Introduced Forages- Grasses - IB

Grasses- Festuceae Tribe Continued

113. Poa pratensis disclimax- Former true prairie grassland of the wheatgrass-needlegrass rangeland cover type (SRM 607) that was a type conversion into a Kentucky bluegrass disturbance climax. There were some adult and juvenile plants of Wyoming big sagebrush (Artemisia tridentata subsp. wyomingensis). The type conversion had been effected by either seeding to Kentucky bluegrass, overgrazing, or a combination of these two practices. Kentucky bluegrass is naturalized throughout the Northern Great Plains (much of North America for that matter) so it is possible that this highly rhizomatous and efficiently self-seeding species had established even without being planted.

Whatever the mechanism(s) this was an example of range vegetation that should have been recognized by the Society for Range Management (Shifllet, 1994) as a rangeland cover type-- for better or for worse.

There were some plants of the naturalized, weedy annual brome, Japanese chess (Bromus japonicus) along with a natrualized, annual, Eurasian forb known as dandelion (Taraxacum officinale). Otherwise, there were few other weedy range plants. (KIentucky bluegrass is estremely effectively at excluding other plant species, especially naturalized ones).

This was a highly productive Subirrigated and Wetland range site association, which somehow was left out of the eastern sedimentary plains range site (Ross and Hunter, 1976). These two photographs were taken during a record wet winter and spring. It was little wonder that Kentucky bluegrass was doing so well.

Garfield County, Montana. Mid-June; ripening stage of caryopses. Units of potential natural vegetation were not appropriate for this type conversion. Northwestern Great Plains-Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

114. Depends on one's point of view- This disturbance climax of Kentucky bluegras had replaced (perhaps permanently in human time scale) the climax (pre-Columbian) Northern Mixed Prairie. This man-made vegetation could be regarded as either 1) degraded ("destroyed") mixed prairie or 2) improved pasture depending on the perspective of the producer or whoever else cast eyes on it. An ancient Greek sentiment expressed in its present form by Margaret Wolfe Hungerford summed it up: "Beauty is in the eye of the beholder".

The climax wheatgrass-needlegrass mixed prairie had through overgrazing, fire cessation, and introduction of the exotic Kentucky bluegrass (almost assuredly all three had been and continued to be in force) and through tillage or farming (perhaps) became a de facto type conversion to a near-monoculture (single-species stand) of Kentucky bluegrass. One of the few remaining native plant species was silverleaf scurfpea or silverlead Indian breadroot (Psoralea agrophylla) several specimens of which were visible in immediate foreground of the first of these two slides. About the only other plant species present was yellow sweet-clover. Very few plants of the native dominant or associate range species remained. Most of these were in microsites of large rocks that were glacial erratics (treated in next slide-caption set). Extirpation of the native flora was almost complete on numerous Kentucky bluegrass type conversions such as this one.

The second slide showed sward and homogenity of a typical Kentucky bluegrass type conversion. Not all such monocultures were as "pure" or "solid stand" of Poa pratensis as this one. A similar anthropogenic grassland range that had somewhat more cover and quite a higher density of some native forbs was just across a highway from the one featured here. That other range was shown and described below.

The range featured in these two photographs and the next three slide-caption sets was being grazed currently by beef cattle. It now appears that once Kentucky bluegrass comprises roughly 40% of plant cover or biomass the range site is degraded to a permanent to semi-permanent Kentucky bluegrass state as viewed from the state and transition model (L. O. Voigt, area range specialist, North Dakota Natural Resources Conservation Service, personal communication). This man-made seral stage was labeled herein as a "stuck steady state" (based on the state and transition model of plant succession employed by the NRCS).

Emmons County, North Dakota. Late June-peak standing crop and grain-shatter stage of Poa pratensis. No units of natural vegetation were appropriate and, unfortunately, the Society for Range Management (Shiflet, 1994) had omitted this widespread rangeland cover type. Clayey range site degraded to Kentucky Bluegrass successional state. Northwestern Glaciated Plains- Missouri Coteau Slope Ecoregion 42c (Bryce et al., Undated).

115. Except for the rocks- Glacial erratics of granite in a former climax wheatgrass-needlegrass mixed prairie that had been converted to a monoclimax of Kentucky bluegrass, a disturbance climax effected through introduction of Kentucky bluegrass by whiteman, overgrazing with livestock, cessation of natural fire, and (possibly) former tillage for production of field crops. Piles of such erratics in numerous farm fields where they were hauled and dumped by farmers attested to the fact that presence of glacial erratics did deter growers from plowing virgin sod of the Northern Mixed Prairie. Many of the glacial erratics on the land surface were, however, natural and remained where they had been deposited by the Kansan, Iowan, or Wisconsin Ice Age series.

Piles of erratics (regardless of whether placed by human hands or glaciers) sometimes provided some protection from grazing to plants growing among such stones. Under other situations, such as presence of only a few larger stones (up to boulder size), rocks encouraged visitation by wildlife and livestock which rubbed on them. Locals often referred to such large erratics as "buffalo boulders", and the smooth surfaces of such boulders was proof of their having been rubbed by larger animals. Range vegetation around the group of glacial erratics shown in this photograph included more plants of native species than in less rocky areas where Kentucky bluegrass grew ranker, denser, and with greater cover which effectively excluded native species. It was possible that cattle which were grazing this range had been attracted to these rocks (as to use them for rubbing posts perhaps) and had reduced cover of Kentucky bluegrass so that native plants had more opportunity for survival. It was also also possible that cattle avoided the rockier terrain and with less grazing pressure the native plants were able to compete with (or out-compete) Kentucky bluegrass. According to the satate and transition scheme for this range type/range site Kentucky bluegrass increases under both undergrazing or nonuse and underburning (Natural Resources Conservation Service, North Dakota, on-line Field Office Technical Guide). Green needlegrass, Junegrass (Koleria cristata), and silverleaf scurfpea or silverleaf Indian breadroot were the most abundant native range plants in this accumulation of glacial erratics.

Sidebar: It is amazing what gems one can find, especially with the marvelous www. Pertinent to this (and some subsequent) slides is the bulletins, Geology of Emmons County, North Dakota (Bluemle, 1984)and The Geology of the Southern Part of Morton County, North Dakota (Laird and Mitchell, 1942).

116. Sward of Kentucky bluegrass disturbance climax- Two nested photographs (farther- and closer-focal points; latter insider inside former) showing simple (and uniform) structure and composition of a single-species stnd or monoculture of a wheatgrass-needlegrass mixed prairie converted (through a combination of overgrazing, underburning, plant introduction, and, perhaps, a series of very wet years) into a Poa pratensis disclimax.

This was standing crop (peak stage) inside the degraded range shown and described in the two immediately preceding slide-caption sets. This herbage was over 99% Kentucky bluegrass material. Wavelike irregularities in height of bluegrass was the result of some standing (fully erect) shoots combined or mixed in with lodged shoots. Lodging is the breaking and/or bending of shoots under force of wind, gravity, weight of accumulated rain or snow, animal traffic, etc. Lodging is more apt to occur when shoots are comparatively tall (longer than typical), a condition more common when shoot growing conditions are extremely favorable such as with abundant soil water, soil nitrogen, optimum temperature aand/or light, and in absence of grazing (ie. limited grazing or mowing when plants were immature and could develop a more prostrate growth form under defoliation).

Kentucky bluegrass presented in this set of slides had grown under the wettest winter and spring growing seasons that the oldest residents of the area could remember.

Emmons County, North Dakota. Late June-peak standing crop and grain-shatter stageof Poa pratensis. No units of natural vegetation were appropriate and, unfortunately, the Society for Range Management (Shiflet, 1994) had omitted this widespread rangeland cover type. Clayey range site degraded to Kentucky Bluegrass successional state. Northwestern Glaciated Plains- Missouri Coteau Slope Ecoregion 42c (Bryce et al., Undated).

117. Grazing made no difference now- Degree of use (utilization) and appearance of sward of Kentucky bluegrass disturbance climax under light grazing in the current growing season by beef cattle as compared to zero degree of use in the outside fencerow of this degraded pasture. The first of these two slides presented the grazed sward which in this "photoplot" included a more heavily grazed plant of yellow sweet-clover (slightly left of center in photograph).

The second photograph presented the grazed sward of the degraded Kentucky bluegrass range including a typical path made by cattle walking the fenceline (left side of fence) as compared to the ungrazed sward of degraded Kentucky bluegrass range immediately outside the pasture (right side of fence). This right side was highway right-of-way which served as a "ready made" exclosure or "control or check plot". Comparison of left with right side of fence permitted a quick estimate (cursory visual observation) from which differences in degree of grass utilization and species composition could be determined. There were a few plants of smooth brome in the fencerow immediately coutside the pasture (in the "control plot"). There was also an ungrazed plant of yellow sweet-clover that could be compared to the grazed plant of yellow sweet-clover presented in the second slide in the preceding slide-caption set.

The immediately discernable lesson was that exception for the few plants of smooth bromegrass species composition was the same inside and outside the Kentucky bluegrass disclimax grazed by cattle. The rangeland that was previously a wheatgrass-needlegrass mixed prairie had already been so depleted by overgrazing (and, perhaps, some nongrazing), cessation of fire, introductions of Eurasian grasses, and so forth that the current degree of use (light utilization in the present grazing season) made no difference (ie. it was an irrelevant management factor).

The rangeland throughout this area had already deteriorated to the lowest (and, perhaps, permanent) man-made seral stage, at least the lowest successional state shy of annual weeds (just above the pioneer plant seral stage) The net result and the fundamental concept of non-dynamic or unchanging vegetation was identical whether applying the time-honored and first-in-predcident Clementsian term of disturbance climax (Clements, 1936; Weaver and Clements, 1938) or the contemporary term of steady state in the state-and-transition model as widely employed by various groups (eg. Natural Resources Conservation Service). In cowboy lingo, the range went to Hell and it is likely to stay there barring major investments of time, capital, knowledge, and inputs to the land (eg. reseeding, specialized grazing management).

A replicate range in this same "steady state seral stage", another Kentucky bluegrass disclimax, existed on the opposite side of this highway (ie. contiguous except for the highway). Next set set of slides, please.

118. Across the road- Another Kentucky bluegrass disclimax on a former climax wheatgrass-needlegrass mixed prairie immediately across a US highway from the Poa pratensis disclimax presented and described in the preceding four slide-caption sets. The state-and-transition model used by the Natural Resources Conservation Service in its on-line National Range and Pasture Handbook showed this state of departure from the historic climax plant community as permanent to semi-permanent in human time scale (ie. a steady state of range vegetation at a successional stage below the historic climax plant community (Natural Resources conservation Service) or potential natural vegetation (Kuchler, 1964, 1966). This simply the Clementsian disturbance climax (Clements, 1936; Weaver and Clements, 1938) given a new name (and a more detailed version of succession/retrogrsssion): state-and-transition mode. Rosiere (herein) periodically dubbed such long-term seral stages as "stuck steady state(s)".

Kentucky bluegrass did not completely overwhelm the native range plant species on this disturbance climax to the extent that it did on the disclimax range across the highway. There were some surviving (though sparsely dispersed) plants of needle-and-thread which was the only climax dominant represented to any extent. There were even fewer plants of Junegrass, a climax (decreaser) associate. Forbs included silverleaf scurfpea or silverleaf Indian breadroot (probably the most prominent forb in view), upright prairie coneflower or Mexican hat (Ratibida columnifera), purple prairie-clover (Petalostemon purpureus), prairie rose (Rosa arkansana), leadplant (Amorpha canescens), pale false dandelion or pale agoseris (Agoseris glauca), and Lambert's crazyweed or purple locoweed (Oxytropis lambertii).

Height and overall size of Kentucky bluegrass as well as denseness of bluegrass turf was not nearly as great as that of the across-the-road disclimax range. For instance, there was very little lodging of bluegrass shoots on the range shown here. A more open grass sward was undoubtedly a major reason for the greater species diversity on this disclimax range.

119. Disclimax structure and composition- This set of two nested slides (the second photograph being a smaller "photoplot" inside the larger "phoptoplot" of the first photograph) showed the ever-so-slight layering and various plant species of the "steady seral state" of a Kentucky bluegrass disturbance climax on degraded mixed prairie the potential natural vegetation of which was a wheatgrass-needlegrass combination. There were a few widely scattered plants of needle-and-thread and Junegrass (a dominant and associate species, respectively; both decreasers of the Pre-Columbian Northern Mixed Prairie for this range site). Forbs were mostly legumes or composites. More common of these were silverleaf scurfpea or silverleaf Indian breadroot, upright prairie coneflower or Mexican hat, purple prairie-clover, pale false dandelion or pale agoseris, and Lambert's crazyweed or purple locoweed, leadplant. Prairie rose was also present.

