Tallgrass Prairie (Interior) - IB

Dominant, Prominent, Major, and Otherwise Interesting
Range Plants of the Tallgrass Prairie

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Grasses of Tallgrass Prairie

1. Texas bluegrass (Poa arachnifera) in the Texas Western Cross Timbers- Throughout the southern two-thirds (or more) of the North American tallgrass, true, mixed, and shortgrass prairies and plains warm-season grasses are far more pre-dominant and important than cool-season species on basis of species number and biomass production. This is more the case the farther south grassland vegetation extends because numbers of panicoid and eragrostoid species increase while festucoid grasses decrease with southward progression (conversely the proportion of festucoid species increases with northward progression in the continental grasslands). Usually, however, there are some warm-season grasses in the north and some cool-season grass species in southern grasslands. Texas bluegrass is one of the these southern cool-season grasses. It is one of the more conspicuous and locally dominant festucoid grasses in the southern latitude grasslands. This dioecious rhizomatous species sometimes forms exclusive colonies at microsite scale especially on deep sand sites as seen here in the Cross Timbers of northcentral Texas. These colonies can be bisexual or largely monosexual suggesting that like buffalograss they are monoclonal (consisting of one genotype or genetic individual as a clonal organism). Two things are certain about Texas bluegrass: 1) it is a species that is highly palatable to grazing animals and 2) it is perceived by human eyes to be an extremely attractive plant. In regards the second point, many native plant fanciers rank Texas bluegrass among their favorite prairie species which gives this grass value for natural landscaping. The rancher and landscaper find common cause in this unique grass.

 

2. Texas bluegrass (Poa arachnifera)- A climax cool season species of the tallgrass prairies and Cross Timbers of central Texas. Prairie hay meadow, Erath County, Texas, April.

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Gals' group- A large colony (almost assuredly, clonal) of female Texas bluegrass. This was an unusually large assemblage of shoots (plants?) of a single sex of Texas bluegrass. Although it might be suspected that apomixis occurs in dioecious species like Texas bluegrass, there has never been documentation of this form of asexual reproduction in Poa arachnifera (Casler et al., 2003, p. 61).

Hunewell Ranch, Errath County, Texas. Mid-April; peak exertion of stigmas.

 

3. Girls' git-together- Colony of female sexual shoots of Texas bluegrass. This was a local group of clonal units of the same female plant (one genetic individual) growing in the West Cross Timbers of northcentral Texas.Erath County, Texas, Late April; peak bloom stage.

 

4. Soparno duet- Two sexual shoots of female Texas bluegrass in the colony introduced in the immediately preceding photograph. Can there be any doubt that this is, simply, one of the beautiful native grasses of southern portions of the Tallgrass Prairie Region?

The author has always tried to encourage the wise-use conservation, restoration, establishment, and enjoyment of range plants, especially the most valuable or readily enjoyed native species. This includes such ancillary uses as prairie landscaping and planting of natives as ornamentals. There are few, if any, native prairie grasses in the areas where Texas bluegrass naturally grows that fit the bill any better than this cool-season, dioecious grass. In addition to such obvious uses as forage for livestock and wildlife, Texas bluegrass makes a beautiful lawn species such as the examples shown here. If ya'll are so fortunate as to have in your lawn take note of the obvious: DO NOT MOW IT UNTIL IT GOES DORMANT. Why would any rational person opt to shred off such lovely panicles? It would be the equivalent of a woman with beautiful flowing locks shaving her head.

Erath County, Texas, Late Aprill; peak bloom stage.

 

5. Inflorescence of female Texas bluegrass- The female panicle of Texas bluegrass is larger than the male, but both are quite attractive. Texas bluegrass shares the common diagnostic feature of the florets of Poa species: the "cobwebby"lemma of the pistillate florets due to presence of cottony like hairs on the callus of the lemma. The specific epithet, arachnifera, refers to arachnid or spider.

Erath County, Texas. April.

 

6. Showy Texas ladies- Female spikelets of Texas bluegrass. A view that preented the beauty of Texas bluegrass in bloom. The pistillate spikelets are bigger and more showy. Girls are supposed to be showier (sweeter, too). Erath County, Texas. Mid-April.

 

Moth on a male-Panicle of male Texas bluegrass and a lepidopteran visitor (first slide) with closeup view of adult grass tubeworm moth (Acrolophus cressonii) on spikelets of a panicle branch (second slide). A study in microhabitat. Somebody's whole world on a grass flower. Life is an amazing set of phenomena.

Erath County, Texas. Late April; late anthesis phenological stage.

 

7. Male plant of Texas bluegrass- Flowering of male Texas bluegrass just before anthesis. Vernal aspect, April. Young County, Texas.

 

8. Junegrass- Prairie hay meadow, Burkhart Prairie, Newton County, Missouri. June.

 

9. Inflorescences of Junegrass- Tallgrass prairie hay meadow. Newton County, Missouri. June.

 

10. June-bloomers- Junegrass (Koleria cristata= K pyramidata= K. macrantha) growing with porcupinegrass (Stipa spartea) and broomsedge bluestem (Andropogon virginicus) on tallgrass prairie in western Ozark Plateau. Occurrence of Junegrass with porcupinegrass on this tallgrass prairie indicated the ecological affilitation of tallgrass prairie with true prairie. Junegrass is festucoid species in of the oat tribe (Aveneae) and has a semi-contracted or narrow panicle.

Junegrass is a much more important species in the true and mixed prairies than in tallgrass prairie. Clements (1920, ps. 121-124) interpreted the true prairie as the Stipa-Koleria Association with Junegrass one of five species forming consociations (though this was less common for Junegrass). It was Junegrass' widespread distribution and association with most other common grass species that distinguished K. cristata as a dominant of true prairie which, even at the time Clements (1920) described it, had been largely eliminated by farming. Clements (1920, ps 131-134) and Weaver and Clements (1938, ps.520-521) regarded the bluestem-Indiangrass (Andropogon species as then interpreted)-dominated grasslands eastward of true prairie as subclimax prairie, the Adropogon associes. Today, Clements' subclimax is described as the tallgrass prairie (Weaver and Clements, 1938, 520-521). Junegrass has also long been regarded as a dominant of the mixed prairie (Clements, 1920, ps. 135-138; Weaver and Clements, 1938, p. 523) in which it forms consociations. Clements (1920, p. 138) regarded Junegrass as about mid-range in the array of dominants with regard to water requirements.

In essence, Junegrass is one of the most widely distributed native grasses theoughout North America, occurring from Canada's Northwest Territorities to Quebec and south and east to Alabama, but it almost never abundant other than as local consociations.

Newton County, Missouri. Early June; peak bloom stage.

 

11. Narrow pyramids- This three-slide sequence presented at consecutively closer camera range examples of the narrow or semi-constricted panicle with two-flowered spikelets of Junegrass at anthesis. These panicles were on the same tallgrass prairie as in the preceding slide. This meadow had been used for production of prairie hay for close to a hundred years.

The arthropod atop the first panicle was a female American dog tick (Dermacentor variabilis) patiently waiting for her victim. Vegetation in humid and subhumid regions is often literally crawling with ticks such as this specimen throughout much of the summer. The photographer left this one hungry, but she should have not been completely disappointed as she got this publicity shot.

Newton County, Missouri. Early June; peak bloom stage.

 

12. Done for the year- Fully mature and sensecent plant of Junegrass with panicles full of ripe grain. Junegrass is a perennial bunchgrass, a cespitose (tufted or bunched) species. The first slide presented five separate plants while the second photograph showed a single individual to emphasize the bunched habit of this member of the oat tribe. All of these plants were growing on a relict of tallgrass prairie in the western Ozark Plateau that was within the Prairie Peninsula. An oak-hickory forest was just behind the small glade-like opening of prairie that was prime habitat for these two and a half foot tall plants.

These photographs were taken five days--and on a drier microhabitat--later than those in the two immediately preceding slide sets.

Newton County, Missouri. Early June; grain-ripe and early dormancy stage.

 

13. Ripe and ready for shedding- Narrow panicles of Junegrass with two-floret spikelets at grain-ripe/shatter stage. The group of panicles shown in the first photograph were on the plant presented in the last preceding photograph immediately above. The second photograph was a close-up of a portion of one of these semi-contracted panicle.

Small tallgrass prairie situated within an oak-hickory forest in the Prairie Peninsula (western Springfield Plateau). Newton County, Missouri. Early June; grain-ripe/shatter phenological stage.

 

14. Another cool member of the oat tribe- Prairie wedgegrass or wedgescale(Sphenopolis obtustata) on a tallgrass prairie in the Prairie Peninsula of the western Ozark Plateau. This approximately 60 acre prairie had been used as a hay meadow for nearly a century or, perhaps even longer as its history of haying was lost with the ghosts of haymakers past. Big bluestem was the dominant of this meadow vegetation. The cool-season prairie wedgescale could survive with the larger, more competitive, more water use-efficient members of the bluestem tribe (Andropogoneae) by completing its life cycle before the tallgrass species got much more than started on the rapid- growth phase of their annual cycles.

This striking and conspicuous plant is an uncommon--though a widespread--festucoid species. prairie wedgegrass is most at home on the virgin sod of central prairies, but it grows from Maine across to British Columbia and from the Sonoran and Chihuhuan Deserts across to northern peninsular Florida.

Newton County, Missouri. Early June; late anthesis.

 

15. Cool-season panicles on the prairie- Sexual shoots with panicles (first slide) and a single (individual) panicle of prairie wedgescale or wedgegrass growing on a tallgrass prairie in the Prairie Peninsula. Grasses are by-and-large determinate bloomers meaning that flowering begins with the uppermost and outermost spikelets (in contrast to lowermost and innermost flowers in indeterminate species). In panicles seen here flowering had advanced to the lowermost spikelets with the most distal (uppermost or "highest") spikelets having completed flowering and ferttilization and progressed to soft-dough grain stage.

Newton County, Missouri. Early June; late anthesis.

 

16. Wedged spikelets- Details of a branch of the panicle (first photograph) and opened florets on this branch (second photograph) of prairie wedgescale or prairie wedgegrass. This was a lower branch off of one panicle such as the one presented immediately above. The open (blooming) florets of spikelets on this panicle branch were the upper- or outermost ones indicating that this particular unit of the inflorescence had just begun anthesis. This was a closer-in look at determinate flowering.

Newton County, Missouri. Early June; early anthesis.

 

17. Sexually explicit- Details of florets in spikelets on a branch of a panicle of prairie wedgescale. Anthers and stigmas were exerted (ie. "letting it all hang out" in the vernacular of The Hombres). Newton County, Missouri. Early June; early anthesis.

There are a number of wildryes (Elymus spp.) that are important (even locally dominant) cool-season perennial grasses on tallgrass prairie. Three of the more common and widespread species in the heart of "tallgrass country" and adjoining hardwood forests and savannahs (including the famed Prairie Peninsula) were presented below. Elymus species ae in the wheat or barley tribe (Tritaceae or Horedae).

 

18. Stand of Canada wildrye (Elymus canadensis)- Large, dense population of Canada or nodding wildrye growing on a floodplain of a branch of the Bosque River in the Western Cross Timbers-Grand Prairie vegetational area of northcentral Texas. Many shoots in this beautiful, luxuriant stand were bent over from rush of recent flood waters. "Pure" (single-species) stands of this size are quite rare because this species does not usually form populations or colonies over large areas as do some species on tallgrass parairie (eg. big bluestem, little bluestem, prairie cordgrass, purpletop).

Erath County, May; peak standing crop.

 

19. Nodding along an ehemeral stream- Several plants of Canada or nodding wildrye with characteristic features and growing in its "prime habitat", moist to wet soil in a fairly cool, shaded locale. These plants were part of the large stand of this species that was shown immediately above growing on a floodplain of the Bosque River. This is the typical habit of this species with the large spike inflorescences "nodding" (drooping or bending over at peduncle, the juncture of inflorescence and rest of shoot).

Erath County, May; peak standing crop.

 

20. Starting to nod over- Examples of spikes of nodding or Canada wildrye that were beginning to bend down (or over) at stage of anthesis. These flower clusters will continue to bend farther downward as the sexual shoots of these plants get ever drier near end of their annual cycle and as spikelets in these spikes get heavier with growing, ripening grin.

Erath County, May; anthesis.

 

21. Virginia wildrye (Elymus virginicus)- A few plants of Virginia wildrye had grown to large size on a protected microsite of a tallgrass hay meadow on the Springfield Plateau of the Ozark Mountains. These plants were growing on the backside of a motte of persimmon where hay-making equipment could not enter. Late haying (August or September) over a span of nearly half a century had greatly reduced decreaser tallgrass species on this meadow, but dormancy (usually) or low-growing, early shoots (sometimes) at late summer harvest permitted plant of this cool-season species to thrive amid fierce grassland competition.

Ottawa County, Oklahoma. June (early summer); late anthesis to milk stage of grain.

 

22. Spikes of Virginia wildrye- Sexual shoots of the Virginia wildrye plants that "paraded" in the immediately preceding photograph. Awns of Virginia wildrye are conspicuously shorter than those of other Elymus species typically found on tallgrass prairie and Prairie Peninsula ranges. Compare relative lenghts of these awns to those of the species introduced before and after E. virginicus.

Ottawa County, Oklahoma. June (early summer); late anthesis to milk stage of grain.

Some more examples- Shown immediately below were more photographs of shoots and spikes (whole-inflorescence shots) of several individuals of Virginia wildrye growing in an oak-hickory savanna in the Ozark Plateau just east of the Cherokee Prairie of the Central Lowlands physiographic province.

Ottawa County, Oklahoma. July.

 

23. Study of an understorey grass- Three-slide sequence showing sexual shoots of Virginia wildrye (Elymus virginicus) in forest understorey in western Ozark Plateau. Peak standing crop with grain in mid-dough phenological stage. Ottawa County, Oklahoma. July.

 

24. Spikes in the shade- Examples of spikes of Virginia wildrye in understorey of western Ozark Plateau forest at mid-dough stage of phenology. Ottawa County, Oklahoma. July.

Another wildrye- Presented next was silky wildrye (Elymus villosus), another locally abundant cool-season species on tallgrass prairie. Specimens shown in this short section were growing on the same range as the examples of Virginia wildrye presented immediately above. These two very similar Elymus species can be distinguished based on curved vs. not curved glumes (Kucera, 1961, p.73; Kucera, 1998, p. 111), but they can be differentiated even more quickly on the range based on complete emergence of the spike from the boot in E. villosus in contrast to completely exerted (and generally larger) spikes in E. virginicus. (It is remarkable how often specialists overlook obvious, stark differences and instead "split hairs" on minutae.)

 

25. Silky wildrye (Elymus villosus)- When discussing the central grasslands of North America most students of the range typically refer to Canada or nodding and Virginia wildryes as the major Elymus species. A lesser known or recognized wildrye growing over much of the tallgrass prairie, Prairie Peninsula, and open understorey oak- hickory forest is E. villosus, silky wildrye. With the longer-awns and larger spike of nodding wildrye and the vertical or "straight" spike of Virginia wildrye silky wildrye looks very much like a hybrid of the two more commonly recognized prairie species. Silky wildrye is, however, a species "in its own right" and one that is often much more common than the two better known Elymus species. These plants and those presented immediately below were growing in a part of the Prairie Peninsula at the western perimeter of the Ozark Plateau.

Ottawa County, Oklahoma. June (late spring); peak anthesis.

 

26. The beauty of reproduction- To the agricultural producer nothing is as exciting as the promish of reproduction, be it a developing, pregnant female or blooming "ears" of grain. Infloresecences of silky wildrye at peak anthesis are a vivid example of the sentiment expressed by Senator Ingalls of Kansas when he wrote that the "homely hue" of grass was "more enchanting than the lily or the rose".

Ottawa County, Oklahoma. June (late spring); peak anthesis.

 

27. Another native, cool-season, prairie grass- Winter bentgrass (Agrostis hyemalis) on moist to wet local habitat on tallgrass prairie in western Ozark Plateau. This is a minor species other than in local mesic microsites. It was included here to give an idea as to the great species diversity of tallgrass prairie. Even in southern areas native, winter, grasses grow alongside warm-season, perennial, panicoid grasses. Winter bentgrass has generally been regarded as a short-lived perennial (Gould, 1975, p. 140). It grows on both interior and coastal tallgrass prairie.

Ottawa County, Oklahoma. Late May, maturing grain to early grain-ripe phenological stages.

 

28. Dainty panicles on tallgrass prairie- Panicles and spikelets of winter bentgrass. These flowers were on some of the same plants shown in the immediately preceding slide. Ottawa County, Oklahoma. Late May, early grain-ripe stage.

 

29. Two examples of little bluestem colonies- Little bluestem, prairie beardgrass, broom beardgrass, or common beardgrass (Andropogon scoparius= Schizachyrium scoparium) was probably the single most important range grass across pre-Columbian North America. This was likely true based on geographical range of the species, the total quantity of animal feed it produced (say, measured as Animal Unit Months), and the land area (acreage) on which this species was the dominant or co-dominant plant species. Hitchcock and Chase (1951, p. 754) showed little bluestem as occurring in every state of the Union except Nevada, California, Oregon and Washington. Little bluestem is found in the three Canadian Prairie Provinces (Looman and Best,1987, ps. 96-98) and as far eastward in Canada as Ontario. It occurrs in the Chihuhuna Desert, but not in the Sonoran Desert.

With loss of most of the original tallgrass, true, and mixed prairies (and the more favorable sites therein) to the plow much of the original cover of little bluestem was destroyed. Less of the virgin range dominated by blue grama (Bouteloua gracilis) was plowed so a shortgrass species that was substantially less widespread and lower-yielding has now likely surpassed the bluestem and other tallgrass prairie species in importance as a range plant. If the contemporary interpretation of "little bluestem" is accepted such that this is a complex of taxa that were formerly interpreted as being distinct species the enlarged S. scoparium may still be the most important range plant in North America based on the criteria specified above. Examples of bluestems or beardgrasses formerly treated as distinct species by agrostologists like Hitchcock and Chase (1951) that were reinterpreted as subspecies or varieties and "lumped" in the enlarged, "umbrella" S. scoparium by more recent taxonomists like Gould (1975) and Allen (1992) included pinehill bluestem (Andropogon divergens= S. scoparium var. divergens), seacoast bluestem (A. littoralis= S. scoparium var. littoralis), New Mexico little bluestem (A. neomexicanus= S. scoparium var. neomexicanum).

Little bluestem often forms naturally occurring single species stands, the Clementsian consociation, over large areas and is a co-dominant (or at least a major species) on many range sites. Dense populations of this least mesic of the Four Horsemen of the Prairies species often occur as vast natural "fields". Such stands are extremely productive of biomass and serve as examples of why defoliation of herbage whether by grazing animals or fire is so important to mainteance of grassland. The heavy yields of biomass and high density of tillers in these two populations will result in excessive accumulations of dead plant material unless reduced by grazing and/or fire. The build-up of excess mulch will effective exclude light from basal portions of the little bulestem plants the following spring and delay "green-up" and retard plant growth and herbage production during the ensuing growing season.

Both stands of little bluestem shown in these photographs were in the Western Cross Timbers and Prairies vegetational area of northcentral Texas (Erath County) and at peak standing crop. The stand in the first slide was on a Bottomland range site in late fall (the latter part of October). The stand in the second slide was on a Rolling Prairie range site in anthesis during early fall (late September).

 

30. Little bluestem plants- The cespitose habit of little bluestem was illustrated by these plants growing on a Rolling Prairie range site in a portion of the Grand Prairie in the Western Cross Timbers and Prairie land resource area of Texas. Unlike big bluestem, Indiangrass, and switchgrass (the other three of the Four Horsemen tallgrass species) little bluestem is not rhizomatous. All secondary shoots are intravaginal (vertical shoots designated as tillers or often, among farmers and stockmen, "stools"). As such little bluestem is never a sod-forming grass although individual plants (genetically distinct individuals) frequently grow so close together as to form a prairie sod or dense sward.

 

31. Single plant of little bluestem- Little bluestem is the one common dominant grass of the tallgrass prairie that is strictly cespitose (ie. a bunchgrass or tussock grass). Older individual plants form large tussocks like this one which had a basal cover of over two and a half feet feet. This growth is in effect asexual reproduction by increase in numbers of tillers (intravaginal, hence upright, shoots). Little bluestem also flowers prolifically (shown in slides below), but this sexual reproduction is generally not as efficient as asexual reproduction. This is especially the case in established swards. Most of the tillers of this plant had advanced phenologically to become flowering shoots.

Portion of Grand Prairie on Tarleton State University Hunewell Ranch, Erath County, Texas. September.

 

Full fall glory- Large plant of little bluestem (four foot tall) on a bottomland range site in the Edwards Plateau. This is an example of the size (both height and diameter as determned by number of tillers) to which little bluestem can attain under grazing deferment management, a productive range site, and a favorable growing season. As part of grazing managent plan this large plant had been ungrazed in the current growing seaaon.

Grass plants of this size and stage of maturity can become wolf plants in the next grazing season. A proper prescribed fire will remove this dead, unpalatable herbage and return its nutrients to the soil to start a fresh grass crop. This is especially important practice when running stocker cattle on range where high levels of individual animal performance are essential for profitable rate of gain under negative market pricing structure (where larger cattle bring lower prices per pound than lighter ccattle which is the usual situation).

March Ranch, Coke County, Texas. Mid-October.

 

32. Flowering shoots of little bluestem- Both anthers and filaments of stamen as well as stigma were visible on the sexually reproductive tillers of little bluestem presented in these two slides. Tarleton State University Hunewell Ranch, Erath Cpounty, Texas.. September.

 

33. Anthesis in little bluestem- This close-up view of staminate and pistillate organs in little bluestem was presented as another example of flower structure in the Gramineae. Tarleton State University Hunewell Ranch, Erath County, Texas. September.

 

34. Inflorescences of little bluestem- Sexually reproductive shoots of little bluestem at seed-ripe stage. Inflorescences of the bluestem or sorghum tribe (Andropogoneae) have typically been interpreted as racemes. Chase (1964, p. 82) described the arrangement of the Andropogon flower cluster as having racemes "borne on numerous slender leafy branches arising in the axils of leaves on the main culm or branches, the whole forming a compound inflorescence". Other workers like Highnight et al. (1988, p. 8) labeled this inflorescence type as a spicate raceme. The rachis is jointed and falls apart upon maturity resulting in the shattering of grains. Tarleton State University Hunewell Ranch, Erath County, Texas. September.

 

35. Raceme of little bluestem- This spicate raceme had advanced to the seed-ripe stage just before the next phenological stage of seed-shatter. The spikelets are paired: one perfect and sessile, the other sterile and pedicellate, with two florets per fertile spikelet. These two florets (in the fertile spikelet) are as follows: one floret is perfect and terminal and the other (the lower) floret is present only as a sterile lemma. The paired spikelets fall entire and together, the sterile pediceled spikelet attached to the fertile sessile spikelet, from the jointed rachis of the compound spicate raceme. The individual branches of this compound raceme are often defined as a rame, an inflorescence branch which bears some pediceled and some sessile spikelets. Two entire rames and the basal part of a third rame were displayed in this slide.

Tarleton State University Hunewell Ranch, Erath County, Texas. September.

 

36. Basal portion of a shoot of little bluestem- It was explained above that little bluestem is strictly a cespitose species (a bunchgrass) the tillers (upright, intravaginal shoots) of which form a tussock habit. In spite of the rank or relatively coarse and large size of little bluestem the individual tillers are rather easily broken off at the ground level (ie. at or just above the root crown) of little bluestem plants. This was shown in these two photographs. In other words, even though this species evolved under heavy grazing and is well-adapted to defoliation even it can be damaged-- remarkable easy-- by imporper grazing, especially overuse (overstocking). In time, prolongued overuse leads to overgrazing and a change in species composition of the range plant community. Increasers and invaders are much less easily damaged than the tallgrasse species including little bluestem.

On a portion of the Grand Prairie on Hunewell Ranch, Erath County, Texas. September.

 

37. Colony of big bluestem- Big bluestem is the dominant tallgrass species across much of the tallgrass (= bluestem) prairies that once extended from western Ohio and the Canadian Prairie Provinces to the central Dakotas and south through the Blackland Prairie and coastal prairie region of Texas, specially on soils having calcareous parent materials like limestone and dolomite. Big bluestem produces more sexually reproductive shoots under spring burning regimes as shown hereon the Konza Prairie (Riley County, Kansas, July).

 

38. Color namesake of big bluestem- This is an example of spring coloration for which bluestem (big bluestem in particular) was named. Technically the blue pigmentation is at tips of leaves not stems but blueleaf just does not have the poetic ring of names so characteristic of the frontier and rural folk in general. Big bluestem is generally the number one dominant tallgrass species (first overall among the Four Horsemen of the Prairies in the Tallgrass Prairie Region). Obviously dominance, abundance, and related ecological characteristics are range site-specific with different climax grass species being dominant on different habitats. Commonness, relative abundance, palatability, adaptation to fire, etc. of big bluestem are responsible for this species often being regarded as "king of the tallgrass prairies".

Ottawa County, Oklahoma. Estival aspect, June.

 

Where "bluestem" comes from- Five plants of big bluestem that had high degrees of "blue" coloration, especially on mid-growing season leaves. The pronounced reddish purple coloration of these big bluestem of leaves was due to presence of anthocyanins, red, purple, blue pigments in plant tissue.

These plants were usually highly pigmented--more so than in previous (and, later, future) years--perhaps due a unique combination of growing conditions (whatever that might have been). In other words, typically leaves of big bluestem are not quite this purplish or redish. The foliage in this particular spring provided an unusual opportunity to demonstrate why the common or vernacular name is "bluestem" (even if it is more in blue leaves than in blue stems).

Ottawa County, Oklahoma. Late June.

 

Bluestem leaf- Leaves of big bluestem showing a common reddish purple color phase due to anthocyanins, the red and urple pigments in plant leaves. These were basal leaves prior to elongation of shoots. They were on one of the plants presented in the immediately preceding five-slide/caption set.

Yees, of course, the leaves still had enough chlorophyll, the more abundant plant pigment, to conduct photosynthesis which in the bluestem tribe (Andropogoneae) of the panicoid grasses (Panicoideae) is the C4 photosynthetic pathway.

Ottawa County, Oklahoma. Late June.

 

Purplish at the bottom- Base of big bluestem plant in late spring with pigmentation of that species in this season. There was more purple coloration than is typical, but it was a nice example of the base of the King of the Tallgrass Prairie (at least here in the western Springfield Plateau at the southwestern edge of the Prairie Peninsula.

Ottawa County, Oklahoma. Early June.

 

39. Basis of the king-grass species- Characteristic basal shoots of big bluestem on tallgrass prairie in western Ozark Plateau. Pubescence and semi-flattened culms that are typical of big bluestem could be seen in detail in these three photographs. Hitchcock and Chase (1950, p. 757) described the pubescence on basal leaves of big bluestem as being occasionally densely villous (long, soft hairs).

Hitchcock and Chase (1950, ps. 749-751) divided the Andropogon genus into three sections based on numbers and other features of the spikelet-bearing branches which they termed racemes. Big bluestem was included in the Arthrolophis section which included such highly unpalatable and invader species as broomsedge bluestem (A. virginicus), splitbeard bluestem (A. ternarius), and bushy bluestem (A. glomeratus). Contemporary treatment in Barkworth et al. (2003, ps. 652-664) limited Andropogon to the Arthrolophis section of Hitchcock and Chase (1950, ps. 750-751) and divided this restricted Andropogon genus into two sections: 1) Andropogon which included only big bluestem and sand bluestem (A. hallii) and 2) Leptopogon with all the other Andropogon species of the traditional Arthrolophis section. Andropogon and Leptopogon sections were distinguished/segregated based on features of pedicellate spikelets (in effect, well-developed versus vestigial or wanting, in the respective sections).

Big bluestem is what has been called variously a "culmless", "short-shoot", or "short-culmed" grass. This king of the tallgrass prairie--overall the dominant, single most important, and most palatable of the Four Horsemen of the Prairies--is a tallgrass species that does not elevate its "growing point" (apical meristem) until relatively late in the warm growing season. Big bluestem blooms about as soon as the sexual shoots have elongated. This is in contrast to Indiangrass and switchgrass ("culmed", "long-shoot", or "long-culmed" grasses) which elongate their shoots early in the growing season and, therefore, much earlier than onset of flowering.

Ottawa County, Oklahoma. Late June; young vegetative state with apical meristems still low.

 

Shoots of the king species- Tillers (vertical grass shoots) of big bluestem growing in part of the Prairie Peninsula (in the Springfield Plateau of the Ozark Uplift). Tillers in the first slide were mostly sexual shoots that had progressed to the early flowering stage (onset of anthesis). They showed a pronounced purplish tinge due to anthocyanins, the red/purple pigments in plant tissue. (The influence of anthocynains in big bluestem was shown above.)

The second slide presented a principal tiller with two secondary tillers arising from this main, sexual shoot that had progressed to the early flowering stage.

Big bluestem is almost unquestionably the single most important species in the tallgrass prairie region of central North America. The first slide while he second slide showed

Ottawa County, Oklahoma. Late July; sexual shoots fully extended, early flowering stage of phenology.

 

King shoots at closer distance- Tillers of big bluestem with seccondary tillers (=shoot branches) arising from the principal tillers. A phytomer (the node-to-node unit of the grass shoot including internode and single leaf) was shown in the center-left foreground. That part of the purplish-colored internode not covered by the leaf sheath was due to fairly high levels of the pigment, anthocynain (see above slide/caption sets).

Ottawa County, Oklahoma. Late July; sexual shoots fully extended, early flowering stage of phenology.

 

Stems more green than blue, purple or red- Group of elongated tillers in big bluestem that had the more typical and characteristic green color of this king-of-the-tallgrass-prairie species. The shoots on this particular plant had lower levels of anthocynains (pigments resonsible for the purplish or reddish colors that were discussed above).

The second slide was somewhat of an insert view of tillers presented in the first slide so as to show details of the leaf axil of this marvelous species, which generally is the single most important species across much of the tallgrass prairie.

Although big bluestem is a tallgass species (reaches roughly the same heights as other tallgrasses like Indiangrass and switchgrass), it is categorized as a short shoot or culmless grass. The tillers of big bluestem elongate later in the growing season so that growing points of tillers remains lower (shorter in height) than its gracefully arching leaves. By contrast, Indiangrass and switchgrass tillers elongate (= shoots extend their growing points of meristematic tissue) earlier in the growing season so as to be categorized as long shoot or culmed grass.

At the stage of tiller development seen in these two photographs this short/long or culmless/culmed distinction cannot be seen.

Ottawa County, Oklahoma. Late June; tillers elongated, immdiate early flowering phenological stage.

 

Magnificant specimen of magnificant species- One immense plant of big bluestem at peak standing crop showing the vast number of tillers and the height of these. This plant, the tillers of which was senescing at end of their completed life cycles, was right at seven feet tall. It was the result of a reseeding project on a sandy bottomland that had been lightly grazed during the current growing season.

Tarleton State University Hunewell Ranch, Erath County, Texas. Early September.

 

40. Inflorescence of big bluestem with the characteristic three rames (inflorescence branches bearing some pedicellate and some sessile spikelets) from which arrangement is derived another common name of turkey track. See next set of slides for greater detail. Next slide, please.

First slide: Ottawa County, County Oklahoma, August. Second slide: Tarleton State University Hunewell Ranch, Erath County, Texas. Early September.

 

41. Rame of big bluestem- The anthers and stigmata were shown fully exerted on these spikelets of big bluestem at anthesis. Newton County, Missouri. September.

 

42. Autumn coloration of big bluestem- This colony of big bluestem at peak standing crop and seed-shatter stage illustrated the phenomenon of seasonal colors of the prairie. The grasslands-- most notably, tallgrass prairie-- of North America undergo color changes with progression of the seasons and phenological stages of the plants. This is the equivalent of seasonal color change in the deciduous forests that were often (sometimes still are) conterminous and intermixed with tallgrass-dominated grasslands and savannas. These are the seasonal aspects that have been noted periodically in this segment of the publication.

Autumnal aspect of tallgrass prairie was pronounced in this stand of big bluestem that was growing near the western extent of it's geographical range in central Texas. The ecotype shown here was scarcely four feet in height (four foot woven wire in immediate background) which was no taller than stands of little bluestem adjacent to this colony. (The far background was woody invasion of live oak, post oak, and mesquite on overgrazed tallgrass Grand Prairie range).

Mills County, Texas. October.

 

43. Sexually reproductive shoots of big bluestem- Grassses are regarded as reproducing by both sexual and asexual means. The latter is essentially multiplication of secondary shoots (tillers, rhizomes, or stolons) by vegetative growth (hence asexual reproduction is also known as vegetative reproduction). Perennial grasses typically reproduce more prolifically (and more effeciently/effectively) by vegetative reproduction than by sexual (grain production) means.

This is most pronounced in many of the prairie grasses native to North America. Flowering and fruit (caryopsis) production in these species is essentially a "luxury" action or physiological response. Going back to an early classic sutdy by Branson (1953) various authors have placed different grasses species into various groups based on certain characteristics. One of the more common of these groupings was a scheme based on location and timing of elongation of the apical meristem in combination with proportion of shoots that became sexually reproductive (Heady and Child, 1994. p.23). These authors placed big bluestem in the group described as having "infertile apices numerous and in or near the soil".

The relatively high proportion of big bluestem shoots that were developed infloresences and produced grains in this colony resulted from a summer that was considerably cooler and slightly wetter than was typical.

Mills County, Texas. October.

 

44. Inflorescences of big bluestem- Rames of big bluestem at the seed-ripe stage and in autumn coloration. Individual spikelets along the rachises were conspicuous in these unusually fertile and heavy yielding shoots. Mills County, Texas. October.

 

45. King and Queen of the Prairies- Big bluestem (right) and little bluestem (left) on a portion of Grand Prairie in the West Cross Timbers and Prairies vegetational area of northcentral Texas. This was at the western edge (the least subhumid or least mesic part) of the species range for big bluestem in this land resoruce area (ie. habitat was marginal for big bluestem). Consequently big bluestem, King of the Prairies, had already matured and was at the seed-ripe (and fast approaching seed-shatter) stage while little bluestem, Queen of the Prairies, was still in the soft dough phenological stage.At this geographic location and on this range site (Laomy Prairie) the range environment was "prime habitat" for little bluestem, regional dominant of tallgrass prairie in central Texas, and this species was slightly later (slower) in it's annual cycle and lagged behind big bluestem, the general dominant tallgrass species across or "averaged over" the entire bluestem-Indiangrass prairie (such Kuchler units as K-66 and rangeland cover types as SRM 710, both designated Bluestem Prairie).

Astute observers will have noted that the ecotype of big bluestem growing on habitat marginal for this species produced shoots no larger (taller) than those of little bluestem. This phenomenon was not the general or typical condition relative to respective sizes of these species on tallgrass prairie. The usual size dimorphism was embodied by common names of the two species.

Mills County, Texas.

 

45. King Grass in winter- Two plants of big bluestem with dead shoots from the preceding growing season. The plant in the second slide had a number of sexual shoots (right center of photograph) whereas the plant in the first slide had not produced any sexual shoots in the preceding summer of Extreme Drought (Palmer Index). Both plants grew within a few feet of each other on a mexic upland tallgrass prairie in the western edge of the Ozark (Springfield ) Pl.ateau that was dominated by big bluestem with tall dropseed (Sporobolus asper) as local associate species.

The dried, dead herbage of big bluestem in this condition is quite palatable to livestock, especially cattle. When taken in human hands in this state it is remarkably "soft" and readily broken up by slight twisting (presumedly this herbaceous material would be even more readily fragmented by ruminant teeth). Perhaps this explains the notable palatability of dead foliage of big bluestem in comparison to that of the other three of the Four Horsemen grasses (little bluestem, Indiangrass, switchgrass) not to mention broomsedge bluestem (Andropogon virginicus) which is in the same section of Andropogon as big bluestem (Hitchcock and Chase, 1950, p. 750).

Ottawa County, Oklahoma. Early January.

 

46. Indiangrass- The State Grass of Oklahoma is the major co-dominant with the bluestems, especially big bluestem, of the tallgrass prairie range type. Indiangrass is more abundant and the dominant grass on soils derived from sandstone parent material and is thus relatively more common south of the zone where big bluestem is the major dominant. Indiangrass is typically the tallgrass dominant over much of the Osage Questas and particularly the Chautauqua Hills portions of the Central Lowlands physiographic province south of the Flint Hills section. As shown here both species are herbaceous dominants in the forests and savannas of the Ozark Plateau section. Newton County, Missouri, October.

 

47. Colony of Indiangrass- These five to six feet-plus shoots of flowering Indiangrass at peak standing crop in a fencerow show the remarkable drought-tolerance of native tallgrass species. This biomass yield was at summer's end in one of the most severe droughts in Texas history. It grew during the fourth year of a protracted drought, the last two growing seasons of which were drier than any two of the great drought of the 1950s. Shoots in this colony remained green throughout the entire duration of each growing season in the prolonged drought of the 1990s. Grand Prairie vegetation. Erath County, Texas. October.

 

48. Inflorescence of Indiangrass- This panicle (in anthesis) illustrates the resemblance of the flowering shoot of a dominant prairie grass to the arrow atop the head of an Indian brave and thus origin of the name Indiangrass. (By the way, the author finds it obvious that the first letter in this common name should properly and always be capitalized given that Indian is a proper noun. Incidentally, Indian— and not that horribly offensive, sickeningly affected, and incorrect invention "Native American"— is the correct, precise name for American aborigines.)

 

49. Graminaceous headress- Panicle and flag leaf (first photograph) and spikelets on branches (second photograph) of yellow Indiangrass. The more specific common name of yellow Indiangrass is often aplied to Sorgastrum nutans in contrast to slender Indiangrass (S. elliottii) and lopsided Indiangrass (S. secundum). The latter occurs only sporatically from North Carolina to Louisiana and Arkansas except in Florida where it is widespread.

Branches with spikelets arising from the cental axis is the pattern or arrangement of main floral units is a panicle which strictly speaking is the only form of compound inflorescence in the Gramineae. The flag leaf is the leaf immediately subtending (below) the inflorescence in grasses.

Hunewell Ranch, Erath County, Texas. October; peak-bloom stage (and almost simultaneous anthesis in all spikelets).

