Southern and Central Forests - IC

Forest Range Types of Eastern North America

Sugar Maple-Dominated Forests

Sugar maple-dominated forest in the Ozark Plateau constitute a more mesic phase(s) of the Oak-Hickory Forest Region (Braun, 168-170). Specifically, sugar maple-dominated forests in the Oak-Hickory Forest Region were maple climaxes (climax sugar maple-dominated forests) the same as (at least, equivalent to) the maple climaxes that define the Maple-Basswood Forest Region of Braun (1950, ps. 327-336). The quote from Braun (1950, p.34) cited in the introduction to this chapter bore reciting: "Each of the several climaxes, although characterizing a specific region, nevertheless occurs in other regions". Braun (1950,p. 164) also stated: "The forest of the most mesophytic slopes usually contain sugar maple and an admixture of other mesophytic species". In other words, sugar maple is a defining, dominant species of some upland, slope forest in the oak-hickory forests. Thus there were the climax sugar maple-dominated forests in the Ozark Plateau that were presented in this portion of the Oak-Hickory Forest chapter.

Three forest tracts dominated by sugar maple were described in this portion of Oak-Hickory Forests-II. These separate tracats of forest range were similar and comparable to published descriptions of sugar maple forests in the Ozark Highlands (Ozark Plateau). Two forms, phases, variants, (or whatever they would be designated) of climax sugar maple were recognized for the Ozark Mountains by forest ecologists. These were the Acer saccharum-Quercus alba associes of limestone slopes and the Acer saccaharum-Carya cordiformis associes" that Braun (1950, ps. 168-169) recognized from the preceding work of Steyermark (1940). Both Steyermark (1940) and Braun (1950) used the Clementsian monoclimax vegetation system (Clements, 1916) in which associes "is the developmental equivalent of the association" so "used where the community is not permanent" but seral (Weaver and Clements, 1939, p. 99). In the geologic time scale of monoclimax theory the limestone bluffs and hills of the Ozarks would be worn down to a peneplane so as to eventually become the regional climax (monoclimax) of oak-hickory forest. Forests of sugar maple (with co-dominant tree species) on more favorable, mesic sites (eg. north and east slopes) are in monoclimax theory, postclimax in oak-hickory regional (climax) forests. In the polyclimax theory of Tansley or climax pattern theory of Whittaker the Clementsian associes of Steyermark (1940) would be associations.

The Acer saccharum-Quercus alba associes and Acer saccaharum-Carya cordiformis associes would be interpreted as variants of the Society of American Foresters (Eyre, 1980, p. 30) forest cover type, Sugar Maple (SAF 27). The forest cover type description of SAF 27 in Eyre (1980, p. 30) included four subtypes, two of which corresponded to the two forms of the moist slope Ozark Highlands forest: the sugar maple-bitternut hickory "restricted to deep soils in the southernmost part of Quebec" and the sugar maple-basswood-white ash subtype "found in the lowlands of the St. Lawrence Valley." Obviously authors of the sugar maple cover type description were unaware that the same subtypes (the associes of Steyermark [1940]) also existed in the Ozark Mountains of Missouri.

This sugar maple-dominated forest range type three variants or forms) was described as mesic-limestone forest by the system of classification and designation in Nelson (1987, p. 28; 2005, ps. 122-125). Two of these variants of the sugar maple cover type had developed on limestone bluffs along (above) Modoc Creek and one was on a steep north slope (limestone parent material though not near a stream) at the western boundary of the Springfield Plateau of the Ozark Highlands. These were presented and described below as sugar maple-bitternut hickory (north slope), sugar maple-white ash- northern red oak (east slope), and sugar maple-northern red oak (north slope).

