Tallgrass Savanna - IB

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204. Representative of another woody family- Rhamnaceae (the buckthorn family) is represented on oak-hickory forest-tallgrass prairie savannas by New Jersey tea (Ceanothus americanus). This species is much more common on tallgrass prairies than in the understorey of forests, but it is locally commonly in both and the savannahs between these two range types. by way of example, this specimen was growing at edge of a tallgrass parairie and a black oak-dominated forest in the western border of the Ozark Plateau.

Ottawa County, Oklahoma. Early June; peak bloom.

 

205. Bristly greenbriar, catbriar, bullbriar, hellfetter, and a host of other common names (Smilax tamnoides var. hispida)- Inflorescences and leaves of bristly greenbriar. The parallel veins that are characteristic of monocots were obvious in this photograph. An indication of the importance of bristly greenbriar in the oak-hickory forest and oak-hickory-tallgrass savanna is provided by the fact that this is the most common and widely distributed Smilax species in Missouri (Steyermark, 1963, 450-454). None of the Smilax species are as apparently common throughout the region of these major range plant communities as woody vines like poison ivy and Virginia creeper, but Smilax species are locally more common in and typical of woody climbers in the Cross Timbers.

This specimen was found in the western edge of the Ozark Plateau in the post oak-black hickory-big bluestem dominated savanna that is a transition between the oak-hickory forest and tallgrass prairie grassland of the Cherokee Priairie section . Ottawa County, Oklahoma. May.

 

206. Mature fruit and upper portions of shoots of bristly greenbriar- Ripened black fruits and the catclaw-like prickles of this species were shown here. This is a very commmon Smilax species from western portions of the Midwest west- and southward throughout the Cross Timbers and into the Edwards Plateau of Texas. This was a sequencial slide of the same plant that was shown in the preceding slide. Ottawa County, Oklahoma. November.

 

207. Basal part of shoot in bristly greenbriar- The adjective, "bristly" comes from the feature of the bristle-like (vs. catclaw) prickles, the appendages on lower parts of the shoot, that readily distinguish this from other Smilax species of this area that have only the rosebush type of prickle. Catclaw prickles are present on upper portions of the shoots of bristly greenbriar. This was the base of the stem of the plant shown in the two preceding slides.

Ottawa County, Oklahoma. December.

 

Let's see if we can ssee more details- Shoot of bristly greenbriar with closer-in views of the bristle-like catclaw prickles. Warning (none should be needed): this thing can slash wayward walkers to pieces.

Ottawa County, Oklahoma. Early January.

 

208. Fiddleleaf, fringed, or sawtooth greenbriar; catbriar; sawbriar; tramps-trouble; and several other common names (Smilax bona-nox)- This is probably the most common Smilax species in the oak-dominated deciduous forest, the ecotones between the oak-hickory forest and the tallgrass prairie (eg. those of the Ozark and Ouchita Mountains), and the savannahs further west like the Cross Timbers.

This plant was growing uphill from a pecan bottoms forest in a vegetational mosaic of West Cross Timbers and Grand Prairie. Locally dominant grasses growing near this particular plant included Canada wildrye, purpletop, Texas wintergrass, and the naturalized Johnsongrass. This particular individual was growing in association with Alabama supplejack or rattanvine (Berchemia scandens) and mustang grape (Vitis mustangensis) in a layer of woody vines that extended from the ground surface to tops of tall pecan trees so as to be component species of all layers of this woodland range vegetation.

This specimen was in full-bloom with details of inflorescence presented in the two succeeding slides.

Erath County, Texas. April.

 

209. Detail of leaves and inflorescences of fiddleleaf or sawtooth greenbriar- Leaves and flower clusers of S. bona-nox were presented in greater detail in this photograph taken in Erath County, Texas. April, full-bloom stage of phenology.

 

210. Inflorescences of fiddleleaf greenbriar- Up-close view of flower clusters in the specimen of S. bona-nox introduced above. Erath County, Texas. April, full-bloom stage of phenology.

 

211. Carolina snailseed (Cocculus carolinus)- This is a widely distributed herbaceous, perennial vine. Both scientific and common names were derived from the seed which resembles a snail shell. Importance of this plant as a valuable feed plant for wildlife, especially deer, has generally not been recognized. Carolina snailseed is an attracative native plant particularily in autumn when plants are commonly loaded with the bright red fruit. This species would seem to be valued for landscaping with native plant species. The herbaceous vines are pose much less of a yard-cleaning task than other twining plants.

This specimen was growing in the West Cross Timbers in a savanna dominated by a post oak-blackjack oak-pecan canopy with an understorey dominated by little bluestem. This plant grew up into the tree canopy such that it was present in all layers of the vegetation of this cover type.

Mills County, Texas. October.

 

212. Leaves and fruit of Carolina snailseed- Mills County, Texas. October.

 

213. Victory vine- Trumpet creeper (Campsis radicans) is one of the widespread, vigerously climbing, characteristic, and showy of all the woody vines in range types of western parts of the eastern deciduous forest formation, Prairie Peninsula, and Cross Timbers. Trumpet creeper prefers moderate to direct light so that it is most at home in fence rows with large trees, open forests or woodlands, and savannahs with abundant trees or large shrubs. This woody vine does, however, often thrive in forest gaps or in the spreading crowns of large forest trees so that it is a frequent neighbor to various grapes and other climbers like Virginia creeper. Trumpet creeper is a long-lived liana that thrives in hot habitats (as long these environments are not too dry) and is generally one of the more distinctive climbing shrubs in savannahs throughout the Ozark Plateau, Cross Timbers, and into the hardwoods-pine forest of the Southeast. In Texas its range extends from the Pineywoods to the Rolling Red Plains, but strangely it is absent from the Edwards Plateau, one of the most diverse savannas in the general southcentral region of the continent.

The beauty, longevity, and tolerance to harsh environments makes trumpet creeper an ideal species for native plant landscaping. It blooms intermittently over much of the summer and early fall. Both flowers and fruits as well as leaves are especially attractive among the native woody climbers. In fact, even the strongly grooved or furrowed bark is picturesque. In homesteading days, pioneers often planted various species of what they called "victory vines" around houses and outbuildings as a symbol of survival in the harsh new land they settled (or attempted to). Trumpet creeper is a native liana that is a natural "victory vine" (whether or not planted by settlers).

Erath County, Texas. September.

 

214. Trumpets in the Cross Timbers- A couple of views of trumpet creeper (at least what that "cap and ball" Epson V700 scanner did not crop off). This often-sprawling, woody creeper is a member of the Bignoniaceae, bignonia or catalpa family. The large, bright orange or even pink flowers and the long, green (later brown when mature) pod-like fruit render this a strinkingly showy, native, woody climber. The fruit is an elongated loculicidal or septicidal capsule The fruit is a "dead give-away" that this species is a member of the catalpa family as fruits of this family are quite distinctive. (Both immature, current-season and last year's open capsules were presented in the second of these two slides.

The plant presented in these photographs was growing happily in the West Cross Timbers of northcentral Texas. It is a common associate of post and blackjack oaks, and on calcareous Grand Prairie soils, live oak as well as pecans and sugarberry or sugar hackberry (Celtis laevigata) the latter two of which this species seems particularily fond. In much of the Ozark Plateau trumpet creeper apparently favors black cherry (Prunus serotina) and elms (Ulmus americana, U. rubra). In all cases, trumpet creeper is strinctly commensal on these trees.

Apparently trumpet creeper is also "neutral" in regards to browse for wildlife there being little discussion of it in that regard. The flowers are regularly visited by hummingbirds and, of course, various insects.

Erath County, Texas. September.

 

215. Bignonia blooms on the vine- A cluster of characteristic flowers of trumpet creeper showed the resemblance of these inflorescences to those of other members of the Bignoniaceae including those of Catalpa species. The ornamental value of this native liana was discussed in the preceding caption. This example was growing among pecans and post oaks in the Western Cross Timbers of northcentral Texas.

Erath County, Texas. September.

 

216. Another striking shrub in the Cross Timbers- Trumpet, evergreen, or coral honeysuckle (Lonicera sempervirens) is another shrub of the Cross Timbers range type. This member of the Caprifoliaceae thrives at edges of Cross Timbers stands where it can get adequate space and sunlight. When in bloom trumpet honeysuckle is clearly one of the most, f not the most, glaringly beautiful of all shrubs native to the Cross Timbers.

The specific epithetic, sempervirens, and common name adjective of "evergreen" indicate the persistence of green leaves through much of the year as, for example, in the relatively mild winters of the Texas Cross Timbers.

Erath County, Texas. April. Full-bloom phenological stage.

 

217. A native shrub ideal for landscaping Cross Timbers style- Details of inflorescences of coral or trumpet honeysuckle growing on a range dominated by post and blackjack oak and pecan. Enthusiasts who insist on using native species for landscaping should definitely settle on this beautiful shrub when housekeeping in the Texas Cross Timbers.Likewise conservation agencies should establish this species near park headquarters and visitors center. In fact, government agencies in general should use this species in prominent places.

Texas settlers made such wide use of trumpet honeysuckle that it is still available commercially as a landscape shrub. Diggs et al., 1999, p. 511) cited earlier work showing that the flowers of trumpet honeysuckle attracted humingbirds. The fruit is eaten by species of song birds (Vines, 1960, p. 957). More reasons why trumpet honeysuckle should be planted around the house, in arboretums, wildlife refuges, and parks.

Erath County, Texas. April for first and third photographs; late March for second photograph. Full-bloom phenological stage

Organization note: White or bush honeysuckle (L. albiflora) is another native shrub in the Texas Cross Timbers and Prairies vegetational (land resource) area, but this species grows on calcareous soils, especially on limestone outcrops, of the Grand Prairie. Thus these two Lonicera species do not usually occur in close proximity to each other except where Cross Timbers and Grand Prairie vegetation are conterminous in this patchwork of grassland and savanna range vegetation.

White or bush honeysuckle was presented with the Tallgrass Prairie (Interior)under the Grasslands biome heading.

 

218. Christmas or hairy mistletoe (Phoradendron serotinum= P. tomentosum)- Christmas mistletoe is one of the few parasitic plants of the oak-hickory forest and the transition zones between these cover types and those of the tallgrass prairie. It is a woody plant of the mistletoe family (Viscaceae) family or, according to some authorities, the Viscacoideae (subfamily) of the enlarged Loranthaceae. Phoradendron species are known as the "true or leafy mistletoes" in contrast to Arceuthobium species which ae the "dwarf mistletoes" that parasitize conifers (true mistletoes parasitize woody angiosperms). The green color (indicating presence of chlorophyll) of the evergreen P. serotinum showed that it is capable of some photosynthesis and is not totally parasitic. In ecosystem terminology Christmas mistletoe is partly heterotrophic, "other-nourishing", and partially autotrophic, "self-nopurishing" (ie. it is both a producer and a consumer). Nonetheless this species does parasitize such species as elms, hackberries, mesquite, and the oaks of the black oak subgenus (Erythrobalanus) to the point of deformity or death, directly or indirectly. Infected trees often survive for years (some branches or even major limbs may die), but they do so with reduced vigor, growth, and tolerance to stresses such as drought.

Like all consumers mistletoe species are selective in the species they use as food. In the Cross Timbers and adjacent prairies of Texas and southern Oklahoma infestations of Christmas mistletoe is sometimes great enough to have an impact on species composition of the range plant community. Populations of the co-dominant blackjack oak are often reduced almost to point of elimination leaving the unparatisized other co-dominant post oak (and live oak on calcareous soils of the adjacent prairies) unaffected. Christmas mistletoe may greatly reduce numbers and density of hackberry (which tend to be locally abundant on the uplands), elm (more common on bottomlands), and mesquite. Infestations and tree mortality appear to be more severe when trees are more concentrated and seeds are more easily transported by fruit-eating birds.

The specimen shown here had parasitized a lone blackjack on virgin prairie dominated by little bluestem. Erath County, Texas. February.

 

219. Details of Christmas mistletoe- Fruit, inflorescence, and leaves of the State Flower of Oklahoma were presented in this photograph of the specimen shown immediately above. The common name of this species was derived from the widespread tradition of using mistletoe branches like the one shown here as Crristmas decorations. This is a poisonous plant, though obviously not to the species that consume it's fruit. Toxicity of the Viscaceae family was covered by Burrows and Tyrl (2001, ps. 1178-1184) who concluded that while members of this family are listed as toxic, cases of animal poisoning have been were quite limited and, in fact, these species comprise part of the typical diet of cervids like deer and elk as well as small mammals and of course birds. The latter in particular spread the seeds either via the gastro-intestional tract or by their feet and bills when they preen themselves and remove the sticky seeds which attached to their bodies by means of viscin the mucilaginous covering surrounding the seed in the fruit (Diggs et al. 1999, p. 1065; Burrows and Tyrl, 2001, p. 1182).

Erath County, Texas. February.

 

220. Stem base of Christmas mistletoe- This is the base of the "trunk" of the parasitic Christmas mistletoe plant at the location where it penetrated the branch of the host or "prey" plant (honey mesquite in this example). The penetrating absorptive organ (for water, mineral nutrients, and photosynthate) of mistletoe species is called the haustorium, which is really a specialized branch or organ of a parasitic plant. The haustorium is a dense system of root-resembling strands of tissue that develop parallel to the cambium layer of the parasitized tree and which then send out in radial direction other organs called sinkers which penetrate into the xylem and phloem, sometimes for considerable distances from the base of the mistletoe (Agrios, 1988, p. 619-620).

Erath County, Texas. March.

 

Are you cocksure on this one?- Leaders at a crown-scale (first slide) and viewed as two leaders (second slide) of cock's-spur hawthorn (Crataegus crus-galli). This specimen was growing in an old fenceow with blackjack oak, post oak, black or Texas hickory, honey locust (Gleditsia tricanthos), and silky wildrye (Elymus villosus). This native species was once more commonly found on tallgrass prairie and in the middle woody layer of tallgrass--oak-hickory savanna than today where taller growing woody plants tend to "overshadow" and crowd it out. Cock's-spur hawthorn also grows at the outer edge of oak-hickory forest and tallgrass prairie.

The Crataegus species are a taxonomic/identification nightmare come true! Crataegus is one of the most complex and complicated genera of woody plants in North America. Vines (1960, p. 329) stated the case well by stating simply that it "...represents a very difficult taxonomic complex". Identification of Crataegus species is often arbitrary and a "best guess" even when all morphologically critical organs are present. There can amazing variation among organs of the same speacies (even on the same plant), yet at the same time great similarity among organs of different Crataegus species. Leaves and fruits are extremely similar among related species. Add to that situation the large number of Crataegus species, the number of hybrids between species, and the fact that some specimens are apomictic clones. For instance, Steyermark (1963, ps. 803-822 passim) showed 18 Crataegus species as native to the general area of southwest Missouri, northeast Oklahoma, and southeast Kansas represented by this example.

(Crataegus has been subdivided into numerous sections or series and there is tremendous variation within any given species with such variation having been shown as taxonomic varieties. For instance, Vines (1960, ps. 354-355) described five varieties of C. crus-galli while Fernald (1950, ps. 775-776) described eight varieties of C. crus-galli. As to subunits within Crataegus, Vines (1960 ps. 354-367) included 19 Crataegus species within Series Crus-galli. Sargent (1933, ps. 400- 422) included 25 species in his natural group, Crus-galli. Steyermark (1963, ps. 808-814) included 14 Crataegus species in his series 7, Crus-galli.

The above difficulty having been underscored, the example species shown here was positively identified as C. crus-galli based on leaves, fruit, bark, and thorn features. C. crus-galli is a fairly definitive species. Cock's-spur hawthorn was also one of the more widely distributed and abundant species in this area which sort of "assisted" with identification of this specimen which the author has observed over several years.

Ottawa County, Oklahoma. Late June; immature fruit stage.

 

Yep, feeling purty cocky- Several leaders (branches) of cock's-spur hawthorn with long thorns that characterize this species. This specimen was growing on degraded tallgrass prairie in the western Ozark (Springfield) Plateau in northeastern Oklahoma. Shrubs or shrub-sized trees of this species commonly grow on infrequently burnt tallgrass prairie (or, alternatively, plants are large enough that they are not top-killed by recurrent [say, annual] prairie fire).

This plant was growing in a fencerow beside blackjack oak, post oak, and Texas or black hickory with an understorey dominated by silky wildrye.

Ottawa County, Oklahoma. Late June; immature fruit stage.

 

At least at the half-cock position- Leaves and unripe fruit of cock's-spur hawthorn on a young shrub-sized plant growing in a fencerow with blackjack oak, post oak, Texas or black hickory, honey locust, and silky wildrye. Hawthorn species are in the Pomoideae (Maloideae= Pyroideae) subfamily of the rose family, Rosaceae, the fruit of which is the pome type. These pomes are frequently important concentrate feeds for all manner of birds and mammals, especially smaller animal species. Seeds in these pomes are often quite effective in establishing new plants of cock-'spur hawthorn (often where they are unwanted by range managers, ranchers, and other landowners).

Cock's-spur hawthorn has been one of the native shrubs more widely used for ornamental purposes, including in Europe (Sargent, 1933, p. 403). This species has a natural biological range that extends from Ontario across to Minnesota south to the Carolinas and westward to east Texas.

Ottawa County, Oklahoma. Late June; immature fruit stage.

 

Cocky in the Cross Timbers- Good-sized specimen of cockspur or cock's-spur hawthorn (Crataegus crus-gallii) growing on a Indiangrass-little bluestem/blackjack oak-post oak savanna in the West Cross Timbers. The Crataegus species are a taxonomic bug-a-bear and a textbook case of "lumpers" versus "splitters". Be that as it may, C. crus-gallii is well recognized with some of the largest plants being included in this species which, by the way, can grow as either large shrub and small tree. Multiple shoots comprised the trunk of the plant picture here so that it was regarded as a shrub.

The Crataegus species have traditionally been separated into three or more Series, including Series Crus-gallli (Correll and Johnston, 1979, p. 734).

Erath County;, Texas. Late April.

 

Cock-spurred leaders- Outer or distal portions of two leaders (first slide) and terminal part of one leader (second slide) of cockspur hawthorn on the specimen introduced in the immediately preceding slide. These outer branches were backdropped against the herbage of co-dominants Indiangrass and little bluestemon a savanna characterized by post oak and blackjack oak and with cockspur hawthorn as the major shrub species. (As indicated above, some specimens of this species grown upwards of a small tree.)

Correll and Johnston (1979, p. 740) described branches of this species as "widepreading" which was certainly descriptive of the examples presented here.

Erath County;, Texas. Late April.

 

Characteristic leaves (first slide) and branching pattern (second slide) of cock's-spur hwwthorn in the West Cross Timbers of northcentral Texas. These organs were on the specimen shown in the two preceding slide-caption sets. A detailed description of this species was provided by Correll and Johnston (1979, p. 740) who regarded leaves as thick upon maturity and shinney on their upper surfaces.

Erath County;, Texas. Late April.

 

Short spurs- Two examples of the thorned branchlet or thorn (depending on wording) of cock's-spur hawthorn on the large shrub that served as an example specimen on an Indiangrass-little bluestem/blackjack oak-post oak savanna in the West Cross Timbers.

As was so often the case regarding woody plants in southcentral and eastern southwestern North America, Vines (1963, ps. 354-355) served as a basic, understandable reference for cockspur hawthorn. In fact, treatment of Crataegus in Texas (Correll and Johnston, 1979, ps. 734-743) followed the treatment of Vines (1963, ps. 329-387).

Erath County;, Texas. Late April.

 

 

221. Colony of highbush or timber blackberry, briar(s), or bramble (Rubus pensilvanicus) in full-bloom at edge of an oak-hickory forest and tallgrass prairie- The Rubus genus is another one of those taxonomic and identification nightmares (once one gets beyond the obvious Rubus taxon) that is easiest left to those specializing in this group. Positive identification usually requires study of both the primacane (first year, vegetative, shoots) and the floracane (second year, sexual or flowering, shoots). Rubus species include blackberries, raspberries, and dewberries. Members of the latter group are the ground-trailing species. Among those species having up-right or vertically oriented canes (often with downward arching or drooping tips) raspberries are distinguished (from blackberries) by having a "hollow" in the "berry" when picked from the bush. Among the Rubus species fitting the blackberry criteria of erect, often nodding, canes and "berries" with a "solid center", R. pensilvanicus is the most common and widely distributed species in the Ozark Plateau where this example was photographed.

The Rubus fruit though called a "berry" is actually an aggregate fruit which, like the fruit of strawberry (in the same subfamily as blackberry), is not an actual fruit but a "false fruit". An aggregate fruit is a type of false fruit formed by the union of the actual fruit with another part(s) of the flower, in this instance the unifying structure comprised of separate fleshy fruits originating from a single flower having numerous free carpels. The carpel is the ovule and the collective carpels comprise the ovary. An ovary may consist of only one carpel, but in aggregate fruits there are several to many carpels. Whatever their botanical excentricies these "berries" are some of the finest human "sweet foods" produced in the woods. They are eaten fresh as well as in some of the most delicious pies, jams, and jelliest known to man.

Contrary to what would seem to be the case given presence of sharp, cat claw-like prickles, the canes (shoots) of Rubus are quite palatable to animals, especially cattle.This situation obtains mostly for the the young primacanes, and mostly early in the season before the shoots become woody and the prickles (thorn-like dermal appendages lacking vascular bundles and attached only to the epidermis of the shoot) grow hard and razor-sharp. That notwithstanding, large concentrations of cattle and even horses will destroy a prized blackberry patch in a single season. When blackberries and raspberries are desired keep livestock away from the plants. Deer also browse the tender primacanes, but their numbers usually are not great enough to kill out blackberries.

The blackberry group of Rubus species are most abundant locally on disturbed but abandoned land like old fields, overgrazed prairie, and forest openings (or small clearings). This would suggest that these are ecological invaders. Specifically they appear to be early seral species in the oak-hickory and oak-hickory-tallgrass ecotone region. However, many of the briar species, of which R. pensilvanicus is a "prime example", are quite susceptible to defoliation like persistent mowing and overbrowsing ind in absence of such stress persist into climax communities of both of these general vegetation types. In fact, the largest fruits are typically find in shade hence the vernacular among berry pickers of "timber berry". The example presented here was in a oak-hickory-tallgrass savanna in the western Ozark Plateau. Ottawa County, Oklahoma. May.

 

222. A Rubus species at two stages of sexual reproduction- Two stages of flower/fruit development were shown here. The first slide was of blackberry flowers at peak bloom. The second slide was at immediate post-bloom stage with tiny aggregate fruits just starting to develop.

Ottawa County, Oklahoma. First photograph taken in May; second photograph in June.

 

223. Himalaya blackberry (Rubus procerus)- This introduced (= horticultural) Asian species of Rubus naturalized in various locations throughout the Ozark Plateau where it's pronounced flavor (and disctinctive canes complete with talon-like prickles) have added another "ingredient" to the fare of berry-picking hillbillies like this author. This species illustrated the fact that identification of Rubus species, which often hybridize "like crazy", was made more difficult by horticultural introductions.

Briar patches like this one provide valuable habitat for many species of wildlife. An immense coachwhip snake (Masticophis flagellum) called this patch home and never expressed the least concern whenever your author helped himself to the plentiful fruits of the naturalized alien Rubus.

Oak-hickory-tallgrass savanna in Ozark Plateau. Ottawa County, Okla;homa. July.

 

224. Aggregate fruit of the famed timber blackberry (Rubus pensilvanicus)- Aggregrate fruit was defined with the first slide that introduced the Rubus genus. Blackberries are one of several types of false fruits. The aggregate fruit is actually a "clump" or "bunch" of several separate fleshy fruits, each containing a hard tiny seed and formed from a separate or single carpel, all of which comprise one ovary and form the whole or collective group (ie. the aggregate). And there is no finer pie in this world than blackberry.

Ottawa County, Oklahoma. July.

 

225. An even sweeter, more delicate aggregate fruit- Western or black raspberry (Rubus occidentalis) is a native to much of the eastern deciduous forest and into the oak-hickory-tallgrass savanna such as that in the Ozark Plateau Region. This Rubus species produces a sweeter tasting aggregate fruit, but also one more easily "squashed" during picking. The feature that distinguishes Rubus species designated as raspberry from Rubus species called blackberry is the cavity or hollow "center" of raspberry fruit. The indivudual fruits (carpels) of the aggreagte frruit type of Rubus species form or insert on a soft receptacle. In blackberries and dewberries this receptacle remains inside the aggregate fruit (the whole "berry") whereas in raspberries the carpels separate from (come off of) the receptacle leaving a hollow (resembling a gastrula) with the individual fruits (mature carpels) barely clinging to each other. This results in a tendency for the individual fruits ( ripened carpels) to come apart in small groups.

Specimens shown here were growing in and aroung water-holding depressions in which sycamore grew interspersed with oak and hickory trees and tallgrasses like big bluestem (ie. a moist savanna).

Ottawa County, Oklahoma. June: peak fruit season (hillbilly holiday).

