal precipitation is usually greatest.

The hop clovers typically grow best on poorer (shallower, rockier, acidic) soils where competition with larger, coarser perennials is less intense. They do frequently thrive on richer, deeper soils (including bottomlands) when density of competiting species has been reduced (as following drought for example). Yellow hop is often extremely abundant in eastern and central areas (eg. Ozark Mountains as for the speciment shown here) on "go-back land" (old-fields) after the sere has progressed beyond the pioneer stage dominated by large annual composites, crabgrass, and other weeds.

Hop clovers (and the other introduced annual legumes) that have completely naturalized could be interpreted as occupying "empty" ecological niches (ie. ones not filled by native plants). The niches now filled by these exotic species have-- according to the philosophical or pictorial concept of niche-- time dimensions or time "volumes" that are different from native species. The time dimension of the hop clover niche(s) obviously encompasses part of the cool-season.

Ottawa County, Oklahoma. June.

186. Red clover (Trifolium pratense)- Red clover is probably second only to alfalfa as a hay species capable of consistently producing high yields of high-quality hay. This Eurasian species has been cultivated as a major hay crop in northeastern North America, the Hay and Dairy Agricultural Region, for upwards of two centuries where it was used as roughage for dairy cattle especially in winter. Red clover was brought westward along the ever-advancing frontier lines and became an important field crop in the Great Lakes Region. With careful management red clover can produce superior hay crops as far south as northern Oklahoma. Under good agronomic practice stands of this biennial or short-lived perennial legume will last indefinitely. The author hauled hay from a field of red clover in northeastern Oklahoma that has remained in production for upwards of three decades.

Red clover is primarily a hay and not a pasture legume. It is difficult, but by no means impossible, to maintain red clover in grass-clover pastures because animals selectively graze the clover. The red clover is this photograph was growing on bottomland used as a hay field in summer and a permanent pasture in fall and winter. The red clover grew with tall fescue as the predominant species and with appreciable proportions of Johnsongrass, common bermudagrass, and crabgrass. Red clover held it's own, but had to be reseeded the spring following a severe summer drought that apparently led to subsequent winter kill (none of the associated grass species were affected).

Ottawa County, Oklahoma. June.

187. Red clover in the Ozarks-A particularily prolific red clover plant at full bloom at edge of an oak-hickory forest in the Springfield Plateau section of the Ozark Plateau. Red clover has naturalized in this region to the degree that in some years it grows and blooms prolifically whereas in other (typically drier, warmer) years plants are considerably smaller. Obviously cultural practices like liming and fertilization are a major factor in this regard. Consideration was laid on naturally occurring red clover that had naturalized as in this instance and not on planted field crops. In this western part of the Ozarks red clover is widely distributed and, as it is self-propagating (initially self-seeded), it persists indefinitely. Interestingly, red clover will appear on clearcuts or appear "suddenly" on go-back lands (old-fields) that have been heavily grazed following abandonment of farming. Under such conditions red clover is not a pioneer or early colonizing species, but it does become established locally early in the sere and persist well into the progression of plant succession if not grazed out.

Ottawa County, Oklahoma. May: late vernal aspect, full bloom stage.

Forage legumes on range are one of those things which “holds out promise” or “has lots of potential”. Range legumes have yet to materialize as forage species other than rarely as in the case of subterranean clover on the California annual type. In addition to sainfoin, research has been conducted attempting to use Cicer milkvetch (Astragalus cicer) and various creeping and rhizomatous Siberian alfalfas (Medicago falcata) as introduced range legumes (See Heinrichs in Campbell and Herbel, 1975, ps. 50-61). Determination of the specific species of Rhizobium bacteria with which to inoculate the specific legume species has been a major problem. Culture of subterranean clover on California range and pasture was unsuccessful until the inoculation problems were solved.

