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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.).
The contribution of crested wheatgrass to diets of native ruminants like pronghorn antelope (Antilocarpa americana) and mule deer (Odocoileus hemionus) is unfortunately all too often unappreciated by those who decry crested wheatgrass seedings as monocultures of introduced species.
2. Crested wheatgrass (A. cristatum) range- Crested wheatgrass seen here is just entering dormancy having set seed and completed it's annual growth cycle. This seeding was in southwestern South Dakota (Pennington County). When it is compared with the one in Idaho and the crested wheatgrass specimens in Alberta the reader can grasp the longitidunal range in North America over which this introduced Eurasian species is adapted. July. No FRES number of Kuchler for this non-native, but naturalized rangeland cover type. SRM 614 (Crested Wheatgrass).
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.
6. Crested wheatgrass (Agropyron cristatum)- Crested wheatgrass includes only A. cristatum in the broadest sense. In a more strict or specific sense crested wheatgrass is the generic name for A. cristatum, A,. desertorum and, less commonly, A. fragile.
The definitive authority on crested wheatgrass (in all aspects) is the symposium proceedings edited by Johnson (1986).
7. Crested wheatgrass- Crested wheatgrass of the species, Agropyron desertorum. According to the flora for Colorado by Weber and Wittmann (2001) and others such as Beidleman (2000) it is A. desertorum and not A. cristatum that is found in the Centennial State. The following specimens from Colorado (Lincoln County) were presented to show readers the other major species of crested wheatgrass introduced on the ranges of western North America. Comparison of the infloresences of these plants to those shown immediately above showed that C. cristatum has spikes that are shorter and wider than those of A. desertorum. June.
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.
10. Appearance of a crested wheatgrass seeding at end of growing-grazing season- This Clayey range site had been put to the plow and was go-back land when it was reseeded to crested wheatgrass. The seeding has persisted for many years and now also supports scattered native grasses including western wheatgrass, blue grama, and green needlegrass . Perhaps this is one of the lasting "legacies" of this marvelous introduced range grass: it prepared the way for at least partial recovery of the native species. This is the process of reaction (later termed facilitation) in the Clementsian model of plant succession in which each seral stage improves the sere (the total environment of a given site complete with the complete sequence of plant communities that will come to occupy it) for the next seral stage (the plant community at that stage of vegetation development) until the termination at climax. If crested wheatgrass persist indefinitely, as seems likely, as a disclimax or anthropogenic climax it will continue to conserve soil and contribute to range restoration while providing valuable forage for livestock and wildlife. That is a most fitting accomplishment for any pasture crop. The introduction of and development of management programs for crested wheatgrass was an example of Man, Manipulator of Ecosystems, at his best. The establishment of crested wheatgrass range was a technological revolution in grassland agriculture. Pennington County, South Dakota. July.
11. A seeding of crested wheatgrass (primarily A. cristatum) and a cultivar of dryland alfalfa- This former cropland was seeded to a mixture of crested wheatgrass and a cultivar of the wild, yellow- or variegated-flowered, branch-rooted alfalfa (Medicgo falcata cv. Ladak). The classification by Vallentine (1990, ps. 7-13) of grazing lands as: 1) long-term, 2) medium-term, and 3) short-term was presented in the Introduction of this publication. Crested wheatgrass is the case exemplar of an introduced species being used as or becoming naturalized (vs. native) range. The Vallentine (1990, p. 10) designation of "seeded range (introduced species)" as a category of "long-term grazing land" with duration of stand in excess of 40 years fits crested wheatgrass "to a T". By comparison, persistence of alfalfa is at best of such a short interval that this seeded introduced species would fall into the Vallentine (1990, p. 10) designation of "permanent pasture" under "medium-term grazing lands" (a period of about 10 years). Those working closely with this grass-legume mixture specified however that the Ladak alfalfa is both less palatable and later-maturing (phenological development is slower) than the crested wheatgrass with the result that the latter is grazed heavier (greater degree of use) and earlier in the growing season (Western Section, American Society of Animal Science annual meeting tour, June, 2001). This favors the alfalfa and enables it to remain competitive with the grass. The net result is that Ladak alfalfa is not grazed out and the seeding is not converted to a single species stand of crested wheatgrass. Alfalfa has also been grown in seedings with intermediate wheatgrass (Agropyron intermedium), Russian wild rye (Elymus junceus= Psathyrostachys juncea ), and in mixtures of these two species plus crested wheatgrass (Heinrichs, in Campbell and Herbel, 1975, ps. 54-56). No FRES or Kuchler designation. Variant of SRM 614 (Crested Wheatgrass). Gallatin County, Montana. June.
12. Dryland creeping alfalfa – An individual plant of Ladak alfalfa (Medicago falcata cv. Ladak). This is a variety of the wild yellow-flowered alfalfa that is native to Eurasia. Ladak was selected from among native populations of M. falcata in the province of Ladakh, India (Heinrichs, 1963, ps. 317-320 passim). Ladak has variegated inflorescences and a branched-root arrangement (vs. the more pronounced or typical pattern of the dicotyledon taproot system). This cultivar (accession may be the more precise designation) was selected for rainfed agriculture in a semi-arid region and it has been used in intercrosses with other M. falcata lines to produce hybrid types having creeping rootstocks (creeping-rooted alfalfa) (Heinrichs, 1963, p. 322-324).
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.
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13. Alfalfa- Leaves and inflorescences of Medicago sativa. Numerous Medicago species are of economic importance as pasture and hay crops. Alfalfa is the most important of all. It is the most important hay species in California, the agricultural wonder of the world. It is probably also the most important hay crop in the Agricultural Region known as the Grazing and Irrigated Crops Region traditionally viewed as extending from the Front Range of the Rockies (western edge of the Great Plains) westward to the crest of the Sierra Nevada and Cascade Ranges. While there are important introduced grass species (eg. timothy, redtop, reed canarygrass) grown for hay in various locations throughout these regions alfalfa is likely the most important hay crop in the vast area known as the Western Range. Development of alfalfa cultivars which are adapted for either dryland pasture or even range has long been a goal of plant breeders in the Western Range. Unfortunately this remains a largely unfulfilled goal, but some breeders have developed cultivars of the creeping (= creeping-rooted) alfalfa. Whereas the typical M. sativa varieties have a prominent tap-root system the wild yellow-flowered alfalfa (M. falcata) has a "branched-root" form of the fundamental dicot taproot system. This morphological shape or pattern of root arrangement permits this Medicago species to exhibit a "creeping" or "spreading" habit rather than the bunched or tufted habit of typical M. sativa plants. This creeping rootsystem permits expansion by asexual reproduction (and consequent better survival under grazing) in M. falcata. Alfalfa breeders have developed several M. falcata X M. sativa hybrid (sometimes identified as M. media) cultivars including Rhizoma, Rambler, Teton, Nomad, Roamer, and Drylander with the latter being the one best adapted to dryland pasture production. Some of these have creeping rootstocks or even rhizomes under certain conditions (eg. Rhizoma, Nomad, Rambler). Unfortunately these are often low-yielding and relatively restricted in their range of adaptation (see review by Heinrichs, 1963 and Heinrichs, in Campbell and Herbel, 1975, p.54-57). Overall these cultivars are relatively drought-tolerant and persistent on seeded range, at least by legume standards, but the goal of legumes having persistence on range like that of introduced grasses is not yet a reality.
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. Spikes of desert crested wheatgrass (Agropyron desertorum)- Close-up views of spike and flag leaf of desert crested wheatgrass naturalized in the sedimentary Northern Great Plains. Garfield County, Montana. Mid-June; immediate pre-anthesis; phenological stage.
21. 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.
22. 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.
23. 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.
24. Another example of ideal habitat on an old seeding- This seeding of crested wheatgrass in the northern San Luis Valley had been reinvaded by Wyoming big sagebrush. The sparse cover and density of sagebrush probably closely resembled that of virgin semidesert grassland. The savanna or, at least, savanna-like nature of this naturalized (man-made) range was another textbook case where re-establishment of Wyoming big sagebrush combined with relatively heavy cover of crested wheatgrass provided ideal habitat for wildlife as well as outstanding forage for livestock (cattle in this case).
Through experience, and in conjuction with some shift of socioeconomic values, it is now realized that "pure stands" of crested wheatgrass are less desirable than wheatgrass seedlings that become re-occupied by some cover of native shrubs like this Wyoming big sagebrush.
Saguache County, Colorado. Late June.
25. Another introduced wheatgrass- Representative plants of Siberian wheatgrass (Agropyron fragile= A. fragile ssp. sibericum= A. cristatum ssp. fragile) seeded on degraded range in the northern San Luis Valley. Siberian wheatgrass is another introduced Agropyron species that is closely related to A. cristatum and A. desertorum. Siberian wheatgrass is more drought-tolerant than these more widely seeded species ( Smoliak et al., 1990; Natural Resources Conservation Service, 2003). In addition, Siberian wheatgrass is better adapted for reclamation of drastically altered land such as strip mines. Furthermore, it is probably more palatable to both livestock and big game species than the related Old World wheatgrasses reflecting, perhaps, finner shoots, culms leaves, and spikes of Siberian wheatgrass (Natural Resources Conservation Service, 2003).
This exotic species is especially aggressive and competes well against cheatgrass (Bromus tectorum), but it does not have weed features that allow it to escape beyond seeded areas and it can be used in mixtures (Smoliak et al., 1990). Historically, Siberian wheatgrass has been planted as single-species stands like crested and desert crested wheatgrasses, and like these more comonly planted Agropyron species Siberian wheatgrass withstands heavy grazing.
By standards of introduced grass species Siberian wheatgrass has been around for quite a while having been seeded on western ranges since the 1920s (Smoliak et al., 1990). There are two main cultivars of this species, P27 and Vabilova, developed through efforts of Soil Conservation Service Plant Materials Centers, Agricultural Research Service, and Utah Agricultural Experiment Station (Natural Resources Conservation Service, 2003).
Saguache County, Colorado. Late June.
26. Siberian spike- Spike and spikelets of Siberian wheatgrass. Saguache County, Colorado. Late June, immediate pre-bloom (first slide) and anthesis (second slide).
27. 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.
28. Former cropland now stabilized- A new (two-year-old) seeding of intermediate wheatgrass on highly erodible land that was retired from farming and put under the protective cover of an introduced perennial grass through the Conservation Reserve Program. Yuma County, Colorado. Mid June, prebloom phenological stage.
Organization Note: Examples of conservation and restoration of grassland in the Mixed Prairie-Shortgrass Plains Region through the Conservation Reserve Program were included at the end of the chapter, Mixed Prairie-I.
29. 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.
30. Spike of intermediate wheatgrass- An elongated inflorescence (relative to most other wheatgrasses) is a characteristic of this species. Lincoln County, Colorado. June.
31. 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.
32. Spikelets of intermedite wheatgrass- Spikelets on this spike of intermediate wheatgrass were in anthesis as were thos in an above photograph of an intermediate wheatgrass spike. Ochoco National Foirest, Wheeler County, Oregon. June.
33. 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 favorably (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 purebred 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.
34. Robust individual of intermediate wheatgrass- The cespitose morphology of intermediate wheatgrass was shown to good advantage in this "hale and hearty" specimen that was growing on the dry-land pasture presented in the immediately preceding photograph. Whitman County, Washington. June, early hard dough stage of grain.
35. Spikelets up close- Spikelets of intermediate growing on the non-irrigated, bottomland pasture showed and described above. Whitman County, Washington. June, early hard dough stage of grain.
A plant breeding- Spikelets at anthesis on spike of intermediate wheatgrass. Big Horn County, Wyoming. Late June (early summer); anthesis.
36. 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 its 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 that 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).
37. Tall wheatgrass- One individual plant of Jose tall wheatgrass showing the cespitose habit and the large, rank shoots that are characteristic of this species. Phenological stage of peak development; grain at soft dough stage.
38. Spikes of tall wheatgrass- All slides from Otero County, Colorado. June.
39. Quackgrass, quichgrass, or couchgrass (Agropyron repens= Elytrigia repens= Elymus 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 obscure 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 understory 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.
40. Examples of quackgrass- Specimens of quackgrass pulled from the stands shown in the previous slides. Total shoot height was between three and four feet. Readers should observe the large inflorescences (spikes) which have potential for heavy grain production. Viewers should also note the organs of asexual reproduction which were shown in a detailed close-up shot in the next slide. Agricullture Canada Research Centre, Lethbridge, Alberta. July.
41. 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.
42. Something to quack about- Local stand, a colony, of quackgrass that had invaded a marsh on the glaciated Northern Great Plains. The highly productive--to chagrin of famer and delight of grazier--quackgrass or couchgrass performs and survives best in moist to wet soil.
Quackgrass was described as a "'Dr. Jekyll and Mr. Hyde' grass" in the always ready Pasture and Range Plants (Phillips Petroleum Company, 1963; p. 4). Quackgrass is an insidious weeed on cropland and described as such in Weeds of the West (Whitson et al, 1992, ps.410-411), but this aggressive perennial with its sprawling rhizomes is not a "major league" weed on a global or even a difficulty of control basis. Although quackgrass was included in group 2 of the "world's worst weeds" by Holm et al., 1991, ps.153-168) group 1 (by far the worse category) included (in the following rank of weed importance): bermudagrass (Cynodon dactylon), barnyard grass (Echinchloa crusgallii), goosegrass (Euleusine indica), Johnsongrass (Sorghum halapense), hairy crabgrass (Digitaria sanguinalis), and wild oats (Avena fatua). So as weeds--in particular, the grasses--go quackgrass is not exactly on the most wanted list, at least other than as a farm-to-farm situation. Quackgrass has proven not to be as important weed as it once was (Heath et al., 1973).