216. Postclimax of the Oak-Hickory Forest Region- A mesic, north slope of limestone bluffs above Modoc Creek supported a sugar maple-bitternut hickory (Acer saccaharum-Carya cordiformis) forest that was--when viewed from the Clementsian perspective--postclimax for oak-hickory (hardwood, in general) forests of the Ozark Highlands. On this limestone bluff forest sugar maple was the tree species with most regeneration, but this was followed closely by bitternut hickory. Associated tree species were (in this general order) white ash (Fraxinus americana), basswood or American linden (Tilia americana), hackberry or, sometimes, western hackberry (Celtis occidentalis), northern red oak (Quercus rubra= Q. borealis var. rubra), chinquapin (=chinkapin) oak (Q. muehlenbergii), black walnut (Juglans nigra), slippery or red elm (Ulmus rubra), honey locust (Gleditsia triacanthos), Kentucky coffeetree (Gymnocladus dioicus), and sycamore (Platanus occidentalis) .

There were two shrub zones: 1) upper (higher on bluffs; farther from creek) and 2) lower (farther down on bluffs; next to creek). The higher or upper zone had one major shrub layer that was dominated by eastern dogwood (Cornus florida) with eastern redbud (Cercis canadensis) as the associate shrub. The lower zone on bluffs had two major or prominent shrub layers or strata: 1) taller shrub stratum made up almost exclusively of pawpaw (Asimina triloba) and 2) lower (although still relatively high) shrub stratum composed solely of American bladdernut (Staphylea trifolia). There was some buckbrush or coralberry (Symphoricarpos orbiculatus), but this otherwise widespread Ozark Plateau shrub was present only in sub-trace quantities in this mesic, relatively cool, north slope forest.

Locally common to dominant forbs included Jack-in-the-pulpit (Arisaema atrorubens), green dargon (A. dracontium), wake robin (Trillium sessile), trout lily (Erythronium americanum), columbine (Aquiegia canadensis), dutchman's breeches (Dicentra cucullaria), bloodroot (Sanguinaria canadensis), and false Solomon's seal or false spikenard (Smilaceina racemosa). Rue anemone (Aneomella thalictroides) and false rue anemone (Isopyrum biternatum) were the earliest vernal forbs in this forest. Mayapple (Podophyllum peltatum) was present at extremely low cover (it was "barely there") which was "sign-significant" for this climax forest vegetation given the nearly ubiguitous presence of this forb in forest, even savannahs, of the Ozark Plateau. Other forbs were mosses and ferns, including walking fern (Camptosorus rhizophyllus= Asplenium rhizophyllum) and Christmas fern (Polystichum acrostichoides). Ferns were not as abundant (general density and cover parameters) as on an east slope sugar maple forest farther downstream (see below).

Dominant grass overall was silky wldrye (Elymus villous) though locally broadleaf woodoats (Uniola latifolia) was common to dominant. Also local stands of Virginia wildrye. Naturalized Kentucky bluegrass (Poa pratensis) was also common at local scale. Higher up on bluffs the dominant climax grasses were Canada brome or hairy, wood brome (Bromus purgans= B. pubescens ) and woodland bluegrass (Poa sylvestris).

Two photographs provided a longer glance (first slide) followed by a shorter glance (second slide) of the range vegetation of this mesic-limestone forest (Neldon, 1987, p. 28; 2005, ps. 122-125). Snag (right side near margin in first slide; right of center in second slide) was sugar maple. Large tree to its left with trunk injury or damage blaze (both slides) was white ash. Sapling to left of blazed big white ash with missing crown and profuse generation of long shoots was white ash. Sprouts in front and slightly to left of blazed big white ash were American elm with a few individuals of slippery elm. Shrubs were pawpaw (taller; more sparse) and American bladdernut (forming colonies; shorter that pawpaw). Smaller trees (not shrubs) at edge of dense forest stand with conspicuous leaves (partial crown in first slide; only one or two limbs in second slide) was sugar maple.