On this range in contrast to first range described, which was straight across a highway, Kentucky bluegrass plants were of relatively smaller size with shorter tillers and a lower density of shoots so that the turf was more open (less thick or lush). This condition of a sparser sward apparently resulted in less competition between the exotic Poa pratensis and native range plants so that the latter were comparatively more abundant (less sparse) than on the other degraded range.

There was also less lodging of Kentucky bluegrass tillers on the range shown in these two photographs.

120. Another view of disclimax structure and composition- A second set of two nested "photoquadrants" of a Kentucky bluegrass disturbance climx or steady seral state in state-and-transition interpretation. While most of the herbage was that of the Eurasian Poa pratensis two native grasses, needle-and-thread and Junegrass (a climax dominant and an associate, respectively) were also present (and distinctly visible in the second photograph). Most of the native forbs visible in one of the other of these photographs were primarily composites including upright prairie coneflower or Mexican hat, Flodman's thistle (Cirsium flodman), and Maximillian sunflower (Helianthus maximilliani).

121. Still yet another view of composition and structure- A third set of two nested "photoquadrants" of a Kentucky bluegrass disclimx or seral range vegetation at a steady state as viewed from the state-and-transition perspective. This set of "photosamples" was in a part of this degraded mixed prairie range that supported an array of herbaceous legumes and composites. It was explained above that this second replicate of a degraded, steady state Kentucky bluegrass range had a less dense turf or sward of Poa pratensis that permitted more native plant species than in the other Kentucky bluegrass range (the other replicate) across the highway that had a more tiller-dense and closed sod.

Forbs in these two "phytoscenes" included silverleaf scurfpea or silverleaf Indian breadroot, Lambert's crazyweed or purple locoweed , purple prairie-clover, Flodman's thistle, and prairie rose (this latter species could be described as either forb or low shrub).

The most abundant native grass in this sample of the disclimax plant community was Junegrass, a common associate species in the wheatgrass-needlegrass mixed prairie which is the potential natural (climax) vegetation for this range site.

122. Rocks (and glaciers) again- Range vegetation of a Kentucky bluegrass disclimax ("stuck steady state" or "frozen" successional stage) that developed around a granite glacial erratic. Kentucky bluegrass did not so completely overwhelm the native plant species to the extent that it did on the disclimax range across the highway (the other replicate) which, for whatever reason, had a greater density of bluegrass tillers and a denser turf.

It was shown that there were more species of forbs and more plants of climax grasses on the more open turf of the disclimax range described immediately above (last four slide-caption sets). In addition to several plants of needle-and-thread and Junegrass, native composites and legumes present in this erratic-centered shot were silverleaf scurfpea or silverleaf Indian breadroot, Lambert's crazyweed or purple locoweed, purple prairie-clover, and upright prairie coneflower or Mexican hat.

Glacial erratic or, simply, erratic was defined by the American Geological Institute (Gary et al., 1972) as: "A rock fragment carried by glacier ice or by floating ice, and deposited when the ice melted at some distance from the outcrop from which the fragment was derived. Generally of boulder size, though the fragments vary from pebbbles to house-sized rocks". Granite erratics such as smaller boulder and the ones shown above were transported by glaciers from what is today Canada in the Pleistocene epoch of the Quarternary period, specifically the Kansan, Iowan, and Wisconsin Ice Ages (Fenneman, 1931, ps. 72-76; Fenneman, 1938, ps. 566-569).

123. Down to Kentucky bluegrass- A climax wheatgrass-needlegrass mixed prairie that had been converted into a farm field which was at some point seeded to smooth brome for use as pasture (an improved permanent pasture) and/or hay. Perhaps the land was planted to a tame pasture mix that included Kentucky bluegrass. Regardless of agronomic practices, a stand of smooth bromegrass was established only to be overgrazed to the extent that Kentucky bluegrass eventually replaced most of the smooth brome. That grazing disclimax (disturbance climax) was the pasture being grazed by this commercial cow-calf herd.

There are thousands of acres of Kentucky bluegrass disclimax throughout the Northern Great Plains. Kentucky bluegrass range is a a rangeland cover type, but it was not described by the Society for Range Management (Shiflet, 1994). Exclusion of a Kentucky bluegrass rangeland cover type was perhaps an oversight (maybe there was no author to take on the task of writing it up), but Kentucky bluegrass range (much of it resulting from abuse of existing native range (eg. Northern Mixed Prairie) or mismanagement of introduced (tame) pasture such as was the case presented here. The Kentucky bluegrass range (whether or not it was recognized and published by the Society for Range Management) is a rangeland cover type comparable to those of other introduced Eurasian grasses such as Crested Wheatgrass (SRM 614) and Tall Fescue SRM 804).

Man-made Kentucky bluegrass range such as that shown here can be quite productive, almost assuredly more productive on some sites than some of the mixed prairie range types, but Kentucky bluegrass disclimax range is also considerably less productive than properly managed smooth brome. Performance of cattle seen in these two photographs would have been greater (and probably more profitable) if these animals were grazing the pasture of smooth brome that had, under mismanagement, deteriorated to about the lowest state of forage production possible (short of becoming nothing but a stand of annual grasses like cheatgrass or downy brome and Japanese chess).

This was a pretty or pleasant pastoral scene only to uninformed viewers seeing nothing but sleek, muscular cattle standing on a green graminaceous carpet.The quality of pasture did not match that of the beef cattle.

Campbell County, South Dakota. Late June-peak standing crop and grain-shatter stage of Poa pratensis. No units of natural vegetation were appropriate and, unfortunately, the Society for Range Management (Shiflet, 1994) did not recognize (or describe) this widespread rangeland cover type. Northwestern Glaciated Plains- Southern Missouri Coteau Slope Ecoregion 42f (Bryce et al., Undated).

124. Not there quite yet, but on its way- A former smooth bromegrass pasture that with past and current overgrazing was being degraded into a Kentucky bluegrass disturbance climax. This pasture had not reached the point of being a monoculture of Poa pratensis, but it was approaching that state of deterioration. There were still quite a few individual plants of smooth brome, but these "few and far between" bromegrass plants comprised only a small proportion of total grasss cover and herbage yield of total biomass.

Appearance of standing crop and relative (proportional) foliar cover of the sward was presented in this "photoquadrant" which was representative of the this overgrazed and deteriorated pasture of smooth brome. In this "photographic sample" one large and very conspicuous plant of green needlegrass (slightly left-of-center foreground) had survived the heavy grazing and was representative of one of the dominant species of the climax wheatgrass-bluestem-needlegrass mixed prairie that had been replaced by the monoculture of Kentucky bluegrass.

125. More than differences in degree of use- Fenceline contrast with a former pasture of smooth brome overgrazed to a degraded stand of Kentucky bluegrass on the left and a sward of smooth brome in the outside fencerow that served as the "control plot" on the right. The big difference in herbage was in species composition and not in current degree of use (although any grass in the grazed pasture would be shorter than ungrazed grass outside the pasture). Instead, the difference in height of grass shoots was a product of different species inside and outside the grazed pasture: taller and larger sshoots of mooth brome in the exclosure (fencerow outside the pasture) in contrast to the lower-growing (shorter shoots) of Kentucky bluegrass in the pasture from which smooth brome had been eliminated by longterm overuse which is overgraizng.

Past degree of use (utilization over course of previous grazing seasons) had changed species composition of the pasture with differences seen in this fenceline contrast being the result of elimination of smooth brome due to overgrazing and not atrtrubutable solely to current overuse.

Clearly distinguishable panicles of smooth brome on the right in contrast to panicles of Kentucky bluegrass on the left are undeniable proof of the stark difference in species composition between herbage of the overgrazed pasture (left) and that of the ungrazed outer fencerow (right).

126. The other big forage bluegrass- Canada bluegrass (Poa compressa) is the second most important agronomic Poa species in North America, but although it out-distances the other minor bluegrasses by considerable degree it is still a distant first runner-up to Kentucky bluegrass. Canada bluegrass was given a brief treatment in the Iowa State University Forages (Hughes et al., 1951, p. 266; Hughs et al., 1962, p. 227; Heath et al., 1973, p. 251; Heath et al., 1985, p. 195). Most of these were extremely short pieces (about two paragarph's) and consistently written by the same chapter author. By the fifth and sixth edition of Forages (Barnes et al, 1995, 2007) Canada bluegrass did not receive any coverage in the standard text for Forage Agronomy.

Canada bluegrass merits the treatment accorded it because it is a Eurasian grass introduced by the white man about the same time as was Kentucky bluegrass and one that often grows in association with the far-more important, introduced Poa species. Canada bluegrass is just one more exotic forage grass that gets little more than the proverbial "footnote" in the history of agronomic forages. Like the other, now-minor, introduced pasture grasses Canada bluegrass has naturalized to be of local importance.

Canada bluegrass is now widely distributed in North America with a range from Newfoundland to Alaska and south to Louisiana and the Carolinas. There are at least three improved (artificially selected) varieties of Canada bluegrass: 1) Canon (out of University of Guelph), 2) Foothills (genetic selection from Natural Resources Conservation Service Plant Materials Center, Bridger, Montana), and 3) Reubens (phenotypic selection by Jacklin Seed Company, Idaho [named after Reubens, Idaho]).

These three photographs presented a local stand of naturalized Canada bluegrass at progressively closer camera range until habit and whole-shoot features were featured in the third slide.

Arapahoe County, Colorado. Mid-July; milk to soft-dough phenological stages.

127. Time for anthers out-Canada bluegrass at peak standing crop and peak of anthesis. Appearance of shoots prior to grain formation and set after which shoots begin to dry and lower leaves die as shoot senesecence and plant dormancy begins about same time cryopses begin to form.

Ottawa County, Oklahoma. Early June (in an extremely hot and dry spring).

128. Sex end of the species- Upper portions of shoots including panicles (first slide) and sexual units, panicles on peduncles (second slide) of Canada bluegrass.

Ottawa County, Oklahoma. Late June; grain-ripe, immediate pre-shatter phenological stage.

129. Purplish Canadian blue- General or overall view of panicles of Canada bluegrass. Arapahoe County, Colorado. Mid-July; soft-dough stage of phenology.

130. The small up close- Spikelets of Canada bluegrass. First slide: Arapahoe County, Colorado. Mid-July; soft-dough stage of phenology. Second slide: Ottawa County, Oklahoma. Late June; grain-ripe, immediate pre-shatter phenological stage.

Another study of Canada bluegrass- Three progressively closer views of panicle and spikelets of Canada bluegrass going from upper part of panicle (first slide), branches in portions of that panicle (second slide), and spikelets on the branches (third slide).

Ottawa County, Oklahoma. Early June; peak anthesis.

131. Above and below- A shoot of Canada bluegrass including rhizome and tiller portion. Canada bluegrass is a strongly rhizomatous and, thus, turf-forming species, but the tillers are sparse in density such that the truf is open in appearance and lower in herbage yield. Ottawa County, Oklahoma. Mid-June.

132. Bowed green- Geniculate shoots of Canada bluegrass. The strongly bent culm with swollen or tightly compressed nodes are a readily identifiable feature of this introduced species. Ottawa county, Oklahoma. Late June.

133. The two big blue crops- A fortituous find was the side-by-dide growth of Kentucky bluegrass and Canada bluegrass on a local area of floodplain of the Platte River. In the first photograph a plant of bottlebrush squirreltail (Sitanion hystrix) in center foreground marked separation of a turf of Canada bluegrass (left) and Kentucky bluegrass (right). In the second and thrid slides the turf of Kentucky bluegrass was on the left and that olf Canada bluegrass was on the right. There was even a natural "no man's land" dividing line between these two species in the third photograph. Both of the Poa species are strongly rhizomatous which explained the physical separation; in fact, each of the two turfs were possibly two clonal plants each of its single species.

Fergus (in Hughes et al., 1951, p. 266) noted that "... Canada bluegrass is essentially coextensive with Kentucky bluegrass and is found to some degree in many Kentucky bluegrass pastures", but Canada bluegrass is dominant over Kentucky bluegrass only on harsher habitats (droughtier, more acidic, less fertile, etc.). Canada bluegrass is inferior in forage production and recommended over Kentucky bluegrass only for more marginal lands. Duell (in heath et al., 1985, p. 195) noted that stands of Canada bluegrass were was less dense, stemmier, and considerably less productive (lower yields) than Kentucky bluegrass due (at leaat in part) shorter shoots and fewer leaves especially near shoot bases.

The observation of Fergus (in Hughes et al., 1951, p. 266) that Canada bluegrass is later maturing than Kentucky bluegrass was strikingly evident in these three photographs.

Arapahoe County, Colorado. Mid-July with phenological stages of grain-ripe in Kentucky bluegrass and milk to soft-dough in Canada bluegrass.