Weed status is no reason to condem this one-time agronomic forage species. Comparatively low productivity of this Mediterranean native is, and it is most likely the main reason quackgrass lost out to higher-yielding Eurasian species such as smooth bromegrass and reed canarygrass. Quackgrass has proven to be quite adaptable (much like bermudagrass, Johnsongrass, wild oats, etc.) now growing in North America from Greenland to Mexico. For this reason quackgrass is regarded as and has legal status as a noxious weed so that in many states this species cannot be legally planted or sprigged (Barnes et al., 2007, p.229). In 2001 the Minnesota Agricultural Experiment Station released Everette cultivar of quackgrass. Amendment of Minnesota seed laws were required for release of this rhizomatous synthetic clone (Barnes et al., p. 226, 229)
Quackgrass does have the proverbial "potential" or "promish" as an improved agronomic forage, but to date its greatest use as been a a parent for hybrid grasses such as those between bluebunch wheatgrass (Agropyron spicatum) and crested wheatgrass (Heath et al., 1985, ps. 174-175; Barnes et al., 1995, ps. 384-385) It is mostly in Alaska where quackgrass has also naturalized that this species has become an important hay crop (Heath et al., 1973, p. 495). Time and events will prove forage and conservation merits of Everette (perhaps other cultivars?) quackgrass. Sheaffer et al. (2004) reported comparative forage yields and nutritive value of quackgrass relative to more accepted agronomic perennial grasses.
Burleigh County, North Dakota. Late June; early anthesis phenological stage..
43. Between prairie sky and wet soil- Sexual shoots of quackgrass with spikes entering anthesis. These tillers were on plants in the colony introduced in the preceding photograph.
Burleigh County, North Dakota. Late June; early anthesis
44. Spike of "mixed views"- Overall spike (first slide) and central portion of the same spike of quackgrass growing in a marsh in glaciated Northern Great Plains. Quackgrass spikelets typically bear three to five (rarely up to eight) florets. Burleigh County, North Dakota. Late June; early anthesis
So important that another set of examples was justified. Quackgrass is to northrn North America what Johnsongrass is to southernNorth America-- weed or Godsend; trash or treasure depending on rangeman looking at it. So it was that this aurhoring rangeman included another short set devoted to Agropyron repens.
45. Another local quack patch- Numerous shoots (tillers) on a local, small group of several plants or a single, large, rhizomatous plant of quackgrass growing in the floodplain of Musselshell River in southcentral Montana.
Wheatland County, Montana. Mid-June (late spring); hard dough phenological stage.
46. Lower down view- Lower shoots (tillers) of two plants of quackgrass in bottomland of the Musselshell River in southcentral Montana.
Wheatland County, Montana. Mid-June (late spring); immediate pre-bloom to early bloom stage of phenology.
47. Sibling shoots- Local population of tillers produced by one plant of quackgrass. Shoots of quackgrass include both tillers, intravaginated or vertical shoots, and rhizomes, underground eaxtravaginal or horizontal shoots. This growth/asexual (vegetative) reproduction is coupled with sexual reproduction (seed production) as shown in the next two sets of slides.
Park County, Montana.Late June (summer solstice); soft-dough to hard-dough phenological stage.
48. The sexual optionn- Spikes of quackgrass emerging from their boots (the enveloping or invaginating sheath of the leaf immediately subtending (below) the peduncle of the spike. Caryopses (grains), the fruit of most grass species, in which the seed is attached to the inner layer of the ripened ovary is the form of sexual reproduction in the introduced forage grasss.
Wheatland County, Montana. Mid-June (late spring); hard dough phenological stage.
49. Overall object of the sexual option- Spike of quackgrass. Spike is the unbranched or simple (versus compound) inflorescence or flower cluster of members of the Tritaceae or Hordeae (wheat or barley tribe of the Gramineae). On spikes, the spikelets are sessile (versus pedicelate which is a structure that is borne between spikelet and rachis).
Park County, Montana.Late June (summer solstice); soft-dough phenological stage.
50. Sexual option, old and new- Current season's spike (live, green) and last year's spike (dead, light tan or bleached amber) of quackgrass growing on the floodplain of Musselshell River in southcentral Montana. The reasons, the morphological features, that enable quackgrass to be such an aggressive species (hence, a widespread, common, and highly successful weed or forage crop depending on one's perspective) are the options of either or both 1) asexual reproduction (tillers and rhizomes as shown above and below) and 2) sexual reproduction (grain production) as presented in above photographs.
Johnsongrass (Sorghum halapense), a warm-season perennial of the bluestem grass tribe has the same form of sexual and asexual reproduction. Southrns have Johnsongrass and northerns have quackgrass.
Wheatland County, Montana. Mid-June (late spring); hard dough and mature (grain-ripe) phenological stages.
51. Alien phytomers- Portions of two photomers of quackgrass. Phytomer is a single repeating unit of the grass shoot consisting of 1) the upper portion of one node and the 2) lower portion of the next node, 3) the internode between these two halves of two nodes and 4) the one leaf (sheath and blade) associated with these organs.
Wheatland County, Montana. Mussselshell River. Mid-June (late spring).
52. Creepy crawlies- Rhizomes of quackgrass, an introduced, cool-season, perennial grass, growing on bottomland of the Musselshell River, southcentral Montana. These horizontal (extravaginated), subterranean shoots comprise one "weapon" in the "arsenal" of this horribly aggressive weed or, alternatively, wonderfully productive forage crop.
Wheatland County, Montana. Mid-June (late spring); active rhizome-growth stage.
53. An introduction for and on the northern plains-Russian wildrye (Agropyron junceus= Elymus junceus= Psathyrostachys juncea) was one of the first of the non-native, perennial, range (non-domesticated) grasses of the Hordeae or Tritaceae, barley or wheat tribe, brought to North America having been introduced into Saskatchewan in 1926 and into the United States a year later according to the record as presented in Barnes et al.(2007, p. 230). Russian wildrye, the common name by which this species has long been known, is native to the steppes from Russia across Mongolia to China, but it has been in North Amrica where much plant breeding of this species was done resulting in a number of cultivars and accesssions of this semi-agronomic grass (Barnes et al., ps. 230-232).
In North America Russian wildrye has been planted and evaluated as a non-native range grass as far south as southeastern Colorado (Svejear and Rittenhouse, 1982). Lodge (1971) and Smoliak and Slen (1974) stated that Russian wildrye and crested wheatgrass were the two main grasses used for reseeding range in the Northern Great Plains. Russian wildrye is also well-suited to pasture mixes with common alfalfa (Medicago sativa) as found in a study conducted by Lodge (1971) in Saskatchewan. In addition, Russian wildrye is tolerant of saline soils and was found to be productive under irrigation (Currie et al. (1986). Smoliak and Slen (1974) found that pastures of Russian wildrye had three to five and a half times the grazing capacity of native range. Use of Russian wildrye for pasture in the western Prairie Provinces was discussed in some detail in a publication by Grazing and Pasture Technology Program (2009) in which it was noted that Russian wildrye was "less productive than most introduced forages", slow to become established and prone to stand deterioration without adequate soil nitrogen.
In a somewhat contrasting summarization North Dakota workers (Sedivec et al., 2011, p. 8) concluded that although Russian wildrye was slower to establish it was long-lived and tolerant of drought, salinity, and alkalinity. Sedivec et al. (2011, p. 8) noted that Russian wildrye was fiercely competitive and recommended that it be grown in pastures as single-species stands with grazing reserved for autumn. Smoliak and Slen (1974) made the same recommendation for fall-grazing.
In essence, Russian wildrye is more of an introduced range grass than a tame (agronomic) pasture species. Even when this introduced (naturalized?) bunchgrass is used under more extensive (versus intensive) production systems proper management of grazing and other husbandry practices is critical. That basic precept applies to all crops.
Fort Keogh Livestock and Range Research Station, Custer County, Montana. Mid-June; varying phenological development from anthesis to soft-dough stages.
54. Tall, rank, and high-yielding- Outer appearance and structure of a stand of Russian wildrye at peak standing crop. Some of these shoots were at or, at least, approached five feet in height in what was a record wet year (wintr through spring). Any student of forage quality knows that herbage the nutritive value of roughage at this advanced stage of plant maturity is the lowest (or near the lowest) of that at any stage of living plant tissue. This was an ungrazed sward that would probably be best harvested for the seed crop. These photographs showed the degree to which Russian wildrye could produce phytomass (plant biomass) in the northern semi-arid zone.
Palatability (at least in a relative sense) Russian wildrye is somewhat of a reversal of the usual pattern of declining acceptability to livestock as plant shoots mature (get older). Compared to other introduced, Eurasian, cool-season, perennial grasses (eg. crested wheatgrass, smooth brome) Russian wildrye becomes comparatively more palatable with advancing maturity (Phillips Petroleum Company, 1963 and Nicholson, 2006, p. 36). Given the phenomenon of animal preferences (ie. this order of grazer preferences) a good grass/grazing program should include separate pastures of crested wheatgrass and smooth bromegrass along with single-species stands of russian wildrye with all of these introduced (and, now, mostly naturalized) pasture grasses grown to complement native forage (range) species.
A good "all-around" reference on this species was NRCS Plant Guide- Russian Wildrye (Ogle et al., 2004).
Fort Keogh Livestock and Range Research Station, Custer County, Montana. Mid-June; varying phenological development from anthesis to soft-dough stages.
55. A study in spikes- Spikes of Russian wildrye growing in Northern Great Plains. The spike in this species is enlogated or exerted a considerable distance above the flag leaf. The spike inflorescence with a typically disarticulating rachis is the key distinguishing feature of species in the barley or wheat tribe (Hordeae or Triticeae).
Fort Keogh Livestock and Range Research Station, Custer County, Montana. Mid-June; caryopses at milk stage of phenology.
56. Sex on a spike- Views of three spikes (or parts of spikes) of Russian wildrye at anthesis. There are typically three (sometimes only two) florets per spikelet with three spikelets per node of the disarticulating rachis. Fort Keogh Livestock and Range Research Station, Custer County, Montana. Mid-June.
Note on nomenclature: Russian wildrye is yet another case of "musical chair" names. This introduced species was long known and shown (over several decades) in agricultural journals, circulars, experiment station bulletins, etc. as Agropyron junceus. Then taxonomic investigations, especially those based on detailed, brilliant, and revolutionary work led by D.R. Dewey (Dewey, 1982, Dewey, 1983a; Dewey, 1983b; Deweey and Hsiao, 1983; Barkworth and Dewey, 1985) as made the case for transferring (renaming) Agropyron junceus to, first, Elymus junceus then to Psathyrostachys juncea which is where things stood--as of this writing--in Barkworth et al. (2007, 372).
The matter of naming and aligning genera and species based on genomic analysis is--again, as of this writing--far from settled. Baum et al. (1987) evaluated genome-based classification in the Triticeae and concluded that "genomes cannot and should not be used for generic delimitations" but instead "...must be used with other attributes for circumscription of genera and for their meaningful classification". Baum et al. (1987) concluded: "Genome analysis in the classical sense doe not provide more conclusive evidence of affinities than other techniques and approaches". Within Triticeae "genome analysis does give useful information about the phylogenetic relatships of taxa, but it can be used only to supplement other criteria for the elamboration of a system of relationships among genera".
Stussey (2009, p. 168) specifically used the case of Agropyron and Elymus (including citing Dewey [1983b]) and pointed out that there were several possibilities and, thus, explanations to crossability (crossing studies) between, say, two separate genera. These possibile results range from merging the twogenera into one genus, one genus should be omittted and moved into the other genus, or the, say, two genera should be put together in their own genus. Stussey (2009, p. 168), however, took the cladistic approach to task based on their misconception that crossing revealed nothing as to phylogenetic or evolutionary relationships. Finally, "ease of intergeneric crossability may reveal more about weak isolating mechabnisms than degree of relationship".
Even the non-taxonomist current author could conclude the obvious: names of genera and species and phylogenetic and/or systematic relationships among them is far from settled. Names and phylogenetic arrangements of Triticeae in, for example, Barkworth et al (2007) may be "official", at least in their minds, but that does mean these attributes are taxonomically or nomenclaturally correct (or that they are "official" in the minds of other observers). Readers, you have your choose of published plant names.
57. Smooth bromegrass seeding- Smooth or anwless bromegrass (Bromus inermus) is another cool season Eurasian grass introduced into the pastures and ranges of North America. Its introduction pre-dates that of crested wheatgrass and some other non-native species that are managed as range plants. Smooth brome is not as important for "non-native range" as many of these species. Furthermore, most of the vast acreage planted to this productive and nutritious grass is managed agronomically as tame pasture and/or a hay crop. Nonetheless, smooth bromegrass is frequently seeded and managed as an introduced range plant. On more fertile soils smooth brome has potential alternatively as a tame pasture or range grass depending on economic factors ranging from price of fertilizer and availability of labor to value of hay. Smooth brome is extremely useful in reclamation of drastically disturbed land such as stabilization of road cuts and establishment of vegetative cover on mine spoils. Smooth brome has naturalized and become a common species in northern portions of the the Western Range especially as an understory species in forests where it is associated with other introduced European grasses such as timothy and orchardgrass, and domesticated legumes like white clovef (Trifolium repens).