Along Modoc Creek, Ottawa County, Oklahoma. Late May (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

217. Another view the next year- Another vantage point provided this farther-off exterior view of the same mesic-limestone forest introduced in the preceding two-slide sequence. This sugar maple-bittrnut (pignut) hickory-basswood-white ash-hackberry-northern red oak-chinquapin oakhad developed on a north-slope of limestone bluffs just above Modoc Creek, a smaller though typical Ozarks Plateau stream. This photograph was taken one year after the two in the immediately preceding set..

The large, tall tree in left foreground was the same white ash with blazed scar on upper trunk. One year later the crown of the sugar maple snag (tree just to right of white ash) had lost another major limb. Most of the maller trees (not shrubs) at perimeter of dense forest to left of big white ash were sugar maple. Shrub layers as seen in foreground (as under fallen sugar maple limb) included American bladdernut, pawpaw, and spicebush (in that order of cover and general importance or dominance). Saplings to right and slightly behind (higher on the bluffs) were basswood (larger sapling) and American and slippery elm (farther to right).

Along Modoc Creek, Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

218. From stream bank to tree tops- Ground to crown "photo-transect" of the mesic-limestone mixed hardwoods (sugar maple, bitternut hickory, white ash, hackberry, basswood, black walnut, northern red oak, chinquapin oak, american and slippery elm) Ozark Plateau forest. American bladdernut, pawpaw, and spicebush made up two (or three) distinct shrub layers while sugar maple (biggest, leftmost tree with holes used as dens by coon; many saplings in background), American elm (tree immediately right of maple), and chinquapin oak (background), which were most of the larger trees presented here, made up most of the uppermost canopy.

Most regeneration was sugar maple with that of bitternut hickory, a distant runner-up.

Along Modoc Creek, Ottawa County, Oklahoma. May (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

219. Tall hardwoods that like it cool and moist- A mesic-limestone forest on bluffs above a creek in the western Ozark Plateau. A fairly comprehensive list of major species (trees, shrubs, forbs, and grasses) in this tract of forest was given in the immediately preceding caption. Coverage in this caption was limited to specific featured species. Tree at left margin was black walnut. Sugar maples were to the close-by right and also behind the black walnut. Tall tree in foreground left of center was basswood. A young Kentucky coffeetree was to the immediate right of tall basswood and with crowns of the young tree passing in front of the bsswood. The three trees in right foreground were (left to right): American elm, hackberry, and white ash.

Shrub layer (mostly in foregroune) were mixture of pawpaw, American bladdernut, and some flowering dogwood. The only herbaceous species visible at this distance were a large plant of silky wildrye and some leaves of the otherwise scarce Mayapple.

Along Modoc Creek, Ottawa County, Oklahoma.Late May (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

220. Tall hardwoods later (and drier) in the summer- Two views of the same tract of north-slope, mesic-limestone mixed hardwood forest that developed along Modoc Creek, a stream typical of those in the Ozark Plateau. Overall dominant, including of regeneration, was sugar maple with bitternut hickory, basswood, white ash, northern red oak, chinquapin oak, American elm, and hackberry local associates. Sycamore and black walnut, pioneers of freshly denuded stream channels and banks, had persisted into this climax forest along its exterior.

Forest range vegetation was presented extending from the creek bank to top of limestone bluffs in both of these slides. In the first of these two slides sugar maple (biggest tree; right-center), northern red oak (big tree left-center), basswood (big tree in right margin; background), and American elm (pole to left of big maple) were represented. In the second photograph northrn red oak (biggest tree; right background), chinquapin oak (foremost tree; left-center foreground), white or American and red or slippery elm (one sapling of each left and in front of northern red oak), and bitternut hickory (seedlings in right foreground), and sugar maple (most regeneration in foregoround) were represented (one way or the other).

These photogrphs did not feature shrub layers, but American bladdernut was present and visible in second slide.