 

50. Beauty in flowers of Oklahoma State Grass- Close-up "shots" of Indiangrass spikelets in anthesis.As chance or Divine Intervention would have it, this and the preceeding pair of photographs showed almost all spikelets in anthesis at once. This flowering sequence is unusual in most grass species because grasses are determinate bloomers (flowering proceeds from upper and outer spikelets downward and inward).

Indiangrass is one of the most prolific grain-producers of the native tallgrasses and, as such, is one of the better native species for reseeding of tallgrass, true, and mixed prairie ranges (Phillips Petroleum Company, 1963, p. 62; Leithead, 1971, p. 157; Tyrl et al., 2008, p. 163). In fact, Indiangrass, big bluestem, and switchgrass have, in recent years, been "rediscovered" as warm-season species for use in tame (agronomic) pastures where extensive management is possible and desirable.

Hunewell Ranch, Erath County, Texas. October; peak-bloom stage (and almost simultaneous anthesis in all spikelets).

 

Asexual and sexual options- These two slides presented the panicle inflorescence and shoot bases, including rhizomes, of common or yellow Indiangrass. This demonstrated sexual reproduction (panicle inflorescence) and asexual reproduction (several tillers and rhizomes) in this major dominant grass of the climax tallgrass prairies of North America.

Most perennial grasses, especially native or non-agronomic species, reproduce primarily asexually, that is by growth and development of new shoots (tillers, stolons, or rhizomes; rarely bulblets). Sexual reproduction (creation of new genotypes by recombination of genes) is a "viable" (as in viable or live seed) option, but typically not a "day-to-day" or "meat and taters" mode. Grain production that leads to generation and establishment of genetically distinct plants (new genotypes; novel genetic individuals) is the primary means of evolution or natural selection (or manmade artificial selection for that matter) that permits adaptation of species to environmental changes. Sexual reproduction via grain production in perennial grasses can also be extremely effective, even essential, in recovery of species following extreme disturbances (drought, catastrophic fire, overgrazing, tillage, etc.).

The combination of sexual and asexual reproduction has enabled Indiangrass to become and remain a dominant species of the climax tallgrass prairie, one of the Four Horsemen Grasses of the Prairies, and associated woodlands and savannas. Indiangrass is a mainstay of the tallgrass prairie range being extremely palatable to grass-prefering herbivores and is well-adapted to drough. It thrives under spring fires with rhizomes and rootcrowns being protected from flames. Indiangrass is ideal for stocker cattle range and has value as a native ornamental grass. Although it was probably due mostly to the common name, Oklahoma nonetheless got it right to select Indiangrass as its State Grass.

The color of these two slides was not reproduced accurately/correctly by an Epson Perfection 600 scanner. Somehow the scanner picked up on the yellow color of the panicle of yellow Indiangrass and incorrectly used that yellow to give a reddish or yellowish cast for soil color. Some other slides on the same roll of Provia 100F film taken within a few minutes of these slides but without the panicle and scanned in the same scan group (one after the other) had the correct color produced by the Epson scanner. One of these slides with only Indiangrass rhizomes (without the yellow-colored panicle) was presented in the next slide-caption set.

Erath County, Texas. October; hard- or ripe-grain stage of phenology.

 

Rhizomes (and right color)- Rhizomes of yellow Indiangrass. Rhizomes are horizonal, subterranean shoots. Horizontal shoots, both stolons and rhizomes, are extravaginated or extravaginal shoots with the shoot apex piercing the sheath rather than comming up through this invaginating tissue in which case the shoot would be intravaginal or intravaginated and vertical (a tiller).

Indiangrass (all the Sorgastrum species) are rhizomatous, meaning having rhizomes. Thus, Indiangrass is not a strictly cespitose (tufted or bunched habit) or a bunchgrass species but instead can be a semi-sodforming grass. It is capable of more lateral spread than if it reproduced asexually only by means of tillerss. Indiangrass obviously has tillers as the major type of shoot and characteristically has a more-or-less bunchgrass habit though not exclusively so.

With this morphology, Indiangrass not only has sexual and asexual reproduction, but within its asexual (= vegetative) mode it has the option of tufted expansion by tillers or sodforming increase by rhizomes. This is a major reason by Indiangrass commonly does so well in range reseeding projects or restoration of tallgrass prairie.

Erath County, Texas. October.

Details of two of the Four Horsemen grasses- Close-up photographs of big bluestem and Indiangrass were presented immediately below. Both of these tallgrass species are in tribe Andropogoneae and have the tribal characters of: 1) paired spikelets on a jointed rachis with one spikelet perfect and sessile and the other spikelet being sterile and pedicellate, 2) the fertile spikelet, in turn, has one perfect floret that is terminal and a sterile (empty) lemma below the perfect floret, 3) the fertile spikelet (with its paired florets; one perfect, one sterile) is shed along with the rachis joints, and while 4) the sterile and pedicellate spikelet remains attached (Chase, 1964, p. 99). In other words, there are paired spikelets with the fertile one of these spikelets having paired florets with the uppermost one of these paired florets bing fertile (perfect). Both florets (one fertile, one sterile) of the perfect and sessile spikelet are shed as a unit (the spikelet) while the sterile spikelet remains on its pedicel which stays attached to the inflorescence.

Examples were growing side-by-side on an oak-hickory-tallgrass savanna on the western edge of the Ozark (Springfield) Plateau. Details of these two members of the Andropogoneae were presented together for consistency in describing fetures of this panicoid tribe.

 

51. Next genetic generation of big bluestem- Details of sexual propagules of big bluestem at scale of spikelets. Close-up vies of rames of big bluestem showing individual spikelets of this champion species of the tallgrass prairie and tallgrass-eastern hardwood savanna. Rame is the traditional term applied to branches of the panicle of Andropogoneae with there being both pedicellate and sessile spikelets on such branches. (Pohl, 1968, p. 242). (Similarly,the adjectives ramose, meaning branching or having many branches, and rameal, in reference to oroccurring on a branch, are used in standard taxonomic talk.)

Paired spikelets--both perfect and sessile and sterile and pedicellate--were visible upon close examination of rames in both of these photographs. The appendages that appear as (look like) little branches with missing units are the sterile, pedicellate spikelets and not an empty space where a spikelet or floret was attached. However, to add to confusion (as if it was needed), some pedicels were without sterile lemmas (ie. some pedicellate spikelets were present only as pedicels). There were also some whole pedicellate spikelets that did have their one sterile lemma.

Generally, intact fertile (perfect) and sessile spikelets were still attached. Some of all floral units were present so that the complete arrangement of paired spikelets and paired florets of perfect spikelets were still attached. Phenological stage was grain-ripe not grain-shatter.

Stoney Point Savanna, Ottawa County, Oklahoma. January; grain-ripe stage.

 

52. Next genetic generation of Indiangrass- Details of sexual propagules of Indiangrass at scale of spikelets. Sections of panicle branches with mature, grain-filled, fertile spikelets. Paired spikelets--both perfect and sessile and sterile and pedicellate--were visible upon close observation of these two photographs. Sterile, pedicellate spikelets were visible as pedicels with pilose pubescence. Fertile, sessile spikelets were much more obvious in these taken-on-the-range shots.

Stoney Point Savanna, Ottawa County, Oklahoma. January; grain-ripe stage.

 

53. Silver bluestem (Andropogon saccharoides= Bothriochloa saccharoides= B. saccharoides var. torreyana= B. laguroides subsp. torreyana) at anthesis - This midgrass is typically an increaser on most mixed prairie range sites and an invader on tallgrass prairie range sites, but it is a decreaser on some of the range sites of the arid semidesert grassland or shallower sites in the western Edwards Plateau adjoining the Chihuhuan Desert. Nomenclature of this species seems to be a never-ending source of debate, revision, followed by debate and further revision (ie. another name), none of which adds anything useful or practical to management of this or associated range plants. Erath County, Texas. July.

 

54. Silver bluestem at seed ripe stage- Characteristic autumn coloration of this prairie midgrass. Note that most of the shoots are sexually reproductive. Erath County, Texas. July.

 

55. Partly in and out of the boot- The four apices of silver bluestem shoots in these two photographs presented four degrees of emergence of the inflorescence (panicle) from the boot. Panicle atop the left shoot in second photograph was fully emerged or exerted. Details of fully expressed panicles were shown in the next set of two slides. West Cross Timbers near end of a summer of severe drought.

Erath County, Texas. September, emergence of inflorescence from boot.

 

56. Silvery spectacle- No, not necessarily spectacular but a conspicuous spectacle nonetheless was this appearance of showy inflorescences of silver bluestem in the Western Cross Timbers. Terminology applied to inflorescences of Andropogon, Bothriochloa, Schizachyrium, and Dichanthium species has about as many interpretations as does taxonomy of these taxa. Contemorary description of this inflorescence type is panicle with primary racemose branches (Gould 1975, p. 591; Hignight et al., 1988, ps. 8, 23; Hatch and Pluhar, 1993, p.43). These specimens were growing in the West Cross Timbers near end of a long, hot sumer of severe drought and following two mowings. This species is survivor if given half a chance (come to think of, even if not given half a chance).

Erath County, Texas.September, fully emerged inflorescence, at 1) pre-anthesis stage (first photograph) and 2) early dough stage (second photograph).

 

57. Broomsedge bluestem (Andropogon virginicus)- Although broomsedge is closely related to big bluetem (Hitchcock and Chase [1951] placed them in the same section of Andropogon) the former has nothing of the ecological status or forage value of the latter. In fact, broomsedge is an invader throughout the tallgrass prairie (if not everywhere it grows) and it is one of the least palatable of all native perennial grasses throughout its biological range. Individual broomsedge plants form large conspicuous bunches and it is a strictly cespitose species whose shoots are all intravaginated (ie. tillers), but it reproduces very effectively both vegetatively and through abundant seed production. Ottawa County, Oklahoma. Seed-shatter stage; autumnal aspect, early December. Ottawa County, Oklahoma.

 

58. Tillers of broomsedge bluestem at seed-shatter stage- Spikelets in the specialized inflorescence (a spicate raceme) of broomsedge. Ottawa County, Oklahoma. Early December.

 

59. Colony of broomsedge bluestem- Broomsedge appears to be a classic example of a ruderal species, a plant associated with man-made disturbances such as waste places or minimally managed land (ruderals are usually viewed as weeds). More specifically, broomsedge was interpreted as a dominant competitive-ruderal that exerts phytotoxic effects (Grimes, 1979, p. 144). Students will get an instructive lesson by studying Grimes' "primary strategies", in this instance of the "low stress-high disturbance strategy" (Grimes, 1979, ps. 7, 39-45).

This population of broomsedge was growing in a 40-year old planting of "Kentucky 31" tall fescue. Fescue is a cool-season bunchgrass so it and the cespitose, warm-season broomsedge co-exist (often as co-dominants) on extensively managed permanent pastures of the introduced tall fescue and on old-fields where tall fescue became established as a volunteer crop. This is the Ozark Fescue-Broomsedge Cover Type category of Tall Fescue (SRM 804), an example of which was shown here. In portions of winter through early summer tall fescue will appear as the dominant until it goes into summer dormancy at which time the broomsedge bunches appear as dominants-- at least as an aspect dominants as is also the situation for tall fescue in it's growing season.

Broomsedge often forms immense consociations like the one shown here on abandoned farmland and cut-over forests from the tallgrass prairie and oak-hickory uplands of the Ozarks to the Piedmont along the Atlanic Ocean. Broomsedge frequently exists as a midseral stage for prolonged periods on old-fields ("go-back-land") that were depleted by soil erosion and overcropping.

Broomsedge is one of the most unpalatable species of native grasses in the tallgrass prairie and oak-hickory-tallgrass savanna. The resemblance (under superficial and careless exmination) of broomsedge to big and little bluestem and it's presence as an invading associate on deteriorated tallgrass prairie was a combination duped many an unsuspecting stockman. At one time many of the large steer operators leased-- and occasionally even bought-- "go-back land" dominated by broomsedge thinking it high-quality bluestem pasture. Jawhawker and Okie landowners had a nice laugh (all the way to the bank) at the expense of steermen (many from Texas). It did not take two such experiences before cattle-rasisers learned that plant identification was not just a hobby for botanists at the local "cow college".

Ottawa County, Oklahoma. Autumnal aspect, early December.

 

60. Basal part of autmunal-hibernal broomsedge shoots- One key part of the specific dominant competitive-ruderal "strategy" (Grimes, 1979, ps. 39-51) of broomsedge bluestem is maintnenace and/or production of green shoots at ground level during what is mostly (= "more-or-less") it's dormant season. Panicoid grasses like the Andropogon species are quite obviously warm-season plants that typically flower and set grain in late summer to fall. The photographs presented here illustrated that broomsedge bluestem reaches seed-ripe and seed-shatter stages in autumn on into early winter. Closer insepection showed that broomsedge maintained (or grew new) live shoots throughout much of the supposed dormant phase of the annual cycle of this perennial grass. This pattern differs from other bluestems such as big and little bluestems and panicgrasses like switchgrass whose shoots die back at or soon after the first heavy fall frosts. In fact, shoots of these decreaser panicoid grasses often enter dormancy before frost.

Persistence of live shoots of warm-season species into and throughout much of winter theoretically allows photosynthesis to be extended over a longer period. Location ot this green tissue at the base of the plant where it is largely covered by dead herbage certainly offers it some protection from cold temperatures and freeze damage by infrequent snows, but such coverage also limits photosynthesis. Perhaps the main survival or competitive advantage provided by this growth and pattern of resource allocation is to give this ruderal and ecological invader a "headstart" over the native dominant bluestems, Indiangrass, and panicgrasses (decreasers).

The specific function(s) of hibernally green tissue in A. virginicus would be a good research project, and one with practical applications that could help reduce cover and density of broomsedge in both introduced permanent pastures like tall fescue and native tallgrass prairies.

Ottawa County, Oklahoma. Early December.

 

61 Splitbeard bulestem (A. ternarius)- This is another native Andropogon species (in the same section as big bluestem and broomsedge) that is an ecological invader. Splitbeard bluestem does not form immense colonies and occur as a dominant species over such an extensive geographic area as broomsedge, but it is an indicator species on depleted range and reflects past or present mismanagement like overgrazing.

This specimen was growing in a 40-year old stand of "Kentucky 31" tall fescue. Ottawa County, Oklahoma. Seed-shatter stage, early December.

 

62. Tiller of splitbeard bluestem at seed-shatter stage- The annual phenological cycle of splitbeard bluestem coincides with that of broomsedge bluestem. Ottawa County, Oklahoma. Early December.

 

63. Rames of splitbeard bluestem- Spikelets on two branches of the spicate raceme of A. ternarius. Ottawa County, Oklahoma. Early December.

 

64. Grampaw grass of the tallgrass prairie-Local stand of eastern gamagrass or, rarely, corngrass (Tripsacum dactyloides) on a bottomland range site in the western Ozark (Springfield) Plateau. Physiogonomy and structureof a consociation of this monecious member of tribe Maydeae (or of the enlarged bluestem tribe, Andropogoneae). With exception of some of the plumgrasses (Erianthus species), eastern gamagrass is the grass species that has the largest individual plants in the interior tallgrass prairie. In fact, eastern gamagrass often produces cespitose plants of larger basal diameter than the much more rare Erianthus species.

With its immense plant size,. large (both long and very wide) leaves, and extremely high palatability eastern gamagrass has received much study, especially in regards to reseeding range as well as use as a domisticated native (see herein end of chapter, Introduced Forages- Grasses).

This example of a consociation of eastern gamagrass had developed on an old field that almost a hallf century before had been farmed. In his youth this author hauled hay of cowpea (Vigna unguiculata) and foxtail millet (Setaria italica) off of this field. Within three decades after abandonment from tillage eastern gamagrass appeared on this go-back ground "from out of nowhere". Individual plants of eastern gamagrass have continued to increase in basal area (asexual reproduction by tillers) and more (younger and smaller) plants have become established, presumedly from caryopses of established parent plants. Where did the propagules from which these parent or first re-established plants originated come from? Had grains or even rhizomes of eastern gamagrass survived under years of tillage? Was grain carried in by birds or from flood water? Saint Paul would call it a mystery and be done with it. This author could add nothing of scientific substance and for the time being simply designate the re-establishment phenomenon a "successional miracle". The ghosts of Frederick Clements and John Weaver were heard rustling the gamagrass shoots.

It is notable that apparently there are not detailed overall treatments of eastern gamagrass as a range plant or native species. Aside from the standard Phillips Petroleum Company (1963, p. 71) Pasture and Range Plants and the newer and highly recommended Field Guide to Oklahoma Plants (Tyrl et al., 2008, ps. 182-183) discussion of eastern gamagrass from a forage plant perspective included Roberts and Kallenbach (1999), Henson (2012), and Shadow (2012). Most research papers devoted to eastern gamagrass have dealt with its taxonomy such as the paper by deWet et al. (1982) and, especially, in relation to the Zea species as for instance deWet and Harlan (1974), Harlan and deWet (1977), Doebley (1982), and Goodman et al. (1983) with much less study having been made of other aspects like plant growth (Anderson, 1987).

Ottawa County, Oklahoma. Early June; peak anthesis.

 

65. Overall view of a specialized inflorescence- Examples of the flower cluster of easstern gamagrass. The inflorescence of eastern gamagrass was described by Barkworth et al. (2003, p. 693) as "panicles of one to several subdigitate to racemose rames, rames with pistillate spikelets proximally and staminate spikelets distally" and by Shaw (2012, p. 979) as a "spikelike raceme or series of two to a few spikelike racemose branches bearing staminate skikelets above and pistillate spikelets below". Further details of this unique inflorescence (eg. two-flowered, paired staminate spikelets, single pistillate spikelets partly embedded in the cob or rachis) are available in these references. The format for such descripions of the Gramineae inflorescence were first provided by Highnight et al. (1988).

Ottawa County, Oklahoma. Early June; peak anthesis.

 

66. Inflorescence of eastern gamagrass at full anthesis with separate staminate and pistillate spikelets- This resemblance to the tassle of corn or maize (Zea mays) was both obvious--both of these species are in tribe Maydeae or Tripsaceae--and the tasslelike monecious inflorescence is origin of the other common name of corngrass. Work by Li et al. (1997) and Orr et al. (2001) regarding hereditary mechanisms and organ development in Tripsacum and Zea genera revealed a common genetic basis to sex dtermination strongly suggesting that flowers and fruit of corn or maize was derived from a Tripsacum genomic basis (Orr et al., 2001) and that abortion of pistils was a genetic process occurring in species of both genera. (Li et al, 1997). These state-of-the-art scientific studies verified a basis of the common name of corngrass applied to T. dactyloides by Ozark hillbillies was a neat example of the validity and vision of folk wisdom.

Ottawa County, Oklahoma, June, peak anthesis.

 

67. 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.

 

68. Staminate florets of eastern gamagrass- Tridens Prairie, Lamar County Texas, June.

 

69. Pedicellate florets of eastern gamagrass- Tridens Prairie, Lamar County Texas, June.

 

70. Switchgrass- Specimen of a bottomland ecotype of Panicum virgatum along the banks of the Bosque River.in northcentral Texas Switchgrass occurs as both a bottomland form, as featured herein, as well as an upland form (upland ecotypes). This is the most mesic of the four dominant tallgrass species (Four Horsemen of the Prairies). Switchgrass characteristically grows in lower habitats than the other three dominants, but upland ecotypes hold their own on upland slopes.

As of this writing switchgrass is the only one of the four magnificant dominants of the tallgrass prairie that has not been designated a State Grass. Some state proud of its prairie heritage needs to pick it up.

Erath County, Texas. September, peak standing crop and immediate pre-bloom stage.

 

71. Switchgrass- An old field reseeded to Cave In Spring accession of switchgrass released out of Missouri by the Soil Conservation Service. Native warm season, perennial tallgrasses like switchgrass, big bluestem, and Indiangrass have been “discovered” by forage agronomists to be dependable, drought-tolerant, and palatable pasture and hay species which require judicious grazing management but not the expensive intensive inputs of fertilizer, irrigation water, etc. Of course much of this “natural advantage” would be lost if these “wild” species were to be domesticated. Newton County, Missouri.

 

72. Panicles confront the blue- Panicles of a big specimen of bottomland switchgrass at seed-shatter stage stood in beautiful exposure against the cobalt-blue sky of a Texas "blue norther". A "norther" in traditional lingo refers to massive cold fronts that originate as Arctic air masses and move southward often following the Rocky Mountain chain and dispersing over the Great Plains eastward across the prairies of the Central Lowlands physiographic province. Initially the sky of a 'norther" is gray, often a dark charcoal gray or bluish-black, as it meets the warmer, moister air moving northward out of the Gulf of Mexico. After a day or so of battling air masses the colder, drier, northern air prevails over its southern cousin and under dominion of this clear Arctic air the sky turns to azure blue.

The "yucky" air so typical of Texas skies in the contemporary atmosphere filled with pollution from the likes of a "zillion" automobile exhausts is "scrubbed" clean as crisp cold air moves out the haze. Such are Kodachrome moments, and when switchgrass stands tall against the air of a "Texas norther" the Lone Star sutterbug captures the meeting of great grass and grand sky.

Tillers of this particular switchgrass plant exceeded eight feet in height and were adorned with panicles comprising two feet of that length. Many, perhaps most, caryopses from these compound inflorescences had already shed. Perhaps new plants (genotypes) will join this mighty sexual specimen.

Erath County, Texas. Late November, peak standing crop at grain-shatter, late maturity stage of phenology.

 

 

73. Big clump- Basal shoots (tillers) of a large specimen of the bottomland form of switchgrass. Switchgrass has relatively long scaly rhizomes, but it forms large bunches or tufts comprised of many robust tillers. Plants of bottomland switchgrass are much larger in contrast to the upland switchgrass form so that the cespitose habit (bunchgrass morphology) of this species is more pronounced, especially under ideal growing conditions. This plant was approaching pre-dormancy stage (note many senescing leaves) in late autumn following an unusually wet period from late summer through end of warm growing season. It showed the immense size of which this species is capable of attaining.

Erath County, Texas. Late November, peak standing crop at grain-shatter, late maturity stage of phenology.

 

74. Big switch- Large panicle on the plant of bottomland switchgrass featured immediately above. It is not uncommon for this expanded panicle to reach lengths exceeding two feet and a half feet and to be nearly a foot across at the base. There are many spikelets along the numerous branches of this inflorescence. Like most of the tallgrass species switchgrass reproduces primarily by asexual means, but it certainly keeps its reproductive options open. his fine crop of florets was produced in an extremely wet late summer and an equally impreseive crop of tiny grain resulted from this profuse blooming.

Erath County, Texas. Late September, full-flower stage.

 

75. Little flowers- Spikelets of the bottomland form of switchgrass. Two views of numerous spikelets in full flower. These floral units were taken off of the large specimen described above. Both anthers and stigmas were exerted, but stigmas were more obvious.

Erath County, Texas. Late September, full-flower stage.

 

76. Stand of beaked panicgrass- Panicum anceps is often a locally dominant species in the tallgrass prairie and, especially, the tallgrass savanna and Cross Timbers. It is a palatable and productive species and often classed as a decreaser depending on range site. This colony grew on a small natural opening (a glade) in the western Ozark Highlands. Newton County, Missouri. September.

 

77. Up closer- Part of a local stand of beaked panicgrass on mesic to wset tallgrass prairie showing overall features of shoots at boot stage. Western Ozark Plateau (Springfield Plateau section).

Ottawa County, Oklahoma. August; boot (immediate panicle-emergence) stage of phenology.

 

Champion specimen- One very large plant (almost as much area as that of a conventional pick-up truck) of beaked panicgrass growing on an old field in the Springfield Plateau. The soil in which this specimen was growing had been that of a tallgrass prairie before it was "broke out" and farmed for over three-fourths of a century (both row crops and small grains). Then the field was abandoned 25 years ago and secondary plant succession had taken the recovering grassland vegetation back to climax or almost climax (near-climax) stage.

Surrounding range plant species were switchgrass (left), Indiagrass (upper right corner), and a rosette panicgrass (center background) known variously as hairy rosettegrass, hairy panicgrass, or wooly rosettegrass, (Panicum lanuginosum var. fasciculatum= Dichanthelium acuminatum subsp. fasciculatum).

Ottawa County, Oklahoma. Early August; approximate mid-maturity grain stage of phenology.

 

Peeking at the beaked- Four progressively closer views of beaked panicgrass growing on a wet prairie in the western Ozark (Springfield) Plateau with switchgrass, two-flower (two-flowered) rush (Juncus biflorus), slim-pod or diffuse rush (J. diffusissimus), both Florida and field paspalums, and fox caric-sedge (Carex vulpinoidea) as neighboring species.

Ottawa County, Oklahoma. Early August, peak stnding crop, early (immature) grain phenological stage.

 

78. Sexual shoots with inflorescences of beaked panicgrass- The flowering shoots of P. anceps on a tallgrass prairie glade in the Ozark Plateau (first slide) and on a wet prairie (second slide). Newton County, Missouri. September; Ottawa County, Oklahoma, August (first and second photograph, respectively) .

 

79. Beaked spikelets- Arrangement and details of spikelets on beaked panicgrass growing on a prairie in the western Ozark Plateau. Ottawa County, Oklahoma. July: anthesis and pre-anthesis stages.

 

Not beaked down here- Associated species of graminoids (the prevalent categories of range prairie plants on this habitat) were listed in the immediately preceeding caption.

 

80. Vine mesquite- Dense stand of vine mesquite showing characteristic sward of this valuable mid-grass. This panicgrass is adapted to a diversity of habitats ranging from understory species in the Texas Cross Timbers through to the rock-strewn shallow ranges of the Edwards Plateau and the floodplain and bosques of semidesert grasslands of the Southwest. Vine mesquite is usually a decreaser and fairly productive of palatable and reasonably nutritious forage when green but when mature it cures with lower nutritive value than associated shortgrasses like buffalograss or blue and black grama.

Vine mesquite produces both stolons and rhizomes such that is a pronounced sod-forming species. This sward was on a lowland site of Windthorst soil, Hunewell Ranch, Tarleton State University, Erath County, Texas, June.

 

81. Less viney than some- A single plant of vine mesquite growing in the West Cross Timbers of northcentral Texas during one of the most one-year droughts on record (at least 14 weeks of Exceptional Drought, the highest or worst category on the Palmer Drought Severuity Index). This plant was growing on a range that had been (was being) grazed by white-tailed deer as the only large mammalian grazer. Deer had not fed on vine mesquite. This was an example of the state of a native, decreaser grass when undefoliated by animals in the worst of drought.

This example demonstrted that if ranchers could destock and move livestock to ranges less impacted by drought the native range plants could survive quite handily. This one appeared to be thriving even under Exceptional Drought. Some readers might object to this hypothetical situation alledging that such an emergency measure would be impractical because everyone else was out of feed and there would be no range to rent at any price. OK, so what to any such objectors propose? Eat what grass is left into the ground and then have to destock anyway when that is gone? Talk about impractical! All that all-too-common mismanagement amounts to is absence of feed even when rains return (as they always do). That foolishness just leaves ranchmen without forage and livestock. Stockgrowers are, at the most fundamental level of animal production, grass growers. Grass (generic, poetic for forage) not animals is the basis of ranching. Animals (livestock or wildlife) are the commodity, but grass is the crop, the primary product. Animals are the secondary product, and that commidified secondary product can be only as good as the fundamental crop.

Returning to the crop species featured here, it was remarked that vine mesquite is often a major forage species on various range sites from tallgrass prairie and savanna through mesic habitats on semidesert grassland in both the Chihuhuan and Sonoran Desert Regions. Vine mesquite extends from the prairies of Illinois westward to Utah and south into Mexico. Vine mesquite is widespread--on mesic environments--thoroughout much of the vast Basin and Range Physiographic Province beginning with the Trans Pecos poration.

This particular vine mesquite plant had not produced long stolons, but it had produced a high yield of caryopses that were in the early hard-dough stage. Again, this grain crop was in the worst one-year drought in Texas history (as of this writing).

Tarleton State University, College Farm, Erath County, Texas. Late September.

 

82. Look what I did in record drought- Panicles filled with ripening grain of vine mesquite growing in the West Cross Timbers of northcentral Texas in the worst one-year drought on record (more than 14 weeks of Exceptional Drought, D4 on the Palmer Drought Severity Index). In the Paniceae (panicgrass tribe of the Panicoideae) spikelets are solitary (unpaired) and consist of two florets, one perfect or fertile and the other, which subtends (is below) the perfect, is sterile. The sterile floret is reduced to an empty lemma which resembles a glume. The lemma and palea of the fertile (perfect) floret are indurate ("hard").

In vine mesquite the entire inflorescence is a racemose panicle. Gould (1975, p. 452) described this flower cluster as a "narrow, contracted panicle or raceme". The inflorescence was regarded as a panicle with "... main branches appressed and sparingly rebranched" by Great Plains Flora Association (1986, p. 1203). These features were visible in these two photographs.

All-in-all, a very pretty--and an extremely drought-tolerant--species.

Tarleton State University, College Farm, Erath County, Texas. Late September; early hard-dough stage.

 

83. Spikelets of vine mesquite- Hunewell Ranch, Erath County, Texas, June.

 

84. Paniced again- Redtop panicgrass (Panicum agrostoides= P. rigidulum ) growing in a local depression on virgin tallgrass prairie. Redtop panicgrass is one of the lesser-known Panicum species of tallgrass paririe and adjoining savannas (eg. relative to switchgrass, beaked panicgrass, vine mesquite). Even the common witchgrass (P. capillare), an annual species, is better-known (even to most seasoned rangemen) than redtop panicgrass. This is undoubtedly a reflection of the rather restricted habitats (local, low-lying, wet environments) on which this species can survive.

On microsites where redtop panicgrass grows this bunchgrass forms small-size consociations with such nearly complete foliar cover that the stand produces a sod-like sward as seen in these two slides. Redtop panic can grow to the approximate heights of upland switchgrass growing on similar habitats. These two species as well as beaked panicgrass all were well-represented on this meadow used for production of prairie hay. This meadow had been improperly hayed (mowed in late summer) for years and had been returned to proper management so as to be in recovery (secondary succession), but management was irrelevant to this local stand of redtop panicgrass that was growing on a swale which ponded water and was to wet to hay. This land had been mowed for hay two weeks prior to time of these photographs. Wet ground permitted the photographing of this by-passed parcel.

Ottawa County, Oklahoma. Mid-July, peak standing crop and full-bloom stages.

 

85 Ya mean this is a bunchgrass?- Redtop panicgrass is a cespitose species (all shoots are tillers), but it commonly grows as such dense colonies in its restricted mesic microhabitats that it appears--at first flush anyway--to be a sod-forming (= stoloniferous and/or rhizomatous) species. The second of these two photographs showed the bunchgrass habit or growth form of this panicoid grass.

Ottawa County, Oklahoma. Mid-July, peak standing crop and full-bloom stages.

 

86. Growing out of the boot- Emergence of immature panicle of redtop panicgrass from the flag leaf, the leaf subtending the infloresecence. Grass inflorescences are pushed out through the sheath of the enveloping last leaf of the shoot as the culm grows (internodes elongate) grow up through it.

Ottawa County, Oklahoma. Mid-July, early bloom (emerging panicle) stage of phenology.

 

87. Top of redtop panicgrass- Entire panicle (first slide) and upper one-third of panicle (second slide) of Panicum agrostoides= P. rigidulum. The branches (secondary shoots) of the panicle of redtop panicgrass occur in whorls around the rachis (sometimes off of just one "side" of the central axis).

Ottawa County, Oklahoma. Mid-July, full-bloom (anthesis) stage of phenology.

 

88. Panicle and spikelets of redtop panicgrass- Upper portion of a mature panicle with nearly ripe grain (first photograph) and of an immature panicle and branches from another maturing panicle at pre-anthesis stage (second photograph). Basis of the common name adjective, "redtop", was apparent in the second slide.

By the way, this panicle and these branches were on a rock of chert "growing" on the hay meadow. Chert is a sedimentary rock made up of quartz of the fine granular type in the microcrystalline group (Pough, 1960, p. 231). Chert is the most common of the fine-grained siliceous rocks which when freshly broken has a shiny appearance 470). This stone had been broken and weathered for a number of years in the humid zone of the Ozark Plateau. It resided for decades in a fence row where someone had tossed it to protect sickle-bar mower knives and sections from its hard surface and "just the right size" for causing mower trouble.Chert and related li

mestones are the main parent material for soils of tallgrass prairie in this region.

Ottawa County, Oklahoma. Mid-July.

 

Upper and lower parts- General views of 1) mostly upper portions of shoots (first slide) and 2) lower to middle portions of redstem panicgrass growing on a wet prairie in the extreme western part of the Springfield Plateau of Ozark Plateau physiographhic province.

Ottawa County, Oklahoma. Late July.

 

 

All in a shoot- Upper portion of a shoot of redtop panicgrass complete with a panicle newly emerged from the boot. The boot is the term used by grassmen for that part of the shoot encclosing the infloresecence. The flag leaf is the leaf subtending (imediately below) the inflorescence (flower cluster).

This specimen grew on a wet prairie in the far-westen Springfield Plateau of the Ozark Plateau physiographic province.

Ottawa County, Oklahoma. Late July, immediately pre-bloom stage. i

 

Re-redtopped- Another example of redtop panicgrass. This specimen, along with several others, was growing on a wet prairie in the far-western Ozark Plateau. It was growing in association with Florida paspalum (Paspalum florida); several caric sedges including Carex vulpinoideae, C. bicknellii, C. frankii; , and rushes especially Juncus interior, J. biflorus, and J. diffusissimus. These associated species were presented and discussed below.

Ottawa County, Oklahoma. Late July; boot to milk-grain stages of phenology.

 

Not red yet- Panicles of redtop panicgrass at developmental stages of emergence from boot (first slide) and fully expanded with immature grain (milk stage) growing on a wet prairie in the far-western Ozark (Springfield) Plateau.

These two flower clusters were growing on the large plant presented in the immediately preceding slide. These two panicles at two different stages of development showed that phenological stage of panicle/flower development varied from shoot-to-shoot (tiller-to-tiller) on the same plant.

Ottawa County, Oklahoma. Late July; emergence from boot to milkly grain stages of development.

 

Axillary area- Area of shot of redtop pancigrass at site or location of grass culm and leaf with leaf at union of blade and sheath. Shown in first slide at abaxial (facing away from stem, underside of leaf blade) view and in second slide at adaxial (facing toward stem, upper surface of leaf blade) view.

This specimen was growing on wet pariarie in the western Springfield Plateau.

Ottawa County, Oklahoma. Late July; peak foliage with panicles at boot to milk-grain stage.

Rosette panicgrasses- Panicum is the largest genus of the grass family, at least as clasically traditionally treated. In a comprehensive review of the Gramineae, Clayton and Renvoize (1986) generalized that Panicum species have relatively uniform features in spikelet morphology, but otherwise have considerable variability in general morphology (eg. of culms ), anatomy, and physiologiy.Perhaps most fundamental in this context is that as conventionally interpreted Panicum includes both C3 and C4 species; that is, there are. some Panicum species with the Calvin-Benson C3 photosynthesis pathway whereas other Panicum species have the Hatch-Slack C4 pathway of photosynthesis. These species are usually regarded as cool-season and warm-season species, respectively.

Hitchcock and Chase (1951, p. 626-636), admittedly consummate "lumpers" at the genus level, sectioned the classic Panicum genus (based on spikelet morphology) into three subgenera based on shoot characteristics. Each of these subgenera was subdivided into untitled sections or subunits. Dichanthelium was one subgenus of Hitchcock and Chase (1951, ps. 627-633). Dichanthelium was divided into 17 sections. In most contemporary treatments of the Gramineae the subgenera of Hitchcock and Chase (1951) were elevated to seperate genera.

Dichanthelium--as subgenus or genus--is morphologically characterized by winter rosettes of short, broad leaf blades (though not all Dichanthelium species have this morphological charaacteristic) and secondary branching of the main shoot. These taxa have C3 photosynthesis and lack Kranz anatomy. They are cespitose though some have rhizomes. The main or principal shoot usually terminates in a primary panicle most spikelets of which are chasmogamous (chasmogamous is the adjective that refers chasmogamy, the condition in whith anthers and stigmas are exposed outside of their own flower so that there is potential for cross-pollination). Off of the main shoot there are typically several branches (secondary shoots) which have secondary panicles at least some spikelets of which are cleistogamous, the condition of natural self-pollination (Barker et al., 2003, p. 406). Members of Dichanthelium typically have vernal and autumnal phases (shoot forms), but vernal panicles set "little, if any" fruit. Instead, most grain is borne in the cleistogamous spikelets of the lateral, autumnal panicles (Gould, 1975, p. 478). It could be hypothesized that this cleistogamous flowering (autogamy) is responsible for the many different morphological forms (subspecies, varieties, or whatever) of the different Dichanthelium.

Notwithstanding the pronounced differences between Dichanthelium and Eupanicum (the main subgenus in Hitchcock and Chase [1951, ps. 627, 633-636] that does not have rosettes and shoot branching, and does have fertile spikelets throughout) or, in contemporary systematics, Panicum it is by no means clear that these are seperate genera. Even the distinction between cool-season versus warm-season is arbitrary from standpoint of flowering as vernal phase (shoots) of Dichanthelium members actually flower in summer not spring as is the flowering season for typical cool-season species like festucoid grasses.

The individual reader can reach his own conclusions. More importantly, it makes no difference from a practical standpoint other than scientific name (and related confusion). There are rosette panicgrasses--Dichanthelium or Panicum--and they are distinct from those related taxa that do not form basal rosettes and branches off of the shoot.

There is one thing that all who have worked with these plants can agree upon: they are an identifiction nightmare come true. Plus, they do provide palatable forage and certainly add biological diversity or--and no pun intended--variety to tallgrass prairie range and hay meadow.

As such, one of the major species of rosette panics was included below.

Note on organization: several rosette panicgrasses (Dichanthelium taxa) were included in the chapter, Tallgrass Savanna. Most of these examples grew within two to five miles of the example species shown immediately below. In fact some or, even, all of these species might well have grown nearer to each other in this area but the author simply did not find such situation. As found, the species treated below was growing on tallgrass prairie whereas those under Tallgrass Savanna were found growing in a black oak-mixed hickory-tallgrass savannah in the western Ozark (Springfield) Plateau.