191. Bur-clover, or burclover, (Medicago hispida= M. polymorpha= M. denticulata)- There are numerous Medicago species known variously as burclover (with or without adjectives) or medic. These are distinct from the Medicago species known as alfalfa that were covered above. Distinction among the various Medicago species known as burclover or medic can be somewhat tedious (and tedious). Bailey (1949, p. 582) listed four Medicago species in addition to alfalfa for the United States and Canada, but Correll and Johnston (1979, p. 805-806) described two other Medicago species for Texas that were different from those of Bailey, including the common button medic or button clover (M. orbicularis).

All of these burclovers, button clovers, or medics were introduced from the Medterranean Region and are self-seeding winter annuals (Martin et al., 1976, p. 768). They have naturalized to varying degrees across much of the South, especially the historic Cotton Belt, and the Pacific Slope where cool and usually moist winters allowed these aliens to find apparently "empty ecological niches". Medics tend to be especially well-adapted to the Mediterranean climatic region of California where they are often components of California annual grassland and grass-oak woodland. They are also locally abundant under the mild winters of southern and central Texas.

The burclovers are nutritious and highly palatable, but they do not provide high yields of dry matter. Improved nutritive value and increased concentrations of protein and minerals are their greatest contribution to range animal diets.

The species shown in this slide is known variously as California or toothed burclover and, sometimes, California medic. This vigerous specimen was thriving in northcentral Texas.

Erath County, Texas. March.

193. Common lespedeza (Lespedeza striata)- This naturally reseeding summer (warm-season) annual is one of two annual lespedeza species introduced to the southern United States from Southeast Asia. The other species is Korean lespedeza or Korean clover (L. stipulacea). These relatively low-yielding but nutritious annuals grow over much of the same general region and have much the same agricultural role as the medics and, even more so, the hop clovers presented above . The big difference is that the annual lespedezas do it in the summer (ie. their "empty ecological niches" included the time dimension or "volume" of warm-season). Introduced annual lespedeza species do best on the harsher sites of shallow, rocky, drier soils and full sunlight. They do compete with larger perennials but their cover, density, and herbage yield are greatest when they grow alone as single-species stands (or nearly so).

These introduced annual lespedezas are not as reliable forage producers as are the hop clovers. Common and Korean lespedeza germinate and mature during the some of the hottest parts of the warm-growing season. This is when soil moisture is typically lowest due to high evapotranspiration and when precipitation is less than at any time except for some of the winter season. Also, droughts so frequent to hop clover-lespedeza areas typically do not develop until after mid-spring. Hop clovers can survive as drought-avoiders, but lespedezas are forced to be drought-tolerators. Once annual lespedezas become established a soil seedbank persist that can produce a new crop for years thereafter. A lespedeza crop does not occur each summer. There are good lespedeza years and years when one has to search-- hard-- to find a spindly lespedeza plant. This is but one example of where perennial species are usually more dependable forage plants.

Annual lespedezas are some of the least expensive forage species to establish. Lespedeza is planted by broadcast seeding often on small acreages using hand-crank Cyclone seeders. The author once overseeded almost 20 acres of go-back land using a Cyclone seeder and obtained an excellent stand. Hand-seeding is easiest when broadcasting on top of a light cover of snow. Broadcasting lespedeza seed is one exception to the nearly iron-clad rule to never broadcast seed but instead prepare a clean, firm, friable seedbed. Planting common lespedeza is a "poor boy" range and pasture improvement practice.

Besides producing good quality pasture for livestock and native ruminants like deer at minimal cost, annual lespedeza furnishes excellent feed for bobwhite quail. Adult birds are often observed bringing their broods onto stands of the short common lespedeza through which the young chicks can easily move and forage.

Ottawa County, Oklahoma. September.