221.. At the base of a bluff- By the banks of Modoc Creek in the Springfield Plateau section of the Ozark Plateau (Mountains) some large trees of a mesic-limestone forest created a peaceful sylvan scene that belied the extreme competition for light (and undoubtedly, other resources) in this postclimax forest range vegetation. Tree in left foreground was an American or white elm (no umbrella-shaped crown on this forest dweller). The tree behind (and largely hidden by) the tall American elm with its upper bole leaning right was basswood. To the right (and behind at some distance) of the right-leaning basswood was a smaller sugar maple. Largest tree in this photograph (center foreground) was white ash. Smaller trees to left and right of large white ash were hackberry, including the large tree in right background. (Incidentially, real woodsmen would have noticed the black spot of a varmit in the right background hackberry, and we didn't even need our Black and Tans or Blueticks to tree it.) Smaller trees at right were sugar maple. Most of the young shoots in the understorey were also sugar maple. Regeneration in dense shade is a trademark of this Very Tolerant hardwood.

Most shrubs were American bladdernut. Some pawpaqw were present. The spike of silky wildrye (lower right corner) was conspicuous (and a rangeman's signature on this slide).

If any greenhorns were viewing this who could not locate the coon you might have better luck finding the large rotting log, the species of which could not be determined, that was close by the large white ash (in right center midground). Downed timber, especially big trunks and limbs that require long time periods to rot away, are part of the forest. A tree does not cease to be part of the forest range ecosystem just because it dies. There is life after death in the forest.

222. A logger's view of trees at the base of a bluff- Vertical "photo-transect" of the larger trees introduced in the preceding slide and caption. The greenhorns' log served as focal point. Largest tree to left of log was the large white ash previously noted. Small tree to immediate left of large white ash was hackberry. Foremost tree (near lower left corner) was American elm. Tree at far left margin (only upper bole in photograph) was basswood. Smaller tree with dark trunk between (and behind) the American elm and the upper bole of basswood was sugar maple-- as were most of the seedlings and saplings in the understorey.

Also visible in understorey were large plants of silky wildrye. Most shrubs (foreground) were American bladdernut.

Along Modoc Creek, Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

223. As much light as you get in this woods if you're limited to ground-level- This poorly (barely) lite photograph showed the most light that this immediate location (local habitat) receives once trees have fully leafed-out in this north slope, sugar maple-dominated forest. Biggest tree (21 inches DBH) that was slightly left of center was a dandy sugar maple. Tree to its left (lower trunk at lower left margin) was northern red oak. Small sapling to left of northern red oak was white ash. Dominant shrub was flowering dogwood with eastern redbud the associate shrub species. Main herbaceous (not visible) was the lebuminus forb pointed-leaf or cluster-leaf ticktrefoil or tick clover (Desmodium glutinosum).

224. Regeneration at the bottom- Two side-by-side views of lower layers of the climax range vegetation of a mesic-limestone mixed hardwood (sugar maple, bitternut hickory-white ash-basswood-northern red oak-chinquapin oak) forest that developed along limestone bluffs above a typical Ozark Plateau stream (Modoc Creek). Understorey of the same tract of climax forest treated above and below.

Tree trunk in these slides (upper left corner in first photograph; center of second photograph) as well as smaller trunk in second slide and fallen limb in first slide were sugar maple. Almost all sapling and tree seedlings in both "photo-plots" were either sugar maple or bitternut hickory with maple the more abundant of these generally co-dominant tree species. A sapling of American elm was present in upper left corner of second photograph.Trumpet creeper (Campsis radicans), a native liana, and Virginia creeper had notable regeneration.

The broadleafed forb conspicuous in foreground of both slides was slender nettle (Urtica dioecia= U. gracilis). Its "excort" in lower left corner of first photograph was brown-eyed Susan (Rudbeckia triloba). It is only the lowerrmost leaves of this species that are three-lobed , and these were not visible in this photograph. The other forb in this sample of forest understorey was columbine (Aquiegia canadensis). Columbine was shown to better advantage in photographs below. Many of the other native forest forbs (see list in introduction to this forest range type) were present earlier in the spring growing season, but these had either entered dormancy or were not readily visible at time of photographs.