 

226. Details of Ozark raspberries- Close-up shots of aggregate fruit of western or black raspberry. The unequivocal identifying feature that distinguishes raspberry from blackberry species within the Rubus genus is the separation--when ripe--of the aggreagte fruit from its central receptacle in raspberry in contrast to continued attachment of fruit and receptacle in blackberry (Steyernmark, 1963, p. 834). The result in raspberry of this separation (non-retention of receptacle) is the gastrula-like hollow within the aggregate fruit or the remaining, empty space around which the individual, small, seed-containing fruits are still held together.

Newton County, Missouri. Early June.

 

One way to spread- Tips of primocanes of western or black raspberry developing adventitious roots in the early stages of layering, forming new canes asexually. This pattern of forming new asexual shoots has often been designated as simple or tip layering. Horticulturalists routinely use this method of vegetative propagation.

"In the wild" layering is much more frequent in eastern or black raspberry than in most other Rubus species found on prairie, forest, or savanna. The example seen here was occurring naturally not by horticultural manipulation.

Ottawa County, Oklahoma. Early August.

 

227. Bittersweet- Bittersweet or American bittersweet (Celastrus scandens) is probably the best known member of the Celastraaceae (bittersweet or staff-tree family). This liana (woody vine) can be found on tallgrass prairie, particularily where associated with larger woody plants on which it can twine and in protected habitats (brush fencerows are a bittersweet favorite). More commonly bittersweet grows in open oak-hickory forest and the oak-hickory forest-tallgrass savanna such as in the Prairie Peninsula.

The individual shown here (first slide) was growing on a persimmon sapling, and without inflicting damage on its supporting host. Fruit presented in the second slide was immature.

This woody vine was one of the first to be planted as an ornamental pioneers and second-stage farmers because wives of badkwoodsmen could dig them up or plant cuttings "free for the taking" throughout much of eastern North America plus the flaming red or bright orange fruit persist throughout much of the winter to add a touch of color in the bland season.

Vines (1960, p. 660) mentioned that many kinds of wildlife, including some of the most valuable upland game and song bird species consumed the brillantly colored fruit which, incidentially, is usually interpreted as a capsule. Steyermark (1963, p. 1010) reported that white-tailed deer readily ate the leaves of this liana.

Springfield Plateau portion of Ozark Plateau (Highlands), Ottawa County, Oklahoma. July.

 

228. Bittersweet fruit- Mature fruit of climbing or American bittersweet. Bittersweet fruit is a globose capsular fruit that breaks open as three valves exposing fleshy seeds covered with arils (Steyermark, 1963, p. 1010). This brightly colored covering of the seed, the aril, "an outgrowth of the hilum which takes the form of a partial covering around a seed" (Smith, 1977, ps. 161, 289). This detail was visible in the second of these two slides.

Springfield Plateau portion of Ozark Plateau (Highlands), Ottawa County, Oklahoma. October.

 

229. An uncommon showy one- Shrubby St. Johnswort (Hypericum spathulum= H. prolificum) in a local mesic microsite in an ecotone between a chert glade variant of tallgrass prairie and oak-hickory forest in the Springfield Plateau in southwest Missouri. There are quite a number of Hpericum species on North American range. Steyermark (1963, ps.1055-1064) listed 13 Hypericum species for Missouri while the Great Plains Flora Association (1986, ps. 237-239) described seven Hpericum species (of which H. spathulum was not one) for a geographical area extending from northern Oklahoma, through much of the Osage Plains in western Missouri and Iowa, a good part of the Great Plains except in Texas and New Mexico, then north to the southern Canadian Prairie Provinces.

Some of these Hpericum species are woody while others are herbaceous. The forbs (hrerbaceous species) are either annuals or perennials. Likewise, while most of these are species are native, others are introduced and naturalized. As implied by the common name of shrubby St. Johnswort, this species is a subshrub (Steyermark 1963,p. 1059). Shrubby St. Johnswort is a semi-woody species of riparian areas/zones or, at least of low, mesic habitats more than upland areas (other than those upland sites or microsites that for various reasons are more favorable soil moisture-wise).

While some of the Hypericum species are poisonous to livestock, H. spathulum is not one of these (Burrows and Tyrl, 2013, ps. 710-716)

Newton County, Missouri. Late July; full-bloom phenological stage.

 

230. Botanical pinwheels- Flower clusters of shrubby St. Johnswort are borne singularly (ie. they are regarded as solitary). Can there be any doubt that this native species has tremendous ornamental value for those wishing to landscape with native plants and natural-like local landscapes?

Hpericum species have been included as members of the family known variously as the Guttiferae (= Clusiaceae) or Garciniae (= Hypericaceae) (Steyermark, 1963, p. 1055; Smith, 1977, p.118 ; Great Plains Flora Association, 1986, p. 237; Diggs et al, 1999, p. 544). Hpericum species are in their own subfamily, Hypericoideae.

Newton County, Missouri. Late July; full-bloom phenological stage.

 

Another example- Leader of shrubby St. Johnswort featuring charactristic leaves and overall arrangement or pattern of the inflorescence of this woody member of a an otherwise group of range forbs. This shoot was on the same plant that introduced this species though in a later year.

Newton County, Missouri. Early July; full-bloom phenological stage.

 

In case you missed them earlier- More examples of flower clusters and leaves of shrubby St. Johnswort on a plant that was growing on a transition zone between tallgrass prqairie and a tallgrass-oak savanna in the Ozark (Springfield) Plateau in southwestern Missouri. These flowers and leaves were on the same plant (above) that introduced this species. These organs were produced on that plant the following year.

Newton County, Missouri.Late June; full-bloom phenological stage.

 

King grass- Big bluestem (Andropogon gerardii= A. furcatus) at end-of-growing-season peak development and at mid-stage of caryopsis maturity. Although these three images were at progressively closer camera distance the first two slides were of two different plants while the third was of tillers on the plant presented in the first slide.

Some of the tillers on these plants exceeded seven feet in height. They were growing on a part of a restored tallgrass prairie/oak-hickory forest savanna. savannah at the western edge of the Springfield Plateau in southwest Missouri.

Big bluestem is not only the overall regional dominant of the tallgrass prairie it is also one of the most fire-adapted grass species in North America. Spring fire tends to initiate or in some way induce more sexual(flowering) tillers.

Big bluestem arguably produces the highest quality forage of any of the tallgrass species. "Few, if any, of the prairie grasses can equal big bluestem in quality or quantity of forage produced" (Phillips Petroleum Company , 1963). The literature on this magnificant tallgrass species is as abundant and, some of it anyway, as high-yielding as big bluestem itself. Interested students will have no trouble finding literature dealing with big bluestem.

Big bluestem is the State Grass of Missouri and State Grass of Illinois.

Wildcat Glades, Newton County, Missouri. Early August; peak standing crop, mid-maturity grain stage of phenology.

 

Now that's a sexy grass- Sexual (= flowering) shoots of big bluestem at mid-stage of grain (caryopsis) maturity. These shoots were some of those shone in one of the lants in the immediately preceding three-slide/photo caption unit. They were produced on a tallgrss prairie-oak/hickory savanna in the western Ozark (Springfield) Plateau in southwest Missouri.

Wildcat Glades, Newton County, Missouri. Early August; peak standing crop, mid-maturity grain stage of phenology.

 

Humble beauty; inspiration to a prairieman- Entire inflorescence of big bluestem (first slide) and rames (defined lower in this caption) of big bluestem (second and third slide), all obviously at anthesis. Specifically, the stage of flowering is mid-anthesis with earliest spikelets (distal portion of rames) having already bloomed and shed anthers while lower (proximate to culm) portions were still in pre-bloom stage. This is determinate blooming: uppermost and outermost flowers bloom first while innermost and lowermost flowers bloom last.

Highnight et al. (1988, p. 8) illustrated and described the flower cluster (inflorescence) of big bluestem as a panicle of subdigitate racemose branches. These racemose branches have long been called rames where a rame is "an inflorescence branch which bears some stalked and some sessile spiklets (Pohl, 1968, p. 242).

The arrangement of rames in the flower cluster of big bluestem is a pattern termed furcatus meaning "forked" from the Latin furca ("fork"), hence an earlier specific epithet used for big bluestem was furcatus: Andropogon furcatus. The epithet furcatus is part of the binomial name for organisms as diverse as plants in the rose family to birds and fish among the vertebratse. Sometimes the more descripive specific epithets have not had priority. Big bluestem is a case in point. The commenrtaive name of gerardii (for Loius Gerard, a French botanist [Diggs et al., 1999, p.1238]) is less descriptive than furcatus for the forked inflorescence.

A "really neat" aspect of grass flowering is that flowers (florets) are so small that only folks who care enough to slow down and marvel at their beauty can appreciate them. Not exactly a rose flower as big as a teacup that any goon can catch a glimpse as he barrels along at 80 mile an hour. The subtlest, humblest beauty is the grandest of all, especially when it is the beauty of reproduction to perpetuate a species.

Wildcat Glades, Newton County, Missouri. Early August; near-peak standing crop, full-anthesis stage of phenology.

 

231. 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.

Of the "Four Horsemen of the Prairies--Big Bluestem, Little Bluestem, Indiangrass, and Switchgrass--", which are all mesophytes, little bluestem is the most xeric followed by Indiangrass. Most often it is big bluestem and Indiangrass that vie for dominance over more mesic range sites In the tallgrass prairie and adjoining oak-hickory forests along with savannas of these two major communities. Indiangrass typically prevails--even if ever so slightly--over big bluestem on slightly less mesic sites such as those on sandstone and shallower soils, south slopes, etc. By contrast big bluestem rates the advantage on range sites with limestone soils and on north slopes.

Grand Prairie vegetation. Erath County, Texas. October; peak bloom.

 

232. 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 especially those tribes that entered into treaties with the United States of America which includes all of those in Oklahoma and Indian Territories.)

Erath County, Texas. October; peak-bloom stage.

 

233. 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).

 

234. 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).

 

235. Tallgrass in the timber- "Timbergrass" was in former days the common name for big bluestem (Andropogon gerardii) commonly used by hill folk in the Ozark Mountains. This descriptive, though nonstandard, common name reflected the most widespread--often the only remaining--habitat of big bluestem in these ancient mountains. Aside from a few "bald knobs" (glades), the only remaining refuge for big bluestem was "in the timber", in the herbaceous understorey of what few oak-hickory forests had not been overgrazed. Fertile grassy and forested bottomlands were quickly converted to cropland, particularily corn. Even many of the rocky forested hillsides were cleared by logging to become strawberry ( Fragaria virginiana) patches. (Strawberry is native to the Ozarks and was covered below with other forbs.) This left second-growth forest as the last stronghold of big bluestem. Ergo, timbergrass to the 40 acres-and-a mule hillbillies.

This large individual of big bluestem was at peak stqanding crop and anthesis. Some shoots approached eight feet in height. It was growing in a small clearing of a second-growth forest and was surrounded by post oak, blackjack oak, Texas or black hickory, and black cherry. The herbaceous associates of the climax tallgrass were the annual composites, giant ragweed (Ambrosia trifida) and horseweed or marestail (Conyza canadensis) and the perennial composites hairy sunflower (Helianthus hirsuts) and Baldwin ironweed (Vernonia baldwinii). A shoot of green-briar (Smilax bona-nox) was the main shrub of this interesting local community. In this region, the two annual composites are first- and second-season pioneers or colonizers on abandoned farm fields whereas ironweed is commonly regarded as an invader due to disturbances like overgrazing. Obviously big bluestem is a member of the climax community. Typically it is the dominant decreaser tallgrass species over much of the oak-hickory region. An unusual combination or aggregation of plants ranging from earliest pioneer to climax species. What combination of biotic and abiotic factors produced such local community?

The second-growth forest shown here was almost assuredly some degree of forest encroachment or afforestation (due to combination of fire reduction or cessation, overgrazing, and high-grade logging) on what had been under pre-white man times part of the Prairie Peninsula, the climax savanna of the regional-scale deciduous forest-tallgrass prairie ecotone. Two old buffalo wallows remained less than a quarter mile from this interesting local range plant community. In the author's boyhood days, this patch of timber burnt off every two or three spring times and big bluestem, Indiagraass, and beaked panicgrass were widely dispersed in an understorey of widely spaced oaks and hickories. In the ensuing four or five decades "setting the woods afire" was but a memory while the woods grew thick with scrubby and sapling trees while a relict individual of tallgrass became cause for celebration and a Kodachrome moment. This photograph was placed here to share the pleasure and pass the knowledge of the "real vegetation" of the Ozark Highlands to those never blessed with the experience. Range yesteryears.

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

 

236. Sharing sex in the brush- Sexual shoots of the big bluestem plant introduced in the preceding slide. This plant was at peak standing crop and height of anthesis. Height of most shoots was from seven to eight feet. The turkey track inflorescences were silhouetted against a backdrop of leaves on post and blackjack oaks and Texas or black hickory.

Ottawa County, Oklahoma. July; peak stand crop and anthesis.

Ecological orientation on two "kind of" ecological equivalents: The major four species of tallgrasses--the "Four Horsemen of the Prairies"-- throughout and across much of the humid zone of the tallgrass prairie, open oak-hickory forest, and ecotone between these two major range types the tallgrass-oak-hickory savanna are 1) big bluestem, 2) Indiangrass, 3) switchgrass and 4) little bluestem. Most of this once vast interior region now has most of its virgin vegetation destroyed, highly modified, or, at best, fragmented, but the original relationships among the dominant grasses and their ceological niches remain the same.

Big bluestem is typically "first among equals" of the Four Horsemen of the Prairies in terms dominance on acreage occupied as well as area and number of range sites on which it is a major species. Switchgrass is "first" on the more mesic to wet range site while little bluestem is the major dominant on the more xeric, shallow range sites. Big bluestem and Indiangrass are closest together in terms of moisture, fertility, etc. requirements (ie. most similar of the "big four" in ecological niches). Indiangrass is somewhat less mesic in its requirements and tends to rate a slight advantage over big bluestem on slightly drier habitats. Indiangrass tends to outrank big bluestem on sandstone-derived soils whereas big bluestem ourtanks Indiangrass on limestone and chert-derived soils.

Big bluestem is often the slight to overwhelming dominant and Indiagrass the weaker dominant to associate species on oak-hickory-tallgrass savannahs such as those in the Ozark Plateau. This is an interesting situation given that Indiangrass is a long-shoot grass, one that elevates its apical meristem early in its annual growth cycle and reaches a greater portion of its total shoot height sooner, while big bluestem is a short-shoot species with the opposite annual patrtern of shoot elongation and growth. Nonetheless, such was the relationship between big bluestem, the dominant grass, and Indiangrass, the associate grass species, on Stoney Point Savannah described above. In the next two slide-caption sets inflorescence units and spikelets of these two tallgrasses, growing on Stoney Point Savanna, were shown and described.

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.

 

237. 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.

 

238. Next genetic generation of Indangrass- 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.

 

239. Two invaders on degraded grazing ground- Broomsedge bluestem (Andropogon virginicus), left, and splitbeaard bluestem(A. ternarius), right, on degraded oak-hickory-tallgrass savanna in Ozark Plateau. Both of these highly unpalatable bluestems are in the same section (Arthrolophis) of Andropogon as big bluestem (Hitchcock and Chase, 1950, p. 750). Likewise, broomsedge and splitbeard bluestems are invaders in contrast to the decreaser status of big and little bluestems as the former are mostly native ruderals of old fields (= go-back land), cutover forests, overgrazed ranges, and improperly managed fields of introduced forages such as Eursian grasses.

These two plants were growing on a savanna pasture that had a history of abuse going back at least 60 years, but which had not been mowed or grazed during the preceding spring through autumn growing season.

Ottawa County, Oklahoma. December; grain-ripe stage and peak standing crop at dormancy.

 

240. Parted pubsecence on spread shoots- Numerous shoots of the splitbeard bluestem growing on a degraded Ozark Plateau savanna. Unique arrangement of spikelets on the plant of Andropogon ternarius that was introduced in the prededing photograph. Another widely used generic common name for the bluestems is beardgrasses. This photograph showing some of the spikelets on some of the shoots on a single plant and particulars of spikelets of this species in the next two-slide set illustrated the apt designation, beardgrass.

Ottawa County, Oklahoma. December; grain-ripe stage

 

241. Split spikelets- Detail of spikelets of splitbeard bluestem. Spikelets are paired (one fertile and sessile, the other being sterile and pedicellate) with there being paired florets (one fertile and the other sterile) in the fertile spikelet. These photographs were taken on the same plant introduced above that was growing beside a plant of broomsedge bluestem on a degraded oak-hickory-tallgrass savanna range..

Ottawa County, Oklahoma. December; grain-ripe stage.

 

242. 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.

 

243. 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.

 

244. Inflorescences of beaked panicgrass- The flowering shoots of P. anceps on a tallgrass prairie glade in the Ozark Plateau. Newton County, Missouri. September.

 

245. 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.

There are a number of species of rosette panicgrasses in the oak-hickory-tallgrass savanna that extends throughout much of the Ozark Plateau Region and on into the Cherokee Prairie of the Central Lowlands. Some of these species were presented below following an explanation of the taxon traditionally regarded as rosette panicgrasses or rosette Panicum species. This introduction included a brief chronology and note on sources in regard to the elevation of this group to a genus of its own, Dichanthelium.

Most grasses, with remarkably few exceptions other than the bamboos, have the typical--at least tthe more common--grass habit consisting of "regular" annual shoots (only rarely do herbaceous shoots survive more than one growing season) that are of the same morphological strucrture throughout growth except for reaching larger size. In most grass taxa typically there are not shoots of one form (morphology) or arrangement (pattern; general physical organization) in one season or stage of phenology and then another morphological structure and pattern in another season or stage of phenological development. There is an exception to this typical grass habit within the panicgrass group. In addition to the panicgrasses that have the same general form and arrangement of shoots from one growth or phenological stage to the next (eg. beaked panicgrass presented above), there is a taxonomic group characterized by development of a basal rosette of lower leaves in winter and, typicallly, a flowering period in both spring and autumn. In addition to this variation from standard grass shoot morphology, some of the rosette panicgrasses develop branches off of nodes on upper parts of the central shoot so as to have a dendritic form. Hitchcock and Chase (1951, ps.627-633) placed this latter taxon in the Panicum subgenus designated as Dichanthelium. Species in subgenus Dichanthelium have traditionally been known as the rosette panicgrasses, sometimes as rosette panics, and recently, with removal from Panicum to Dichanthelium, simply as rosette grass (= rosettegrass).

Gould (1975, ps.477-498) and Gould and Shaw (1983, ps. 229-230) split out species in the traditional Dichanthelium subgenus of Panicum and elevated them to a new and separate genus: Dichanthelium. This reinterpretation of Dichanthelium was not adopted subsequently by all agrostologists. For example, the International Symposium on Grass Systematicvs and Evolution (held 27-31 July, 1986 at the Smithsonian Institution) retained Dichanthelium as a subgenus of Panicum (Soderstrom et al., 1987, p. 299). Almost two decades later in the Flora of North America, ...Poaceae, part 2, which is likely to be regarded as the authorized version of the "bible" of North American grasses, Barkworth et al. (2003, ps. 406-451) adopted the elevation of Dichanthelium to genus level "based on recent molecular data" (Barkworth et al., 2003, p. 406) as well as the morphological features of winter rosette and biseasonal flowering relied on by Gould (1975) and Gould and Shaw (1983).

Whether or not one accepts the standards of cladistics over traditional phyletics in Plant Taxonomy (and this author does not), the two grass volumes in Flora of North America (Barkworth et al., 2003; Barkworth et al., 2007) are invaluable references if for nothing other than the outstanding line drawings and descriptions that permit users to distinguish and identify grass species better--though certainly not easier--than any other available manual. Species names of rosette panicgrasses were shown as the synonyms of Panicum (Hitchcock and Chase, 1951) and Dichanthelium (Barkworth et al., 2003), at least to the extent permitted by two references that are not completely commensurable. Names were also checked against those in both editions of The Grasses of Missouri (Kucera, 1963, 1998), the grass manual for an area that was in closest proximity to where most of these specmens were photographed (extreme northeastern Oklahoma). In the following binominals the first listed is Panicum from Hitchcock and Chase (1951) and Kucera (1963, 1998) and the second is Dichanthelium from Barkworth et al. (2003). Although Hitchcock and Chase (1951) and Kucera (1963, 1998) all used Panicum rathern than Dichanthelium some Panicum specific epithets changed from Kucera (1963) to Kucera (1998). These latter changes were shown using "[Kucera, 1963]" and/or "[Kucera, 1998]".

 

246. Scribner's panicgrass (Panicum scribnerianum= P. oligosanthes var. scribnerianum [Kucera, 1998]= Dichanthelium oligosanthes var scribneriaum)- This is one of the most widely distributed rosette panicgrasses in North America. It extends from Maine across to British Columbia and southeastward to Florida. The Natural Resources Conservation Service, United States Department of Agriculture (www.plants.usda.gov/java/profile) showned this species as present in all but two of the "lower 48" USA states. In addition to this immense species range, Scribner's panicgrass is frequently the most common of several species of rosette panics such that all such Dichanthelium members are "lumped" as "Scribner's panicgrass" (for example this is common practice for FFA and 4H range contests).

The first specimen of Scribner's panic presented here (first slide) was growing in a black oak-big bluestem-dominated savannah on a rocky soil with tripoli as parent material in the Springfield Plateau part of the Ozarks Region. In a commonly observed situation, this plant was growing beside a plant of slimleaf or linearleaf panicgrass (P. linearifolium), a physical association presented in more detail shortly. The second specimen of Scribner's panic was growing on a sandy soil in the West Cross Timbers of northcentral Texas.

First photograph: Ottawa County, Oklahoma, June (milk stage just past anthesis). Second photograph: Erath County, Texas, May (graain-ripening stage).

 

247. Shoot apex of Scribner's panicgrass- Panicle in a shoot of Scribner's pancigrasss in the West Cross Timbers of northcentral Texas. This panicle was not as fully exerted out of the boot as some others, but this afforded a better view of the shoot apex than in cases of fuller expansion. Erath County, Texas. Early May.

 

248. Velvet panicgrass (P. scoparium [Kucera, 1963, 1998]= Dichanthelium scoparium [Barkworth et al., 2007])- This somewhat decumbent plant had a shoot that was over a yard long (tall). It was growing on a tripoli-derived soil in the Springfield Plateau of the Ozark Mountain Region. This specimen was associated with blackberry and big bluestem in the understorey of a black oak-dominated savanna. Abundant precipitation from the preceding winter and spring undoubtedly explained some of the robust size and vigor of this plant as shown here in early summer. This plant was protected from livestock and subject to grazing by white-tail deer and other wildlife species, none of which had seen fit to partake of the available feast.

This is a readily identifiable species once one knows its key distinguishing feature, long slilky pubescence all over the shoot. "The villous character of this species is distinctive and easily recognized" (Kucera, 1961, p. 185). Styermark (1963, p. 221) remarked that P. scoparium is not likely to be confusesd with another Panicum species (at least not in Missouri) givne the velvety pubescence along with a glabrous band subtending hairy nodes.

Ottawa County, Oklahoma. July (grain-ripening stage of phenology).

 

249. Shoots and inflorescence of hairy panicgrass- General view of a vigerous though typical shoot of hairy panicgrass (first photograph) followed by detail of the large panicle (second photograph) on this shoot. First slide was instructive in showing branches (secondary shoots) coming off of the main shoot of this individual plant. It was explained above in the introduction of rosette panicgrasses that some species of the at outer edge of a oak-hikcory/tallgrass savanna taxon have the unusual shoot morphology of secondary shoots branching off of nodes on upper parts of the main shoot (versus tillering from the rootcrown which is a typical feature of most grasses). This deviation from standard grass shoot morphology is otherwise restricted primarily to the bamboos (Bambusoideae).

Details of the panicle, including branching pattern and ripening spikelets, were presented using the larger inflorescence (second photograph) of this individual plant that was first presented in the immediately preceding slide.

Ottawa County, Oklahoma. July (grain-ripening stage of phenology).

Given the uniqueness of velvet panicgrass (Panicum scoparium) the following section was included for greater detailed treatment of this cool-season, C3 photosynthetic, panicoid grass.

 

250. Velvet colony- Colony of velvet panicgrass shoots produced on a pile of tossed field rock (mostly chert) on the outer margin of an oak-hickory/tallgrass savanna in western Ozark (Springfield) Plateau. Ottawa County, Oklahoma. Velvet panicgrass is not rhizomatous, but in long-established plants, such as this large plant, there can be many tillers arising from the large rootcrown of a single individual (one genetic plant or genet). There was also a large plant of Canada or nodding wildrye growing in this rock pile.

Ottawa County, Early July.