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.
58. Close-up of the smooth brome seeding seen in the previous view- This reseeded range has a bunchgrass structure in contrast to the usual sod form when smooth bromegrass is managed as domestic pasture and hay field.Harvey Valley, California. July, late vernal-early estival aspect.
59. 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.
60. 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.
61. Properly managed smooth brome- A pasture of smooth brome under proper (wise use) management in the Northern Great Plains. Commercial cows and calves (Red Angus and some black baldies) were grazing this single-species stand (monoculture) of Bromus inermis, almost unquestionably the single most important agronomic (tame= introduced= domestic) forage crop in the more mesic (especially, subhumid) Central and Northern Great Plains. Viewers should note the amount of herbage (biomass) left standing and the absence (more-or-less) of spotgrazing and wolf plants. There were almost no plants of weedy annual bromes such as the regionally common Japanese chess (Bromus japonicus) or of the naturalized Kentucky bluegrass (Poa pratensis). Other would-be interlopers such as weedy dicototyledons were "conspicuous by their absence". It was obvious or self-evident that this pasture had been managed correctly for many years. (The only other possibility was that the pasture was a new seedling just now subjected to grazing; a was highly unlikely situation.)
Unfortunately, smooth brome, (a highly productive, Eurasian, perennial grass) continues to be abused by overstocking and, thus ultimately, converstion to a grazing disturbance climax of Kentucky bluegrass. This latter species (another Eurasian festucoid grass) is also extremely valuable as a pasture grass, but Kentucky bluegrass is not a hay crop species. Furthermore, it is much less productive even as a pasture (not to mention as a hay crop) species than smooth bromegrass. Most importantly, from standpoint of stand maintenance smooth brome is much more palatable and much less tolerant of grazing abuse than is Kentucky bluegrass even though both agronomic species are "strongly rhizomatous" (McGregor et al., 1986). Under the far too widespread practice of overgrazing, pastures planted to smooth brome become degraded Kentucky bluegrass patches.
These two slides and the one immediately showed premier management of pasture and cattle. These photographs showed a "success story" when growers play by the rules or game play imposed by Mother Nature. This producer got it right.
Walworth County, South Dakota. Late June; grain in milk to soft-dough stages. Northwestern Great Plains-Southern Missouri Coteau Slope Ecosystem 42f (Bryce et al., Undated).
62. Good grass management= good cattle husbandry=profitable beef production- High-quality herd of Red Angus cows and calves on a pasture of smooth bromegrass that was being properly grazed: correct stocking rate for grazing capacity of this tame pasture. Much--in some areas, most--of the Northern Great Plains smooth brome stands have been abused through overgrazing and converted into disturbance climaxes of Kentucky bluegrass. Such was definitely not the case in this smooth bromegrass pasture that was being managed by the principles of proper grazing management. (It would probably be a safe bet that the soil was being fertilized "to test".
Good husbandry is sound business practice; in this case, of land, forage crop, and cattle. A well-earned tip-of-the-hat to this wise steward who was true to that in which he was entrusted: "Well done, thou good and faithful servant" (Matthew 25:21).
Walworth County, South Dakota. Late June; grain in milk to soft-dough stages. Northwestern Great Plains-Southern Missouri Coteau Slope Ecosystem 42f (Bryce et al., Undated).
Location note: an example of a smooth bromegrass pasture in this same general region, but that was mismanaged by overgrazing and converted into a Kentucky bluegrass disclimax was shown below. That example of a (former) pasture of smooth brome stood in stark contrast to the above example of proper pasture management.
A short section that could have gone several places in this chapter: shown immediately below were three slides-caption sets of a naturalized range plant "mix" that could have been located at various points in this chapter as well as in the one entitled, Introduced Forages-Legumes. Given that smooth bromegrass was the predominant plant species, the lesson and example of this range mixture was situated here. Anyplace, it was still fine forage.
63. Three-way range mix- A combination of 1) initial seeding (by grass or forage drill) and 2) subsequent self-seeding established this range plant community made up of three introduced (and, later, naturalized) forage species: smooth bromegrass, crested wheatgrass (Agropyron cristatum) and alfalfa. Naturalized range vegetation seen in these three photographs was at the edge of an interstate highway and an adjoining allotment of human-modified grassland in the unglaciated Northern Great Plains. With grains and seeds arising from plants seeded along berms and ditches of this major thoroughfare (and, probably, later from transport by motor vehicles along it ) as well as wind, wildlife and livestock, water, etc.this grass-legume mixture established and spread outward from the transportation corridor into mixed prairie that had been degraded by the usual causes (ie. overgrazing, cessation of fire, perhaps tillage, dust storms).
There were some plants of such native grasses as western wheatgrass (Agropyron smithii), thickspike wheatgrass (A. dasystachyum), needle-and-thread (Stipa comata), green needlegrass (S. virida), blue grama (Bouteloua gracilis), and buffalograss (Buchloe dactyloides) as well as naturalized annuals like Japanese brome (Bromus japonicus) and cheatgrass (B. tectorum) plus the naturalized perennial, Kentucky bluegrass. There were also plants of the naturalized yellow sweet-clover (Melilotus officinalis). All of these natives and naturalized exotics combined made up only a fraction of herbage yield (biomass), cover, and general abundance of the three main agronomic forages.
This three-way naturalized range mixtue (compliments of the whiteman) had successfully replaced the severely depleted native Northern Mixed Prairie. This grass-legume combination was included here to illustrate how plant communities comprised largely of naturalized alien species of plants can supplant the potential natural (climax) range vegetation. Maintenance of such a mixture under grazing regimes that maximize profit for livestock might well be problematic. The range plant community seen here was part of cattle allotment and appeared to have been grazed the previous year (no cattle were present at time of photographs).
Buffalo Gap National Grasslands (Interstate 90 right-of-way), Pennington County, south Dakota. Mid-June (late spring with vrious phenological stages in the major introduced forage species). Units of potential natural vegetation (climax) were not appropriate.Northwestern Great Plains- River Breaks (of Cheyenne River) Ecoregion 43c (Bryce et al.; Undated).
64. Line-up of usual suspects- Smooth bromegrass (taller grass with panicles), crested wheatgrass (shorter grass shoots with spikes still in the boots), and alfalfa (bush-appearing plants with broader, trifoliate leaves)--three whiteman-introduced forages--overwhelmed native grass species of Northern Mixed Prairie resulting in this disturbance climax. Under existing management conditions this man-made (and admittedly simple) plant community consistuted the range vegetation.
This was a naturalized range version of the grass-legume field crop of smooth brome and domestic alfalfa that has become the standard tame pasture and hay mixture from subhumid to semiarid portions of the Great Plains eastward into the Corn-Soybean Belt of the Upper Midwest (in some areas as far east as Ohio). The smooth brome-alfalfa mixture is difficult to maintain. When the standard smooth brome-alfalfa mixture is utilized as pasture livestock (especially sheep) tend to graze out the more palatable and less grazing-tolerant alfalfa. To overcome (at least partially) this threat to the legume soil fertility and pH must be closely monitored and skillfully manipulated (phosphorus and calcium nutrition of alfalfa become very critical). When the smooth brome-alfalfa crop is used for hay production the nitrogen-fixing alfalfa has the advantage over smooth brome so that nitrogen levels favoring the grass become more important. On the other hand, alfalfa is a short-lived perennial that usually does not regenerate (alleochemicals produced by parent alfalfa plants retard establishment of alfalfa seedlings which in combination with short longevity of existing adult alfalfa plants effectively prevents perpetuation of the legume).
In this three-way-combination forage crop, crested wheatgrass added another variable and at first flush appeared to make maintenance of the half-again increased mixture even more problematic. When alfalfa plants are as widely dispersed as in this man-made plant community regeneration of alfalfa is more likely so that the short-lived legume can maintain itself in the plant community, at least at the widely spaced density and dispersal pattern of this range vegetation.
Native grasses--both cool- and warm-season species had been effectively excluded from this naturalized range plant community. The climax wheatgrass-needlegrass-gramagrass mixed prairie had been degraded by standard mismanagement (overgrazing, elimination of natural fire, perhaps tillage) under the heavy hand and ignorance of Anglo-European man such that introduced plant species had the upper hand so to speak. Under present conditions (including everything from climate to livestock grazing of this allotment) the smooth brome-crested wheatgrass-alfalfa community appeared permanent, a disturbance climax.
This naturalized range provided a beneficial forage crop, especially when wisely used in conjuction with the native range of Northern Mixed Prairie. The three-way mixture of two introduced, cool-season grasses and a nodulated legume added biological diversity. Yellow sweet-clover, a biennial, and Japanese brome and downy brome or cheatgrass, two annual (and Eurasian) grasses, also added biodiversity. These three naturalized exotics were so widely scattered and also so overwhelmed that they added little by way of herbage or foliar cover.
Buffalo Gap National Grasslands (Interstate 90 right-of-way), Pennington County, south Dakota. Mid-June (late spring with vrious phenological stages in the major introduced forage species). Units of potential natural vegetation (climax) were not appropriate.Northwestern Great Plains- River Breaks (of Cheyenne River) Ecoregion 43c (Bryce et al.; Undated).
65. Three-species sward- Details of a classic grass-grass-legume (smooth brome-crested wheatgrass-alfalfa) mixture that developed and maintained itself as naturalized range in the Northern Great Plains. It was explained in the two preceding captions that this grass-grass-legume mixture was range vegetation more than a simple community of tame (domestic) pasture. It was not only the presence of a third forage species, but also a function of several variables (including the sparse density and dispersion of alfalfa, an allelopathic and short-lived perennial, critical balance of soil nutrients with correct pH, and preferences of grazing animals) that increased the difficulty of maintaining a mixture of forage grasses and legumes.
These three "photoplots" showed the variation in species composition and structure of this simple range plant community over comparatively short distances. They also showed the predominance of smooth bromegrass (first among the three dominants) and the relatively wide spacing (low density and cover) of alfalfa , but at this point in the growing season crested wheatgrass (the spikes of which were still deep down in the boots) was underrepresented in these photographic presentations.
It was also emphasized that this range vegetation had not been grazed this year (current growing season) though it appeared to have been grazed by cattle the previous year. Foliar cover and overall size of the more palatable alfalfa was undoubtedly greater in comparison to that of the two major grasses under the exiting harvest schedule (grazing management). Given that this range appeared to have been grazed in the previous growing season (as evidenced by weathered dung and deep hoof prints in mud) it had to be assumed that alfalfa was maintaining itself in this plant community.
This smooth brome-crested wheatgrass-alfalfa at its ideal was an example of introduced forage species comprising naturalized range. As a domestic grass-legume mixture that "went wild" or had "gone native" this plant community served as an example of extremely productive grazing land that was used and maintained with extensive, low-imput management.
Buffalo Gap National Grasslands (Interstate 90 right-of-way), Pennington County, south Dakota. Mid-June (late spring with vrious phenological stages in the major introduced forage species). Units of potential natural vegetation (climax) were not appropriate.Northwestern Great Plains- River Breaks (of Cheyenne River) Ecoregion 43c (Bryce et al.; Undated).
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).
66. 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.
67. Stand of smooth bromegrass at peak standing crop (just after anthesis)- This old field reseeded to smooth brome is an example of the tremendous herbage yield and soil conservation that is possible with this introduced European tame pasture grass. Gallatin County, Montana. June.
68. Panicle of Smooth Brome- Large inflorescences of smooth brome. Gallatin County, Montana. June.
69. Several anicles of smooth brome- Panicles of smooth brome at peak anthesis. Ottawa County, Oklahoma. Late May.
70. Sexy smooth brome- Flowering in smooth brome with both anthers and stigmas fully exerted. The anthers in these spikelets has already senesced (died) and were in process of being shed.This was an example of protandy, the flowering pattern or sequence in which anthers mature (and, in this instance, die) before appearance of stigmas. This insured cross-pollination among different plants (genotypes) because the protanndrous pattern took place in all florets of an entire panicle (each plant), but at different days within the brief flowering period in this local stand of smooth brome.
Ottawa County, Oklahoma. Late May, immediate-post anthesis.
71. As close as this photographer could get- Living, pollen-producing anthers in smooth brome. Photographs of organs were taken on standing shoots in the field. Ottawa County, Oklahoma. Late May.
Ecological note: over much of the Northern Great Plains there are hundreds of thousands of acres of native true and mixed prairies that were converted (through tillage and direct seeding, overgrazing, modification of fire regimes, road-building, farming of row crops and small grains, etc. ) into anthropogenic (man-made) grasslands made up of 1) varying mixes of Kentucky bluegrass (Poa pratensis) and smooth brome (Bromus inermis) and, at the ultimate state of disturbance (primarily overgrazing) climax, 2) monocultures of Kentucky bluegrass. These two artificial (human-induced) range types became naturalized types much like the California annual grassland type and cheatgrass or downy brome (Bromus tectorum) annual ranges in the Intermountain Region. All such type conversions (natural grasslands converted into grasslands of domestic plant species through human action) are disturbance climaxes.
Examples of disclimaxes of Kentucky bluegrass were included below.
72. 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.
73. 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.
74. Spikelets of rescuegrass- Rescuegrass is usually a prolific grain-producer. This is the necessary condition for most short-lived rennials. 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.