There were two climax decreaser grasses that formed an almost exclusive herbaceous layer: 1) Canada brome or hairy, wood brome (Bromus purgans= B. pubescens ) and 2) woodland bluegrass (Poa sylvestris). These two species were clearly key indicator species of this entire climax forest community.

225. Deep inside a bluffs forest- Wide-angle view of the lower layers of a mesic-limestone forest dominated by sugar maple with bitternut hickory, white ash, northern red oak, basswood, and chinquapin oak local associates. Dominant shrub was American bladdernut with pawpaw the associate shrub. These two shrubs formed two distinct though discontinuous lower woody layers. Virginia creeper was quite common and occupied almost all layers of this forest including lower tree canopy.Flowering dogwood was present and, while conspicuous during "dogwood and redbud time", confined to upper reaches of bluffs and not common at this level on the bluffs..

There was some regeneration of all of tree species, but sugar maple outdistanced all others. Bitternut or pignut hickory was second in regeneration. Regeneration was both sexual and asexual from basal sprouting.

Columbine (right foreground) was the most prominent herbaceous species at this time and from this camera station. Other common forbs in this photograph (though not visible) were Jack-in-the-pulpit, green dragon, dutchman's britches, rue anemone, false rue anemone, and bloodroot. The more abundant (and less showy) herbaceous species were Canada or hairy, wood brome and woodland bluegrass. Virginia wildrye and broadleaf woodoats formed locally dense stands.

On limestone bluffs above Modoc Creek, Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

226. Climbing the bluffs- Climax vegetation of this mesic-limestone mixed hardwood forest (sugar maple dominant with bitternut hickory, white ash, northern red oak, basswood, and chinquapin oak "swapping places" as local associate species) changes over short vertical distances in progression up limestone bluffs. At this near-to-the-top zone sugar maple dominated the canopy and tree regeneration (various age/size classes as, for example, the sapling featured here). American bladdernut was still the dominant shrub in in this sample of vegetation (foreground of both photographs, especially prominent in lowr left corner of the second).

Canada or hairy wood brome and woodland bluegrass were the dominant and assocaite species, respectively, of the grass(the taller) layer of the herbaceous understorey. Columbine was conspicuous in the second of these slides.

Moss- and fern-covered ledges of the limestone bluffs to "top-off" this calendar cover-like view of pristine Ozark Plateau sugar maple-dominated forest range.

227. These already climbed the bluffs- At ledges atop the limestone bluffs of this mesic-limestone, mixed hardwood forest Canada brome and a northern red oak seedling (lower left corner) had staked their claim to space, moist and shaded soil, and what little sunlight filtered through or fleetingly blazed at full intensity on their small spots. They were joined by columbine and wind flower or thimbleweed (Anemone virginiana).

At top of limestone bluffs above Modoc Creek, Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

228. In its own shade, and more some- Two "photo-quadrants" of the understorey of a mesic-limestone, north slope forest dominated by sugar maple (with bitternut hickory, white ash, basswood, northern red oak, hackberry, and American elm other locally important tree species). There was no doubt as to which was the dominant on this forest range (and most range hungry animals had better like it or go elsewhere). The adult tree (center), saplings (small and large), seedlings, and even root or trunk sprouts were sugar maple.

There was some American bladder, distinguised by its compound (largely trifoliate) leaves, buckbrush or coralberry, and silky wildrye (prebloom). Otherwise, this was an "all-ages gathering" of sugar maple.

The second slide featured shoot or stump sprouting in a sugar maple sapling that lost its crown in a severe icestorm in February (three to three and a half months before time of photograph). This never-say-die small sapling simply started over at the bottom as it were. The ability to reproduce (sexually and/or asexually) in "its own shade" is the defining feature of a species with high tolerance. Sugar maple has a tolerance rating as high as it goes: Very Tolerant. This was according to the Society of American Foresters (Wenger (1984, p. 3) which gave ratings of Tolerant for basswood while white ash, bitternut hickory, hackberry, northern red oak, and American elm were Intermediate.