 

251. Patch of tillers in a rock pile- Closer-in views of some of the numerous shoots of one plant of velvet panicgrass growing in a rock pile (this was the plant that was introduced immediately above). Shoots of velvet panicgrass are often referred to as "stout" (Greaat Plains Flora association, 1986, p. 1160) which, though a relative adjective, was an apt description. These tillers were also comparatively tall (by standards of the rosette panicgrasses) with some approaching a yard in height. The ascending and somewhat zig-zag morphology of culms is one of several defining and distinguishing features of this species of robust individuals.

Ottawa County, Early July.

 

252. Branches off the main shoot- Secondary shoots (shoot "branches" or, more descriptively, "limbs") growing out of intercallary meristem (in leaf axils) of the primary shoot in velvet panicgrass. These shoots were growing at outer edge of a oak-hickory/tallgrass savanna in the western Ozark (Springfield) Plateau. There was a terminal panicle at apex of each secondary shoot. These were the vernal flower clusters that were approaching maturity in early summer. The rosette panicgrasses (Dichanthelium taxon) produce both vernal and autumnal sexual shoots (weather permmitting).

Ottawa County, Early July.

253. Axillary outgrowths- Secondary shoots ("limbs off of the main stem" so to speak) arising or coming out of the f leaf-culms axils in velvet panicgrass. These secondary shoots produced grain-filled panicles in the late spring-early summer period. These are the vernal sexual shoots and flower clusters (versus autumnal counterparts) of these two-flowering season, cool-season (C3 photosynthesis) grasses. These plants were growing on the exterior of an oak-hickory/tallgrass savanna in the westrn Ozark Highlands.

Ottawa County, Early July.

 

254. Velvet flowers and fruits in the Ozarks- Vernal sexual shoots and panicles of velvet panicgrass. These shoots/inflorescences were on some of the same shoots that were featured above. The rosette panicgrasses (Dichanthelium subgenus of Panicum) typically bear both vernal and autumnal sexual shoots. The first of these three slides showed a panicle emerging from the boot (the leaf sheath that envelops, surrounds, the developing inflorescence that will subsequently grow and elongate out of this part of the shoot). After emergence of the inflorescence the leaf that previously enclosed, enveloped, this inflorescence will become the flag leaf (the leaf subtending, immeditely below, the flower cluster). This emerged stage of development (complete with flag leaf) was shown in the second slide. The third slide showed spikelets on seondary branches at base of the panicle (that portion of the flower cluster immediately above the flag leaf).

Another phenomenon seen in these photographs was the inward folding (ie. folding up from sides or lateral surface) of velvet panicgrass leaves in response to exceptional drought. The preceding winter and early spring of the current growing season had been wet, but early onset of the usual late-summer high pressure system brought extreme heat and dryness (drought) and severe stress on plants. Closing up of the broad leaves of velvet panicgrass was an apparent adaptation of this C3, cool-season species that demonstrated natural selection over generations (evolution). With abundant reproduction, both asexual and sexual, there was no doubt that the velvet panicgrass featured here was a fit survivor--where fitness is defined as producing enough progeny to perpetuate its species. What lessons God's garden teaches to those who are inclined to learn their range lessons.

Ottawa County, Early July.

 

255. Soft as velvet- Detail of shoots, with emphasis on culms and leaf sheaths, of velvet panicgrass. The velvety pubescence of culm and leaf was visible in these three photographs. Also clearly visible in these slides was the characteristic (in fact, a nearly tell-tale feature) of a glabrous, glandular subunit of the internode immediately below atttachment of leaf sheath.

Ottawa County, Oklahoma. Early July.

 

256. Leaf-tails- Abaxil (upper) surface of a leaf of velvet panicgrass accompanied by a folding-up leaf showing response to early season drought. The "notch" removed from the outer edge of the large leaf was made by a feasting katydid.Ottawa County, Oklahoma. Early July.

 

257. A convenient lineup that featured slimleaf or linear-leafed panicgrass (P. linearifolium = D. linearifolium)- Two plants of slimleaf panicgrass (two center cespitose plants with dense shoots) with one plant of Scribner's panicgrass on either side. All four of these individuals were in fruit stage ranging from milk to grain-ripening phenology. This opportune "Kodachrome moment" exemplified the fact that various species of the rosette panics frequently grow in close spatial association with each other. It is less common to find two or more of these obviously different species with such drastically different leaf features as to afford a striking contrast and show to such advantage this niche phenomenon. OK, it was not Darwin's finches in the Galapagos Islands, but it did shown speciation in the oak-hickory-tallgrass savanna of the Ozarks. That should get honorable mention by hillbilly standards.

Ottawa County, Oklahoma. July (milk to grain-ripening phenological stages).

 

258. Two fine plants- The two individual plants (geneotypes) of slimlieaf panicgrass introduced in the immediately preceding slide were viewed at closer distance in these two photographs. The seocnd photograph focused on the plant shown on the right in the first photograph in order to present the entire shoot complex including the panicle. Leaves in background of this second slide were those of Scribner's pancigrass as portrayed in the "lineup" of the immediately preceding slide and its caption.

Ottawa County, Oklahoma. July (grain-ripening phenological stage).

 

259. Basal details- Basal shoots of the plant of slimleaf panicgrass shown in the right in the immediate last three slides.

Ottawa County, Oklahoma. July (grain-ripening phenological stage).

 

260. Fat fruits of a slimleaved parent- Panicle and spikelets of slimleaf panicgrass. This was one of several panicles on the slimleaf panicgrass plant shown on the right in preceding slides. This panicle was removed and placed on a dead leaf of black oak, the dominant tree species of the oak-hickory-tallgrass savanna that this nifty specimen was a member of. First photograph presented overall panicle features while the second showed spikelet arrangement and features.

Ottawa County, Oklahoma. July (grain-ripening phenological stage).

There are also Paspalum species in the oak-hickory-tallgrass savanna that includes much of the Ozark Plateau Region and extending into the eastern Cherokee Prairie of the Central Lowlands. There are not as many Paspalum species as there are rosette panicgrasses in the oak-hickory-tallgrasss savanna, and they are certainly less confusing and controversioal in this range type. Two of these paspalum (the name is used as both noun in the common name and genus of the binominal) species were included below.

 

261. Florida paspalum (Paspalum floridanum)- This rhizomatous species is one of the larger, more robust paspalums in the central savannah, including the Prairie Peninsulah. This, like most Palpalum species in North America, is not typically common, occurring instead as scattered individual plants. Florida paspalum is far from common in the Ozark Plateau Region, but at its wesern margin where it contacts the Cherokee Prairie P. floridanum is a frequently encountered decreaser grass.

The plants shown in these two slides were growing in associagtion with big bluestem in the understorey of black oak-dominated savannah oa tripoli-based, shallow, rocky soil. This was in a year of above average precipitation with consistently moist soil from winter through spring and early summer. Tallest sexual shoots on these plant exceeded a yard in height.

Ottawa County, July: anthesis.

 

262. Basal shoots of Florida paspalum- A plant of Florida paspalum of typically large size with an exceptional number of tillers. The wider-spaced shoots at outer edge of the plant (foreground) had arised from short rhizomes that are characteristic of this species. Purplish-tinged leaves comprise another common characteristic of Florida paspalum. This coloration frequently "tricks" even experienced rangemen who on seeing it from considerable distances mistake it for big bluestem until they examine it at "close-quarters". This specimen was growing with beaked panicgrass, field paspalum, and big bluestem as well as naturalized tall fescue (Festuca arundinacea) on a prairie that adjoined a black oak-tallgrass savannah.

Ottawa County, July:

 

263. Hair to spare- Axillary portion of shoot of Florida paspalum. Plentiful pubsecence on this individual clearly marked it as a plant of Paspalum floridanum var. floridanum in contrast to P. floridanum var. glabratum (Kocera, 2003, p. 245). This shoot was on one of the plants shown in the photographs before the preceding one.

Ottawa County, July.

 

264. Growing out of the boot- Panicle of Florida paspalum emerging from the enveloping leaf sheath known as the boot. The boot is the uppermost leaf sheath on the culm (ie. it subtends the inflorescence). This uppermost entire leaf is the flag leaf. The newly exposed inflorescence will be ready to assist is and receive cross-pollination as anthers and stigmas are expressed.

Ottawa County, Oklahoma. Late July, still pre-bloom stage.

 

265. Out of a hairy boot- Spikelets of florida paspalum at base of inflorescence as it was emerging from the boot. This was one of the lower branches on the inflorescence presented in the immediately preceding two-slide set.

Ottawa County, July: anthesis.

 

266. Flowering P. floridanum in the Ozarks- Inflorescence on a plant of Florida paspalum growing in understorey of black oak-big bluestem dominated savannah in the western Springfield Plateau section of the Ozark Plateau physiographic province. This was one of the several plants featured in the first two slides that introduced this species.

Hitchcock and Chase (1951, p. 599) described the Paspalum inflorescnce as consisting of "one to many spiikelike racemes, solitary, paired, or several to many on a common axis" while Kucera (1998, p.239) interpreted the inflorescence as "...consisting of one to several spicate racemes or branches along the flower stalk, or sometimes in conjugate pairs at apex..."and Barkworth (2003, p. 566) described the Paspalum inflorescences as "... panicles of one-many spikelike branches, these digitate or racemose on the rachis". In an interesting, revealing, and/or ironic twist in sytematics and identification of grasses, it was the description in Barkworth et al. (2003, p. 566) that most closely fit the description of the Paniceae inflorescence written by Agnes Chase (1937, p. 98; 1964, p.98) in her First Book of Grasses: "Inflorescence a panicle or of one to many racemes, these digitate or racemose on the main axis". Unquestionably, the description in Barkworth et al. (2003, p. 566) was an almost verbatim copy of that written by Chase (1937, 1964) almost seven decades earlier. The description contained in Barkworth (2003) was a slightly rephrased version of and certainly a description much closer to that of Agnes Chase than that the description of the Pasopalum inflorescence in Hitchcock and Chase (1951) which did not provide a description of the inflorescnce of Paniceae.

Ottawa County, July: anthesis.

 

2567. Flowering and fruiting in the Ozarks- The first of these two photographs showed anthers and stigmas on a branch of the inflorescence of Florida paspalum presented in the last two-slide set. It was upper part of the lower branch on the right side of the inflorescence in both of those slides. The second of these photographs showed spikelets at somewhat closer distance. These spikelets, with only stigmas exerted (on left branch), were one to two days older than those shown at pea anthesis in the first slide.

Ottawa County, July: anthesis.

 

268. 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.

 

269. 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.

 

270. 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.

 

271. 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) .

 

272. 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.

 

273. Foolproof feature- 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.

 

274. 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.

 

275. Fringed to its base- Basal shoots of thinseed or thin paspalum with typical features of culm and leaf including some wavy leaf margins which--along with hairs on margins--is the basis for common name of this abundant Cross Timbers grass. This same feature of crinkled or wavy leaf margin was also visible in most succeeding slides of fringeleaf paspalum. For whatever reason, Barkworth et al.(2003, ps. 589-592) did not make note of these features, but these characteristics were clearly shown in line drawings of P. setaceum var. ciliatifolium.

Erath County, Texas. Early September; early growth stages (two to four leaf progressions).

 

276. Shoot specifics- Details of upper shoot of fthinseed paspallum including the full-grown phytomer immediately below the upper-most node. The phytomer is the fundamental unit of the grass shoot including the upper half of the lower node, lower half of the upper node, the internode in between these two nodes, and the one whole (entire) leaf of this node/internode area. The shoot apex contains the infloresence of this sexual (sexually reproductive) shoot. The upper-most, young (immature) leaf that is rolled inward is the flag leaf, the leaf immediately below (subtending) the inflorescence. This arrangement or grass units (organs) was presented in the two slides immediately below.

Erath County, Texas. Early September; immediate pre-bloom stage of phenology.

 

277. Coming up and going out- Two stages in exertion of inflorescences in thin or thinseed paspalum. The envaginating unit of culm surronding the flower cluster is known as the boot. The leaf that will be below (subtend) the entire inflorescence is the flag leaf. It is the upper-most leaf in both of these photographs. Students should observe that anthesis had already commenced even before the inflorescence was fully exerted (second of these photographs).

Erath County, Texas. Early October; early bloom stage of phenology.

 

278. Fringes on the top- Inflorescence of thin or thinseed paspalum at anthesis. Chase (1937, p. 98; 1964, p.98) simply and eloquently described the Paspalum infloresecence as "a panicle or of one to many racemes" with these racemes (or racemose branches) being digitate or racemose on the central axis of this entire flower cluster. A brief review of the various interpretations of the Paspalum inflorescence was given above in conjuction with Florida paspalum.

Erath County, Texas. Early October; anthesis.

 

279. Purpletop (Tridens flavus) in understorey of West Cross Timbers- Whereas purpletop is an increaser species on most tallgrass prairie range sites it is characteristically a decreaser tallgrass on many prairie-oak/hickory forest savannas or ecotones. Purpletop readily populates open grasslands in humid and subhumid regions, but it excels as an understorey species in savannas and open-understorey deciduous forests where it often has a competitive advantage over otherwise dominant tallgrasses like big bluestem, Indiangrass, and little bluestem all of which have relatively greater light requirements. Purpletop, however, does have greater demands for sunlight than such forest understorey grasses as woodreed grass and beakgrain (these latter two grass species were presented under Oak-Hickory Forest and Southern and Central Forests sections).

The purpletop specimen shown here was growing at outer edge of a West Cross Timbers community on a site comprised of deep sand (Sandy Loam range site). On such sites where purpletop is often the dominant grass, little bluestem is commonly the associate species with sand lovegrass and nodding wildrye also common. A large clump of little bluestem accompanied the purpletop speciment seen here (former to left-rear of latter). In the background common greenbriar had ascended into branches of post oak treaes.

Purrpletop is widely distributed throughout the savannahs of the Ozark Plateau Region and into those of the eastern Cherokee Prairie of the Central Lowlands physiographic province. Purpletop is also common in the Blackland Prairie (on what tiny remnants remain protected from conversion to cropland or urban sprawl) and the Pineywoods of east Texas. The height of purpletop (usually in excess of a yard in stature and, frequently, exceeding five feet) is such that it is, strictly speaking, a tallgrass species.

General references for purpletop included the Phllips 66 Pasture and Range Plants (Phillips Petroleum Company, 1961; Nicholson, 2006, p. 70) and Tyrl et al. (2008, ps.178-179). Purpletop is an Increaser on most range sites, but it does provide forage of Good to Excellent feed value when young and is a Decreaser on certain forest, woodland, and savannah sites including some in the Ozark Plateau, Prairie Peninsula, and Pineywoods regions and areas. Purpletop is also one of the most prolific species on old fields ("go-back land") at mid- to late seral stages.

Hunewell Ranch, Erath County, Texas. October.

 

280. Purpletop panicle- Panicle of right-most purpletop shoot shown in preceding slide. Hunewell Ranch, Erath County, Texas. October.

 

281. Old-field dominated by purpletop- Former (=abandoned) cropland adjacent to an oak-hickory forest in Ozark Plateau supported essentially a single-species stand of purpletop. This was roughly advanced mid-seral stage on go-back land in tallgrass prairie that was a large glade within an open-understorey oak-hickory forest or perhaps oak-hickory-tallgrass sananna cover type. Approximately 20 to 30 years post-farming (small grains) this old-field vegetation had developed (progressed) to a community composed almost exclusively of purpletop (purpletop population would probably be a more accurate description). Stands of purpletop of this "purity" and vigor were (are) not common (peak standing crop in one of wettest summers on record), but they are far from rare in the Ozark Mountains. This example demonstrated the extent to which purpletop can dominate certain go-back pastures once the seral community dominated by broomsedge bluestem has been succeeded. Continued overgrazing of such go-back land will maintain the broomsedge stage indefinitely. For practical purposes this is likely the most successionally advanced vegetational stage possible in realistic human time scale. Such beautiful stands should be cherished and protected from future abuse. This was a lucky and forgiven landowner.

Newton County, Missouri. September (purpletop should NOT be cut for hay or grazed heavily at this advanced phenological and late seasonal stage as such severe defoliation will not allow adequate replinishment of reserve nutrients for overwintering survival).

 

282. Even in drought- Two representative examples of purpletop growing on the floodplain of a small river in the West Cropss Timbers of northcentral Texas under conditions of Extreme Drought (Palmer Index). Drought conditions recently moderated by light showers had permitted these plants to progress to the flowering stage of their annual cycle. Four plants growing in close proximity were presented in the first of these two slides whereas one plant was shown in the second slide.

Purpletop is a cespitose species, but it does have rhizome-like structures at its rootcrown (Gould, 1975, p. 205) and described by Barkworth et al. (2003, p.39) as "knotty, shortly rhizomatous".

Terrace of Bosque River, Erath County, Texas. Early November; late anthesis to immediate post-anthesis stage of phenology.

 

283. Upper purple shoots- Phytomer--the fundamental shoot unit of node and internode of culm and associated leaf--of shoot (first photograph) and closer view of two partial phytomers (second photograph) of purpletop growing on a floodplain in West Cross Timbers of northcentral Texas. Phytomers seen here were immediately below base of panicle. These shoots were produced in Extreme Drought (Palmer Index).

Terrace of Bosque River, Erath County, Texas. Early November; flowering phases of phenology.

 

284. Basal purple shoots- Basal portions of shoots (first photograph) and details of basal leaves (second photograph) of purpletop growing on a floodplain in the West Cross timbers dring an Extreme Drought (Palmer Index).

Terrace of Bosque River, Erath County, Texas. Early November; flowering phases of phenology.

 

285. Purple-shooted details- Closeup photographs presenting sideview (first slide) and dorsal view (second slide) of axillary area of shoot and specifics of node/internode portion and sideview of leaf sheath (third slide) of purpletop on a floodplain in the West Cross Timbers.

Terrace of Bosque River, Erath County, Texas. Early November; flowering phases of phenology.

 

286. Purple-topped panicle- Overall or general view of a panicle (first slide) and primary/secondary branches with flowering spikelets (second slide) of purpletop on a floodplain in the West Cross Timbers. These spikelets quickly developed to anthesis during Extreme Drought (Palmer Index) following rain showers that brought moderate amonts of precipitation in mid-autumn.

Terrace of Bosque River, Erath County, Texas. Early November; first slide, late anthesis to immediate post-anthesis stage of phenology and peak-anthesis, second slide.

 

287. Will have to do- Two "make do" views of the tiny, flittering spikelets of purpletop at peak-bloom on the floodplain of a small river in the West Cross Timbers of northcentral Texas. Spikelets had achieved flowering status during Extreme Drought (Palmer Scale) and shortly after "showers of blessing" gave plants a "shot in the shoot" that permitted a brief flowering period.

Admittedly these are not particularily revealing images, but they were taken at 1/15 second (using ASA 100 ProviaF film) under a bottomland forest canopy. In short, images were the best this photographer could do photographing small organs under "in-the-field", "on-the-land" conditions. Fortunately for survival of this eragrostoid species, plants of purpletop are much more adapt at doing their thing than the author was at doing his.

Terrace of Bosque River, Erath County, Texas. Early November; peak-anthesis stage of phenology.

 

288. Another approach- Branch (upper slide) and group of spikelets (lower slide) of purpletop at anthesis. This specimen was growing in the West Cross Timbers. Photographed on a hot (100 F), sticky (60% humidity) afternoon; conductions conducive for pollination, but not the best photography. (Though certainly not enjoyable such an abiotic environment does make photogaphers appreciate the blessing of cool water).

Erath County, Texas. Early September (with a lot of summer left to go down Texas-way).

 

289. Sand lovegrass (Eragrostis trichoides)- The adjective "sand" is the defining one for this tough and delicately beautiful native grass that is found almost exclusively on the sandier sites . Sand is also usually the defining soil seperate of the Cross Timbers. Sandstone is a major parent material of soils that support Cross Timbers vegetation. It addition, sand allows more favorable soil moisture conditions because high percentages of sand allow greater rates of water infiltration and consequently more mesic plant communities than on adjoining soils with less sand.

This is the chressard (soil water available for plant use) concept of Clements ((Clements, 1920, 26-28; Weaver and Clements, 1938, p. 203). According to Clements' comprehensive succession model certain savanna vegetation like the Cross Timbers is postclimax to true climatic climax. Dominant tree species have been able to persist only because of the greater chressard of sandy soils (Weaver and Clements, 1938, ps. 509 and, especially, 515 which was accepted and quoted by Dyksterhuis, 1948, p. 328).

The sandy soil-requiring sand lovegrass indicates presence of soils high in sand and this species is thus a textbook example of another Clementsian conept, indicator plant or species (Clements, 1920, especially ps.19-25, 270-335). In this context, sand lovegrass is an indicator (hence, indicator plant) of edaphic conditions (soil indicator) in contrast to other factors or stresses such as grazing (grazing indicator). Sand lovegrass is also sensitive to heavy grazing (perhaps partly in response to competition from accompanying, shade-casting trees on sandy soils) and is categorized as a decreaser. This showy grass is thereby an indicator species in at least two senses of the indicator plant concept.

Astute students should also have picked up on the fact that Dyksterhuis accepted and applied both Clements' chressard concept and his indicator species concept that institutionalized range condition and trend analysis based on plant responsees to grazing (Dyksterhuis, 1949).

See what lessons can be learned in Plant Ecology as well as in the sociological aspects of professions from one grass plant growing in the sand!

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

 

290. Panicles of of sand lovegrass- The flower clusters of sand lovegrass often make up over half of it's total shoot length. Sand lovegrass is one of the most showy plants that can be used as an ornamental when landscaping with native species. To the point of this discussion, sand lovegrass is quite sensitive to improper grazing (namely overuse) and it's common occurrence on ranges such as those of Cross Timbers vegetation indicates careful management and concerned husbancry (or protection from grazing).

Tarleton State University Hunewell Ranch. Erath County, Texas. October (at anthesis stage).

 

291. Dense enough- Local assembly (population) of several plants and a lot of shoots of sand lovegrass in the West Cross Timbers. Strictly speaking the population of sand lovegrass growing here was at the edge of (a local ecotone between) the West Cross Timbers and the Grand Prairie, but a large microsite of deep sand provided the defining habitat of this decreaser grass species.

Tarleton State University, Hunewell Ranch. Late October (and still over 90 degrees Fahrenheit); peak standing crop with maturing caryopses.

 

292. Sterling citizens- Some unusually large and robust plants of sand lovegrass in West Cross Timbers. Sand lovegrass was highly recommended for those who enjoy landscaping with native plants. Ornamental value of this native, perennial, warm-season grass is most obvious at full plant maturity and peak standing crop when panicles of this essentially xerophytic species can reach lengths/heights of over a yard. These grain-laden panicles typically droop over rather than reaching heights corresponding to panicle length, but this just makes sand lovegrass all the more showy and graceful. The upright plant in the upper (vertical) photograph extended to the 42 inch mark on the Lufkin folding rule.

Unfortunately, this relatively grazing-tolerant grass is so palatable that it has been greatly reduced overgrazing so sadly common on ranges in Texas and southern Oklahoma.

Tarleton State University, Hunewell Ranch. Late October (and still over 90 degrees Fahrenheit); peak standing crop with maturing caryopses.

 

293. Shoot details- Basal and lower portions of shoots (upper slide) and particulars of a phytomer (lower slide) of sand lovegrass in the West Cross Timbers. The pilose pubescence in the leaf axil (collar) is diagnostic in identification of various grass species, but the large, almost-plumose hairs of sand lovegrass are particularly distinctive.

Tarleton State University, Hunewell Ranch. Late October (and still over 90 degrees Fahrenheit); peak standing crop.

 

294. Spikelets on sandstone- Tiny spikelets of sand lovegrass lying against the parent material of Cross Timbers soils. Sand lovegrass usually has four to 18 florets (Gould, 1975, p. 185). These spikelets were gathered from the plants shown in the preceding slide.

Bosque River, Erath County, Texas. October, grain-ripe stage.

 

295. Texas cupgrass (Eriochloa serica)- This leafy, panicoid bunchgrass is extremely palatable to livestock and is another example of a decreaser. Texas cupgrass is a prairie grass, but it also grows in savannas such as the Cross Timbers and Edwrds Plateau. Texas cupgrass is usually not a common or major species even in relict sites, but it's presence is evidence of proper grazing management (or exclusion from grazing). This is a grazing indicator species.

Fort Bowie. Brown County, Texas. May.

 

296. Inflorescence of Texas cupgrass- Set of three photographs to show the arrangement and details of spikelets in the contracted panicle of this panicoid grass. In this species, spikelets are arranged along unbranched primary branches that lie closely --almost adnate--to the central axis of the panicle (Gould, 1975, p. 433). Hitchcock and Chase (1950, p. 588-589) interpreted the inflorescence of Texas cupgrass as consisting of racemes along a rachis. The interpretation by Gould (1975, p. 433) was consistent with recognition of six subfamilies of Gramineae and more phylogentically precise than the more artificial approach of only two broad subfamilies used by Hitchcock and Chase (1950).

The unique floral feature of this species is the small indentation or depressed portion of the branch in which the caryopsis is situated, hence the common name of cupgrass (ie. depression in which the grain lies is a "cup").