134. Ryegrass pasture- The Hordeae or Tritaceae tribe of Gramineae traditionally included the genus Lolium known generically (genus-wide) as ryegrass based on the commonality of the spike type inflorescence. Gould (1975, p. 16) was one of the first agrostologists to include Lolium in Festuceae or Poeae tribe. Recently Barkworth et al. (2007, p. ) in the latest "version" of the "bible" for North American grasses also reinterpreted Lolium as being in Festuceae based on cladistic indicators. The current author adopted the revised interpretation of Lolium based on gross morphology including interpretation of the Lolium flower cluster as a spikelike panicle and obvious close genetic relations with some unquestionable members of the Festucoideae (eg. hybridization between Lolium and Festuca species ). While acceptance of Lolium in Festuceae and not Hordeae relied on Gould (1975) and Barkworth et al. (2007) rather than Hitchcock and Chase (1951) other conclusions reached by Barkworth et al. (2007) such as movement of some Festuca species to new genera was not adopted herein. Likewise, interpretation and distinction between the two taxa of perenne and multiflorum remained an unsettled matter in the mind of the current author.

Wheteher Italian and/or perennial ryegrass is one (or two) species seemed destined for controversy beginning with its name (their names). Hitchcock and Chase (1951, ps. 274-277) recognized five ryegrass (Lolium) species in the United States. All of these were introduced from Europe, four of the five are annuals, and two (or one) are important agricultural grasses. Hitchcock and Chase (1951) distinguished between the two agronomically important species of perennial ryegrass (L. perenne) and the annual Italian ryegrass (L. multiflorum) on obvious basis of life cycle and morphological features, namely "more robust habit" of the latter. This distinction was retained by all major forage agronomists (see for eg. all editions of Forages under authors of Iowa State University [1951, 1962, 1973, 1985, 1995] and the American Society of Agronomy and Crop Science Society of America [Jung et al. in Moser et al. 1996]). Barkworth et al. (2007, ps. 455-456) also retained these two species of Lolium.

Gould (1975, p. 107)) argued that ryegrass in Texas exhibited so much phenotypic variation in plant and spikelet size and in presence or absence of awns that there was only the one species of L. perenne. Gould's departure from traditional nomenclature was not surprising given his radical treatment of so many genera and species (treatments frequently disregarded by other agrostologists), but it does indicate the variation in morphological features and longevity of ryegrass (whichever and however many species there are) in Texas and other southern states. Most importantly is whether ryegrass responds to its environment as an annual, weak perennial, or long-lived perennial. Even this is usually a "mute point" because ryegrass reproduces adequately to persist as permanent pasture. Kucera (1998, p. 46) followed the general interpretation of Gould (1975, p. 107) for Missouri taxa and recognized ryegrass as L. perenne pointing out that" this species intergrades with closely related L. multiflorum". Consequently Kucera (1998, p. 46) separated perennial ryegrass and Italian ryegrass as L. perenne var. perenne and L. perenne var. multiflorum, respectively.

Hitchcock and Chase (1951) stated that L. perenne was the first distinct grass species to be cultivated specifically as a forage crop. Jung et al. (in Moser et al., 1996, p. 605) stated that L. perenne and L. multiflorum are the most important forage and turf grass species in the world and Riewe and Mondart (in Iowa State University, 1985, p. 241) claimed that Lolium "probably is the most widely used of all grasses". L. perenne and/or L. multiflorum are (is) certainly major pasture grasses in the Southeast and in California and Oregon. This use is primarily as an agronomic crop (= tame pasture). However, the species have (has) naturalized over much of the geographic range in which it is planted. They (it) frequently function(s) as naturalized range grasses much like the many Bromus species, wild oats, Johnsongrass, common bermudagrass, King Ranch bluestem, tall fescue, etc. Like many of these (all of those named) ryegrass has become a weed, especially of small grains as far north as southern Kansas.

Ryegrass was included in a publication on range cover types because ryegrass as domestic pasture is often complementary forage to natural pasture. This is most notable in southern range states from Oklahoma and Texas eastward through the humid region where this introduced festucoid grass is adapted. On ranches throughout the southcentral and southeastern states introduced, cool-season, perennial grass species are grown as permanent pasture that enhances or complements the range-forage base of the livestock (or wildlife) firm during late fall through early spring. It was discussed immediately above how tall fescue serves in this capacity. Both of these festucoid grasses are domesticated species and have high economic as well as biological responses to agronomic inputs like fertilizer, lime, intensive harvest, genetic selection, etc. Both species have however, as noted in the preceding paragraph, at least semi-naturalizied such that they frequently persist indefinitely and still produce substantial quantites of forage without the intensive inputs of agronomic management (ie. they have become naturalized range or at least naturalized species on range much like timothy, orchardgrass, smooth bromegrass, and other perennial Eurasian grass species have done in northern portions of the Western Range Region).

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

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

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

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

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

Hamilton County, Texas. May; peak standing crop.

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

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

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

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

137. Ryegrass- Italian or perennial ryegrass (Lolium multiflorum or L. perenne), take your choice. Even as far south as Texas this cool-season grass provides forage through late spring to early summer given adequate soil moisture. Erath County, Texas. June.

138. Flowering shoots of ryegrass- Erath County, Texas. April.

139. Spike of ryegrass- Erath County, Texas. April.

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

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

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

Erath County, Texas. May; anthesis.

141. A follow-up sequential study- Sequence of increasing closeness (or magnification) of Italian or annual ryegrass (Lolium multiflorum) from most of one spike (first slide) through to several individual spikelets on the spike (second and third slides) at peak anthesis. According to Gould (1975, p. 106) there are from five to twelve florets per spikelet in L. perene (the taxon interpreted as including both L. perenne and L. multiflorum).

These spikes were on annual plants growing in a field of winter wheat (Trisetum aestivum) so these annual plants were L. multiflorum according to Steyermark (1963, p. 134) or L. perenne var. aristatum (= L. multiflorum) by Great Plains Flora Association (1986, p. 1189).

Ottawa County, Oklahoma. Late May; peak snthesis.

Grasses- Oat Tribe (Aveneae)

142. Timothy (Pleum pratense) meadow- If there is any domesticated European grass that can lay claim to the historical antiquity and agricultural importance of ryegrass it is timothy. This cool-season member of the oat tribe (Aveneae) remains one of the most important hay grasses in North America. At one time or another many agronomists have stated that it is the number one hay grass in America. For example: "Timothy is by far the most important hay grass in America" (Piper, 1939, p. 140). The Range Plant Handbook stated: "Timothy is by far the most important perennial grass cultivated in North America" (Forest Service, 1940, p. g94). This is clearly the case for the Hay and Dairy Agricultural Region of North America, but overall timothy likely is less important than it's former dominance of the grass hay culture and market. Timothy is still "one of the most winter-hardy cool-season forage grasses" (McElroy and Kunelius in Iowa State University, 1995, p. 305) assuring its continued importance. Timothy is not drought-tolerant so it cannot be grown or persist very far south, but it has naturalized in those parts of North American to which it is adapted. Timothy's value in Range Management is it's inclusion in reseeding mixtures for regeneration of disturbed land (eg. logging road cuts, reseeding old crop fields in northern regions), occasional use as hay crop in range regions, and most of all the fact that it has naturalized and become a permanent part of North American range communities (cf. the Black Hills ponderosa pine type under Forest and Woodland slides). Timothy is one of the 200 species on the Society for Range Management contest list and, as indicated above, was included in the USDA Range Plant Handbook. Timothy is one of the most important Eurasian grasses to become a North American range plant.
Seen in this slide is an old timothy hay meadow that once provided winter feed for livestock in the northern Rocky Mountains and that has persisted without further husbandry. The old hay field is surrounded by forest communities of locally dominant trees like quaking aspen and lodgepole pine with an occasional Englemann spruce indicating the elevational adaptation of timothy.This illustrates that this cool-season introduced species is well-adapted to montane forest range. Teton National Park, Teton County, Wyoming. July.

143. Timothy pasture- Timothy (Phleum pratensis) is one of several Eurasian grass species in the oat tribe (Aveneae) that has been introduced into North America. Timothy is probably the most important hay grass in North America. Timothy also furnishes nutritious grazed forage in permanent pasture in northern latitudes where, as is the case for hay, it is produced by rainfed agricultural practices. This dryland timothy pasture in the Idaho Panhandle was being grazed by stocker cattle and, though at peak standing crop,this festucoid grass was still providing forage of relatively high nutritive value. This high-yielding Eurasian grass complements range and farm feedstuffs and can greatly increase ranch income and profitability. No FRES, Kuchler, or SRM designations for this introduced, agronomic, monoculturally grown, cool-season grass. Latah County, Idaho. June.

144. Timothy at peak standing crop (dough stage)- Ungrazed mature timothy immediately across the fence from the timothy pasture shown immediately above. Latah County, Idaho. June.

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

Jackson County, Colorado. June.

146. Cohort of contracted panicles- A number of contracted panicles on one plant of common timothy. This plant was at the stage of peak anthesis in the western Ozark (Springfield) Plateau. Occasional plants are found in this area, but they do not persist with any grazing or short mowing. The author has seen some timothy plants at edges of prairie hay meadows (here there is never any mowing) that have persisted for over 50 years. Whether these are the same fifty-year old plants or new genotypes that are progeny of older plant could not be determined.

Ottawa County, Oklahoma. June.

147. Inflorescence (a contracted panicle) of timothy (Phleum pratense)- Overall view of entire panicle of timothy. Newton County, Missouri (about the southern and western limit at which this grass can survive summer drought). June.

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

149. Anthers aplenty- Upper portion of contracted panicle of timothy at peak anthesis. Stamen were still held out and up by their filaments. Ottawa County, Oklahoma. Mid-June.

150. Naturalized timothy on prairie bottomland- Timothy appears to have naturalized on this Overflow range site to become a man-made associate to the dominant prairie cordgrass. Both species are doing well and the buffalo do not seem to mind that one is as native as they are while the other is an introduced alien like all the races of Homo sapiens that ever hunted the buffalo.

Custer State Park, Custer County, South Dakota.

151. Redtop (Agrostis alba= A. stolonifera= A. gigantea)- This rhizomatous-stoloniferous member of the Aveneae (oat or timothy tribe) is another European cool-season perennial grass introduced into North America as a domestic forage species. It, too, has naturalized on range especially in the coastal grasslands of northern California where, as discussed under that range type, it and other Aveneae members crowded out the native perennial grasses to create a disturbance climax managed as such.

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

Redtop's story is the same as that of Johnsongrass and, to a lesser degree, species like bahiagrass (Paspalum notatum), the sweet clovers (Melilotus spp.), and the vetches (Vicia spp.). These were the best agronomic forage crops for certain areas at the time of their introduction, but gradually superior forage species were found and the earlier introductions were more or less replaced by later, more productive, introduced species. The "prototype" agronomic forages either died out or, worse, became major weeds (eg. Johnsongrass). Even bahiagrass is a weed where Coastal bermudagrass can be grown because the former has pretty much naturalized while the latter requires intensive farming, including control of bahiagrass, in order to persist. Like the rhizomatous Johnsongrass and quakgrass (Agropyron repens), redtop can become a weed on cropland. It can also be used as a hay crop that requires little management beyond proper haying practices. Redtop hay production is still common in restricted locations especially where redtop is one component of meadow hay.

Redtop can also be very important in riparian vegetation where its rhizomes are outstanding for stablization of creek banks. The Forest Service (1940, ps. g8-g12) Range Plant Handbook a time-honored, always relevant reference that should be a part of every rangeman's library stated that while cattle preferred almost any other cultivated grass to redtop it was "highly regarded as range forage plant" that was "extensively raised as a meadow hay and pasture grass in the valleys of the West" (p. g9). In certain areas of the Western Range nothing has changed since those words were written. As water becomes an ever-scarcer resource and more expensive commodity, crops like redtop that require minimal management will likely become more commonplace perhaps replacing premium hay crops like alfalfa (Medicago sativa) that require more intensive cultural practices and costly inputs of water and fertilizer. The old standbys like the old virtues seem never to be completely replaced.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

154. Red verdure on wet sand- Redtop at peak bloom on a freshwater marsh, a subirrigated meadow, in heart of the Nebraska Sandhills. Redtop is one of the most widely distributed of the Eurasian imports to the North American range. Across the immense expanse of its new home, however, redtop thrives best on its preferred habitat of mesic to wet habitats. This is so much the situation that redtop has been regarded as (or, at least, included in the group of) aquatic plants as, for example, in the manuals by Fassett (1957, p. 115) and Godfrey and Wooten (1979, ps. 93-94).

In the Sandhills of Nebraska redtop commonly grows in association with such wetland graminoids as Scirpus, Eleocharis, and Typha species. In marshes and wet meadows redtop lays claim to mesic more than hydric habitats preferring to let the native grasslike plants claim open, deeper, and longer-standing water. Redtop is not a hydrophyte, one of a group of plants characterized by growing in wet (hydric) environments, but it does best where its roots stay in wet soil and rooting material.