89. Example of the rosette panicgrasses- Local colony of fascicled panicgrass (Panicum lanuginosum var fasciculatum= Dichanthelium acuminatum subsp. fasciculatum) This is one of the principle rosette panicgrass species of tallgrass prairie though less common and widespread than some of the others. It is, however, one of the most distinctive bearing, as it does, one of the most pronounced fascicles in the Dichanthelium group.

Ottawa County, Oklahoma. Late July, mid- to near maturity of grain (see below).

 

90. Vernal fascicles-Apices of the main shoots (not branches) of fascicled panicgrass bearing primary panicles. Actually these leafy units of the shoot are both panicles and clusters or bunches of leaves which by convertion are termed fascicles. Most, if not all, of the Dichanthelium taxa bear such fascicles. Some of the most distinctive and pronounced fascicles are on this species, Panicum lanuginosum var fasciculatum.

Ottawa County, Oklahoma. Late July, mid- to near maturity of grain.

 

91. Fancy fascicles- Details of fascicles on thee specimen of fascicled panicgrass shown in the two preceding slide sets. The somewhat pilose pubescence is a distinctive characteristic of this unusual species of the rosette panicgrass group.

Ottawa County, Oklahoma. Late July, mid- to near maturity of grain.

 

Another year, another specimen- Another example of fascicled panicgrass as it appeared in late spring (first slide) and, as presented at closer camera distance, some representative vernal (spring) fascicles of this individual plant (second slide).

Ottawa County, Oklahoma. Mid-June; immature grainstage of phenology.

 

Spring foliage- Vernal or spring fascicles of fascicled panicgrass. These organs were less mature vernal fascicles than those of mid-summer that were presented above. This growth was produced the next growing season after the grass growth shown above. Leaves of facicles had not reached their typical morphology and more clumped arrangement typical of maturity (see again, above).

Ottawa County, Oklahoma. Mid-June; immature grainstage of phenology.

 

Spring grains- Caryopses produced on vernal fascicles of fascicled panicgrassThere are also autumnal fscicles with their fall-borne caryopses.

Ottawa County, Oklahoma. Mid-June; grain in hard-dough phenological stage.

 

92. A fancy, fascicled, and favored grass- This fenceline contrast provided a dramatic example of the extreme palatability of most of the rosette panicgrasses. On the left side, under, and as far outside the fence as cows and calves could stick their tongues they had eaten (nearly to ground level) shoots of fascicled panicgrass. At the farther right, just beyond tongue reach, shoots of fascicled panicgrass remained untouched.

This dichotomy was a striking illustration of grazing selectivity and intensity, two of the important facets of range plant defoliation. Although this short-growing, cool-season, C3, panicoid grass has relatively high palatability it is not so palatable or vulnerable that it will be greatly reduced on ranges that receive proper grazing management. In this example, the tame pasture to the left which was dominated by common bermudagrass (Cynodon dactyledon), an introduced eragrostoid grass of moderate or lower palatability, and subjected to heavy utilization. Little wonder that cattle had reached as far under the fence as they could to get every bit of fascicled panicgrass within reach.

Ottawa County, Oklahoma. Late July in a fairly typical moisture year. (Note that this native, cool-season species was still green one third of the way through summer.)

 

92. Rounded out- Roundseed panicgrass (Panicum sphaerocarpon) on tallgrass prairie at western edge of Ozark (Springfield) Plateau. "Phytoneiighbors" include big bluestem, winter bentgrass (Agrostis hyemalis), tall nut-grass (Scleria triglomerata), and numerous forbs including ragged orchid (Habenaria lacera), yuccaleaf eryngo or button snakeroot (Eryngium yuccifolium), and lousewort (Pedicularis canadensis).

Roundseed panicgrass is one of numerous species of the rosette panicgrasses, the Dichanthelium subgenus of Hitchcock and Chase (1950) or Dichanthelium genus of some authorities including Gould (1975) and Great Plains Flora Association (1986). These are the panicgrasses with the Cs photosynthesis pathway and cool-season leaves arranged as a basal rosette. Distinction among species in the Dichanthelium group is a rangeman's nightmare. Roundseed panicgrass cannot be readily distinguished from some of the other rosette panicgrasses, but short and prouncedily purple spikelets and lack of ligule are key characteristic features.

Ottawa County, Oklahoma. Early June.

 

93. Dash of purple coloration- Upper portions of vernal sexual shoots (first and ssecond slide) and spikelets in a diffuse panicle of roundseed panicgrass (third slide) on a tallgrass hay meadow in the far-western Ozark Plateau. Roundseed panicgrass was one of several of the rosette panicgrasses, the taxon (whatever level a given author chose to put it) of Dichanthelium. These cool-season, C3, panicoid perennials produce both vernal and autumnal sexual (flowering) shoots, at least weather permitting.

Ottawa County, Oklahoma. Late May; soft dough phenological stage.

 

A drought can be deceiving: shown below were some examples of relatively large--heck, huge in relative terms--specimens of roundseed rosette panicgrass that grew in the late spring in the fifth consecutive year of drought that varied from Severe Drought to Extreme Drought on the Palmer Index. Several heavy rainfall events earlier in the current spring (especially two multi-inch thundershowers within the last month) resulted in shoots of roundseed panicgrass that grew to lengths of over four feet. Thereafter, it turned hot and dry and many shoots of big bluestem did not exceed three feet in stature.

When it rains at the right time- Dense stand of roundseed panicgrass plants of which grew to comparatively immense size after a brief repreive of thundershowers during a Severe Drought. Details of shoots (some over four feet in length) were shown in slides immediately below.

Western Springfield Plateau, Ottawa County, Oklahoma. Mid-June; grain fast reaching maturity.

 

Yep, they really did it- Sexual shoots of roundseed panicgrass that grew to lengths of slightly over four feet. These panicle-bearing shoots, which typically reach lengths of eight to roughly eighteen inches (at most), were so long that they "laid down" (ie. were in decumbent position).

What is more, these plants were growing on an old field that had been in tall fescue (Festuca arundinacea) for fifty plus years until killed out by a combination of factors during the prolongued five-year drought.

Western Springfield Plateau, Ottawa County, Oklahoma. Mid-June; grain fast reaching maturity.

 

What they really did- Increabily long panicles on sexual shoots--the combined length of which reached four feet--on roundseed panicgrass produced during the fifth copnsecutive year of protracted drought (Severe to Extreme Drought state on Palmer Index) after two heavy thunderstorms brought temporaty relief. This is an example of the opportunistic ability of range plants to utilize limited resources with incredible efficiency and speed when these resources become available.

This crop of native grain was produced on "go-back land" that had been in tall fescue for over a half century. A combination of drought and mismanagement killed out the tall fescue (see disdcussion of tall fescue in the Grassland chapter, Introduced Grasses), but this native rosette panicgrass got the same improper management--and it thrived instead of dying (= being killed) out.

Western Springfield Plateau, Ottawa County, Oklahoma. Mid-June; grain fast reaching maturity.

 

A little more common (and still impressive)- Two panicles of roundseed rosette panicgrass growing on an old field in late spring during the fifth consecutive year of Severe Drought to Extrmen Drought, but following recent heavy rains. These inflorescences were more typical panicles in contrast to the extraordinarily long panicles of some other plants of roundseed rosette panicgrass growing in the same general area at the same time.

All the rosette Panicum species are cool-season species with the C3 photosynthetic pathway.

Western Springfield Plateau, Ottawa County, Oklahoma. Mid-June; grain fast reaching maturity.

Location note: several other species (in addition to those shown here) of rosette panicgrass, Dichanthelium subgenus of Panicum, included in the grassland chapter, Tallgrass Savanna.

 

94. Fall witchgrass (Leptoloma cognatum= L. cognatum var. cognatum= Digitaria cognata)- Two photographs from the exact same focal point with the first taken under a shaft of full-sun light and the second, only minutes later, under an overcast sky.

Fall witchgrass is either one species of a two-species genus (Hitchcock and Chase (1950, p. 585), a single species consisting of two varieties that comprise a monotypic genus (Gould, 1975, p. 419), or two species among several in an expanded Digitaria genus (Barkworth et al., 2003, 362-363).

Classification of this species has long-been problematic. In Texas Grasses Silveus (1933, ps. 494-495) recognized L. cognatum as the only Leptoloma species in Texas and noted that it differed from Digitria species in having long-pediceled spikelets on a loose panicle in contrast to short-pediceled spikelets on thin racemes as in case of Digitaria species. In this vein, the other common name given by Silveus (1933, p. 494) for L. cognatum was diffuse crabgrass. In Botany of Western Texas (Coulter, 1891-1894, p. 508) had included fall witchgrass within an immense Panicum genus as P. autumnale closely aligned with such species as P. capilliare, P. hallii, P. virgatum, and P. bulbosum. Incidentially, the adjective "fall" almost assuredly was derived from the specific epithet autumnale.

Fall witchgrass is a locally important, perennial, forage species on Texas prairies and throughout the Edwards Plateau. Beginners in these areas often confuse fall witchgrass with the annual species, called witchgrass or common witchgrass (P. capilliare). Common association of these two species is one reason for this confusion even though they are distinctly different when campared hand-in-hand. There again, Coulter (1891-1894, p. 508) aligned these two species closely so calling one for the other in understandable.

Fall witchgrass (and common witchgrass for that matter) is a shortgrass or, possible interpreted as a midgrass species with panicles comprising one-third to one-half the height of the shoot. Hatch and Pluhar (1993, p. 127) described fall witchgrass as having fair feed value for both livestock and wildlife. This species is widespread in . It favors somewhat disturbed microsites and fits the classic description of an increaser. Its biological range extends from Ontario south to Florida and west to Texas northward to central Nebraska. Fall witchgrass is found in all ten vegetational (land resource) areas of Texas and in Oklahoma is absent primarily from the southeastern forest area.

Crinkled or wavy margins of leaves (at least on one edge) is a distinguishing feature that is sometimes--though by no means always-- present (Hatch and Pluhar, 1993, p. 127). This feature seems to be a more widespread characterfrom southern Oklahoma through central Texas (Coffey and Stevens, 2004, p. 73) than farther to the north and east in the Tallgrass Prairie Region.

Ottawa County, Oklahoma. Mid-July; full (though immature)-panicle stage.

 

95. Composite view- General view (under an overcast sky) of a portion of one plant of fall witchgrass showing features of leaves, culms, and panicle. This and the next two photographs showed that roughly a third to a half of total height of mature plants is in the comparatively large panicle.

Ottawa County, Oklahoma. Mid-July; full (though immature)-panicle stage.

 

96. Sexy increaser- Detailed composite views of sexual shoots of fall witchgrass showing most of the panicle and subtending shoot (first slide) and particulars of plant features at axillary region and point of panicle emergence from boots (second slide). Fall witchgrass is an increaser on most range sites with its natural increase by both sexual and asexual reproduction with vegetative increase by both tillers and rhizomes (Hatch and Pluhar, 1993, p. 127). Although semi-cespitose in habit, fall witchgrass develops into small colonies with daughter plants (modular units) arising from the short rhizomes.

The forb in the upper left of the first photograph (and also visible in the two preceding slides) was a species of tick-clover or tick-trefoil (Desmodium sp.).

Ottawa County, Oklahoma. Mid-July; full (though immature)-panicle stage.

 

97. Big leaves on a small grass- Details of leaves, including axillary area (collar), of fall witchgrass. Ottawa County, Oklahoma. Mid-July; full (though immature)-panicle stage.

 

98. Maximum development- Fall witchgrass with ripening caryopses on the panicle as seen from a more-or-less topdown view. Portion of a fairly large colony with several sets of shoots that arose from short rhizomes. This was a good example of both sexual and asexual (vegetative) reproduction in this increaser species that is widely distributed across southern grasslands.

Ottawa County, Oklahoma. Mid-July; full--and mature--panicle stage.

 

99. Bewitched panicle- Fall witchgrass panicle (first and second slide) and spikelets (third slide) growing on a tallgrass prairie-Cross Timbers transition range. Erath County, Texas. Early May (first two slides) and early October (thrd slide).

 

100. Paspalum plants- Two individual (two genetically unique) plants of field paspalum (Paspalum laeve) on degraded prairie in the Ozark Plateau. Individual shoots were discernible, but details of shoots were reserved for succeeding slides. These two plants provided typical examples of the habit (including the pattern of wide spacing of shoots) of this prairie and savanna grass species that usually grows to middle heights (usually not more than two and a half feet with roughly a foot and a half being more common on habitats typical of this region). This species has short rhizomes from which ascending tillers arise resulting in a somewhat at sprawling plant with widely spaced tillers. This habit was best seen in the first of these two photographs.

Field paspalum is frequently found (mostly as isolated plants or a few local concentrations of plants) on oak-hickory forest-tallgrass prairie savannahs, especially in the Prairie Peninsula, in the Ozark Plateau as well as on tallgrass prairie in the Central Lowlands physiographic province. Field paspalum appears to be adapted to a wide array of microhabitats from pristine to disturbed. The two plants in these photographs were both growing on roadsides subjected to infrequent heavy mowing. They were growing about a mile distant from each other.

There are a number of Paspalum species on tallgrass prairie and related mesic grasslands, but none of these are major (ie. dominant, abundant, critical or indicator) species. The paspalums are generally quite palatable, especially to cattle and horses, but they do furnish preferred forage in late maturity through dormant stages.

Ottawa County, Oklahoma. Mid-July early bloom stage.

 

101. The hairy leaf axils of field paspalum have been described by many workers (Coulter, 1891-1894; p. 499; while others such as Hitchcock and Chase (1950, p. 615) described sheaths as "glabrous or nearly so" and included plants similar to P. laeve but having pilose sheaths as P. longipilum. Steyermark (1963, p. 202) in Missouri Flora (the definitive flora closest to location of these photographs) distinguished between P. laeve var. pilosum with "strongly hairy" sheaths from P. laeve var. laeve with glabrous or nearly glabrous" sheaths (and which does not occur in the western Ozark Plateau) and from P. laeve var. circulare spikelets of which were "more or less circular or orbicular". Examples shown here obviously fit the discription of P. laeve var. pilosum. In Texas, Gould (1975, ps. 526-527) recognized three varieties of P. laeve including P. laeve var. pilosum which he interpreted as synomous with P. longipilum. Barkworth et al. (2003, ps. 572-573) eliminated three varieties of P. laeve and the species, P. longipilum listed in their synonymy (Barkworth et al., 2003, p. 748) and lumped all four of these taxa under P. laeve.

Another interesting feature of P. laeve is crinkled or corrugated margins (wavy edges) of leaves. This characteristic was shown in Hitchcock and Chase (1950, p. 618, Fig. 895) and Barkworth et al.(2003p. 573) This character was visible in the second and third slides in this three-slide set.

In essence, the Paspalum genus has survived--as of this writing--onslaughts of cladists, but there has been considerable variation in interpretation of taxa over admittedly relatively minor features such as pubescence of leaf axils.

Ottawa County, Oklahoma. Mid-July early bloom stage.

 

102. Lower parts- Details of basal shoots of field paspalum that was thriving on a disturbed roadside on a bottomland range site in the western Ozark Plateau. Field paspalum produces short rhizomes from which tillers arise.The first of these two slides showed several tillers with bulbous (sacculate or inflated-appearing) nodes. Such tillers frequently produce sister tillers or branches off of the parental shoot at these large, swollen-appearing nodes. Portions of two of these sister tillers (daughter shoots) were shown at close camera distance in the second slide.

Ottawa County, Oklahoma. Mid-July early bloom stage.

 

103. Sexual reproduction from asexual reproduction- In field paspalum older tillers that are still solely in vegetative stages (ie. are asexually reproductive) often have a semi-prostrate morphology (somewhat of a creeping habit) near the ground surface that ascends at higher and younger portions (see again whole-plant shots above). This results in a shoot that resenbles a stolon off of which daughter tillers (modules of vertical shoots) develop. The secondary (daughter) tillers are considerably shorter than primary (parental) tillers and these clonal offshoots frequently produce small infloresecences that usually do not fully emerge from the boot (enveloping sheath). Two such basal inflorescences (still in the boot) on shoot branches were visible in the first photograph.

The second photograph presented two secondary or daughter tillers (vertical sister shoots). The secondary shoot on the left had its nflorescence emerging from the boot (about to the final degree to which it would be expressed). The daughter tiller on the right had its inflorescence still completely encased within the boot. Pictorial representations of basal flower clusters in field paspalum were shown in line drawings in both Hitchcock and Chase (1950, p. 618, Fig. 895) and Barkworth et al. (2003, p. 573).

 

104. Cluster and spikelets- Details of a sexual shoot with inflorescence (first photograph) and separate branches with flowering spikelets (second and third photographs) of field paspalum on a disturbed bottomland range site in the western Ozark (Springfield) Plateau. The inflorescence of Paspalum species has been interpreted variously by different workers. Hitchcock and Chase (1950, p. 599) regarded the flower cluster as as consisting of "... one to many spikelike racemes, solitary, paired, or several to many on a common axis". Hignight et al. (1988, pos. 7, 28) provided one of the most exacting and specific categorizations of Gramineae inflorescence types ever and described the Paspalum inflorescence as a panicle of alternate spicate primary unilateral branches. Barkworth et al (2003, p. 572) interpreted the Paspalum inflorescence as a panicle of racemose (racemosely arranged) branches.

In P. laeve spikelets occur singlely yet strongly overlaping in two rows along the racemose or spicate primary unilateral branch. This arrangement produces a crowded or jumbled apperance of these broad, oval flowers and grains (second slide) on the branch or rachis. The third slide showed one of the unilateral panicle branches along most of its length at peak anthesis. Anthers and stigmas were visible, at least in the photograph prior to it being scanned in an Epson Perfection scanner (slip-shod equipment which is far from perfection in regards to clear, detailed reproduction.of images).

Ottawa County, Oklahoma. Mid-July early bloom stage.

 

105. The other dubious variety- It was discussed above that Steyermark (1963, p. 202) in Missouri Flora (the definitive flora closest to location of these photographs) distinguished between P. laeve var. pilosum with "strongly hairy" sheaths from P. laeve var. laeve with "glabrous or nearly glabrous" sheaths (and which does not occur in the western Ozark Plateau) and from P. laeve var. circulare, spikelets of which were "more or less circular or orbicular". Examples that were obviously P. laeve var. pilosum were presented immediately above. In this pair of slides and in the immediately following three slide/caption sets specimens of P. laeve var. circulare were presented.

It was explained above that Barkworth et al. (2003, ps. 572-573) eliminated the three varieties of P. laeve . By contrast, for Texas, Shaw (2012, p. 752) retained these three taxonomic varieties as did his predecessor Gould ((1975, ps. 526-527). This author, who was not a fan of Barkworth et al. (2003), presented examples of P. laeve that certainly fit the descriptions of P. laeve var. circulare. Whether these varietial distinctions are "real" (differences are consistent at the level of taxonomic variety) or were instead merely examples of genotypic/phenotypic variation within a highly variable species of P. laeve is perhaps a matter known but to the Great Taxonomist.

Clearly there were morphological differences among the specimens presented here that were growing separtely from each other over a distance of about four miles and in three different years. One of the most pronounced differences was the greater diameter of coarser shoots of specimens keying to P. laeve var. pilosum om contrast to smaller diameters of slender shoots of specimens keying to P. laeve var. circulare. It could do no harm to include more examples of field paspalum, which is rather sporadic in abundance across years, and demonstrate to students the morphological variability that can exist within what all taxonomic keys showed to be a single species of range grass.

Ottawa County, Oklahoma. Late July, early bloom (pre-anthesis) phenological stage.

 

106. Widely spaced shoots- Top-down view of field paspalum in the Ozark (Springfield) Plateau of northeastrn Oklahoma. It was explained above that morphological features of field paspalum included relatively wide spacing among shoots that often had a decumbent (or partly so) growth form that gave plants of this species a "sprawling" habit. That morphological pattern was visible in this photograph of a specimen that was at peak standing crop and in early bloom stage.

Ottawa County, Oklahoma. Late July, early bloom (pre-anthesis) phenological stage.

 

107. Field tops- Upper shoots (mostly sexual) and branches of inflorescences of field paspalum (P. laeve var. circulare) on a heavily mowed (until this year)--and so mostly a former--tallgrass prairie in the western Springfield Plateau in northeastern Oklahoma.

Ottawa County, Oklahoma. Late July, early bloom (pre-anthesis) phenological stage.

 

108. Orbited in the field- Three progressively closer images of branches with spikelets of field paspalum of the taxonomic variety P. laeve var. circulare. The perhaps arbitrary (imagined) "orbicular" (Steyermark, 1963, p. 202; Shaw, 2012, p. 752) shape of spikelets (they were slightly wider than long) was evident in these images. These examples were on some of the plants presented above that were growing on a degraded (and, mostly, former) tallgrass prairie in the western Ozark Plateau.

Ottawa County, Oklahoma. Late July, early bloom (pre- to early anthesis) phenological stage.

 

What it can do with more water- Large local stand of field paspalum growing on a wet prairie in the western Springfield Plateau section of the Ozark Plateau physiographic province in northeastern Oklahoma. Rather than large individual plants like those presented above these were views of more plants of smaller size with more tillers under conditions of greater soil moisture.

.Ottawa County, Oklahoma. Late July,

 

More views with more water- Upper portions with inflorescences (first slide) and an entire flower cluster or inflorescence (seond slide) of field paspalum growing on wet prairie in the western Springfield Plateau section of the Ozark Plateau physiographic province in northeastern Oklahoma. Most of the inflorescences produced in this stand had fewer branches thaan the one featured in the second photogrph.

.Ottawa County, Oklahoma. Late July,

 

A confusing infloresecence- The flower cluster of the Paspalum species is confusing at best and open to interpretation by different agrostologist. Your author followed a combination of descriptions including a spikelike raceme (Hitchcock and Chae (1951) or as a panicle of alternate spicate primary unilateral branches in the classification of Hignight et al. (1988, p. 7). The first photograph seen here showed an entire inflorescence with fewer unilateral branches while the second photograph featured one unilateral branch (of a spikelike raceme or of a panicle of alternate spicate primary unilateral branches). A mouthful and continuing confusion no matter what terms aree used.

.Ottawa County, Oklahoma. Late July,

 

109. Ready for another paspalum?- Specimens of fringed, fringeleaf, or silky paspalum (Paspalum ciliatifolium) growing on an abused (closely and frequently mowed) and degraded tallgrass prairie in the Springfield Plateau in northeastern Oklahoma. This species

There are very few references in regards to what are known as the "low paspalums" (in contrast to taller-growing species such as P. floridaum, Florida paspalum). One of the best--albeit brief--reference sources for fringeleaf or silky paspalum (including grazing value and management) was Leithead et al. (1971, p. 321-133).

Ottawa County, Oklahoma. Early July.

 

110. Some more perspectives on a silky one- Oblique sideview of three plants (first or upper slide) and top-down view of one plant of fringeleaf or silky paspalum growing on a degraded (heavily, frequently mowed) tallgrass prairie in the Springfield Plateau in northeast Oklahoma.

The "low paspalums" typically have a "sprawling" or "crawling" habit (outward appearance) which is a consequence of widely spaced (comparatively speaking) tillers that tend to bend over with development of their inflorescences. Also, these tillers are often decumbent at their lower or basal phytomers (shoot units consisting of node, internode, and leaf) which contributes to the appearance of a "sprawling" appearance. .

Ottawa County, Oklahoma. Early July.

 

111. Fringed on top- Shoots, including upper sexual shoots with inflorescences, (first slide) and inflorescences emerging from their boots (second slide) of fringeleaf or silky paspalum. These organs were produced on plants growing on wet tallgrass prairie in the western Ozark (Springfield) Plateau.

Ottawa County, Oklahoma. Early July; early fruit (mostly milk) stage.

.

112. What fringes; and, where's the silk?- Two inflorescences with immature spikelets of silky or fringed paspalum growing on a degraded tallgraass prairie in the western Ozark (Springfield) Plateau.

Ottawa County, Oklahoma. Early July; early fruit (mostly milk) stage.

 

113. Silky (sort of) on the bottom- Part of base of plant (first slide) and basal shoots (second slide) of fringed or silky paspalum on degraded tallgrass prairie in the Springfield Plateau in northeast Oklahoma.

Ottawa County, Oklahoma. Early July.

 

114. Paspalum on the prairie- Local stands of Florida paspalum (Paspalum floridatum) on mesic tallgrass prairie in the western Ozark Plateau. Florida paspalum is typically the largest of the Paspalum species on tallgrass prairie and savannas of tallgrass prairie and oak-hickory forests. On coastal tallgrass prairie such as those of the Gulf coast there are Paspalum speaacies that grow to larger size, but on prairies of the continental interior Florida paspalum generally takes the size prize. It is also the single most important Paspalum species in the Tallgrass Prairie Region. For example, it was the only Paspalum species included in Field Guide to Oklahoma Plants by Tyrl et al. (2008).

Leiithead et al. (1971, p. 137) and Tyrl et al. (2008, p. 147) reported that Florida paspalum is quite palatable, especially to cattle, though forage quality declines rapidly in this succulent species after dormancy (a situation that applies to all Paspalum species). Given this palatability, Florida paspalum is apt to be heavily utilized so as to benefit from periodic deferment during the growing season every second or third year (Leithead et al., 1971, p. 137).

The Paspalum species are similar in general apperarnce and, in case of some of the native species, in morphology of inflorescences and details of spikelets. The color photographs in Coffey and Stevens (2004, ps. 89- 94) are useful in helping beginners (sometimes even "experts") distingush among Paspalum species common in southern Oklahoma and northcentral Texas. Barkworth et al. (2003, ps. ) is the new definitive source for North American paspalums.

Ottawa County, Oklahoma. Late July; peak standing crop and anthesis.

 

Wet soil for big shoots- Two examples of flowering Florida paspalum growing to robust size and rank growth habit on wet prairie in the western Springfield Plateau section of the Ozark Plateau physiographic provincce.f

Ottawa County, Oklahoma. Late July; peak standing crop.

 

115. Passing for a paspalum - Basal shoots of Florida paspalum growing on a wet prairie in the Springfield Plateau. Associated species of graminoids (the prevalent categories of range prairie plants on this habitat) included switchgrass, field paspalum, two-flower (two-flowered) rush, slim-pod or diffuse rush, and fox caric-sedge.

Ottawa County, Oklahoma. Early August, peak standing crop.

 

116. Hard one to categorize- Young inflorescence emerging from the boot (first slide), entire inflorescence (second slide) and uppermost (distal) parts of an inflorescence (third slide) of Florida paspalum produced on wet prairie in the western Ozark (Springfield) Plateau. The flower cluster of the Paspalum species defies "tidy" classification (not to mention description). Diggs et al. (1999, p. 1304) described the Paspalum inflorescence as having a central axis with "...one to many spike-like, unilateral branches". Probably the most thorough descriptions of "specialized panicle inflorescences" remains that of Hignight et al. (88, p. 7). By this descriptions the flower cluster of Florida paspalum is "a panicle of alternate spicate primary unilateral branches".

Technical note- the first slide was a fine 35 mm Provia 100 slide that was overexposed in scanning and converting to a Jpeg by an Epson Perfection 700 scanner. It may be a sorry craftsman that blames his tools, but viewers can see from most other slides that their photographer pretty much knew his craft. It is was an inferior tool, the Epson scanner, that botched the image being discussed. Adobe PhotoShop helped to partially restore the image to what is was before Epson botched it.

Reason for this aside comment was to serve as a warning to would-be buyers of scanners. DO NOT BUY EPSON SCANNERS; TRY A SLIDE-DESIGNATED SCANNER LIKE NIKON. TOO LATE FOR THIS PHOTOGRAPHER. POTENTIAL SCANNER BUYERS BEWARE.

Ottawa County, Oklahoma. Early August, peak standing crop and anthesis phenological stages.

 

117. Part of the flower cluster- Two examples of the "alternate spicate primary unilateral branches" (Highnight et al., 1988, p. 7) of Florida or big paspalum produced on wet prairie in the Springfield Plateau.

The inflorescence of Florida paspalum is extremely attrative in anthesis as seen here and up closer in the next two-slide set ...

Ottawa County, Oklahoma. Early August; peak anthesis.

 

118. Study in grass sex- Anthesis in Florida paspalum with anthers and stigmas exerted simultaneously. Florida paspalum is one of the best examples of a native prairie grass in which to show the sex organs because typicall both female and male organs are expressed at the same time.

Ottawa County, Oklahoma. Late July; peak anthesis.

 

"My baby got something she calls her stingaree"--Stingaree, by Bob Wills and the Texas Playboys .

119. Sexy orthopteran on sexy grass- Female straight-lanced meadow katydid (Conocephalus strictus) on a primary unilateral branch of Florida paspalum. The staight lance (or is it a "stingaree"?) is this "baby's" ovipositor, the elongated tubular organ through which she deposits her eggs (= oviposits). The straight (versus curved) ovipositor of the straight-lanced katydid is unmistakable both as to 1) the species of meadow katydid and 2) the sex of the individual insect. Sort of an application of the song and song line line by Georgia White, "I just want your stingaree".

Ottawa County, Oklahoma. Late July.

 

120. Spent panicles in an azure sky- Branches of panicles of Florida paspalum with most spikelets already shed. A contemporary interpretation of the Paspalum panicle is that of "unilateral spicate branches" (Gould, 1975, p. 500) or, more specifically, a specialized panicle of alternate spicate primary unilateral branches (Hignight et al., 1988, p.7) That arrangement was visible with these two panicles set against a late autumn sky. This bright blue sky was the remnant atmospheric phenomenon of a "blue norther", a hugh mass of cold Arctic air (the Siberian or Yukon Express or Alberta Clipper), on the second day of its bitter reign. God's gift to shutterbugs capturing autumn and winter skies.

Ottawa County, Oklahoma. Mid-December, grain-shatter stage.

 

121. Ripe grains- Spikelets of Florida paspalum with mature caryopses just prior to shedding. Readily visible was the feature of dorsally compressed caryopses, an identification feature made famous by Hitchcock and Chase (1952). This example was from a small tallgrass prairie in the western margin of the Ozark (Springfield) Plateau.

Ottawa County, Oklahoma. Mid-December, grain-ripe to grain-shatter stages of phenology.

 

122. Field paspalum (Paspalum laeve)- Local colony of field paspalum on a tallgrass prairie at western edge of Ozark Plateau. P. florida and P. laeve are the two Paspalum species with the largest plants on the prairies and savannas of the Ozark Region. Field paspalum is smaller than Florida paspalum in all features, but both are among the largest Paspalum species native to the mid-continent of North America.

The Great Plains Flora Association (1986, p. 1205) recognized two taxonomic varieties of P. laeve for the general Great Plains Region while Steyermark (1963, p. 202) described three varieties in Missouri and Gould (1975, ps. 526-527) recognized three varieties in Texas. Two of the three varieties described by Steyermark (1963, p. 204) were recorded for the county contiguous with the Oklahoma county in which these speciemens were photographed.

Ottawa County, Oklahoma. Late June; maturing to grain-ripe stage.

 

123. Tops in the field- Top-sown view (first photograph) and side view (second photograph) of apical parts of shoots of field paspalum on a tallgrass prairie in western Ozark Plateau. This phytomass was produced in late spring after a wet (and extremely cold) winter followed by severe drought (at time of photographs).

Ottawa County, Oklahoma. Late June; grain-ripe/shatter stage (see spikelets on racemes, racemose branches of panicles, in three slides below).

 

124. Field paspalum in an Ozark prairie- Specimen (first photograph) and inflorescence (second photograph) of field paspalum on a prairie situated among stands of black oak-dominated tallgrass savanna in the western part of the Springfield Plateau portion of the Ozark Plateau Region. Few of the grasses are photogenic, especially when trying to feature one plant of a species that was growing amid other plants of other species. This was the best this photographer could pull off for one field paspalum keeping company with beaked panicum, flowering paspalum, big bluestem, and broomsedge bluestem.

Ottawa County, July: milk-stage right after anthesis.

 

125. Some more samples- Three more specimens (actually more than that given that there were two or three plants in the first two of these three slides) of field or smooth paspalum growing on a moist tallgrass prairie in the Springfield Plateau of the Ozark Uplift. This area was part of the Prairie Peninsula (Transeau, 1935).

Ottawa County, Oklahoma. Late June; early ripening grain stage.

 

126. Deserves closer looks- Field or smooth paspalum on tallgrass prairie in the western Ozark Highlands (Springfield Plateau) in early summer. These plants were in part of the Prairie Peninsula first described by Transeau (1935). The interesting part of the story regarding these plants is that they were growing at edge of a tallgrass prairie that had been closely mowed (shredded with a rotary lawn mower) as part of a yard for over sixty years. Upon death of the rural couple who had kept this yard turf closely cropped, mowing was much less frequent and with higher mowing heights so that in about four years smooth paspalum was able to development to stages of maturity seen here.

The question that begs answering was, did a) smooth paspalum establish from a soil seed bank after decades of severe defoliation or b) had it persisted as closely mowed plants over all or most of the heavily defoliated time span and been able to develop to this grain production stage only with less frequent and intensive defoliation (ie. fewer shreddings and higher shreddding heights)? The question must remain unanswered given lack of observations under the more severe defoliation regime.

Ottawa County, Oklahoma. Late June; early ripening grain stage.

 

127. Starting to ripen in the field- Grain (= spikelets) maturing or ripening on racemes of field or smooth paspalum on tallgrass prairie in the western Ozark Uplift (Springfield Plateau). Recent elaborate description of grass infloresences interprets that of the Paspalum species as being a panicle of racemose branches (first or upper slide) and with these branches regarded as racemes (second and third slides).

Spikelets seen here were produced on what had been a domestic or household yard subjected to extreme defoliation (repeated close shredding with a rotary lawn mower) over more than 60 years. Roughly four years after replacement of the extreme defoliation regime with a less frequent, higher mowing height management, field paspalum was able to produce spikelets as seen here. It cannot be known if smooth paspalum had persisted in vegetative (nonsexually reproductive) stages over course of the extreme yard-mowing regime or if plants had reestablished from a soil seed bank with less frequent, less intense defoliation.

Ottawa County, Oklahoma. Late June; early grain-ripe stage (see spikelets on racemes, racemose branches of panicles, in three slides immediately below).

 

128. Foolproof feature- Ripe grain (spikelets) of racemes or racemose branches of a panicle of field or smooth paspalum. Field paspalum has one feature that differs from all the paspalums indigenous to the tallgrass prairie: spikelets are arranged/attached singly or individually in contrast to paired spikelets in other Paspalum species (Steyermark et al. 1963, p. 200; Great Plains Flora Association, 1986, p. 1204; Diggs et al., 1999, p. 1305). This field as well as herbarium identifying characteristic was visible in these three slides. Spikelets are aligned opposite each other in two rows, one row on each of the two sides of the rachis.

Ottawa County, Oklahoma. Late June; grain-ripe/shatter stage (spikelets on racemes, racemose branches of panicles).

 

129. Scattering some- Shattering spikelets of field or smooth paspalum. These spikelets (= grain or grains) are interpreted as being on racemes or, more precisely, as on racemose branches of panicles. These organs were produced at edge of a moist tallgrass prairie in part of the Prairie Peninsula (Transeau, 1935) in the Springfield Plateau of the Ozark Uplift.

Ottawa County, Oklahoma. Late June; grain-ripe/shatter stage.

 

130. Side-by-side paspalums- Florida paspalum (Paspalum floridatum) and smooth or field paspalum (Paspalum laeve) growing and with inflorescences overlapping on a wet tallgrass prairie in part of the Prairie Peninsula within the Springfield Plateau of the Ozark Uplift. The racemose-branched panicles of these two Paspalum species were presented and described variously above.

In these three photographs panicles of Florida paspalum were still in the boot (inside the flag leaf, the leaf immediately subgtending or below the panicle) whereas the smaller panicles of field paspalum were fully expressed and, in the second and third slides, seen to the immediate right of the larger, still-enclosed panicle of Florida paspalum.

Details of these panicles, racemose branches, and spikelets were presented in the very next two-slide/caption set.

Ottawa County, Oklahoma.

 

131. God-given arrangement- Closer-in, detailed views of panicles of field or smooth paspalum (one with three racemose branches) and Florida paspalum (larger one whose racemose branches were still still pressed together along the rachis, the central stalk of the panicle). These were the same panicles introduced in the immediately two preceding slides of the last slide/caption set.

These organs had been produced on wet tallgrass prairie in the western portion of the Prairie Peninsula within the Springfield Plateau of the Ozark Highlands.

This slide/caption set was entitled "God-given arrangement" because their intertwined, touching-each-other positions were naturally occurring and so educationally fortunitious. God is always a good pardner to have around when taking range photographs (or doing any other honorable activity for that matter).

Ottawa County, Oklahoma.

 

132. Panicoid pals- Co-dominant genera of Panicum and Paspalum were represented on a wet prairie in the western Springfield Plateau by beaked panicgrass (Panicum anceps) and Florida paspalum (Paspalum florida) at early (anthesis) peak bloom and peak standing crop.

Ottawa County, Oklahoma. Late July.

 

133. Thin paspalum or thinseed paspalum (P. setaceum var. stramineum= P. setaceum)- Thin or thinseed paspalum is arguably the single most common Paspalum species in the West Cross Timbers of northcentral Texas. It is propably an increaser on range sites of lower production potential, but Dyksterhuis (1948, p. 355) plainly described fringeleaf paspalum as an invader. By whatever name or taxonomic treatment this species, variety, (or whatever) is interpreted as, the grass is well-adapted to disturbance. Forage value is fair primarily due to relative low herbage yield. It is relatively palatable to cattle and horses when immature and of considerably lower forage value for deer. Dyksterhuis (1948, p. 355) described fringeleaf paspalum as " the principal perennial grass of the Main Belt" of the Western Cross Timbers and specified that it was a large component of livestock diets, it being grazed preferentially at some seasons and avoided at others.