Flower cluster and spikelets of a decreaser and long day plant (species that flower when day length is getting longer or, often more importantly, when night length is getting shorter; usually corresponds to more than 14 hours of daylight).

Edwards County, Texas. October, soft to-mid dough phenological stage.

 

297. Three-flower melic (Melica nitens)- This member of the small Meliceae (melic tribe) is one of the largest cool-season native grasses in the tallgrass savanna and the ecotone between the oak-hickory forest and the tallgrass prairie. Three-flower melicgrass is common on limestone outcrop habitats and where it can get "full sun" as at forest edges and openings and on glades as well as within the savanna. It does well under disturbances like fire, forest windthrow, and shredding of small trees and shrubs under powerline corridors like the one that this robust four-foot specimen called "home".

Ottawa County, Oklahoma. June.

 

298. Panicle of three-flower melic- Ottawa County, Oklahoma. June.

 

299. Spikelets of three-flower melic- Ottawa County, Oklahoma. June.

 

300. Nimblewill (Muhlenbergia schreberi)- This eragrostoid grass (Eragrosteae tribe) is another widely distributed grass in understories of oak-hickory forest ranges. Nimblewill grows on a diversity of habitats ranging from fairly dense, mesic woods to open prairies on shallow soils and as a component of stremside vegetation to a common species on abandoned fields (= go-back land) or even old city lots and schoolyards. It is generally accepted that nimblewill is probably the best-adapted of any muhly in the southern oak-hickory forests and savannahs, including adaptation to disturbances like overgrazing and trampling. Nonetheless, nimblewill was interpreted by Tyrl et al., 2002, p. 109) as "... characteristic of the mid to late stages of plant succession".

Although nimblewill is neither rhizomatous nor stoloniferous it frequently grows as localized colonies especially in forest clearings, lanes, old barnyards, etc. This habit is due to adventituous rooting at nodes of the low, decumbent tillers from which short-branched, largely unspread panicles arise. Nimblewill has a unique feature (unique among North American Muhlenbergia species) in that morphology or growth form differs greatly from the early season, short plants with broad leaf blades to the late-season form with ascending to even sweeping shoots.

Nimblewill specimens presented here were growing in shade of post and blackjack oaks at extreme western edge of Ozark Plateau where relatively large-sized tallgrass prairies and oak-hickory forests developed in a complex vegetational mosaic. This patchwork of range vegetation was part of the southern extension of the prairie peninsula.

Ottawa County, Oklahoma. December: hibernal aspect, dormancy.

 

301. Shoot tips of nimblewill- Photograph of shoot apices of nimblewill with remnants of mature inflorescences (seed shatter stage) many of which were either reduced to their bare floral axisis or still partially to largely enclosed in the boot,. These are common or even characteristic late-season features of the spike-like panicles of this species.

Ottawa County, Oklahoma. December: hibernal aspect, dormancy.

 

302. Local stand of nimblewill- This small colony or population of nimblewill shoots (along with a few shoots of Carex molesta) was growing along the edge of an oak-hickory Ozark forest. This provided an example of typical habit and morphology of this eragrostoid grass. Nimblewill is not a heavy or high herbage-producer, but it is one of the best adapted native grasses to stress caused to a combination of defoliation and shade.

Ottawa County, Oklahoma. July; immediate pre-bloom stage of phenology.

 

303. Shooting in the shade- - Shoot of nimblewill growing in its common and, apparently, "prime habitat" in the Ozark Plateau. In shade of post and blackjack oaks, black cherry, and hackberry and amid blackberry canes this nimbleweill plant had produced a robust (for this species) shoot. Shoots of nimblewill are, as revealed here, distinctive among grasses.

Ottawa County, Oklahoma. July; immediate pre-bloom stage of phenology.

 

304. Shooting a panicle- Sexual shoot of nimblewill with largely exerted panicle. Typically the panicle of nimblewill remains partly enveloped in the boot (encased in the leaf that subtends the inflorescence) so that branches off of the central floral axis appear appressed and ascending so as to resemble a semi-condensed panicle. Traditionally the fruit of Muhlenbergia species has been interpreted as an achene rather than a caryopsis. Spikelets in this panicle were pre-anthesis.

Ottawa County, Oklahoma. July; early summer.

 

305. Midgrass in hardwood-tallgrass savanna- Local colony of western wheatgrass (Agropyron smithii) growing in shade of post oak (Quercus stellata) in an oak-hickory-big bluestem savanna in extreme western edge of Ozark(Springfield) Plateau. Presence of a native midgrass species that is a dominant in mixed prairie in a humid 40 inch precipitation zone in the Ozark Highland struck this author as unusual so this example of western wheatgrass was included here.

Another interesting phenomenon (again, at least to this rangeman) was the fact that this colony of a cool-seaon, rhizomatous species "just suddenly appeared from out of nowhere" on the boyhood home of the author. Just a short distance from the barn on this small farm western wheatgrass emerged and produced a grain crop (see immediately below) on land where the grass-wise author, after having walked over this local spot for over half century, had never found it. The simple explanation was that this barnyard had been heavily grazed by cattle, sheep, and horses for decades. When the last of the livestock were sold the author's father mowed this parcel with fervor and zeal so that the grass community was primarily Japanese chess (Bromus japonicus) with some Kentucky bluegrass (Poa pratensis) and tall fescue (Festuca arundinacea) in the cool-season and hairy crabgrass (Digitaria sanguinalis) with some common bermudagrass (Cynodon dactylon) in the warm-season. When the author's 91-year-old father reduced frequency of lawn-mowing this colony of wheatgrass "magically appeared" and crowded out (out-competed) exotic annual and perennial grasses. This was within one year after having heavily grazed and mowed (mostly overgrazed and overmowed) for three-fourths of a century. Obviously, some western wheatgrass had persisted under some or all of this abuse, but when did propagules arrive? How long had there been some western wheatgrass on this land before it could finally could reproduce sexually?

"Banished from the thoroughfare and the field, it bides it time to return, And when vigilance is relaxed, orthe dynasty has perished, it silently resumes the throne from which it has been expelled, but which it never abdicates." ---Senator John James Ingalls, 1872

Ottawa County, Oklahoma. Mid-June; late anthesis.

 

306. From out of nowhere- The local colony of western wheatgrass that was introduced in the immediately preceding slide shown two years after the above view. This colony of western wheatgrass (asexual or sexual origin of individual shoots was unknown) had about tripled in cover in this two-year time period--and all of those years had been from Severe to Extreme Drought with a short period of Extraordinary Drought (Palmer Index), but shoots had not been grazed during this time. This local population of western wheatgrass spread outward from limestone rock outcrops into an area that had been mowed by a riding lawnmower for about 30 to 35 years prior to first notable appearance of western wheatgrass (ie. mowed for a third of a century until about four years ago) . When routine close mowing ceased, western wheatgrass could survive beyond the rock outcrop habitat so that it spread into the adjoining area as shown here.

Appearance of this population of western wheatgrass was a mystery (or a miracle) because the location had been a barnyard, just a manure shovel-throw away from a cattle barn, until it became a pasture for trail-ride horses, and, finally, an extension of a domicile yard. This western wheatgrass was growing at the edge of shade cast by senescing post oaks and close to blackjack oaks on what appeared to be an oak-hickory-tallgrass savanna. Prior to the land being a barnyard, it had been along the eastern edge of the the famed cattle trail known most commonly as the Sedalia Trail or, sometimes, the Kansas Trail.

In the late 1930s an old man who had trailed cattle up from Texas on the Sedalia Trail stopped by this author's grandmother's house and asked if there was a spring close by. The trail drover remembered the spring as being just west of the Missouri state line in Indian Territory. Was there a spring around here? Yes, Grandma pointed to where an old spring had been until it dried up with hand-digging of a shallow well some years before. There was the remains of an old springbox there. The old man got excited and asked Grandma if he could go over and look for the spring. Yes, certainly, and she directed him to a low hillside beyond her house. When the old man returned he exclaimed that it was the spot where they had watered Texas cattle on the drive up into Missouri. Grandma asked the man if they were trailing cattle to the Joplin Stockyards. No, those yards were not there then. They drove their cattle to Sedalia, Missouri.

Grandma then asked the old trail driver why a spring was so important when there were all these creeks around. Trail drovers tried not to water cattle in creeks he answered. Grandma asked the drover why not. Too many bluffs above creeks and hard to find ways to get cattle down through the bluffs. Also, cattle could sometimes get bogged down in creek mud. Besides, trail hands could lose cattle in the trees. Well, how did the trail drivers keep from losing cattle in all the trees around here Grandma wanted to know. Weren't many trees then other than some in little patches, stands, groves (whatever name he called those small groups of trees). It was all prairie then. Grandma said he called it "prairie".

Obviously most of the surrounding trees--some of them big ones--had established after closing of the Sedalia Trail and turning the tallgrass prairie here on the western Ozark (Springfield) Plateau into stock farms (initially dairy farms) that soon had so many hardwood trees the land resembled a savanna (at best) and, frequently, oak-hickory forests.

That dried-up, trail-drivers spring was about 125 yards from where this patch of western wheatgrass grew. Miraculously, a remnant of that midgrass had persisted amid the limestone rocks that had once been part of tallgrass prairie through trail driving days to conversion into small dairy operations (and overgrazing), horse-pasturing, to lawn-mowing. When the pattern in which one regime of heavy defoliation following another regime ceased, the western wheatgrass regained its rightful place in the former Ozark prairie. An ecological tale in time and place.

Ottawa County, Oklahoma. Mid-June; grain in milk stage.

 

307. An evidentiary species (or Not one expected in a tallgrass savanna)- Shoots of western wheatgrass in the local colony featured in the preceding two slide/caption sets that had re-established after a historical march of the whiteman went through cattle trail, stock farming, past-time horse grazing, and lawn-mowing. In earliest oral accounts this land had been tallgrass prairie that was first used by whiteman for trailing cattle to market (eastern edge of Sedalia Trail). Current vegetation had gone to an oak-hickory savanna with most of the native grass and forb species extirpated.

In absence of grazing and machine mowing for about three years western wheatgrass had returned--during Severe to Extreme Drought--to its former place spreading from a remnant that had been protected from extreme defoliation for decades in a rock outcrop.

Readers may recall that investigators were eventually able to solve the terrorist case of airline sabatoge that came to be known as the Lockerbie Scotland bombing from specific traces of evidence found on the debris field of Flight 103. Presence of western wheatgrass on this pasture served as ecological evidence. The trace of western wheatgrass was not so much a mystery as the solution to something of an ecological mystery. Western wheatgrass occurrence was unequivacal proof that the pre-settlement vegetation on this land had been tallgrass prairie or, perhaps enough scattered trees for a tallgrass oak-hickory savanna. Western wheatgrass is not widespread in the Ozark Plateau, but is is native to this area (Steyermark, 1963, p.122). Neither is western wheatgrass typically a common species on tallgrass prairie, but natural regeneration of this cool-season midgrass species in this tallgrass prairie area under conditions of heavy grazing over the last century confirmed that native grassland or, maybe, savanna--and not oak-hickory forest--was the climax plant community, the virgin vegetation before it was became a casualty of the whiteman's advance.

Ottawa County, Oklahoma. Mid-June; grain in milk stage.

 

308. Throne reclaimed; reign resumed- Spikes with anthers of western wheatgrass. These inflorescences were on tillers in the local colony of a Ozark Plateau savanna that was presented and explained in the immediately preceding caption. After a span of three-quarters of a century of extremely heavy defoliation by grazing and mechanical mowing yet with only one year for recovery this plant flowered and produced caryopses. The above quoted Kansas Senator put if forthrightly, "[G]rass is immortal.".

A typical spikelet of western wheatgrass has between three to, sometimes, fifteen florets (Great Plains Flora Association, 1986, p.1126).

Ottawa County, Oklahoma. Mid-June; anthesis.

 

309. Silky in a savannah- Silky wildrye (Elymus villosus) in understorey of a range plant community dominated by black oak (Quercus velutina). This range vegetation included both an open canopy oak-hickory forest and an oak-hickory savanna with three herbaceous layers. These three layers were:1) tallgrass of big bluestem (Andropogon gerardii), Indiangrass (Sorgastrum nutans), and silky wildrye; 2) midgrass of Canada or nodding wildrye (Elymus canadensis) and velvet panicgrass (Panicum scoparium); and 3) low-growing grass, especially poverty oatgrass (Dantohona spicata), and woodland forbs.This vegetation also included at least two shrub layers, including one co-dominated by highbush blackberry (Rubus pensilvanicus) and smooth or scarlet sumac (Rhus glabra). This plant was growing in a local transition zone between these two range plant communities.

Shoots shown here were on a single plant. These tillers approached seven feet in height.Tillers of silky wildrye were shown from mid-shoots to spikes (first slide) and from immediately below flag leaf up through spike (second slide). The shoots of this cool-season member of the barley tribe had retained basal leaves and short culms throughout a winter that had temperatures of at least -20 degrees Fahrenheit. This "twenty below minus some" temperature was the all-time cold record for this area. Obviously silky wildrye is is well-adapted to its environment..

Foliage around these shoots was that of highbush blackberry and smooth or scarlet sumac.

Ottawa County, Oklahoma. Early July; peak standing crop, anthesis.

 

310. Coarse silk- Spike (first slide) and spikelets (second slide) of silky wildrye. There are usually only two (less commonly, one or three) spikelets, each of which most commonly bears one or two (rarely three) florets, per node of the rachis joint (Great Plains Flora Association, 1986, p.1169-1170). Spikes, like the shoots they adorn, of silky wildrye are considerably larger than those of Canada wildrye and Virginia wildrye (Elymus canadensis, E. virginicus) when these three species are found on the same or similar ranges within a general area. This obviously variable comparison combined with the smaller, shorter spike with often awnless to short-awned lemmas of Virginia wildrye and the arching over to nearly horizontal spike of Canada or nodding wildrye that contrast with the vertical to vertically arching spike with long lemma awns of silky wildrye make for handy distinction among these three species when on the range.

Ottawa County, Oklahoma. Early July; anthesis.

 

311. Silky later- Four ( you have to count carefully) sexual shoots each topped with a large spike of sliky wildrye growing in an oak-hickory/tallgrass savanna in the western Ozark (Springfield) Plateau. Compound, broad leaves surrounding wildrye shoots were those of highbush blackberry which occurred as occasional thickets (colonies) on both savanna and in understories of surrounding oak-hickory forests.

Caryopses of were approaching maturity (grain-ripe stage of phenology).

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

 

312. Nodding off- Spikes of Canada or nodding wildrye (growing with velveet panicgrass) growing within a dumped pile of field rock at edge of an oak-hickory/ tallgrass savanna in the western Ozark Plateau. This plant was growing about 250 yars away from the silky wildrye featured immediately above. When these two species (and sometimes along with Virginia wildrye for three Elymus species) the larger silky wildrye mature slightly (a few weeks) later.

Ottawa County, Oklahoma. Early July; anthesis.

In the "dead of winter": the following short section presented two cool-season (C3) grass species growing on tallgrass-oak/hickory savanna in the western Springfield Plateau.

Two that can take the cold- The winter rosette of a Panicum species in the Dichanthelium subgenus (Dichanthelium taxon levated to its own genus by numerous workers) known variously hairy panicgrass, hairy rosettegrass, wooly rosettegrass, wooly panicgrass, western rosettegrass or western panicgrass (Panicum lanuginosum var. fasciculatum= Dichanthelium acuminatum subsp. fasciculatum) in immediate foreground and Virginia wildrye (Elymus virginicus) immediately behind the rosette in the western Ozark Plateau on the first day of winter.

Both grass species are perennials, obviously (well some viewers might have taken them for biennials, but they are perennial species).

Ottawa County, Oklahoma. Late December; early winter (early growth phenologicl stage).

 

Cold, but comforted by leaves- Local population of Virginia wildrye (first slide) and two individual plants of Virginia wildrye (second and third slides) growing in a tallgrass-mixed hardwood savanna in the western Ozark (Spingfield) Plateau. Leaves around these wildrye plants were of chinquapin oak (Quercus muhlenbergia), northern red oak (Q. rubra= Q. borealis), black oak (Q. velutina), American elm (Ulmus americana), and common hackberry (Celtis occidentalis) in that order of leaf cover.

Ottawa County, Oklahoma. Late December (early winter); early growth, mature lower leaves.

 

Two of a similar--but different-kind- Spike of silky wildrye (front) and apike of Virginia wildrye growing in understorey of blackjack oak (Quercus marilandica)-post oak (Q. stellata)-tallgrass savanna in the western Springfield Plateau. Thee were also widely scattered trees of common persimmon (Diospyros virginiana), sassafras (Sassafras albidum), black walnut (Juglans nigra), and American elm near by, but mostly various forbs in background.

Ottawa County, Oklahoma. Mid-June (late spring); immediate post-anthesis stage of phenology.

Coming out (sort of ) party- Three progressively closer views of a spike of Virginia wildrye emerging (about as much as it is going to) from the boot at anthesis in a tallgrass-forb-mixed hardwood savanna in the western Ozark (Springfield) Plateau. The leaf immediately behind the spike is the flag leaf. The spike is emerging from below the flag leaf sheath which is the boot.

Incomplete emergence of the spike from the boot is a fairly reliable, field-indentification feature of Virginia wildrye.

Ottawa County, Oklahoma. Mid-June (late spring); immediate post-anthesis stage of phenology.

 

They do better when it warms up- Adult shoots of the same two plants introduced immediately above. Hairy rosettegrass, wooly rosettegrass, wooly panicgrass, western rosettegrass or western panicgrass in front and Virginia wildrye immediately behind. Both plants at peak standing crop in late spring on a "rock pile" of a microsite on the Springfield Plateau in northeastern Oklahoma.

Ottawa County, Oklahoma. Mid-June (and both plants at maturing grain stage of maturity).

Another one that stands the cold of winter- Shoots from last year (standing and lying dead staw) and current growing season (green and short) of Canada wildrye (Elymus canadensis) growing on a tallgrass-oak/hickory savanna on the Springfield Plateau in northeastern Oklahoma in early winter. This native perennial of the wheat or barley tribe (Tritaceae or Hordeae) is a cool-season C3 grass species.

Yes, those are patches of snow which, by the way, furnished God's own stage prop just for emphasis.

Ottawa County, Oklahoma. Early January; early growth stage of phenology.

Back to warm weather:

 

Disturbancee pioneer- Old-field threeawn, old-field wiregrass, prairie threeawn or, simply, annual threeawn- (Arist;ida oliganth;a) is a first or second seral stage pioneer on abandoned cropland, old homesites and schoolyards, and overgrazed pasture in the humid precipitation zone such as tallgrass prairie and the West Cross Timbers.annual threeawn. The dense stand of old-field threeawn seen here was growing on a sandy soil of an old cotton (Gossypium hirstum) field in the West Cross timbers. This soil of this poor old field had lost most of its A horizon which had been replaced with sand eroded in from adjacent land.

Tarleton State Universxsity Hunewell Ranch, Erath County, Texas. Mid-September, peak standing crop.

 

Little pioners- Several uprooted (pulld up by the author) plants of old-field threeawn at the stage of maximum plant deveelopment (amaturity). These were some of the plants of the local stand introduced in the two slides in the immediately preceding slide set. Forage value of this small annual species is poor except when the plants are young and before their inflorescences (with their troublesome awned lemmas) have emerged. Thee awns usually are not mechanically injurious, but occasionally they can be problematic to eyes of grazing animals.

Tarleton State Universxsity Hunewell Ranch, Erath County, Texas. Mid-September, peak standing crop.

 

Next year's ccrop- Inflorescences of old-field threeawn, a pioneering species on degraded raangee. This annual is one of the native "six-weeks grasses", but while ephemeral it takes longer to complete its life cycle than some other six-weeks species such as six-weeks threeawn (A. adscensionis) of the Chihuhuan Desert Region. Plants shown here had very succesfully completed their annual life cycle and produced grain for next year's crop. The inflorescence of Aristida species is complex enough that usually it has not been described though it can be regarded as a form of panicle.

Tarleton State Universxsity Hunewell Ranch, Erath County, Texas. Mid-September, mature seed-set stage of phenology

313. Grasslike plant of the oak-hickory savanna- Broadleaf or common cattail (Typha latifolia) growing in an old stone quarry excavated on a black oak-black hickory big bluestem savannah in the Ozark (Springfield) Plateau. Typhaceae, along with Juncaceae and Cyperaceae, is a family of monocotyledons tht rangemen describe or "lump" into the practical category or group they call grasslike plants.

Many, though certainly not all, grasslike plants are wetland plants. The cattail species clearly are wetland plants. T. latifolia was included at this "clearing in the woods" as a representative of both grasslike plants and wetland plants indigenous to the oak-hickory-tallgrass savanna. Though by no means a common plant, and restricted as it is to wet habitats, broadleaf cattail is a widely distrubuted native species and one that showed the diversity of plant life in the hardwoods-tallgrass savannah range type.

Newton County, Missouri. Late-July, fruit-ripening stage.

 

314. One gone, one stayed behind- Fruit cluster of common or broadleaf cattail. The brown-colored, oblong-cylindrical structure is the female inflorescence matured into a column of tiny fruits. The bare "stalk" or "stick" is the uppermost part of shoot that was the staminate part of inflorescence from which the male flowers were shed. The "cattail" is the male part (the assembly of staminate flowers) of the entire flower cluster (monecious arrangement) whereas the female portion (collection of pistillate flowers) is the "cat". In this mature sexual shoot the "tomcat" left while the "pussycat" remained behind.

Much of the plant of Typha species was used as food by the Indians and, later, adopted by frontiersmen. The most abundant food portion was the starchy rhizome, but young shoots, the flowers, fruit-bearing portion portion of shoot, and even the copius quantities of pollen were eaten one way or the other. Leaves and/or leaves and whole shoots of cattail were importnt house building materials including thatch for roofing. A good review of cattail uses (besides any feed value of the herbage, including rhizomes for fur-bearers) was that of Turner, 2009, ps. 284-288).

Newton County, Missouri. Late-July, fruit-ripening stage.

 

315. Rushed crowd- Stand of poverty, path or slender rush (Juncus tenuis) in understorey of open stand of black oak. There are relatively few species of grasslike plants in tallgrass savanna, but poverty rush is one that is locally abundant where overgrazing, trampling or heavy traffic (hence, "path"), or shallow soil (thus, "poverty") create a habitat fit mostly annual grasses and forbs( or, in some conspicuous locations, dense stands of slender rush).

Ottawa County, Oklahoma. Late May.

 

316. The impoverished are prolific- Just like humans (and other organisms) in poverty and hard times, poverty or path rush produces many progeny. In the first of these photographs three or four plants of Juncus tenuis were at vried phenological stages of sexual reproduction (fruit production). The second slide gave a general view of fruit-bearing shoots (inflorescences had become both flower- and fruit-bearing organs).

Ottawa County, Oklahoma. Late May.

 

317. Poverty prolificacy- Details of sexual shoots of poverty or path rush. Both flowers and fruit were in these sexual clusters of a species of grasslike plant that is tolerant of defoliation (general disturbance). Poverty or path rush is a textbook example of a perennial that is an r-selected species. It is self-evident to the observant rangeman that poverty rush is an ecological invader in the Clementsian concept as applied by Dyksterhuis (1949). This species produces fruit/seed like an annual weed.

The fruit of Juncus species is a loculicidal capsule, "a capsule which dehisces by means of openings into the locules, about midway between the partitions". Locule in this context is a cavity or space within the ovary. Capsule is a dry dehiscent fruit formed from a pistil having two or more fused carpels (Smith, 1977, ps. 66, 229, 291, 300).

Ottawa County, Oklahoma. Late May.

 

318. One flower flat-out- One flower flat- (or umbrella-) sedge (Cyperus retroflexus) on an ecotone between West Cross Timbers and Grand Prairie. One-flower flat- sedge was the major grasslike plant on this little bluestem-post oak-blackjack oak savanna in northcentral Texas. Numerous species of forbs accompanied this species, several of which were displayed and described below.

The second of these two slides presented the entire inflorescence of the plant shown in the first slide.

Erath County, Texas. Mid-May; immature fruit (achene) stage of phenology.

 

319. Another one flat-out- A second plant of one-flower flat (umbrella-) sedge growing on a little bluestem-post oak-blackjack oak savanna in northcentral Texas. This specimen was growing a short distance from another individual of the same species (the specimen showed in the preceding two-slide/caption set). The second of these photographs preented the entire inflorescence of the plant of one-flower flat-sedge seen in the first photograph. Units of individual spikelets (the smallest floral units) are "heads" or spikes (ie. spikes are compirsed of a number of individual spikelets). Five spikes are visible in the inflorescence seen in the second photograph.

Erath County, Texas. Mid-May; immature fruit (achene) stage of phenology.

 

320. Straight-fruit or rich woods caric seddge (Carex oligocarpa) at edge or ecotone of tallgrass prairie and oak-hickory forest in the western Ozark (Springfield) Plateau. Straight-fruit caric sedge is more a species of forests or, at least, savannahs than of grasslands. The specimen seen here and featured immediatelly below were growing on an outcropping of chert beneath an old-growth and senescing post oak (Quercus stellata).