194. Sericea lespedeza (L. sericea= L. cuneata)- Sericea lespedeza on an old-field in the Ozark Plateau. "Sericea", as it is known to farmers and stockmen, is the major introduced Lespedeza species that is a perennial. It was widely planted for hay and permanent (tame) pasture, soil-building, and wildlife feed, especially for bobwhite quail. It was commonly seeded during the 1950s and 1960s on land enrolled in the Soil Bank Program like that shown in this slide. In areas having lots of infertile soils (either natural or "farmed out"), such as the general region encompassing the Ozark Plateau and Cherokee Prairie, fields formerly in row crops or small grains were seeded to sericea or sericea-perennial grass mixtures, commonly sericea and tall fescue or sericea and bermudagrass. Sericea really "took aholt" and quickly naturalized. It will "grow on a rockpile" and persist through severe drought to produce high-quality hay and pasture.

Unfortunately sericea eventually became a weed over vast areas of the Central Lowlands (again, the Cherokee Prairie for example) and became a threat to native vegetation and other agronomic crops when it invaded tallgrass hay meadows and permanent pastures. Sericea was either declared or generally regarded as a noxious weed in states like Kansas and Missouri, at least in certain counties.

This is an old story in which only the plant names change (mostly new species are added to the noxious list). Many of the exotic "conservation species" that were introduced and established-- through cooperation of a well-meaning United States Department of Agriculture and conscientious agricultural producers-- for erosion control, increased forage production, and wildlife habitat became major pests. Other examples include kudzu (Pueraria lobata), Johnsongrass, and multiflora rose (Rosa multiflora).

This does not mean that sericea lespedeza has no benefit to agriculture and conservation and should therefore be eradicated (assuming for argument that such is possible). It means only that sericea must be managed more carefully, including standard weed control practices (perhaps even eradicated in localized areas). On many smaller farming and ranching operations sericea remains the "poor man's alfalfa". Hay made from immature sericea and baled properly so as to retain its leaves is on par with alfalfa (which may require irrigation even in humid zones with frequent summer drought) and red clover (which requires richer soils like bottomland). Sericea is still valuable for deer forage and bobwhite quail feed (sometimes cover) though perhaps inferior to common and Korean lespedeza for upland game species. With decline of the bobwhite throughout the Southeast sericea may have increased value, but any such conclusions should be based on findings from research that verify value of sericea to wildlife including song and game birds.

Ottawa County, Oklahoma. September.

196. Sweetclovers or melilots (Melilotus species)- Yellow sweetclover (M. officinalis) and white sweetclover (M. alba) growing together on rangeland in Alberta. Martin et al. (1976, p. 645) indicated that these natives of Asia Minor were used as green-manure and bee plants 2000 years ago and that they had naturalized in parts of Virginia over 30 years before the Declaration of Independence. Like so many crops, including introduced forage species that later naturalized (eg. Kentucky bluegrass, white clover), dispersion and, ultimately, invasion of the sweetclovers corresponded with the predominant westward movement of the frontier. Eventually the melilots became most common in the Great Lakes and Upper Midwest Regions, but the two most common species are forbs on range and pasture lands as far south as Texas.

The Melilotus species present in North America are biennial and, rarely, annual plants. The biennial life cycle is, of course, far less frequent than annual and perennial cycles being particularily uncommon on range except among some groups like certain of the Compositae. The two major melilot species are white sweetclover and yellow sweetclover. Basic biology of these species was given by Martin et al. (1976, ps. 645-649) and in the various editions of Iowa State's Forages text beginning with the first edition (Hughs et al., 1951, ps. 166-171).

From their high rankings as some of the major forage legumes, white and yellow sweetclover became consistently less important as field crops until currently they barely merit mention (less than one page) in Forages (Barnes et al., 1995, p. 277). The sweetclovers, like many of the earliest introduced forage species, in effect became obsolete (at least under intensive, agronomic management). Ironically (sadly perhaps) some of the early introductions that were major crops eventually proved to be less productive than newer introductions or improved cultivars of existing species. As noted earlier in this section, some of these agronomically obsolete species even became weeds because though less productive of forage they had superior survival and aggressiveness as ruderals or as reversions back toward the wild types.