75. Tall fescue (Festuca arundinacea) pasture- "Tall fescue is the predominant cool-season, perennial grass in the United States" (Buckner, R.C. and L. P. Bush, 1979, p. xi). Tall fescue includes numerous cultivars (eg. Kentucky 31, Alta) and has been one of the most studied, domesticated, and planted tame pasture grasses in the United States. It is perhaps the agronomic forage species, particularly among perennials, most persistent under abuse and poor management. It is therefore extremely popular among cattlemen because it is easy to establish and to maintain under all manner of management (ie. it is hard for them to kill it out by overgrazing, failure to fertilize, and so on). Most tall fescue cultivars are infected with the endophytic (an endophyte is any plant such as a fungus or algae that lives within another plant) fungus (Acremonium coenophialum) which makes the host fescue quite resistant to insect pests and perhaps some diseases. Some agronomists have proposed that the endophytic fungus-fescue mutualism contributes to fescue's ability to recover from grazing abuse, drought, etc. The cool-season tall fescue is a drought avoider because in most areas where it is grown winter drought is not a major threat and tall fescue completes it's annual growth cycle by late spring and avoids drought through summer dormancy. Overall then tall fescue is remarkably adapted to survival and to moderate production with minimal management yet capable of considerably more production with optimal agronomic inputs.
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 trade-offs and tall fescue continues to be planted and maintained as the major cool-season permanent pasture grass in the USA. In this vien, the definitive treatment (as of this writing) of tall fescue as an agronomic crop is Fribourg et al. (2009).
On the Southwest Missouri Research Station in the Sprinfield Plateau portion of the Ozark Plateau physiographic province the standard stocking rate for tall fescue managed as permanent pasture (managed more intensively including with liming and fertilization as required) has been three to four acres yearlong (in a typical year) for a 900 to 1100 pound Aberdeen Angus or black baldy brood cow (David Cope, superintendent of the Southwest Missouri Research Branch Station; personal communication). Larger (greater metabolic body size) brood cows such as crossbred cows with Continental beef breeding and/or those of considerably greater milk production (not to mention their larger calves) would require considerably more pasture (more acres of tall fescue per cow).
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.
76. Tall fescue plant at full flower to seed-set stage- Although tall fescue was developed for a pasture and not a hay grass it is almost as widely used for hay as for grazing. Most hay-makers (balers more than growers) in many regions of the upland South (eg. Ozark Plateau) bale tall fescue at the stage seen here to get maximum numbers of bales (dry matter more than nutritive value). To reduce labor and storage costs most of this fully mature tall fescue hay is put up in "big round bales" and stored outside uncovered where in the humid precipitation zone the outer third of the bale is nutritionally worthless. As with pasture the emphasis is on "the cheap". Tall fescue is not quality forage, but it is an amazing pasture grass. It propelled Missouri to the number two cow/calf state behind the giant Texas.
Ottawa County, Oklahoma. Mid-May; early onset of anthesis.
77. One plant and a "gazillion" pollen grains- A single cespitose plant of tall fescue at phenological stage of maximum pollination. This plant literally had a yellowish hue as a result of thousands of exerted anthers dangling from its panicles. There was one panicle per tiller and several hundred tillers arising from the poraxis of this one genetic individual that approached yard-height stature. At this phenological stage and mature plant size herbage of tall fescue provides poor-quality forage. Plants of tall fescue should not be permitted to reach such rank size and with so many tillers advancing to sexual maturity. Biomass from wolf plants like tthe one shown here is very unpalatable, even in an open bale on snow-covered ground.
This individual was growing in the outside fencerow and was presented here to show students an example of mature size to which tall fescue can attain. Likewise, when this large plant was viewed with the next slide-caption sets viewers can understand why tall fescue causes hay fever and related allergies.
Ottawa County, Oklahoma. Late May; peak anthesis.
78. How tall fescue "conquers"- A typical panicle on one tiller of tall fescue (first photograph) and one floral unit of several spikelets (several florets per spikelets) on one part of one branch of a tall fescue panicle (second photograph). These two photographs should demostrate to viewers the amazing potential for sexual reproductive in tall fescue. This cool-season, perennial Eurasian grass became a naturalized range and pasture gras primarily through its seemingly limitless capacity for seed production in combination with its tolerance to mismanagement (abuses of overgrazing and under- or non-fertilization). This latter phenomenon is, as was explained above, due to the mutualism of tall fescue and an endophytic fungus.
Tall fescue is an invaluable forage grass because it produces moderately high yields of herbage even though this is inferior quality forage. Tall fescue enabled areas of "poor land" such as the Ozark Plateau (where these photographs were taken) to become rich in beef production--or "cow-poor" as the case may be--and at the same time to restore organic matter and fertility to "cropped-out" soil. Thus was the human standard of living and quality of life increased through crops of tall fescue.
Tall fescue is, however, also a dreaded weed on tallgrass prairie, especially bluestem-Indiangrass cattle ranges. In this regard tall fescue puts other weeds like Johnsongrass on par with "Ned in the third reader" while tall fescue threatens the integrity (or the very existence) of native grasslands. This threat and, all too often, ruination of prairies and herbaceous understories of oak-hickory-tallgrass savannahs is reached via the tremendous grain production of tall fescue.
Tall fescue is such a prolific seed producer that in a relatively few years (or, perhaps, a few decades) a field, pasture, or range having some widely scattered plants of tall fescue becomes a single-species stand (or nearly so) of tall fescue. Other species of grasses and forbs, weeds, are crowded out by the fiercely competitive and aggressivly propagating tall fescue. This self-seeding feature, and combined with asexual reproduction of tillers (most of which will bear heavy grain crops), is ideal when there are thin stands of tall fescue as the intended forage crop. Even if a poor stand was established initially there can be a complete and vigerous stand of tall fescue and an acceptable permanent pasture in short order (maybe five years or so).
The converse applies to ranges of tallgrass prairie and open grazable forests. Good pasturemen utilize the remarkable reproductive capacity of tall fescue or, alternatively, they limit impact of this feature to the advantage of their operations and the natural resources on which these firms are based.
Ottawa County, Oklahoma. Late May; peak anthesis.
79. Pollinating its world- Slides of tall fescue spikelets at maximum anthesis.Florets number from to per spikelet of tall fescue There are between three to six, rarely nine, florets per spikelet of tall fescue.
Ottawa County, Oklahoma. Late May; peak-bloom phenological stage.
80. Invasion of tall fescue on tallgrass prairie range- Overgrazing of this big bluestem-dominated range (Indiangrass, upland switchgrass, prairie dropseed, Canada wildrye, eastern gamagrass, leadplant, Illinois bundleflower were other major species) allowed establishment of Kentucky 31 tall fescue by seed dispersal from adjoing pastures and highway right-of-way that ultimately resulted in competitive displacement (under combined-- and probably synergestic --influence of overgrazing) of native species. Net result was a grassland type conversion: conversion of range (native range) into tame pasture (or perhaps viewed as naturalized range): SRM 601, the larger and more general type, or 710 converted into SRM 804 by improper range management. Woodson County, Kansas. June.
81. 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.
Organization note: slides showing cold (winter) hardiness (or tolerance, whichever) of tall fescue as compared to orchardgrass (Dactylis glomerata) in side-by-side plants was presented below at the end of the treatment of orchardgrass.
The Society for Range Management (Shiflet, 1994, ps. 112-113) named and described as a rangeland cover type, Tall Fescue, SRM804. Beyond doubt tall fescue (Festuca arundinacea) is (and has been for a half-century) one of the most important introduced (= agronomic, tame, domesticated) forage grasses in North America (Fribourg and Hannaway, 2009). This does not imply anything with regard to tall fescue being a naturalized range plant with its own rangeland cover type as presented in Shiflet (1994, ps. 112-113).
Tall fescue deserves a place on par--or maybe quite a bit higher--with other agronomic Eurasian grasses including those in the Festuceae tribe such as smooth brome (covered above) or orchardgrass (Dactylis golmerata) and Kentucky bluegrass (Poa pratensis) which were covered below. Those species did not have their own rangeland cover type as did tall fescue (Shiflet, 1994, ps. 112-113). That tall fescue, orchardgrass, smooth brome, Kentucky bluegrass and a myriad of other Eurasian grasses have naturalized in North America to the point of having individual plants or even local populations of their species surviving in scattered locations and microsites is not a contested point. That is profoundly different, and of an entirely lower magnitude of scale regarding adaptation, from surviving via naturalization to the extent as to form a range cover type of its own.
In this short section, your author used his experience, extending over 50 years, with tall fescue in the Cherokee Prairie of the Central Lowlands and Springfield Plateau to document loss of tall fescue in a drought that ranged from Severe to Extraordinary (Palmer Index) over a period of five years involving parts of or entire tall fescue-growing seasons.
This "botanical interlude" served as more of an introduction to native species of pioneer plants that replaced tall fescue when Clementsian denudation set the sere of tallgrass prairie--that had been converted to tall fescue pastures and hay fields--back to initiation of secondary plant succession. This proved to be an on-going experiment, but one in which there was some, though not much, evidence to indicate that tall fescue was recovering from drought damage.
In this author's ecological judgment and experenicial opinion, this "drought-induced experiment" was conclusive proof that on many soils and under some growing/management conditions tall fescue is not adapted enough (not thoroughly naturalized to the point) that tall fescue is deserving of a designated rangeland cover type. In fact, other introduced forage grasses, especially common bermudagrass (Cynodon dactyledon) and Johnsongrass (Sorghum halapense), were found to be much more apt to survive (even increase) under prolongued drought than was tall fescue. Much, if not most of this increase, particularily of bermudagrass and Johnsongrass, was due to loss of competition from tall fescue.
In this "heartland" area of the Central Lowlands and Ozark Plateau physiographic provinces--the intersecting Four State Area of Missouri, Kansas, Oklahoma, and Arkansas--it had been a much talked about question for about a half-century among farmers and stockmen as to whether tall fescue or bermudagrass would be more likely to survive a devestating drought like those of the 1930s and 1950s. Both bermudagrass and tall fescue had been introduced into this area in the late 1950s and especially the early 1960s. Johnsongrass had been introduced decades earlier and had already thoroughly naturalized and become established. Johnsongrass was (and still is) commonly regrded as more of a weed of roadsides and tilled fields. Value of Johnsongrass was recognized as an emergency forage, especially for hay, but it was feared for its acute livestock poisoning (more prussic acid than nitrate toxicity though both occurred) that struck rapidly "without warning". For this reason Johnsongrass was not planted. It seeded and spread quite well "on its own". (Problems of toxicity with tall fescue came much later; this was a chronic toxicity that was less dramatic and typically did not result in animal deaths.)
Following introduction of common bermudagrass (both by seeding and, mostly, sprigging) and tall fescue shortly after the drought of the 1950s there had not been a prolonged (multi-year) drought or even two consecutive harsh drought years (two back-to-back droughts extending throughout most of the warm-growing, the frost-free growing, season). Then beginning near the end of the first decade of the Twenty-First Century (2008 or 2009 depending on exact location) the equivalent of the 1930s and 1950s drought set in. From then through 2013 drought rating varied locally, but ranged from Severe to Extraordinary (Palmer Scale) Drought. Technically this multi-year, prolonged drought extended throughout 2014 which was locally a Severe Drought, but a spring-summer season of cool temperatures puctuated with fortituous showers minimized drought impacts.
This latter set of conditions was a mute point for purpose of this discussion because stands of tall fescue had been wiped out at least two years earlier (by 2011 or 2012 depending on local conditions). Some growers reseeded tall fescue; some of these seedings resulted in re-establishment while others produced nothing. Concurrently, Johnsongrass survived quite well in spite of heavy infestations of differential grasshoppers (Melanoplus differentialis) which had an obvious preference for Johnsongrass over about everything else, including native grasses like big bluestem (Andropogon gerardii). Grasshoppers grazed selectively and severly overutilized patches of Johnsongrass which had reached the rank-growth vegetative stage and grain production before 'hopper populations peaked at development of adult insects. Most Johnsongrass was "riddled" by feeding grasshoppers. Johnsongrass that grew along roadsides was frequently mowed, often to near ground surface with no living shoots remaining aboveground. Never mind, Johnsongrass survived, even rebounding quickly. (This topic was covered below in this chapter, the Johnsongrass section.)
At the same time, in this multi-year drought common bermudagrass spread slowly even with cattle grazing and/or hay harvest. In fact, bermudagrass did well in overmowed lawns (actually these were yards as there was no grass management practice --other than overmowing). This author observed numerous cases where bermudagrass spread and did better in yards during the extended (multi-year) drought than in wet summers because there was much less hairy crabgrass (Digitria sanguinalis), the main competitor of bermudagrass in overmowed yards, during this drought. (Apparently, there was not adequate rainfall to germinte seeds of the annual crabgrass whereas the perennial bermudagrass regrew from rhizomes and some lower-growing stolons passed over with overmowing.)
By contrast, tall fescue died like differential grasshopers smashed in roads by passing vehicles. Most telling of all was the phenomen of Johnsongrass "springing up like weeds" (pun intended) in what had been nearly "solid stands" of tall fescue. Less dramatically, common bermudagrass advanced slowly though steadily into bare areas that previously had been the home of tall fescue. With partial alleviation of drought in 2014 some tall fescue recovered--another partial condition. Full recovery of tall fescue had not yet taken place with moderation of the on-goining drought.