Under such conditions redtop can produce trementous tonages of fairly nutritious hay, but it is often difficult to harvest it on wetlands, especially with contemporary heavy hay-making machinery. Redtop is even more valuable on wetlands--whether riparian zones, irrigation ditches, flooded fields, or wet meadows--as a "foolproof" sod-forming species that provides the maximum protection against erosion. Redtop's extensive system of tough rhizomes and stolons (there is considerable morphological--genotypic and/or phenotypic--variation as regards these shoots) along with its root system permits development of a nearly unerodible sod. This combination of features makes this naturalized perennial grass quite tolerant of trampling and comparatively heavy defoliation.

Cherry County, Nebraska. Mid-June; phenological stage of anthesis.

155. Still to be fully expanded- Young (incompletely expanded or exerted) panicles of redtop. These sexual shoots were growing in the stand seen in the immediately preceding photograph. Cherry County, Nebraska. Mid-June; phenological stage of anthesis.

156. Redtop flowers- Details of redtop spikelets at anthesis. Redtop has naturalized aross much of the Westrn Range Region in more mesic environments such as on river floodplains which was the habitat on which these floral units grew. Larimer County, Colorado. Mid-June.

157. Another redtop set- Series of views of three different units of spikelets on open panicles of redtop. Immature redtop panicles (the compund inflorescnece of grasses) produced on a marsh in the Nebraska Sandhills were presented above. These three slides showed details of branches of those panicles on which there were numerous spikelets each spikelet in turn bore several florets that were in anthesis. There are, on average, three stamens per floret of redtop.

Cherry County, Nebraska. Mid-June; and could the phenological stage of anthesis be doubted?

158. Lower particulars- Details of culm and leaves of redtop shoots that were produced on a Nebraska Sandhills marsh. (A stand of redtop that grew on this marsh was shown above.) Drops of precious rain were beaded on leaves of this introduced--and, now, naturalized--Eurasian perennial member of the Aveneae, oat tribe.

Redtop is one of the older domesticated forages grasses. As could be seen from this photograph redtop produces high yields of leafy hay with a relatively high leaf to stem ratio; that is, when proper hay-making practices are used. If not, it was no fault of this time-honored and true agronomic forages.

Cherry County, Nebraska. Mid-June.

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

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

160. Panicle of reed canarygrass- Phalaris species belong to the Aveneae, the oat or timothy tribe, whose genera have panicle inflorescences. Some of these festucoid genera such as Phleum and Phalaris have contracted or even spikelike panicles. Some panicles of reed canarygrass bear rather long secondary branches coming off of the rachis resulting in regular panicles (as in oats), but typically most are contracted panicles such as these seen here. Idaho County, Idaho. June.

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

Whitman County, Washington. June, early summer.

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

Whitman County, Washington. June, early summer.

163. Spikelets of reed canarygrass- General view of spikelets on reed canarygrass growing on the pasture featured above in the Paouse Prairie region. Spikelets of Phalaris species consist of one perfect floret that is terminal and one or two lower sterile florets reduced to scale-like structures. Whitman County, Washington. June, early summer (late anthesis to post anthesis stage).

From Pacific Northwest to Midwest:

Semi-wild mixed pasture- Pasture od mostly reed cnarygrass with "a taste" of timothy and meadow foxtail (Aloecurus pratensis) along with "a tad" of some caric-sedges (Carex sp.?) and flat-or umbrella-sedges (Cyperus sp.?). This soil was saturated and this about half-deserrt rat rangeman wondered in these cattle would have foot problems.

Sain Louis County, Minnesota. Late June (and summer, but still spring up in these parts).

Outside fencerow- A part of the stand of reed canarygrass presented in the immediately two preceding slides that was on the outside of the pasture showing a population of mature 9floering) reed canarygrass. it was not known if this reed canarygrass was native or if it was of numerous cultivars of introduced (hence, agronomic) Eurasian reed canarygrass. Alhough introduced genotypes or lines of reed canarygrass constitute a major tame pasture grass, the species is alson native to North American prairies and aside from the fact that iti is nearly impossible to distinguish among native and introduced forms or taxa (varieties, cultivated varieties, whaever) thesehave hybridized plus even the cultivated forms have at least partly naturalized.

Anyway, here is what reed cnarygrss can do when it has more water than it kinows whaat to do with.

Saint Louis County, Minnesota. Late June (and summer, but still spring up in these parts).

Mddle America examples Two or more plants of reed canarygrass (first slide) and lots of sexual shoots of reed canarygrass (second slide) growing in a field (used as both pasture and, alternatively) hay in the Red River Valley of southeast North Dakota.

Cass County North Dakota. Late June; late-bloom stage.

Late blooms- Two panicles of reed canarygrass at late-bloom phenological stage produced in a field in the Red River (of the North) Valley.

Cass County North Dakota. Late June.

Reed canarygrass culms- Details of nodes and internodes of reed canarygrass grown as an introduced forage crop in the Red River Valley in southeast North Dakota.

Cass County North Dakota. Late June.

Axillary details- Leaf axil and ligule of reed canarygrass growing in a field used as pasture or, alternatively, hay in the Red River Valley.

Cass County North Dakota. Late June.

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

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

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

Butte County, California. June.

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

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

Hopland Field Station, Mendocino County, California. June.

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

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

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

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

167. Panicles of tall oatgrass- Overall morphology and structure of the panicle inflorescence of tall oatgrass was presented in these two photographs. Latah County, Idaho. June (early summer), hard dough stage of caryopsis.

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

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

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

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

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

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

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

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

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

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

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

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

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

Creeping and foxy-Several plants and their contracted panicles of creeing foxtail (Alopecurus arundinaceous) in a field used as both pasture and hay crop in the Red River Valley in southeast North Dakota.

Cass County, North Dakota. Laate June, peak standing crop.

Subterranean stuff- Root crowns and rhizomes of creeping foxtail growing as both pasture and harvested forage in a field in the Rred River Valley of southeast North Dakota. This is the asexual form of reproduction in this introduced forage grass.

Cass County, North Dakota. Laate June, peak standing crop.

Ripe and shattering- Contracted panicles from which ripe grain was being shed and scattered in the wind of the Red River Valley in southeast North Dakota. This is the sexual form of reproduction in this introduced forate grass. This species was engineered--even without contemporary biotech technique.--probably by both natural and artificial selection.Charles Darwin would be impressed (this author certainy was).

Cass County, North Dakota. Late June, seed ripe stage of phenology.

173. 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.

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

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

175. Flood-irrigated tame pasture of mixed species- This "duke's mixture" of introduced cool-season grasses is characteristic of many spring-flooded bottomland pastures in the northern Rocky Mountain-Cascade Region. Flood meadows such as this once supported such diverse species as willows and cottonwoods along streams and outward a short distance from the riparian zone to native bunchgrasses varying in size from the large Great Basin wildrye along watercourses down (and outward from riparian zones with increasingly less mesic moisture regimes) to Idaho fescue and Sandberg bluegrass with tules or bulrushes and cattails in between. With the coming of pale-faced cattlemen and their white-faced cattle these flood meadows were plowed and planted to first one species (or one mixture) of Eurasian grasses and then to another. Perhaps timothy or redtop was seeded first as a hay grass. Then this was turned under and reseeded to the latest "newest and improved" species like reed canarygrass, smooth bromegrass, or meadow brome (Bromus erectus). Then perhaps the hay field was converted into tame pasture by seeding a third or fourth time to a pasture grass like tall fescue or creeping foxtail (Alopecurus arundinaceus). In many areas of the northern Rocky Mountains flood meadows were seeded to various standard mixtures developed and recommended by Agricultural Extension agents or range conservations with the Soil Conservation Service. A standardized pasture mixture might include varying proportions of two or three grass species like timothy, smooth or meadow brome, or creeping foxtail and a pasture legume like white clover (Trifolium repens). Such mixtures were often specific as to cultivars such as Alta tall fescue, Manchar smooth bromegrass, Climax or Clair timothy, and Ladino white clover. Over time many of these agronomic forage lands (such as the one seen here) became diverse communities consisting of several domestic species plus scattered individuals of natives like Great Basin wildrye and Idaho fescue. Such derived pastures or hay lands appeared stable or even naturalized as long as the soil moisture regimes are maintained.

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

Grasses - Grama or Cordgrass Tribe (Chlorideae)

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

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

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

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

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

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

Erath County, Texas. July.

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

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

Erath County, Texas. September.

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

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

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

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

Erath County, Texas. September.

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

Erath County, Texas. September.

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

Erath County, Texas. September.

182. Inflorescences of bermudagrass (dorsal-view)- A top-down look at the spikelike racemes of bermudagrass in anthesis. Erath County, Texas. September.

For the record: To southern stockmen bermudagrass (in all its cultivar as well as unimporved and naturalized forms) is an outstanding forage crop without peer for cetain environments and stockraising operations. On the other hand, it should be included "for the record" that bermudagrass was ranked as man's second worst weed on a global basis and is perhaps "the most serious weed of the grass family" (Holm et al., ps. x, 7, 25-31). "One man's poison is another man's bread".

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Grasses - Lovegrass Tribe (Eragrosteae)

188. Weeping lovegrass (Eragrostis curvula) pasture- This native from Africa was often planted in Texas and Oklahoma beginning around the late 1930s to early 1940s. Weeping lovegrass is well adapted to dry, sandy soils, drought prone areas, and heavy grazing. Quite a bit of research was conducted on weeping lovegrass to determine best management practices and to develop ways to integrate it into ranching and farming operations in the southern prairies and Great Plains regions. Weeping lovegrass is able to produce so much fair quality forage that by reseeding abandoned fields and depleted ranges to it many growers were able to substantially increase the carrying capacity of their operations. Reseeding a fourth or a third of their land to weeping lovegrass enabled some stockmen to double the grazing capacity of their ranch. What is more, they were able to do this while sparing their native pastures when these were most apt to be damaged by grazing. This increased production and flexibility comes at price of replacing haphazard, lackadaisical grazing management with planned, more labor-intensive management including burning and periodic heavy stocking of weeping lovegrass pastures. These practices are often necessary to maintain forage at juvenile stages and encourage regrowth. Mature clumps of weeping lovergrass are unpalatable and poor quality forage. This is a stand of Ermelo weeping lovegrass that was a type conversion of sandrough scrub to grassland. It is on deep sand and it is obviously a sporadic stand. As explained in the Shrubland Slides sandrough is climax vegetation and it is a marginal range site for weeping lovegrass (ie. there are two lessons to be learned from this scene). Erath County, Texas. July. This introduced species has no FRES, Kuchler or SRM designation.

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

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

Wheeler County, Texas. June.

190. Weeping lovegrass (Eragrostis curvula)- The cespitose habit and downward curvature of leaves for which this species was named were apparent in these photographs. Erath County, Texas. April.

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

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

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

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

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

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

Yavapai County, Arizona. June.

193. Panicle of Lehmann's lovegrass- Coconino County, Arizona. June.

194. Spikelets of Lehmann's lovegrass- Eragrostis lehmanniana is closely related to E. curvula. Gross comparison of spikelet and panicle features of these two species can be compared herein. Coconino County, Arizona.

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

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

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

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

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

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

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

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

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

198. Lehmann's lovegrass at full-bloom stage. This plant was also growing on the Longhorn range presented above.

Presidio County, Texas. June.

199. Thriving in a salt desert- Robust plant of Lehmann's lovergrss growing on a saline, alkaline site in Great SaltLake Desert. This introduced species is tolerant of aridity, random drought, severe soil environments, and heavy defoliation over a broad geographic range.

Tooele County, Utah. June.

Yet another example of Lehmann's lovegrass- One vigerous sprawling plant (first slide) and some of its panicle-bearing sexual shoots (second slide) that was growing in a disturbed understorey of a second-growth ponerosa pine (Pinus ponderosa) forest in the Sierra Blanca Mountains of southcentral New Mexico.

Under certain conditions this sort of invasion of the non-native Lehmann's lovegrass can be amajor weed problem. Yes, a lot of forage, especially for cattle, has been produced, but this can be inferior forage for wildlife species like mule deer (Odocoileus hemionus). The aggressive, robust plants of Lehmann's lovegrass can then out-compete more desirable native range plants including some of those tht are valuable for livestock.

Lincoln County, New Mexico. Late June; peak standing crop and grain-ripe stage of phenology.

Laid overa Lehmann's- Grain-heavy panicles of Lehmann's lovegrass that were bent over with their "burden" of their high yield. Some of the sexyal shoots/panicles of this specimen were over four feet in heigght so it did not take an extremely heavy yield to bowl them down.