P. setaceum var stramineum (= P. stramineum) as considered herein is a confusing species including in Texas up to four varieties (Gould, 1975, Natural Resources Conservation Service, 2003) or several taxa recognized by some authors as separate spceies including such better known ones as P. ciliatifolium, P. stramineum, and P. debile, (Gould, 1975, ps. 523-526; Diggs et al., 1999, p. 1308). Barkworth et al. (2003, ps. 589-592) recognized a total of nine varieties of P. setaceum for North America (north of Mexico). The taxon or form of this Paspalum species in the West Cross Timbers has historically been designated as P. ciliatifolium with the common name of fringeleaf paspalum (see for eg. Dyksterhuis, 1948, ps. 3490, 355). Barkworth et al. (2003, p. 590) specified that fringeleaf paspalum was "the most vriable and widespread" of the nine varieties of P. setaceum. This range of this variety extends as south as Panama and Bermuda. This group of grasses with varieties interpreted as separate species have been known generically as the low paspalums (Leithead et al., 1971, p. 133).

Erath County, Texas. Early September; early grain (milk) stage.

 

134. Done for the year- Dead shoots of knotroot bristlegrass (Setaria geniculata) shortly after this plant entered dormancy. Knotroot bristlegrass is a widely distributed panicoid grass that prefers mosit habitats such as those along streams or seeps (Gould, 1975, p. 546). It is a common, characteristic species on the Fort Worth or Grand Prairie in northcentral Texas especially on disturbed sites (Tyrl et al., 2008, p. 161) such as overgrazed ranges where the scarcity of climax little bluestem and Indiangrass results in reduced competition.

Erath County, Texas. November, grain-shatter at dormant stage.

 

135. Opposite ends- Two dense panicles (first photograph) and the characteristic swollen nodes on short rhizomes (second sphotgraph) of knotroot bristlegrass. This species has one of the more descriptive common names of grasses, but one does have to pull a specimen up to see the source.

Erath County, Texas. November, grain-shatter at dormant stage.

 

136. New cords and old cords- Prairie cordgrass growing in the Platte River Valley with green shoots of the current growing season along with dead shoots . from last summer's growing season. Adams County, Colorado. Early July; pre-boot stage of phenology.

 

137 Collar and ligule- Two views of the axillary (leaf axil) portion of shoots of prairie cordgrass. Serious and comprehensive presentations of grass features should include (when possible) details of collar, ligule, sheath, blade, node, internode, veination, and so on.

Adams County, Colorado. Early July; pre-boot stage of phenology.

 

138. Bottom portions- Lower portions of shoots of prairie coregrass. The first slide was of shoot bases of both dead (from the previous warm-growing season) and live (current spring andsummer-growing season) Spartina pectinata shoots growing on the floodplain of Platte River. The second photograph included rootcrowns and a short (young) rhizome of this strongly rhizomatous, native, eragrostoid grass. This C4 grass reproduces asexually via rhizomes and tillers so that it is often found large and (apparently or presumedly) clonal stands.

Prairie cordgrass usually forms relatively extensive colonies in more mesic areas such as prairie sloughs (see above where vegetation of range types was featured). Preference of prairie cordgrass for sloughs is basis for another common name of sloughgrass (Shaw, 20008, p. 183), but this common name should be reserved for Beckmannia syzigachne which has the preferred (published) common name of American sloughgrass. This latter species frequently occurs sympatrically with prairie cordgrass, but is an annual and does not form expansive colonies.

Adams County, Colorado. Early July; pre-boot stage of phenology.

 

139. Crowned up and starting to crawl- Rootcrown and rhizome of prairie cordgrass on the floodplain of Platte River. These two photographs presented the the scaly phytomers (tough bract [sheath]-covered node-internode units) of shoots--both tillers and rhizomes--at or just below soil surface (O horizon). The first slide showed a rhizome growing outward from two tillers (top of slide) with a new tiller (daughter or sister shoot) arising from the rhizome and then either the rhizome growing still outward or, alternatively, the daughter shoot producing a rhizome (depending on interpretation). The second slide showed the large number of basal phytomers of a tiller and another tiller (sister or clonal unit; another module) from which a short (immature) rhizome was growing. The rhizome is also a sister or sister shoot although horizontal and completely subterranean in contrast to daughter tillers. The vertical tillers are intravaginated shoots (ie. the outward growing shoot does not perforate or pierce the invaginating sheath) whereas the horizontal rhizomes are extravagated shoots which do perforate of burst through the enveloping sheath(s).

Observe the organic matter-rich alluvium of the Platte River floodplain that served as the natural medium for growth of this rank, perennial, tallgrass species.

Adams County, Colorado. Early July; pre-boot stage of phenology.

 

140. Crawling quite a way- A long rhizome of prairie cordgrass against a backdrop of tillers growing in the fertile alluvium of the Platte River floodplain. Obviously, this specimen (like those in preceding photographs) was dug out and exposed for photographic purposes.

Adams County, Colorado. Early July; pre-boot stage of phenology.

 

141. Cluster of racemes- Inflorescences of prairie cordgrass growing on a plains cottonwood-mixed prairie savanna that developed along an intermittent stream in the Colorado Piedmont. The inflorescences of Spartina species are confusing. Traditionally the branches with units of spikelets aligned along them were interpreted as racemes or, maybe, spicate racemes, but as these spikelets are sessile this view was at least somewhat inconsistent with standard definitions. Hitchcock and Chase (1950, p. 508) regarded the Spartina flower cluster as being made up of "... appressed or sometimes spreading spikes racemose on the main axis". Later, more detailed (sophisticated) descriptions of grass inflorescences recognized specialized panicle inflorescences (Highnight et al., 1988). Using this scheme, Stubbendieck et al. (1992, p. 107) described the inflorescence of prairie cordgrass as a "panicle of 6-40 spicate pirmary unilateral branches". Shaw (2008, p. 183) described the inflorescence of prairie cordgrass as a panicle with racemosely arranged, appressed, alternate branches.

East Bijou Creek, Elbert County, Colorado. Mid-July; peak-bloom stage (full anthesis).

 

142. Anthers, anybody?- Two close-up views of racemes in full anthesis on prairie cordgrass. The first photograph presented a standard side-view perspective of the raceme while still on the plant whereas the second photograph gave a ventral view of a raceme that was possible only by removal from the shoot. (The latter was laid on cattle dung in situ on a floodplain cottonwood-mixed prairie range that developed along an intermittent stream.)

There are often so many spikelets (one-flowered and sessile) on the racemose, unilateral branches that these floral units overlap (ie. imbricate spikelets). The Spartina species often produce some of the greatest number of anthers of any grass genera native to North America which in combination with imbricate spikelets produces an extremely showy array of exerted anthers. That characteristic was seen in these photographs.

East Bijou Creek, Elbert County, Colorado. Mid-July; peak-bloom stage (full anthesis).

Sporobolus complex- A group of closely Sporobolus species, varieties of one species, or varieties of two or more species with a wide biological range have long been recognized as important on tallgrass prairie. Tall dropseed, as a catchall common name, has been treated as including up to three varieties of the species S. asper or, as a synonym, S. compositus. . The specific epithet asper means "tall". More recently it was discovered that the author of S. asper used as basis of his scientific name an earlier designation that by international botanical rules was "illegitimate" (Tyrl et al., 2008, p. 173) so that the binomial now regarded as correct is S. compositus.

If this confusion was not enough there has been the long-standing view that S. compositus (S. asper) is so "morphologically variable" that perhaps it is "conspecific" with S. clandestinus (Tyrl et al., 2008, p. 173). All told this mess amounts to what taxonomists regard as a "complex". The Sporobolus compositus complex is a difficult assemblage of forms, perhaps affected by their primarily autogamous breeding" which is largely responsible for these plants being able to "perpetuate unique character conbinations" (Barkworth et al., 2003, ps. 121-122 ). Autogamy refers to self-fertilization within a flower. Much of this autogamy occurs in cleistogamous spikelets, those having "self-contained" fertilization (Gould, 1975, ps. 303, 631).

By whatever name(s)--at species or variety level--there are at least three taxonomic forms under the scientific "handle" for a generic tall dropseed. Presented immediately below were two of these: 1) "regular" tall dropseed (S. compositus var. compositus= S. asper var. asper) and 2) meadow dropseed (S. compositus var. drummondii = S. asper var. drummondii).

Weaver and Clements (1938. p. 520) regarded tall dropseed as a midgrass and while a dominant of true prairie a frequent member of tallgrass prairie. On tallgrass prairie tall dropseed commonly achieves heights of four to five feet, a stature roughly equal to that of little bluestem growing on tallgrass prairie Given this fact, it appeared to the current author that tall dropseed could be either a tall- or midgrass species depending on range site, especially edaphic features of it range environment.

 

143A. Autumn attire- Stand (first and second slide) and single cespitose plant (third slide) of tall dropseed in its late autumn state of foliage under dry, mild atmospheric conditions following a comparatively mist late (late summer-early autumn) growing season. These plants were growing in an outside fencerow on a mesic tallgrass prairie in the western Ozark Plateau.

Tall dropseed was regarded by Weaver and Clements (1938, p 520) as a mid-grass dominant of true prairie that commonly occurred on tallgrass prairie.

Ottawa County, Oklahoma. Mid-December.

 

143B. Staying inside their shoots- Two progressively closer views of upper portions of sexual shoots of tall dropseed growing in an outside fencerow on a mesic tallgrass prairie iin the western Ozark (Springfield) Plateau. The contracted panicles of this tallgrass species characteristically remain partly to almost completely enclosed within the boot much like the more common and widely distributed sand dropseed (Sporobolus cryptandrus), a characteristic species of mixed prairie. Details of the enclosed contracted panicle of tall dropseed were featured immediately below.

Ottawa County, Oklahoma. Mid-December.

 

144 . Tall tops- Sexual shoots of tall tropseed (Sporobolus asper var. asper= S. compositus var. compositus) with spikelike panicles retained within th r boots (enclosing sheath). The "dead giveaway" of tall dropseed is the "buggy whip"-appearing extension of the panicle-enclosing sheath. Unfortunately this very distinctive feature was often missed by botanical artists so that illustrations of this species (variety) frequently did not show the most foolproof feature for field identification. For example, this 'buggy whip" flagellate extension of the enclosing sheath was not shown in Hitchcock and Chase (1952, p. 416) nor in the popular Phillips range and pasture plant book (Phillips Petroleum Company, 1963; Fort Hays State University, 2006, p. 65). One can pretend that the prolonged extension of the sheath had been grazed or weathered off in these publications. Fortunately, more accurate renditions were shown in Barkworth et al. ( , p. ) and Tyrl et al. (2008, p. 172).

Ottawa County, Oklahoma.

145A. Grraminoid buggy whips- Two inflorescences of tall dropseed growing on degraded tallgrass prairie ranges: the first from the Ozark Plateau and the second from the West Cross Timbers. The flagellate extension of the sheath that encloses the contracted panicle of Sporobolus asper var. asper (= S. compositus var. compositus) is a very distinctive feature of this variety. This morphological characteristic tends to be more commonplace in wet years. Both of these sexual shoots were produced in a widespread, ususually wet late summer through autumn

First slide: Ottawa County, Oklahoma. Mid-December. Second slide: Erath County, Texas, Early November.

 

145B. Pushed and bleached out- Basal part of a sun-faded spicate (contracted) panicle of tall dropseed within its enveloping sheath. This partial exertion (or, same difference, retention) of the panicle within the boot is a feature of several of the Sporobolus species. The fruit (achenes in strict definition) had shed from this contracted panicle which was the same one presented in the immediately preceding photograph.

Erath County, Texas, Early November, fruit-shatter stage.

 

146. Bunches on the meadow- Several plants of meadow dropseed (Sporobolus asper var. drummondii= A. compositus var. drummondii) Although perennial dropseeds are generally increaser species, meadow or Drummond's dropseed is a member of the climax grassland community in some types of tallgrass prairie. Such is the case for specific range sites of the Grand or Fort Worth Prairie in northcentral Texas. There are several varieties (or whatever taxa is used by this or that authority) that comprise a Sprorbolus asper (S. compositus) complex.

At this stage of development all leaf and most of culm growth had already taken place. Flowering would not be until late summer.

Erath County, Texas. Early May.

 

147. Upper end- Mid-portion of sexual shoots (first slide) and largely enclosed, contracted panicle (second slide) of meadow of Drummond's dropseed on the northcentral Texas Grand Prairie.

Erath County, Texas. Late October; fruit (achene)-shatter phenological stage.

 

148. A less common dominant of tallgrass prairie- Two plants of prairie dropseed (Sporobolus heterolepis) on a dry-mesic chert tallgrass prairie in the western Ozark (Springfield) Plateau. It was explained in the grassland chapter, True Prairie, that prairie dropseed is a key species of that Clementsian association. As a general rule on tallgrass prairie, prairie dropseed is far less important than the Four Horsemen of the Prairies (big bluestem, little bluestem, Indiangrass, and switchgrass). In fact, prairie dropseed is sparse to typically absent from most of the actual tallgrass (bluestem-Indiangrass) range type. It is pretty much the same situation with porcupinegrass (Stipa spartea), these two speciess being more indicative of true prairie.

There is, however, a zone of tallgrass prairie to the east of the main body of tallgrass prairie (the Osage Questas, Flint Hills, Smoky Hills complex) that occupies parts of the Ozark Plateau and Osage Plains (Cherokee Prairie) as though this grassland is a frontier between the oak-hickory forest and the main body of tallgrass prairie. It is in this vegetational transition that prairie dropseed (and, somewhat less so, porcupinegrass) occur as co-dominants with big bluestem, the more pronounced dominant (larger, taller, more pituresque plants). Nelson (2010, p. 282) showed this. An idea of importance of prairie dropseed could be gained through such sources as well as from the species distribution map (2103) in Atlas of the Flora of the Great Plains (Great Plains Flora Association, 1977, p. 526).

Prairie dropseed is one of the most readily identified and conspicuous of the prairie grasses in the non-flowering stages of growth and development. Steyermark (1963, p. 166) described prairie dropseed as forming "conspicouous large clumps of handsome narrow, almost hairlike, long blades, which are mostly basal and curl or arch gracefully backwards".

Yellow composites in the second photograph were black-eyed susan (Rudbeckia hirta). There were several composite species in both photographs.

Diamond Grove Prairie (Natural Area), Newton County, Missouri. Mid-June.

 

149. The cespitose habit- Several palnts (first slide) and a single plant of prairie dropseed on a small chert glade in the Springfield Plateau. Behind the single prairie dropseed plant in the second slide was a plant of prairie dock (Silphium  terebinthinaceum). These two species are typically associated with one another.

Prairie dropseed is typically more characteristic and indicative of true than tallgrass prairie, but this species is found on grasslands of both of these Clementsian associations-- when they are climax and higher seral stages (eg. subclimax).

McDonald County, Missouri. July; pre-bloom phenological stage.

 

150. Short with a puffy boot- Puffsheath dropseed (Sporobolus neglectus) on a shallow habitat of a deteriorated range. Typical example of the small, annual species. Most of the Sporobolus species are perennials that are most abundant on abused ranges, especially overagrazed pastures. The same is true of puffsheath dropseed, but it is even more common on abandoned fields in early seral stages (ie. go-back land or old fields). Puffsheath dropseed is not so much a pioneer or initial colonizing species as it a second-stage invader although it is commonly associated with colonizing annual grasses and forbs.

Tyrl et al. (2008, p. 177) explained that S. neglectus along with two or more other Sporobolus species constitute an "interbreeding comples". This integrading group has been intrerpreted variously by different authorities, but to this authors eyes S. neglectus is noticably distinct due to its diminutive size and short inflorescences.

Tarleton State University Hunewell Ranch, Erath County, Texas. Late October, fruit-ripe phenological stage.

 

151. Puffed but not out- Details of sexual shoots of puffsheath dropseed. The small, contracted panicles of this annual species were still largely retained within the boot, the enveloping sheath surrounding the flower cluster. Thus, basis of the adjective of the common name. It was remarked above that this partial retention of the panicle is a characteristic of many of the Sporobolus species.

Tarleton State University Hunewell Ranch, Erath County, Texas. Late October, fruit-ripe phenological stage.

 

152. Heavy infestation of common grassbur or sandburr- Cenchrus incertus is one of the most abundant grasses in Texas and south into Mexico. It is a weed of overgrazed pastures and ranges, suburban yards, and abandoned city lots (in short, on any disturbed soil surface). It is usually a short-lived perennial or, sometimes, an annual. If ever there was a meek to inherit the earth of Texas it would be this spiny weed, though for that reason some would make the case it most certainly is not meek. (In fact, it is a pretty good match for the personality of many Texans who fashion themselves as half-hoss and half-gater just not quite wild enough to eat grassburrs.)

Erath County, Texas. July.

 

153. Involucres on grass or sand burr- The sharp-spined spikelets on this "black sheep" of the prairies are a real pain in the you-know-what (and about everywhere else). Grass burrs are a textbook example of zoochory, dispersal of plant disseminules by animals, including rangemen.

 

154. Another native, cool-season, prairie grass- Winter bentgrass (Agrostis hyemalis) on moist to wet local habitat on tallgrass prairie in western Ozark Plateau. This is a minor species other than in local mesic microsites. It was included here to give an idea as to the great species diversity of tallgrass prairie. Even in southern areas native, winter, grasses grow alongside warm-season, perennial, panicoid grasses. Winter bentgrass has generally been regarded as a short-lived perennial (Gould, 1975, p. 140). It grows on both interior and coastal tallgrass prairie.

Ottawa County, Oklahoma. Late May, maturing grain to early grain-ripe phenological stages.

 

155. Dainty panicles on tallgrass prairie- Panicles and spikelets of winter bentgrass. These flowers were on some of the same plants shown in the immediately preceding slide. Ottawa County, Oklahoma. Late May, early grain-ripe stage.

 

156. White tridens (Tridens albescens)- White tridens is another cool-season native grass common on prairies and prairie-forest edges, especially in local moist habitats such as depressions and edges of imponded water. Unlike the preceding wildryes and Junegrass that usually respond as decreasers, white tridens is typically an increaser in response to disturbances like overgrazing. This species is particularily common and conspicuous on mesic Texas grasslands including the Grand, Blackland, Fayette, and North Central Prairies as well as adjoining savannahs like the Cross Timbers and Post Oak Savanna.

White tridens often forms large stands on depressions, along banks of ephemeral stream and ponds, and other places where soil moisture can accumulate or even inundate the land surface for brief periods. By nature of its preference for a wet rooting medium white triden often grows on clay or generally heavy textured soils. The stand of white tridens seen here was growing on a depression just upslope from a shallow stream in the Western Cross Timbers area, a made-to-order home for white tridens.

Lee Ray Ranch, Stephens County, Texas. May (and a wet one).

 

157. Customary stance- Typical habit of white tridens. This cespitose grass of intermediate size (a mid-grass species) usually has a sprawling or "spread-open" physical appearance as shoots come off from the basal part of the plant at angles less than vertical.

Lee Ray Ranch, Stephens County, Texas. May.

 

158. Panicles of white tridens- The "tight-packed" spikelets of numerous florets on short pedicels along a central floral axis make up a contracted panicle inflorescence type in white tridens. The panicles in this frame are just past anthesis with the tiny grains in the milk to soft dough stage.

Lee Ray Ranch, Stephens County, Texas. May.

 

159. Tinge of purple in white tridens- Viewed at close distance the coloration pattern of white tridens is quite pleasing with traces of dark blue, purple, or bluish red splotched across the lemmas.Some of the other Tridens species have similar colors (see immediately below). Hitchcock and Chase (1950, p.216) described white triden spikelets as having eight to twelve florets whereas Gould (1975, p. 209) put the number of flowers at four to eleven.

Lee Ray Ranch, Stephens County, Texas. May.

 

160. A second tridens top- Spikelets on contracted panicles of rough tridens (Tridens elongatus= T. muticus var. elongaatus). This and the succeeding specimen have been interpreted as two species and also as two taxonomic varieties of the same species, Tridens muticus. Mills County, Texas. October: hard-dough stage to approaching grain-ripe stage.

 

161. A third tridens top- Spikelets on contracted panicles of slim tridens (Tridens muticus= T. muticus var. muticus). Mills County, Texas. This and the preceding specimen have been interpreted as two species and also as two taxonomic varieties of the same species, Tridens muticus. October: hard-dough stage to approaching grain-ripe stage.

Set for another tridens- Example plants of longspike or, if preferred, long-spike tridens (Tridens strictus) that grew on a deteriorated (overmowed) big bluestem (the potential climax dominant)-dominated tallgrass prairie meadow in the Ozark Highlands immediately to the east of the Cherokee Prairie. Longspike tridens is the second-most abundant tridens (a distant second to purpletop) over most of the tallgrass prairie. In less mesic parts of the tallgrass country and in much of the Cross Timbers, especially in Texas, longspike tridens is uncommon. Longspike tridens is mostly an increaser to, even, an invader (depending on range site) on tallgrass prairie dominated by big bluestem, Indiangrass, and/or switchgrass. Longspike tridens is not the invader to the same degree as that of, say, broomsedge bluestem and the threeawns (Aristida spp.), but it appears to be an increaser to a lesser degree than is purpletop.

Longspike tridens grows on a variety of environments in the heart of tallgrass prairie. Habitats tha this species thrives on range from wet prairie such as local potholes to rock outcrops that funnel runoff into deep, between-rocks crevices. For whatever reason(s) longspike tridens has received "less press" given its prominence and local abundance than just about any grass species of the tallgrass association. (Put this author in mind of several rangemen he has known.)

These dead shoots were of plants that had gone into dormancy about six to eight weeks ago. Most of their caryopses had not fallen from their condensed panicles. Longspike tridens is a bunchgrass (cespitose habit) with large, fairly thick shoots (culms).

Ottawa County, Oklahoma. Late December; dormancy.

 

162. Cycle all over again- Several plants of longspike tridens at peak (or near it) development of herbage and sexual organs in the summer that followed the winter in which the above photographs of longspike tridens were taken.. These specimens were growing on the edge of a wet prairie in the western Ozark Plateau in an Extreme Drought (Palmer Index).

Plants had made this growth during the worst single summer drought in 60 years and it was the second consecutive summer of Extreme Drought. Obviously these individuals had used effeciently what little rain had fallen, but it was also obvious that edaphic moisture conditions of this wet prairie habitat were favorable to this species. It was also apparent that the very limited precipitation had come at time periods and stages of plant growth that were favorable for longspike tridens.

Ottawa County, Oklahoma. Late July; height of anthesis.

 

163. A species summarized- Summary photographs of two plants (and outer portions of some others) of longspike tridens at peak plant biomass and phenological development of shoots. These specimens were growing on the edge of wet prairie in the western part of the Springfield Plateau. Associated plant species included Florida paspalum, switchgrass, and several species of Carex and Juncus, most of which were included in this chapter (see below).

Purple top (Tridens flavus) is a far more common Tridens species over most of the tallgrass prairie, but purpletop does best on drier (less mesic) environments than does longspike tridens. It appeared to this author that longspike tridens was an increaser in its response to disturbances and as an indicator of successional stage. On heavily pastures adjacent to the ungrazed wet prairie on which these specimens grew, longspike tridens was grazed quite closely by beef cattle except where it was protected by larger (and "well-armed") woody plants.

Ottawa County, Oklahoma. Late July; height of anthesis.

 

164. Upper part- Upper portion of sexual shoot with lower panicle and flag leaf (first slide) and closer-in view of the contracted panicle (second slide) of longspike tridens. The common name notwithstanding, the flower cluster of longspike tridens is a contracted panicle not a spike. The short branches of this compound inflorescence were visible in these two slides as well as in the immediately succeeding set of three slides.

Ottawa County, Oklahoma. Early July; pre-anthesis phenological stage.

 

165. Contracted details- Three close-up views of the contracted panicle at peak anthesis of longspike tridens growing on wet prairie in the western Ozark (Springfield) Plateau. Short branches coming off of the rachis of panicles were readily shown in these three slides as well as the two slides of the immediately preceding slide-caption set. Detailed shot of the axil of the flag leaf (leaf subtending--immediately below--the inflorescence) was shown in the first of these three slides. The second slide, taken at one-half life size (1:.2 scale), gave an idea as to how many caryopses are produced by longspike tridens. There are four to six florets crowded ("closely imbridate") per spikelet in longspike tridens (Hitchcock and Chase, 1950, p. 215). Longspike tridens is a prolific sexual reproducer as well as a species that excels at tiller production.

The third photograph in this set presented the uppermost portion of the contracted panicle of longspike tridens with a goldenrod crab spider (Misumena vatia) on the prowl at the pinnacle of its home. Morse (2007) wrote an ecological monograph of M. vatia which is one more than has been devoted to T. strictus, a species for which there remains remarkably little knowledge.

One general (and rather short) reference for this species was that prepared by Shadow (not dated). This author concluded that longspike tridens was "very drought tolerant". This was by standards of east Texas, but that conclusion was consistent with observations by the present author in eastern Oklahoma. Likewise, Shadow (not dated) stated that longspike tridens was moderately palatable and indicative of Fair range condition. These apparent features were consistent with observations made by Rosiere who suspected that longspike tridens was, as noted herein (see above captions), an increaser species and one adapted to marginal and disturbed , though usually moist, microsites.

Ottawa County, Oklahoma. Late July; peak anthesis stage of phenology.

 

166. Drawn out and condensed- Two contracted panicles of long-spike tridens on degraded tallgrass prairie in the western Ozark (Springfield) Plateau. The spikelets in these panicles were ripe but still retained their caryopses (ie. this was late seed-ripe stage just prior to grain-shatter stage).

Ottawa County, Oklahoma. Late October.

 

167. Long spikes (well, kind of) on a condensed version- Spikelets on contracted panicles of long-spike tridens.Ottawa County, Oklahoma. Late October; grain-ripe phenological stage (just before fruit-release stage).

 

168. Tumblegrass (Schedonnardus paniculatus)- This distinctive grass is a native perennial that is regarded as an invader or, depending on range site, an increaser at best. It is a characteristic member of early seral stage communities and thus is more common on deteriorated ranges, sacrifice areas, etc. It is one of the earlier perennials to become established on go-back land (ie. a component of seral stages that are slightly more advanced than the pioneer stage). Consistent with early seral species, tumblegrass has poor forage value. Texas A&M University Agricultural Experiment Station (Stephenville branch station), Erath County, Texas.April.

 

169. Inflorescences of tumblegrass- The inflorescence of tumblegrass is a raceme which is characteristic of the Chlorideae tribe of which this weedy grass is a member. This "seedhead" is quite distinctive. It is of a relatively large size and frequently breaks off from the basal part of the plant and rolls along the ground surface much like a tumbleweed (hence it's common name). Erath County, Texas. June.

 

170. Local stand of tumble windmillgrass (Chloris verticillata)- This dense population of tumble windmillgrass with a typical pattern of inflorescences was growing on a disturbed site-- the most common category of habitat for the Chloris species-- in the Cross Timbers of northcentral Texas. Chloris species are not major forage producers nor are they usually climax species. Some Chloris species are interpreted as ecological invaders while others are classified as increasers. Erath County, Texas. July.

 

171. Inflorescences of tumble windmillgrass- The inflorescences of the Chloris species has been a subject of semantical debate among agrostologists. Traditionally the inflorescence type of Chloris (and some related genera) was called a raceme (an unbranched inflorescence with spikelets pedicellate-- pediceled-- on the rachis) as typical for members of the Chlorideae tribe (eg. Bouteloua, Buchloe, Spartina). In this view the individual racemes are each separate inflorescences. Alternatively the individual racemes can be viewed as part of a general compound raceme or, as another alternative, as a number of one-sided spikes that are either solidary, digitate, or racemose on a main axis. More recently, some workers came to interprete the entire inflorescence as a panicle of of verticillate (arranged in whorls) spicate (ie. spikelike) primary unilateral (one-sided) branches. A nice debate about a genus of relatively little consequence relative to so many others.

In Cross Timbers vegetation, Erath County, Texas. July.

 

172. Perennial threeawn (Aristida purpurea) on deteriorated tallgrass prairie range - What is most likely "purple threeawn" was thriving and adding spring color to an area heavily impacted by motor vehicle traffic.This was typical "prime habitat" for threeawns which are some of the most abundant perennial grasses on the most abused areas (eg. sacrifice areas, "stomp lots", ranch roads). The other most common species on this deteriorated site was the naturalized alien, rescue bromegrass. The threeawns or "wiregrasses", as they referred to in the southeastern pine forests, are typically classified successionally as invaders. As a general rule Aristida species are characteristic of vegetation at some stage of retrogression and therefore indicator plants of deteriorated ranges (or regenerating pine forests in less advanced seral stages). One of the general features of species responding as ecological invaders is low feed value. This is most pronounced in annual species, but some perennial species (in fact, entire genera) have this feature. Aristida is one of these (with some rare exceptions). In fact, there is a high proportion of Aristida species that are annuals, even short-lived annuals known as ephemerals.

As if all this were not enough, Aristida includes several taxa that are a taxonomist's worst nightmare. Some of these integrade or hybridize (or whatever grass plants do "to make life harder for agrostologists") with the result that various authorities have interpreted such taxa as species, subspecies, and varieties in different ways which has led to confusion and frustration (especially because the taxonomic arrangements seem to change with every new treatment of Aristida). The term that has often been invoked in describing and arranging these taxa into groups is "complex". An example of such includes such closely related and intergrading threeawns as red threeawn (A. longiseta), purple threeawn (A. purpurea), wright threeawn (A. wrightii), roemer threeawn (A. roemeriana), and a few others depending on which of these species grow in association in any given geographic location. In Texas the complex is often known simply as "perennial threeawn" or, sometimes, "purple threeawn" after A. purpurea which is viewed as the species, with what have previously been interpreted as separate species ranked as varieties therein. Thus "perennial or purple threeawn" includes A. purpurea var. longiseta, A. purpurea var. purpurea, and A. purpurea var. wrightii. Other agrostologists still designate these threeawns known by the various common names as separate species. There was bound to be some sort of lesson in this example of "where the experts cannot agree".

 

173. Examples of perennial threeawn- These photographs displayed some individual plants that were typical of "perennial threeawn" (A. purpurea complex) in the Cross Timbers and Prairies vegetational area of northcentral Texas. Aristida species are strictly cespitose (= bunchgrass or tufted) species that produce only intravginated shoots (tillers) and no extravaginated shoots (stolons and rhizomes). There are some species in Sporobolus, Muhlenbergia, Festuca, and Andropogon (among others) that also have the cespitose habit, but in North America Aristida is probably the most extreme case of this. The tufted habit was very obvious in these examples.

 

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174. Old-field threeawn (Aristida oligantha) on an old-field- Dog-hair stand of the annual old-field threeawn or annual wiregrass on small parcel of go-ck land of Fort Worth or Grand Prairie in noethcentral Texas. Most stands of this r-selected or pioneer species are not this dense, but this demonstrated the physiogonomy and structure of such populations of this species that was performing the essential function of protecting soil made bare by the Clementsian process of denudation (in this case, plowing).

Tarleton State University Hunewell Ranch, Erath County, Texas. October.

 

175. Plants of old-field threeawn- Numerous individual plants (genotypically unique individuals) of the annual Aristida oligantha. Zoom-in view on some of the plants presented in the immediately preceding slide to show habit of this pioneer species. While old-field threeawn is an ecological invader it is valuabel-- often locally essential to prevent soil erosion and facilitate modifications in its habitat that prepare the way for species of higher successional order. This improvement in the environment (often edaphic modifictions like addition of organic matter with this species) is the Clementsian process of reaction or, in contemporaty terminology (and same thing), facilitation.

Tarleton State University Hunewell Ranch, Erath County, Texas. October.

The dominant plant species on tallgrass (and true) prairie are, obviously, native perennials. The perennial life cycle is even more common among grass than forb species on the grasslands, especially mesic ones or those in humid and subhumid zones, of North America. There are, however, native grass species that are annuals. These species almost invariable have as their habitats and ecological niches disturbed environments such as those created by overgrazing, tillage, mining activity, traffic lanes, and out-of-season or unusually intense fires, etc. Nonetheless, these annual grasses of primarily disturbance microsites are a natural part of tallgrass prairie vegetation. Annual grasses, especially native annuals, provide essential roles (functions) as pioneers in facilitating plant--usually secondary--succession.

Some of the more common and widespread native annual grass species of tallgrass (and true) prairie(s) were included below. Students of Range Manageament should fully understand the importance of native annual grasses on grasslands.

(Incidentially, none of this is meant to imply that these native annual grasses are never weeds because there are situtions where they clearly are weeds. Nor should it be assumed that only native annual grasses can perform essential roles in facilitation of plant succession when, in truth, naturalized exotic annual grasses frequently perform the same range ecosystem goods and services.) Students of Range Management must always have open minds and be ever ready to learn new things about our remarkable grazing lands.

 

176. Old-field threeawn (A. oligantha)- There are many annual Aristida species. Some of these are, as noted above, even epemeral (ie. the so-called six-week grasses"). Other annual species take somewhat longer to complete their annual life cycle, but are nonetheless very opportunistic. Old-field threeawn was appropriately named for its common occurrence as a pioneer species on abandoned farm fields ("old-fields"). In fact, it is probably the most common native annual grass colonizing disturbed areas like old-fields in the the central prairies prairies region (Central Lowlands and eastern Great Plains). Old-field threeawn is an example of an r-selected species (those in which natural selection has been for traits which maximize the intrinsic rate of increase such as to allow members to rapidly colonize newly or recently created habitats) covered in standard Ecology texts. For practical purposes the annual threeawns have no forage value for livestock or big game species.

The specimen, the sporophytic generation, shown here had just completed its short life cycle tht streatched from early spring or late winter to early summer. Erath County, Texas. July.

 

177. Annuals on an old field- Old-field threeawn (Aristida oligantha) growing on an old crop field in the West Cross Timbers of northcentral Texas. This land had been farmed roughly 15 years ago and was not reseeded but simply left alone to "heal itself". This "go-back land" had local populations of this annual threeawn, which is the classic, the single most common or abundant annual native grass on such old fields. Old-field threeawn is the textbook native pioneer grass on denuded grasssland, forest, and savannah over eastern North America including the vast Eastern Deciduous Forest, Prairie Peninsula, and Tallgrass Prairie Regions.

A. oligantha is even more widespread than Sporobolus neglectus, another annual grass of disturbed areas, in the first seral stage of secondary plant succession on such range vegetation, both forest and grassland. Old-field threeawn is Mother Nature's advance guard, her spermatophytic "first responder" to bring successional first aid to heal the land--to restore vegetation lost by some factor(s) of denudation. Old-field is the major grass species making up the botanical scab on such "wounded land".

Yes, indeed old-field threeawn is an ecological invader, one of the lowest of the seral plant species, and as such it is just as valuable as the climax (the decreaser species) as tallgrasses like big bluestem and Indiangrass, palatable forbs including those among composites and legumes, and defining tree species of the savanna including the oaks and hickories.

Tarleton State University Hunewell Ranch, Erath County, Texas. Top (first) slide: late October, fruit (achene)-ripe phenological stage. Second, third, and fourth slides: mid-September fruit (achene)-ripe stage of phenology.

 

178. Tops of an annual- Middle to upper parts of shoots of old-field threeawn growing on an abandoned crop field in the West Cross Timbers of northcentral Texas. The threeawned spikelets of this invading and seral (successional) revegetating species were visible against a backdrop of shoots of the rest of this annual grass population.

Tarleton State University Hunewell Ranch, Erath County, Texas. First (top) slide: late October, fruit (achene)-ripe phenological stage. Second and third slides: mid-September, fruit (achene)-ripe phenological stage.

 

179. Little 'un on the prairies, plains, and Cross Timbers (and a lot of other range to boot)- Single plant (first slide) and two plants (second slide) of little barley (Hordeum pusillum), a native, cool-season annual grass locally common on tallgrass prairie and tallgrass savannah. This species is found frequently on the prairies of northcentral Texas and adjoining Cross Timbers (where these plants were photographed) and extending westward throughout the Great Plains clear across to British Columbia and northeastward to Maine and down south to Florida.

Given its tremendous species range, nativeness, annual life cycle, and local commonness little barley was included on the list of 200 plant species for the Society for Range Management (SRM) International Range Plant Identification Contest (Stubbendieck et al., 1992). Little barley is also included in the Texas Intercollegiate Range Plant Identifiction Contest sponsored by the Texas Section, SRM as well as on both the range plant list for Texas 4H and Future Farmers of America Contests (Hatch and Pluhar, 1993).

Little barley is classified as an invader what with being an annual--even if a native one--on range where almost every one of the dominant plant species is a perennial. Little barley typically invades disturbed areas such as overgrazed ranges and "go-back land" where it undoubtedly provides some soil protection against the agents of soil erosion. A concise yet informative description of little barley, including practical aspects, was in both editions of Pasture & Range Plants (Phillips Petroleum Company, 1963; Nicholson, 2006), although the author of Range Types disagreed that little barley is "essentially worthless" for livestock and wildlife. In point of fact, this short-statured species sometimes provides forage for cattle, sheep, horses, and deer. On dormant tallgrass prairie where the herbage of bluestems, Indiangrass, sideoats grama, etc. is weathered after three months of dormancy this C3, festucoid grass frequently provides some green feed--the quantity varying substantially--thart is richer in carotene, higher in crude protein, and better in flavor than that in the herbage of the decreasers and increasers. This could be especially to the smaller ruminant species.

Erath County, Texas. Mid-May (first slide); anthesis and March (second and third slides); just past anthesis to early milk stage. (Note variation from plant-to-plant as well as year-to-year (first slide of separate plant taken in a different year from the plant and year of second and third slides).

 

180. Basal properties- Shoot bases of little barley at phenology ranging from anthesis to milk stage. Erath County, Texas. March.

 

181. Barley spiked- Spike of little barley (H. pusillum) growing in West Cross Timbers. Spikelets of little barley are one-flowered (one floret per spikelet). Erath County, Texas. March; milk stage of phenololgy.

 

182. Next little example- Shoots and spike of little barley growing on a locally disturbed microsite in the West Cross Timbers. These photographs presented examples of the flag leaf (the leaf immediately subtending the inflorescence in grasses) and spikes in anthesis.

Hunewell Ranch, Tarleton State University, Erath County, Texas. Mid-April; anthesis.