So the half-century question of whether bermudagrass or tall fescue was more likely to survive long-term drought had been answered, and answered dramatically. Tall fescue was observed to be highly inferior to common bermndagrass for survival in multi-year droughts. It should be specified that these drought conditions included the cool-seasons as well as the warm-seasons (frost-free periods) of these drought years. These were not summer droughts that could benefit cool-season species or autumn-winter droughts that would competitively benefit warm-season species. Instead, the Grim Reaper's drought scythe cut across all climatic seasons throughout most of this five- to six- year drought. Again, it required only three or four consecutive drought years to wipe out stands of tall fescue whereas up to six consecutive years had not resulted in loss of bermudagrass or Johnsongrass.
The decades-long debate among growers had always hinged on the question as to whether the warm-season bermudagrass (and Johnsongrass) could survive a summer drought with their drought tolerance or whether tall fescue could survive a summer drought due to its drought avoidance. Noone doubted the superiority of warm-season grasses to grow (if at all) during warm-season drought, but producers also knew that the cool-season tall fescue would be dormant (or mostly so) during the peak time period of summer drought. Again, would 1) ability to withstand drought impacts while growing or 2) growing in the cool-season and being dormant in the warm-season so as to avoid drought (ie. an annual growth cycle of dormancy during summer drought) be the superior growth pattern, the superior survival "strategy" or "allocation of resources" in words of theoretical ecologists. This argument missed the point of, What about year-long drought, several years in a row? The obvious likely drought condition was somehow missed, at least by most who raised this question. Now we know.
Mr. David Cope, superintendent of the Southwest Missouri Research Branch Station, told this author that he felt like on some rocky hillsides in the Springfield Plateau tall fescue (most of it overseeded on such stony habitats) will "be there to the end of time". Maybe so, but (as the slides presented immediately below attest) on some arable land in the Springfield Plateau where mowing or grazing can be closer (remove more of the grass shoot biomass) tall fescue was, in effect, eradicated (locally exterminated) during a multi-year drought varying from Severe to Extreme to, eventually for better part of one warm-growing season, Extraordinary Drought. Mr. Cope specified that tall fescue should not be mowed to a stubble height shorter than four or five inches. The example of a destroyed--by a combination of mowing to ground level, heavy grazing by sheep, record low winter temperatures , and perhaps other factors--"wiped out" hay field of tall fescue that had been established for a half century illustrated that tall fescue is not necessary a permanent forage crop "to the end of time". Now we know.
We also known that when the millions (tens of millions) of dollars and scientist years (in some instances, lifetimes) of research devoted to tall fescue and with the fortunes of cattlemen being tied to production of tall fescue in states like Missouri or Kentucky are considered, the conclusion cannot be avoided or the statement denied that much of this effort would have been better, more wisely invested in warm-season species like bermudagrass than the more toxic, and less drought-adapted tall fescue. The travesty was not the commitment in research and development given over to tall fescue, but rather that there was nothing even apporaching such commitment to introduced warm-season grass species. At least that was the case in many states. Other states such as Oklahoma used public research funding for a balanced approach, a combination of warm-season and cool-season forage species, to permanent agronomic pasture (and range) management.
Inevitably, experience trumps speculation, promotion, even theory. Experiment prevails over argument. So where was the experiment with warm-season agronomic grasses in states like Missouri, Tennessee, and Kentucky? That is the scandelous and revealing question that was raised by the the first major and prolonged drought following introduction of species like tall fescue.
** See below under the Eragrostoideae and Andropogoneae tribes the clear superiority of common bermudagrass (Cynodon dactyledon) and Johnsongrass (Sorghum halapense) over tall fescue for drought adaptation, survival to heavy defoliation, and herbage production potential during multi-year droughts in Severe to Extraordinary Drought ratings.
82. Once in a half century stand- An across-the-field view of a dense population of wooly, prairie, or Pursh's plantain (Plantago patagonia= P. purshii) with some large-bracted plantain (P. aristaus), these two together forming an upper herbaceous layer, and a population of rough buttonweed (Diodia teres) that developed into a lower herbaceous layer (and which later in the warm-growing season became the sole living herbaceous layer). This plant community that was comprised of native, annual, pioneering forb species, replaced a medicore stand of tall fescue that had persisted on this field for fifty years. A prolongued drought that extended over parts of six consecutive and most of the growing seasons for five consecutive years in conjuction with over-mowing, overuse, plus absence of lime and fertiliztion application, wiped out this fifty year-old stand of tall fescue. The exact abuse of the tall fescue along with years of, plus duration of, drought were detailed in the above introduction and in longer captions below.
The only tall fescue that remained at time of these photographs was in shaded fencerows, especially where tall fescue grew close enough to fences that it was not mowed to ground level with a disk mower. Almost all of the former field of tall fescue looked like the scenes of plant life seen here. Total loss of the forage crop.
At time of the photographs presented here the drought had reached high stages of Severe Drought to beginning stages of Extraordinary Drought (as measured by the Palmer Drought Severity Index).
Ottawa County, Oklahoma. Early-July; mature plant, ripening fruit phenological stage.
83. From the top down- Topdown view into a "solid" or "nearly pure" stand (a population) of wooly plantain in an upper herbaceous layer and with an equally "nearly pure" stand of rough buttonweed that formed a lower herbaceous layer on what for fifty years had been a field of tall fescue used primarily for hay and secondarily for seasonal pasture. A prolongued drought that extended over almost all of the growing season for six consecutive years, coupled with human abusive management resulted in what amounted to a total loss of this fifty year-old stand of tall fescue. By the time these slides were taken the protracted drought measured at the top of the Severe Drought category to early stages of Extraordinary Drought ( Palmer Drought Severity Scale).
Ottawa County, Oklahoma. Early-July; mature plant, ripening fruit phenological stage.
A descriptive and explanatory note on botanical composition of the degraded (devestated is a more apt--though less objective sounding--adjective) tall fescue stand was inserted here. The pioneer plant community (the initial seral stage of secondary plant succession) of the lost pasture consisted not so much as a homogenous stand of the three major native, annual, pioneering forbs but rather of patches of plant life dominated by first one and then another of these three colonizing forb species. This was shown in the preceding two slides that featured wooly or Pursh's plantain and the next two slide-caption sets of large-bracted plantain. Rough buttonweed came into dominance later in the summer season and replaced the two Plantago species in many of the patches they dominated previously in spring and early summer. In other local areas rough buttonweed did not grow to dominance and, instead, these remained patches of dead plants of the two Plantago species. Completely missing were any plants of tall fescue.
84. Drought replacement- A fifty-year old stand of tall fescue completely eliminated by four years of continuous drought that ranged in severity from Severe to Extraordinary (Palmer Scale or Index) and with old-field (secondary) plant succession set back to the initial or pioneer stage. Pioneer plants in this field were large-bracted plantain (Plantago aristaus), wooly or prairie plantain (P. patagonia= P. purshii), and rough buttonweed (Diodia teres). The green color was cover of woolly plantain and rough buttonweed whereas the golden amber was large-bracted plantain.
This stand of tall fescue had been abused over a span of fifty years. Except for application of lime and fertilization to soil test at time of seeding and one application of lime subsequently there had been no soil management. For about 35 years it had been mowed for hay once a year (usually in late spring or early summer; the wrong time for quality hay, but after grain shatter). Then for about 15 years prior to stand loss this field had been mowed twice: once in late spring and then again in mid to late summer.
This view was in early summer of 2013. Spring and summer of 2009 was a drought reaching Severe Drought status for a brief period. It was mowed for hay during that time. In autumn 2009 through spring 2010 re-growing tall fescue was overutilized by sheep. It was mowed twice for hay in spring-summer 2010, another drought year. Winter of 2011 was aone of record cold. The record low temperature for the town of Seneca, Missouri (three-fourths of a mile distant) was -20 deegrees Fahrenheit (Guinan, February, 2011). Temperatures were so low that the native shrub, spicebush (Lindera benzoin) a half mile from this stand was completely topkilled and had to regrow from rhizomes and rootcrowns the following growing season. Tall fescue survived the record low winter temperatures. It did not make much growth in the following spring-summer season (2011), but it was mowed for hay twice. It was mowed for hay twice during spring and summer 2012. The mowing over the last decade or so had been with disk-mowers set so low that patches of bare soil frequently remained after mowing.
In summer 2013 the culmination of this long-term mismanagement (yea, abuse) was shown in the above photograph. The year of 2013 was the worst year of the drought now in its fourth or fifth year, depending on criteria used to measure and determine drought.
The only tall fescue that remained in this field was in tree shade along fencerows. Some of these shade survivors had been subjected to all treatments: mowing and sheep grazing as well as drought and low winter temperatures. Some fencerow tall fescue that had not been grazed or mowed survived, and this included some plants that had benefit of shade and some plants that had no shade. The only tall fescue that had been grazed (overused) and mowed (overmowed; overused) was in the shade. Some plants of tall fescue that had been neither grazed nor mowed died anyway, but these undefoliated plants were all in full-sun and never had shade. Empherically, it was obvious that shade was the single most important observed factor related to survival of tall fescue.
With this confounding it was impossible to determine which factor--drought, absence of shade, close-grazing, low-mowing, extreme winter temperatures, or even absence of proper soil management--or combination of factors led to death of tall fescue and loss of the stand.
At this juncture, the conclusion could be drawn that perhaps with proper management the stand might have persisted. One cannot prove a negative. Absence of evidence is not evidence for or against. Several possibilities exist, but none of these can be proven. There is no conclusive evidence.
That is irrelevant. It is irrelevant because other grass species growing in this field did not die out (or were not killed out) in spite of abuse. Plants of warm-season species including common bermudagrass, splitbeard bluestem (Andropogon ternarius), switchgrass (Panicum virgatum), and purpletop (Tridens flavus) were subjected to the same treatments as tall fescue--except for absence of grazing during their growing seasons. Other grass species in this field included dwarf rosette panicgrass (Panicum depauperatum) and broomsedge bluestem (Andropogon virginicus). Dwarf rosette panicgrass is a cool-season species while broomsedge bluestem, though primarily a warm-season species, retains considerable numbers of live shoots, along with last-season's dead shoots, at base of the cespitose plant. These two native grasses were thus subjected to grazing during much of their growing seasson the same as with tall fescue. It was observed that Dorper sheep grazing this pasture heavily grazed these two green, growing, native species as they did tall fescue.
In the year of 2014 (still a drought year) following the year of this photograph of the devestated stand (July 2013, and the worst year of the multi-year drought) dwarf rosette panic grass had recovered to point of producing grain-loaded, sexual shoots that measured over four feet in length. Broomsedge bluestem was still struggling to survive, but it had made noticeable recovery. The four warm-season species listed above had not only recovered, but had expanded their coverage. (The disk-mowers returned in 2014 to leave patches of bare soil so fate of the long-shoot switchgrass remained unknown.) Purpletop made a post-mowing bumper crop. Bermudagras slowly spread its cover. Tall fescue was gone.
Gone also was the bumper crop of the Plantago species and rough buttonweed. There was no doubt that plantain and buttonweed had produced massive crops of "live-forever" seeds that could await other opportune times to produce future bumper crops. The big question--the overarching question--was, Could tall fescue, with 50 years of seed crops, re-establish itself on this field? That is the ultimate measure of naturalization. Is tall fescue naturalized as, say, Johnsongrass is? Or is tall fescue merely an opportunistic species that can survive in good times yet be wiped out permanently by consecutive years of drought? If answer to the first question is Yes, then tall fescue is naturalized, including as a naturalized range grass. If the answer is No, then tall fescue is only a waif the same as cereal grains volunteering in a field or roadside. Tall fescue can only return to field seen above (which is representative of tall fescue stands established on many upland pastures of comparatively infertile soils) by self-seeding. Almost all pre-existing (adult) plants of tall fescue died during the drought, and with improper management practices (= abuse). Self-regenerating ability under natural conditions--including multi-year drought-- (ie. naturalization) of tall fescue depends upon its capacity to produce a viable seed (a persistant grain) bank in the soil from which it grew and reproduced.
The correct categorization of tall fescue as a range grass and, thus, tall fescue as constituting a rangeland cover type (Shiflet, 1994, ps.112-113) depends on its naturalization, the ability or capacity of tall fescue to recover from drought and re-establish itself following drought (and other natural disturbances such as extreme low winter temperatures) as does Johnsongrass and, even, common bermudagrass not to mention various crabgrass species and introduced annual legumes. Time will tell.
Note on content: the following slide-caption sets showed plants of pioneer species that grew in the wake of tall fescue annihilation and in the fourth (or fifth) year of prolongued drought. They are all native species. They were functioning as pioneers or first colonizers of denuded land. They were not weeds in this situation. These species were healers of land laid bare. They were Mother Nature's "first responders" to a natural--or was it natural?--disaster. Rangemen need to know the the pioneer plant species the same as the climax plant species. All species have an ecological niche--a place, a role in range and pasture ecosystems. Populations and vegetation change. Nature is dynamic.
"Pioneers! O pioneers!", Leaves of Grass, Walt Whitman- This pioneer seral stage had it seasonal societies (socies, the seral equivalent of the climax societies in Clementsian terminology) with two Plantago species (P. aristaus and P. patagonia= P. purshii) dominating the spring/early summer community (socies) and rough buttonweed or poor joe (Diodia teres) dominating the mid-summer/autumn community.