Reversion or atavism (being or becoming like a remote ancestor, rather than like the parents) though a "vice" among cultivars, inbred lines, etc. often becomes the essence of persistence "in the wild". The very characteristic that dooms species as domesticated crops and makes them weeds to farmers and agronomists can make them invaluable to ranchers and rangemen. To some extent this is the case for the naturalized sweetclovers. The two biennial sweetclovers remain locally ("spot") dominants and important sources of forage as range forbs. It was explained above that the major limitation of legumes on range was survival, longevity of stands, persistence, you provide the term. Any legume --native or perennial-- that persist on range is desirable as superior livestock and wildlife feed, soil nitrogen increaser, bee plant, or wild flower unless it has noxious qualities like competitiveness with more desired plants (the bad aspect of "weediness") or toxic properties.

In the latter of these there was a response from the "raw nerve". Yellow and white sweetclovers are, under certain conditions, one of the classic poisonous plants. They contain a glycoside that is an ether of glucose and coumarin. When coumarin is metabolized by molds (ie. moldy sweetclover hay) the compound dicoumarol results and in turn causes a Vitamin K deficiency. Out of this discovery and subsequent research by the University of Wisconsin th rat poison Warfarin (for Wisconsin Alumni Research Foundation) was developed and patented. This interesting story, that is surely relayed in every poisonous plant course in every School of Veterinary Medicine, was given by Cheeke and Shull (1985, ps. 186-190). And as always readers are referred to Kingsbury (1964, ps. 342-346) and Burrows and Tyrl (2002, ps. 590-594).

Sweetclover poisoning is largely a thing of the past as Melilotus species were phased out with more and improved cultivars of other hay crops, notable alfalfa, and-- likely in no small part-- to the threat of livestock toxicity. The two sweetclovers remained common range legumes and can be found, sometimes locally in great populations, from the Allegheny Mountains across the Sierra Nevada. They were recognized as having naturalized in California by the time Jepson published the first comprehensive California flora in 1925. Sweetclovers have generally had a positive impact on the range vegetation of North America.

Habitats of white and yellow sweetclovers are so similar that the two species often grow side-by-side as seen in this slide. Cardston Municipal District, Alberta. July.

Vetches (Vicia species) comprise another group--albeit it a small one--of introduced forage legumes that are sometimes important, especially at local scale, on North American ranges. There are Vicia species that are native to parts of North America, but generally the vetches that are most important as forage plants are the self-seeding annual species. All of these are cool-season plants that most commonly are only component species in mixtures with grasses. Unlike most introduced grasses and many legumes the introduced vetches are not usuallly used as or develop into single-species pastures. Rather these cool-season annuals are component and, commonly, secondary species with grasses. At one time certain vetch species (especially V. sativa and V. villosa) were used as a complementary species on small grains pasture (particularily winter wheat and rye). This works well if the small grain species (say, wheat or rye) is used striictly as pasture and "grazed out" rather than removing livestock and allowing the grain production which is subsequently harvested. Vetch seed, especially of V. sativa, is close enough in size to wheat or rye grain that it is a major contaminant of these crops resulting in severe discounts or even rejection of wheat or rye at the elevator. For this (and various other) reasons some of the introduced vetches are regarded as weeds in some states and regions.

Traditionally the major use of vetch species has been as nutrient-dense, high-protein sources in pastures that are made up primarily of introduced grasses. This varies from small grains pasture to permanent pastures of introduced perennial grasses, both warm-season species (eg. green, actively growing vetch in dormant herbage of bermudagrass) and cool-seson Eurasian species (eg. a cool-season legume complementing the lower nutritive value herbage of tall fescue, ryegrass, or orchardgrass). Aside from the main agronomic use of vetches (as a domestic forage crop) the value of vetch on range is as naturalized species that furnish greater concentrations of nutirients and add some stability to forage production through species diversity. The vetches also provide some cover as well as high-quality feed for species of wildlife, many of which are highly selective feeders (eg. deer, turkey, quail). Vetchs can also serve as valuable bee plants and furnish soil cover for watersheds. Overall, vetches with their leafy sprawling habit provide protection against soil erosion. Some Vicia species have been routinely planted as winter cover crops, especially in southern states like Oklahoma. Under certain conditions these nodulated legumes can fix atmospheric nitrogen thereby increasing soil nitrogen which in turn benefits their grass competitors.