 

183.. Another native annual- Part of a local stand of sixweeks fescue (Festuca octoflora= Vulpia octoflora) growing aroung the entrance to a red harvester ant (Pogonomyrmex barbatus) colony or nest. This was an example of the typical habit of this annual festucoid (subfamily, Festucoideae; tribe, Festuceae) grass which, as implied by the common name, completes its life cycle in a single growing season (or even faster). The adjective "sixweeks" has been widely applied to a number osf "sixweeks grasses" including sixweeks fescue. While these are annual species, often the more apt descriptor is ephemeral, meaning to complete the life cycle of emergence from seed to production of seed in an unusually short period of time (as in one or two months; hence, "sixweeeks"). Species of "sixweeks grasses" like sixweeks fescue can produce more than one generation in a year.

Sixweeks fescue is a native annual (ephemeral) as is little barley and prairie or old-field threeawn which were described immediately above.

Hunewell Ranch, Tarleton State University, Erath County, Texas. Early April; peak shoot development, caryopses mostly at soft-dough stage.

 

184. Yep, its an annual- An entire plant--most of it anyway--of sixweeks fescue that was growing by a red harvester ant mound and then pulled up and laid across the bare area to show the entire body of this ephemeral (see caption immediately above) species. While most of the length (though not necessarily biomass) of its fibrous root system remained in the soil this plant was shown in most of its entirety, at least enough that viewers could see relaive proportions of an adult plant of an annual grass.

The "bent" (decumbent) culm, a common feature in some individuals of sixweeks fescue (Gould, 1975, p.96), was shown in this specimen.

Hunewell Ranch, Tarleton State University, Erath County, Texas. Early April; peak shoot development, caryopses mostly at soft-dough stage.

 

185. Panicles in six weeks- Sevral panicles of sixweeks fescue at peak development with spikelets clearly visible. Gould (1975, p.96) stated that spikelets of sixweeks fescue had from five to seventeen florets.

Hunewell Ranch, Tarleton State University, Erath County, Texas. Early April; peak shoot development, caryopses mostly at soft-dough stage

Technical note: Festuca octoflora has gone by various epecific epithets over the years as shown in numerous synonymies (see for eg. Hitchcock and Chase, [1950, ps.875-876]). Gould (1975, ps. 95-99) elevated to genus rank the Vulpia section of Festuca genus used by Hitchcock and Chase (1950, p. 58) for annuals (vs. section Eufestuca for perennials). Gleason and Cronquist (1991, ps. 748, 750) explained that Vulpina species are "only marginally distinct" from Festuca but that Vulpia is generic rank for mostly annual species that are cleistogamous and usually with only one or two stamens per floret whereas Festuca species aare open-pollinated and have three stamens per flower.

A section on naturalized, Eurasian annual bromes- Several (exact number varying with location) annual Bromus species have naturalized and become locally abundant on disturbed sites and microsites across the Tallgrass Prairie Region. Abundance of and, varying from year-to-year, mere presence or absence of these species differs considerably. The section immediately below dealt with four annual Bromus species that periodically grew in an area of northeast Oklahoma within the Prairie Peninsula in the western portion (Springfield Plateau) of the Ozark Uplift.

Photographs were all taken in late spring of one year (one cool-growing season) in which, for whatever reasons, plants of the four naturalized, annual bromes grew in close proximity (often side-by-side and within inches of each other). This was the only year in the photographer's memory that this phenomenon took place. It provided an unusually opportune moment in ecological time to present these weedy or ruderal Eurasian annuals growing on grassland for which the potential natural vegetation is a big bluestem-dominated prairie with local stands of oak-hickory forest.

 

186.Three of a genus- Plants with abundant panicles and spikelets of three species of naturalized annual bromes growing on a disturbed bar-ditch in the western Springfield Plateau in northeastern Oklahoma. These three species were: 1) Japanese chess or Japanese brome (Bromus japonicus), 2) cheatgrass or downy brome (B. tectorum), and 3) poverty brome (B. sterilis). These three Bromus species can be distinguished by the following colors for the respectively listed speccies in the previous senteence: 1) green, 2) faded or light tan, and 3) purple.

Presence of these three species in close quarters side-by-side spacing was a remarkable phenomenon. This photographer had not ever witnessed this (before or after) this unique occurrence.

Ottawa County, Oklahoma. Late May; hard-dough stage (japanese bome), ripe-grain to grain shatter stage (downy and poverty brome).

 

187. Three of a genus side-by-side- Panicles of ripe, shedding spikelets of three naturalized, Eurasian, annual bromes: 1) Japanese brome, 2) chetgrass or downy brome, and 3) poverty brome. All plants were growing (had grown and matured) in situ (ie. grew in exact location as shown). Habitat was basically a pile of chert with shallow soil in a county road barrow ditch.

This location was in part of the Prairie Peninsula in wesstern Springfield Plateau of extreme northeastern Oklahoma.

Ottawa County, Oklahoma. Early June; grain-ripe, shattering phenological stage.

 

188. Three of a genus at another place- Plants of japanese brome (largest, green plant; center foreground), cheatgrass or downy brome (reddish plant; far right foreground), and poverty brome (purple plant; center and right background) growing on a "rockpile" (chert rocks and shallow soil) habitat of a barrow ditch in the western Springfield Plateau. This was once part of the Prairie Peninsula with tallgrass prairie and stands of oak-hickory forest growing in a patchwork pattern.

All plants photographed where they grew naturally (ie. all plants still rooted in situ).

Ottawa County, Oklahoma. Early June; hard-dough (Japanese brome) to floret/grain shatter (cheatgrass or downy brome and poverty brome) stages of phenology.

 

189. Tree busom-buddy bromes- In situ (growing naturally as shown; not transplanted) plants of three Eurasian (and naturalized) species of annual bromes or bromegrasses growing on disturbed tallgrass prairie in the Spingfield Plateau portion of the Ozark Highlands (part of prairie Peninsula). Plants of the three species were: 1) downy brome or cheatgrass (one plant at far left), 2) poverty brome (center and tallest plant) and 3) Japanese brome (plants to immediate left and right of poverty brome).

These plants were growing on relatively deep, fertile soil.

Ottawa County, Oklahoma. Early June; soft- to hard-dough stage of phenology.

 

190. Three kinds of annual brome- Downy brome (plants with large, drooping panicles; center foreground), Japanese brome (plants with more upright panicles to left and right of center downy brome), and rye brome (Bromus secalinus) seen as tallest plants both light green (left-center midground) and tan-colored (right midground) in the first slide.

In the second slide, panicles of these three species were presented as Japanese brome (green spikelets at left to center), rye brome (tight panicle with tan spikelets in center foreground), and downy brome or cheatgrass (loose, fan-shaped panicle with tan spikelets in center foreground).

Ottawa County, Oklahoma. Late May; hard-dough (Japanese brome) to grain-shatter (downy brome and rye brome) stages of phenology.

 

191. Brome bouquet; four kinds of annual brome- Panicles and spikelets of 1) Japanese brome, 2) downy brome or cheatgrass, 3) poverty brome, and 4) rye brome all growing side-by side on a disturbed upper part of a barrow ditch in far-western Springfield Plateau of the Ozark Uplift in what was, before the whiteman's legacy, part of the Prairie Peninula.

These plants were growing and still rooted in situ. They were not transplanted; however the photographer did gather panicles together for purposes of photographing this Bromus "bouquet".

Some rare combination of growing conditions known only by God was responsible for this essentially "once in a lifetime" phenomenon of four species of naturalized, annual, Eurasian bromegrasses to grow and mature (complete their life cycles) in immediately proximity to one another. Your author grew up in this area and continued to visit it for decades. This was the only time he witnessed this array of weedy, annual Bromus species. Unique teaching opportunity.

Ottawa County, Oklahoma. Late May; hard-dough to grain-shatter stages of phenology.

 

192. Two panicles: one each of two annual Bromus species- One panicle with spikelets (still green-colored ) of Japanese chess or Japanese brome and one panicle with spikelets (purplish-colored) of poverty brome. Disturbed roadside in Prairie Peninsula of western Ozark Uplift (Springfield Plateau), extreme northeastern Oklahoma.

Ottawa County, Oklahoma. Late May; ripe, shedding spikelets in sterile brome, hard-dough stage of spikelets in Japanese brome.

 

193. Big 'uns of the two main species- One plant of cheatgrass or downy brome (left foreground) and one plant of Japanese brome or Japanese chess (right midground) growing on disturbed roadside in western Springfield Plateau in northeastern Oklahoma. This area was originally (prior to European settlement) part of the tallgrass prairie/oak-hickory forest ecotone known as Prairie Peninsula.

The plant of cheatgrass was 31 inches tall with a 24 inch panicle, one of the largest specimens (among tens of thousands of individual plants) observed by this author. The plant of Japanese brome was about 22 to 25 inches tall. It, too, was one of the largest ever encountered by this photographer. Speaking of thousands, there had to be several hundred up to maybe a thousand florets (grains each with its enclosed seed and attendant lemma and palea or chaff) in both of these plants. Safe to say that the soil seed bank of these annual bromes was well-stocked in this year.

Ottawa County, Oklahoma. Early June.

 

194. No cheating these tops- Sexual shoots of cheatgrass or downy brome topped with roughly 18-inch panicles produced on a disturbed roadside in part of the original Prairie Peninsula that developed in the western Springfield Plateau of extreme northeastern Oklahoma.

The next two-slide/caption set presented closer-in views of some of the spikelets produced on these panicles ...

Ottawa County, Oklahoma. Late May.

 

195. Not cheated cheated tops- Spikelets on panicles of cheatgrass or downy brome produced in the western part (Springfield Plateau) of the Ozark Highlands. First slide presented the upper main branch of a panicle while the second slide featured spikelets being shed from part of the panicle.

These plant parts were on one of the panicles introduced in the immediately preceding slide. They were produced on a disturbed roadside in a remarkably favorable growing season.

Ottawa County, Oklahoma. Late May.

196 Perfect view not, but ...- Portions of three shoots pof cheatgrass or downy brome showing the "swollen" node characteristic of this species (at least of ecotypes in this area of tallgrass prairie).

This quite clear, well-composed shot was marred by over-exposure while being scanned by an Epson Perfection 700 scanner. (This author advises against Epson products, but this is about the only game in town--at least realistically-- thus leaving photographers with little choice.)

Ottawa County, Oklahoma. Late June.

 

197. From Japan- Local stand (first or upper slide) and one unusually large plant (second or lower slide) of Japanese chess or Japanese brome (Bromus japonicus) growing on a locally disturbed habitat in the western Springfield Plateau in extreme northeastern Oklahoma.

Almost all of these plants were from about 18 to over 25 inches tall having grown in an extremely favorable (high soil moisture and moderate temperatures) winter-spring season. The individual plant seen in the second slide was "a good two foot tall". Whether this size and stage of development was due to genetics (a genotype of large growth potential) and/or remarkable growing conditions could not be determined from this simple field observation. One thing for sure: it (and a "gazillion"others) produced ample viable seed to insure crops of Japanese chess for decades to come. (Not that this was needed given that every year ample grain was produced by many plants of this naturalized Eurasian annual).

In "good years" (favorable conditions for the cool-season of growth) plants of the various anual brome species produce more tillers (vertical, intravaginated shoots) that in turn porduce larger, higher grain-yielding panicles. This phenomenon was illustrated by samples in the next two-slide unit.

Ottawa County, Oklahoma. Early June.

 

198. Japanese spikelets- Panicle with numerous spikelets (first slide) and details of spikelets (second slide) of Japanese brome or Japanese chess. These plant parts were produced under cool-growing season conditions that resulted in amazingly high yields of grain of this weedy, naturalized, Eurasian annual.

Some of these individual spikelets six to eight bore florets, and, in numerous plants, there were a few hundred spikelets per panicle with from four to more than dozen tillers per plant.

Ottawa County, Oklahoma. Early June.

Lesson note: students should have obsrved that their teacher repeatedly used the proper term of grain (= caryopsis or the dry fruit type of most grass species) when referring to the annual Bromus species discussed herein. Some students, especially farm-raised ones familar with wheat, corn, sorghum, etc., may find this to be somewhat misleading. It need not be when one thinks of the grain-filled panicles of oats and the spikes bearing kernals of wheat, rye, and barley.

In this regard, it is common practice for stockmen, especially sheepmen, to collect the spikelets/florets of Japanese brome and related annual weedy grasses at grain elevators where this is one type or component of what is known as elevator screenings, including wheat screenings. In the cleaning (= screening) of small grains such as wheat and rye at grain elevators this trash accumulates. Screenings are sometimes sold at low prices or, frequently, given away to get rid of the unwanted residue or trash.

Such spikelets/florets of annual weedy grasses (Japanese brome is the big one in the Wheat Belt) are a by-product feed. This by-product feedstuff is a combination of grain and attendant chaff. This is not the chaff of wheat or rye (both naked grains or naked caryopses), which is separated out at combining, but the lemma and palea of the covered or husked grain of Japanese brome. There is a high-fiber content in the by-product grain of elevator screenings much like that of the covered grain of oats or barley.

In essence, grain (= caryopsis) is the proper agricultural term for the fruit of the annual Bromus species.

 

199. Japanese fuzz- Lower shoots of one plant (first slide) and two phytomers of Japanese chess or Japanese brome. The velvet-like pubescence was obvious on the leaf sheaths in the second slide. Phytomer is the repeating unit of the shoot including the upper portion of the lower node, the internode, and the lower part of the upper node. There are numerous phytomers in a tpical grass shoot.

There were nine or ten shoots (tillers: vertical, intravaginated shoots) of the single plant presented in the first slide.

Ottawa County, Oklahoma. Early June.

 

200. One chess- Single plant of rye brome (Bromus secalinus) that grew on a disturbed local area in the Springfield Plateau portion of the Ozark Highlands.

Ottawa County, Oklahoma. Early June; peak standing crop, grain-ripe phenological stage.

 

201. Rye-like up close- Phytomers (repeating node-internode units) on basal shoots, which were tillers, (first slide) and the fairly compact panicle with prominently displayed spikelets (second slide) of rye brome. These organs had developed on overmowed, degraded tallgrass prairie in the western Springfield Plateau. Land on which this rye brome grew had been heavily defoliated with rotary lawnmowers for over 60 years up until three years before these plants had grown and completed their life cycle. The current year's crop of rye brome had grown in an exceptionally favorable cool-growing season in which Japanese brome, downy brome, and poverty brome also produced super-abundant yields of grain.

Big bluestem was the dominant grass in neighboring (adjoining) fencerows that had not been subjected to prolonged heavy shredding.

Ottawa County, Oklahoma. Early June, first slide; late May, second slide.

Cool-Season versus Warm-Season Species: All North American tallgrass prairie has both cool-season and warm-season species among grasses, grsslike plants, and forbs. This is most pronounced among the grasses, if for no other reason, simply because grasses are the dominant and definng group (family in this instance) of range plants on grassland. An example of cool-season and warm-season grasses followed.

202."Some like it hot; some like cold"- Canada wildrye (Elymus canadensis), a climax cool-season grass, on the left and sideoats grama (Bouteloua curtipendula), a climax warm-season grass, on the right at winter solstice (first day of winter) on tallgrass prairie that was an ecotone between a chert glade and an oak-hickory forest in the western Ozark (Springfiield) Plateau. The same two plants were in both slides only taken from two different direction (two directional orientations). Neither plant had been grazed and instead had remaining herbage of grass shoots from peak standing crop of the previous growing season still present. Current cool-season growth of Canada wildrye was growing on only one side of last year's standing dead shoots and therefore was visible only in the second (lower) slide. Sideoats grama, the warm-season grass representative, had no new growth.

Canada wildrye is in the fescue or bluegrass subfamily (Festucoideae or Pooideae) and sideoats grama is in the grama or dropseed subfamily (Eragrostoideae or Chlorideae) of the grass family (Gramineae or Poaceae). Both of these climax grass species were associates (to dominants, switchgrass and big bluestem) and, at that status, decreasers on this savanna range site.

Wildcat Chert Glades Natural Area, Newton County, Missouri. Winter solstice.

Grasslike Plants of Tallgrass Prairie

Tallgrass prairie includes or, at least, is adjacent to floristically associated wetlands, most of which are marshes. Most rangemen would probably interpret such marshes as part (a range site) of the overall grassland community, most logically as another range cover type within the general tallgrass prairie. In some locations these wetlands are formed from springs that drain directly into nearby creeks which are corridors of the Eastern Deciduous Forest, especially forms of the Oak-Hickory Association. In some such range vegetation, marshes abutt both tallgrass prairie bottomland and bottomland forest dominated by species like sycamore (Plantanus occidentalis), (Acer segundo), and ashes (Fraxinus spp.).

In these marsh habitats there are various species of bulrush or tules (Scirpus spp.). A few tules or bulrushes were included below. Taxonomy of these remians a "mess". For example, the most useful manual for the tallgrass prairie marshes of eastern Oklahoma and Kansas was the Missouri flora. Treatment of Scirpus spp. changed dramatically from the original Flora of Missouri (Steyermark, 1963) and Steyermark's Flora of Missouri (Yatskievych, 1999). This included changing some former Scirpus species to Schoenoplectus species which was consistent with some other treatments (eg. Diggs et al. (1999). This author did know if the dead Dr. Steyermark would have approved of these change, but the non-taxonomist author of this publication did not. Scientific names reflect that opinion.

Other range plants common in-- and, indeed, often comprising dominants and associates of-- marshes are various grasslike-plants especially species of. Carex, Cyperus (Cyperaceae), Juncus (Juncaceae), and Typha (Typhaceae). Several of these lesser-known, but often locally important grasslike-plant species were included below.

 

203. Soft rush (Juncus effusus)- This is a widely distributed rush in the tallgrass prairie region. It was found to be an important forage species to the American muskrat (Ondatra zibethicus). Craig County, Oklahoma. June.

 

204. Grasslike neighbors-Two species of grasslike plants on a wet tallgrass prairie in the western Ozark Plateau. Short-fruit or whiteroot rush (J. brachycarpus) and Frank's caric sedge (Carex franki) growing in association on a local ponded area of big bluestem-dominated prairie. The rush is in front of the sedge in both of these slides.

Ottawa County, Oklahoma. July; early fruit-ripening stage.

 

205. Short-fruit or whiteroot rush (J. brachycarpus)- On a local depression with periodically ponded water and slowly drained soil this rush was growing in association with beaked panicum, broomsedge bluestem, and several species of caric sedges including fox sedge (Carex vulpinoidea) which was presented below. Whiteroot rush is not usually a common species, but it is frrequently present as individual to small numbers of plants in particular on areas of wet prairie.

Palatability of whiteroot rush was unknown to this author, but it is a common observtion that generally speaking none of the Juncus species are very palatable. Economic value of Juncus species is undoubtedly in soil protection and improvement through addition of organic matter.

Ottawa County, Oklahoma. July; early fruit-ripening stage.

 

206. A distinctive topknot- The spherical inflorescence of whiteroot or short-fruit rush makes identification of this Juncus species a fairly straightforward endeavor. Ottawa County, Oklahoma. July; early fruit-ripening stage.

 

207. Inland or interior rush (Juncus interior)- These two specimens were growing on the same wet paririe (though a different local depression) as many of the other grasslike plants presented in this section. This degraded tallgrass wet prairie was in the western Ozark Plateau. The first specimen (first photograph) was in mid-flowering (just post-anthesis with some stigmas still alive). Second specimen (second photograph) was at fruit-ripe to fruit shatter stage. An interesting and, to a plant photographer, obvious variable about this species (at least at this location) was the variation in time of flowering and fruitset of different plants growing within a few feet of each other. Phenological stages were similar overall, however. The less mature plant (first slide) was photographed two weeks earlier than the more mature plant (second slide). Both plants fulfilled their role: they recombined genes, yielded fruit, and helped perpetuate their race.

These plants were growing in association with two-flower rush (Juncus biflorus), diffuse rush (Juncus diffusissimus), fox caric sedge (Carex vulpinoidea), Bicknell's caric sedge (C. bicknellii), Frank's caric sedge (Carex frankii), and Florida paspalum (Paspalum florida).

Ottawa County, Oklahoma. Mid-June and late June.

 

208. Inland inflorescences and fruit- Panicle and fruit of inland or interior rush. The fruit of Juncus is a locusidal capsule). These organs were on plants growing on a small derpression--nothing much but a mud hole--on wet prairie.

Ottawa County, Oklahoma. Late May, fruit-ripening (just pre-maturity) stage.

 

209. Ripe inland- Inflorescence with mature loculicidal capsules of inland rush. Ottawa County, Oklahoma. Late May, fruit-ripe phenological stage.

 

210. Two-flower (two-flowered) rush (Juncus biflorus)- Two-flower rush growing on a wet prairie in western Ozark Plateau. These shoots were growing in association with interior rush, diffuse rush (Juncus diffusissimus), fox caric sedge (Carex vulpinoidea), Bicknell's caric sedge (C. bicknellii), Frank's caric sedge (Carex frankii), redtop panicgrass, and Florida paspalum.

Ottawa County, Oklahoma. Late June, onset of fruit-ripe phenological stage.

 

211. Capsuled panicles- Panicles of two-flower rush with loculicidal capsules at beginning or onset of fruit ripening. First slide of these two slides presented . several panicles in their entirity. The second slide showed one panicle at close-enough distance tha individual capsules could be distinguished.

Ottawa County, Oklahoma. Late June.

 

212. Just strting to ripen- Loculicidal capsules of two-flower rush that were still in wet-dough fruit stage. Ottawa County, Oklahoma. Mid-June.

 

213. Bottom of a rush- Bases of shoots of two-flowered rush growing on a wet prairie in the western Ozark Plateau. Ottawa County, Oklahoma. Mid-June.

 

Slim and diffuse- Slim-pod or diffuse rush (Juncus diffusussimus) gowing on wet prairie in the western Springfield Plateau at mid-bloom stage and essentially peak standing crop. The immediately following two slide set shows slim-pod rush at a more advanced stage of maturity.

Ottawa County, Oklahoma. Late May.

214. Plenty of water and still slim- Slim-pod or diffuse rush (Juncus diffusissimus) growing on wet prairie in the western Ozark Plateau. A plant was shown at flowering stage and peak standing crop (first slide) and then (the same plant) at mature-fruit stage and onset of dormancy (second slide). This is one of the most unique--hence, one of the most readily identifiable--of the Juncus specieas it has a "widely spreading and much branched" inflorescence bearing comparatively long and slender capsules (Steyermark, 1963, p. 414).

"Phytoneighbors" of slim-pod rush on this marshy habitat included two-flower rush, inland or interior rush, fox caric sedge (Carex vulpinoidea), Bicknell's caric sedge (C. bicknellii), Frank's caric sedge (C. frankii), redtop panicgrass, and Florida paspalum.

Ottawa County, Oklahoma. Early June (first photograph); late June (second photograph).

 

215. Branches and fruits- Portions of the much-branched infloresence and tiny loculicical capsules of slim-pod or diffuse rush produced by the shoots introduced in the two immediately preceding slides.These organs were produced on a wet prairie at the western edge of the Ozark (Springfield) Plateau. Ottawa County, Oklahoma. Early June; immature-fruit stage of phenology.

 

216. Branches and fruits a little closer- Branches and loculicical capsules of slim-pod rush presented at closer camera distance to give a zooming-in sequence following preceding photographs. The tiny and thin capsules that are basis of the common name were visible features in both of these photographs.

Early June; immature-fruit stage of phenology.

 

217. Slim and ripe- A progressiveling zooming-in three-slide sequence of the tiny and narrow/slender loculicical capsules of slim-pod rush produced on a wet prairie in the western Springfield Plateau.

Early June;fruit-ripe stage of phenology.

The following Carex species were all growing on tallgrass prairie within a five-mile radius in the western portion (Springfield Plateau) of the Ozark Highlands or Ozark Uplift.

218. Fox caric sedge (Carex vulpinoidea)- Fox sedge is but one of numerous carices found in wet prairies and marshes of the central tallgrass prairie and prairie peninsula region in the interior of North America. These plants were growing in association with inland or interior rush, two-flower rush, diffuse rush, Bicknell's caric sedge (C. bicknellii), Frank's caric sedge (Carex frankii), redtop panicgrass, and Florida paspalum.

Ottawa County, Oklahoma. June.

 

219. The distinctive inflorescence of fox sedge- This is a common wetland species in the marshes and poorly drained grasslands of the Cherokee Prairie and Ozark Plateau. This one was on a wet prairie used as a hay meadow, but at the edge where it was too wet for haying equipment. Ottawa County, Oklahoma. June.

 

210. Details of fox caric sedge inflorescence- These two inflorescences of Carex vulpinoidea showed the namesake and unmistakenable taillike (and extremely long) bract at base of the flower cluster. Fruit (achenes) on these two sexual shoots were approaching maturity and the shatter stage. These shoots were on the same wet prairie hay meadow as those presented immediately above.

Ottawa County, Oklahoma. Mid-June.

 

211. Troublesome sedge (Carex molesta)- The carices are a troublesome genus with many "look-alikes" and positive identifiction problematic, often unless plants can be collected at various stages of phenological development. Troublesome sedge is one of numerous carices that is adapted to various habitats varying from nearly aquatic to dry prairies and open deciduous forests. This "nifty" local population was in the western Ozark mountains of the Prairie Peninsula on a tallgrass prairie located between oak-hickory forest vegeetation. Climax dominants were big bluestem and Indiangrass, but the varied immediate range plant plant community also included the invaders broomsedge and Baldwin ironweed (Vernonic baldwini).

Ottawa County, Oklahoma. June, hard-seed phenological stage.

 

212. Inflorescences of troublesome sedge- This Carex species is in the subgenus Vignea the species of which are characterized by having two stigmas and perfect (bisexual) flowers in contrast to subgenus Eucarex whose species have three stigmas and separate staminate and pistillate flowers (unisexual).

Though not of high palatability or high on animals' "preference list" carices like troublesome sedge are consumed by livestock and wildlife like deer. All the grasslike plants are valuable for soil protection and formation.

Ottawa County, Oklahoma. June, hard-seed phenololgical stage.

 

213. Another caric sdege of wet prairie- Bicknell's caric sedge (Carex bicknellii) growing on the marsh-like drainage of a wet prairie used as a hay meadow. This was the same portion of meadow on which fox sedge (above) and Frank's sedge (below) were also important local species. These plants were growing in association with inland or interior rush, two-flower rush, diffuse rush fox caric sedge, Frank's caric sedge (Carex frankii), redtop panicgrass, and Florida paspalum.

Ottawa County, Oklahoma. Late May, early fruit development stage.

 

214. Distinctive top- Flower cluster and individual floral units of Bicknell's sedge. This is another Carex in the Vignea subgenus. Ottawa County, Oklahoma. Mid-June, ripening to mature fruit stage.

 

215. A caric sedge of wet prairie- Frank's caric sedge (Carex frankii) in extreme western Ozark Plateau. Several other Carex species, including C. vulpinoidea, fox sedge (shown above), and C. bicknellii , Bicknell's sedge (shown below), were growing on this wet prairie that was used as a hay meadow. These plants were growing in association with interior two-flower rush (Juncus biflorus), diffuse rush (Juncus diffusissimus), fox caric sedge (Carex vulpinoidea), Bicknell's caric sedge (C. bicknellii), Frank's caric sedge (Carex frankii), redtop panicgrass, and Florida paspalum.

Ottawa County, Oklahoma. Mid-July.

 

216. Boy groups and girl groups on wet prairie- Close-in view of Frank's caric sedge showing both pistillate (lower) and staminate (upper) flower clusters. This Carex species is in the Eucarex subgenus which was described in the caption of troublesome sedge above.

Ottawa County, Oklahoma. Mid-July.

 

217.Another (and larger) example- An extremely large specimen of Frank's caric sedge. This cespitose plant had an oblong outline or shape and was over three feet across.

Ottawa County, Oklahoma. Early June.

 

218, Important enough (at least locally) for another example- Another specimen of Frank's caric sedge (first slide) and further examples of the bi-sexual flower clusters (second, third, and forth slides) of this member of the Eucarex subgenus.

Ottawa County, Oklahoma. Mid-June.

 

219. Easily identified- The very distinctive achenes of Frank's caric sedge. These dry fruits were produced in the female flower clusters presented in the immediately preceding slides.

Ottawa County, Oklahoma. Mid-July. Ripe (or nearly so) fruit stage of phenology.

 

220. One that likes it wet- Individual plant (first slide)and its numerous shoots (second slide) of eastern narrowleaf caric sedge (Carex amphibola= Carex grisea var. amphibola ) at edge of tallgrass prairie and oak-hickory forest in the Springfield Plateau in northeastern Oklahoma.

This species, which is in the Eu-carix subgenus, is adapted to wet and moist edaphic conditions. It is a sprawling plant that reaches considerable size The specimen seen here was a yard across in a nice, moist spring.

Ottawa County, Oklahoma. Late May.

 

221.Sexual end of a moisture-loving one- Male and female flower clusters immediately above the flag leaf on a shoot of eastern narrowleaf caric sedge produced on the Springfield Plateau of the Ozark Highlands.

Ottawa County, Oklahoma. Late May.

 

222. In winter and in summer- Another caric sedge of moist, though not wet, prairie of tallgrass prairie in the Ozark Plateau and Cherokee Prairie was Leavenworth sedge (Caraex leavenworthii). It was shown here in its apparel of winter (first photograph) and of spring-summer (second photograph). Note that in its winter turf Levenworth sedge was beginning to green up by early winter. Obviously this is a cool-season species (at least in this area).

Ottawa County, Oklahoma. First slide, early December (just breaking dormancy); second slide, mid-June.

 

223. Another distinctive cluster- Inflorescences of Leavenworth's caric sedge. These two flower clusters were on plants shown in the summer sward of this species (second slide immediately above). This species of Carex is in subgenus Vignea.

Ottawa County, Oklahoma. Mid-June, early fruit development.

 

224. Composite view in full-fruit- General or overall view of Leavenworth's cqric sedge. This was a particularily large specimen that showed the "laid-over" habit of many individuals of this species due to the decumbent morphological feature of sexual shoots, especially when spikes were full of ripening achenes as was the case seen here.This particular plant was growing in the same pasture as examples presented in the immediately preceding two slide-caption sets.

Ottawa County, Oklahoma. Mid-June, late-fruit developmental stage.

 

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Few and rch- Few-fruited or richwoods caric sedge (Carex oligocarpa) growing on a tallgrass-hardwood savanna in the southwestern Prairie Peninsula.

Ottawa County, Oklahoma. Early June, early to mid-fruit stage.

 

Not rich in number- Seual shoots of few-fruited or richwoods caric sedge in the southwestern Prairie Pniinsula of the Springfield Plateau..

Caric sedges of adjoining areas in Springfield Plateau of the Ozark Uplift to the Osage Plains of the Central Lowlands physiographic province.

 

225. Heavy-laden for a little one- Inflorescence and ripening fruit (achene) of heavy or heavy-fruit caric sedge (Carex gravida) on tallgrass prairie in the Osage Ouesta part of Cherokee Prairie. The foliaceous base subtending (beneath) the spike or cluster of achenes is characteristic of severalcarex species and was of diagnostic value in identification of C. gravida. On this hay meadow heavy sedge was associated with longspike tridens (Tridens strictus) on local depressions (ponded in spring and wet autumns) within an overall consociation of big bluestem. This is another Carex species in subgenus Vignea

White Oak Prairie, Kelly Ranch, Craig County, Oklahoma. Late May; ripening fruit stage.

 

226. Drought-stricken stand but it reproduced- Microsite (small local) population of Mead's cariic sedge (Carex meadii) on dry-mesic chert tallgrass prairie in western Ozark (Springfield) Plateau. These plants (somewhere between eight to ten or, perhaps, as many as thirteen) were growing as a colony of different genotypes (not as a clonal organism). The plants surrounding these caric sedges was mostly prairie dropseed (Sporobolus heterolepis), a local co-dominant along with the Four Horsemen of the Prairies tallgrass species. Mead's caric sedge is an associate species on this climax tallgrass prairie community (Nelson, 2010, ps. 282). In this context and in this climax vegetation Mead's caric sedge is an indicator species, in this case an indicator of 1) the potential natural vegetation and 2) characteristically dry, shallow, stoney soil in the ancient, highly eroded Ozark Mountains.

All plants and their organs shown in this sequence of slides (Diamond Grove Prairie, Late July) were growing in the second consecutive summer of Extreme to Extraordinarry Drought (Palmer Drought Severity Index). The highest state of drought, Extraordinary Drought, had been in effect for over a month at time of these photographs.These plants had made most of their growth and development prior to the most damaging impacts of drought. These plants were concluding their annual life cycle at this time, undoubtedly hastened by drought and now entering drought-induced dormancy (roughly six to eight weeks prematurely). Mead's caric sedge fared much better than the dominant four tallgrass species.

.Diamond Grove Prairie, Newton County, Missouri. Late July; fruit-shedding stage of phenology.

 

227. Drought-suffering but still representative- Three plants or, in other words, three genotypic specimens (first slide) and synopsis view of one plant (second slide) of Mead's caric sedge growing on a dry, chert Ozark Plateau tallgrass prairie in the middle of an Extraordinary Drought. It was explained in the preceding caption that Mead's caric sedge naturally matures earlier than most of the other principal plants in this climax range vegetation. so that C. meadii "came out in better shape" than the other climax dominants.

Furthermore--and as shown in subsequent photographs--Mead's caric sedge successfully completed sexual reproduction and produced an abundant crop of achenes, most of which had already been shed at time of these photographs.

.Diamond Grove Prairie, Newton County, Missouri. Late July; fruit-shedding stage of phenology.

 

228. Drought-stricken stand but it reproduced- Inflorescence (a spike) of Mead's caric sedge with a full crop of mature achenes that were produced in an Extraordinary Drought (Palmer Drought Severity Scale) on a dry chert, upland tallgrass prairie.

C. meadii is in Carex subgenus, Eu-Carex.

Diamond Grove Prairie, Newton County, Missouri. Late July; fruit-shedding stage of phenology.

 

229. Mostly shed out- A few mature and not yet-shed achenes on spike of Mead's caric sedge. This author had been in the Sonoran Desert for two weeks immediately prior to taking this series of photographs of Mead's caric sedge. (Incidentially, the desert that was in a less pronounced drought than this prairie did not show water stress nearly as bad as this mesic grassland.) Before the needed-to-be-in-two-places-at-once photographer could arrive on this prairie scene most of the ripe achenes had been shed to the stoney soil of the Ozark Plateau prairie.

The miracle shared with viewers of this series was the fact that Mead's caric sedge bore fruit and successfully sexually reproduced in the most horrible of droughts. Prairie plants are survivors in the environment in which they evolved and to which they are adapted.Students must always remember that fitness in contest of "survival of the fittest" means only one thing: reproduction adequate to perpetuate an organism's species. With sexual reproduction (in contrst to asexual reproduction described above) even in Extraordinary Drought, the genotypes of Mead's caric sedge seen here maintained their option of genetic improvement, continued adaptation to their droughty environment, through genetic recombination.

Diamond Grove Prairie, Newton County, Missouri. Late July; fruit-shedding stage of phenology.

 

230. One of the more common ones- Two local populations of cedar caric-sedge (Carex planostachys) growing on a shallow calecareous soil in the Grand Prairie of northcentral Texas. Cedar caric sedge is one of the "dry land" Carex species. Many of the Carex species are either wetland plants or, at least, adapted to mesic habitats. By contrast, C. planostachys grows on some of the more shallow and drier environments within its bilogical range.

Carex is typically the largest genus (or, at least, one of the largest genera) of vascular plants in many range areas and regions. Caricologists (scientists devoted to study of the crices, Caricology) have historically divided this species-rich genus into two subgenera: Carex: 1) Vignea in which inflorescences (spikes) have perfect flowers (both male and female organs in the same flower) with two stigmas in the pistil and 2) Carex or Eu-Carex in which inflorescences consist of terminal staminate spikes beyond or above pistillate spikes (monecious flowers) and typically with three (rarely two) stigmas per pistil (Steyermark, 1963, ps.313-316; Hipp, 2008, ps.ix-x). C. planostachys is Eu-Carex.

Cedar caric-sedge is often found growing in local populations as seen in these two examples, but it also grows as isolated individual plants as shown in the following examples ...

Tarleton State University Hunewell Ranch, Erath County, Texas. Late March; ripening fruit (achene) phenological stage.

 

231. A sedge for shallower soils- Four specimens of cedar caric-sedge growing on a shallow calcareous soil in the Grand Prairie of northcentral Texas. Cedar sedge is one of the more common and widespread of the Carex species in the prairies and savannahs of northern and central Texas. Cedar sedge is about as common in the Edwards Plateau and some eastern portions of the Trans-Pecos Basin and Range provinces as in the tallgrass prairies and savannahs of northcentral Texas, and, coincidentially, cedar sedge is primarily a Texas species although it has been found as far west as southeast Arizona.

Cedar caric-sedge got its common name from its frequent occurrence or association with various "cedars" (Juniperus species) many of which, like cedar sedge, grew on shallow or rocky soils (habitats less apt to grow herbaceous fuel capable of burning with enough intensity to kill those Juniperus species that do not resprout such as J. ashei, J. virginiana). Cedar sedge also grows where there are no Juniperus (or other woody) species (though typically this is still on comparatively shallow, edaphically harsh, range sites and microsites therein).

Similarily, cedar sedge is an increaser and is more commonly found with other increasers than decreasers as for instance more with perennial or, even, annual dropseeds (Sporobolus species) rather than with little bluestem, a common climax dominant in areas of the Grand Prairie where cedar sedge is widespread.

Cedar caric-sedge is a cool-season species with fruit maturation in early to, at latest, mid-spring. In this author's observation, growth and annual variation in shoot numbers and shoot height of cedar caric sedge is quite varied from one year to the next. Empirically, there is greater annual variability in size of cedar sedge plants than in most associated grasses. This would most likely be attributable to variation in winter and spring moisture conditions.

Tarleton State University Hunewell Ranch, Erath County, Texas. Late March (specimens in first two slides), Mid-April (specimens in last two slides); ripening fruit (achene) phenological stage.

 

232. Preparing to bear much fruit- Sexual shoots with spikes at anthesis in a plant of cedar caric-sedge. This plant was growing on a shallow, calcareous soil in the Grand Prairie of northcentral Texas. There was an nusually high number of inflorescences, the result of an unusually wet spring with mild temperatures.

Tarleton State University Hunewell Ranch, Erath County, Texas. Late March.

 

233. Cool-season fruit- Sexual shoots with panicles and ripening fruit (achenes) in a plant of cedar caric-sedge in the Grand Prairie of northcentral Texas. This cool-season graminoid typically has ripe achenes by mid-spring, if not earlier (depending on temperature and moisture conditions). As typical for many plants of this species, this specimen was growing on a shallow, rocky, calcareous soil.