The author had observed this field with its stand of tall fescue since it was established in 1963 and cannot remember ever seeing P. patagonia= P. purshii anywhere in this area. He had observed an occasional plant of P. aristaus and a few patches (roughly less than 25 square yards) of Diodia teres in this field about 20 years earlier. This was one of those textbook cases of the situation in which a faarmer truthfully claims, "I never saw that stuff before and now its everywhere! What is it?"
Seed of these pioneer species--and they are native forbs not whiteman-introduced alien weeds--had evidently been in the soil seed bank for decades and, with elimination of cover and competition from tall fescue, their "moment in the sun" had arrived. And boy, did they exploit their rare opportunity!
It was rare indeed because the year following this pioneer stage of secondary (= old field) plant succession none of these three species other than a few small patches of rough buttonweed remained. The phenomenon of a plantain/rough buttonweed "outbreak" was like a swarm of bees or the mating of a queen wasp with a group of male inseminators: over, done with, and gone in a flash of Mother Nature's time, a rare phenomenon to be savored but for a moment.
85. An annual healer, bristles and all- Large-bracted plantain or bottlebrush Indianwheat (Plantago aristaus) growing as a dense stand in a field that, prior to a multi-year drought and mismanagement, had been made up mostly of tall fescue in the Ozark (Springfield) Plateau. Following extirpation (or essentially so) of tall fescue from this field large-braceted plantain became the dominant plant species over much of this former fifty year old stand of tall fescue. Overall associate species of the plant life (the pioneer or initial seral stage community) were wooly plantain and rough buttonweed at this late spring season, the spring/early summer community or socies. There were local patches where wooly plantain was dominant. In the mid-summer/autumn community or socies--and with the saprophytic generation dead and the sporophytes wainting for the next need for pioneer species--rough buttonweed ascended to seasonal dominance.
Large-bracted plantain does have conspicuous floral bracts, but it is one of the smaller Plantago species in this region. An outstanding treatment of this small, native, annual forb was provided by Tyrl (2008, ps. 416-417) who described its ecological as well as morphological features. The fruit of large-bracted plantain is a circumscissile capsule which according to Tyrl (2008, p. 417) is eaten by about all species of grassland birds including the greater prairie chicken (Tympanuchus cupido).
At maturity stage as seen here the senescing or dead shoots of bottlebrush Indianwheat turn a honey gold or orangish-amber. And, yes this was the color as captured by Fujichrome Provia 100F (of course the photographer would not vouch for this being the natural color the way he would have had the film been Kodachrome).
Plantago species are in their own family, Plantaginaceae (p,. 1380-1386; Smith, 1977, ps. 201-202; Diggs et al., 1999, ps. 882-886).
Ottawa County, Oklahoma. Early July; fruit-ripe/fruit-shatter stages.
86. Another healer, this one wooly- Wooly or prairie plantain (P. patagonia= P. purshii) growing on denuded land left practically bare when a fifty year-old stand of tall fescue in the Ozark Highlands (Springfield Plateau) died out in a multi-year drought under human-induced abuse. In the spring/early summer season of this initial stage of secondary succession wooly plantain was the overall associate species to large-bracted plantain although it was the dominant in patches (see slide above that introduced this section).
The author had observed the stand of tall fescue (since its establishment in 1963) on which these examples of wooly plantain grew and cannot ever recall seeing a single plant of P. patagonia on this field, or anywhere else in this area of the western Ozark (Springfield) Plateau. A few years prior to development of this pioneer or initial seral stage on a former stand of tall fescue your author had to photograph specimens of wooly plantain on the Central Plains Experimental Range in northeastern Colorado to be able to bring images of this species to viewers. A few years later there was an entire field of wooly plantain or wooly Indianwheat in the author's "own backyard".
Ottawa County, Oklahoma. Early July; immediate post-bloom phenological stage.
87. Other wooly views- Entire aboveground shoot, sexual shoot with inflorescence, and close-up of inflorescence (first, second, and third slide, respectively) of wooly plantain as the overall associate pioneer species in the initial seral stage of old field (secondary) plant succession on a field that had been a fifty year-old stand of tall fescue in the Springfield Plateau until it died out (or was killed off) by a mulit-year drought and long-term abusive management.
This condition was described perfectly in Weeds of the West (Whitson et al., 1992, p. 407): "Wooly plantin is a common invader of abused native ranges". Whitson et al. (1992, p. 407) followed with: "This native warm-season forb is not a highly competitive species with native rangeland perennials". Nor, apparently with introduced perennials like tall fescue. The noncompetitive annual species has to be invited. Mismanaging man extends the invitation.
Wooly Indianwheat or wooly plantain is important and widespread enough that it made the famed list of 200 species used in the International Intercollegiate Range Plant Identification Contest sponsored by the Society for Range Management. See North American Range Plants (Stubbendieck et al., 1992, ps. 386-387) for treatment in that regard . Wooly plantain was also given brief coverage in Notes of Western Range Forbs (Hermann, 1966, p. 271) as well as in the Range Plant Handbook (Forest Service, 1940, W150) and Pasture and Range Plants (Phillips Petroleum Company, 1963, p. 146). An important little guy.
Ottawa County, Oklahoma. Early July; immediate post-bloom phenological stage.
Note on location: other coverage of wooly or prairie plantain (P. patagonia) in Range Types of North America was provided in the Grasslands chapter entitled, Mixed Prairie- IA, Southern and Central Plains.
88. As opportunistic as its feed source- Larvae, caterpillar, of buckeye butterfly (Juanania coenia) feeding on wooly or prairie plantain that "came from out of nowhere" to pioneeer or initially colonize the bare soil of a former stand of tall fescue that was devestated by a combination of Extreme Drought and mismanagement. Your author observed this former tall fescue pasture for fifty years and cannot recall ever seeing wooly plantain growing on it. Likewise, this author never noticed buckeye butterfly caterpillars on this pasture. Caterpillars of buckeye butterfly feed on various plant species, but species of the Plantaceae are one of their major feed sources. Apparent absence of these caterpillars reflected the absence of their host or prey plants. The native, annual, pioneering wooly plantain became available only as a result of prolongued drought and simultaneous managerial abuse, and it was only under that combination of conditions that the caterpillars also appeared "from out of nowhere". Presence of host plants and phytophagous insects was a result or an outcome--a symptom as it were--and not a cause of disappearance of tall fescue.
Note that under these conditions any attempt at human control of "weeds" or insect "pests" would be stupid. Under these, at least partly anthropogenetic, set of circumstances the plants and insects are neither weeds nor pests. They are Nature's response to severe disturbance that, again, was at least partly man-made.
Caterpillars of the buckeye butterfly are specialized phytogophagous larvae that sequester iridoid glycosides from herbage of their host plant species (Bowers, 1984; Bowers and Collinge, 1992). Plantago species are primary among these plant hosts. This is an evolutionary adaptation against would-be predators, especially insects, of buckeye butterfly larvae (Stamp, 1992; Camara, 1997). It is an adaptation that involves interrelations among herbage-eating insects and their host plants (Stamp, 1997; Quintero et al., 2014). These plantains and their hosted herbivores obviously caught the attention of observers besides your Range Management professor. Ain't Nature grand?
Ottawa County, Oklahoma. Early July.
89. A rough character for rough times- Rough buttonweed poor joe (Diodia teres) was the late summer/autumn dominant of the pioneer or initial colonizer stage (first seral stage) on "go-back land" that prior to a multi-year drought and "piss-poor" management had been a stand of tall fescue for 50 years in the Ozrk Highlands (Springfield Plateau). Drought accompanied by overuse and total lack of soil management wiped out the tall fescue except for local areas of shade. In the first year (growing season) after near-estirpation of tall fescue the initial seral stage of old field (secondary) plant succession was a community of two Plantago species, large-bracted and wooly plantain, and rough buttonweed.
Under this condition these three, native, annual forbs were Nature's healers of degraded plant life and bare soil vulnerable to soil erosion. These were the first three "first responders" to render aid to threatened land. They were not weeds, and this author will go on the prod if they are so labeled. It was man that made them this way. There were some native perennial plants tht manged to "hang on" in this devestated plant life at much smaller cover and density (see above introduction to this section), but in the first year of "land laid bare" it was these three annual species that protected the bare breast of Mother Nature.
This is a classic example--and a perfect one for teaching this concept to beginning rangemen--that so-called "weeds" are all too often invited to "crash the climax party" when man as range manager, farmer, forester, or wildlifer abuses range, pasture, or forest. "Mother Nature abhors a vacuum" as the old syaing so poignantly put it.
qrffollowing Tyrl (2008, ps. 304-305) and Phillips Petroleum Company (1963, p. 125)
Ottawa County, Oklahoma. Early July; peak-bloom phenological stage.
90. Tiny but rough- Details of rough buttonweed or poor joe growing and blooming profusely on bare soil in the western Ozark Plateau following loss of a stand of tall fescue in a multi-year drought and extreme mismanagement.
Rough buttonweed is a native, annual forb and obviously an r-selected species. It is the ultimate disturbance-adapted species. The self-compatable rough buttonweed has received some study from evolutionary adaptation as to ecotypes (Jordan, 1992) and variation among populations (Hereford and Moriuchi, 2005). Hereford (2013) found that there were no differences between inbred and outbred plants of rough buttonweed in regards fitness to foreign areas within this species' native habitat. Althought outcrossing is generally beneficial to plants introduced into foreigh habitats, Hereford (2013) reported that inbreeding depression in rough buttonweed did not result in lowered fitness (adaptation) within the native habitat of this the highly selfing, annual, pioneer species. From an evolutionary as well as an ecophysiological perspective, poor joe is highly adapted to pioneering disturbed areas.
Ottawa County, Oklahoma. Early July; peak-bloom phenological stage.
91. Rough and getting ready- Views of whole shoots (first slide) and of sexual shoots from flowering to fruit production (second, third, and fourth slides) of rough buttonweed or poor joe growing on bare land following loss of a tall fescue crop in a multi-year drought (ranged from Severe to Extraordinary on the Palmer Index) and horrid mismanagement. Almost no cover or even presence of this species was observed in this stand of tall fescue until it was wiped out and an opportunity afforded for colonization by rough buttonweed.
In an observational context, rough buttonweed had not been seen before and it was not seen after this one year on this land. Such is the nature of pioneering species. Rough buttonweed made a fantastic seed crop, and those "I'll-be-here-forever" fruits (technically speaking, indehiscent carpels according to Diggs et al. [1999, p. 962]) will await another opportunity (a man-made or natural; or, an and/or successional moment) for the species' next appearance.
Rough buttonweed is in the mader or coffee family (Rubiaceae) Good coverage of rough buttonweed from ecological as well as morphological and taxonomic perspectives was provided by Tyrl et al. (2008, ps. 304-305). For taxonomic/morphological treatment of poor joe in this Ozark Plateau-Central Lowlands region the following manuals were recommended (Steyermark, 1963, ps. 1395-1397 passim; Great Plains Flora Association, 1986, p. 617; Diggs et al, 1999, ps. 962-963).
Ottawa County, Oklahoma. Early July; late-bloom/early to mid-fruit phenological stage.
Prairie succession delayed by tame pasture- Two views of a stand of broomsedge bluestem at peak standing crop still standing in late December. The old field show in these two slides had been in cultivation (for winter wheat) for about 20 years when it was seeeded to tall fescue (Festuca arundinacea). A poor to marginal stand of tall fescue eventually grew to become a good stand which, after 38 years, was wiped out in an Extreme Drought six years prior to these two photographs. Broosedge bluestem, which had invaded following cessation of winter wheat farmng and seeding to tall fescue, had persisted at modest cover with the tall fescue which was used as a hay field. Four years following Extreme Drought the old field became a single-species stand of broomsedge bluestem as seen here.
This vegetational development stood in stark contrast to an example of secondary plant succession on an old field with essentially the same soil that was a mile and a half away. That old field had nothing done to it. "On it's own" (through secondary plant succession) it went back to climax tallgrass (big bluesem, switchgrass, Indiangrass, and hairy panicgrass) prairie in less than a quarter century (23 years) following a period of tillage.that had lasted for over three-fourths of a century. That was a period of farming that was almost four times longer than the wheat field that became a tall fescue hay field--only to revert to broomsedge bluestem when Extreme Drought killed out the tall fescue.
If the field that is now in broomsedge bluestem had just been abandoned, and had nothing done to it, it would now be in climax big bluestem, switchgrass, Indiangrass prairie. Seeding to tall fescue prevented (at least retarded) secondary plant succession. Then after Extreme Drought wiped out the tall fescue the broomsedge bluestem seral stage developed.
Ottawa County, Oklahoma. Late December.
92. Once it was tall fescue; now bermudagrass- A former mixed pasture of Kentucky 31 tall fescue and common bermudagrass (tall fsecue the slight predominant species; first among co-dominants) that had been severely overgrazed by beef cattle (cows and calves ) and mules (Equus caballus mare X E. asinus jack) during the five-year Severe to Extreme Drought that began in late first decade of Twenty First Century. This was early summer of the first year after end of the drought.
The combination of pasture abuse (mostly overgrazing; no fertilization or weed control) and drought all but eliminated the tall fescue while common bermudagrass had thrived and so increased its cover that bermudagrass was now sole dominant grass of the former mixed cool-season and warm-season introduced perennial grass pasture.