A major drawback with any annuals (even self-seeding ones) is their "feast-or-famine" nature of production. In years favorable for germination of vetch seed these species produce "bumper crops", more than animals can begin to eat. At the other extreme there are years with little or no vetch is produced. Between these extermes there is enough variation in herbage production that grazing capacities cannot be estimated with much reliability. This is the problem inherent with any annual forage crop, cool- or warm-season (except under unique situations), especially of legumes. Self-seeding annual clovers and lespedeza confront the producer with the same problem. Annual grasses are much less variable in germination and herbage production than are legumes. Even with grasses, however, there are years when this (these) annual species or that predominate.

The vetchs were never major forage species (other than locally or on individual operations) and their relative importance as forages declined over the last half century or so as evidenced by converage in the classic undergraduate-farmer reference Forages by Iowa State University Press (compare first to latest edition). Most treatments of vetch culture were in various USDA and state agricultural experiment station publications (most of these long out of print). Hairy or winter vetch (V. villosa) is common and valuable enough that it was recognized as an important range plant (even though an introduced species) by authorities in Texas (Hatch and Pluhar, 1993, ps. 218-219) and Oklahoma (Tyrl et al., 2002, ps. 210-211).

Regardless of noxious features, highly vriable forage production, and primary use as field crops the vetches are frequently valuable naturalized range legumes. Some examples werer included below.

202. Not sweet peas or string beans, but pulses nontheless- Immature though ripening legumes (as well as a flower) on a shoot of naturalized common or field vetch. The fruit type of legumes is a legume or, as it is also known, a pulse. Pulse crops are legumes raised for their pulses (legumes) that are used as human food or animal feed. The pulses of forage legumes on range and pasture are an frequently important sources of critical nutrients, especially protein and amino acids. Pulses constitute concentrate feeds in contrast to other plant parts like stems and leaves which are roughage or forage feeds. It is important to remember this basic fact when considering the nutritive value of legumes growing on grazing lands.

Erath County, Texas. April.

204. Woolypod or smooth vetch (Vicia dasycarpa)- This showy specimen is an example of one of at least eight Vicia species grown as field crops in North America, but there are as many as three dozen species of native or naturalized vetches here (Martin et al., 1976, p. 760). Most vetches are annuals, rarely biennials, and most of the agronomically important ones are grown as winter annuals. Obviously these are best adapted to southern areas, especially the southcentral and southeastern states. Vetches are sometimes overseeded on fields of warm-season perennial grasses such as bermudagrass. Upon, or shortly after, their introduction vetches were seeded with wheat to improve diets of wheat pasture cattle. It was quickly discovered that many of the wheat pasture vetches were weeds because the closeness in size of vetch seed and wheat kernals allowed contamination of grain in the combine. In some cases the cool-season vetch grew to such size and abundance as to compete with winter wheat. When wheat is grown solely for pasture and will be grazed out the combination of vetch and wheat is pasture without peer.

On range, vetch usually grows as either isolated plants or small stands and typically amounts to little more than increased diversity in plant community and animal diets.

The Eurasian V. dasycarpa shown here was readily identified by the infloresence that has all of the papilionaceous flowers arranged along one side of the rachis or central stem. It lacks the pubescent leaves of the very similar hairy vetch (V. villosa). By the way, some of these vetch species share the same common name, depending on authority,.which can be a source of confusion. Smooth vetch is a common naturalized species in isolated spots on tame pasture (eg. bermudagrass, tall fescue), tallgrass prairie and, as with the speciemen shown here, oak-hickory-tallgrass prairie savanna.

Ottawa County, Oklahoma. June.