The capacity of Lehmann's lovegrass to produce small, readily dispersed sexual propagules like that shown here is one of the major chartacteristics that allowed lehmann's lovegrass to naturalize--and in many cases to become a major wed of the range, especially from standpoint of wildlife habitat.

Lincoln County, New Mexico. Late June; peak standing crop and grain-ripe stage of phenology.

One of its ecophysiological secrets- Spikelets of Lehmann's lovegrass bearing a myraid of viable caryopses. High yields--such as shown in these two macrolens views-- of tiny grains has been one of the principal means by which Lehmann's lovegrass naturalized over a vast region extending from the Rio Grande Plains of Texas across to the Pacific Ocean.

Lincoln County, New Mexico. Late June; grain-ripe stage of phenology.

Another (though lesser-run) introduced lovegrass- Examples of Wilman lovegrass (Eragrostis superba) growing on and stabailizing a highly eroded old field. Wilman lovegrass was introduced from South Africa and has the advantage of producing vigerous seedlings and, therefore, establishing quickly and effectively in range reseeding projects. In fact, that is probably the main use of this short-lived perennial.Wilman's lovegrass does not, however, produce as uniforn and abundant grain yields as do the other two main introduced Eragrostis species: weeping and Lehmann's lovegrass.

Wilman's lovegrass works admirably as a companion species to native grass species in range reseeding operations. Rapid establishment of Wilman's lovegrass stabilizes the partly exposed soil of reseeded areas yet does offer excessive competition to co-reseeded species. In such seedings, Wilman lovegrass is a cover crop not a nurse crop.

Given the limited usefulness of Wilamn's lovegrass (mostly as a cover crop in reseeding of perennial grasses and forbs) as an important forage, this species received limited treatment in standard references and textbooks. Nonetheless, numerous reputable seedsmen offer Wilman lovegrass seed.

Tarleton State University Hunewell Ranch, Erath County, Texas. Late September (late summer); ripening grain phenological stage.

Ondeep sand and ungrazed- Part of stand of Wilman lovegrass growing on deep sand and not grazed (cattle excluded and white-tailed deer did not touch it) showing both how rank it can become and what an effective soil cover it can provide. This was a local area that had eroded and erosion had been stopped in its tracks". Yes, there was some wastage (unused) feed here, but it could be seen as stockpiled and aa a hedge agaiinst dry autumn conditions. .

Tarleton State University Hunewell Ranch, Erath County, Texas. Early September (mid-summer); mid-grain phenological stage.

Ungrazed bunches- Basal portions of ungrazed Wilman's lovegrass. This was another part of the local stand introduced immediately above. This unused herbage would serve as stockpiled forage as isurance against the-always-and-ever-possibility of drought. This slide also showed what an effective soil cover (in this case, deep sand) this bunchgrass provides.

Tarleton State University Hunewell Ranch, Erath County, Texas. Early September (mid-summer); mid-grain phenological stage.

Grain aplenty (and in cute containers)- Panicles with sparsely produced spikelets (first slide) and "nests" of spikelets (second slide ) of Wilman's lovegrass. These organs were on plants that were introduced in the preceding slides.

Spikelets of Wilman lovegrass are some of the more pronounced in lateral compression of spikelets, a key morphological feature in grass identification (Hitchcock and Chase, 1950).

Tarleton State University Hunewell Ranch, Erath County, Texas. Late September (late summer); ripening grain phenological stage.

Grasses- Panicgrass Tribe (Paniceae)

200. Kleingrass (Panicum coloratum) pasture- Kleingrass is another introduced forage species, and one that can be managed as tame pasture or naturalized range. It is probably the most widely reseeded introduced species in Texas. Kleingrass is easily seeded and is quickly established. It produces more and better quality forage than other introduced grasses managed as range (eg. King Ranch bluestem). Time will tell if kleingrass will ultimately become a naturalized introduction that becomes as much, or more, a weed than a desirable crop as is the case for KR bluesetem, Johnsongrass (Sorghum halepense), bahiagrass (Paspalum notatum), and sericea lespedeza (Lespedeza cuneata). Mason County, Texas, June. No FRES, Kuchler, or SRM designations.

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

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

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

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

San Saba County, Texas.October.

202. Field of kleingrass being grown for seed- Erath County, Texas, May.

203. Kleingrass- Example of a typical kleingrass plant in full-flower (peak standing crop). Erath County, Texas. May.

204. Panicle of kleingrass- Erath County, Texas. May.

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

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

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

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

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

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

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

Another exotic panicgrass-Two large specimens of blue panicgrass (Panicum antidotale) represented this rank-growing, warm-season exotic that was introduced from India by way of Australia by the United States Department of Agriculture first in the second decade of the Twentieth Century. The literature devoted to blue panicgrass is very sparse. This was especially so given the considerable acerage reseeded to this introduced species. This was for all practival purposes next to nothing about it in the text, Forages-The Science of Grassland Agriculture. The old standby, Phillips 66 (1963, 2006; p. 47) Range and Pasture Plants remains about the overall best coverage.

Probably the heyday of planting this introduced grass was in the 1950s and early 1960s when it was widely used in range reseeding in the Texas Rio Grande Plains and Edwards Plateau (Gould, 1975, 458). Since that period kleingrass (Panicum coloratum) has largely replaced blue panicgras.

The germplasm of blue panicgrass on record in the United States stems from original grain in Texas and farther to the southwest. This is an accession that was released in 1950 through cooperation by the Arizona Agricultural Experiment Station and the Soil Conservation Service Plant Materials Center in Tucson (Alderson and Sharp, 1994).

McMullen County, Texas. Early October; peak standing crop, ripening grain stages of phenology.

"Blue" shoots- Characteristic shoots of blue panicgrass seen at progressively closer distances to reveal the swollen nodes and prominent midrib (= midvein) of the leaves of this native of India. Blue panicgrass can grow to heights of six to seven feet. Shoots shown here had grown to heights of between four and a half to five foot levels.

Leaves of this exotic, warm-season perennial have a characteristic bluish-gren tinge so the common name is apt.

Examples shown here were growing in their "adopted home" of the south Texas Rio Grande Plains where blue panicgrass can be an amazingly productive forage crop. While blue panic has at least semi-naturalized in the Tamaulipan Plains, the potential productivity of this introduced species is most commonly achieved only with fertilization. (ie. blue panicgrass is a "semi-range" plant grass).

By comparison the exotic panicgrass called guineagrass, which was the preceding introduced grass discussed here, appears to have more thoroughly naturalized in the Rio Grande Plains. "Time will tell."

Blue panicgrass is better adapted and more productive on lighter-textured (clay and silt) soils.

McMullen County, Texas. Early October; peak standing crop, ripening grain stages of phenology.

Shoots galore- Upper parts of sexual shoots with panicles (first slide) and details of some of these upper portions (second slide) of blue panicgrass growing in the Rio Grande Plains of south Texas. The prominent midrib or midvein of the leaf blade is a prominent characteristic of this species. More importantly from a practical agricultural standpoint is the comparatively large leaf blade, a major feature responsible for high herbage yield and nutritive value of blue panicgrass.

La Salle County, Texas. Early October; peak standing crop, ripening grain stage of phenology.

Blue panicles- Several panicles of blue panicgrass were presented to indicate the high proportion that the inflorescence makes up of the shoot in this introduced forage species. Of course, once blue panicgrass (or any forage species for that matter) advances to this state of maturity (full-sexual development stage of phenology) forage nutritive value of the growing plant is at it lowest.

Caryopses in these large panicles (see next slide/caption unit) can add considerably to ntritive value if grazing animals also eat some of them.

McMullen County, Texas. Early October; peak standing crop, ripening grain stages of phenology.

Exotic antidote for the blues- Panicles with nearly ripe caryopses in the blue panicgrass growing in the Rio Grande (= Tamaulipan) Plains of south Texas. As could be expected panicles and spikelets of blue panicgrass are quite similar to those of switchgrass and kleingrass, much more common (and important) range grsses of the Panicum genus.

Heavy seed (grain) yield--at least in "good years"--is a characteristic of blue panicgrass. This has been a major factor in semi-naturalization of this introduced (from India) species in areas like south Texas and restricted parts of the Edwards Plateau. Even with this feature, however, blue panicgrass is not as adapted or as commonly reseeded on range as kleingrass.

McMullen County, Texas. Early October; peak standing crop, ripening grain stages of phenology.

Conclusion: blue panicgrass was another one of those supposed exotic "wonder grasses" that was "discovered" by plant explorers (largely from the United States Department of Agriculture) and imported to the United States of America to "revolutionize" range and/or pasture productivity. Though each of these promising "miracle grasses" was touted as the "botanical Holy Grail" ("this is the one we wanted all along") for range revegetation, livestock forage, soil conservation, etc., most introduced species either "faded from the scene" to persist only as scattered plants as in the case of blue panicgrass or 2) went on to become major weeds like Johnsongrass (Sorghum halapense).

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

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

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

209. Stands of dallisgrass- These two colonylike stands of Dallisgrass are typical of the density and cover of this species at local or microsite scale. Dallisgrass produces its characteristic ascending, hanging shoots with prostrate basal leaves from short rhizomes, but it is a tufted not a sod-forming grass. At densities and foliar cover like that seen here Dallisgrass appears to have a sod-like physiogonomy and growth form. It produces herbage of higher palatability and greater nutrient concentrations than bahiagrass (below), the other major introduced Paspalum species.

210. Dallisgrass panicles- Two panicles atop the drooping or suberect shoots of Dallisgrass.

211. Dallisgrass panicle- The inflorescence of Paspalum dilatum is interpreted as a racemose panicle. The drooping archlike pattern of secondary branches coming off the rachis as racemes (or, more precisely, racemelike branches) is typical of this species. Panicles of other Paspalum species have different patterns including erect racemelike branches. Spikelets of Dallisgrass are frequently infested with various ergots (Claviceps spp.). Erath County, Texas. Mid-July.

212. Preparing for fuit production- Anthesis in dallisgrass. Exertion of anthers was shown on two (racemose) racemelike branches. This specimen was growing in extreme northeastern Oklahoma, but this introduced panicoid species has adapted to local areas as far north as central Missouri, southern Illinois, and southeastern Oregon. Ottawa County, Oklahoma. Late July.

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

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

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

214. Stand of Argentine bahiagrass (P. notatum var. notatum)- Argentine is a cultivar of bahiagrass sometimes known known as "Argentina Bahia" that is slightly superior to Penascola bahiagrass, the traditional "old stand-by", from standpoint of forage digestibility and slightly higher herbage yield. On the other hand, Argentine is much less cold tolerant than Pensacola bahiagrass. Also the Argentine cultivar sometimes causes toxicity due to ergot (Claviceps paspali).

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

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

215. Argentine selection of bahiagrass- Two views of adult plants of the bahiagrass cultivar, Argentine. Even with the more prostrate (vs. upright) growth habit this cultivar has a semi-cespitose feature.

Henderson County, Texas. September; anthesis.

216. Another general view of bahiagrass- Another view of the cultivar, Argentine, of bahiagrass showing--about as good as could be expected--general morphology and characteristic features of Paspalum notatum var. notatum.

Henderson County, Texas. September; anthesis.

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

Henderson County, Texas. September; anthesis.

218. Beauty to a grassman- Details of spikeletes and general features of the inflorescence of Argentine (a cultivar) of bahiagrass. Henderson County, Texas. September; anthesis.

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

Henderson County, Texas. October.

Organization note: An example of an agrosilvopastoral system based on naturalized bahiagrass (Panola cultivar) was presented and described in detail in chapter, Loblolly Pine Forests, while an example of a naturalized Pineywoods woodland or savannah range with a bahiagrass-dominated understorey was included in the chapter, Texas Pineywoods-II (both chapters under Woodlands and Forests).

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

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

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

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

Erath County, Texas. September, peak standing crop

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

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

Erath County, Texas. September, spikelets in anthesis.

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

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

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

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

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

225. What the hay?- Large hay field in a location ectonal between Texas Pineywoods and Coastal Prairies nad Marshes vegetational areas that was a mixture of bahiagrass and Vaseygrass. The history of this cropland was almost certainly known but to God. Row crops, probably including upland cotton(Gossipium hirsutum), and small grains had more than likely been grown on this field in the past. At present it was used for production of hay, quite possibly following use for pasture. A typical forage cropping history of land in this locality was to plant bahiagrass and then, at a later point in time, replant to Vaseygrass (or reverse order), each species of which at the time of establishment was the latest introduced "miracle grass" that promished to revolutionize grassland agriculture and put more "gingle" in the pockets of stockmen. Over time improvead cultuvars of bermudagrass made these two agronomic forages at least partly obsolete and outdated only to eventually naturalize and become weeds.