The now-dominant bermudagrass was continuing to be overused by mules while occasional plants of the narrow-leafed whorled milkweed (Aasclepias verticillata) were untouched (that is, for a limited period of time).
Ottawa County, Oklahoma (Springfield Plateau portion). Late July.
93. Mules left the toxic one (at least for a while)- Several plants of whorlded milkweed growing on a severely overgrazed (by beef cattle and , mostly, mules) pasture that had been a mixed stand of tall fescue and common bermudagrass. but that was now almost exclusively bermudagrass after a five-year Severe to Extreme Drought (Palmer Scale) in conjuction with overgrazing. At time of these slides, mules were the only livestock on this deteriorated mixed pasture (now existing as a bermudagrass stand).
Whorled milkweed has long been recognized as a stock-poisoning species (Burrows and Tyrl, 2006, 2012). For ever longer, mules and asses have been known to more intelligent (and stubborn) than horses. On this deteriorated and overgrazed pasture of species stand of common bermudagrass mules had demonstrated their intelligence and/or instinct and avoided whorled milkweed. Currently mules were cultivating the milkweed plants--at least at this time. (Of course the mules' owner was the one who was really doing the "cultivating", but he was not personally grazing the depleted pasture.)
Ottawa County, Oklahoma (Springfield Plateau portion). Late July.
94. Next day mules changed their minds- The next day your author discovered that the mules had decided that they should eat the toxic whorled milkweed. Actually mules did not eat the milkweed plants they just bit them off at ground level and left the gnawed off shoots of the poisonous forbs on the ground. Were mules just playing (muling around)? Were they bored and engaged in crib-like behavior? Or did mules intend to consume milkweed herbage and think better of it? Was the herbage bitter or off flavor for mules' taste? Only God and the mules know the answer, but for sure mules cut off milkweed plants at shoot base level.
A degraded mixed pasture formerly of Kentucky 31 tall fescue and common bermudagrass that was converted into a single species stand of bermudagrass during a five-year Severe to Extreme drought combined with overgrazing by beef cattle and mules in the western Ozark Plateau. First the mules "cultivated" the whorled milkweed by severely defoliating the bermudagrass while leaving the whorled milkweed untouched. Then mules bit off the milkweed which might seem to be biological control of this toxic forb. No, let's take a closer look...
Ottawa County, Oklahoma (Springfield Plateau portion). Late July.
95. Jist bein' mules-- and milkweeds- Shoots of whorled milkweed had ripe fruit (capsules) when they were bitten off and spit out by mules on an overgrazed pasture dominated by common bermudagrass. Yes, the mules appeared to apply biological weed control, but it was too late. Whorled milkweed had already made seed for its next crop, plus this perennial forb might regrow enough to manufacture adequate food reserves to survive the dormant season. Even if these plants were killed by mules there was still...
Ottawa County, Oklahoma (Springfield Plateau portion). Late July.
96. Baied out to safety- Ripe capsules with mature--and parachute-equipped--seeds of whorled milkweed as they occurred lying on the ground after mules bit off and spit out shoots of whorled milkweed on an overgrazed pasture dominated by common bermudagrass. Mules and beef cattle in conjuction with a five-year Severe to Extreme Drought killed out the former dominant tall fescue leaving the lesser co-dominant bermudagrass and accompaning weeds such as whorled milkweed. After helping propagate the milkweed by reducing competition around milkweed plants the mules eventually bit the milkweed off--after milkweed plants had set viable seed.
Now milkweed seeds can germinate under conditions of reduced competition brought about by continuing overgrazing by mules. Whose side are these mules on anyway?
Ottawa County, Oklahoma (Springfield Plateau portion). Late July.
97. Besides the mules missed a few (and we might as well enjoy the flowers)- Sexual (flowering) shoot, details of inflorescence, and close-up of flowers, top to bottom, respectively, of whorled milkweed (Asclepias verticillata). This was one of the plants that grew on an overgrazed, recovering-from prolonged-drought pasture that had become dominated almost exclusively by common bermudagrass after tall fescue died out from the combined stresses.
This milkweed plant was growing close to a barbed wire fence rather than out in the middle of the pasture and mules did not bit off this individual. For whatever reason, this plant was less mature and still flowering rather than maturing seed as was the case fo plants the mules bit off. This plant was never bitten off and later set and spread plumose seeds. There was not going to be any shortage of whorled milkweed plants on this overgrazed, degraded pasture--with or without tall fescue.
Ottawa County, Oklahoma (Springfield Plateau portion). Late July.
Stubbling back to the introduced fescue species...
98. Another (and minor) introduced fescue- Two plants of meadow fescue (Festuca elatior= F. pratensis) shown as examples of anothere introduced Eurasian Festuca species. Meadow fescue has naturalized in North America, including over eastern parts of the general Great Plains Region (Great Plains Flora Association, 1986, p. 1181). In comparison to its "kissing cousin", tall fescue, meadow fescue is generally an inconsequential species. It was included here due to closeness morphologically and taxonomically to the extremely important tall fescue, and because both species have been used as agronomic forages with each frequently included under the banner of tall fescue.
Sleper and West in Moser et al. (1996, p. 472) included both F. pratensis, a diploid species, and F. arundinacea , a hexaploid species, in the tall fescue forage category noting that they were "[t]he two species of primary interest for cultivation". These authors remarked that Carolus Linnaeus confused these two morphologically similar species and included both in his species, F. elatior. For Missouri, the heart of tall fescue empire , Steyermark (1963, p. 92) distinguished between F. elatior var elatior and F. elatior var. arundinacea, but this was revised in Steyermark's Flora of Missouri (Yatskievych, 1999, ps. 835-836, 838-839) to F. arundinacea and F. pratensis.
Meadow fescue so closely resembles tall fescue that the Great Plains Flora Association (1986, p. 1179) distinguished between the two species based on auricle features. These authors also described panicles of meadow fescue as being "relatively narrow" whereas panicles of tall fescue are typically open though they, too, are sometimes narrow (Great Plains Flora Association, 1986, ps. 1180-1181).
A more recent developmen was reinterpretation of tall fescue as Lolium arundinaceum based on work by Darbyshire, but by the time the two-volume treatment of Gramineae appeared in the Flora of North Americca (Barkworth et al., 2007, ps. 445-448) Darbyshire had already moved tall fescue from Lolium to Schedonorus. Wonder how long that latest taxonomic fad will survive. Perhaps as long as it takes to get another publication, which is mostly what "musical chairs" of plant names is about. Meanwhile thathad been a half century of research with tall fescue labeled as Festuca arundinacea (or, perhaps, F. elatior; one way or the other with genus Festuca) such that "musical names" created confusion and hindered, rather than facilitated, scientific communication and knowlege dissemination.
Anyway the biological range of naturalized meadow fescue (F. pratensis) in North America is essentially the same as that of tall fescue (F. arundinacea) so examples of both species were presented to viewers. An example of these overlapping species ranges in Missouri was shown in Yatskievych (1999, ps. 830, 836).
Abandoned lot in Hays, Kansas, Ellis County. Late June-grain-ripe phenological stage.
99. A narrower panicle- Panicle of meadow fescue showing panicle appressed branches (pressed against the central floral axis) . This example should be compared back to those of tall fescue presented above. Abandoned lot in Hays, Kansas, Ellis County. Late June-grain-ripe phenological stage.
100. 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.
101. Orchardgrass or, chiefly British, cocksfoot (Dactylis glomerata)-
This Eurasian cool-season grass of the Festuceae or Poeae tribe is another agronomic
forage species that does "double duty" as a sometimes range plant.
Orchardgrass is primarily a pasture and not a hay grass though it is often valuable
for both. The usefulness of this tame pasture grass in a range context is similar
to that of timothy. While orchardgrass is not as cold-tolerant as timothy it
is more drought- and heat-tolerant than timothy or Kentucky bluegrass and less
drought- and heat tolerant than smooth bromegrass and tall fescue (Jung and
Baker in Iowa State University, 1985, p. 225). Given the "agronomic niches"
of these various introduced cool-season perennial grasses it is not surprising
that orchardgrass also became a naturalized range grass and persist as a locally
important plant on the native grazing grounds of North America. Understandably,
orchardgrass was included on the Society for Range Management national range
plant contest list.
As a tame pasture and/or hay grass, orchardgrass is grown over geographic
regions similar to those of other cool-season agronomic grasses, especially
tall fescue and ryegrass (eg. northcentral North America and the Pacific
Northwest), except that, as indicated by its adaptation, orchardgrass
tends to be a more northern pasture plant. For example, orchardgrass
is locally important in Missouri but not in Oklahoma or Arkansas. As
an irrigated pasture grass orchardgrass is grown from the Rio Grande
Valley and High Plains of New Mexico south into Mexico. Orchardgrass
naturalized over a geographic range similar to that of timothy with
which it often grows as a fellow naturalized associate (eg. mixed prairie-ponderosa
pine savanna and westward as an understory species in forests through
the central and northern Rocky Mountains). Both of these domesticated
festucoid grasses provide nutritious and palatable forage for native
ruminants including, at certain sessons and prior to grass maturity,
browsers like deer. Their lasting value as livestock forage is attested
to by their cultivation for hundreds of years.
Likewise, drought-tolerance of these two species is extremely low or poor. One veteran cow-calf operator remarked wryly (and wily), "Orchardgrass is named right; the only place it can do well here (southwest Missouri) is under trees in an orchard". His play-on-words comment had much wisdom in the empherical observation that in dry spells (prolongued drought in paticular) several species of introduced perennial grasses, especially the cool-season Eurasian species, do better under moderate shade than in full sun. This certainly includes tall fescue, timothy, and orchardgrass. Where these Eurasian species have naturalized individual plants survive best and at greatest density, cover, etc. in shade at edges of forests, under isolated trees (such as shade trees), on north and east slopes, and even under light cover of shrubs.
In much of the 1960s and 1970s throughout the Ozark Highlands and Boston Mountains Region there was much cost-sharing under the Agricultural Conservation Program, administered through the Agricultural Stabilization Conservation Service, for clearing (by bull-dozing) native oak-hickory forests and type-converting these forests, woodlands, and savannahs to permanent pastures of introduced species like tall fescue and orchardgrass. Federal rules (United States Department of Agriculture) for cost sharing permiitted leaving only about one tree per acre in such pasture-establishment projects because allegedly the "experts" asserted that adequate light for these introduced grass species necessitated removal of all but one tree per acre. The stupidity of this misplaced (and research/empherically baseless) rule became only all too evident in the protracted Extreme Drought of the 2010-2012 grass-growing seasons. Then the major and most common microsites on which orchardgrass and tall fescue survived in pastures on rocky hillstops and shallow environments was under shade, especially where grazing had been moderate to heavy. Such is the folly of half-educated, self-assertive fools with bureaucratic authority.
Ottawa County, Oklahoma. May.
102. Another composite (and cespitose) example- A single plant and two panicles (first and second slide, respectively) of orchardgrass in a dry period (early on-set of drought) in late spring. This solitary plant--though smaller than some individuals like the one immediately above--provided readers with a good example of the overall habit of this species which usually consist of a large clump of mostly basal leaves with large sexual shoots bearing panicles. Under moderate grazing only a few shoots "escape" defoliation with the result that there are few shoots that develop into sexually reproductive ones. This individual plant was growing in an ungrazed pasture and provided a textbook specimen to show a growing on-the-land "herbarium mount" to beginning students, a few of whom might even appreciate the effort and be able to recognize orchardgrass in a pasture.
Ottawa County, Oklahoma. Late May; peak biomass of shoots, anthesis.
103. 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.
104. Dense cluster- Groups of spikelets blooming on a branch of the panicle of orchardgrass. These floral units were on the plant presented in the two photographs of the slide-caption set before the last (two sets immediately above).
Ottawa County, Oklahoma. Late May; peak anthesis.
105. Inflorescence of orchardgrass (Dactylis glomerata) at anthesis-
Ottawa County, Oklahoma. May.
One of the more recent and comprehensive works on cool-season forage grasses,
including for rangeland, is that by Moxer et al. (1996). It is well worth
the investment in money and reading time.
106. With snow all around on the ground- Old (dead, last year's) shoots and young (green, slowly growing) shoots of orchardgrass in early winter with snow-covered soil. All species in the fescue or bluegrass subfamily (Festucoideae= Pooideae) are cool season species with the Cs photosynthetic cycle. Theses grasses (both annual and perennial species) grow when it is the cool season (as in "winter time"). Actually this growth may be quite slow or intermittent (take place only when temperatures are relatively warm) within the cool season yet remain green (maintain live tissue) throughout the cool season so as to appear to be growing.
Furthermore, even this living tissue of cool-season species can be winter-killed (frost-killed) especially if comparatively cold temperatures occur before current living shoots have "hardened off". "Hardening off" refers to the series of complex biochemical/physiological processes taking place in plants that adjust their living tissue to cold temperatures. This "hardening off" process includes increasing concentration of solute in plant cells.
The next pair of slides presented examples of two species of festucoid grasses (one plant of each species) growing side-by-side in early winter, both plants of which had some winter-killed or frost-killed leaves.
Ottawa County, Oklahoma. Early January.