Tarleton State University Hunewell Ranch, Erath County, Texas. Mid-April; ripening fruit (achene) phenological stage.

 

234. Mating in the sedge- Three examples of inflorescences (spikes) at anthesis atop sexual shoots of cedar caric-sedge growing on shallow, calcareous soil in the Grand Prairie. This species, Carex planostachys, is in subgenus Carex or Eu-Carex species of which have a monecious floral arrangement with terminal or apical staminate spikes produced above (at least to the side or separate from) pistillate spikes.

This monecious feature was shown to better advantage in the "very next" two-slide/caption unit ...

Tarleton State University Hunewell Ranch, Erath County, Texas. Late March, peak anthesis.

 

235. OK, boys and girls- The monecious spike inflorescence(s) of cedar caric-sedge shown at tip of a sexual shoot (first slide) and removed from it then laid on a log (second slide), a log of what else but redberry cedar (Juiperus pinchotii). The male (staminate) spike at anthesis can be seen as distinct from the female (pistillate) spike (who appears "ready, willing, and able" to bear much fruit).

Tarleton State University Hunewell Ranch, Erath County, Texas. Late March, peak anthesis.

 

236. Green flatsedge (sometimes written as flat sedge) or umbrella sedge (Cyperus virens= C. pseudovegetus)- This is typically an aquatic plant or hydrophyte that is one of the more common Cyperus species on wet to moist soil of the tallgrass prairie, openings in the western oak-hickory forest, and the prairie peninsula ecotone between these two major range communities. That it is readily eaten by cattle was indicated by the fact that in a pasture adjoining the prairie hay meadow in which this photograph was taken no inflorescences on shoots of umbrella sedge could be found. Ottawa County, Oklahoma. June.

 

237. Manyflowered unbrella sedge (Cyperus lancastriensis)- This species or flat or umbrella sedge grows on various local habitats though it usually pre8ers moister soils. This specimen was growing on an upland slope at edge of prairie and oak-hickory forest. Many of the Cyperus species resemble each other closly in morphology so that identification to species level is difficult. Many species display considerable variability in features in infloresence, spikelet and floret. Frequently there is intergrading and hyrbridization among similar species. Treatment frequently varies greatly among authorities.

Ottawa County, Oklahoma. July; pre-bloom phenological stage.

 

238. Flowering in manyflowered umbrella sedge- Inflorescences on a plant of manyflowered unbrella sedge growing in close proximity to the individual shown in the preceding slide. This plant was more advanced phenologically and had fully exerted anthers and stigmas.

Cyperus species are not as difficult to identify accurately as are many Carex species, but both are difficult enough. Much of this difficulty is due to hybrids of closely related species. By way of example, the plant shown in this and the plant presented in the precding slide fit the description and were in the near vacinity of the hybrid specimen described by Steyermark (1963, p. 267) as that between C. lancastriensis and C. filiculmis, C. ovularis, or C. strigosus. Lower spikelets of the plant presented here did not have its lowest spikelets reflexed in the manner typical for C. lancastriensis, but it clearly was of this species. An example of differing treatments of Cyperus species among authors was provided by this case. In the expanded Steyermark's Flora of Missouri (Yatskievych, 1999, p. 372) did not allude to the hybrid that Steyermark himself (Steyermark, 1963, p. 267) gave a detailed description of.

Ottawa County, Oklahoma. July; anthesis.

 

239. Hedge-hogs (so to speak) on tallgrass prairie- Number of spikes in the compound inflorescence (first slide) and details of one spike (second slide) of hedgehog club-sedge (Cyperus ovularis). These organs had been produced in the western portion of te Ozark Highlands (Springfield Plateau) in extreme northeastern Oklahoma.

Ottawa County, Oklahoma. EArly July; immediate-past stage of anthesis.

 

240. Sand spikerush (Eleocharis montevidensis)- Eleocharis is another genus in the Cyperaceae. There are several spikerush species on tallgrass prairie. Sand spikerush is a represnetative species from the Grand Prairie of northcentral Texas. It is typically a short-statured plant. For example, the shoots in these photographs varied from six to nine inches in height.

Tarleton State University Hunewell Ranch, Erath County, Texas. March (peak anthesis).

 

241. Abdandant stand- "Solid" or "pure" stand of blunt spikerush (Eleocharis obtusa), an annual species, growing in an old farm pond that no longer held water other than on a temporary basis (for a short period of about a week or less depending on amount of rain caught). This man-made more mesic habitat than that of surrounding environments enabled this comparatively short-lived annual to prosper on a very infrequent basis (not more than one year out of four) when rainfall came "just right".

An interesting phenomenon of grazing animal behavior was that the plants in this local population were grazed exclusively by nursing beef calves (roughly 350 to less than 500 pounds). Mothers of these calves did not graze plants in this stand; did not touch them. The author observed this grazing ehavior on almost a daily basis; plus as conclusive proof of this animal feeding pattern, there were no tracks of cows in the mud in which these plants were growing. There were planty of calf tracks (as shown in the next slide...).

Ottawa County, Oklahoma. Mid-July; early bloom (anthesis) stage of phenology.

 

242. One by itself- A top-down view of a single cespitose plant of the annual spikerush commonly called blunt spikerush. This plant had been grazed by nursing beef calves. Hoof prints encircled this individual. plant. Grazing mothers of these calves did not touch these plants. This was conclusively proven by absence of cow hoof prints in mud of this local microsite. It seems that the young of species experiment more than their mammas. Nor could this different behavior be attributed to cows being reluctant to stand in mud given that a much larger water-holding pond,which existed less than forty yards away, was quite an attraction to brood cows who "wanted to cool their heels" and stood hock-deep in water of this impondment.

Tales told on the range.

Ottawa County, Oklahoma. Mid-July; early bloom (anthesis) stage of phenology.

 

243. A couple from the side- Side-view of two conspicuously cespitose plants of the annual species blunt spikerush. The first or upper slide had half of a plant at left margin.

Ottawa County, Oklahoma. Mid-July; early bloom (anthesis) stage of phenology.

 

244. Tops of the tips- Inflorescences (spikes) on sexual shoots of blunt spikerush. These shoots had somehow excaped grazing by beef calves in an old farm pond that did not hold water on a tallgrass prairie in the western Ozark (Springfield) Plateau.

Ottawa County, Oklahoma. Mid-July; early bloom (anthesis) stage of phenology.

 

245.. Woolgrass (Scirpus cyperinus)- The example of this distinctive Scirpus species was growing in a water-filled deep depresion on a virgin hay meadow in the Blackland Prairie vegetational area of northcentral Texas.

Grayson County, Texas. June, soft-grain stage.

 

246. Pendant bulrush (Scirpus pendulus)- Typical plant of pedant bulrush growing in a roadside ditch on wet prairie in the eastern portion of Cherokee Prairie, a major section of the Central Lowlands physiographic province. Craig County, Oklahoma. Late May.

 

247. Pendulus clusters- Sexual shoot with numerous panicles (first slide) and two units or clusters of spikelets off of one panicle (second slide) of pendant bulrush on wet prairie. The shoot with its entire panicle (and with numerous units or clusters) was one of the tillers on the plant presented in the immediately preceding photograph (Craig County, Oklahoma). The panicle units had spikelets, the distal most floral units of which were in anthesis (Ottawa County, Oklahoma). Both photographs taken in Late May.

 

248. Great bulrush, softstem bulrush, or giant tule (S. validus= S. tabernaemontani). The example of this species was growing in a slough at edge of a tallgrass bottomland prairie just above Lost Creek in western portion of the Ozark Highlands (Mountains). According to Steyermark (1963) this species is the common bulrush of the Ozark Plateau. The terminal end of the culm that extended beyond the inflorescence in this specimen closely resenbled that of S. acutus, but this latter species does not occur in within 150 miles or more of this location (Steyermark, 1963, p. 292-293).

Ottawa County, Oklahoma. July, hard dough-grain stage.

 

249. Common bulrush (S. atrovirens)- This Scirpus species is probably the most common one in prairies of Missouri and eastern Kansas and Oklahoma. S. atrovirens is an extremely variable species. Steyermark (1963, p. 296) divided it into three varieties which Yatskievych (1999, ps. 429-432) elevated to three species. These plants were growing in a tallgrass prairie slough that drained into Lost Creek at the western edge of the Ozark Plateau.

Ottawa County, Oklahoma. June, hard dough-grain stage.

 

250. Same plant; another year- The same perennial plant of common bulrush as shown in the preceding two-slide /caption set in another year. This plant was growing in a semi-permanent (usually yearlong) spring that formed a slough a tallgrass prairie in the Springfield Plateau.

Ottawa County, Oklahoma. Late May; soft-dough phenological stage.

 

251. Shoot details- Sheath at base of flower cluster of common bulrush. This was one of the shoots on the plant shown in the preceding slide.

Ottawa County, Oklahoma. Late May.

 

252. Burdened over- The entire flower cluster or inflorescence on one sexual shoot (first slide) and groups of flowers/achenes (second slide) on the plant of common bulrush shown above.

This plant, which was growing in a spring-fed tallgrass prairie slough, had produced a "bumper crop" of flowers and consequent achenes in this part of the Springfield Plateau in northeastern Oklahoma.

Ottawa County, Oklahoma. Late May; soft-dough phenological stage.

 

253. Goes nuts with tallgrasses- Local population (first photograph) and inflorescence (second photograph) of tall nut-rush (Scleria triglomerata). This grasslike plant is generally a weedy or seral species on improperly managed tallgrass prairie used as prairie hay meadows. Tall nut-rush florishes when big bluestem, Indiangrass, eastern gamagrass, prairie cordgrass, and other tallgrass dominants (depending on range site) are reduced by imporoper harvest practices such as overmowing, short mowing (leaving of inadequate stubble cover and height), or late mowing (mowing too late in the growing season; haying too close to end of frost-free period). There are, however, small local populations of tall nut-rush even on pristine tallgrass communities, including those not subjected to haying or livestock grazing, indicating that this species is an occasional component of tallgrass prairie in Excellent range condition class.

Such was the case of the local population or colony shown in the first photograph that was growing on an upland tallgrass prairie co-cominated by big bluestem and eastern gamagrass in the Osage Ouesta region of southestern Kansas.

First slide: Woodson County, Kansas, late June; late-bloom to late-fruit phenological stages. Second slide: Ottawa County, Oklahoma, Late May; early to mid-bloom phenological stages.

 

254. Flowers and fruit of a nutty one- Inflorescences (first two slides) and maturing fruit in flower cluster (third slide) of tall nut-rush on tallgrass parairie. The inflorescence of Scleria species consist of several spikelets partially enclosed within subtending leaves so that the entire unit resembles a foliaceous cluster (first and second slide). The fruit of Scleria species is a spherical to egg-shaped, hardened (bonelike) achene. In S. triglomerata the achene (third slide) is whitish and reminiscent of a tiny "pearl" (This fruit is a "pearl of great price" to the parent plant). The image of the bluish-white or somewhat bone-colored achene was produced in the colony presented in the first photograph of the immediately preceding slide-=caption set.

First and second slide: Ottawa County, Oklahoma, Late May; early to mid-bloom phenological stages. Third slide: Ottawa County, Oklahoma, Early June; immature fruit stage. Fourth slide: Woodson County, Kansas, late June; ripening fruit stage.

 

255. 204. Common or broad-leaf cattail (Typha latifolia)- This is one of three Typha species (as commonly interpreted by most authors) found on wetlands of the central grasslands of North America. Along with members of the rush and sedge family (and some minor ones like the burr-reed family) cattails constitute the groups of range plants commonly known as grass-like plants. This group or category of range plants carries no official taxonomic recognition in the lexicon of Plant Systematics, but is used primarily by rangemen and foresters to designate those species of monocots which do not produce flowers with showy or conspicuous petals and yet which are not grasses. These include several orders of monocotyledons. Vascular spore-bearing plants such as the horsetails or scouring rushes (Equisetum spp.) and monocots with obvious petals (eg. the orchids) are classified as forbs by natural resource practitioners like rangemen and wildlifers. foresters. The grasses and grass-like plants are, in turn, known collectively as graminoids.

Cattails are some of the most common and conspicuous species of grass-like plants on both marshes and wet microsites on grasslands across North America. Cattails grow not only in natural depressions like potholes and wet draws but also around man-made structures that impond water such as ditches, lakes, sewage lagoons, and farm and ranch ponds such as the one shown here on Kelly or Lone Oak Prairie, Craig County, Oklahoma. June.

 

256. Inflorescence of common cattail- Cattail gets its common name from its prominent flower cluster composed of distinctly separate groups of staminate and pistillate flowers. The upper structure (seen here as the gray, curved portion) of the inflorescence consist of male flowers and is called the "cattail"; the lower structure (the brown, cylindrical part) of the inflorescence is composed of female flowers known as the "cat".

 

257. Narrowleaf cattail (T. domingensis)- Local stand of narrowleaf cattail which on this savanna wetland in the Kansas Smoky Hills.

Ellsworth County, Kansas. Mid-July.

 

258. Thin cat- Shoots of narrowleaf cattail (Typha domingensis) in a local colony on a drainage in the West Cross Timbers. The staminate--upper-- inflorescences (the "cattails") and pistillate--lower-- inflorescences (the "cat") were both present.

Erath County, Texas. Late-May; post-anthesis phenological stage.

 

Forbs of Tallgrass Prairie

For openers will begin with the best, the most highly derived (and, arguably, the most beautiful) of all the forbs: the orchids.

259. Early or spring lady's tresses (Sprianthes vernalis) on a badly overgrazed tallgrass bluestem prairie- It would appear that this little orchid is particularily palatable. According to most taxonomic accounts the Orchidaceae is the largest family inthe Angiospermae (flowering plants). Most of these orchid species are in tropical forests (forest generally for that matter) some are grassland species. Two of the more widespread orchid species on tallgrass prairie were included here to provide coverage of more grassland plant taxa. Besides, no other group (family) of "wild flower" or forb can rival the Orchidaceae for sheer diversity of beauty. Ottawa County, Oklahoma. June.

 

260. Inflorescence of spring or early land's tresses- This dainty orchid inflorescence was on an individual that was a neighbor to the specimen presented in the preceding slide. No other taxon of plants so captures the fancy of wild flower followers than the orchids. None surpass their striking diversity and beauty. The crossover crooner Eddie Arnold sang of sending red roses to a blue lady, but (rose lovers take note) "... if that does the trick I'll hurry back and pick your best white orchid for her wedding gown". Ottawa County, Oklahoma. June.

 

261. Tiny male in his territory- Several flowers of spring or early land's tresses one of which served as the "throne" for a male celer craab spider (Mecaphesa celer) proudly guarding his "kingdom" and waiting for a female to serve as his "queen". This entire photographic subject seemed worthy of writing by Solomon, a philosopher king who appreciated things of value, large and small.

Ottawa County, Oklahoma. Early July, first slide; early June, second slide.

 

262. The not-so-showy part- Basal shoot with senesced leaf of spring or early lady's tresses at full-bloom stage. This was the shoot that bore the inflorescence shown in the immediately preceding slide.

Ottawa County, Oklahoma. Early June.

 

263. A llonger paart of not-so-showy- Another basal shoot of spring or early lady's tresses from a greater camera distance to show remnants of the two leaves of this prairie orchid. In this population of Sprianthes vernalis individual plants had from one to three leaves with two being the most common number.

Ottawa County, Oklahoma. Early July.

 

264. Beauty in a botanical spiral- Details of the inflorescence of spring or early lady's tresses on a degraded tallgrass prairie in the western Ozark (Springfield) Plateau. These examples were of the same population as the examples presented earlier (above) though in a different year The year in which the specimens exhibited here were produced was a spring/early summer with daytime temperatures that were eight to twelve degrees Fahrenheit below mean (arithmetic average) temperatures. The flower clusters is this substantially cooler year were produced from three to four weeks later than was typical for this location.

The genus name Spiranthes is from the Greek speira, meaning a spiral or wound or twisted around, and anthos Greek for flower; hence spiral flower (Allred and Ivey, 2012, p. 616).

Ottawa County, Oklahoma. Early July; peak-bloom phenological stage.

 

265. Inflorescence of nodding lady's tresses (Spiranthes cernua)- Some like it early; some like it late. This species of Spiranthes is best distinguished from the preceding S. vernalis while on the range simply by knowing spring from autumn. Ottawa County, Oklahoma. Nodding lady's tresses is a late bloomer while spring lady's tresses comes of age at an earlier season. This specimen was on an old-field that had once been tallgrass prairie. Ottawa County, Oklahoma. November.

 

266. A ragged but rare find- Ragged orchid (Habenaria lacera) on a tallgrass prairie hay meadow in the western Ozark Plateau. This first pair of photographs presented this rare prairie forb with its neighbors that included wood betany (Pedicularis canadensis) dead fruit stalks; prairie blazing star or prairie gayfeather (Liatris pycnostachya), greatest number of linear leaves; ashy sunflower (Helianthus mollis), rosette panicgrass (Panicum lanuginosum), winter bentgrass (Agrostis hymenalis), and tall or or littlehead nut-sedge (Scleria oligantha). Yes, all these species are visible (at least they were--and clearly--before scanning the slides) if you are good enough at sight identification to find them.

The potential dominant of this tallgrass prairie vegetation was big bluestem with switchgrass as the associate. Unfortunately, this meadow had been degraded by delayed haying (any time from late summer to early autumn) for at least 40 years (probably for over a half century). Hence, invader speacies like littlehead nut-sedge and numerous forbs such as prairie blazing star were much more plentiful than would have been the cse on virgin prairie. However other forbs, especially cool-season and spring-blooming species like wood betany and ragged orchid, were not impacted by late mowing. In fact, if anything mowing late in the warm-growing season benefitted these species via reduced competition from early growth of tallgrasses that had been weakened and reduced by improper hay making.

The entire shoots of these specimens of ragged orchid were visible in these two slides.

Ottawa County, Oklahoma. Late May; peak-bloom stage.

 

267. Ragged and raw beauty- Upper shoots complete with inflorescences of three plants of ragged orchid on a degraded tallgrass hay meadow in the western Springfield Plateau.Other plant species visible in these two photographs included tall or or littlehead nut-sedge, wood betany, rosette panicgrass, ashy sunflower, and catclaw sensitivebriar.

Ottawa County, Oklahoma. Late May; peak-bloom stage.

 

268. Ragged flowers and contrast in light- Two photographs of the same flower cluster of ragged orchid taken within a minute (or less of each other). The photographer took these paired shots with two goals in mind: 1) get as nearly perfect photograph as possible of the subject and 2) compare full-sun to light overcast for quality of photograph. Both photographs were taken at shutter speed of 1/15th second. The first and obviously most crisp photograph was taken in full sun. The second and, unfortunately, wind-blown photograph was taken under cover of a fair-weather cumulus cloud. The comparison was confounded by the wind-blown inflorescence, but obviously greater depth-of-field (higher f stop; approximately 8 vs. 5.4) was possible under the brighter light of a full-sun sky. The prickled stem and unopened floral buds beside and slightly behind the orchid inflorescence were of catclaw sensitivebriar.

There is a common idea that slightly dim light, such as that of early morning sunlight, is superior for wild flower photography. The stated logic for this view is that the brightness of direct, full sunlight "bleaches out" the more subtle colors often found in petals.This is sometimes true particularily for pastel shades such as pink, light blue, pale yellow, etc. Nonetheless, it has been this photographer's experience over more than three decades of photographing native plants on range, forest, and field and at all seasons that the absolute best light conditions for more than the proverbial "nine times out of ten" is bright, full, glaring sun at high noon without one single puff of a cloud in a summer sky.

The reason for this is simple: more detail due to greater depth-of-field. That says it, period. This is the case with slower film which, is of course, essential if one wants highest quality photographs that are free from grainness and portray color as the human sees them. Slow-speed Kodachrome is in a league by itself in this regard. None of the E-6 process films come close to capturing the yellows, reds, oranges, browns, and, even, buff-colored greens of most grasses like Kodachrome does. The other Kodak films are clearly inferior to those of Fujichrome, especially Provia 100F which is the film least different from Kodachrome. End of statement as to superior film.

With slow-speed, superior-quality films (Kodachrome 64 to Provia 100F) slow shutter speeds (1/30th down to 1/15th second) are necessary to get deep enough depth-of-field to show details of intricate flowers such as those of rgged orchid. That means a requirement of bright, direct, sun light (ie. when artificial lighting is not used).

The myth of dim, morning light is just that: a myth or misconception. That is, unless one just wants pictures of pretty petals and does care about other details of inflorescences. The latter is the equivalent of expensive wedding pictures that just show smiling faces while lace on the bridal gown is as much a blur as the second-place suitor.

Ottawa County, Oklahoma. Late May; peak-bloom stage.

 

2 69. Ragged details- Three views of increasingly closer and more detailed views of flowers of ragged orchid. Orchids are the darlings of flower fanciers, including connoisseur of wild flowers. The most cursory gander at these beauties shows why.

Orchid flower vary greatly in this the Orchidaceae, third largest family of angiosperms, but they have some features in common. Pollen grains are most commonly clumped together in a unit known as pollinia (singular, pollinium). The androecium or male unit usually is made up of only one or two stamens and these are united with stigmas and styles forming a gynandrium or column. Typically there are three stigma, the later two of which are fertile while the central third stigma is sterile and designated the rostellum. Pollen is deposited on the sticky end of this rostellum. Students desiring more details were referred to Smith (1977, p. 259) from which this brief description was taken.

These beautieswere growing on a degraded tallgrass prairie in the western Springfield Plateau.

Ottawa County, Oklahoma. Late May; peak-bloom stage.

 

270. A striking prairie beauty- Wild hycanith, eastern camas, or Atlantic camas (Camissia scilloides) growing on a consociation of big bluestem in the Osage Cuestas section of the Central Lowlands (more commonly known as Cherokee Prairie). This member of the lily family (Scilloideae subfamily of Liliaceae) is found widely and variously on tallgrass prairie and glades within the oak-hickory forest as well as ecotones between these. The biological range of this species is quite large extending from Ontario and the Great Lakes Region through Pennsylvania to the east and southward through Iowa and Missouri and on to Georgia and Texas.

Plants of this prairie forb were not super-abundant on the virgin sod of this tallgrass prairie, but it was present as a healthy population in the minst of a nearly single-species stand of big bluestem. Shoots of Atlantic camas in this stand were from one and a half to over two feet in length although, being the top-heavy with these beautiful inflorescences, they were bent over with ascending shapes. Wild hycanith is one of the more common--though never abundant---of the monocotyledonous forbs on tallgrass of prairies in climax or sub-climax state. As was the case for other Camissia species American Indians ate the bulbs (raw or cooked; boiled or roasted) of C. scilloides.

White Oak Prairie, Kelly Ranch, Craig County, Oklahoma. Late May; peak-bloom phenological stage.

 

271. Drummonded beauties- Specimens of Drummond's onion (Allium drummondii), native perennial forb, growing on a degraded tallgrass prairie range (Fair range condition class) in the Grand Prairie of northcentral Texas These plants grew in an unusually wet late winter and early spring period in association with native species ranging from annuals like sixweeks fescue (Festuca octoflora) to the dominant decreaser, little bluestem (Andropogon scoparius) and naturalized species like redstem filaree (Erodium cicutarium) and Japanese chess (Bromus japonicus). These plants of Drummond's onion were unusually large and vigerous in comparison to plants growing in a more typical (drier) growing period. Inflorescences of this year's plants were also larger and more colorful than usual ones in drier years. (The author could not resist the temptation to photograph them and subsequently share them with those less blessed to see them on the land.)

It is doubtful if any Allium species are palatable to grazing animals. Any Allium foliage eaten (and the photographer did not find any plants with foliage removed) would probably have been unintentional and instead gathered unavoidably with consciously selected forage. Foliage of Drumond's onion does not have as pungent an odor (at least not to humans) as some of the larger Allium species, but it almost a given that if eaten onion odor would have been secreted in the milk. Schmidt (1971, p. 296) explained that most off flavors in milk come from feed, including onions. These off flavors (odors) in milk are imparted through the blood via the digestive system, but some of the digestion-derived odor comes through the lungs from eructated gases of fermentation (Schmidt, 1971, p. 296). It is also possible that some of the odors from plants, such as onion, could come directly from the atmosphere surrounding plants on range or pasture. The author remembers vividly--though not particularily fondly-- drinking raw milk (or trying to) from the family cow grazing spring pasture with wild onions and garlic bloomingly beautifully amid green grass.

Tarleton State University Hunewell Ranch. Mid-April; full-bloom stage of phenology.

 

272. Lowly beauty- Bright and unusually large flower clusters of Drummond's onion, a native perennial monocot growing on a deteriorated tallgrass prairie range in the Grand Prairie of northcentral Texas. These inflorescences, like the rest of the shoots on which they were produced, were larger and more colorful than usual because they grew in an unusually wet late winter and early spring, the growing season for these cool-season perennial forbs.

Tarleton State University Hunewell Ranch. Mid-April; full-bloom stage of phenology.

 

273. Beautiful false dragon's-head (Physostegia pulchella)- Yes, that was the long common name listed for this species by the definitive manual for the area (Diggs et al. 1999, p. 770) which called this "the most common Phyostegia of the Blackland Prairie". Another common name for many of the Physostegia species is obedient-plant which originated from the feature whereby individual flowers remain in the position in which they were placed by human hands (or other forces like wind).

Physostegia species are members of the mint family (Labiatae) which is often one of the families best represented on prairies and adjoining forests and savannas. This dense stand (which was grown under cultivation for ornamental purposess) illustrated the rhizomatous characteristic of this genus.

 

274. Inflorescence of obedient-plant (P. virginiana)- This is another Physostegia species found on the Blackland Prairie (Diggs et al., 1999, p. 770). Like the preceding species this one was grown under cultivation.

The Leguminosae is second only to the Compositae in total number of species on North American ranges. A much higher proportion of legumes than composites are valuable forage plants. The nodulated or papilionaceous legumes (generally only members of the subfamily Papilionoideae host nitrogen-fixing Rhizobium bacteria) are the more nutritious and palatable Leguminosae species. With rare exceptions nodulated legumes are forbs and not shrubs. Legumes are higher in energy and nutrients like protein and minerals (especially calcium) than grasses and most other forbs. Native papilionaceous legumes are desirable members of range plant communities and are often important indicator plants. They are generally decreaser species in which cases their presence on range indicates proper grazing management and higher range condition class. The extent to which the legume-nitrogen-fixing bacteria symbiosis provides soil nitrogen for other range plants like the grasses has not been determined, but undoubtedly it is of some key role.

 

275. Heavy on the legume- Lead plant (Amorpha canescens) is one of the more common native legumes on tallgrass prairie. It typically occurs as individual to only a few plants yet is is one of the most characteristic legumes of this range type. It is also quite nutritious and is a decreaser in response to heavier grazing (as in intensive early stocking). The specimen shown here was over a yard in height.

The second slide showed the shoot apex including the entire flower cluster or assembly.

Ottawa County, Oklahoma. Mid-June; early-bloom stage.

 

276. Purple Pb blooms-Spikelike racemes of the leadplant introduced in the immediately preceding slide. Fuzzy pubescent leaflets (basis of the epithet, canescens) the compound leaf were also visible.

Ottawa County, Oklahoma. Mid-June; early-bloom stage.

 

277. Sorta woody- Leaves and inflorescences of indigo bush (Amorpha fruticosa), a subshrub This is a grandpaw version of lead plant (Amorpha canescens) introduced immediately above. A. fruticosa was thus included here rather than with woody plants of tallgrass prairie below. This was an arrangement made strictly for continuity purposes and to facilitate student learning of related species.

Newton County, Missouri. Late May; peak-bloom phenological stage.

 

278. Indigo in detail- Leaflets and flower clusters of indigo bush. Shoot apex with a closer views of leaflets and inflorescences of this shrub in tribe Galegeae. Nowata County, Oklahoma, May.

 

279. Inflorescence of purple prairie clover (Petalostemon purpureum)- Several Petalostemon species occur on tallgrass prairie and the ecotone (a transition zone) formed where the borders of the tallgrass prairie and oak-hickory forest merge. The prairie clovers are papilionaceous legumes with the individual five-petal corolla of each flower arranged on a column the entire structure of which is designated a head or, sometimes, a spike. Prairie clovers are quite palatable and relatively sparse compared to the greater relative abundance and herbage production of grasses. The Petalostemon species thus often serve as indicator plants. Prairie clover is viewed as an ice cream species on some range sites where it is able to persist only under light or no more than moderate grazing.

Top two (first two) slides were from Springfield Plateau, Newton County, Missouri. Third slide was from Hunewell Ranch, Erath County, Texas. All slides taken in May.

 

280. Gold on Grand Prairie- One good-sized and two smaller plants of golden prairie clover, golden silkthumb, or golden dalea (Dalea aurea) and details of the inflorescence of this papilionaceous range legume (first and second photographs, respectively) growing on a degraded little bluestem-Indiangrass tallgrass prairie in the Grand Prairie of northcentral Texas. D. aurea is a widely distributed, native leguminous forb that has a biological range that extends from northern Mexico northward to South Dakota and westward to New Mexico with an outlier population in central Arizona, a good portion of the Western Range Region.

Needless to say across, golden prairie clover is an extremely showy species with its large heads of papilionaceous flowers adding a striking presence on many a well-managed range. It was remarkable that plants presented here were growing on a dreadfully overgrazed tallgrass prairie (at least formerly) cattle range. Flowery neighboring species included Wright's skullcap (Scutellaria wrightii), white compassplant or white rosinweed (Silphium albiflorum), Missouri evening-primrose (Oenothera macrocarpa subspecies macrocarpa), among other forbs (most species of which were included later in this chapter), yet almost no decreaser and very few increaser grass species.

Tarleton State University Hunewell Ranch. Late May; and no doubt at peak bloom stage.

Note on location: other examples of Dalea aurea were included in the chapter, Mixed Prairie -IA.

 

281. Rounded out on a grazed-out prairie- Topdown view of roundhead prairie clover (Petalostemon multiflorus= Dalea multiflora) on a contest staging area for Future Farmers of America contests on a college ranch. What range vegetation hads not been destroyed by cattle overgrazing was probably finally annilated by school buses. Nonetheless, this hearty specimen of roundhead prairie clover had miracelously survived and, in fact, appeared to be prospering.

Palatability of Petalostemon species is regarded as comparatively high. It appeared that cattle finally had abandoned this grubbed-out part of the college ranch as having so little left to eat that it was not worth their effort looking for what no longer existed. This plant showed evidence of grazing by white-tailed deer albeit rather sparingly. A neighboring plant of roundhead prairie clover had not been so fortunate. Please stay tuned...

Tarleton State University Hunewell Ranch, Erath County, Texas. Mid-September; peak-bloom stage of phenology.

 

282. Round heads on rounded (and lightly grazed) shoots- Shoots of roundhead prairie clover with its rather dainty leaves and clusters of papilionaceous flowers in a cluster regarded as a head. Some of the leaves on these shoots had been eaten by white-tailed deer.

Tarleton State University Hunewell Ranch, Erath County, Texas. Mid-September; peak-bloom stage of phenology.

 

283. Round heads- Inflorescences of roundhead prairie clover. The numerous papilionaceous flowers of this species were arranged in a circular pattern called a head tht was frequently described as globose. Various insects were attracted to this arrangement. According to the range classic, Botany of Western Texas (Coulter, 1891-1894, p. 79) P. multiflorus was commonly found on prairie throughout Texas.

Tarleton State University Hunewell Ranch, Erath County, Texas. Mid-September; peak-bloom stage of phenology.

 

284. Good place to talk about selective grazing- A single plant of roundhead prairie clover prairie clover (Petalostemon multiflorus= Dalea multiflora) had been "hammered" (overused) by beef cattle and white-tailed deer while several shoots of the rhizomatous western ragweed (Ambrosia psilostachya) adjacent to the legume had not been touched. The prairie clover was to the left of western ragweed in the first photograph and to the left of western ragweed in the second photograph. The first photograph was a topdown view whereas the second photograph was a side view of these same range plants. (The photographer changed camera focus direction and orientation to provide different views of this selective defoliation.)

Actually there was also a group of smaller, shorter shoots of western ragweed to the opposite side of the roundhead prairie clover (western ragweed shoots to left and right of roundhead prairie clover) in both photographs, but the set of two or three larger, taller ragweed shoots so dwarfed the lower-growing ones that the latter were irrelevant. The photographer did know reasons for smaller shoots of ragweed, but they had not been grazed either. Given the rhizomatous morphology of the perennial western ragweed it was possible that the smaller shoots were younger offshoots (clonal units) of the same genetic plant as the larger, taller ones. Anyway, this was a classic example of selective grazing by two ruminant species (one native; one domesticated). It was also an example of overutilizaton. The remains of the poor specimen of roundhead prairie clover shown here was barely visible due to its having grazed to nearly ground level. An example of utilization that was much less severe was presented in the next slide (and it was heavy use bordering on overuse).

This was an area of tallgrass prairie in the Grand Prairie of northcentral Texas that had been degraded by human action, in particular overgrazing by cattle and excessive vehicular traffic in a shcool bus parking area for Future Farmers of America Range Contests.

Tarleton State University Hunewell Ranch, Erath County, Texas. Mid-September.

 

285. Utilization on the heavy side- Defoliation of roundhead prairie clover by beef cattle and white-tailed deer on a severely deteriorated area of tallgrass prairie on the Grand Paririe of northecntral Texas. Roundhead prairie clover is a forb, but it is not strictly herbaceous and instead falls into one of those suffrutescent or suffruticose categories. Great Plains Flora Association (1986, p, 443) described this species as a perennial herb with a "branching, woody, subterranean caudex"

Almost all of the aboveground parts of roundhead prairie clover are annual tissues so that parts seen in the above plant were not perennial (ie. there could not be vegetative regeneration from these shoots). Though this plant had been hedged to considerable degree and had no infloresences remaining (hence sexual reproduction was impossible), it had not been "abolished" to the degree seen in the overutlized speciment discussed above. This plant should survive through winter dormancy and grow back from its woody caudex next growing season. The case could be made, however, that utilization of this plant was excessive (plant was over-used) because there was no possibility of sexual regeneration from it this year.

Great Plains Flora Association (1986, p. 443) specified that roundhead prairie clover was selectively grazed by all species of livestock and readily grazed out on abused ranges (ie. it is a decreaser).

Tarleton State University Hunewell Ranch, Erath County, Texas. Mid-September.

 

286. Blue wild indigo or blue false indigo (Baptisia australis var. minor)- There are three species of Baptisia native to the grasslands of the North American Great Plains and Central Lowlands. Blue wild indigo is one of these. It is represented here by a robust specimen blooming in early morning light on an old horse trap in the Cherokee Lowlands physiographic unit or cherokee Prairie region in southeastern Kansas.

The Baptisia species (another one was included as the next succeeding ranged plant) are but one genus of papilionaceous legumes (Papilionoideae subfamily of the Leguminosae) on tallgrass prairie. This example was flourishing in the early morning light of southeastern Kansas.

None of the wild-indigaos are very palatable although individual plants bear sign of having been "nibbled on" by grazing animals. In this rangeman's observations even cattle will take small portions of these species, but they are not highly preferred like some of the other legumes such as catclaw sensitivebriar or Illinois bundleflower.

Labette County, Kansas. Mid-May.

287. Papilionaceous flower- Detail of inflorescence of blue wild-indigo. Individual papilionacaeous flowers off the central stalk of an inflorescence of the plant shown immediately above.

Labette County, Kansas. Mid-May.

288. White wild or false indigo (Baptisia leucantha)- This is another wild or false indigo indigenous to the tallgrass prairies. Like most native legumes it occurs as scattered individuals in dense grassland swards, including the sod of introduced grasses like that of tall fescue (Festuca arundinacea) seen here. As the stand of tall fescue grew older (more time elapsed since last tillage of the land) native prairie plants like white wild-indigobegan to reappear on this former tallgrass prairie.

Newton County, Missouri. Mid-June; peak-bloom phenological stage.

 

289. White wild indigo- These leaves and flowers are on a specimen growing over 6 � feet tall on a Chert Savanna range site of big bluestem and black oak in the Ozark Plateau. Newton County, Missouri. June.

 

290. A little later- Details of leaves and legumes of white wild or false indigo. Ottawa County, Oklahoma. Late July, ripe fruit stage.

 

291. A long sexual unit- Entire inflorescence of white wild-indigo with baby legumes to not-yet-open flower buds on the extended sexual portion of the primary shoot of this plant. This is conspicuously an indeterminate inflorescence, one which matures from the "bottom up" and/or "outside in" (ie. oldest flowers are the most basal and/or exterior) in contrast to a determinate inflorescence on which individual flowers first from the top or appex and/or from the interior of inflorescence (most distal and/or innermost flowers bloom first). This is indeterminate versus determinate flowering. Besides being a papilionaceous legume this provided a good example of flowering pattern.

Based on this author's field observations, the flowers of Baptisia species are the most palatable part part of the shoot. One can find intact inflorescences like the one shown here only on local areas protected from livestock (namely cttle) grazing. This pretty example was on an upland tallgrass prairie-- and in a fence row.

Ottawa County, Oklahoma. Late June.

 

292. Closer in on the sexual unit- Flowers of white wild-indigo or white false-indigo. Specimen was growing on a tallgrass prairie in the western Ozark Plateau. Ottawa County, Oklahoma. Early June.

 

293. Stalked beans on the prairie- Mature legumes of white wild-indigo on a relict area of tallgrass prairie in the western Ozark Plateau. This example was growing in a fence row where cattle could not get at it. None of the Baptisia species are overly palatable, but animals (including cattle) exhibit enough preference for them that most wild-indigos plants bear evidence of animal defoliation especially of sexual shoots (flower stalks). A nice sample of mature legumes like this is typically rewstricted to animal excluded areas like fencerows.

Ottawa County, Oklahoma. September; fruit-ripe stage of phenology.

 

294. Shoot details of a so-called false one- Close in views of main stem and branches (first slide) and of compound leaves (second slide) of white wild-indigo or white false-indigo on a tallgrass prairie in the western Ozark Plateau. This plant was growing on the outside of a fencerow where it was protected from beef cattle that were being forced to overgraze the inside of the fencerow. Note a few immature legumes in the first photograph.