Ottawa County, Oklahoma. Early June; ripening-fruit stage of phenology.

 

321. Straight shoots- Inflorescences of straight-fruit caric sedge with staminate and pistillate clusters present in two of three inflorescences. This is a cool-season species with more-or-less green shoots throughout winter. C. oligocarpa is in sub-genus Eu-Carex.

Ottawa County, Oklahoma. Early June; ripening-fruit stage of phenology.

 

Whe it's really full of fruit- A dense group of immature achenes of straight-fruit or richwoods caric-sedge on a specimen growing on a chert outcrip under crowns of an old-growth and senescing post oak.

Ottawa County, Oklahoma. Early June; ripening-fruit stage of phenology.

In winter- Three plants of the cespitose straight-fruit or richwoods caric-sedge (first slide) and one plant of straight-fruit caric-sedge at closer camera distance (second slide) in the "dead of winter". Leaves on all these neighboring plants, that were growing on a chert outcrop under the bare canopy of an old-growth and senescing post oak, had been frost-burned after being exposed to ambient temperatures as low as -8 degrees Fahrenheit early in the season before the plants had time to "harden-off" (chemically adjust to freezing temperatures).

There was no doubt that richwoods caric-sedge is a cool-season species. There are more species of Carex than of any other genus of vascular plant in the oak-hickory-tallgrass savanna of Missouri and Oklahoma (than in those entire states for that matter). Carex species occupy as many niches as there are Carex species (Gause's Competitive Principle: one niche; one species). It is only to be expected that there would be both cool-season and warm-season species of Carex.

Ottawa County, Oklahoma. Early January; early vegetative growth stage of phenology (and already a lot of leaves, "frost-bit" or not).

 

322. An interesting (and confusing) interloper in the Cross Timbers- Simpleleaf ticktrefoil (Desmodium psilophyllum) in West Cross Timbers. Geographic distribution of the Desmodium species in Texas is in a serious state of disarray with the number of collections apparently insufficient to firmly fix county-by-county occurence. The mostly single-leaf arrangement of this specimen and intense process of elimination tentatively identified this species of tick clover or ticktrefoil which often occurs as one to only a few plants in more mesic habitats of the Western Cross Timbers..

Along Richardson's Creek, Tarleton State University's H:unewell Ranch. Erath County, Texas. September; full-bloom to immature fruit stages.

 

323. Some details of a tick clover that did not play by the rules- A Desmodium species tentatively identified as simple-or sessile-leaf ticktrefoil (D. psilophyllum) based on the primarily single-leaf arrangement of this individual. The tick clovers or ticktrefoils (with these nouns variously sritten as one word, two words, two words hyphenated, etc.) are a commonly occurring but confusing lot throughout much of the oak-hickory forest region and the various savannas, woodlands, and grassland-forest mosaics thereof. This is one frequently found in the West Cross Timbers of northcentral Texas.

Along Richardson's Creek, Tarleton State University's H:unewell Ranch. Erath County, Texas. September; full-bloom to immature fruit stages.

 

324. Pink tips in the Cross Timbers- Details of the papilionaceous inflorescences of simple-or sessile-leaf ticktrefoil growing in understorey of Western Cross Timbers. Along Richardson's Creek, Tarleton State University's H:unewell Ranch. Erath County, Texas. September; full-bloom to immature fruit stages.

 

325. Another sorta sneaky--though strong--interloper- Sampson's snakeroot (Psoralea psoralioides) is a papilionaaceous legmue that is equally at home on tallgrass prairies, oak-hickory forests, and savannahs of these two formations. It blooms and produces its legumes early in the warm-growing season.Wide leaves of Sampson's snakeroot bespeak adaptation to low-light yet mesic habitats. Other Psoralea species, especially varieties of P. tenuiflora, are better adapted to drier and less light-restricted environments such as those of tallgrass, mixed and, even, shortgrass praires far to the west of the Ozark Plateau on which the attractive "customer" was photographed.

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

 

326. Giant forbs at the edge- Colony of cup plant (Silphium perfoliatum), a strongly rhizomatous composite with large, clasping leaves on tall shoots, some of which grew to heights greater than eight feet, growing on a north slope at the edge of a sugar maple (Acer saccharum)-pignut hickory (Carya cordiformis)- white ash (Fraxinus americana)-basswwood (Tilia americana) mesic forest where it contacted a bottomland tallgrass prairie dominated by eastern gamagrass (Tripsacum dactyloides). Cup plant is the largest and most mesophytic of three Silphium species that can be found on the oak-hickory forest--tallgrass prairie savanna in the Ozark Plateau amd Osage Plains-Cherokee Prairie (Central Lowlands) physiographic provinces.

The seemingly outlandishly large plants seen here were growing on an ecotonal, mesic microhabitat in the second consecutive summer of Extreme Drought (Palmer Severity Index).

Butterfly species on center capitulum (head inflorescence) in the second slide was a nice specimen of great spangled fritillary (Speyeria cybele).

Newton County, Missouri. Late July; peak standing crop, peak bloom phenological stage.

 

327. Cupped and visited- Cup plant derived its common name from a cuplike depression formed where its large leaves clasp (more or less envelop) a massive herbaceous, square or cubular stem. The big heads (capitula) of this native composite of the Heliantheae produce high yields of nectar and pollen that are attractants to many insects. The composite inflorescences of this cup plant colony attracted insects from three orders: Lepidoptera, Hymenoptera, and Diptera. Two colorful butterfly species were conspicuous in the second of these two photographs: eastern tiger swallowtail (Papilio glaucus) at lower left-center and spicebush swallowtail (P. troilus) at center. (A spicebush swallowtail was also visible at lower right capitulum in the first photograph.)

Newton County, Missouri. Late July; peak standing crop, peak bloom phenological stage.

 

328. Cupped capitulum- A capitulum (head inflorescence) of cup plant on a shoot from the colony shown in the two immediately preceding slide-caption sets. 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).

Silphium species are members of the large Heliantheae tribe. Newton County, Missouri. Late July; full bloom.

 

Brown eyes under the trees- One plant of brown-eyed susan (Rudbeckia triloba) growing on an oak-hickory-tallgrass savanna in the western Ozark (Springfield) Plateau in northeastern Oklahoma. The principal oak tree species were black oak and northern red oak. In this immediate area, however, the most common tree was black walnut (Juglans nigra) represented by a number of young sapling-sized trees.

There are six Rudbeckia species indigenous to this part of the Springfield Plateau (Steyermark, 1963, ps.1554-1560). The most common of these six species is black-eyed susan (R. hirta) a species that is usually an annual (though it can be biennial or, even, a short-lived lerennial) and generally consist of substantially smaller plants than those of R. triloba, a perennial. The plant shown here had been living for at least six years at time of photograph.

Rudbeckia is another genus of composite forbs that are members of the huge Heliantheae tribe.

Newton County, Missouri. Late July; full bloom.

 

Where it got its name- A shoot with tripartite leaves (first slide) and an example of one such three-lobed or three-parted leaf (second slide) of brown-eyed susan growing on an oak-hickory-tallgrass savanna in the Springfield Plateau of northeastern Oklahoma. This was one of several shoots (each of which bore two to six heads) belonging to the plant introduced in the preceding slide/caption unit.

Stoney Point Savanna, Ottawa County, Oklahoma. Late July; full-bloom stage.

 

Now for the sexual end- Capitulum (head) of brown-eyed susan growing on an oak-hickory-tallgrass savanna in the western Ozark (Springfield) Plateau of northeastern Oklahoma. This was one of several dozen heads produced on the plant introduced two slide/caption sets above.

Palatability of brown-eyed susan to ungulates was unknown (at least to this author), but it is generally found without evidence of having been grazed. This plant was on the outside of a fencerow of a pasture grazed by beef cattle. There were no plants of brown-eyed susan in this savanna pasture. Part of one brown-eyed susan plant growing right next to the fence had been grazed down heavily by young (roughly four months of age) suckling calves which also heavily grazed neighboring plants including leaves of sapling black walnut! Obviously, these calves were inexperienced and were clearly not representative of older cattle which had developed more selective tastes.

Stoney Point Savanna, Ottawa County, Oklahoma. Late July; full-bloom stage.

 

Many heads at the edge- An immense, multi-shooted plant of grey (gray)-headed coneflower (Ratibida pinnata) growing in an oak-hickory savanna just below a chert glade in the Springfield Plateau of southwest Missouri. Grey-headed (gray-head) coneflower appears to be as much at home in forest openings and along forest edges as on tallgrass prairie in this region as well as in the adjoining Cherokee Prairie and Osage Plains of the Central Lowlands physiographic province.

This member of the huge Heliantheae tribe grows to heights of seven (rarely, eight) feet and produces an almost unimaginable number of capitula (heads) for a perennial species. This author was not aware of any assessments of the palatability of gray-head coneflower, but it is clearly not found on overgrazed ranges and, instead, is much more abundant on virgin sod such as that of tallgrass prairie hay meadows.

Newton County, Missouri. Late June; clearly at full-bloom phenological stage.

 

They don't look grey (or gray) to me- A large number of capitula (heads) on shoots (first slide) and two heads (second slide) on the plant of grey-headed coneflower presented in the immediately preceding slide. Basis for the adjective "grey-headed", "gray-head", etc. was unknown and appeared meaningless as the disks are typically some shade or brown or rarely brownish or even purplish, but certainly not gray (or grey). Disks would be something of a grayish color upon death of the shooot as, of course, are almost all shoots, including flower disks, of the forb species.

Newton County, Missouri. Late June; clearly at full-bloom phenological stage.

 

The ox of my eye- Ox eye (oxeye), oxeye sunflower, or false sunflower (Heliopsis helianthoides) is one of the showier composites (and an obvious member of tribe, Heliantheae) growing in the savannahs of the tallgrass prairie-oak/hickory forest regions such as the aptly named Prairie Peninsula. This native perennial so closely remembles sunflowers, the "true" sunflowers (Helianthus spp.), that it commonly causes confusion to observers new to areas in which it grows. Both genus name and specific epithet of this species allude to this resemblance.

Ox eye (probably the more common name, and not to be confused with the Eurasian perennial Chrysanthemum leucanthemum called oxeye daisy) is a highly rhizomatous species that forms colonies which frequently cover considerable area. Such local colonies are more typically found on undisturbed areas such as ungrazed or only lightly grazed savanna or tallgrass prairie ranges. Outside fencerows are the more common habitats for this perennial composite. This phenomenon strongly suggest that ox eye is palatable to livestock. (In this rangeman's observation oxeye is extremely palatable to cattle.)

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

 

Ox eyes and leafy horns- Capitula (head inflorescences) of oxeye or false sunflower growing in an outside fencerow in the tallgrass-oak/hickory forest savanna in the western Springfield Plateau of northeast Oklahoma. A distinctive feature of this apparently palatable, native perennial is a pair of opposite leaves just below the capitulum that extend horizontally from the shoot. This morphological feature was shown to good advantage in the first of these two slides and in the first slide in the immediately succeeding slide-caption set.

Ottawa County, Oklahoma. Late July.

 

"The better to see you, my dear"- Details of the head or capitulum of ox eye or false sunflower growing in an outside fencerow in the western Springfield Plateau of extreme northeastern Oklahoma. Ox eye is a member of the huge tribe Heliantheae, the heads of which species very closely resemble each other.

The head in the first of these two slides had been fed on by the small katydid (species unknown to the photographer) still seen on a ray (ligulate) flower. Two more examples of a different form of insect herbivory were presented in the next two slide-caption set.

Ottawa County, Oklahoma. Late July.

 

For honey and sweat- Capitula (heads) of ox-eye being visited by hymenopterans. The head in the upper slide was hosting a brown-belted bumblebee (Bombus griseocollis) while the head in the lower slide was serving nectar to a metallic sweatbee (Augochlora pura). This species of sweatbee is a solitary insect whereas the brown-belted bumblebee is a eusocial (social) species. Eusociality is interpreted as the most highly organized society among animals outside of man. Eusociality is particularily common within the Hymenoptera (order of insects having membraneous wings including bees, ants, wasps).

Ottawa County, Oklahoma. Late July.

 

329. A widespread composite of the tallgrass-hardwood savanna- The goldenrods (Solidago species) comprise some of the more common (not to mention, showest) of the forbs to make their home on the broad ecotone between tallgrass prairie and oak-hickory forest. One of the more frequently found goldenrods (especially on mesic habitats) is S. gigantea, known by a variety of common names including giant, late red-stem, glaborous goldenrod. The specific epithet, gigantea, is appropriate as this species can attain large size not infrequently reaching heights of eight feet or more. Like many of the other Solidago species S. gigantea produces sister plants from creeping root stocks (rhizomes) so that it commonly forms extensive colonies which also contributes to the gigantic proportions attained by this forb.

Giant goldenrod can abe found on more moist and otherwise favorable sites on both grassland,more open woodlands, and savannas of these. The examples shown here were growing in a narrow transition zone between the Grand Prairie and West Cross Timbers on a low-lying local habitat.

Giant goldenrod is one of the more widely distributed of all the Solidago species perhaps explaining the array of common names.Range of S. gigantea extends from extreme peninsular Florida to Nova Scotia and British Columbia and the Northwest Territories south to the southernmost tip of Texas.

Erath County, Texas. Early September; peak-bloom stage of phenology.

 

330. Higher parts of giant goldenrod- Upper shoot (first slide) and details of one branch of the panicle (second slide) of giant, late, glaborous goldenrod. These were some of the shoots of the colony introduced in the immediately preceding photographs. Giant goldenrod is one of the showier forbs commonly found on prairie, marsh, and open oak-hickory forest. Given the huge geographic (speceis) range with tremendous adaptability, larage mature size, and showy inflorescence giant or late goldenrod is an ideal species for native plant enthusiasts, especially for those landscaping with prairie plants.

Forage value and response to grazing intensity (stocking rate) of this species was unknown to this author, but he has found it growing beside switchgrass and big bluestem.on moist environments including some in the Nebraska Sandhills.

Erath County, Texas. Early September; peak-bloom stage of phenology.

 

331. Prairie forb in a savanna- Three spindly plants (or possibly only two plants with the two shoots on the right being of the same plant?) of Missouri goldenrod (Solidago missouriensis) growing in a tallgrass/black oak-mockernut hickory (Carya tomentosa) savanna in the far western Ozark (Sprinfield) Plateau. Other tree species were post oak and blackjack oak (eg. trunk in center backghround). Major asociated herbaceous species were mostly grasses including purpletop (Tridens flavus), poverty oatgrass (Danthonia spicata), Japanese chess (Bromus japonicus) in the cool-season, and big bluestem as a potential decreaser, but here about killed out by repeated shredding.

Missouri goldenrod is one of the shorter-growing Solidago species. Missouri goldenrod produces both rhizomes and short stolons yet with "solitary" or widely speced shoots (Fernald, 1950, p.1399). The Solidago inflorescence has traditionally been regarded as a panicle, but some author regarded the branches of the inflorescence as racemes which in this species are usually one-sided recurving racemes (Fernald, 1950, p. 1399) as seen here.

Ottawa County, Oklahoma. Early August; early bloom phenological stage.

 

332. Typical Show-Me stem and leaves- Two views of a shoot of Missouri goldenrod (one of those presented in the immediately preceding slide). The stem of this species is frequently reddish to purplish in color and leaves are acute, characteristically serrated (prominently so here) with a prominent midrib (= midvein) and two parallel lateral (often somewhat obscure) veins.

Ottawa County, Oklahoma. Early August.

 

333. Show-Me flowers- Flowers of Missouri goldenrod on one of the shoots shown in the first slide that introduced Missouri goldenrod. Flower clusters of Missouri goldenrod (one of the shorter Solidago species) are typically smaller and more sparingly branched than inflorescences of many of the other goldenrods.

Ottawa County, Oklahoma. Early August; early bloom phenological stage.

 

Camphored away- Upper portion (the crown) of a plant of camphorweed or camphor daisy (Heterotheca subaxillaris) growing on sandy soil of a disturbed microsite in the West Cross Timbers of northcentral Texas. This is an annual or, less commonly, a biennial member of the Astereae (aster tribe). Camphorweed is, in so many words, a weed and in parlance of Range Ecology, an eccological invader. Camphor daisy thrives on harsh microsites where it receives less competition from perennials, especially those of higher (more advanced) successional status. Diggs et al. (1999, p. 372) remarked that the aormatic camphor-like odor of camphorweed was responsible for it being avoided by livestock (and probably most wildlife species as well).

Erath County, Texas. Early September; peak bloom stage of phenology.

 

Camphor close-up- Upper shoot with flower cluster (first slide) and two capitula or heads (second slide) of camphorweed. These slides were taken on the same plant crown that was seen immediately above. Camphor daisy is probably one of the most unpalatable forbs of the Cross Timbers. Natural selection has favored an early seral-stage, annual composite that contains higher concentrations of a chemical deterrant which aids in its individual survival (and high probability that it will successfully reproduce) on hostile habitats.

Erath County, Texas. Early September; peak bloom stage of phenology.

 

Did he eat or not?- A male differential grasshopper (Melanoplus differentialis) resting on a branch of camphorweed or camphor daisy in the West Cross Timbers of northcentral Texas. This 'hopper was on the same plant shown in the two immediately preceding slide/caption sets. It was explained there that camphorweed is quite unpalatable to livestock, but the palatability to insects of this annual member of the aster tribe was not known (at least to this author).

There was no evidence of insect feeding (eg. grasshopper bite marks) on this plant so it seemed likely that this ole boy simply found this a convenient place to light, or perhaps there was an instinctive behavior that predators would be less abundant on the foul-smelling camphorweed. Of course, the odor might not be offensive to insects; perhaps the odor was even attractive if through evoutionary instinct or learned experience predators suspected their prey in the "camphor patch". This grasshopper seemed safe for the time being. (By the way, some insects adjust very quickly to human presence that they do not perceive as threatening. This individual allowed the photographer to get extremely close to him. This photograph was taken with a macro- (micro-) lens just a few inches away from the nonchalant subject. Cooperative cuss, hugh?)

Erath County, Texas. Early September.

 

331. Little patch- A small stand of prickly lettuce (Lactuca serriola= L. scariola) growing near a north-slope mesic forest that had been clearcut (highgraded) in western Ozark Plateau. This was a sugar maple-northern red oak-pignut or bitternut hickory-white ash-basswood climax community. Prickly lettuce is widely abundant in this general locality being found in overgrazed pastures, road ditches, cutover forest, parking lots, school yards, and you get the idea.

Prickly lettruce is in the Liguliflorae, the subfamily of Compositae (tribe Cichorieae) whose members produce a milky (latex) sap. This sap does not deter consumption of the plant-- either by beast or man. This photographer enjoys a good bait of tender "spring greens" of which prickly lettuce is a "prime target species". Sow thistle (Sonchos oleraceus), another member of the Cichorieae, also eats well. Throw in lambsquarters (Chenopodum album), which is probably best pot herb overall, and poke salad (Phytolaca amricana), which should always be included, and any self-respecting hillbilly (eg. your author) is ready to devour a pot of "good eats". For pot herbs these forb species must be used earlier in the spring when shoots are relatively small, say less than a foot and a half tall (and then just take the first (top or upper) half foot or a little more).Not even deer would eat prickly lettuce at adult stages of maturity such as the plants shown in this slide. Cooking note: always pour out the first pot of water that such "weed greens" have been boiled in and then add fresh water and finish boiling to tenderness.

Some of the Lactuca species are cool-season biennials with overwintering rosettes, but L. serriola is a cool-season annual (Diggs et al., 1999). L. serriola does have a basal leaf rosette, but this develops and persist through only one winter versus two winters in biennial Lactuca species.

Most members of the Liguliflorae growing in forests of the Ozark Plateau are Eurasian species that have naturalized. This includes Lactuca and Sonchos species such as those featured in this section. oak-hickory. Species of these genera are certainly weeds--in both agronomic (cultural) and ecological (successional) meanings--that can be locally abundant, but they require fresh disturbances for establishment and, other than locally, do not form such extensive populations as to be serious pests. On clearcuts, such as the one presented here, Lactuca and Sonchus (treated shortly below) species are beneficial from standpoint of soil protection against erosion, recycling soil nutrients, providing green feed for grazing animals during the "off-season" (they are cool-season species), and even serving as less-than-choice bee plants.

Ottawa County, Oklahoma. Early July.

 

332. Prickled shoots- Shoots of two plants of prickly lettuce growing near a clearcut north-slope mesic forest in western Ozark Plateau. Prickly lettuce is a rank-growing biennial. These shoots had the diameter of posthole digger handles and stood over ten foot tall. These plants were in peak-bloom stage with lower leaves already senescing and dying as this sporophytic generation was completing its phase of this species' life cycle. At this stage shoot are usually as hollow as a length of water pipe.

Also at this stage of plant maturity, shoots of prickly lettuce have lost their palatability. For example, in this vicinity white-tailed deer feed on prickly lettuce shoots with the more apical portions being cropped. At the stage of "rank-maturity" seen here prickly lettuce is no longer taken by deers or other herbivores (ie. it has grown past its stages of vulnerability, at least to vertebrate grazers.

Ottawa County, Oklahoma. Early July.

 

333. Prickly particulars- Details of stem and, especially, leaves of prickly lettuce. Organs seen here were on plants in the stand shown above that introduced this naturalized composite. These are full-grown and fully mature leaves. Leaves of Lactuca species such as prickly lettuce are extremely vriable in shape depending of taxonomic form (Steyermark, 1963, p. 1638).

Ottawa County, Oklahoma. Early July.

 

334. How it grew back- New shoots growing from leaf axils of prickly lettuce in response to removal of apical meristem by grazing white-tailed deer. These axillary branches will go on to become sexual shoots (ie. produce a multi-headed flower cluster). This plant was in the same location as that of other slides in this section (near a clearcut mesic mixed forest in the Ozark Plateau). Deer frequently consume upper parts of prickly lettuce shoots. The author has yet to see one such grazed prickly lettuce that did not regrow and produce a plentiful crop of achenes, the dry fruit of composites. Speaking of which kindly advance the carousel.

Ottawa County, Oklahoma. Early July.

 

335. Sexy lettuce- Sexual shoot of annual prickly lettuce with unopened heads, blooming heads, and heads of achenes all on the same cluster of sexual units. Upper portion of a floral assembly (= all heads of total flowering shoot) on one plant of prickly lettuce.

Ottawa County, Oklahoma. Early July.

 

336. Impressive in numbers- Three tiny heads of prickly lettuce on one limb of the floral assembly shown immediately above. Any one of these heads or capitula may not strike the typical, citfied field-walker as being all that impressive, but such neophytes should observe just how many of these little heads are on one plant. To complete this mental calculation one should bear in mind just how many seeds (inside of achenes) are produced by each smallish capitula. For that part of the exercise please press the remote Forward button...

Ottawa County, Oklahoma. Early July.

 

337. Loaded and ready to fire- Mature achenes (the fruit of composites) on the former heads or capitulae of prickly lettuce growing near a clearcut forest in the Ozark Plateau. There is one seed per achene (think sunflower "seeds"), and at least ten achenes per head. (Well, if you don't beleive me count them yourself!) This author has counted from 80 to over 200 heads per prickly lettuce plant. (If you think this is a Ozark talltale push the remote Reverse button and count the number of achene clusters, ripened heads, on just the upper portion of one flowering shoot.Damn fureners, anyway!)

Ottawa County, Oklahoma. Early July.

 

Now for another or follow-up lesson, including more on the "business end" of prickly lettuce:

338. Alien among the natives- Prickly lettuce (Lactuca serriola= L. scariola) in an oak-hickory-tallgrass savanna in western margin of Ozark Plateau. This annual member of the Liguliflorae, the Compositae subfamily whose members produce a milky (latex) sap, like the red and white man now in North America, originally hailed from Eurasia. Both species came to the New World from the Old, liked their newly adopted land, and naturalized enough to stay.

This is one of three Lactuca species known generically as "wild lettuce" that became well-established in the Ozark Highlands. All these species of "wild lettuce" do best in newly and frequently disturbed ground where they commonly thrive. In so doing they provide both forage from their foliage and small concentrates from their tiny achenes. The herbage is valuable for livestock and wildlife ranging from rabbits to white-tailed deer whereas the achenes are used by birds. Mere restriction of light by a woody canopy is frequently enough of a stress for native plants that "wild lettuce" can outcompete--at least co-survive with--other herbaceous species. This is frequently most obvious on steep, rocky land.

Some of the Lactuca species are cool-season biennials with overwintering rosettes, but L. serriola is a cool-season annual (Diggs et al., 1999). L. serriola does have a basal leaf rosette, but this develops and persist through only one winter versus two winters in biennial Lactuca species.

The first of these two slides showed a robust (eight foot tall) plant at edge of a forest stand dominated by black oak. The second slide was of upper portion of inflorescence with phenological stages varying from newly opened flowers to maturing achenes.