107. Side-by-side cold hardiness- Orchardgrass (left) and tall fescue (right) growing in early winter and showing differences in frost (freeze) damage to leaves in these two festucoid grasses (in fact, both species are in the same tribe, Festuceae or Poeae). The single plant of each species illustrated the strictly cespitose (bunched or tufted) habit of these two bunchgrass species. The orchardgrass plant--as was typical for its species--retained more dead herbage from last year's shoots whereas the tall fescue plant--as was typical for its species--had experienced earlier death of sexual shoots so that these shoots had already been decomposed rather than being retained as dead herbage. Neither of these two plants had been grazed by vertebrate or insect species.
It should be obvious that the second (lower) slide was a closer-in view of the two plants whose aboveground herbage was shown in its entireity in the first (upper) slide.
What is less obvious was the much greater degree of winter-killing (frost-caused death) of shoots in orchardgrass (left) than in tall fescue (right). It was readily apparent that there were only a few leaves of tall fescue that had sustained frost injury (appeared as brown, dead leaves) whereas the terminal portions of all green leaves on orchardgrass had some dead tissue due to freeze injury. What was less obvious (at least to viewers not familar with orchardgrass) was that much of the dead (brown, withered, "wrinkled") leaves near or in the center of the orchardgrass plant were current year's growth that had been killed by cold temperatures. The taller, full-grown shoots of orchardgrass (that were dead) were produced the previous growing season (last year), but all the low dead leaves were previous growing season (this year) growth that had been killed by cold. Tall fescue suffered much less winter-killing or frost injury of its current growing season (this year) growth.
Ergo: tall fescue is much more (dramatically so) winter-hardy or cold-tolerant than orchardgrass. Such winter-hardiness or cold-tolerance has traditionally been attributed--at least, anecdotally--to endophyte symbiots (Neotyphodium coenophialum) living inside tissue of tall fescue, a less obvious yet just-as-real mutualism as that of papilionaceous legumes and their nitrogen-fixing, Rhizobium bacteria root symbiots. Over the last several years much has been written about the tall fescue-endophyte mutualism, but the author of this caption deemed it sufficient to refer readers only to the most up-to-date (at this time) treatise on tall fescue, Agronomy Monograph No. 53 (Fribourg et al., 2009, passim). Much more has been written about the host plant-endophyte mutualism in regards to drought tolerance than to cold tolerance or winter-hardiness. (As was shown above, much of the hype--including that in scientific literature--regarding drought tolerance of tall fescue was grossly overstated.)
Ottawa County, Oklahoma. Early January.
108. 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.
109. 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.
110. Type conversion in the Palouse Prairie- Conversion of a former Idaho fesuce (Festuca idahoensis) consociation in the Palouse Prairie into a monoculture of Kentucky bluegrass. Grassland composition in fencerows adjacent to this pasture served as relict or reference areas showing that this Northern Great Plains grassland had once been an Idaho fescue variant of the Palouse Prairie bunchgrass grassland (ie. an Idaho fescue-dominated range plant community).
It was most likely that heavey grazing had converted this native bunchgrass prairie range into a Kentucky bluegrass pasture.
Cascade County, Montana. Late June. No FRES, Kuchler, or SRM units.
Kentucky bluegrass- Kentucky bluegrass is beyond doubt one of the most important cool-season grasses in North America from both the standpoint of an agronomic forage (pasture species) and low-input, more ecologically managed pasture (naturalized range species). 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. In the most recent encyclopedia of North American grasses (Barkworth et al., 2007, ps. 522-527) specified that there are six subspecies of P. pratensis in North America, but the only taxa native to this continent are those of Arctic and closely adjoining regions. The agronomic P. pratensis (forage and turf forms) in North America, of which there are over 60 cultivars, are essentially of subspecies pratensis. These are all Eurasian exotics derived directly through plant introduction or indirectly through artificial selection (plant breeding) from imported plants. P. pratensis subsp. pratensis is clearly a highly domesticated group of grasses.
Kentucky bluegrass is most ceretainly was not native to southeastern and central 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. Barkworth et al. (2007, p. 522) described P. pratensis as "highly polymprphic" and concluded that it most likely has "the most extensive series of polyploid chromosome numbers of any species in the world". They further specified that P. pratensis is a hybridogenic species meanint that it is made of of "numerous lineages with the same basic maternal genome, different paternal genes (Barkworth et al., 2007, p. 522).
With this diverse assemblage of germ plasm it is little wonder that Kentucky bluegrass has become so widespread in its species range and adapted to such an array of habitats. These environments include numerous range sites to which this species (again, probably P. pratensis subspecies pratensis) has naturalized. Kentucky bluegrass is now as much a North American range grass as the native Poa species. Treatment of Kentucky bluegrass herein was primarily that for a naturalized range grass and not as an agronomic (tame pasture) forage.
For instance, besides grasslands of true and mixed prairies in the Northern Great Plains there are immense acreages of grazing disclimax grassland comprised of 1) varying mixes of Kentucky bluegrass (Poa pratensis) and smooth brome (Bromus inermis) and, at the ultimate state of disturbance (primarily overgrazing) climax, 2) monocultures of Kentucky bluegrass. These two disclimax range types are naturalized types much like the California annual grassland type and cheatgrass or downy brome (Bromus tectorum) type. The Kentucky bluegrass and Kentucky bluegrass-smooth brome types were not recognized as such by authors in Shiflet (1994). These two disclimax grassland types are comprised of exotic (Eurasian in origin), perennial grass species that were intentionally introduced as agronomic crops by the white man in contrast to Eurasian annual grasses that were inadvertently brought over by Christopher Columbus and his enlarged clan from Europe and the British Isles. Eurasian disclimax grasslands as well as skilfully managed pastures and hay fields of smooth bromegrass and Kentucky bluegrass were covered under the chapter, Introduced Forages, Grasslands.
111. Typical turf- Characteristic sward of Kentucky bluegrass as it appeared under light grazing. This Kentucky bluegrass sod was of plants that had naturalized and covered patches of soil that were largely shaded in the Ozark Plateau. These shoots had not been grazed for several months. They were at full-bloom phenological stage. While this was a thick local stand of Kentucky bluegrass it was also representative of the growth and development (as to density, cover, etc.) of which this introduced pasture species is capable under favorable growing conditions.
Ottawa County, Oklahoma. Mid-June (late spring).
112. Mature grain and still green leaves- Typical plant of Kentucky bluegrass at grain-ripe stage. Sexual shoots had matured, died and turned to straw while lower leaves and culms were still bluegrass-green. This is one (of several) features that renders this agronomic--and now naturalized range--species such a good grazing grass. In many other grass species maturation of sexual shoots coincides with (perhaps influences) senescence of many or most other shoots, both culms and lower leaves.
Ottawa County, Oklahoma. Late May; grain-shatter stage of phenology.
113. Kentucky bluegrass form under heavy grazing- In contrast to the robust almost bunchgrass-habit of the preceding specimen growing in the American south the closely cropped specimen shown here grew in the foothills of the northern Rocky Mountains (Bitterroot Range). Note that in both individuals of this rhizomatious species reproduction was both asexual and sexual. Kentucky bluegrass is one of the textbook examples of a short shoot grass that produces a relatively high proportion of sexually infertile shoots but that does not not elongate it's shoots very far from the soil until late in the annual growth cycle (thereby providing maximum protection for it's propagules-- both vegetative and sexual diaspores-- from defoliation). Flying D Ranch, Gallatin County, Montana. June.
114. 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.
115. 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.
116. 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.
117. 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.
118. Kentucky bluegrass in full-flower- Panicle of Kentucky bluegrass with spikelets at anthesis. This panicle with four or five whorls of secondary branches was more typical than the extremely elongated panicles shown in the preceding slides. Students should take note of the geographic range over which photographs of Poa pratensis shown in this "chapter" were taken. Jackson County, Colorado. June, peak bloom.
119. Rank growth with plenty of water- In one of the wettest winters and springs on record (and following a previous wet year) this Kentucky bluegrass grew to heights of over two and one-half feet. Their production of anthers and stigmas must have also set some kind of record. These over-sized plants were obviously atypical and herbage of this degree of rankness was obviously of inferior feed quality. These plants were ungrazed (and the shoots had not been mowed). They were presented here to show students the comparatively large size (maximum height and rankness ever seen by this rangeman) and high herbage yeild to which this introduced species is capable under ideal growing conditions (especially of soil moisture).
Garfield County, Montana. Mid-June; peak anthesis.
120. Hanging small- Various views of spikelets with their florets at anthesis of Kentucky bluegrass. Garfield County, Montana. Mid-June; peak anthesis.
Across much of the Northern Great Plains naturalized Kentucky bluegrass (offspring of the introduced Eurasian forage crop that have "gone native") forms vast acreages of disturbance climax consisting almost exclusively of this one species. The extent of this anthropogenic naturalized range is readily apparent to any observant sojourner who has traveled widely in this region. The total area covered by Kentucky bluegrass disclimax range almost certainly comprises millions of acres or, more generally, hundreds of square miles. The Kentucky bluegrass disclimax includes both degraded range (perhaps mostly atrtributable to overgrazing, past farming, commercial and residental development, transportation, military activities, and mining, especially fossil fuel development) as well as tame pasture. The latter includes both purposely seeded monocultures (single-species stands) of Kentucky bluegrass and improperly managed permanent pasture mixes. Examples of these degraded permanent pastures include mixtures of such agronomic crops as smooth bromegrass (Bromus inermis) and alfalfa (Medicago sativa) that under varous combinations of heavy grazing, incorrect hay making practices, inadequate fertilization, etc. became "solid stands of nothing but Kentucky bluegrass".
Throughout much of the Northern Great Plains countless acres of native grasslands that formerly consisted of miscellaneous communities of needlegrass, bluestem, grama, and/or wheatgrass species like Society for Range Management (Shiflet 1994) rangeland cover types 606, 607, 608, and even 610 have been converted to Kentucky bluegrass naturalized range. These drastic and profound changes in range vegetation constitute de facto type conversions.
The cause(s) of these unintended type conversions is apparently unknown and remains controversial. This author hypothesized that much of this was due to overuse/overgrazing of native grasses that resulted in degraded native range and development of a grazing disturbance climax of Kentucky bluegrass. Such an ecological transformation would be much the same as conversion of native bunchgrass prairies in California into anthropogenic grrasslands made up of Eurasian annual grasses and forbs (Weaver and Clements, 1938, Burcham, 1957). In contrast to the standard "overgrazing leads to invasion by introduced species" theory, Sedivec and Printz (2012, ps. 10, 12, 13, 14, 15, 16, 19, 20, 23, 24, 25, 27) stated (hypothesized) that on some North Dakota range sites (eg. Saline Lowland, Subirrigated, Sandy Claypan, Clayey, Thin Loamy, Loamy Overflow) extended periods of light use to nonuse resulted in dominance by Kentucky bluegrass and/or Kentucky bluegrass with native grasses and forbs of lower successional status. These same authors, however, pointed out that on other range sites (eg. Limy Subirrigated, Subirrigated, Wet Meadow, Claypan) Kentucky bluegrass comes in as a dominant with overgrazing as well as extended nonuse (Sedivec and Printz , 2102, ps. 8, 11). According to Sedivec and Printz (2012, p. 10, 27) underuse or nonuse resulted in accumulations of mulch (dead herbage) that excluded enough light to give competitive advantage to and result in dominance by "more shade-tolerant grasses" like Kentucky bluegrass and smooth bromegrass. Another factor in the increase of Kentucky bluegrass invasion was the recent series of wet years (Sedivec and Printz, 2012, p. 5).
Thus, cause(s) of dominance by Kentucky bluegrass (the Kentucky bluegrass disclimax; Kentucky bluegrass range type) in the Northern Great Plains are multi-faceted and range site-specific.
Such "man-made" grasslands constitute what Clements (1936, ps. 265-266) described as disclimax (disturbance climax), a plant community of such permanency or indefinite duration as to have the general features of a "true" or "natural" climax (ie. of the previous climatic or edaphic climax; of the potential natural vegetation). Therefore, such anthropogenic disclimaxes should be interpreted as the "climax of now and for foreseeable future".
While the Society for Range Management (Shiflet, 1994) recognized rangeland cover types of intoduced grasses such as crested wheatgrass (SRM 614; for the Northern Great Plains Region at that) and tall fescue (SRM 804), their treatment did not include a Kentucky bluegrass cover type which occupies a far greater land area than that of the tall fescue type. For that matter, if total area of cover types could be measured it is would almost assurredly be found that Kentucky bluegrass as a naturalized rangeland type occupies a considerably greater acreage in the Northern Great Plains than does crested whetgrass. Future editions of Rangeland Cover Types should include a Kentucky bluegrass cover type.
It should be remarked also that naturalized ranges of Kentucky bluegrass disclimax might well be more productive--and with less judicius management--that the ranges of native grasslands (climax or seral) that were replaced by this disclimax. At least this appeared conceivable under existing socioeconomic conditions in which ranches and farms consisted of comparatively small land holdings on which the main consideration was the capacity to graze--at a profit or, at least, with minimal financial loss and/or input--beef cattle under relatively high stocking rates (ie. fewer acres per animl unit or animal unit month). Obviously, livestock (primarily beef cattle) have not grazed out the rhizomatous, grazing-tolerant Kentucky bluegrass-- at least not at stocking rates which permit continuation of cattle operations. Livestock grazing was obviously a major factor in replacement of native range cover types by Kentucky bluegrass across much of the Norther Great Plains
Some examples of naturalized Kentucky bluegrass range (disturbance climax) in the Northern Great Plains were presented below under Introduced Forages- Grasses- IB.
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