Ottawa County, Oklahoma. Early June.

 

295. Close relative- Portions of crowns of three plants of long-bracted wild (false) indigo (Baptisia leucophaea) growing on a shallow upland habitat that was tallgrass prairie-savanna (oak-hickory) in the western part of the Springfield Plateau in southwest Missouri.

Newton County, Missouri. Early July; fruit-ripe stage of phenology.

 

296. Long-bracted legumes- Fully ripe (even dehiscing) legumes of long-bracted wild (false) indigo growing on a tallgrass prairie savanna in the western Ozark (Springfield) Plateau in southwest Missouri.

Newton County, Missouri. Early July; fruit-ripe stage of phenology.

 

297. Fruit of a related species- Legumes of long-bracted wild indigo or long-bracted false-indigo (Baptisia leucophaea). Tallgrass prairie-oak-hickory forest edge or narrow ecotone in western Ozark (Springfield) Plateau.

Newton County, Missouri. Late May; immature--but ripening fast--stage of phenological development.

 

298. Trailing bush-clover or trrailing lespedeza (Lespedeza procumbens)- There are several Lespedeza species that are native to the North American tallgrass prairie. Several of these were included herein. Some of these are upright whereas others creeping or trailing in haabit. Trailing lespedeza is one of the more obvious of the creeping species. It is also one of the more 'frail" appearing of the so-called bush-clovers.

Ottawa County, Oklahoma. September;full-bloom phenological stage.

 

299. Native legume amid native grasses- Roundhead lespedeza or roundhead bush-clover (Lespedeza capitata) is one of the upright and cespitose native le5pedeza species. This robust specimen was growing on restored tallgrass prairie on which tall dropseed was the dominant (mostly a consociation of tall dropseed) and big bluestem the associate species. Switchgrass was the almost exclusive grass species in background.

Homestead National Monument, Gage County, Nebraska. Mid-August; immediate pre-bloom stage.

 

300. Heads lined up- Dead sexual shoot of roundhead lespedeza with numerous flower clusters arranged as heads or capitula (capitulum, singular) groups of sessile flowers arranged on a common receptacle called a disk. This is a more common arrangement in the Compositae, but several genera of the Leguminosae have this feature also.

Ottawa County, Oklahoma. Early January; dead shoot stage of winter dormancy.

 

301. Dead heads- Dry (dead) flower clusters (capitula) of roundhead lespedeza. Legumes in these heads were very much alive and represented the next generation.

Ottawa County, Oklahoma. Early January; dead shoot stage of winter dormancy.

 

302. Slender bush clover or slender lespedeza (Lespedeza virginica)- This is one of several Lespedeza species native to the central and southern portions of the Prairie Peninsula and tallgrass prairie region. This large legume with its delicate leaves and inflorescences is extremely palatable and generally found only under light grazing and the outside of a fence row.

Ottawa County, Oklahoma. September.

 

303. Dead but alive- Dead sexual shoots with flower cluster of slender bush-clover or slender lespedeza. Viable seeds in the plump legumes in these inflorescences were the propagules or diaspores of the next generation of this palatable native legume.

Taxonomic Note: legumes of tribe Hedysareae, which includes Lespedeza and Desmodium species, open transversely in contrast to the more common opening (dehiscing) of legumes in the longitudinal plane.

Ottawa County, Oklahoma. Mid-October; dead shoot stage of winter dormancy.

 

304. Wooly or purple locoweed (Astragalus mollisssimus)- Not all papilionaceous, herbaceous legumes are "good guys". This showy character looks good enough to eat, and so it is thereby creating a poisonous plant problem. Wooly loco causes the classic loco poisoning. The poisonous principle (= toxin) is an indolizidine alkaloid (Cheeke and Shull, 1985, p. 200). This is one of the most costly and spectacular forms of range livestock toxicity. Readers are referred to the timeless work of Kingsbury (1964, ps. 306-311). Remarkably little is known of this historic form of range livestock poisoning as evident by its near absence from the standard veterinary manuals. Wooly loco is widely distributed ranging from Texas to Wyoming eastward to the Dakotas and Kansas. Rocky Mountain National Park, Colorado. Early August.

 

305. Illinois bundleflower (Desmanthus illinoensis)- Shoots of Illinois bundleflower on a prairie remnant in western Springfield Plateau of Ozark Mountains. This plant was at peak bloom, but in this species blooming is indeterminate and progresses from lower portions of shoot to shoot apex.

While Illinois bundleflower is not a papilionaceous legume (it is in subfamily Mimosoideae not Papilionoideae) in some respects this species represents tallgrass prairie legumes as much as any other member of the Leguminosae. Illinois bundleflower is clearly one of the most important native herbaceous legumes in tallgrass prairie across much of the Central Lowlands and eastern Great Plains. The western extreme of this species range extends to Alaska while it is found naturally as east as the Great Lakes Region. Its palatability, large size, wide geographic distribution, drought tolerance, distinctive features, high seed yield, relative ease of propagation, regrowth potential, and competitiveness have made Illinois bundleflower a major species for range reseeding and prairie restoration projects. Soil (Natural Resources) Conservation Service plant materials centers have developed and released numerous accessions of this species over the decades of their diligent work.

For descriptions of Illinois bundleflower from perspective of range management and grassland imporvement readers were referred to such basic works as Phillips Petroleum Company (1963, p. 82) or Nicholson (2006, p. 82), Hatch and Pluhar, 1993, ps. 192-193), and Tyrl et al.(2002, ps. 180-181).

Ottawa County, Oklahoma. July (early summer); full bloom stage.

 

306. Beauty and utility- Compund leaves and the infloresence of Illinois bundleflower. These are bipinately compound leaves. The flowers of the Mimosoideae are interpreted as heads of several small flowers each with several stamen. In the Mimoseae tribe of which Desmanthus is a member there are as up to twice as many stamen as petals (Smith, 1977, p. 152). These two slides presented details of floral and foliage features on one of the shoots presented in the two preceding photographs.

Ottawa County, Oklahoma. July (early summer); full bloom stage.

 

307. Young but on their way- Immature cluster or "bundle" of new legumes on one of the shoots of Illinois bundleflower introduced above. It is actually the "bundle" of fruit not flowers that give this important range legume its common name.The dried stigma and stamen are still visible in this cluster of legumes.

Ottawa County, Oklahoma. July (early summer); early fruit stage.

 

308. Nearly there- Clusters of legumes of Illinois bundleflower at midway stage and approaching ripe stage. This plant was a neighbor of the blooming specimen shown in previous slides of this species. All these plants were on a tallgrass prairie remnant in the western part of the Ozark Highlands (Springfield Plateau) in what was formerly a portion of the Prairie Peninsula.

Ottawa County, Oklahoma. July (early summer); approaching fruit-ripe phenological stage.

 

309. There- Photograph of a section of a plant of Illinois bundleflower at fruit-ripe (seed-shatter) phenological stage followed by a photograph of a ripe dehising legumes on this same plant. This particular plant was one of several that were thriving in a field of bermudagrass (Cynodon dactylon) on the flood plain of the Red River. More on this aspect in the next slide ...

Cooke County, Texas. July (early summer); peak standing crop and fruit-ripe to seed-shatter stages.

 

310. Several stories (well, two good ones anyway)- Two plants of Illinois bundleflower in a field of bermudagrass on floodplain of Red River.The competitiveness and relatively large size of Illinois bundleflower was noted previously in this section. Both of those ecologically and economically important characteristics were shown in this photograph of two plants of Desmanthus illinoensis thriving in a dense stand of bermudagrass. Bermudagrass under proper management grows so dense and aggressively that weeds (monocot or dicot) are seldom a major problem with this field crop. The author found it revealing that the only "weed" (defined agronomically as any unwanted plant growing in the intended crop) in this patch of bermudagrass on Red River alluvium was Illinois bundleflower, except for Johnsongrass growing at field edge bordering on the river (background of this photograph).

The second story in this slide was a lesson in grazing defoliation from perspective of 1) grazing animal selectivity and 2) morphological/physiological response of plants to defoliation. Close observation of the Illinois bundleflower in this slide revealed not one but two plants. A taller, ungrazed, sexually reproductive plant was growing in front (more forward toward camera) of a shorter, closely grazed (by whitetail deer), sexually immature plant (to rear of tall, fruit-bearing plant). Deer had cropped the one plant of Illinois bundleflower on several feeding bouts as evident from age of scars and production then grazing of regrowth on several partly defoliated shoots. There had been zero feeding on the other (larger, taller, fruit-bearing) plant. This deer-feeding behavior kept the grazed bundleflower in a sexually immature state with more shoots that had less mature stems and leaves (hence, undoubtedly more nutritious forage). Deer converted the grazed bundleflower from a sexually reproductive to an asexually reproductive plant. Both plants survived in their respective states of reproduction and in their morphological forms that were induced by defoliation by deer (or permitted by non-defoliation).

Why did deer choose to feed on the one and totally "defer" the other bundleflower that was only inches away from the frequently cropped plant. Was one more nutritious, larger, or of a different plant form at time of first feeding? Did one smell differently? Were any differences due to different genetypes or did a canine (or deer) mark its territory on one bundleflower and not the other? Or was it simply "luck of the draw", a strictly random event?

By the way range students, was defoliation or non-defoliation of this palatable prairie forb (herbaceous legume) by free-ranging native ruminants the "natural condition"? Asked in an applied sense, would grazed or ungrazed plants serve as the "control" plant in an experiment (say, for instance, of plant response to drought stress)? Don't jump to conclusions now.

Cooke County, Texas. July (early summer); peak standing crop and fruit-ripe to seed-shatter stages.

 

311. Coming back- Regrowth on an Illinois bundleflower about 16-17 days following total defoliation (no leaves left; stubble height of shoot of six to eight inches) by mechanical shredding (road crew). An unfortunate (?) Illinois bundleflower with a height of slightly less than two feet "got in the way" of a rotary shredder in late June. The bundleflower was stripped of every leaf, young bloom, etc. leaving a spindly, split stalk. Undetured, the brave little bundleflower which, it should be underscored, was rooted in soil moist from recent rains, regrew as shown here in slightly more than two weeks. The shattered basal shoot of this plant died so that there was no living tissue on this above the ground surface. All regrowth was from subterranean tissue.

Ottawa County, Oklahoma. July (early summer).

 

312. Coming back even more- After approximately 35 days following more-or-less total defoliation (except for about a half-foot length of splinttered stalk) by mechanical shredding the never-say-die Illinois bundleflower had regrown densely foliated shoots ass shown in this slide. This and the preceding photograph demonstrated the potential for regrowth (given adequate soil water) of one of the most valuable, native, herbaceous legumes on tallgrass and mixed prairie ranges.

Under most systems of proper grazing management Illinois bundleflower would not be completely stripped of aboveground living tissue. On tallgrass prairie meadow that are mowed for hay plants are cut (severed neatly) not twisted off and splintered (ie. beat to death) as with shredding. Hence, injury to mowed plants is much less than damage inflicted by beating and twisting. In other words, regrowth on the Illinois bundleflower shown in these photographs had to make a greater recovery than that which would be required of plants grazed on properly managed ranges or mowed for prairie hay on a commercial basis.

Regrowth potential and tolerance to severe (if infrequent) defoliation in Illinois bundleflower makes this an extremely valuable, native prairie legume.

Ottawa County, Oklahoma. July (early summer).

 

313. Unmolested- Sexual shoots of Illinois bundleflower that had received no substantive defoliation (by man or beast). In a very moist growing season these shoots were of nearly unbelieveable heights (between five to over six feet) with both flowers and immature fruit.

Newton County, Missouri. Western Ozark (Springfield) Plateau.

 

314. Another--though littlier--bundle of range joy- Velvet bundleflower (Desmanthus velutinus) on a calcareous soil on Texas' Grand Prairie. This species is considerable smaller (hence less productive) than its "ksssing cousin" Illinois bundldflower. On the other hand, this smaller Desmanthus species is better adapted to less favorable environments (eg. those with shallower, rockier, and generally drier soils). These two Desmanthus species frequently grow in close proximity on the same range as here on this Purves-Dugout complex soil mapping unit of a degraded Rolling Prairie range site (Soil Conservation Service, 1973).

Tarleton State University Hunewell Ranch, Erath County, Texas. September. Full-flower phenological stage.

 

315. Another specimen of velvet bundleflower- Another (of several others) plant of D. velutinus with a more upright form that was growing near the specimen introduced in the preceding photograph presented a more advantageous view of the shoot and inflorescence of this fairly small but palatable prairie legume.

Tarleton State University Hunewell Ranch, Erath County, Texas. September. Full-flower phenological stage.

 

316. Flower and fruit bundled on tallgrass prairie- Two legumes nearing maturity on the apex of a shoot of velvet bundleflower while with an inflorescence lower on the same shoot this specimen looks with anticipation toward even more sexual reproduction.

Tarleton State University Hunewell Ranch, Erath County, Texas. September. Full-flower and ripening fruit stage simultaneously on the same plant.

317. Catclaw sensitivebriar (Schrankia uncinata)- This palatable herbaceous legume is in the Mimosoideae subfamily along with numerous “outlaw” woody shrubs that were discussed under the South Texas or Rio Grande Plains savanna type. This decreaser is an indicator species whose presence bespeaks solid stewardship of native tallgrass prairie range. The common name of this palatable and beautiful forb comes from the phenomenon of thigmotropism (= haptotropism), the tropic response of a plant to touch. More precisely it is thigmonasty (nastic movement to touch; nastic movements are those in which the direction of movement is not related to the direction of the stimulus). Whatever it is called the leaflets of the compound leaves fold up under the slightest touch. That condition as viewed here was caused when the plant was inadvertantly brushed 2 feet from the portions pictured.

The sensitivebriar in this shot is co-dominant with big bluestem on a Chert Savanna range site in the Ozark Plateau. Ottawa County, Oklahoma. May.

 

318. Sensitive leaves and purple puffs- Two closer views of catclaw sensitivebriar growing as co-dominant with big bluestem on a Chert Savanna range site in the western Ozark (Springfield). Leaves in the first (upper) slide demonstrated the phenomenon of thigmonasty (a trophic response to pressure of touch) when the dainty leaflets "drew up" when accidentially brushed by a somewhat less sensitive photographer. These "insulted and sulking" leaflets can be compared to "happy ones" that the photographer did not touch.

By the way, these plants did not wait four decades before "coming forward" about inappropriate touching.

The sensitivebriar in this shot is Ottawa County, Oklahoma. May.

319. Flowers to fruits sensitively- Inflorescences (first and second photograph) and legumes (third photograph) of catclaw sensitivebriar. Each inflorescence of this member of theMimosoideae is a head consisting of numerous small flowers with protruding stamens of bright filaments (Smioth, 1977, p. 152). The legumes are covered with stiff, but not sharp spines.

Ottawa County, Oklahoma. Late May for inflorescences; Late June for legumes.

 

320. Wild senna (Cassia marilandica)- This leguminous forb is a member of the Caesalpinioideae, the smallest legume subfamily in North America. This individual is growing on a bottomland prairie in the Ozark Plateau that is dominated by peaked panicgrass (Panicum anceps). This large specimen is almost a yard tall.

Ottawa County, Oklahoma. May.

321. Showing its wild side- Upper shoot (first slide) and inflorescence (second slide) of wild senna growing on tallgrass prairie in the western Ozark (Springfield) Plateau.

Ottawa County, Oklahoma. Early August; peak-bloom stage of phenology.

 

322. Inflorescence and fruit of the wild senna seen immediately above- Ottawa County, Oklahoma. July.

 

323. Nature's bounty of native dry beans- Two shoots of wild senna bearing abundant crops of legumes, the fruit of legumes. The legume fruit type is the common feature that unites the three subfamilies of Leguminosae. These fruit-laden plants were growing at the edge (local ecotone) of a black oak (Quercus velutina)-dominated upland forest and a big bluestem-dominated tallgrass prairie in a more southern part of the Prairie Peninsula that extends into the Ozark (Springfield) Plateau physiographic province.

In contemporary pre-white man times, lightening-ignited and Indian-set fire maintained an understorey of tallgrass species including prairie shrubs like redroot and New Jersey tea (Ceanothus ovatus and C. americanus, respectively) along with native grasses and forbs. Currently, understorey spceis such as these can be found only on small relicts of more-or-less virgin vegetation. These two Ceanothus species were presented under the woody species section of Tallgrass Prairie, Interior.

Ottawa County, Oklahoma. Mid-Jaunary. Fruit-ripe stage.

 

324. Fruits of wild senna- Opened (dehisced) and unopened legumes (first slide) and dehisced legumes showing seeds (second slide) of wild senna. These specimens provided "fruitful" examples of the legume fruit type which is legume. It is the fruit and not the inflorescence that unites the three subfamilies of Leguminosae. Legumes are fruits that dehisce (open) along two sutures (lines of union or fusion and, later, of division for each fruit half) in both longitidunal sides of this dry, unicarpellate (one ovule bearing structure) fruit (Smith, 1977, ps. 67, 291, 299).

Another name for legume(s) is pulse(s). In some strict applications plulse is limited to the dry seeds of legumes used as food or feed (eg. dry beans and peas) and excludes legume seeds used as oil crops (Glycine max, soybean) and entire legumes used as vegetable crops (Phaseolus vulgaris, green beans). In general usage, however, pulse and legume are synomyms.

It is axiomatic that the fruit--and certainly, the seeds--of wild senna is a native concentrate feedstuff of great value to smaller wildlife. Common sense would deem it almost certain that these seeds would be highly palatable and nutritious for birds, especially ground feeding species such as northrn bobwhite quail.

Ottawa County, Oklahoma. Mid-Jaunary. Fruit-ripe stage.

 

325. Two-leaf or twin-leaf senna (Senna roemeriana= Cassia roemeriana)- Most of the senna species are smaller than the large, showy species presented immediately above.

Tarleton State University, Hunewell Ranch, Erath County, Texas. Mid-April, full-bloom phenological stage.

 

Another angle- Topdown view of win-leaf senna growing on the Grand Paairie of northcentral Texas.

Tarleton State University, Hunewell Ranch, Erath County, Texas. Mid-April, full-bloom phenological stage.

 

326. Pretty but poisonous- Single plant and infloresecence (two flowers) of twin-leaf senna. Tarleton State University, Hunewell Ranch, Erath County, Texas. Mid-April, full-bloom phenological stage.

 

327. An annual member of the senna or cassia clan- Showy partridge-pea (Chamaecrista fasciculata= Cassia chamaecrista= C. fasciculata) growing on a recently graded road berm in the western Ozark (Springfield) Plateau. At an earlier time in botanical history this annual member of the tribe Cassieae was included in genus Cassia (Coulter, 1891-1894; Fernald, 1950, p. 886). Later, the specific epithet of this species was "upgraded" to its own genus. Regardless of nomenclatural changes, this "showy" species is, as always, a pioneer plant of disturbed sites or microsites such as road cuts, as in this instance, abandoned farmland (=old fields), cutover forests, and severely overgrazed ranges. In other words, showy partridge-pea is a native weed (weedy annual). There is an ecological twist, however, showy partridge-pea is so valuable as a wildlife feed plant (ideal for northern bobwhite and whitetailed deer [Martin et al, 1951, p. 400; Tyrl et al., 2008, ps. 200-201]), facilitator of plant succession, and aesthetically pleasing wild flower that it is not a weed at all (other than in crop fields). Interestingly, showy partridge-pea can induce digestive upsets in domestic ruminants, but fortunately this occurs only rarely (Tyrl et al., 2008, p. 201).

Given these valuable roles and the prominence of showy partridge-pea on degraded and recovering tallgrass prairies (eg. former sacrifice areas, old fields) this native legume was included with other senna or cassia members. Besides, it is a pretty and cute little botanical delight that is one of the author's favorites so it was included here for numerous reasons.

Ottawa County, Oklahoma. Mid-July, full-bloom stage.

 

A few more- Three more photographs of showy partridge-pea trying to show clearer views with more details that were less botched by an Epsom Perfection 800 scanner. Once this mcchine fouls up images it is all Adobe Photoshop can do to halfway save them. These three were nearly flawles, crisp slides taken using Provia 50 film. Enter Epsom. What the Epsom scanners do is underexpose the slide images so that the level and exposure (and often brightness and contrast) of Adobe Photoshop have to be used to restore the proper light of the original image.

Ottawa County, Oklahoma. Mid-July, full-bloom stage.

 

 

328. Showy resident- A gravid Carolina praying mantis (Stagmomantis carolina) residing on a senescing shoot of showy partridgge pea on tallgrass prairie in a vegetational mosaic of Western Cross Timbers and Grand Prairie in northcentral Texas. This iconic predator of the insect wold was "ready to pounce" on careless prey, although Mrs. Mantis did not seem to be very well hiddden. Perhaps she was in preparation for oviposition (egg deposition, the egg-laying--and final--phase of her life cycle).

These photographs were taken at close range with a macrolense. Mrs. Mantis seemed to be as interested and intent in the glass of the lense as the phtographer was in her. Good luck, ole girl.

Tarleton State University Hunewell Ranch, Erath County, TeEarly September.

 

329. Wild alfalfa or scurfy pea (Psoralea tenuiflora)- This is one of the most widely distributed papilionaceous legumes on North American grasslands. It grows from the understory of the western oak-hickory forest of the Ozark Mountains to the Great Plains shortgrass country. Kelly Prairie, Craig County, Oklahoma. June.

 

330. Flowering shoot of wild alfalfa or scurfpea- Inflorescences of a leguminous forb that is a frequently a local dominant on tallgrass, true, mixed, and shortgrass prairies. This one was in the understorey of an oak-hickory and tallgrass savanna. Ottawa County, Oklahoma. June.

 

331. Wild cluster on top- Dorsal view of cluster of papilionaceous flowers of wild alfalfa or scurfpea growing just outside the fence of an Osage Questas stocker range. The Osage Questas province is adjacent to and east of the better-known Flint Hills physiographic unit. Coffee County, Kansas. Mid-June.

 

332. Tall-bread scurf-pea (Psoralea cuspidata)- One "heap big" specimen of tall-bread scurfpea (over two and a half feet tall and a good yard across) growing in a consociation of little bluestem on a tallgrass prairie range in the Grand Prairie of northcentral Texas. This range was in Good range condition class yet with a rich array of plant species, including a small palnt of Texas pricklypear (Opuntia engelmannii). There were several other tall-bread scurfpea plants on this range, but they were not as impressive as this large and gorgeous individual

Tall-bread scurfpea has historically been classified as a decreaser. McGregor et al. (1986, p. 475) specified that although cattle only rarely grazed tall-bread scurfpea the species did decrease "under grazing". (This obviously begs the question, Grazing by what?).

Tarleton State University Hunewell Ranch. Mid-April; full-bloom phenological stage.

 

333. Scurfing for bread- Upper flowering shoot with entire inflorescence shown (first slide) and details of individual papilionaceous flowers on this inflorescence (second slide) of the large, showy plant introduced in the immediately preceding slide. Everything about this species and this individual plant is impressive.

Tarleton State University Hunewell Ranch. Mid-April; full-bloom phenological stage.

 

334. A snakey species- Sampson's snakeroot (P. psoralioides) on a relict of tallgrass in the western Ozark Plateau. Sampson's snakeroot is a more mesic (hence, more easterly) Psoaralea species. This papilionaceous legume is well-distributed on both tallgrass prairies, oak-hickory forests, and savannahs of (transization zones between) these two formations. Size of leaves of this species compared to the preceding Psoralea species revealed distinctive differences in plant morphology as adaptations to quantity of precipitation and soil moisture conditions through natural selection. Interestingly, however, these two Psoralea species are frequently growing in close proximity on range vegetation of any of the three range types just mentioned. Of course, similar spatial relations exist for Darwin's finches.

Ottawa County, Oklahoma. Late May; peak-bloom phenological stage.

There are several legmes in the Desmodium genus that are known generally by such common names as tick-clover and tick-trefoil. These species are found variously on grasslands, savannahs, forests glades, etc. throughout the general Tallgrass Prairie Region, including mountains like the ancient Ozark Plateau and Arbuckle Mountains. For example, Styermark (1963, ps. 912-925) described 18 Desmodium species for Missouri A few of these species were shown below.

Desmodium species are extremely palatable to livestock and, as such, are much more common on prairies and forests from which livestock have been excluded as well as outside fencerows. Desmodium species on tallgrass prairies in North America are perennial forbs. Desmodium is from the Greek desmos for chain or bond in referrence to the linear or chain-like arrangement of legumes (Fernald, 1950, p. 915). By the way, seeds of Desmodium species have a distinct peanut taste. These species and the domestic peanut (Arachis hypogaea) are in the same tribe, Hedysareae (Smith, 1977, p.153).

 

335. Narrower than most- Narrowleaf tick-clover (Desmodium sessilifolium) growing in the a sandy soil of a tallgrass prairie in the West Cross Timbers of northcentral Texas. First slide was a topdown view of a local colony of narowleaf tick-trefoil. Second and third slides were of a shoot of singlg plant shown as a side view and topdown perspective, respectively.

Narrowleaf tick-trefoil is a widely distributed species with a biological range extending from Massachusetts across to eastern Kansas and in the south from Virginia to east Texas (Fernald, 1950. p. 919).

Erath County, Texas. Late April; late vegetative growth and pre-bloom stages of plant development.

 

336. Not so narrow up here- Inflorescences of narrowleaf tick-trefoil growing on tallgrass prairie in the Springfield Plateau in northestern Oklahoma. The Desmodium inflorescence has traditionally been interpreted as a raceme.

Ottawa County, Early August; late-bloom phenological stage.

 

337. Not narrow flowers- Papilionaceous flowers of narrowleaf tick-clover growing on a tallgrass prairie in the western Ozark (Springfield) Plateau in northeastern Oklahoma.

Ottawa County, Early August; late-bloom phenological stage.

 

338. Stiff, but flexible- Stiff tick-clover or stiff tick-trefoil (Desmodium. rigidum) growing on the sandy soil of a relict tallgrass prairie in the West Cross Timbers of northcentral Texas. These three images obviously presented stiff tick-clover at progressively closer views ending with the apical portion of a single shoot.

Plants shown here were growing in close proximity to plants of narrowleaf tick-trefoil presented above. In fact the biological (= species) ranges of these two Desmodium species are very similar and overlap throughout most of native habitat (Fernald, 1950, p. 920).

Erath County, Texas. Late April; late vegetative growth and pre-bloom stages of plant development.

 

339. Commenerating the Dominion- Canada tick-trefoil (Desmodium canadense) on a relict tallgrass prairie in the Springfield Plateau in northeast Oklahoma. These maximum-height, pre-bloom shoots were presented in a topdown view of a colony (first slide) and as a sideview of a single shoot (second slide).

The uppermost portions of some shoots had been lightly browsed by white-tailed deer (Odocoileus virginianus). The Desmodium species are so palatable to livestock that plants of these species are generally found only on more lightly grazed tallgrass prairies, savannahs, and forests. Specimens seen here were growing in a fencerow that was inacessible to livestock.

This is yet another Desmodium species with an extensive biological range that extends from northeast Canada (NovaScocia) south to West Virginia and west to eastern Oklahoma (Fernald, 1920, p. 921).

At home on prairie or savanna- Hoary tick-trefoil or hoary tick-clover (Desmodium caanescens) is one palatable legume common (on relict sites, not typical abused range or go-back ground) on both tallgrass prairie and tallgrass-hardwood savanna in the Prairie Peninsula first described by Transeau (1935). The specimen presented heree was growing on the outside of a fencerow of an old field in the wesstern Springfield Plateau in northeastern Oklahoma.

Ottawa County, Oklahoma. First slide, early August; second slide, late July.

 

Ticked flower- Two examples of the lovely flower of hoary tick-trefoil growing in a spot protected from livestock and yet lightly grazed by white-tailed deer in the southwestern Prairie Peninsula (Transean, 1935).

Ottawa County, Oklahoma. First slide, late July; second slide, early August.

 

 

340.. Heads to the ground- Characteristic compound leaves and numerous small heads of Nuttall's or little-flower milkvetch (Astragalus nuttallianus) growing on a degraded tallgrass prairie range in Texas' Grand Prairie. There are a "gazillion" Astragalus species over the North American continent, frequently with numerous species over a relatively small geographic area. Furthermore, there are some Astragalus species that include several taxonomic varieties. For example, Diggs et al. (1999) described five--count 'em--varieties for A. nuttallianus. (By the way, this marginally taxonomically literate rangeman did not dare to distinguish which of these five varieties was pictured here.) According to Diggs er al., 1999) the range vegetation of northcentral Texas includes ten Astragalus species.

Tarleton State University Hunewell Ranch, Erath County, Texas. Early April; full-bloom phenological stage.

 

341. Land of milk and ants- Details of papilionaceous flowers arranged in heads of Nuttall's milkvetch on degraded tallgrass grassland of the Forth Worth or Grand Prairie of northcentral Texas. Ants found whatever secretions were produced by these tiny but showy heads to be quite appealing. This was another example of selective grazing (= preferential defoliation) by range animals (selectivity in herbivory).

Tarleton State University Hunewell Ranch, Erath County, Texas. Early April; and no doubt as to full-bloom stage.

In addition to the leguminous forbs there are also a number of woody legumes--both trees and shrubs--on tallgrass prairie. These woody legumes are in all three subfamilies of the Leguminosae and examples of these were shown below in the section devoted to woody plants. This author elected to organize vascular range plants of tallgrass paririe as to grasses, grasslike plants, forbs, shrubs, and trees with woody species arranged as to families under the more general groups as viewed by rangemen, foresters, and wildlifers.

Woody legumes (trees and shrubs of the Leguminosae) were treated below following the Salicaceae.

 

342. No regrets from this photographer- Goat's-rue, hoarypea, Devil's shoestring, or catgut (Tephrosia virginiana) growing on a tallgrass-oak/hickory savanna in the western Ozark Plateau. This papilionaaceous, herbaceous legume is an uncommon forb in prairie communities when compared to most other species (such as those included in this treatment). The author has seen locally abundant populations of goat'srue, but isolated individual plants are the typical pattern of dispersion. This particular plant was photographed periodically over course of a quarter century, and as these slides showed it was still "goin' strong").

Most authorities placed Tephrosia species in their own tribe, Tephrosieae. Goat's-rue does have a beauty all its own. It is also somewhat unique in having high concentrations of rotenone (along with related compounds lincluding tephrosin) in its roots s well as being used as a medical plant by American Indians (Diggs et a., 1999, p. 700). Gard (2010) reported documentation showing that all of the Five Civilized Tribes of Indians in Oklahoma used roots of goat's-rue or Devil's shoestring as a piscicide for fishing. In her own laboratory tests Gard (2010) found that extracts from the T. virginiana rootstocks (woody rhizomes) were toxic to small fish under laboratory conditions.

Newton County, Missouri. Late May; peak-bloom phenology.

 

343. No room for rueful- Sexual or flowering shoots (first slide) and inflorescence (second slide) of the goat's-rue plant shown immediately above. Inflorescences of goat's-rue were described by Diggs et al. (1999, p. 700) as spike-like racemes.This particular plant bloomed each year for over a quarter century on a tallgrass-oak/hickory savanna in the western Ozark Plateau. Its habitat was a relatively shallow, rocky soil on a south slope that it shared with big bluestem, the locally dominant range plant (God's own grass-legume mixture, and both decreasers).

Newton County, Missouri. Late May; peak-bloom phenology.

 

344. Goat's fruit- Upper sexual shoot of goat's-rue bearing legumes (first slide) and details of some of these legumes (second and third slide). Immature, still-developing eeds inside the podlike shell of the legume can be seen due to bright light behind the subjects. This was the same plant as that seen immediately above (though photographs were taken one year apart of a plant that the author had observed over span of a quarter century).

Newton County, Missouri. Late June; intermediate phenological stage of fruit development.

 

345. Little pea on the prairie- Shoots (stolons) with trifoliate leaves (first slide) and closer in view of these trifoliate leaves (second slide) of trailing wild or wooly bean, running fuzzy bean, (Strophostyles helvula or, sometimes, helvola) on a big bluestem-mixed oak (Quercus velutina-Q. marilandica) savanna in the western Springfield Plateau. This annual, papilionaceous, native legume does best on disturbed microsites such as roadsides, recently abandoned cropland, and overgrazed ranges (after livestock have been removed from the abused area). The trailing shoots of these plants can be quite long (in excess of 20 feet is not uncommon) with organs of this ground-hugging legume seldom being more than six to eight inches off of the soil surface, unless the plant is climbing objects like trees or fences as it is wont to do.

(This low growing species makes for a difficult photographic subject. While obtaining these photographs on a road berm the author was laying on his side when a motorist stopped in shock fearing that she had killed a pedestrian. Motorist, her passenger, and the photographer shared a good laugh. One takes his pleasures where he finds them in this line of work.) Pleasant little bean afterall.

This species was previously interpreted as being in genus Phaseolus (P. helvula). It is still taxonomically located in the bean tribe, Phaseolae.

Ottawa County, Oklahoma. Late July; plant with widely varying maturity (ie. freshly opened blooms, tiny legumes, maturing legumes).

 

346. Little flower and little fruit- Clusster of three papilionaceous flowers (first slide) and a single remaining flower with a tiny immature legume (second slide) of trailing wild bean or trailing wooly bean on a local disturbed microhabitat (a county road berm) in a tallgrass prairie-mixed oak savanna. In addition to adjoining cover of Johnsongrass (Sorghum halapense) other neighboring range plants included big bluestem, purpletop, catclaw sensitive-brier, dewberry (Rubus flagellaris) and blackjack oak.

These plant parts were on the specimen introduced in the preceding two-slide/caption set.

Ottawa County, Oklahoma. Late July; plant with widely varying maturity (ie. freshly opened blooms, tiny legumes, maturing legumes).

 

347. Dimunitive flowers and snap beans- Small papilionaceous flowers and legumes of trailing wild bean on a local disturbed area beside big bluestem, catclaw sensitive-brier, dewberry, and and a senescing blackjack oak in the western edge of the Springfield Plateau of northeastern Oklahoma. These organs were on the plant that was introduced two slide-caption units above.

Steyermark (1963, p. 951) described this species as being adapted to habitats varying from rich woods and alluvial soil to gravel bars, fallow farmland, roadsides, and railroad tracks. He described this little bean as doing well on acidic, infertile soils yet enriching them with its nitrogen-faxing capacity (Steyermark, 1963, p. 951). Given that trailing wild bean is in the bean tribe (Phaseolae) and so closely resembles morphologically the beans that it was formerly included in the bean genus, Phaseolus, it is nearly a given that the same species of Rhizobium bacteria inhabits (innoculates) this low-growing wild bean. One such species commonly associated with trqiling wild bean has been shown to be Rhizobium leguminosarum (Lynd and Odell, 1982), but numerous other likely nitrogen-fixing bacterial species include R. etli, R. gallicum, R. giardinii, and R. tropici based on their symbiosis with common bean (Phaseolus vulgaris) (Martinez-Romero, 2003).

Coverage received by this little plant in the botanical literature (even in wild-flower guides) pretty much matches its size: scant, miniscule, overlooked. Some efffort was herewith expended to rectify this oversight.

Ottawa County, Oklahoma. Late July; plant with widely varying maturity (ie. freshly opened blooms, tiny legumes, maturing legumes).

 

348. Little vine on Texas prairie- Little peavine (Lathyrus pusillus) growing on a Sandy range site (Soil Conservation Service, 1973) in the West Cross Timbers of northcentral Texas. This native member of the bean tribe (Fabeae) of the legume family (Leguminosae= Fabaceae) is a rather inconsequential species, but its presence added species and biological diversity to the range plant community of which it was a part. Plus, little peavine undoubtedly is a palatable range plant that has comparatively high nutritive value based on nutrient concentration even though its forage value would be limited by its relative sparcity on the range.

As with trailing wild bean or wooly bean or running fuzzy bean, the previous plant species presented above, little peavine is typically a species of disturbance and/or harsh microhabitats.

Tarleton State University Hunewell Ranch. Early September, late-bloom/early fruit phenological stages.

 

349. Smallish flowers and little peas- Papilionaceous flower and legumes of little peavine (on the same plant as shown in the preceding two slides) growing on a Sandy Range site (Soil Conservation Service, 1973) in the West Cross Timbers of northcentral Texas.

Tarleton State University Hunewell Ranch. Early September, late-bloom/early fruit phenological stages.

The Compositae is the largest family of range forbs. Composites are especially plentiful on grasslands. Some are extremely palatable species (decreasers) while others are ecological weeds indicative of disturbances such as overgrazing and drought (invaders).

 

350. Dense population of compass plant (Silphium laciniatum) at peak bloom- The common name of compassplant refers to the alleged or perceived north-south orientation of the pointed vertical leaves of this composite. This characteristic was elaborated on in captions below.

Frisco Railroad right-of-way, Ottawa County, Oklahoma, July.

 

351. Flowering shoot of compass plant- Typically a decreaser so as to be most abundant on properly managed tallgrass ranges. This hale and hearty specimen of over eight feet in height was growing on a highway right of way. (Note: see immediately succeeding caption for discussion of palatability and response of S. laciniatum to heavy grazing and/or overgrazing.) Osage County, Oklahoma. July.

 

352. Pointing in the same direction- Capitula (heads) of common compass plant at edge of oak-hickory forest and tallgrass prairie in Springfield Plateau of southwest Missouri..

Newton County, Missouri. Late July.

 

353. Compasses on an overgrazed range- Two plants of white-flowered compassplant or white-flowered rosinweed (Silphium albiflorum) growing on a. depleted range on which retrogression was induced primarily by past and present overgrazing. S. albiflorum is a Texas endemic that is commonly abundant on Grand Prairie grasslands of northcentral Texas (Diggs et al., 1999, p.). White-flowered compassplant is generally restricted to calcareous soils such as those of the Grand Prairie. This in contrast to a pronounced absence from the noncareous soils of the often contiguous West Cross Timbers savanna. The biological range of white-flowered rosinweed extends west to the Rolling Red Plains and south into the Edwards Plateau.