"Wild lettuces", like prickly lettuce, are now as much a part of plant communities in open forests, savannas, and forest glades in regions like the Ozark Mountains that these naturalized exotics must be included as part of the range vegetation. The fact that they provide useful feed to range animals just "levens" the range loaf.

Ottawa County, Oklahoma. First slide, mid-July and early fruit stage; second slide, early July and flowering to fruit-set stages.

 

339. Flowers to fruits at a distance- Blooming heads and unopened floral buds (first slide) and clusters of achenes (second slide) of prickly lettuce. These were on plants on the same oak-hickory-tallgrass savanna as shown in the immediately preceding photograph. One or two branches of one entire inflorescence (flower cluster) on a prickly lettuce were displayed.

Ottawa County, Oklahoma. First slide, mid-July and flowring stage; second slide, early July fruit-ripe stage. Note that a week's difference was irrelevant as some branches of one inflorescence was more advanced phenologically than adjacent or closely neighboring branches, all on the same plant.

 

340. Flowers to fruits in detail- Flowering heads and just emerging achenes (first slide) and the intricate pattern of ripe achenes just as they were being shed (second slide) in prickly lettuce.

Ottawa County, Oklahoma. Early July.

 

341. Replacement parts- Plant of blue lettuce (Lactuca canadensis) that had its primary shoot (= its main shoot that originated from the embryo) removed by white-tailed deer (Odocoileus virginianus) and then subsequently exhibited regrowth as two younger shoots arising from intercalary meristem immediately below point of original (primary) shoot removal and a third shoot from intercalary meristem at the next lower shoot node below point of defoliation. These younger shoots that branch off from the primary shoot are designated as secondary shoots.

The phenomenon at work in regrowth of this blue lettuce plant was apical dominance. Apical dominance is the process whereby apical meristem (meristematic tissue produced by/in the shoot apex or shoot tip, shoot terminus) produces growth-regulating hormones, especially auxins such as indole-acidic acid, that regulate or dominate increase in size of plant cells thereby controlling growth of the plant. When the apical meristem is removed as, in this case by white-tailed deer, meristematic tissue in intercallary portions of the shoot below the former apical meristem are now released from previous dominance by apical hormones (auxins) so that they can develop in the fashion of the former primary shoot.

In this example, the exact point at which the primary shoot was grazed off by deer was conspicuously indicated by an assassin wheel bug (Arilus cristatus) who was using the shoot stump as a perch from which to prey on ("assissinate") careless, unsuspecting insects (or to inflict a painful bite with its piercing mouthpart to clueless photographers who were not as alert as the one who captured this instructive image). Yes, several lessons in this slide.

Ottawa County, Oklahoma. Early July; regrowth at late pre-bloom stage.

Lactuca as well as Sonchos (see immediately below) species are in Liguliflorae, the lataex-secreting subfamily of Compositae. The best known species in this subfamily is common dandelion (Taraxacum officanale). It was featured in the Alpine chapter of Range Types. Another similar species is goat's beard (Tragopogon dubius) that was treated in the Palouse Prairie chapter. Compare these species to note the extremely similar disks and plummed achenes of these composites, all of which are alien to North America and yet all very much naturalized in their new "Home on the Range".

 

342. Another naturalized commoner- Common sow thistle (Sonchos oleraceus) on an oak-hickory-tallgrass savanna in Ozark Plateau. These plants (two plants in first slide; upper shoot with inflorescence of right plant in sescond slide) were on the edge of the same black oak-dominated forest in photographs of prickly lettuce immediately above. It is common for several species of Lactuca and Sonchos to grow side-by-side (or nearly so) in the Ozark Highlands, especially on harsher habitats such as those of shallow, rocky soils.

Like prickly lettuce (and other "wild lettuces"), sow thistles (there are at least two species in the Ozarks Region) are commonly eaten by white-tailed deer as well as cattle, sheep, goats, and eave horses on occasion.

Ottawa County, Oklahoma. Mid-July.

 

343. Pale, but pretty- The pale yellow heads of common sow thistle. These flowers were on plants growing on a tallgrass-oak-hickory savannah in the western margin of the Ozark Plateau. Ottawa County, Oklahoma. Mid-July.

 

Bearing flowers and fruit in its adopted home- Shoots of one plant of tall sow thistle (Sonchus asper) growing on a disturbed area (frequently shredded railway right of way) in the West Cross Timbers of northcentral Texas.

S. asper and S. oleraceus are very similar species and grow together in the same areas. Diggs et al. (1999, p. 416) in their flora of northcentral Texas distinguished between these two species using the following: "leaves stiff and painfully prickly..." versus "leaves softer, not painfully prickly..." for S. asper versus "S. oleraceus", respectively. That is a cowboy story: "No shit, this really happened". In his Missouri flora, Steyermark (1963, p. 1635) distinguished between these two Sonchus species using rounded auricle leaf lobes and smooth-surfaced achenes versus pointed auricle leaf lobes and rough-surfaced achenes for S. asper and S. oleraceus, respectively. Again in cowboy parlance, these two sow thistle species are "purty damn close" in their morphological appearances. (Maybe a cladist could look at some chemical compound and put the two species in two separate plant families.)

Erath County, Texas. Early May.

 

Flowers and fruits following some grazing- Capitula (heads) at both flowering and fruit (achene)-bearing stages in tall sow thistle on a railroad traack in the West Cross Timbers of northcentral Texas. There had been some grazing of this plant by white-tailed deer. These organs were on the same plant that was shown in the immediately preceding photograph/caption set.

Any of the standard flora treat this species descriptively and taxonomically (two such flora were quoted in the preceding caption), but for coverage as a weed (and in comparison to other weedy Sonchus species) the Agricultural Extension publication, Weeds of the West (Whitson et al., 1992, ps. 180-181) was recommended.

The sow thistles (regardless of Sonchus species) are quite palatable to certain animal species, such as white-tailed deer (see immediately below).

Erath County, Texas. Early May.

 

344. Several smaller where there had been one larger- A plant of sow thistle that had been grazed by white-tailed deer resulting in removal of the primary shoot and its apical meristem. The resulting pattern of plant growth following release from apical dominance was proliferation of secondary shoots each of which had its single apical meristem (and subtending--situated below--intercalary meristems). Further deer grazing that might eliminate these terminal or apical meristems would result in prolific production of new shoots (from intercalary meristem) from (off of) the defoliated secondary shoots. Such shoots arising from secondary shoots would then be designated as tertiary shoots. With further defoliation by deer (eg. production of quarternary shoots, and so on) the growth form (morphology) of the plant would become consecutively bushier and relatively shorter. The highly grazed plant could eventually have a highly "hedged" appearance.

The phenomenon of apical dominance and plant regrowth (production of secondary shoots) was described above under treatment of Lactuca species.

Details of growth of this "de-topped" sow thistle were presented in the next two-slide/caption set. Before scrolling sown, however, students should find the "stump" of this shoot (point at which deer grazed off the primary shoot).

Ottawa County, Oklahoma. Early July; regrowth at late pre-bloom stage.

 

345. Several for the cost of one- A plant of sow thistle that had its apical (terminal) meristem removed (apical dominance eliminated or overridden) by white-tailed deer resulting in production of numerous secondary shoots. The "stump" of the primary shoot (point at which deer grazed off the main shoot that originated from the embryo) can be seen in both of these slides as well as in the immediately preceding slide. (It is harder to locate the "stump" in the first of the two current slides, but rangemen need continuing practice in "reading sign".)

An age old question in rgards to secondary, tertiary, etc. shoot production following defoliation (eliminating, or more correctly, relocating apical dominance) has been whether or not such response is increased plant production. Even if there was no increase in plant yield (not greater biomass production) the change in plant growth pattern and morphology did have regrowth that was higher in nutrients than in a maturing primary shoot. Thus, grazing (defoliation) by white-tailed deer had resulted in greater net yield of forage nutrients (energy, crude protein, etc.) for these deer. Eventually there were more flower clusters on this plant (most secondary shoots bore inflorescences) rather than a single and much larger inflorescence. It was not known if there was any difference in fruit yield (ie. any difference in numbers of achenes produced by/on this particular plant of sow thistle).

This was clearly an example that showed regrowth as a survival adaptation (evolution) to defoliation. This genotype of an annual composite did reproduce its kind, and it did so with formation of numerous new genotypes (any one of which might, in theory, increase the fitness of this species for defoliation).

Ottawa County, Oklahoma. Early July; regrowth at late pre-bloom stage.

Culinary note: Properly cooked (boiled with two rinses of water and maybe fried thereafter) the leaves and even young shoots of immature plants and shoot tips of mature plants of Lactuca and Sonchos make good greens for hillbillies with connoisseural tastes (your author for example). Mix in some lambs quarters (Chenopodium album) and poke (Phytolacca americana) and the eats are "larruping good".

Interesting diversity on a West Cross Timbers-Grand Prairie ecotone: the following forb species were growing and blooming concurrently and intermixed among themselves on a Cross Timbers-tallgrass prairie savanna in northcentral Texas in late spring. In addition to grasses (Gramineae) and grasslike plants of Cyperaceae or sedge family there were forbs in the following families: Compositae, Leguminosae, Labiatae, Onagraceae, Asclepediaceae, Polygonaceae, Rubiaceae, and Nyctaginaceae.

First (and following the preceding plants) for the other (the big) subfamilly of Compositae, Tubluiflorae .

 

346. A composite that likes the edge- A few spindly plants of prairie fleabane (Erigeron strigosus) growing on a relict savanna in the West Cross Timbers of northern Texas. There are many species of Erigeron native to North America. Diggs et al. (1999) named and described six Erigeron species for northcentral Texas. As with fleabane species elsewhere those of northcentral Texas are not easy to distinguish as there are "morphological intermediates" at "species boundaries" (Diggs et al, 1999, p. 350). The fact that three of these Erigeron species are annuals simplified the matter. Even then it can be complicated as there appears to be "introgressive hybridization" between E. strigosus and the very similar E. tenuis (Diggs et al., 1999, p. 352).

In the area where these can-be-confusing characters called home they are equally at home on tallgrass prairie or Cross Timbers. The edge of a local ecotone (narrow transition zone) between these two range types resulted in a savanna that proved ideal for these plants. They liked living on the vegetational edge.

Erath County, Texas. Mid-May; peak-bloom phenological stage.

 

347. Tops of one on the edge- Pair of topdown nested views of prairie fleabane on the ecotonal edge between tallgrass prairie and West Cross Timbers in northcentral Texas. These heads (capitulae) were on the plant seen in the immediately preceding photograph.

When perpetuation of their species depends on sexual reproduction, annual plants like these shown here will produce the gametophytic generation if given half a chance. This wild bouquet attested to that fact of botanical life.

Erath County, Texas. Mid-May; peak-bloom phenological stage.

 

348. Plainsman in the Cross Timbers- Old plainsmen (Hymenopappus scabiosaeus var. corymbosus) on a local ecotone between tallgrass prairie of Grand Prairie and the savanna of West Cross Timbers (ie. a savanna within a savanna). This species with often robust individuals is a biennial of the Heliantheae, a composite tribe primarily of annual and perennial species.

Old plainsman is one of the earliest native composite to bloom in the spring.It is a showy plant in spite of a rather lackluster inflorescence.

Erath County, Texas.Mid-May; peak standing crop and peak-bloom stage of phenology.

 

349. Not false to itself- Entire shoots (first slide) and upper shoot (second slide) of plant of false boneset (Kuhnia eupatorioides var. corymbulosa) growing at edge of a sand post oak-fiddleleaf gerrn-brier dwarf forest in the West Cross Timbers of northcentral Texas. The specific epithet, eupatorioides, was in reference to the genus Eupatorium and/or the tribe Eupatorieae known by such common names as boneset, thoroughwort, Joe Pye weed, and several others. K. eupatorioides is in the Eupatorieae and, in addition to tribal features, presumedly or supposedly bore enough resemblance to Eupatorium species to get the specific epithet, eupatorioides.

Some taxonomic treatments included Kuhnia in genus Brickellia as in Brickellia eupatoroides (Diggs et al., 1999, ps. 328-329), but Kuhnia eupatorioides is the traditional binominal (Coulter, 1891-1894, p. 180; Fernald, 1950, ps. 1370-1371; Steyermark, 1963, ps. 1468-1469; Great Plains Flora Association, 1986, ps. 966-967).

Erath County, Texas. Early November; peak flowering phenological stage.

 

350. Not falsely blooming- Upper shoots and flower clusters (first slide) and details of individual heads (second slide) of false boneset growing at the outer edge of a sand post or margarettia oak-fiddleleaf green-brier dwarf forest in the West Cross Timbers.

This species is just another one among "jillions" of composites growing on ranges in northcentral Texas. Falseboneset has a biological (species) range that extends from Florida to Texas in the south northward to New Jersey and Pennsylvania and then westward to Illinois and Missouri. There was sufficient morphological variability in this species to recognize three varieties in the greater Great Plains Region (Great Plains Flora Association, 1986, ps. 966-967).

Relative palatability of false boneset is apparently not known, but it is doubtful that grazing animals have much preference for it.

Erath County, Texas. Early November; peak flowering phenological stage.

 

351. Mint for the breath of the Cross Timbers- Upper shoot of spotted beebalm or yellow horsemint (Monarda punctata). There are at least four Monarda species in the general area of the West Cross Timbers of northcentral Texas. These are all similar, especially under field conditions. As this was not confusion enough there are three varieties of M. punctata (Diggs et al, 1999, ps. 764-768). The student of Cross Timbers vegetation need not despair, however, as the other three Monarda species are annuals whereas M. punctata can grow as a short-lived, colony forming perennial.

Erath County, Texas.Mid-May; peak standing crop and peak-bloom stage of phenology.

 

352. Let's make it double mint- Basil beebalm or horsemint (Monarda clinopodroides) on fine sandy loam bottomlaand in the West Cross Timbers. This another one of the four (or more) Monarda species in the West Cross Timbers. Basil beebalm is one of three annual Monarda species in this general area. M. clinopodroides is a widely distributed forb species (Diggs et al., 1999, p. 765) the petals of which vary from white to pink to purple. The inflorescence of Monarda species was interpreted by Diggs et al.(1999, p.764) as an interrupted spike.

Greater floodplain of Bosque River, Erath County, Texas. Mid-May; peak-bloom stage.

 

353. Tall forb in Cross Timbers- Tall or smooth four-o'clock (Mirabilis glabra) at peak standing crop and peak bloom on a transition between Western Cross Timbers and Grand Prairie. This specimen was over seven feet tall. It was growing in the sward dominated by little bluestem with red lovegrass (Eragrostis secundiflora subsp. oxylepis= E. oxlyepis), fall witchgrass (Leptoloma cognatum), small-head or short-leaf rush (Juncus brachyphyllus), and one-flower flat- (umbrella-) sedge (Cyperus retroflexus) as its principal neighbors.

Erath County, Texas. Mid-May; full-bloom phenological stage.

 

354. Taking time to admire- Upper portion of inflorescence (first photograph) and details of two flowers (second photograph) of tall or smooth four-o'clock. These organs were on the plant seen immediately above.Mirabilis species like this M. glabra are members of the Nyctaginaceae, four-o'clock family. It was explained in introduction to this short section that the botanical diversity on this Cross Timbers-tallgrass savanna included forb members of Compositae, Leguminosae, Labiatae, Onagraceae, Asclepediaceae, Rubiaceae, Polygonaceae, and Nyctaginaceae.

Erath County, Texas. Mid-May; full-bloom phenological stage.

 

355. Diamonds on the savanna- Diamond-point or four-pointed evening-primrose (Oenothera rhombipetala) on an ecotone of West Crosss Timbers and Grand Prairie (ie. an ecotone within a broader ecotone). The plants seen here of this biennial species were growing on a locally disturbed (parking and maneuvering of road-construction equipment). This was an example of patch dynamics on a little bluestem-post olak-blackjack oak savannah. Apathetic abuse of the land resulted in loss (at least temporarily) of litttle bluestem and the prospering of these native biennial forbs.

Erath County, Texas. Mid-May; (and could there be any doubt as to plants being mostly at full-bloom phenological stage).

 

356. Four points to ponder- A single plant of the biennial four-pointed or diamond-pointed evening-primrose (first slide) featuring its large inflorescence that was in early stages of flowering. These two photographs highlighted the indeterminate flowering pattern (progression from lower or bottom to upper or top and from inner to outer portions) of the Oenothera species.

Erath County, Texas. Mid-May; early bloom stage of phenology.

 

357. Pointed ponderings- Finer details of flowers of diamond-point or four-point evening-primrose. These flowers were on one of the plants introduced above that were growing on a tallgrass-post oak-blackjack oak savanna in a transition zone between West Crosss Timbers and Grand Prairie tallgrass prairie.

This whorl of flowers was near bottom of an inflorescence of the indeternminate flowering pattern (see immediately preceding caption).

tThe common name of "evening- primrose" was derived from the phenomenon of these species opening their petals--often quite quickly--near time of sunset or evening. Basis of the adjective of "diamond-point" or "four-point" was also apparent in these slides

Erath County, Texas. Mid-May.

 

358. Now for a little one- Mat bluets (Houstonia humifusa) on a transition zone between West Cross Timbers and tallgrass prairie of Grand Prairie. A number of these little folks were growing with the forb species presented above. This species one had the honor of being the one that prduced the smallest plants (and fruits; see below).

Houstonia species are in Rubiaceae, the madder family.

Erath County, Early May; ripening-fruit phenological stage.

 

359. Capsules in the Cross Timbers- Capsules on floral branches of mat bluets. These structures were on one of the two plants shown above.

Erath County, Early May; ripening-fruit phenological stage.

 

360. Bound for Texas- Part of shoot (first slide) and close-up of leaves and three flowers (second slide) of Texas bindweed, gray morning-glory, or Texas morning-glory (Convolvulus equitans) growing in a disturbed area (railway right of way) in the West Cross Timbers of northcentral Texas. This native perennial of the morning-glory family (Convolvulaceae) is widely distributed throughout Texas with a species (biological) range that, though sporadic, extends from Oklahoma an Texas westward to California.

Texas morning-glory is often quite palatable to and heavily fed on by grazing animals, even grass-grazers like cattle. The fact that the plant shown here and the plant specimen presented immediately below both grew on fenced railroad property and away from any deer feeding area might explain why these individuals were so vigerous and without any evidence of defoliation.

Erath County, Texas. Mid-August; full-bloom phenological stage.

 

361. Texas enough for another view- Another plant of Texas morning-glory or Texas bindweed featuring a section of shoot with leaves and flowers (first slide) and details of two flowers (second slide) growing on a railroad right of way in the West Cross Timbers of northcentral Texas. This plant (parts of) and the plant (parts of) shown in the preceding slides/caption unit illustrated the morphological variation present among individual plants of this species, especially within flowers and leaves. This pronounced, phenotypic variation can occur among genetically different individuals (genotypes) within close distance of each other (about 25 yards in instance of these two plants).

Like most Convolvulus species (in fact, many perennial members of the Convolvulaceae) Texas bindweed grows a relatively large, food-storing taproot Diggs et al., 1999, p. 552).

Erath County, Texas. Mid-August; full-bloom phenological stage.

 

362. Wild strawberry (Fragaria virginiana)- This was one of the few plant species native to North America that was domesticated as a horticultural crop. The specimens shown in this and the next slide were examples of the wild type (ie. native or nondomesticated genotypes) and not the "tame" varieties some of which have been bred to bear the accessary fruit in grotesque sizes and shapes (only to be hollow in the middle). It appears that the domestic strawberry is a hybrid between F. virginiana and F. chiloensis (Forest Service, 1941, p. W77; Bailey, 1949, ps. 526-527).

The individual seen here was growing in the oak-hickory-tallgrass savanna It was at the edge of a stand of black hickory (Carya texana) in an overgrazed tallgrass prairie dominated by broomsedge (Andropogon virginicus). The stolon or runner of this asexually reproductive member of the rose family (Rosoideae subfamily of the Rosaceae) was obvious. Ottawa County, Oklahoma. April.

 

363. Flower and fruit (mature and immature) of wild strawberry- A comparison of this flower with those of blackberry (next plant in this series) showed characteristics of the rose subfamily of the rose family and the similarity among species within that subfamily. The fruit type of strawberry is accessary fruit which is one type of "false fruit" (those produced by the coalescing, fusion, or uniting of the pistil and other parts of the flower). The accessory fruit involves retention of an enlarged receptacle of the (ie. of one) flower which is covered with achenes, these latter are the true or actual fruits. Achenes are dry, one-seeded, indehiscent fruits having seed and pericarp united only at funiculus. Accessory fruit occurs only in the strawberry and closely related species.

Wild strawberry as a range forb was discussed by the Forest Service (1941, p. W77) and Hermann (1966, ps. 24-25).

Edge of black oak- and post oak-dominated oak-hickory forest, Ottawa County, Oklahoma. Inflorescence photographed in April; fruit photographed in May.

 

364. Rough around the edges- Large, robust specimen of rough-fruit or sulphur chiquefoil (Potentilla recta) growing on an ecotone of tallgrass prairie and oak-hickory forest (a savanna form) in the western Ozark (Springfield) Plateau. This large forb is a member of the rose subfamily (Rosoideae) of the rose family (Rosaceae). Rough-fruit chiquefoil is aparently palatable (at least to some extent) given that it is almost never found on heavily grazed--not to mention, overgrazed--ranges and tame pastures.

This photograph was taken in full sun in contrast to the first slide in the next slide-caption set that was taken under overcast sky.

Ottawa county, Oklahoma. Early June; peak-bloom phenological stage.

 

365. Flowering not foiled, plus four unfoiled chiques- Large specimen of sulphur or rough-fruit chiquefoil in first slide and four flowers in second slide growing on ecotone of tallgrass prairie and an oak-hickory savanna in western Ozark (Springfield) Plateau. This individual was almost a yard tall; somehow it managed to produce this herbage and profuse flowering during Moderate Drought (Palmer Index).

Ottawa county, Oklahoma. Early June; peak-bloom phenological stage.

 

-

366. Leaves not foiled either- Shoots (first slide) and characteristic leaves (second slide) of sulphur or rough-fruit chiquefoil growing on ecotone of tallgrass prairie and an oak-hickory savanna in western Ozark (Springfield) Plateau. These parts were o/of the same plant that was introduced in the first slide in the immediately preceding slide-caption set. It had produced these organs in/under Moderate Drought.

Five-finger is another common name for Potentilla species. Five-finger allludes to five (though these can vary from three up to seven) leaflets comprising one compound leaf of some species as shown in these two slides.

Steyermark (1963, ps. 826-830) described eight Potentilla species for Missouri. Three of these Potentilla species grow in the Ozarks and Osage Plains(of the Central Lowlands) provinces. Chiquefoil or five-finger species are in the rose subfamily (Rosoideae) of the rose family (Rosaceae).

Potentilla species, at least the herbaceous (forb) species, are generally quite palatable. Preference relative to other--especially climax grass--species was not known to this author, but he has observed, over a lifetime, that both P. recta and P. simplex are almost always found only on prairie hay meadows or ungrazed sides (outsides) of fencerows as was the case for this specimen that grew to this fine stage less than ten feet away frrom a dreadfully overgrazed introduced pasture of tall fescue (Festuca arundinacea). This is yet another example that what those landowners who happen to be stupid and greedy (like the one who had the adjoining overgrazed pasture) call "weeds" are not weeds at all except in monoculture agronomic fields. The "little ole ladies in tennis shoes" who call native plants like P. recta "wild flowers" are closer to truth than "weed"-obscessed, amateurish land managers.

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

 

367. Pokeweed, poke salad, poke berry, or poke (Phytolacca americana)- Poke, one of only teo species of this family (Phytolacaceae) in North America, is one of the largest native perennial forbs on the continent. It is a native, rank-growing, drought-tolerant, colorful, herbacaeous species that prefers fertile soil though somewhat bare ground (ie. some degree of disturbance or rich soil). Pokeweed is a weed mostly in name only or in farm field, orchard, vineyard etc. It is usually not a climax species except in the understories of certain forest, woodland, and savannah sites. On tallgrass prairie presence of poke typically indicates degradation, but it is a common plant at edges (points of contact) of forest and grassland. It is, in other words, a widespread and often common plant of savannas.

The first of these photographs was of single five-year-old plant with numerous shoots reaching heights of about nine feet. This individual grew from a seed on a denuded area caused by uprooting of a large honey locust from straight line winds five springtimes ago.This poke plant was growing in the "crater" (cavity in land surface of approximately two foot depth) formed when the large honey locust was uprooted.

The second photograph presented a general perspective of two limbs coming off of one of several shoots of a single poke plant. Flowering/fruiting units varied in phenological development from green fruit to unopened flowers.

The fruit of pokeberry is a favorite of wildlife, especially passerine bird species (Martin et al., 1951, p. 392), which explains much of its rapid population of locally disturbed spots such as that around perimeters of brush piles (planty of bird perches), fires, cleared fencelines, etc.