The result of such cropping adventures and forays into the latest phase of "imporved [intensified] forage production" was naturalized pastures (= naturalized range) that was utilized with minimum inputs, usually mostly exploitative harvest.This is a classic example of that production situation. This beautiful (and remarkably productive) two-species stand was a testimonial to the adaptation of these two naturalized, perennial, panicoid grassees.

Liberty County, Texas. Early October.

226. Naturalized sod- Sward of bahiagrass and Vaseygrass-- number one and number two as to apparent relative cover, respectively--on the large hay field intrduced in the preceding photograph. Crop had been mowed several weeks earlier and shoots shown here were regrowth. Vaseygrass was at grain-shatter stage.

Liberty County, Texas. Early October.

227. Buffelgrass (Pennisetum cillare) range- This species is indigenous from southern Africa to India. It has proven to be the single best introduced species for a combination of good quality forage and successful establishment and maintenance in the Rio Grande Plains and more mesic sites of the Trans Pecos regions of Texas and Mexico.

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

228. Stand of buffelgrass- Close-in view of small part of a buffelgrass stand showing density of shoots and leaf cover that account for characteristic high yields of grazable forage produced by this introduced species.

Cameron County, Texas. October, seed-shatter stage.

229. Buffelgrass plants- Culms, leaves, and infloresences of buffelgrass at maturity and approaching senesence of shoots. The relatively high ratio of leaves to culms-- a key feature of a high-quality forage species --was evident in this ground-level view.

Cameron County, Texas. October, seed-shatter stage.

230. Buffelgrass (Pennisetum ciliare)- The often prostrate form of buffelgrass under heavier grazing (intensive mowing in this example) was presented in the photograph on the left. That on the right was a composite shot with a detached flowering shoot on top of a repeatedly mowed plant. Buffelgrass shoots are tillers, intravaginated shoots, and not stolons, extravaginated shoots, but these tillers are often semi-horizonal with a geniculate (bent like an elbow) culm that has swollen-appearing nodes and that bears strongly compressed or keeled leaf sheaths. This shape and pattern or arrangement of organs results in the shoot appearance shown here.

Pima County, Arizona. June.

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

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

Pima County, Arizona. June.

Location note: images of pastures made up mostly of King Ranch bluestem (Andropogon ischaemum= Bothriochloa ischaemum, often B. ischaemum var. songarica) yet wih bufflegrass at such cover as to vary from local dominant to co-dominant to associate pasture species were pressernted below where discussiing king Ranch bluestem.

Time for an annual- Stand of yellow foxtail (Setaria glauca) that volunteered on a field sprigged to some high-falutin' hybrid bermudagrass which was pretty much a crop failure. The yellow foxtail served as a fortuitous catch crop and the cattleman got some hay. The cattleman had fertilized this field anticipating a"bumper crop" of hybrid bermudagrass. Itstead he harvested a "catch-as-catch-can" hay crop grown from a soil sed bank sown by Mother Nature decades before.

This Eurasian annual is mostly a weed (Whitson et al., 1992, ps. 490-491) in row crops, small grains, and hay fields, but yellow foxtail can be an important forage crop--albeit frequently an emergency or even an unintended one (as shown here). A number of Setaria speccies that are more commonly called "millet" (usually preceded by a descriptive adjective in this common name) have, since antiquity, been grown for their grains which are important food staples in the human diet. Frequently some of these same food-grain Setaria species have been sown as animal fodders, especially as catch crops following drought or late freezes.

A number of the native Setaria species are very important--often major or even dominant--range plants. These are usually designated as this-or-that "bristlegrass" whereas the weedy Setaria species (most of these are annuals and were introduced) are known as such-and-such "foxtail". Even the weedest of the weedy can be important feed sources, both as herbage used as forage and grains that serve as concentrates for birds and rodents (and ungulates when they consume the caryopses during grazing).

Yellow foxtail frequently fits into this "beneficial weed" category or role as it did here in the western Ozark Plateau.

Ottawa County, Oklahoma. Mid-July; peak standing crop and ripening grain stage of phenology.

Roots of an annual adjunct forage- Fibrous roots, root crowns, and basal tillers (vertical intravaginated shoots) of naturalized yellow foxtail growing on a fertilized bottomland field after an intended crop of unadapted bermudagrass failed leaving something Mother Natures abhors-- the vacuum of an empty habitat.

An annual, naturalized range grass--weedy though it was--filled void. The farmer harvested the yellow foxtail for hay, but as shown below there were enough ripe grains to insure the next adjunct hay crop. An example of "sustainability". (Hey (pun intended), got to use a green "buzzword").

Ottawa County, Oklahoma. Mid-July; peak standing crop and ripening grain stage of phenology.

Shoot, details of an annual adjunct- Details of lower shoots of yellow foxtail that grew as a naturalized forage crop on a bottomland field tht had a "spatially interrupted" (sparse or scant) cover of unadapted huybrid bermudagrass. "Swollen" nodes on a somewhat geniculate culm are characteristic of this introduced weed that served as an adjunct range grass.

Ottawa County, Oklahoma. Mid-July; peak standing crop and ripening grain stage of phenology.

Tops of the annual adjunct (or foxy tails)- Contracted panicles of yellow foxtail, a naturalized Eurasian annual, that "magically" appeared on a hay field haaving only a sparse cover of some unadapted hybrid bermudagrass. The sporadic cover (more uncovered than covered) of the intended crop on a rich bottomland soil in the western Ozark Plateau was the proverbial "open invitation" for this Eurasian weedy grass, which under the conditions described here was not a weed at all but instead a happenstance catch crop.

An old farmer's expression was most appropriate: if it weren't for weeds I'd have no crop at all".

Ottawa County, Oklahoma. Mid-July; peak standing crop and ripening grain stage of phenology.

Foxy tails up close- Spikelets on contracted panicles of yellow foxtail, an annual weedy grass, that volunteered on a ferttilized bottomland field with a failed crop of hybrid bermudagrass. The volunteer plants of this Eurasian annual provided the clueless cattleman (a "cattle farmer") with a forage catch crop. Although the yellow foxtail was mowed while most grain was unripe (in early maturity stage) there were enough mature caryopses that an adequate soil seed bank was deposited to insure another crop of this weed--this naturalized, annual range grass--at some opportune time in the future, be it near or distant.

Grain of yellow foxtail adds a concentrate to the forage plant to provide range or pasture animals with a "dietary bonus". These grains are especially valuable to upland game birds like mourning dove (Zenaida macroura) and northern bobwhite quail (Colinus virginianus) which, by the way, are still relatively abundant in this western area of the Springfield Plateau.

Ottawa County, Oklahoma. Mid-July; peak standing crop and ripening grain stage of phenology.

Grasses-Bluestem or Beardgrass Tribe (Andropogoneae)

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

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

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

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

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

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

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

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

Commanche County, Texas. October.

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

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

Mills County, Texas. October.

235. Tame pasture or naturalized range? An old crop field (upland cotton [Gossypium hirsutum] among other field crops) in the central Rio Grande Plains of Texas that had been reseeded to and maintained as a man-made grassland of King Ranch bluestem and (close second) bufflegrass. This was still a field--a hay field--maintained by mowing (harvesting for hay).

In this context, the forage crop could be interpreted as an agronomic stand if it was managed more intensively by fertilization and perhaps noxious plant control. Conversely, the two-grass species stand could be interpreted as an introduced (and, largely, naturalized) range if the only maintenance or management practice consisted of extensive hay harvest without benefit of such agronomic (more-or-less) manipulations as fertilization.

Some students would likely point out that there may have been some genetic selection of the two introduced grass species in this field. This situation does automatically categorize this anthropogenic grassland as tame pasture or an agronomic crop because there have also been many genetic selections (Darwin's artificial seletion) of almost all major range grasses native to North America. (Most of these have been selected and developed by Soil/ Natural Resources Conservation Service Plant Materials Ceners.)

Anyway, the determination was left up to individual viewers. The concept (and conceptual thinking) was the lesson here not some established category (which might alwaysbe somewhat arbitrary).

By the way, hay harvested at this advanced stage of maturity will not be as in high quality as if it had been harvested at some less advanced (less mature) stage of phenology. That certainly includes higher total yield or greater content of harvested nutirients (Total Digestible Nutrients, Digestible or Metabolizable Energy, Crude or Digestible Protein, Vitamin A, etc.).

It is amazing how many producers still have not gotten the message that greater tonage of dry matter does not coincide with greater yield of other nutrients. In far too many cases, 'bragging right" about how many bales rather than how much usable nutrients were harvested prevails. (Same thing of cattle except much more so.) An even more relevant question iwaWhy even harvest hay at all in the mild-climate, essentially year-long, green-grass grazing season of the Rio Grande Plains"? One plausible answer: "Sell hay at exorbitant prices to horse hobbists". These sorts will not know the difference, plus for most sedentary pet hay-burners it will not make much, if any, difference.

McMullen County, Texas. Mid-October; peak standing crop, ripening grain stage of phenology.

236. Pasture (tame or naturalized) mix- Sward of a two-grass species (King Ranch bluestem and bufflegrass) sand that was established on an old field in the central Rio Grande Plains of Texas. The two introduced panicoid grasses were co-dominant in this field of view.

McMullen County, Texas. Mid-October; peak standing crop, ripening grain stage of phenology.

237. Rio Grande Plains version of a verdant pastoral scene- The green of a single-species stand of bufflegrass after a a wet late summer and early fall complete with two sleek Santa Gertrudis cows made for this South Texas Plains rendition of a romantic pastoral landscape. If this naturalized range of bufflegrass had not been surrounded by a horizon-to-horizon brush patch it migh have put the viewer in mind of the emerald pastures of the British Isles and New England, but the brush was present with ever-increasing cover all the time.

OK class stay alert and pay attention (at least stay awake) for the next series of slides...

Just introducing "miracle" grasses is not enough- The following series of six slides presdented five pastures or ranges that had been seeded and were roughly two-thirds King Ranch bluestem and one-third bufflegrass. The first four slides were of four reseeded ranges (pastures) that appeared to have been old fields devoted to row crops like upland cotton (Gossypium hirsutum) that had subsequently been planted to introduced King Ranch bluestem and bufflegrass to be used primarily for beef (mostly feeder calf) production. Establishment of these pastures, which were located "back-to-back" or "in line" with each other (occurred consectively), along a state highway and furnished a visual teaching lesson in which space substituted for time to demonstrate the invasion of honey mesquite into these pastures of exotic perennial grasses.

The fifth range or pasture was located approximately three and a half to four miles away from the four in-line pastures. It was representative of mesquite invasion to the ultimate development of this disturbance climax type. It had become a mesquite woodland. A two-slide/caption set explained this stage of brush invasion.

The exact seeding mixtures of these five cattle pastures probably have been lost from memory because the land appeared to have been sold by cattlemen to absentee land owners for hunting (presumedly for northern bobwhite quail [Colinus virginianus] and/or white-tiled deer [Odocoileus virginianus] or perhaps even exotic game animals). Regardless, the five pastures were essentially of the same composition, again being roughly two/thirds King Ranch bluestem and one/third bufflegrass.

The five ranges--five former cattle pastures or grazing units--represented five different stages of invasion by honey mesquite (Prosopis glandulosa), the only brush species to have invaded at time of photographs. It was not known if mesquite control had been practiced on the first four, the four in-line, pastures previously or if the four pastures had ever been used for hay production. What was obvious was the increasing cover and density of honey mesquite in these four stands of King Ranch bluestem and bufflegrass. The fifth grazing land unit of King Ranch bluestem and bufflegrass was so far gone that it could not even speculated as to its past management, including what--if any-- brush control measures had been used. The undeniable conclusion was that this fifth pasture had become a mesquite woodland, a state of range or pasture deterioration that was far beyond the point of obvious cost-effective brush control measures. This land had undergone a range or forest cover type conversion, an anthropogenic type conversion that could be called a man-made forest.

It is common knowlege that most wildlife--certainly bobwhite quail and, and even more, white-tailed deer--benefit from some woody cover more than do beef cattle. (Of course "everyone" beneficially uses shade and wind blockage afforded by larger plants including that of woody species.) Given this understanding, landowners devoting ranges primarily to--or at least more to--wildlife rather than livestock (at present, mostly beef cattle in the Rio Grande Plains) desire some more woody cover.

The obvious question is, How much more? What has become all too prevalent in the Rio Grande Plains (and everywhere else where wildlife has become the major crop) is such an increase--so much more--of brush invasion that the maintenance or, even, very existance of forage crop is threatened.