Silphium species, especially common compassplant (S. laciniatum), have historically been interpreted as decreasers due to apparent high palatabilities, but this author has observed--over a span greater than a quarter century--that plants of S. albiflorum are almost always untouched on Grand Prairie ranges grazed by cattle and white-tailed deer. Even more relevant to the subject of palatability and persistence of white-flowered compass on Grand Prairie grasslands is the fact that sometimes there appears to be as many--if not more--plants of this species on grazing-degraded ranges. Plants presented in these two photographs grew to peak standing crop and full-bloom stage with zero defoliation even though beef cattle and white-tailed deer had been on this range for quite some time during the growing season. Nor was this the first time for this phenomenon to have existed as was unquestionably attested to by this former little bluestem-Indiangrass tallgrass prairie that had been degraded largely by decades of overgrazing by cattle to Poor range condition class. (Even when universitiy farms and ranches are improperly managed, which is probably about as common as not, there are lessons that can be taught and learnt from such properties.)

Tarleton State University, Hunewell Ranch, Erath County, Texas. Late May; peak-bloom phenological stage.

 

354. Compassed in all directions- The common name of compassplant is often said to have been derived from the "fact" that the leaves of some species have a "supposedly north-south orientation of the leaves" (Diggs et al., 1999, p. 404). As observed on many plants over the years by the author--and as obvious from the first of these two photographsand the second of the two preceding photographs--such is not necessarily the case. The deeply dissected, vertical leaves of white-flowered compassplant point "in ever which a direction". The capitula (head inflorescences) do tend to be somewhat helitrophic or, more correctly, photrophic. Floral phototropism is a response of inflorescences (heads of some composites provide textbook examples) to sun light such that they (their shoots) bend in the direction of prevailing light rays (second photograph). Shoots of white-flowered compassplant from the previous year can often be seen lying or, less commonly, still standing in close proximity to current season's shoots.This condition was presented in the second slide (left margin) of this set .

Tarleton State University, Hunewell Ranch, Erath County, Texas. Late May; peak-bloom phenological stage.

 

355. Compassed beauty-Two Capitula (heads) of white-flowered compassplant. A capitulum (capitula is the plural) or head is an inflorescence that consist of a dense cluster of sessile (or nearly so) flowers, characteristic of the Compositae (Diggs et al., 1999, p. 1428). More specifically, a capitulum consist of small flowrs known as florets that are on a hemispherical shaped (flattened) receptacle known as a disk with this entire structure surrounded by an involucre of bracts (modified leaves) which are known as phyllaries. Florets that are inserted on the disk are disk or tubular flowers whereas flowers surrounding the receptacle (disk) are ray or ligulate flowers. There are three forms of capitula:1) only tubular flowers, 2) both central tubular (disk) flowers and peripheral ligulate (ray) flowers, or 3) only ligulate (ray) flowers (Smith, 1977, p. 241). Fertile florets are either perfect or, alternatively either staminate or pistillate. In certain Compositae species some of the florets are sterile or infertile (Smith, 1977, p. 214). In Silphium species it is only ray flowers that are fertile (disk flowers are sterile).

Tarleton State University, Hunewell Ranch, Erath County, Texas. Late May.

 

356. Compass bearings- Several plants of white compassplant or white rosinweed (Silphium albiflorum) growing on the shallow soil of a degraded Grand Prairie range. Compassplant is one of seven (maybe, eight) Silphium species native to the central prairies and adjoining oak-hickory-tallgrass savanna. White compass plant is one of the smaller, especially shorter, of these Silphium species. White compassplant is also a Texas endemic.

Compassplant (S. laciniatum) has traditionally been regarded as a decreaser so that its reduction or elimination from tallgrass prairie range is an indicator of overgrazing (Nicholson, 2006, p.152), but according to the Great Plains Flora Association (1986, p.1000) compassplant appears to do best "in areas of mild disturbance" such as the degraded range on which these plants of S. albiflorum were thriving. The latter comment was consistent with observations by this author who never saw any evidence of cattle feeding on white compassplant on ranges, including depleted ones such as that shown here, in northcentral Texas. Conversely, in the Cherokee Prairie and Flint Hills of northeastrn Oklahoma (see preceding slides) the author always found plants of S. laciniatum only in protected areas such as highway and railroad rights of way (above slides).

There are conspicuous differences between compassplant (S. laciniatum) and white compassplant (S. albiflorum) in size of mature plant and color of heads. White compassplant has white ray flowers in contrast to yellow ray flowers of compassplant. White compassplant is compartively short (this author never found any individuals of white compassplant that even approached a yard in height) in contrast to the eight-foot giants of compassplant. The most obvious out-on-the-range similarity between these two Silphium specis is reflected in their common names: both have deeply incised vertical leaves with prominent pointed tips and that tend to orient themselves in roughly north-south directions. See next slides...

Hunewell Ranch, Tarleton State University, Erath County, Texas. Mid-September; fruit-ripe to fruit-shatter phenological stages.

 

357. Onward and upward (even in drought)- A mature plant of white compassplant with heads of ripe achenes (first slide) was growing on the thin, caliche-underlaid soil of a degraded Grand Prairie range in northcentral Texas at the end of the driest summer on record. This plant had produced a "bumper crop" of achenes in several heads (second slide) over course of this spring and summer of Exceptional Drought (rating of D4 on the Palmer Drought Severity Index). It is amazing that some of these native range plants survive, both phenotypically as current shoots (genotype) and sexually reproductive as fruit/seed production (gene recombination and creation of new genotypes). On-going natural selection on the range.

The vertical position or orientation of the deeply incised leaves of white compassplant were featured prominently in the first of these two slides. Most references (Great Plains Flora Association, 1986, p. 1000; Hatch and Pluhar, 1993, p. 263; Diggs et al., 1999, p. 404; Nicholson, 2006, p.152) have remarked as to the general north-south orientation of leaves of S. laciniatum. Close similarity of leaves between S. laciniatum and S. albiflorum are basis of the common names of compassplant.

Hunewell Ranch, Tarleton State University, Erath County, Texas. Mid-September; fruit-ripe to fruit-shatter phenological stages.

 

358. Descriptive leaves- Basal leaves of white compassplant growing on thin, calcareous Grand Prairie soil. Lower leaves of this species point outward from the plant. Upper leaves, by contrast, point upward or are in a vertical position (first of two immediately preceding slides). Leaves of white compassplant offered an opportunity to illustrate the rich terminology of Taxonomic Botany. There is no shortage of descriptive terms for plant leaves that include everything from general outline of simple or compound leaves to leaf bases, tips or apices, veins (veination), margins, surfaces, colors, and sizes.

These leaves of white compassplant could be described as simple (not divided into leaflets), lanceolate (tip), palmately divided (general palm-shaped leaf that was deeply incised [nearly to midrib]), and scabrous (rough--like sandpaper--surface). Leaves of white compassplant (S. albiflorum) do not seem as strongly oriented in a general north-south direction as are those of compassplant (S. laciniatum). Distinctive (and descriptive) leaves nonetheless.

Hunewell Ranch, Tarleton State University, Erath County, Texas. Mid-September.

 

359. Rosin' up the bow (or something)- Rosin-weed (Silphium integrifolium) is a large, rank-growing (coarse), perennial composite that has numerous shoots arising from a short rhizomatous base (frequently the case) or a caudex (always the case). A caudex is a tough, rough, persistent base of an herbaceous plant (Great Plains Flora Association, 1986, p. 1318) or the root-shoot axis often termed proaxisis.

Rosin-weed is a characteristic composite of bottomland habitats. The specimen exhibited here and in the immediately following slides and caption set was growing in asociation with eastern gamagrass on the floodplain of a typical Ozark Plateau stream. The author watched this plant grew progressively larger each warm-growing season over a period of three decades with each of this year's clumps having more shoots and extending with a larger cover (basal and foliar) over more of the rich soil surface. Rosin-weed has to be a decreaser. The only plants this author has ever seen are either in fencerows, ungrazed relict areas, or well-managed prairie hay meadows.

There are several Silphium species, and all are large plants with exception of the Texas endemic, white compassplant (Silphium albiflora), featured immediately above. Essentially all of these Silphium species are members of the climax tallgrass prairie community in which they are conspicuous forbs (due to large size, unique leaves, and showy yellow heads.) . Common compassplant (S laciniatum),is one of the larger-growing Silphium species that is at home on tallgrass prairies. This is why that species was shown above just before S. albiflora herein. Prairie dock (S. terebinathinaceum) .is another forb of prairie, savanna, and woodland. Prairie dock is quite common and diagnostic of true prairie such as that developing on chert and dolomite glades in the Ozark Plateau. Prairie dock was featured with these glades in the Miscellaneous Grassland chapter.

Cup plant (S. perfoliatum) is another large, rank-growing sunflower-like forb in this group. Cup plant is typical of range plants that grow on the edge or ecotone between oak-hickory forests and tallgrass prairies. Cup plant plant was featured in this publication in the chapter entitled, Tallgrass Savanna while, again, prairie dock was included with glades grasslands in the chapter, Miscellaneous Grasslands.

These Silphium species constitute an ecological guild, "a group of species all members of which exploit similar resources in in a similar fashion" (Allaby 1998). The ecological guild has sometimes been compared analagously to a set of box or open end wrenches or stringed instruments within the violin group. Avain guilds include groups of raptors or woodpeckers. The bluestem grasses (Andropogon spp.) and cordgrasses (Spartina spp.) are good examples among the Gramineae. Silphium species provide a fine example of a forb guild.

Ottawa County, Oklahoma. Late July.

 

360. Rosined heads- Capitula (heads) of rosin weed on a bottomland range site in the western Ozark (Springfield) Plateau. "Believe it or not" these flowers were produced in Severe Drought (Palmer Scale). Being on a bottomland range site there was more available soil moisture, plus native plants do not show experience drought stress (or show drought stress symptoms) as soon or to the pronounced degree observed in domestic field crops.

Ottawa County, Oklahoma. Late July.

Sunflowers (Helianthus species) perhaps characterize forbs on tallgrass prairie better than any ther genus of the Compositae. Some of the sunflower species also produce some of the largest of all herbaceous plants of the tallgrass association. Helianthus species are impossible to miss on this marvelous North American grassland. A few of these that are part of the "chrismatic megaflora" were presented below.

A common and very important forb- Whether viewed from perspectives of popular culture as the State Flower of Kansas (or the National Flowre of Ukraine) or of commerce as one of the world's major oilseeds annual sunflower (Helianthus annuus) is an important plant. As a native range plant, common or annual sunflower is one of the most widely distributed of all the prairie forbs. The natural biological range os helianthus annuus clearly included most of the whole of the Americas. The original species range of common sunflower would be impossible to determine given that the species unquestionably has expanded its current biological range following its human-assisted expansion, first, as a common weed and, second, as domesticated field crop. Both extremes are a classic example of adaptation to--perhaps some degree of co-evolution with--man the farmer. Annual sunflower was domesticated by American Indians in Mesoamerica (along with so many other major crop species). The current range of annual or common sunflower is planet-wide or global.

Across ther ranges of North America common sunflower is frequently abundant on all the central grasslands from tallgrass to semidesert grassland. Annual sunflower is "at home" in all North American deserts where it sometimes attains heights of over six feet. In North America common sunflower grows as far north as the Northwest Territories of Canada. Common sunflower does have one stipulation: it does not tolerate shade. Even with that habitat limitation annual sunflower often thrives in forest communities following denudation due to clearcutting, forest fires, wind or ice storms, and disease outbreaks. It is, of course, on the great grasslands where common sunflower is so "common" (sorry about that). This robust annual is particularily abundant on tallgrass, true, and mixed prairies.

Common sunflower is currently used for bioremiadition, specifically phytoremediation, as well as a field crop where uses range from oilseed to bird feed to biodiesel. A good reference for this is the agronomic monograph edited by There is no equivalent treatment for H. annuus as a native plant, but all the floras (manuals) and "wild flower" field guides covered it. It seemed strange to this author that common sunflower was not included in the Phillips Petroleum series, Pasture and Range Plants nor was this species on the Society for Range Management "200 List" (the Master Plant List for the International Range Plant Identifiction Contest) or in the accompanying volume, North American Rnage Plants (yet the SRM included common dandelion [Taraxacum officinale]). In fact, there is not one Helianthus species on the SRM Plant Identification List (Stubbendieck et al., 1992). Damned dandelion and no sunflower? I'd be for changing that situation, pronto!

Hatch and Pluhar (1993) did include Helianthus maximilliani in Texas Range Plants, but still no H. annuus. Leave it to the sod-busting "dirt farmers" to get one up on the rangemen. Grazing animals, including grass-preferring cattle, readily eat annual sunflower. Based on this rangeman's observations, cattle find the leaves of common sunflower particularily palatable.

Common sunflower, as befits annuals, is a colonizer or pioneer species. Some might snort "weed", but this applied ecologists would quickly (and correctly) point out that under secondary plant succession the processes of colonization or, the complete phase, invasion is an essential part of natural revegetation. In such cases, common sunflower is no more a weed than it is when used as an oil cropof phytoremiator. Common sunflower is part of the "crop" in early seral stages of natural revegetation through "old field" succession. Jayhawkers got it right;: annual sunflower was a fitting choice for state flower of a prairie state.

There is tremendous diversity (in size and shape of plants as well as in leaf, stem, and flower features) in H. annuus. "This is a huge, polymorphic complex encompassing numerous wild and weedy races, plus the tall, strict, large-headed cultivated phases" (Great Plains Flora Association, 1986, p. 954).

Bank of Colorado River, Runnels County, Texas. Mid-October; full-bloom to early fruit stage.

 

361. Composite- The flower cluster or pattern of flower arrangement as a whole of the Compositae (sunflower or aster family) is a composite affair, the inflorescence type being a head or capitulum. Smith (1977, ps. 63, 291) described the capitulum as "a dense spherical or rounded inflorescence of sessile [unpediceled] flowers" or "an aggregation of + sessile flowers on a common receptacle". More specifically in case of the Compositae the head consist of numerous small flowers (florets) seated within a common receptacle or disk that is conical, hemispheric or flattened in general shapewith this entire unit encircled gy an invloucre of braxcts termed phyllaries.Phyllaries are arranged in one or more series or in overlapping fashion. The actual flowers of composites (as all members of the Compositae are known) can be of one or two types: 1) disk or tubular flowers which are inserted in the disk (ie. the central part of the inflorescence) and 2) ray or ligulate flowers which surround or encircle the disk and are inserted around the perimeter of the common receptacle. Generally, inflorescences of the Compositae can be of both tubular (disk) and ligulate (ray) flowers or only ray (tubular) flowers. Ray flowers can be of two types. This confusing composite situation was explained by Smith (1977, ps. 214-217).

In tribe Heliantheae (of which Helianthus species are members) capitula can be composed of either: 1) tubular along with one type of ligular (disk) flowers or 2) strictly tubular (ray) flowers. Helianthus species have only one series of ray (ligulate) florets which are pistillate but infertile (yellow in color) while the disk (tubular) florets are prfect and fertile (hence the only source of fruit). In H. annuus these disk florets vary from shades of red to purple to, sometimes, yellow (Great Plains Flora Association, 1986, p.s 953, 954). The fruit of Helianthus species is an achene (a dry, indehiscent, single-seeded fruit).

Bank of Colorado River, Runnels County, Texas. Mid-October; full-bloom to early fruit stage.

 

362.. Various stages- Heads of common sunflower in various phenological stages. In the first slide flower-fruit (maturity) stages were full-bloom, early fruit filling, and fruit shatter stages right to left and front to rear. In the second slide the head at peak flowering stage was to right and front of a head at fruit-shedding stage. This complete array of head maturation was perhaps attributable to recent recovery of sunflower plants after heavy rains broke a summer-long Exceptional Drought (Palmer Drought Severity Scale rating of D4). Native range plants are survivors.

Bank of Colorado River, Runnels County, Texas. Mid-October; full-bloom to early fruit stage.

 

363. Beautiful colony of Maximilian sunflower (Helianthus maximiliani or H. maaximilianii) on a tallgrass prairie range- This species and compass plant (Heliantheae tribe of Compositae) are decreasers that are so palatable even to Bovidae ruminants that they can usually be found only in ungrazed places like rights-of-way and fence rows or on properly grazed ranges. Given forage value and value as an ecological indicator numerous examples of Maximilian sunflower were furnished below.

Wichita Mountains Wildlife Refuge, Commanche County, Oklahoma, October.

 

364. Beauty in fecundity- A robust, multistemed Maximilian sunflower (first slide) and numerous capitula (flowering heads) on this same plant (second slide) on an ungrazed range in Texas' Grand Prairie. This many shooted (vertical shoots or tillers) individual produced this crop of beautiful flower heads under conditions of Extreme Drought (Palmer Scale). Even under severe stress this plant was a particularily fecund specimen.

Fecundity (state of being fecund) refers to the capacity to reproduce abundantly or having extreme fertility as measured by numbers of offspring. Annual plants are generally more fecund than perennials from standpoint of sexual reproduction (ie. as measured by quantity of fruits, seeds, or genetically recombined progeny). The more typical asexual reproduction in plants is, of course, proliferation of shoots such as tillers, stolons, rhizomes, tubers, etc.

The Maximillian sunflower plant seen here had excelled at both asexual and sexual reproduction and, again, under very trying times. The ole natives can just survive. Survival in an ecological/evolutionary context is natural selection as measured solely by successful reproduction relative to demands of environment. This is fitness: capacity to perpetuate the species or continue to occupy its ecological niche. This palatable (decreaser) forb is a survivor.

Erath County, Texas. Late October.

 

365. Pattern of production- Flowering stalks of Maximilian sunflower arising off of a branch of the main shoot (ie. head inflorescences were on secondary branches that grew off of this primary branch which was an outgrowth of the main stem). Maximilian sunflower is a decreaser that is extremely palatable even to grass-preferring grazers like cattle and horses. Better yet, it is also remendous forage producer and not some dainty forb that is an ince cream species. This plant showed the branching pattern of which this species is capable under ideal conditions which, in this instance, was on an ungrazed wet prairie (subirrigated).

Erath County, Texas. Mid-October.

 

366. Inflorescences of Maximilian sunflower- Several heads of Maximmilian sunflower on a single shoot of this rank-growing species that is capable of reaching immense sizes.

Wichita Mountains Refuge, Commanche County, Oklahoma..

 

367. Crown of a king composite- Head (capitulum) of Maximilian sunflower. Capitulum (plural, capitula) is the inflorescence type that has several to numerous sessile flowers inserted into a common receptacle or disk (Smith, 1977, ps. 63, 214, 291). Although this species (of tribe Heliantheae) has outer petals these are infertile pistillate flowers. The disk or tubular flowers are perfect and fertile (Diggs et al., 1999, p. 368) producing abundant crops of achenes, the fruit type of the Compositae.

The arthropod on a petal was a female goldenrod crab spider (Misumenia vatia), Thomisidae. One of the distinctive features of this species is the camoflage cability that enables spiders to change from white to yellow and vice versa (Milne and Milne, 1980, p. 907; Marshall, 2006, p. 606). A detailed explanation of this phenomenon, including the pigments involved, was reported by Wikipedia (Google "goldenrod crab spider")

Erath County, Texas. Mid-October.

 

368. Another head and a male arachid- Details of a capitulum (head type inflorescence) of Maximilian sunflower with a tiny--though adult-size--male goldenrod crab spider. No, not a range romance here. The female crab spider (immediately preceding slide) and male crab spider (these two slides) were on different ranges in different years. They did make good screen co-stars, however, and no messy silver screen divorce or marital infidelity.

Heads of members of tribe Heliantheae have only tubular or both tubular and ligulate flowers (Smith, 1977, p. 216-217); Helianthus species have sterile (infertile), pistillate ray flowers (Diggs et al., 1999, p. 368).

Erath County, Texas. Late October.

 

369. Giant forb- Local stand of sawtooth sunflower (Helianthus grosseserratus) with gigantic hoots some of which reached heightsgreater than 11 feet and most of which at least 9 feet. H. grosseserratus is extremely vriable in both size and morphological features, but it is abundantly rhizomatous so that colonies comprised of one to only a few genotypes (genetically unique plants). Phenotypic plasticity is also considerably high. For example, the colony shown here and including parts of two shoots presented in the immediately succceeding two photographs grew on the edge of a deep highway barrow ditch on a very shallow, rocky soil in the western Ozark Plateau. Othe individual plants and local colonies of sawtooth sunflower growing on deeper, more fertile soil within one or two miles radius from the plants shown here were smaller. Obviously it was quantity of available water that explained this difference.

In eastern extremes of the biological range of sawtooth sunflower it hybridizes or integrades with giant sunflower (H. giganteus). Names notwithstanding, H. giganteus attains to smaller size and stature than H. grosseserratus (Fernald, 1950, p.1494 ).

Ottawa County, Oklahoma. September; peak-bloom phenological stage.

 

370. Giant shoots of a fiant forb- Terminal (distal) portions of two shoots of sawtooth sunflower. Although heights of both of these shoots exceeded 10 feet they were so "top-heavy" with heads (capitula) and leaves that they were bent to within two feet of ground level. These shoots were in the colony of sawtooth sunflower introduced in the preceding slide.

Ottawa County, Oklahoma. September; peak-bloom phenological stage.

 

371. More leaves and heads of giant forb- Shoots of sawtooth sunflower of a plant growing on rich alluvial bottomland (of Modoc Creek). A comparison of leaves (and even capitula) on this plant with shoots of plant(s) growing on a rocky, shallow soil (two preceding slide-caption sets) provided an example of the variation in this species. Genotypic, phenotypic, and edaphic sources of variation were all in play in examples seen here.

Ottawa County, Oklahoma. September; peak-bloom phenological stage.

 

372. In name only- Patch of Jerusalem artichoke (Helianthus tuberosus), that is about as far from the Holy City as is possible. Nor is this sunflower (also known as sunroot or sunchoke) the artichoke of commerce which is specifically the globe artichoke (Cynara cardunculus), a major (one of many) field vegetable crop in California. Both "artichokes" are, however, perennial composites. Jerusalem artichoke (origin of the adjective is apparently obscure) is a native of North America (the globe artichoke hails from the Mediterranean Region) and is generally regarded as one of only two agricultural plant crops cultivated in the United States of America that was indigenous to soil that is today that of the USA. The domestic strawberry (Fragaria sp[p].) is the other plant species (combination of several species) native to USA soil, but the commerical strawberry is interpreted as a hybrid in which the native North American strawberry (F. virginiana) was only one contributing species. Thus, Helianthus tuberosus remains the only plant species qualifiying as a field crop that is native to soil that today is part of the United States.This species--like those of maize or "corn", potatoes, squash, punpkins, beans, tobacco among many others--is complements of the dispossed American Indian (mostly tribes of Meso America and South America).

According to Owensby (1980, p. 79) the proper noun, Jerusalem, was derived from an Italian term, girasole, meaning sunflower which itself (as well as the generic name, Helianthus) was derived from the tropic response of these species to turn toward the predominant general direction of direct sunlight .

Readers were referred to Wikipedia for detailed discussion of Jerusalem artichoke, especially brief mention of the "bubble" of this crop. The author was unable to find studies concerning palatability to range animals of shoots of H. tuberosus. The tubers are obviously of considerable nutritional value to certain animals including humans and hogs (see below).

Vegetable garden of D.E. Waggener, Erath County, Texas. September; peak-bloom phenological stage.

 

373. General features of sunchoke- Views of shoots of H. tuberosus showing overall characteristics of leaves and heads (flower clusters) of Jerusalem artichoke. One of the key features--though not readily visible in these photographs--is the semi-winged petioles. First photograph, Newton County, Missouri; second photograph, D.E. Waggener vegetable garden, Erath county, Texas. All plants at peak bloom.

 

374. Crop of Jerusalem artichoke or sunchoke- Tubers of H. tuberosus immediately after digging. While not visible when belowground (except to Xray-vision Superman) the tubers of this sunflower are the most uniquely distinctive feature of this Helianthus species. The principal carbohydrate stored in these belowground shoots is inulin rather than starch. Inulin is largely indigestible in the nonruminant gastrointestional tract prior to site of absorpiton (primarily duodenum) so that energy from inulin is unavailable for most monogastrics like humans and hogs. Inulin is degraded and its energy released by bacterial degradation in the hindgut. In nonruminants with large cecums (eg. equines) much of the energy released from inulin is absorption through the cecal epithelium.

Nutritional as well as agronomic and ecological aspects of Jerusalem artichoke are as interesting as this species is unusual. The definitive source for such knowledge and general treatment of this fascinating plant is the monograph by Kays and Nottingham (2008).

Vegetable garden of D.E. Waggener, Erath County, Texas. October; full-grown tuber stage.

 

Immense colony- A gigantic colony of ashy or, less commonly, soft sunflower (Helianthus mollis). This comparatively vast population was growing on the virgin sod of a tallgrass prairie used as a hay meadow for decades. In western Springfield Plateau in a moist year. Ashy sunflower is one of the shorter, smaller (but no less beautiful) and one of the more palatable Helianthus species.

Ottawa County, Oklahoma. Early August.

 

A purty and a tasty one- Shoots and heads of ashy sunflower. This is one of the more palatable Helianthus species. It is a decreaser on tallgrass prairie. These plants were growing on a wet prairie in the western edge of the Ozark (Springfield) Plateau.

Ottawa County, Oklahoma. Late July.

 

375. Brown eyes on prairie or savanna- Brown-eyed Susan (Rudbeckia triloba) growing on a tallgrass-oak savanna in the western Ozark Plateau. This species is far more common on more moist, often somewhat shaded, habitats though it is frequently found on tallgrass prairie. For this reason and to include Rudbeckia species together R. triloba was included here.

Stoney Point Savanna, Ottawa County, Oklahoma.

 

Close relaive, but darker-headed- Head of black-eyed susan (Rudbeckia hirta) which is generally more common than R. triloba that was presented immediately above. This one was growing on a disturbed tallgrass prairie, during a wet spring, where it was being used as a layover stop by two orthopterans: a grasshopper (left) and a katydid (right). Everybody from black-eyed susan to visiting insects seemed happy and contennted, afterall there was plenty of soil moisture (for now).

Ottawa County, Oklahoma. Early June.

 

376. A showier Susan- Shoots of showy brown-eyed Susan (Rudbeckia speciosa= R. fulgida var. speciosa) on tallgrass prairie in the western Springfield Plateau portion of the Ozark Plateau in southwestern Missouri.

This Rudbeckia species is generally less common than brown-eyed Susan (R. hirta), being more prevalent on moister habitats (Steyermark, 1963, ps. 1558-156).

Diamond Grove Prairie, Newton County, Missouri. Early August; peak-bloom phenological stage.

 

377. Showy details- Upper flowering shoots and a capitulum (head) of showy brown-eyed Susan growing on tallgrass prairie in the western Springfield Plateau portion of the Ozark Plateau.

Diamond Grove Prairie, Newton County, Missouri. Early August; peak-bloom phenological stage.

 

378. Someone else appreciated the composites- A female praying mantis (Stagmomantis carolina) in the folage of a plant of showy brown-eyed Susan (slides two, four, and five of this set) and great ironweed (slides one and three of this set) on a tallgrass prairie in southwest Missouri, at the edge of the Ozark Plateau.

This particular insect flew (yes, the Carolina praying mantis is an apt flyer) back and forth between a plant of showy brown-eyed Susan and one of great ironweed as the photographer placed his normal/marcolens within inches of her. Apparently, she either tired of flying away from the persistent photographer or figured out the camera meant her no harm so that flying away was not worth while (and, ineffective anyway as the camera lens kept coming).

By the way, numerous insects habituate to the camera in relatively short order. This is true for butterflies and bees as well, as can be seen from other slides in this section.

Diamond Grove Prairie, Newton County, Missouri. Early August; peak-bloom phenological stage.

 

379. One that benefits from abuse- Prairie, lanceleaf,or yellow Indian-blanket (Gaillardia aestivalis = G. lanceolata= G. fastigiata) growing on degraded tallgrass prairie in the Grand Prairie of northcentral Texas. This example was growing in its more common habitat, a disturbed microsite (and on deep sand).

Yellow Indian-blanket is typically an annual or short-lived species. This species has no trouble in replinishing its kind, especially on overgrazed ranges.

Tarleton State University Hunewell Ranch, Erath County, Texas. Early September; peak-bloom stage of phenology.

 

Two spring composite cousins- Engelmann daisy (Engelmannia pinnatifida) at left and Mexican hat or upright prairie coneflower (Ratibida columnifera) at right on a Grand Prairie range in northcentral Texas. Both composite species are in the same tribe, Heliantheae.

Tarleton Staate University Hunewell Ranch, erath County, Texas. Laate March.

 

380. Engelmann daisy (Engelmannia pinnatifida)- This is another of the composites that is very palatable to grazing animals, including grazers like cattle, that usually prefer grass over forbs. Engelmann daisy has generally been categorized as a decreaser and, though here are other range plant species that experienced observers regard as being more palatable, Engelmann daisy does decline quickly on overgrazed ranges. It's presence on a range indicates that grazing has not been improper very often or very long.

Engelmann daisy frequently grows in colonies such as the one shown here, especially in areas like central Texas renowed for it's wild flowers. Engelmann diasy blooms very early in the spring and can arguably be viewed as a cool-season species.

Erath County, Texas. (This example was growing in a protected location in the Grand Prairie.) April.

 

381. Shoots from a sideview- Engelmann daisy shown from a sideview to present overall habit (first slide) and upper portions of sexual shoots at peak flowering stage (second slide) for floral details. Diggs et al. (1999, p. 348) described Engelmann's daisy as an "ice cream plant", a species that is extremely palatable to some kinds of animals such that forage of these plants is often overused even under otherwise proper grazing management resulting in loss or reduction of such range plants.

Engelmann's daisy can grow to comparatively large size (eg. upto four foot heights or more) under favorable conditions expecially with moist to wet soil. Engelmann's daisy is a cool-season species.

Tarleton Stte University Hunewell Ranch, Erath County, Texas. Mid-April; peak-bloom stage of phenology.

 

382. Study in Engelmann heads-Composite capitula (heads) of Engelmann ('s) daisy. Heads of Engelmann's daisy have both disk and ray flowers. This is one of many species in the immense tribe Heliantheae.

Tarleton Stte University Hunewell Ranch, Erath County, Texas. Mid-April; peak-bloom stage of phenology.

 

383. Late bloomer- Specimen of Engelmann daisy shown as to details of shoot (first slide) and two less-common forms of capitula (second slide). The rays of this species sometimes curl under in the fashion presented here, but more commonly they are fully extended as in typical sunflower or daisy (composite) habit (compare to above slides).

This specimen was growing at the western edge of tallgrass prairie in early summer where its dominant neighbors were little bluestem and Indiangrass. This time of blooming is a full two months later than in northcentral Texas, for esxample, where Engelmann daisy is a spring-blooming forb.

Black Keettle National Grassland, Roger Mills County, Oklahoma. Early July.

 

384. No fever here- American feverfew or wild quinine (Parthenium integrifolium= P. integrifolium var. integrifolium= P. hispidum). There are a number of Parthenium speciess in North America ranging from annual forbs to desert shrubs. This is still yet another "proud member" of the Heliantheae, a giant of a tribe. This is a very picturesque tallgrass-oak-grass savanna forb. It is usually quite uncommon and a rare pleasure to find such robust specimens as this one.

Newton County, Missouri, Late May; peak flowering phenological stage

385. Fewer fevers- Large--though drought-stressed--plant of American or eastern feverfew or wild quinine (Parthenium integrifolium). This composite forb is one that is usually sparsely populated yet widely distributed with a species range from the Middle Atlantic States south to Georgia and Texas. Eastern feverfew is found in remaining native vegetation of grasslands, especially glades, and savannas across Midwestern States like Missouri. The large specimen (a little shy of five foot in height) seen here was on a chert glade in the western Ozark (Springfield) Plateau. It had grown to this robust size and scale in the second summer of Exterme Drought (Palmer Index).

The ommon names of feverffew and wild quninine are in reference to early use of concoctions made from the flowers of this species for treatment of fevers, especially recurrent fevers such as those from "fever-ague" (malaria).

Newton County, Missouri. Early July; peak standing crop and peak bloom stage.

 

386. Fever fewers- Flowers of American feverfew or wild quninine. This member of the huge Heliantheae tribe has fertile flowers (achenes) only as ray flowers, the disk flowers being sterile. Apparently it was the entire inflorescence that was used by Indians and frontiersmen as a folk medicine (effective or not) for fever, especially sporatic fevers as in "fever and ague" (malaria).Today, eastern feverfew is enjoyed as a large, picturesque prairie forb, and one that has obvious value for landscaping with native plants.

These flowers were onthe same plant only two years apart, the second or lowere slide being taken two years subsequent to the firs or upper slide.

Newton County, Missouri. First slide: Early July; peak flowering phenological stage. Second slide: mid-June; peak flowering phenological stage. (Note the two week difference in peak flowering on the same plant from one year to the next; however the "past-their-prime" heads in the upper slide were obviously several days older than the newly bloomed capitula in the lower slide.)

 

387. Pointed leaves and tick seeds- Lanceleaf (narrowleaf) ticksseed or tickseed coreopsis (Coreopsis lanceolata) growing on a tallgrass hay meadow in the Springfield Plateau. C. lanceolata is one of six Coreopsis species native to the tallgrass prairies on the Osage Plains section of the Central Lowlands physiographic privince (Steyermark, 1963, p. 1580-1584). The undivided leaves that clasp the stem mostly in pairs provides ready field identification of this species.

Ottawa County, Oklahoma. Early June-full-bloom phenological stage.

 

388. Ah chew!- Sneezeweed or purple (purple-headed) sneeezeweed (Helenium flexuosum) on a dry-mesic chert tallgrass prairie (Nelson, 2010, ps. 281-284) in the western Ozark (Springfiield) Plateau. This pernnial member of the Heliantheae tribe prefers moister habitats on tallgrass prairie and tallgrass-oak-hickory savanna. The yard-plus tall specimen seen here was growing at the edge of a patch of hardwood trees and shrubs that had invaded an otherwise virgin tallgrass prairie. This prairie had been burnt about three to four months earlier resulting in topkill of many of these hardwood species. Several plants of purple sneezeweed were growing robustly at edge of hardwood patches and rest of the bluestem-prairie dropseed climax community.

This author could not figure out where the common name of "sneezeweed" came from. It did not affect him thusly. Even more pertinent was basis of the adjective "purple" and "purple-headed". The author never found any plants of H. flexuosum with heads (capitula) that were other than brown in color. Some common names are quite descriptive while others seem to have been based on old wives tales (or maybe their colorblind husbands).

Diamond Grove Prairie, Newton County, Missouri. Late June; peak-bloom phenological stage.

 

389. Bless you- Single plant of purple sneezeweed (first slide) and entire flower cluster of same plant (second slide) on the virgin sod of a chert tallgrass prairie in the western Ozark Plateau. The specimen featured in these two photographs was a different plant from the one in the preceding slide, but it was growing less than its height away from that first individual.

This auuthor remarked in the preceding caption that he had never seen a capitulum of "purple-headed" sneezeweed that was other than brownish or tannish in color. Perhaps "purple" sneezeweed is in the same category as the "purple cow" of children's poety (Gelett Burgess, 1895). The specific epithet flexuosum is fitting as this limber, top-heavy (at flowering stages) composite bends slightly over and sways with mild breezes.

H. flexuosum (by whatever common name) can be seen in local mesic sites (moist to relatively wet) microhabitats on virgin tallgrass prairie as shown here, but it also grows on overgrazed pastures (including those planted to agronomic pasture grasses). Other Helenium species have been reported and verified as being toxic to livestock (Kingsbury, 1964, ps. 409-414), but not H. flexuosum.

Diamond Grove Prairie, Newton County, Missouri. Late June; peak-bloom phenological stage.

 

390. A bitter one- Bitterweed or common or bitter sneezeweed (Helenium amarum var. amarum= H. tenuifolium) on the edge of a severely overgrazed Deep Sand range site in the West Cross Timbers of northcentral Texas.This species has an interrupted biological (species) range over the eastern half of North America. It is also a species of extremely low palatability yet one shown to cause livestock poisoning problems. Those interested in animal toxicity were referred to Burrows and Tyrl (2013, ps. 177-186 passim).

Late October; peak bloom stage.

 

391.. Bitter bloom details- Upper shoot (first slide) and details of capitula or heads (second slide) of bitterweed or bitter sneezeweed. These organs were on plants growing overgrazed range along the edge of a Deep Sand range site in northcentral Texas. This species is toxic, but typically is extremely unpalatable. In addition to Burrows and Tyrl (2013, ps. 177-186 passim) and Kingsbury (1963, p.412-413) a good reference for livestock producers in Texas was Hart et al. (2003, ps. 100-101).

Erath County, Texas. Late October; peak bloom stage.

 

392. What a stand!- An immense, expansive population of pale coneflower (Echinacea pallida) that had developed on a disturbed tallgrass praire in the Springfield Plateau in southwest Missouri. Parking and moving of heavy equimpemtn during road construction left this disturbed area on which a "nearly pure" stand (almost all basal and foliar cover) of pale coneflower had developed. Successional status of pale coneflower is not consistent, but generally it is a member of the climax plant community although it can become established earlier on a prairie sere and then persis into the climax range vegetation.

Jasper County, Missouri. Mid-June; peak standing crop and full-bloom stage of phenology.

 

393. An example of its species- One (or two growing inseperably close together) plant(s) of pale coneflower in the vast population presented in the immediately preceding photograph. One of the major Compositae tribes is Heliantheae to which Echinacea species belong. Pale coneflower is one of the most characteristic forbs on tallgrass prairie communities that are in pristine, virgin, or relict state (climax vegetation; Excellent range condition class).

Pale coneflower has to have relatively high palatability because it is sparse to non-existent on overgrazed ranges while being locally abundant on properly managed tallgrass hay meadows. The author can recall boyhood experiences of feeding prairie hay and finding plants of pale coneflower in the bales. The cows cleaned them up, too (there again, those cattle could not afford to leave much uneaten).

Jasper County, Missouri. Mid-June; peak standing crop and full-bloom stage of phenology.

 

394.. Another example- A single plant of pale coneflower with its last heads in bloom (first slide) and close-up views of heads (second or lower slide) on tallgrass prairie in the Ozark (Springfield) Plateau of southwestern Missouri. In addition to being a common forb on climax tallgrass prairie (hence an indicator species of proper grazing or hay management), pale coneflower is much prized as a native wildflower for landscaping (prairiescaping) purposes.

Newton County, Missouri. Mid-June; full-bloom stage.

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