205. Beauty (and a study) in a naturalized forage crop- General view of woolypod or smooth vetch showing its inflorescences with papilionaceous flowers aligned on one side of the rachis (central stalk) of the flower cluster. Also visible in this photograph were long, twining tendrils characteristic of this species. An addition perquisite of this naturalized legume was the aesthetic value found in the elegant beauty of this economically important plant of field and range.

Ottawa County, Oklahoma. May.

206. Wooly, hairy, or winter vetch (Vicia villosa) in the Oregon Coast Range- Wooly or woolypod vetch has naturalized through much of North America. Whereas the immediately preceding photographs were taken in northeastern Oklahoma this photogarph was of a naturalized grassland in the Coast Ranges of westcentral Oregon. The grasslands in the California's Central Valley and Coast Ranges except for extreme northern California are in the region of Mediterranean climate where naturalized grasslands are of Mediterranean annual grasses and forbs. Farther north in the Coast Ranges Eurasian cool-season, perennial grasses are typically dominant and the important forage species. Wolly vetch is an example of a naturaized Eurasian annual legume that is common and a locally important range forb.

This example was from a forest range of such introduced and now naturalized species. Also present was orchardgrass, tall fescue, perennial ryegrass, and the annuals, soft chess (Bromus mollis) and ripgut borme (B. rigidus= B. diandrus).

Benton County, Oregon. June.

210. Field of kochia (Kochia scoparia)- Kochia is an Eurasian annual that has naturalized over a wide area particularily in the semiarid zone. Kochia is a member of the saltbush or goosefoot family (Chenopodiaceae) which in North America is second only to the rose family (Rosaceae) in number and value of range browse plants. Kochia or, as it is also known, belvedere and summer cypress is obviously a forb; in fact, it is primarily an alien weed much like other chenopod species such as lambsquarters (Chenopodium album). Aside from any value as a successional pioneer species and related aspects such as vegetative cover for soil protection, the main utility of kochia is as palatable forage. As is the case for other herbaceous chenopods (such as lambsquarters for example) these weeds--and they qualify as weeds in both agronomic or cultural as well as ecological definitions--are often highly relished by most kinds and classes of grazing/browsing animals. Even gras-preferring grazers like cattle readily graze immature kochia or, less commonly, more mature plants under certain conditions, and not just when there is nothing else to eat (eg. when dried leaves are softened by rain). Cattle, sheep, even horses as well as deer consume shoot tips and young leaves.

Farmers and stockmen were quick to realize the feed value of kochia (even though it is a major weed on disturbed land such as farm fields and overgrazed ranges) when it became apparent that animals ate it to varying degree. This was most noticable where kochia grew in dense populations in corrals, go-back land, barrow ditches, trails, etc. A few growers in the Great Plains began to manage some of their fields to encourage kochia production. Eventually some producers planted kochia as a crop while others raised kochia for seed. The stand of kochia shown here was being grown as a commercial seed crop though at this time there was not a Certified Seed program for this new crop.

Kochia produces nutrient-dense forage (eg. crude protein levels sometimes exceed 20% although some of this may be non-protein nitrogen) of fairly high palatability, but it is also a poisonous plant under certain conditions. Poisonous principles (=toxic substances in kochia tissue) include alkaloids, nitrate, sulfate, saponins, oxalates, and even liver-damaging agents. Thus, livestock toxicity from kochia can span an array of maladies from nitrate toxicity to alkaloid poisoning to oxalic acid poisoning to photosensitization as a secondary symptom and disease resulting from hepatatis (Burrows and Tyrl, 2001, ps. 351-355 passim; Hart et al., 2003, p. 118). These and other references offered management suggestions for reduction of livestock losses to kochia. All things considered, on most livestock operations animal poisoning due to kochia does not override the feeding value of this naturalized "poor boy or substitute legume". The economic gain from nutrient-rich kochia forage is greater than economic losses from kochia-poisoning. Afterall, in some situations alfalfa (Medicago sativa) contains bloat-causing saponins, induces photosensitization of both primary and secondary types, and accumulates toxic levels of nitrates (Kingsbury, 1964, ps. 33, 43, 57, 341-342). No, kochia is not alfalfa-quality feed; neither does it require the expensive, intensive imputs and demanding culture of alfalfa. Hart et al. (2003, p. 118) provided a pertinent summary of this naturalized forb: "Despite its drawbacks, kochia can sometimes be considered a valuable forage".