Ecological note: invasion was used in the Clementsial meaning of successful plant establishmen and reproduction on land that was "new" or "recent" to that plant species. Invasion was not intended to have some subliminal stimuli (subliminal or hidden message), but instead was employed in a strict scientific and management context. "For the record", King Ranch bluestem and bufflegrass are exotic or introduced (by European man) species; honey mesuite is a native woody species and certainly one with numerous benefits in contexts of Range Management, Wildlife Management, and Watershed Management, this latter expecially in floodplains where honey mesquite is the potential natural dominant (eg. mesquite bosques). Scientific (wise use) management of range resources aims to achieve the proper cover, density, biomass, yield, etc. of all range plants consistent with both economic, cultural, political, and ecological goals. Anytime any plant species exceeds its general abundance beyond these goals (and so as to conflict with scientific management) that plant species is a noxious plant, weedy species, or, most commonly, just plain weed (and weed in a strict scientific management sense). If weedy plants are woody they are brush or brush species.

In the management of pasture species such as King Ranch bluestem and bufflegrassin (the examples presented immediately below) the native honey mesquite either is or is becoming brush. It is brush in a management context because its cover, density, etc. is exceeding wise use levels for management of the ranges or pastures shown. The cover, general abundance, biomass (whatever) is becoming or already is excessive for most wildlife species (other than those which nest in mesquite). Even more relevant in the economic, business aspect of Ranch (the firm) Management, is the fact that honey messquite at brush levels is reducing income from the potential beef cattle enterprise. Ranges or pastures like those described below are adapted to multiple use, use of natural resources for more than one purpose, product, commodity, activity, and so on. It is a waste of the forage resources of these ranges to forgo beef cattle income just to have simplier (lazier) management or a hobby hunting preserve (ie. a tax write-off or tax dodge).

Five ranges or pastures in a case study of honey mesquite invasion of introduced and naturalized grass stands:

238A. Pasture 1; Start of an invasion- Young mesquites almost all of seedling to small sapling size establishing on a King Ranch bluestem-bufflegrass (two-thirds, one third cover, respectively). Saplings were in background whereas mesquite seedlings hardly as tall as the grass shoots were visible in foreground.

It was apparent or seemded plausible that there had been brush control practices on this range or pasture in the not too distant past. Giventhe level nature of this land noxious plant control could have been by mowing or shredding as well as by herbicides. Brush canopy might not yet be at the economic threshold level (stage at which to initiate brush contol practices), but the inevitable was undeniable.

Trouble lies ahead.

Zavala County Texas. Early October; grass herbage at peak standing crop stage.

238B. Pasture 2- Is the grass or mesquite increasing faster? A second pasture or range of two-thirds King Ranch bluestem and one-third bufflegrass on which honey mesquite has invaded. Plants of honey mesquite are all beyond seedling stage and most have progressed to multi-trunk shrubs, some of which were eight feet in height.

Itis time to begin brush control measures. At this cover and density chemical control is most feasible unless there is a favorable time for prescribed burning, including grass herbage being dry enough to burn at a safe period.

Trouble has arrived.

Zavala County Texas. Early October; grass herbage at peak standing crop stage.

238C. Pasture 3- Same question as for the preceding pasture only easier. A third range or pasture and a third "photoplot" of honey mesquite invasion of a grazing land stand that was approximately two-thirds King Ranch bluestem and one-third bufflegass. Brush control is essential while it can still be cost-effective. Both herbicide treatment as well as mechanical removal (grubbing) are still feasible at this degree of cover and plant density. In absence of mesquite control at this or somewhat more advanced stages of mesquite invasion (see next slide) the grazing land will essentially be lost as an economically feasible or financially viable forage management unit.

Trouble getting worse (Double trouble).

Zavala County Texxas. Early October; grass herbage at peak standing crop stage.

238D. Pasture 4- Same question as for the two preceding ranges but reserved for the village idiot to answer. The view ( another "photoplot") of this fourth pasture or range was taken at closer camera distance than in the preceding three pastures to compare size of plants of King Ranch bluestem and bufflegrass to two shrubs or small trees of honey mesquite.

This landowner is on verge of losing his forage crop to brush in the near future. It long past time to invoke brush control measures on this grazing land. Mechanical control is still comparatively favorable to use of herbicides, but chemical control seems more feasible with trees of this size and, more imporantly, extent of tree crown cover at this stage of brush invasion. Prescribed burning would in theory still be an option, but mesquite would almost assuredly be flagged (only lower branches killed while rest of mesquite crown survived). This might be the preferred method if the landowner wanted mesquite like the two close-to-each other plants (shrubs) shown here left for shade.

Trouble big time: major brush control project overdue and needed now. (Maybe nowor never.)

Zavala County Texas. Early October; grass herbage at peak standing crop stage.

238 E. Pasture 5. A whole new set of questions, beginning with what to use this land for, including such considerations as picnicking, hiking in the woods, fire wood harvest, and coon (Procyon lotor) hunting. The ultimate stage of invasion by honey mesquite in a pasture or range that was (past tense; "was" is the operative word) a stand of King Ranch bluestem and bufflegrass (about two-thirds and one third, respectively).

Oh, yes, of course, mesquite control is still possible. Aerial application of herbicide and mechanical control using heavy equipment, including grubbing with a track hoe, could be used to get mesqite control. The practical question is, Would mesquite control be cost effective? This question does not only include the 1) profitability of such practices, but also the 2) opportunity cost involved. Money used for the highly expensive cost of such intensive control could also have been used for any number of other ranch expentures or projects including brush control on land where brush encorachment could be more feasibly treated; other improvements or developments including fencing, water development, and range reseeding, purchase of improved livestock germ plasm, or even sending Susie off to college (to Tarleton Stte University of course).

Furthermore, the state of brush invasion might have reached the stage at which the honey mesquite (now at tree size) might be valuable as, say, for barbeque woodlot, a picnic area, or even bird watching habitat. Of course cattle ranching is still possible, and the mesquite woodland might be used as a movie set for a second version of Rawhide episodes ("Head 'em up, move 'em through [the woods]". Perhaps the mesquite woodland (or, maybe, its even a forest) would be conducive for "wrangling dudes". "Dude ranching" might be too dangerous (litigation-prone), however, given all the mesquite limbs.

The state of range vegetation development could be interpreted as afforestation, "the establishment of a forest or stand in an area where the preceding vegetation or land use was not forest" (Helms, 1998). In other words, it it is an anthropogenic (human-caused) forest or woodland. Such vegetation could be regarded as a disturbance climax, a man-made assemblage of range plants that is basically permanent in practical human time scale. This range plant community is clearly a range or forest type conversion brought about by human action (inaction, that is).

In the words of an old expression your author grew up with, this former pasture or range is a "long gone gosling". Or, in the same context, "Your goose is cooked".

Frio County, Texas. Early October; grass herbage at peak standing crop stage.

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

Carson County, Texas. June.

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

Carson County, Texas. June.

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

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

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

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

Erath County, Texas. September.

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

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

Erath County, Texas. September.

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

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

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

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

Erath County, Texas. September.

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

Erath County, Texas. September.

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

Erath County, Texas. September.

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

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

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

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

249. Now that's a flag!- Johnsongrass at early stage of panicle emergence. Featured in this slide was the typically long (extremely long, in fact) flag leaf of Johnsongrass immediately above the boot from which the upper-most part of panicle was emerging.. The flag leaf is the leaf at base of inflorescence in grasses. The flag leaf seen here was a strong two feet in length.

Ottawa County, Oklahoma. Mid-June.

250. Still in and partly out- Johnsongrass in early bloom stage with one panicle emerging from boot (left shoot in first slide; second slide) and still in boot (right shoot in first slide). Boot refers to the leaf sheath that envelops (surrounds) the unemerged panicle. Typical stand of Johnsongrass.

Ottawa County, Oklahoma. Mid-June.

251. Panicle of Johnsongrass- The great number of exerted anthers in this flower cluster furnished an example of the wind-pollination of grasses in general. Specifically, it illustrated that Johnsongrass is a cross-pollinated species with most flowers producing viable seed.

Erath County, Texas. July.

252. Johnsongrass spikelets in anthesis- A primary branch off of the rachis in a Johnsongrass panicle has three secondary branches with spikelets having exerted anthers.

Erath County, Texas. July.

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

Ottawa County, Oklahoma. July.

254. Important enough for more- Another example of distal (terminal) portion of a panicle branch (first photograph) and spikelets in anthesis (second slide) of Johnsongrass. The color of these spikelets was unique and extremely pretty, at least in the view of this photographer who had looked at Johnsongrass since childhood (six decades as of this photographing).

Erath County, Texas. Late October (illustrating that Johnsongrass will frequently keep producing grain until killed by frost).

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

Erath County, Texas. October.

2562. Rhizomes of Johnsongrass- Rhizomes are belowground extravaginated shoots. "Rootstock" is a confusing synonym for rhizome. (Rhizomes are modified shoots or stems and not roots at all, but the subterranean location and superficial resemblance of these shoots to roots accounts for this old and traditional synonymy.). Daughter shoots come off of the rhizomes by piercing the enveloping sheath such that each rhizome is first a new daughter shoot and subsequently a mother or parental shoot. This is an example of a clonal organism in which each new daughter plant is a clonal unit (= a module) of the same genetic individual. Modular organism is synonymous with clonal organism. The repeating units, the modules, are ramets while the entire genetic individual (the entire plant having the same genotype) is the genet.

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

257. Secrets to one of Earth's most successful weeds (or crops)- Panicles with ripe grain, storage rhizomes, and clumps of roots of Johnsongrass in middle of a cold winter in northern Oklahoma. The Food and Agriculture Organization of the United Nations has long listed Johnsongrass as "one of the world's worst weeds". And guess what, Johnsongrass was the first glyphosate-resistant warm-season grass reported in North America. Does that come as any surprise?

"One farmer's weed is another's crop". To a "poor boy" stockman this naturalized, drought-tolerant, perennial sorghum is a dream-come-true forage crop. The capacity to reproduce--and reproduce copiciously--by both sexual and asexual means is the secret to survivability of Johnsongrass. This photograph showed the "weapons" of survival in the arsenal of this species.

Ottawa County, Oklahoma. Mid-January, dormancy.

258. Rhizomes and roots of a survivor species- Two views of large, reserve food-filled rhizomes and clumps of roots that formed at nodes of short rhizomes wwhich appeared to function more for growth/reproduction than for storage of energy and nutrient reserves. These node-origin roots could be interpreted as adventious roots.

Ottawa County, Oklahoma. Mid-January, dormant phenological stage.

259. One grasses "winter fat"- Plump, food-filled rhizomes of Johnsongrass as they appeared in early winter of the continental climate at degrees latitude. Such rhizomes or, as sometimes labeled, "rootstocks" are organs of both asxexual reproduction and food storage. These underground horizontal shoots are one of the "tricks of the trade" for surviving stressful (eg. hot, often dry physiochemical habitats combined with defoliation).

Ironically one of the most effective ways to kill out stands of Johnsongrass is through frequent, heavy defoliation by mowing or, even more so, grazing.The rhizomes of Johnsongrass are extremely nutrient-rich forage, especially by standards of tropical, panicoid grasses and, typically, in comparison to mid- and later stages of plant maturity. Rhizomes are usually available (other than an occasional pulled-up "plum") only to range animals that are capable of rooting. Hogs, domestic or feral, can devastate dense stands of Johnsongrass quickly and effectively through their grazing, especially of rhizomes. Even heavy grazing by ruminants can be used as a method of biological control against unwanted Johnsongrass. Conversely, maintenance of cherished Johnsongrass pastures requires that this perennial sorghum grazed be grazed judiciously so that it can produce fat rhizomes like the ones shown here in order to survive winters longer and colder than those in which this native of Africa evolved.

Nonetheless, Johnsongrass has naturalized throughout much of the Western Hemisphere (specifically the American Hemisphere or New World) into which it was introduced. Johnsongrass has continued to genetically adapt to its new home in North America (including under conditions where it is a dreaded weed) as, for example, by developing genotypic resistance to the wonder herbicide, glyphosphate. This was but another stage in the "agronomic arms race" between farmer and weeds, a saga tha began when man was driven from the Garden of Eden (Genesis 3:18).

Ottawa County, Oklahoma. Mid-January, dormancy stage of phenology.

Critical clarification: Even though Johnsongrass rightfully earned its reputation as one of the "world's worst seeds" it came in seventh on the list (Holm et al. 1991, ps. x, 7, 54-61) which was a distant runner-up compared to the number two spot held by bermudagrass, (Holm et al., 1991, ps. x, 7, 25-31). Again, "one farmer's crop is another's weed.

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

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

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

Erath County, Texas. May.

261. Johnsongrass at boot stage- The boot stage is the growth or phenological stage of grasses when the "head" (the inflorescence) is enclosed by the sheath of the uppermost leaf. Grassmen refer to the uppermost part of the shoot with the head still inside of the enveloping sheath as the "boot". In the boot shown in this photograph the terminal spikelets of the infloresecence were just emerging from the encasing sheath. Leaves and leaf axils of Johnsongrass were also presented in this slide.