This author has long felt that the utility of pokeweed as a native landscaping plant has been overlooked by too many native plant enthusiasts. Recently more people have come to appreciate the beauty and practical value of pokeberry as a native ornamental for lawn and garden.

In addition (and maybe more importantly to hillbillies) the shoots (both stems and leaves) of young poke make ideal cooked greens. In fact, it is this human food aspect that is source of the word "poke" which in southern parlance means "sack". Hill folk would (some like this author still do) gather tender, spring poke shoots (say less those about six inches to a foot or so in height) of poke which would be put in their pokes, usually old flour and gunny sacks, (ie. poke poke in the poke) and brought back to the kitchen. Shoots should be boiled (just like cooked spinach) until cooked to desired degree of doneness (this author prefers poke "biled" to a firm yet thoroughly cooked state; nothing like raw, if you please). A precaution against any toxic principles in poke tissue is to pour off all water after boiling for several minutes and then add fresh water to finish boiling the shoots.

CAUTION: Phytolacca americana is or can be a toxic plant. Safe consumption of poke salad requires careful preparation. Readers were referred to Burrows and Tyrl (2001, ps. 857-861), the contemporary authority for poisonous plants in North Amreica. These authors specified that poke contains such toxicants as saponins (glycosides of phytolaccagenin, phytolaccinic acid, and pokeberrygenin) as well as oxalates (oxalate salts). The uncooked roots are quite poisonous as can be the berries if eaten in sufficient quantities, but if one is hillbilly enough to know how to cook poke it is "potherb par excellence" as Burrows and Tyrl (2001, p. 858) noted when they quoted Euell Gibbons.

Ottawa County, Oklahoma. First photograph, June; second photograph, July; pre-flowering to early, immature fruit stages.

 

368. Leaves, flowers, and fruit of pokeweed- Closer-in view of leaves and flower/fruit clusters on a single plant of pokeweed in the western portion of the Ozark (Springfield) Plateau. Flowers of pokeweed differ somewhat from the "typical model". Petals are absent from members of the pokeweed family but there are sepals (five sepals in pokeberry). Stamen are borne on a fleshy, plate-resembling structure (Smith, 1977, p. 102). There are ten stamens and styles. Poke fruit is a multi-locular berry with one ovule (seed) per locule produced from a pistil comprised of a number of fused carpels (Fernald, 1950, p. 606; Smith 1977, p. 102).

The overall inflorescence type (and fruit cluster) of pokeweed is a raceme (Fernald, 1950, p. 606), examples of which were presented in the last two of these three photographs. In the second slide a young raceme has phenological stages going from unopened floral buds to green fruit). The third photograph was of a raceme with fruit stages going from shed (already fallen) through "dead ripe" (fully mature) to just formed.

Ottawa County, Oklahoma. First two slides: July; pre-flowering to early, immature fruit stages. Third slide: September.

 

369. Most at home where its more open- Dense population of tall bellflower (Campanula americana) growing in a small clearing on a north slope in the western portion of the Ozark Plateau (Springfield Plateau portion). This widespread annual species is equally at home in grassy glades, clearings or cutover forest, and oak-hickory-tallgrass savanna. The major determinative growth requirement being adequate sunlight. Obviously, recurrent fire (a regime of frequent, and thus light or low-intensity burning) benefits this sciphyte.

It is also obvious that this is an r-selected species, an annual that allocates a tremendous quantity of its resources for production of shoots with heavy biological investment in sexual reproduction. Tall bellflower is a pioneer species that colonizes habitats of recent disturbance. It is one of the showest forbs in the Ozark Plateau Region.

Newton County, Missouri. Early July.

 

370. Beauty at closer distance- Detailed views of inflorsences of tall bellflower growing in the Springfield Plateau of the Ozark Highlands. It was explained in the immediately preceding caption tht this is an annual pioneer plant species that allocates considerable resources to production of immense, rank shoots that expends heavily in sexual reproduction. Even the most weary woods traveler pauses to admire the beauty of this native forest and savanna forb.

Newton County, Missouri. Early July.

 

371. Standing cypress or red Texas star (Ipomopsis rubra)- This biennial forb of the phlox family is regarded by many native plant proponents as the single most striking and brillant wild flower in Texas. Standing cypress is often found in small colonies (such as shown here) in glades, wind throw openings, and edges of the Cross Timbers and adjoining prairies like the Grand Prairie. The biennial life cycle is the least common one. It is found in some of the Compositae (many of the common thistles) and Umbelliferae. The overwintering rosette of standing cypress is quite conspicuous but the first-summer seedling is so small as to go unnoticed. Then the next spring, wow! Even a typical "bat-out-of-hell" Texas motorist is hard put to miss the blazonry of this Cross Timbers roadside beauty.

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

 

372. Inflorescence of standing cypress- Brillant flowers arranged along the shoot apex of Ipomopsis rubra. Tarleton State University Hunewell Ranch, Erath County, Texas. May.

 

Even a showy base- Spring stage of the leaffy base of standing cypress or red texas star. This striking forb is a biennial that forms the "standard" winter rosette which, as sen here, is also quite attractive. Great plant for prairie-scaping and a favorite among Texas wildflower and native plant fans.

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

 

373. One of a big genus- Upper shoots showing overall habit and inflorescence (first slide) and more detailed views of stem and leaves (second and third slide) of annual wild-buckwheat (Eriogonum annuum) growing on a sandy loam soil in the West Cross Timbers in northcentral Texas.

Annual wild-buckwheat is usually most abuncant on disturbed environments (Diggs et al., 1999, p. 899). That was the case for the specimen presented here which was growing on an overgrazed old-field range.

Best reference for annual wild-buckwheat this author could find Tyrl et al. (2008, ps. 314-315) which, by the way, is one of the finest treatments for numerous, major range plants in Oklahoma, southern Kansas and north Texas.

Hunewell Ranch, Erath Couhty, Texas. Late September; peak-flowering stage of phenology.

 

374. "Hey, Buckwheat" (from Our Gang or The Little Rascals)- Infloresceence (first slide) and groups of flowers of annual wild-buckwheat growing in the West Cross Timbers of northcentral Texas. Apparently, the exact origin or literal meaning (if any) for the common name of wild-buckwheat was not well-determined, but it is known that North American Indians used parts of Eriogonum species for food (eg. seeds used as mash) and a tea-like beverage (Moerman, 2010). Eriogonum species, including E. annuum, were used medicinally by North Amereican Indians (Tyrl et al., 2008, p. 315).

There are many Eriogonum species in North America, especially in the Intermountain West. Correll and Johnston (1979, ps. 511) reported roughly 225 Eriogonum species in North America. By contrast, Correll and Johnston (1979, ps. 510-516) gave a key for and descriptions of only 19 Eriogonum species in Texas. Three of these 19 Eriogonum species are found in northcentral Texas,

On Hunewell Ranch of Tarleton State University this author has found all three of these Eriogonum species. One of these species is a perennial, one is typically biennial or annual, and one is an annual. Heartsepal wild-buckwheat (E. mulitflorum), the biennial was treated immediately below beause both heartsepal and annual wild-buckwheats are more characteristically Cross Timbers species. The perennial E. longifolium grows on calcareous soils and is tyopically a prairie plant rather than a Cross Timbers species.

Tarleton State University Hunewell Ranch, Erath County, Texas; early September (first and third slides). Former Brian Edwards Ranch, Wichita County, Texas; early October (second slide).

Location note: E. longifolium was included the chapter on Tallgrass Prairie Plants in Range Types of North America.

 

375. Broad-topped forb in Cross Timbers- Manyflowered or heart-sepal wild buckwheat (Eriogoum multiflorum) in an ecotone between a stand of post and blackjack oak in West Cross Timbers and a virgin, mima mound prairie in northcentral Texas. This annual (rarely biennial) lacks the brilliance of some forbs like standing cypress (but then what doesn't?), but it is showy in its own way-- and it certainly is prolific. This specimen was growing in deep sand.

West Cross Timbers, Erath County, Texas. October.

 

376. Plain but eye-catching- Details of upper shoot (first slide) and then of flower cluster (second slide) of manyflowered wild buckwheat. This was the same plant introduced in the preceding slide.

West Cross Timbers, Erath County, Texas. October.

Moerman, D. E. 2010. Native American Food Plants: An Ethnobotanical Dictionary. Timber Press.

 

377. Not exactly camouflaged- A green lynx spider (Peucetia viridans), a common spider in the Cross Timbers and Prairies vegetational area of Texas, on a flower cluster of heartsepal wild-buckwheat. Why this bright-green and early stage-pregnant female was bringing Christmas color (and attention to herself) at the edge of a sand post oak (Quercus margarettae)-fiddleleaf green-brier (Smilax bona-nox) dwarf forest in the West Cross Timbers was known only to her. Perhaps she did not know (was cluelessly unaware) of her vulnerablity. Maybe she had some misplaced instinct that she was invinicible. Or perhaps she felt her beauty and "charm" would carry her through anything. Kind of reminded this ole professor of a lot of college students encountered over course of a career in the classroom. Some of these self-styled, attractive personalities apparently thought or instinctively felt that their beauty, charm, excuses, and past experiences with gutless instructors made them invinicible and immune to stated expectations in courses taught by this photographer professor.

Hope the spider fares better. At least she passed this Range Ecology course with "crawling colors". Still, she needs to realize that every living thing--including an eye-catching predator--is something's prey. Just takes one serious "professor" to spoil your record.

Erath County, Texas. Early November; peak flowering phenological stage.

 

378. Not false to itself- Entire shoots (first slide) and upper shoot (second slide) of plant of false boneset (Kuhnia eupatorioides var. corymbulosa) growing at edge of a sand post oak-fiddleleaf gerrn-brier dwarf forest in the West Cross Timbers of northcentral Texas. The specific epithet, eupatorioides, was in reference to the genus Eupatorium and/or the tribe Eupatorieae known by such common names as boneset, thoroughwort, Joe Pye weed, and several others. K. eupatorioides is in the Eupatorieae and, in addition to tribal features, presumedly or supposedly bore enough resemblance to Eupatorium species to get the specific epithet, eupatorioides.

Some taxonomic treatments included Kuhnia in genus Brickellia as in Brickellia eupatoroides (Diggs et al., 1999, ps. 328-329), but Kuhnia eupatorioides is the traditional binominal (Coulter, 1891-1894, p. 180; Fernald, 1950, ps. 1370-1371; Steyermark, 1963, ps. 1468-1469; Great Plains Flora Association, 1986, ps. 966-967).

Erath County, Texas. Early November; peak flowering phenological stage.

 

379. Not falsely blooming- Upper shoots and flower clusters (first slide) and details of individual heads (second slide) of false boneset growing at the outer edge of a sand post or margarettia oak-fiddleleaf green-brier dwarf forest in the West Cross Timbers.

This species is just another one among "jillions" of composites growing on ranges in northcentral Texas. Falseboneset has a biological (species) range that extends from Florida to Texas in the south northward to New Jersey and Pennsylvania and then westward to Illinois and Missouri. There was sufficient morphological variability in this species to recognize three varieties in the greater Great Plains Region (Great Plains Flora Association, 1986, ps. 966-967).

Relative palatability of false boneset is apparently not known, but it is doubtful that grazing animals have much preference for it.

Erath County, Texas. Early November; peak flowering phenological stage.

 

380. Wild hyancinth, eastern or Atlantic camassia, camass, or camass-lilly (Camassia scilloides)- This member of the lily family (Liliaceae) is one of the more widely distributed (though seldom abundant even locally) forbs of the glades and open woodlands of the eastern deciduous forest and hardwood-tallgrass savanna types of eastern and central North America. C. scilloides ranges from Ontario to the Great Lakes and south- and westward to the Cross Timbers and Edwards Plateau in Texas.

According to Dayton (1960, p. 28) in Notes on Western Range Forbs there are six Camassia species in North America. All but C. scilloides are "confined to the Far West". Camassia species have a fairly large bulb which with but one exception were used as foods, often at feasts, among the various Indian tribes. Diggs et al. (1999, p.1200) reported that C.scilloides was an "important food source" for both Indians and backwoodsmen. It's leaves could be used as forage by sheep and deer, but based on Dayton's discussion cattle and horses probably do not select Camassia species.

The specimen pictured here was growing on a limestone outcrop at the top of a bluff (east slope) above a creek in a post-oak-big bluestem dominated savanna at the edge of a black oak-black hickory forest in the western Ozark Plateau. Ottawa County, Oklahoma. April.

 

381. One of several kinds- Plateau spiderwort (Tradescantia edwardsiana) growing in the understorey of a post oak-little bluestem dominated range plant community in the West Cross Timbers of northcentral Texas. In this general area (western portions of the Cross Timbers and Prairies vegetational area) there are four widespread Tradescantia species. Plants of T. ewardsiana are distinguishable from plants of the other Tradescantia species by having leaf blades (at least on upper leaves) that are wider than their opened sheaths (Diggs et al., 1999, p. 1101).

Spiderwort and dayflower (Commelina spp.) are two common genera in the West Cross Timbers that are members of the Commelinaceae, commonly referred to as the spiderwort or dayflower family. Spiderwort species make up some of the showiest range forbs ("wildflowers") in north Texas and southern Oklahoma and, perhaps more important in this context, most prolific and hardiest of the native forbs. Furthermore, plants of most of these spiderwort species are relative large (which also increases their value as native "wildflowers" because they are readily seen by "like bats out of Hell" Texas morortists). Tradescantia species are also quite palatable to livestock as evidenced by their restriction to outside fencerows where domestic grazing is heavy to overgrazing.

Plateau spiderwort is endemic to Texas (Diggs et al., 1999, p. 1102).

Erath County, Texas. Late April; peak-bloom stage.

 

382. No spiders (or pollinators) here- Cluster of showy flowers of plateau spiderwort. These flowers were on one of the plants featured in the shoots seen immediately above. It was remarked in the immediately above caption that spiderworts are some of the most strikingly showy forbs ("wildflowers") in the Cross Timbers Region. This photograph should positively reinforce that assessment.

Flowers of spiderworts are, as a rulke, quite attractive to insect pollinators-- and their predators. In this view that was captured on a warm April morning, neither pollinators or their predaceous enemies (shuch as various spiders) were present.

Erath County, Texas. Late April; peak-bloom stage.

 

383. Virginia creeper or woodbine (Parthenocissus quinquefolia)- This member of the grape family (Vitaceae) is likely the most common woody vine (soft woody and slender stems) in western parts of the eastern decicuous forest, especially the oak-hickory portions (forest cover types). It is often abundant in the ecotone of the oak-hickory forest and tallgrass prairie which takes on the form of a savanna. Virginina creeper is easily identified by the whorled arrangement of five distinctive-shaped leaves. It is mistaken for the various poison ivy/oak species only by greenhorns. This vine covers extensive areas of the oak-hickory forest floor as well as climbing to the tops of hardwood (and pine) species. It is frequently one of the most common ground-covering plants in the oak-hickory-tallgrass savanna of both the Cross Timbers and Prairie Peninsula. Virginia creeper is more common in second-growth and seral forests more than in old-growth or climax stage forests, but it is present in nearly all stages of succession that have trees of any size. It thrives on savannas for this same reason: it gets more light.

Virginia creeper is a widely distributed species occurring from Quebec to Florida to Mexico and Guatemala to the Edwards Plateau of Texas.

Virginia creeper is readily eaten by grazing/browsing animals and often provides copious amounts of feed for livestock, including grass-preferring species like cattle and horses. This fact has not generally been appreciated until livestock are removed from the oak-hickory forests after which Virginia creeper quickly becomes a dominant species in the understorey layer(s) as well as being plentiful (though certainly far from dominant) in the canopy. In this context, Virginia creeper has been an unappreciated indicator species in that presence of this vine generally indicates that grazing has not been excessive. This lack of understanding was borne out by the statement by Crawford et al. (1969, p. 202) that on Ozark forest range woodbine "is seldom grazed by livestock". This statement was in direct opposition to this author's experience and observation in the Ozark Plateau for over 40 years. Abundance of Virginia creeper corresponds highly with absence of or light grazing by cattle, sheep, and horses.

Western Ozark Plateau, Ottawa County, Oklahoma. August.

 

384. Inflorescence of woodbine or Virginia creeper- This creeper was especially abundant in a second-growth oak-hickory forest from which livetock had bveen excluded. Five years prior to time of photograph cattle, sheep and/or horses had grazed this forest continuously for 40 years and no Virginia creeper was present except in an adjacent fence row. Ottawa County, Oklahoma. August.

 

385. Fruit and leaves of Virginia creeper or woodbine- Fall coloration of leaves and mature fruit of a dominant woody vine of the oak-hickory forest and oak-hickory-tallgrass savanna. Ottawa County, Oklahoma. September.

 

386. Coons and 'possums acomin'- Conn or 'possum grape (Ampelopsis cordata) is not a grape (Vitis) species, but it is in the grape family, Vitaceae. Coon grape grows into one of many kinds of liana or woody vine in savannahs (transition zones) between the eastern decisuous forest and tallgrass or true prairies. In other words and to regroup, the grape family (Vitaceae) includes such woody vine species as grapes (Vitis spp), Virginia creeper (covered immediately above), and, now, 'possum grape. Other woody vines in savannahs were treated above as represented in such families as Anacardiaceaeby by poison oak/poison ivy, Bignoniaceae by trumpet creeper, Caprifoliaceae by various species of honeysuckle, and as monocotyledons in the superfamily, Liliaceae or restricted family, Smilaceae by greenbriars. All of these species produce mast for range animals as either dry fruits (eg. drupe in poison ivy) or fleshy fruits (eg. berries in grape, Virginia creeper, and coon grape and, also, honeysuckle).

A. cordata was covered in such reference works as Vines (1960, ps. 707-708) and Kurz (1997, ps. 306-307) as well as florae including Steyermark 1963, p. 1031), Gleason and Cronquist (1991, p. 343), and Diggs et al. (1999, ps.1066).

Ottawa County, Oklahoma. Early June; peak flowering stage of phenology.

 

387. In "fur" a closer look- Leader (branch or shoot) with leaves and flower clusters (first slide) and closer view of leaves and inflorescences (second slide) of coon or 'possum grape in the Springfield Plateau on a savanna between oak-hickory forest and big bluestem-Indiangrass tallgrass prairie. The leaf shape of Ampelopsis cordata is cordate, having "the shape of the stylized heart, with the petiole attached betwen the basal lobes" (Smith, 1977, p. 41), thus the specific epithet of this species. The shoot of 'possum grape is soft-woody in older tissues to herbaceous in immature tissues such as that on outer (distal) parts of shoots such as those shown here.

Ottawa County, Oklahoma. Early June; peak flowering stage of phenology.

 

388. Clusters of preparation-Inflorescences of coon or 'possum grape in a oak-hickory-tallgrass savanna in the western Ozark Plateau. Flowers of 'possum grape were described by Diggs et al. !1999, ps. 1065, 1066) as being small, greenish, and perfect in an axillary, peduncled, compound unbel-like inflorescence.

Ottawa County, Oklahoma. Early June; peak flowering stage of phenology.

 

389. Furbearers' appetizer- Individual fruits in clusters of coon or 'possum grape. Fruit type: berry, a fleshy fruit which is a true fruit (fruit derived from the ovary of one flower) the eintire pericarp (the fruit wall) of which is soft and surrounds several seeds; " a multiseeded, indehiscent (non-opening through sutures, pores, etc. fruit in which the pericarp is fleshy throughout" (Smith, 1977, ps. 69, 290, 298, 303).

Furbearers like coons, 'possums, skunks, foxes, and coyotes as well as rodents, birds, and ruminants readily eat berries of this semi-woody vine, although they might prefer larger-fruited mast such as grapes, persimmons, pawpaws, and plums among fleshy mast or, even, acorns, pecans, and dent corn among dry mast. The author has found seeds of coon grape in scat of coons, coyotes, 'possums, and white-tailed deer.

Ottawa County, Oklahoma. Late June; immature but ripening fruit stage.

 

390. Not quite a truffel- Even by hillbilly standards the fruiting body of dye stone, pea stone, or false puffball (Pisolithus arrhizus) is not edible, but this interesting organism still has value for hill folk. Its toxic pigments can be used for home dying of fabric like wool. This interesting member of the Sclerodermataceae (This group of fungi was briefly described above under the Cross Timbers.)

Ottawa County, Oklahoma. September.

 

391. Breakin' up- The fruiting body of pea or dye stone disintegrates by loosing layer after layer of it tissues as it releases its spores into the autumn air of an oak-hickory-tallgrass savanna in the Ozark Plateau.

Ottawa County, Oklahoma. October.

 

392. Likin' ya- Myelochroa lichen or Rock axil- bristle lichen (Myelochroa obsessa= Parmelina obsessa) on post oak limb in a post oak-Texas hickory-tallgrass savanna in the Springfield Plateau section of the Ozark Plateau physiographic province. This is one of the more common lichens in this savanna range type. Lichen and oak bark were wet from recent rains. The lichen was thus shown to good advantage at its full morphologically distinguishable stage (in contrast to the dry, dehydrated stage).

Ottawa County, Oklahoma. June.

For lack of a better place- Before leaving the tallgrass-oak-hickory savanna and moving on to brief coverage of glades (and for want of a more "natural" place to insert the topic) a short inclusion of lightening as an agent of defoliation and a cause of savanna vegetation dynamics followed.

393. Initial impact- An adult (and at the peak of adulthood) post oak struck by lightening in a post oak-blackjack oak-big bluestem savanna in the Ozark (Springfield) Plateau. Response of individual trees to lightening hits varies tremendously. Apparently factors ranging from soil moisture (whether relatively wet or dry) to stage of phenology or annual growth cycle (eg. dormant or active growth) to stage of tree life cycle (eg. young tree or old tree; age/size class) can all be critical to life or death following lightening strikes. Physical impacts (extent of organ or tissue damage) also are highly variable from one lightening strike to another. Some trees receive less visible damage than that seen here whereas others are "blowed to smithereens" even more than this one (and it was blasted "purty hard").

This was the condition of the tree 18 hours after it had been struck by a powerful blast (the author was a quarter-mile away and the strike shook the small frame house he was inside of).

This strike took place in a thunderstorm with heavy rain such that there was no fire ignition (ie. no immediate secondary effects).

Ottawa County, Oklahoma. Early June.

 

394. Outcome of lightening strike- This was the post oak shown immediately above that was the victim of a powerful lightening strike. This was condition of the adult post oak 31 days after it was hit. The tree was clearly dead of its lightening wounds. No stump sprouts of tree regrowth in any form. This tree that had been in peak health as a prime adult was dead.

In this instance, results of lightening was only this one dead tree. Heavy rain in the thunderstorm that spawned the lightening prevented any sort of fire that otherwise might have been ignited by the lightening.

Attention was drawn to the selective nature of fire as an agent of defoliation. Throughout this publication it has been stated that all defoliating agents--natural or man-made; biotic or abiotic--defoliate selectively. This slide was a textbook example of that phenomenon. Lightening struck one tree, and even with relatively wet soil, there was not enough conduction of electricity to affect close neighboring trees. With pin-point accuracy lightening took out one tree and left all the others that grew close-by.

Ottawa County, Oklahoma. Early July.

 

395. Needlesome invasion- Five large seedlings or tiny saplings (about four to five years old which is four to five years too old) of eastern red cedar (Juniperus virginiana) growing at edge of a grove of black oak, blackjack oak, and post oak that was surrounded (more or less) by a savanna of these three tree species and big bluestem on three sides and a small tallgrass prairie dominated by big bluestem on the fourth side. This range vegetation was in the western Ozark Highlands (Springfield Plateau) about 12 miles from eastern edge of the Cherokee Prairie in the Cental Lowlands. Oak leaves and dead herbage of big bluestem, poverty oatgrass, and purpletop provided the litter seen around these plants. (Leaves still retained in the tree crown above these plants were those of blackjack oak.)

There had been no fire in this area for over a half century and eastern red cedar was invading the oak grove, oak-big bluestem savanna, and relict tallgrass prairie. Eastern red cedar is a non-sprouting conifer. Leaves and dead grass herbage were more than enough fuel to carry a fire that would kill these invading native conifers. Invasion by eastern red cedar is a major problem throughout oak-hickory forests, oak-hickory-tallgrass savnnahs, and tallgrass prairies across a vast "super-region" extending from the Ozark Plateau to the mixed prairie hundreds of miles to the west.

These slide were taken in late afternoon on a winter day. Direct light on these eastern red cedar correctly showed the nearly dead needles of these large seedlings or small sapling. These needles have nearly senesced so as to be "about half dead"". In a couple of months, a good fire on a dry, relatively calm day with the fuel seen here (and needles even drier than now) rwould incinerate these noxious, invading conifers. Strike a match and listen to Hank Williams sing "Settin' the Woods on Fire".

Ottawa County, Oklahoma. Early January. Ecotone between FRES No. 15 (Oak-Hickory Forest Ecosystem) and FRES No. 39 (Prairie Ecosystem). Clearly K-73 (Mosaic of Bluestem Prairie [K-66] and Oak-Hickory Forest [K-91]. SRM 801 (Savanna). Chert Savanna range site. Ozark Highlands- Ozark Plateau Ecoregion, 39a (Woods et al., 2005).