The Society for Range Management (Stubbendieck, Hatch, and Butterfield, 1992, ps. 312-315) included both kochia, Eurasian naturalized forb, and greenmolly, perennial summer cypress, or red sage (K. americana), the native shrub, on the list of 200 plant species for its Intercollegiate Range Plants Contest. Consistent with this, the contest list also included the chenopodiaceous Eurasian annual forb commonly known as Russian thistle (Salsola iberica= S. kali-tenui-folia). Both of these forbs are the common "tumbleweeds" of the Western Range. Both species break off near the ground surface and, being spherical in general shape, they roll and bounce wildly across the land disseminating their seed as driven by ever-present winds.

Erath County, Texas. May, early bloom stage.

215. Iuka eastern gamagrass- No, Tripsacum dactyloides (viewed as the grandpappy of North American tallgrass species) is most definitely not an introduced forage crop. As was explained in the introduction to the grassland biome, eastern gamagrass was included in the Introduced Forages section to illustrate a general agricultural prinicple or practice. Native plant species (a range grass in this example) can be domesticated (to increase biological yield and increase economic efficiency) and made dependent upon and responsive to intensive management just as domesticated plant species (say introduced pasture grasses and legumes) can naturalize through natural selection (revert back to, or closer to, the wild type) or be introduced and managed less intensively (extensive management) as if they were native or naturalized species. All of the introduced species in this section fell into the latter category. This last species was included here to illustrate the former category.

The following interesting explanation and interpretation of Iuka eastern gamagrass and the particular 40 acre stand of Iuka presented in these two slides was related to the author by Glen Snell (personal communication, 12 August, 2002; address and telephone at end of this caption) and supplemented by information from the web address of www.sprrs.usda.gov/eggintro.htm. Beginning in the late 1940s to early 1950s various USDA agencies like the Soil Conservation Service (SCS) and Agriclutural Research Service (ARS) began collecting accessions of and breeding eastern gamagrass. At one time there were over 500 native clones of this species. These were eventually narrowed down to the 21 most vigerous, highest yielding, or apparently "best" (most acceptable) accessions which became the basis for a breeding program that eventually lead to development and release of Iuka IV. At some point, the ARS Southern Plains Range Research Station near Woodward, Oklahoma began pioneer breeding work with eastern gamagrass. This was under the research leadership of Chester L. Dewald. In 1988 the eastern gamagrass cultivar known as Pete was released by the SCS and ARS in cooperation with the Kansas and Oklahoma Agricultural Experiment Stations. Pete was developed from composite seedstock selected from 70 native populations from Knasas and Oklahoma following propagation for three generations with open-pollinated grain. Release of Pete followed prerelease of PMK-24 in 1974.

Meanwhile Mott Farms near Iuka Kansas began cooperating with the Southern Plains Range Station and seeded one acre under center pivot irrigation to the final 21 accessions mentioned earlier. At end of the first growing season Mott Farms gathered all the interbred seed produced from the one acre open-pollinated, 21 accessions-parental population. Any dormant seed from the 21 accessions would not have germinated this first growing season so there was no progeny of dormant genotypes harvested. This initial harvest of interbred grain from the 21 accessions was the first filial generation (F1). The F1 grain was planted and grain produced by the F1 generation (plus any grain produced from dormant seed of the parental generation) was harvested. This was the F2 generation. Grain of the F2 generation was seeded and fruit of this F2 (plus any grain from dormant seed of previous generations) was gathered. This procedure stopped with the fourth filial (F4) generation. The F4 progeny served as the basis for subsequent selection in the eastern gamagrass breeding program which culminated with the recognition of the cultivar, Iuka IV, by the National Variety Review Board in April 1995.