Erath County, Texas. July.

262. 'hopers having a heyday- Immature (an advanced instar stage) differential grasshopper (Melanoplus differentialis) feeding on Johnsongrass. In this locality, differential grasshoppers preferred leaves and even younger parts of Johnsongrass culms to any other species of neighboring grass species including bermudagrass, tall fescue, and even big bluestem (Andropogon gerardii) that was at younger stages of maturity than Johnsongrass. Along road rights-of-way where all these grass species grew side-by-side and Johnsongrass was overwhelmingly selected by grasshoppers.

The remarkable thing was that Johnsongrass grew back rapidly and recovered quickly from heavy defoliation by differential grasshoppers.

Ottawa County, Oklahoma. Late July; sub-adult stage of insect development.

263. Remarkable!- Nearly phenomenal regrowth of Johnsongrass in Severe Drought (Palmer Index) and after wildfire in early summer. During Severe Drought--and for no purpose whatsoever other than "busy work"--a road maintenance worker for Ottawa County, Oklahoma shredded a stand of Johnsongrass growing along a county road (East 120 Road). "Busy body" shredded Johnsongrass to such low stubble heights (from two to six inches) that in Severe Drought all Johnsongrass was topkilled with no "green" (live) tissue remaining aboveground.

Then, around Independence Day some clueless, careless, or vandelous bastard set off fireworks that started a wildfire in the great quantity of shredded Johnsongrass litter. Flames "spread like wildfire" into all of the dead plant material, including standing herbage that had been droughted "to a crisp" in an adjoining hay meadow and in the fencerow beyond the shredded zone. Most fuel responsible for spread of the wildfire was the shredded, , bone-dry Johnsongrass. This "unbaled hay" that had been left after shredding by the stupid "make work" county worker was fuel waiting for a fire.

Shredding and "leaving lay" the hay was human-caused fuel loading, the accumulation of readily ignitable materials. The county "busy body" had laid down a fuel line. The "stay busy", "show-the-voters-you're-doing-something" public servant not only wasted tax money by unnecessary use of expensive fuel (diesel) and wear-and-tear on machinery, but by senseless shredding he also went out of his way--again, at taxpayer expense-- to provide enhanced-flammability fuel for the wildfire.

The Ottawa County volunteer fire department efficiently and quickly distinguished the fire leaving "burnt black" areas like that shown here. Fire suppression costs were also a waste of tax funds and annual dues paid by Ottawa County property owners to the volunteer fire department. By the way, failure by property owners to pay for membership in the volunteer fire association would have resulted in all fire suppression costs being incurred by the landowner who called in the wildfire. Who would have paid if the fire emergency was reported by a passerby who was not a member of the volunteer fire association nor the landowner whose property was burning? Somebody would (and did) pay one way or the other, but the somebody was neither the foolish county employee or the firebug. Note that both of these nonpayers were necessary but, by themselves, insufficient actors in the needless drama.

Nineteen days following the wildfire--started by two idiots--a shower of blessing brought one and three/quarters inchs of rain. Twenty hours after the rain shower the Johnsongrass, which had zero shoot growth above the soil surface since shredding and wildfire, had regrown to the extent seen here. Some regrown shoots (tillers) had attained a height of five inches in twenty hours.

Such potential for plant recovery through new growth (regrowth) is the result of evolution by both natural and artificial (human-induced) selection, the two forms defined by Charles Darwin (1859) in his monumentally influencial book, On the Origin of Species by Means of Natural Selection. This evolutionary adaptation in Darwin's Struggle for Existence (part of the book's subtitle) has rendered Johnsongrass capable of tremendous roughage yield. The high-yield potential of Johnsongrass as pasture crop and/or mechanically harvested roughage should not be overlooked in branding this introduced, pereennial sorghum a weed (though weed Johnsongrass certainly is when unwanted by man or ecologically undesirable).

Ottawa County, Oklahoma. Late July; early regrowth stage of phenology.

264. Rain followed by warm-seson regrowth- Shoots of Johnsongrass, that grew from rhizomes, 20 hours after a 1.75 inch rain. Three weeks before the rain all tillers of Johnsongrass had been killed by two-pronged abuse. First, five weeks earlier (prior to time of photograph) a county road worker shredded four to five foot-tall Johnsongrass to stubble heights ranging from two to six inches. There was some regrowth which was mostly from rhizomes though some new tillers grew from the taller (five or six inch height) shoots that were not killed by shredding. Two weeks after close shredding, a passing motorist carelessly or maliciously threw fireworks into the "hay" (dead Johnsongrass herbage) made by shredding. A wild fire started and burned up the hay and top-killed what little Johnsongrass regrowth there was. This was during a Severe Drought (Palmer Drought Severity Index). Three weeks after the wildfire the 1.75 inch rain fell and Johnsongrass had grown new tillers as seen here. How much more abuse could Johnsongrass stand? What if, after this combination of drought, over-mowing, and total kill of tillers by wild fire, regrowth of, say, two- or three-foot tillers was shredded again?

Recovery of Johnsongrass was a marvel to behold-- at least to this point of abuse. This introduced, perennial Sorghum species is one of the most remarkably defoliation- adapted, tall-growing, grass species to be found in North America.

The tan-colored clump of grass stubble in left center was of dormant tall fescue.

Ottawa County, Oklahoma. Late July; early regrowth stage of phenology.

265. Rapid regrowth following rain- Topdown view (first slide) and sideview (second slide) of Johnsongrass 20 hours after a 1.75 inch rain that broke a Severe Drought (Palmer Drought Severity Scale) and that came after a wild fire had killed all aboveground shoots (tillers) three weeks earlier which the fire itself followed by two weeks the close shredding of the Johnsongrass which amounted to a fire line being laid down. The charred culms are all that was left after the wild fire was started by 1) a stupid, "busybody", county road worker who laid down a fire line when he shredded and then left the four to five foot length, dead Johnsongrass shoots and 2) a careless or mischievous motorist who threw fireworks into the fuel line that had been laid down by the county.

Five weeks after 1) close shredding that killed almost all Johnsongrass tillers and yet resulted in limited grass regorwth and 2) three weeks after the over-shredding a wild fire that killed all grass regrowth, Johnsongrass had grown back to the extent seen here within 20 hours after a one and three/fourths inch rain. High temperature had been about 87 degrees Fahrenheit. Heights of Johsongrass shoots averaged four inches (shoots ranged from little over one inch to more than six inches). Leaf lengths ranged from one half inch to over five inches. Remarkable rate of grass regrowth.

Ottawa County, Oklahoma. Late July; early regrowth stage of phenology.

266. Supposed shredded- A stand of Johnsongrass in a fencerow with a strip shredded to ground level by a rotary blade shredder operated by an apathetic, unskilled county road worker. This excessive defoliation (all aboveground grass shoot material was killed and removed or turned into mulch in the shredding operation) took place during Severe Drought (Palmer Scale) in the fifth consecutive year of drought (varied over this five-year period from Severe to, briefly, Extraordinary Drought).

The next slide showed the amazing adaptation and response of this perennial, introduced panicoid grass to total defoliation...

Ottawa County, Oklahoma. Early June; late spring.

267. Rapid recovery under worst of conditions- Regrowth of Johnsongrass shoots 22 days following close shredding that killed all aboveground parts of a stand and left a bare soil surface (shown in the immediately preceding slide) during a Severe Drought (Palmer Index). Amazingly, Johnsongrass regrew (primarily from large rhizomes and, secondly, from rootcrowns or proaxes) during some of the most trying, harshiest growing conditions ever witnessed by this author in over six decades of expereience in this local area.

Tall fescue (mostly endophyte-infected Kentucky 31) was introduced into this part of the western Ozark (Springfield) Plateau over a half century ago (during the early 1960s). Tall fescue thrived and appeared to have naturalized in this region until it experienced the first multi-year drought since its introduction. Throughout this part of the Ozark Plateau almost entire fields and pastures, and (especially) most local populations of Kentucky 31 tall fescue died out or, at least, suffered drastic die-back-- other than those in shaded or small mesic sites. By contrast, common bermudagrass--most of which had not been planted and instead grew from both grain and vegetative sources of natrualized germ plasm--and Johnsongrass--none of which had been planted and, instead, existed as naturalized "weeds"--increased during the four- to five-year drought of the second decade of the Twentieth-First Century.

The image presented in this and the preceding slide showing rapid regrowth and recovery of Johnsongrass following complete herbaceous defoliation during Severe Drought during the fifth consecutive year of drought was a dramatic example of the remarkable survival capacity and production potential of Johnsongrass as a naturalized forage crop under harsh growing conditions.

** See above under the Festuceae or Poaeae tribe heading the relatively poor adaptation of tall fesuce as a naturalized range or pasture plant species in humid areas that are prone to multi-year droughts when compared with the remarkable drought-survival adaptation of Johnsongrass and common bermudagrass as naturalized, perennial forage species.

Ottawa County, Oklahoma. Late June; early summer. Early regrowth stage of development.

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

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

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

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

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

269. "It'll make a turd"-The field of Johnsongrass shown in the immediately preceding two slides in winter dormancy being grazed by cows and their fall calves. This pasture had been sprigged to Greenfield bermudagrass about 45 years prior to time of these photographs. Johnsongrass invaded immediately (more-or less) the intended bermudagrass pasture and over the intervening time the pasture remained a mixture of these two introduced C4 grasses. Cattle obviously preferred the finner-stemed bermudagrass over the coarse shoots of Johnsongrass, but cattle--especially the lactating cows--consumed some of both grass species.

The terse title of this caption is an old cowboy expression that applied most aptly to this roughage. A half century earlier this author ran registered Herefords on a similar Johnsongrass pasture that was on the opposite side of the county road from the pasture seen here. That pasture had even less bermudagrass than the pasture presented here (same soil and approximate same cropping history). Herefords were wintered as dry, pregnant cows on dormant, weathered Johnsongrass. They received their complete Crude Protein requirement in the form of soybean meal, but no other feedstuffs were offered other than a complete mineral mix ad libitum. The author's cattle like these shown here did fine and gave birth to normal calves. The Johnsongrass pasture for registered cattle was grazed yearlong at light or conservative stocking rates (roughly five and a half to six acres per cow) in this part of the humid zone (average annual precipitation was about 40 inches). At this light stocking Johnsongrass plants plants completed their annual life cycle and entered dormancy well before the likelihood of frost so that prussic acid (hydrocyanic acid) poisoning was not a threat.

The roughage shown in these two slides was the dormant regrowth (following haying) shown in the preceding two slides. Johnsongrass is a tremendous biomass producer, but unlike switchgrass Johnsongrass did not appear in the state of the union speech by President George W. Bush. As a member of the Andropogoneae, Johnsongrass has solid culms so that perhaps it would produce more combustible biomass than switchgrass.

One thing for sure, any beast feeding on (and surviving off of) Johnsongrass roughage like that seen here is a fuel-burning brute. It will indeed make turds. We just no longer have the 'buffalo gals' to gather them for fuel.

Ottawa County, Oklahoma. Early January; dormancy stage (rank, mature shoots and weathered to boot-- and panicle).

Lots of branches- Ungrazed/unmowed Johnsongrass with robust shoots that had developed secondary shoots off of the main tillers. These iller branches had developed from intercalary meristem even though there had been no defoliation to remove apical meristem and eliminate apical dominance. Under favorable growing conditions Johnsongrass will exhibit this form of extremely abundant--though relatively uncommon--herbage production. Your author witnessed this highly productive growth pattern as a young man when he ran registered Herefords on Johnsongrass pasture under light grazing. Conservative grazing (low degree of use) was a hedge against drought and eliminated need for hay feeding (even with heavy snowfall).

Ottawa County, Oklahoma. Late July, mid-summer.

A Domesticated Native

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

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

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

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

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

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

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

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

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

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

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

Summary reference- The definitive reference on eastern gamagrass as a domesticated forage species was Springer and Dewald (2004).

272. Boys and girls on the floral playground- Staminate spikelets (above or distal) and pistillate spikelets (below or proximal) on rames or racemose primary branches of inflorescences of eastern gamagrass. This basic inflorescence consist of one to three (usually two) spicate racemes (Hignight et al., 1988, p. 8) or racemose branches (Gould, 1975, p. 622). These racemose primary branches (from the interperetation of the basic inflorescence as a panicle) are often called rames in the bluestem tribe, Andropogoneae in which some workers include Tripsacum and Zea species.Other agrostologists regard the monecious genera that have separate clusters of male and female spikelets in the same flower cluster (inflorescence) as being in their own tribe which is designated as Maydeae or Tripsaceae. Details of this specialized inflorescence were given by Gould (1975, p. 622), Barkworth et al. (2003, p. 693), and Shaw (2012, p. 979).

Ottawa County, Oklahoma. Early June; peak anthesis.

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