 

396. Little needlesome grows into big problem- Seedling (about nine months old) of eastern red cedar growing in a black oak-northern red oak-post oak-big bluestem savanna in the western Ozark Highlands (Springfield Plateau) that had been severely disturbed by rotary shredding. Future shredding make take the top off of this little guy once he grows tall enough to be beaten off, but even that will not kill the tree as it will live as a hedged invader.

Note fleshy seeds (fleshy female cones) at base of this seedling.

Ottawa County, Oklahoma. Late December.

 

397. Start of an invasion- Young (from less than to slightly more than one year of age) seedlings of eastern red cedar in a oak-hickory-tallgrass savanna community at edge of a post oak-black (Texas) hickory-northern red oak forest with a limited herbaceous understorey (Canada and Virginia wildrye were the dominant grass species). Here at the narrow ecotone of these two range plant communities major plant species were broomsedge bluestem, poverty oatgrass, purpletop, and big bluestem (in that relative order of cover and shoot density).

Seed source for these invading eastern red cedar seedlings were several relatively small adult trees that produced extremely bountiful crops of fleshy cones and grew only about 150 yards away from this site of invasion. Examples of fleshy cones from these yound eastern red cedar were presented in the next four-slide/caption unit. Now where's that remote…

Ottawa County, Oklahoma. Late December.

 

398. Seeds of invasion- Fleshy female cones (= fleshy seeds) of eastern red cedar at progressively closer camera distance. The first slide presented the outermost portion of a lower branch to show the density and absolute number of mature seeds that had been produced in just part (a small part) of the tree. Every leader (= branch) of this tree was "loaded" with female cones just like the portion shown here. The second slide was of a part of the larger portion of one branch shown in the first slide. This second photograph gave viewers a better grasp on the number of seeds borne by just one small easter red cedar tree. Details of these simple fleshy cones and needles of eastern red cedar were shown in the two macro-lens shots (third and fourth slides) of these organs.

These fleshy cones were produced on a comparatively small adult tree that had invaded an oak-hickory-tallgrass savanna in the Springfield Plateau in northeastern Oklahoma. This tree was one of the seed sources for eastern red cedar seedlings that had invaded an ecotone between an oak-hickory forest and an oak-hickory-tallgrass savanna in the Springfield Plateau. Part of that invasion was presented in the immediately preceding slide/caption unit.

Ottawa County, Oklahoma. Late December.

 

399. Roots of the invasion problem- Recently pulled seedlings of eastern red cedar showing their well-developed root systems. The most "ecologically impressive" part (meaning the most conspicuous, adaptative morphological feature) of these roots was the long, large laterals. These well-developed lateral roots on such young seedling-stage trees was obviously an adaptation for rapid aquisition of water from the upper soil profile. Long, extensive lateral roots is an evolutionary (natural selection that results in greater fitness) trait which enables "baby trees" of his coniferous species to obtains shallow soil water before such water either evaporates to the atmosphere or percolates downward below the root zone. In regards to getting soil water, it is likely that seedlings of eastern red cedar can use their limited resources to greater advantage by growing longer lateral roots than by growing deeper tap roots. This would be one form of water use efficiency (ie. more water absorbed per unit of root growth by extending outward rather than downward).

Lateral root development is one ecological "trade secret" in survival of eastern red cedar in shallow, rocky soils such as those of the Ozark Plateau and Flint Hills and in semiarid climates like that of the mixed prairie. Compare the relative area from which soil water could be absorbed by these, spreading lateral roots compared to area in tree crown of these eastern red cedar seedlings.

Well-developed, extensive, sprawling lateral roots is a major morphological feature that enables eastern red cedar to invade grasslands such as tallgrass prairie and mixed prairie, forests like those of the oak-hickory association, and savannahs as, for example, the oak-hickory-tallgrass Cross Timbers.

One general adaptation lacking in eastern red cedar is to fire. Given ready invasion of numerous grassland, forest, and savanna range types by eastern red cedar, as facilitated by such morphological features as large lateral roots, fire in range vegetation with high fuel loads counteracts the water-acquiring competitive advantage of this non-sprouting, ecologically invasive conifer. Sic 'em fire. Rangemen and foresters need to prescribe fire as a natural factor for keeping eastern red cedar from invading grasslands and hardwood forests. Incinerate eastern red cedar and singe Smokey Bear's hair.

Ottawa County, Oklahoma. Early January.

 

400. Ozark Chert Glade- Glades makeup one of several kinds of range "barrens". Nestled among the oak-hickory forest and the tallgrass prairies in the Ozark Plateau are small areas of rock outcrops or shallow, rocky soils on hilltops, and other similar kinds of land and its vegetation known as "glades". In other words, ithin this southern portion of the Prairie Peninsula there are locally restricted sites that are edaphic or topographic climaxes within the climatic climax of the western extension of the Eastern Deciduous Forest and the Tallgrass Prairie Region. Nelson (1985, p. 99) defined glades as "rocky barrens dominated by a herbaceous flora with sparse woody vegetation". This fits the standard definition of savanna. In fact, savannas and barrens traditionally have been considered as similar environments (see especiallly Anderson et al., 1999). Nelson (1985, p. 99) further specified that glades " ...are topographically located on moderate to steep slopes in deeply dissected drainages or hilly to mountainous terrain, and usually have a southern or westen exposure". Bedrock is at or near the soil surface and the shallow soils are subject to frost heaving in winter and are very dry during much of the frost-free growing season following soil saturatrion during spring, sometimes fall.

Glades or, as they are more commonly known by the local hillbillies, "balds" or "bald knobs" are one form of "barrens". Glades are classified as to their parent material. Missouri has greater diversity in balds than any state and has limsteone, dolomite, chert, sandstone, shale and igneous glades. The bald seen here in the Ozark Highlands of southwestern Missouri is a chert glade derived from massive brecciated chert of the Elsey Formation (Nelson, 1985, p. 106). An ephemeral (“wet-weather”) stream flows through the center draw in early to mid-spring and there are seeps from under the chert outcrops which support the “runt” or scrub form of blackjack and post oaks (Quercus marilandica and Q. stellata) and common hackberry (Celtis occidentalis). The domiant grass with the “lacy” or "dainty" appearance in the center and left-center foreground is Festuca paradoxica. The dominant yellow composite is tickseed coreopsis (Coreopsis lanceloata). This is the bottom of the bald which supports the vernal, cool-season, most mesophytic species. The surrounding oak-hickory forest was described immediately below. Wildcat Glade Natural Area, Newton County, Missouri. Vernal aspect, May. FRES of the glade vegetation is 39 (Prairie Ecosystem), K-66 (Bluestem Prairie), FRES of the neighboring forest is No. 91 (Oak-Hickory Forest), and FRES of the combined glade and forest vegetation is No. 73 (Mosaic of Numbers 66 and 91). SRM 803 (Missouri Glades). Range site is Limestone Rock Outcrop. Ozark Highlands- Springfield Plateau Ecoregion, 39a (Chapman et al., 2002).

 

401. Mid-slope vegetation on an Ozark Highlands chert glade (= chert barrens)- Above the mesic habitat of the draw in the chert glade seen in the previous slide is mid-slope vegetation dominated by big bluestem (Andropogon gerardii) and an uplant ecotype of switchgrass (Panicum virgatum). The scrub trees are blackjack oak, post oak, and hackberry. These same oak species grow in the surrounding oak-hickory forest which, growing interspersed with tallgrass prairie (the Prairie Peninsula), is the climatic climax vegetation. The forest has as its dominant, however, black oak (Quercus velutina) which besides post and blackjack oak has as associates black hickory (Carya texana), red or sweet pignut hickory (C. ovalis), and mockernut or white hickory (C. tomentosa). Redbud (Cercis canadensis) and flowering dogwood (Cornus florida) are dominant shrubs of the adjacent forest. Glade vegetation has variously four to five layers: 1) tallgrass species like big bluestem and switchgrass, 2) scrub blackjack and post oak trees of single stems but no taller (usually shorter) than the mature tallgrasses such as to be a shrub layer, 3) a mid-height forb and grass layer (eg. composites, festucoid grasses), 4) xerophytic low herb-shrub layer (short forbs, ferns, cactus; especially common on chert outcrop ledges), and 5) xerophytic soil and rock surface layer (cryptogams). Layer 5 is not present at all microsites. Wildcat Glade Natural Area, Newton County, Missouri. Estival aspect, August. FRES of the glade vegetation is No. 39 (Prairie Ecosystem),K-66 (Bluestem Prairie) FRES of the oak-hickory forest is No. 91, and FRES of the combined glade and forest vegetation is No.73 (Mosaic of Numbers 66 and 91). SRM 803 (Missouri Glades). Range site is Limestone Rock Outcrop. Ozark Highlands- Springfield Plateau Ecoregion, 39a (Chapman et al., 2002).

Location note: greater coverage of chert and dolomite glades was presented in the Grasslands chapter entitled, Miscellaneous Grasslands. Glades of these forms are most correctly seen as grasslands rather than as savannahs, but some examples of such glades were included at end of the current chapter to provide some sense of continuity of range vegetation types and to serve as a set of directions to their location based on more precise vegetational criteria.

 

402. Ozark Bald Knob- This is a dolomite glade or dolomite barrens in the Ozark Plateau. Such distinctive habitats and their vegetation are known to the local hillsmen as “balds” or “bald knobs”. In virgin condition dolomite balds were almost entirely grasslands devoid of woody growth except for scattered individuals of yellow-wood or smoke tree (Cotinus obovatus), the most common shrub, smooth sumac (Rhus glabra), skunkbush sumac, red bud, and persimmon (Diospyros virginiana), all of which are in this scene. Dominant plants are the prairie grasses Indiangrass, prairie dropseed, and big bluestem with sideoats grama and poverty dropseed (Sporobolus neglectus) as associates. The dominant forb which is co-dominant with the three dominant grasses is prairie dock (Silphium terebinthinaceum). Forb associates include calamint (Satureja arkansana) and longleaf bluets (Houstonia longifolia). Except for the dominant prairie dock, composites are rare and locally limited to yellow coneflower (Echinacea paradoxa). Lichens are common only on xeric microsites and much less developed than on the chert glade seen immediately above. The tall blue-green grass in the right foreground is Indiangrass, the dominant species of the bald. The tall yellow-flowered composite in center foreground is prairie dock, the dominant forb.

The Ozarks are ancient mountains worn down to their roots. The dolomite geologic strata are of the Cambrian-Ordovician formations, some of the oldest in these "everlasting hills". Note the concentric ring arrangement of the Cambrian-Ordovician formations.

Note also that this is bald is relatively free of invading trees, which unfortunately surround this oasis of pristine vegetation. The dominant invaders are eastern red cedar (Juniperus virginiana) and Ashe cedar or Ashe juniper (J. ashei). This glade has been kept open by prescribed burning which handily controls the non-sprouting noxious cedars yet allows maintenance of endemic shrubs like yellow-wood. Several ecological reviews of Ozark vegetation prior to European settlement have been made based on surveyers records, travelers journals, Army accounts, etc. (Steyermark, 1940; Beilmann and Brenner, 1951; Howell and Kucera, 1956; Steyermark, 1959). All these ecological surveys proved conclusively that throughout the Ozark Mountain Region oak-hickory-pine forests had expanded and invaded the pre-white man prairies and savannas. Cessation of fire along with farming, overgrazing, and commercial activity weakened the prairie sward and allowed the woody invasion that reached proportions of massive afforestation. The Soil Survey of Taney County Missouri, the adjoining county to the west of this glade, cited a mean fire frequency of 3.2 years on some Ozark glades and credited this as “the mian reason for this open grassy uypland landscape” (Natural Resources Conservation Service, 1996, p. 41).

A more colorful testimony of the original openness of these balds is the gripping tale of the “Bald Knobbers”, post-Civil War “night riders” (vigillantes) who held clandestined meetings and lite signal fires atop bald knobs so as to be able to spot any unwelcome visitors and send coded messages to citizen peace officers (at least before vigilante justice degenerated into blood fueds). Today, most of the Bald Knobbers grassland meeting places are forests or cedar thickets.

McClurg Glade, Ava Ranger District, Mark Twain National Forest, Ozark County, Missouri. Estival aspect, July. No specific FRES or Kuchler designations for vegetation at this restricted scale, but it is primarily an “island” of FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie). However, at landscape scale this is K-74 (Cedar Glades, Quercus-Juniperus-Sporobolus) which is contradictory because by definition glades are open areas of herbaceous vegetation so there is no way that oak and juniper could be the dominant genera. SRM 803 (Missouri Glades). Ozark Highlands- White River Hills Ecoregion, 39c (Chapman et al., 2002).

 

403. Bald knob range- Two photo-plots of a dolomite glade in the Missouri Ozarks maintained (more-or-less) by prescribed fire consistent with the natural fire regime. In the first photograph eastern red cedar and Ashe or post cedar were reinvading the bald knob. This is the Kuchler (1964; in map of Garrison, 1977) physiogonomic type designated Cedar Glades which recognized Juniperus virginiana and J. ashei as the dominant woody species of this potential natural vegetation. In absence of fire (and probably aided by improper grazing, especially overgrazing) these two junipers increase to such cover and density to result in a post-climax juniper woodland instead of the climax juniper-tallgrass savanna.

In the second photograph junipers were largely limited to the rock outcrop perimeter of the dolomite glade (where there is insufficient fuel to carry fire that kills these non-sprouting conifers). Instead the invading woody species was smoke tree (smoke bush) or yellow-wood. Also on the perimeter were individuals of smooth sumac, skunkbush sumac, sassafras, and redbud. These shrubs are well-adapted to fire. In fact, smooth sumac typically increases with range burning. Yellow-wood matures quickly and becomes senescent, but fire serves to "rejuvenate" older individuals of smoke tree through resprouting as was case for the young stump sprout in left center foreground of this second photograph.

Major (dominant) grasses were Indiangrass, prairie dropseed, and big bluestem. Numerous individuals of Indiangrass were readily distinguishable in these photographs as photographs as the tallest, bluish-green clumps of grass. Interestingly and for unknown reasons, many of the grass clumps of light-green (a "washed-out" green) color were also Indiangrass. In fact, several of these Indiangrass plants (prominent in center midground of first slide) had shoots of both the regular chalky blue coloration and the uncommon pale green color. Indiangrass is a long-shoot tallgrass that elongates the culm early in the warm growing season whereas big bluestem is a short-shoot tallgrass species that does not elongate the culm until much latter in the growing season (often two months or more after Indiangrass). Large specimens of Indiangrass were visible in front of the yellow-wood sprout in the second slide. Dead shoots topped with prominent panicles were specimens of the cool-season (hence, now dormant) Junegrass.

A nice specimen of prairie dock, the common dominant forb on bald knob glades, was in front of the young cedar in center foreground of the first slide. White-tailed deer (Odcoileus virginianus) bucks had been burnishing their antlers on this sapling juniper resulting in severe damage to the conifer and range imporvement (aid in restoring the range plant community to potential natural vegetation) on this dolomite glade.

FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie) and K-74 (Cedar Glades). Prescribed fire has been used here at intervals of three to eight years. SRM 803 (Missouri Glades). Ozark Highlands- White River Hills Ecoregion, 39c (Chapman et al., 2002).

 

404. Grass and dock on the glade- Details of species composition of the herbaceous layers of this Ozark glade range. This example of bald knob vegetation was a consociation of prairie dropseed. It was a example of Clements' true prairie (in contrast to tallgrass prairie dominated by bluestems and Indiangrass). Tallgrass species of Indiangrass, big bluestem, and little bluestem (in that order) and prairie dock were associates to prairie dropseed. Other grasses included sideoats grama, poverty dropseed, and Junegrass. Other forbs were gayfeather willow-leaf sunflower (Helianthus salicifolius), compassplant, calamint and longleaf bluets.

The sward of this bald knob range had been maintained by three prescribed fires in recent records: 6, 12, and 22 years before time of this photograph.

FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie) and K-74 (Cedar Glades). Prescribed fire has been used here at intervals of three to eight years. SRM 803 (Missouri Glades). Ozark Highlands- White River Hills Ecoregion, 39c (Chapman et al., 2002).

 

405. Detail of the vegetation of McClurg Glade- Prairie dock and prairie dropseed are the dominants; Indiangrass is the local associate. McClurg Glade, Ava Ranger District, Mark Twain National Forest, Ozark County, Missouri. Estival aspect, July. FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie) and K-74 (Cedar Glades). Prescribed fire has been used here at intervals of three to eight years. SRM 803 (Missouri Glades). Ozark Highlands- White River Hills Ecoregion, 39c (Chapman et al., 2002).

 

406. Friends on the glades- Capitulum (head) of prairie dock being visited by an American bumblebee (Bombus pennsylvanicus) on a chert glade in the Ozark (Springfield) Plateau in southwest Missouri. There are a number of Bombus species, and their identification can often be difficult. Identification of the closely resembling species (not to mention different color patterns or markings between the two sexes). Identifiction is usually best accomplished in the field, but photographic identification is frequently possible, especially when slides are projected. The comparatively large and bright yellow patch on thorax of this visitor was ready proof of this example of the American bumblebee.

McDonald County, Missouri. Late July; busy nectar season for bumblebees.

 

407. Longleaf bluets (Houstonia longifolia)- A common forb on the more xeric microsites of dolomite glades or bald knobs. McClurg Glade, Ava Ranger District, Mark Twain National Forest, Ozark County, Missouri.

 

408. An Easterner- Two plants of American aloe or false aloe (Agave virginica) growing at edge of a chert glade in the western Ozark (Springfield) Plateau. This small Agave species is the only kind of its genus to grow east of the Mississippi River and in the more eastern portion of the TransMississippi West such as central Oklahoma and Texas. As indicated by the specific epithet, virginica (meaning Virginian), this Agave species is found from Virginia and West Virginia across to Ohio and Illinois and south to Florida and west to Texas (Fernald, 1950, p. 454).

American aloe is not strictly limited to chert and dolomite glades, but it is clearly a characteristic species of bald knobs and barrens. There is probably not another plant species that is any more indicative of these glade habitats. American aloe is one of the original "bald knobbers".

Another common name for this Agave species is rattlesnake-master, apparently a folk reference to its supposed use by Indians as a treatment for snakebite. This is a source of confusion because that common name (again in reference to use in treating snakebite) was also used for Eryngium yuccifolium in the Umbelliferae.

McDonald Couny, Missouri. Late July; maturing fruit stage of phenology.

 

In its heart- The "heart section" and stalk of capsules of American aloe growing at edge of a chert glade in the Springfield Plateau subsecction of the ozark Plateau physiographic province.

McDonald Couny, Missouri. Mid-July; maturing fruit stage of phenology.

 

409. A unique glade fruit- Cluster of capsules on the sexual shoot (first slide) and closer-in view of capsules (second slide) of American aloe growing on the edge of a chert glade in the western Ozark Plateau (western part of the Springfield Plateau portion of the general Ozark Plateau). These organs were on the two plants introduced in the preeding slides.

McDonald Couny, Missouri. Late July; maturing fruit stage of phenology.

 

410. Invasion of dolomite glades or bald knobs by eastern red cedar, primarily, and with some Ashe cedar. At landscape scale this vegetation was mapped by Kuchler (1966) as K-74 (Cedar Glades, Quercus-Juniperus-Sporobolus) which as shown above was contradictory because grassland is climax and oaks and junipers are at best only aspect dominants. True dominants are Indiangrass, prairie dropseed, and big bluestem. Detailed studies clearly proved that woody invasion occurred and caused (or is the result of) range retrogression due mostly as a result of fire suppression. Fortunately that dim-wit bruin Smokey Bear died (cremation would have been more appropriate than his expensive burial). Now the Forest Service is using prescribed fires to clean up the bald knob country and restore the climax prairie vegetation. It was none-too-soon as seen on these cedar-invested glades over which a fire tower stands sentinel. How ironic, prophetic, and symbolic of good intentions that wrought bad! Fortunately man is a thinking animal.

Three Sisters viewed from Gladetop Trail, Caney Campground, Ava Ranger District, Mark Twain National Forest, Ozark County, Missouri. Estival aspect, July. FRES No. 39 (Prairie Ecosystem), K-74 (Cedar Glades). SRM 803 (Missouri Glades). Ozark Highlands- White River Hills Ecoregion, 39c (Chapman et al., 2002).

 

411. Cedar encroachment on Ozark Glades- Eastern red cedar and Ashe or post cedar with "tag-along" individuals of yellow-wood or smoke tree, sassafras, persimmon, smooth sumac (which increases rapidly and profusly with fire), and chittam-wood or gum-elastic (Bumelia languinosa var. albicans= B. languinosa var. oblongifolia).

This invading woody component was on the edge of the glade presented in the immediately preceding seven photographs where this glade joined unburnt portions. The woody vegetation at this edge served as a control plot from which response to prescribed fire of range vegetation on the prescription-burnt glade could be compared. Patchworks or mosaics of such burned and unburned range vegetation on Ozark Highlands handily fit the concept in Landscape Ecology and Wildlife Management of edge effect. Small tracts of cedar-dominated vegetation were patches within a matrix of glades just as at larger spatial scale glades were patches in an overall matrix of oak-hickory forest (patches inside patches).

Edges between adjoining dolomite glades and cedar-infested glades were small ecotnes that benefitted wildlife species. However, the ultimate state of this Ozark Plateau range type in absence of natural fire or prescribed fire that simulated natural fire regimes is all too often a cedar (juniper) woodland or, ultimately, a cedar forest of closed canopy and no understorey, often with accelerated erosion of soil surface.

The sermon for this range type is a reversal of words, but the same moral as "repent or burn". On range such as this the prophetic message is "repent and burn or regret and perish".

FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie) and K-74 (Cedar Glades). Prescribed fire has been used here at intervals of three to eight years. SRM 803 (Missouri Glades). Ozark Highlands- White River Hills Ecoregion, 39c (Chapman et al., 2002).

Location note: Most coverage of chert and dolomite glades was in the grassland chapter entitled, Miscellaneous. These few slides of glades where inserted here because glades, balds, and barrens within forest regions are closely related to tree-grass savannahs.

 

412. Mountaintop savannah- Here on the top of Mount Scott (elevation 2464 feet) in the Wichita Mountains is a big bluestem-eastern red cedar savanna. Indangrass and little bluestem are tallgrass associates and sideoats grama is the midgrass associate. The dominant forb is broom snakeweed (Gutierrezia sarothrae). Eastern red cedar is wind-pruned thereby having a semi-krumholtz form and forming a tallgrass-juniper savanna or perhaps a mixed prairie-juniper savanna. There is some Ashe juniper (Juniperus asheii) in the Wichitas, but eastern red cedar (J. virginiana) is overwhelmingly the dominant cedar. On dry, rocky hill tops like this the juniper would be less apt to burn and would logically be a component of the climax community such that the potential natural vegetation has to be interpreted as a savanna. As discussed under the previous slide no FRES or Kuchler Units exist at this restricted scale. Likewise there is no SRM rangeland cover type although some of the Texas Edwards Plateau types have a similar floristics and habitat (eg. SRM 733, Juniper-Oak Woodland). Consistent with the granite parent material of this "boulder field" and the inclusion of glades as a form of barrens and primary natural plant community it seemed that the most likely published classification of this unique range vegetation was Cedar Glades (Kuchler Unit 74) or, alternatively, some designation based on parent material (eg. Granite Glade or Granite Rock Outcrop) or perhaps a combination of geologic material and dominant species (eg. Granite Cedar Glades, Granitic Bluestem-Cedar Savanna). The range site designation is most fitting: Hilly Stony Savanna. Central Great Plains- Wichita Mountains Ecoregion, 27k (Woods et al., 2005).

Wichita Mountains National Wildlife Refuge, Commanche County, Oklahoma. Autumnal aspect, October.

 

413. Granite Rock Outcrop- This boulder field resembles the boulder fields of the alpine ecosystem as do the wind-pruned near-krumholtz forms of eastern red cedar. Granite rock outcrop habitats and natural plant communities were discussed variously in Anderson et al. (1999), but none of this dealt with the exact kind of granite outcrop seen here. Big bluestem and Indiangrass are dominant grasses.Shrubs include scrub blackjack oak and Texas red oak or Spanish oak (Quercus shumardii var. texana) which is the larger shrub in the center skyline. Smooth and skunkbush sumac are visible in foreground and eastern red cedar is in far background. There is a diverse lichen community on the granite boulders. This plant community corrresponds closely to that of the chert glades which, as mentioned in the preceding slide, prompts designation of this as a granite glade. The tallgrass-oak-juniper-sumac savanna community is aptly designted as Hilly Stony Savanna range site. Central Great Plains- Wichita Mountains Ecoregion, 27k (Woods et al., 2005). Wichita Mountains National Wildlife Refuge, Commanche County, Oklahoma. Autumnal aspect, October.

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