Iuka IV and Pete are the only two released cultivars of eastern gamagrass (unless the PMK-24 pre-release is counted). Some eastern gamagrass grain from Mott Farms was not isolated and screened yet it can be sold as Iuka. It is only Iuka IV (after the F4 generation) that is the "genuine article", the actual approved cultivar. Iuka IV was selected for higher yield (the 21 accessions were presumedly the most productive) and ease of stand establishment. Seed dormancy is so common in eastern gamagrass that stratification and/or scarification are standard pre-planting seed treatments. The planting and harvest procedure used in the Mott Farms breeding program attempted to eliminate or greatly reduce the problem of seed dormancy in eastern gamagrass by selecting mostly progeny from early germinating, non-dormant parents. Iuka IV has consistently produced better stand establishment in absence of pre-planting seed treatment (still a recommended practice) when compared to plantings from non-treated grain of Pete or PMK-24. Field trials over a four-year period indicated that Iuka IV producead 19% higher yields than Pete.

Eventually Mott Farms established four breeder blocks for Iuka IV. When interest rates for farmers hit double digits (eventually reaching over 20%) Mott Farms folded and sold its eastern gamagrass enterprise to Glen Snell. Three of the IUKA IV breeder blocks were plowed under. The fourth and last remaining breeder block is the 40 acre field shown in these photographs. Shortly after these slides were taken the field was combined and yielded 50-60 pounds of Pure Live Seed per acre. Seed from this field can be sold as Breeders, Foundation, Registered, or Certified Seed.

Management of this stand involves burning off each spring and application of 150 pounds of actual nitrogen per acre. Eastern gamagrass is baled following seed harvest. Soils are the Pratt and Canadian series.

It is an interesting irony that this field was once virgin prairie plowed under and turned into a corn field. Now it has been returned to a prairie grass closely related to corn. In fact, corn and eastern gamagrass (known by early settlers as corn grass) have been hybridized in crop breeding research. More recently, genes from eastern gamagrass were used to confer apomixis (seed production without fertilization; asexual seed production) to corn in order to produce apomictic hybrids in hopes of retaining the heterosis of hybrid corn for more than one generation.

Peak standing crop (soft-dough grain stage). Barber and Harper Counties, Kansas. June.

Address of Glen Snell: 300 N. Adams Street, Medicine Lodge, Kansas 67104; (800) 279-5841.

216. Individual plant of Iuka IV eastern gamagrass- This immense, sprawling tuft is the characteristic habit of eastern gama grass. Each plant (the clump or bunch of each genetic individual) continues to grow by increase in number of tillers. Haymakers claim it is a "rough, wild ride" mowing and baling this giant bunchgrass. But that we should all be so lucky.The 40 acre field photographed here is probably one of the most productive temperate grasslands on Earth. After grain harvest, this field still yielded 1.5 ton of hay per acre-- under dryland conditions in one of the most extreme droughts (2002) in history of Kansas weather records.

One of the most important findings discovered once eastern gamagrass began to be grown as a field crop in single-species stands was how remarkably drought-tolerant this large prairie grass was. Initially producers grew eastern gamagrass under irrigation, especially in the semiarid climate of the Great Plains. Easterrn gamagrass was afterall now a domesticated or agronomic forage crop and it was logically assumed that "solid" stands of this big, broad-leafed "ice cream species" would require supplemental water (and lots of it), especially when managed intensively as tame pasture or a mechanically harvested forage crop. Growers were pleasantly surprised at the remarkable drought tolerance of this deep-rooted, native prairie grass even when grown as a cultivated, "tame" field crop. Now eastern gamagrass-- including cultivars --is being sold as a "drought-buster". And to a rangeman that beats a sod-buster any day. Barber and Harper Counties, Kansas. June.