229. Not dead yet (at least, not quite)- Stump suckering (= coppicing) from an otherwise dead sugar maple on a sugar maple-dominated forest that developed on a bluff above Modoc Creek in the western portion of the Ozark Uplands. These were long or heterophyllus shoots (in contrast to fruit-bearing short shoots). Production of stump suckers is a form of asexual reproduction that is well developed in many hardwood trees and shrubs that are interpreted as having higher levels of Tolerance. Sugar maple has the highest rank of Very Tolerant (Wenger, 1984, p.3). In this tract of north slope, mesic-limestone forest sugar maple had various tree species that were local associates including bitternut hickory, white ash, basswood, northern red oak, and American elm.

Note the Jack-in-the-pulpit and various ferns in front and to side of the trunk and long shoots.

Along Modoc Creek, Ottawa County, Oklahoma. April (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

230. Coppicing in bitternut hickory- Sprouting from two stumps of Carya cordiformis along the banks of Modoc Creek in a sugar maple-dominated forest in which bitternut hickory and white ash were the most consistent associate tree species. This photograph taken in late autumn provided the detail of several bouts of suckering or coppicing in two hickories that had suffered repeated injury (most likely from spring wildfires).

Along Modoc Creek, Ottawa County, Oklahoma. November (autumnal aspect); dormancy in hickory.

231. Jack the ash lost its crown- Two slides showing coppicing (=suckering ; production of long shoots) from high up on trunk of a white ash that lost almost all of its aboveground portion in a severe ice storm. This intermediate-sized tree was growing about mid-way up a limestone bluff above an Ozark Plateau stream in a sugar maple-dominated forest (in which bitternut hickory and white ash were most consistent associaties; other associate tree species being basswood, northern red oak, and white or American elm). A severe ice storm "wrecked havoc" on some trees in this mesic-limestone forest (Nelson, 2005, ps. 122-125). The white ash presented here lost its crown in February, but by late May it had regrown these long shoots (stump or snag suckers). White ash is a strongly coppicing species, at least under certain conditions.

Along Modoc Creek, Ottawa County, Oklahoma. Late May (estival aspect; coppicing in white ash).

232. And Jill the basswood came tumbling after- When the crown of the white ash (discussed with the immediately preceding two slides and caption) crashed in a February icestorm it took an neighoring (and even larger) basswood down with it. In fact the basswood was brought even lower as the additional weight combined with wet soil resulted in complete toppling of the basswood.

The first of these two photographs presented immediate local habitat (microsite) by the toppled basswood (including the stump of trunk from which the landowner had cut firewood). The owner had not worked up the small side shoot of this basswood that was on the ground surface (center foreground to left lower corner). The basswood had sprouted profusely all along this secondary bole. (A morphological feature of basswood is on-going or continued production of secondary shoots off of the main (original) tree trunk.)

Also visible in this first slide were resprouting and seedling plants of white ash, sugar maple, and American elm along with American bladdernut and several pre-bloom-stage plants of silky wildrye, the dominant grass in this mesic-limestone, north slope, Ozark Mountains forest.

The second of these slides showed stump sprouting (=suckering= coppicing) of the basswood in more detail.

Along Modoc Creek, Ottawa County, Oklahoma. Late May (estival aspect; coppicing stage in basswood).

233. Leafy layers- Follow-up "photo-plot" of stump sprouts of basswood in middle of second summer of growth following downing in an ice storm (two winters earlier). This photograph also presented associated herbaceous species including silky wildrye, broadleaf woodoats, and lopseed (Phryma leptostachya). The latter, a forb, is in its own family, Phrymaceae.

On limestone bluffs just above (on upper banks of) Modoc Creek, Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

334. Past-prime autumn coloration (in sugar maples), but still instructive- Late autumn in a north slope, sugar maple-dominated Ozark forest was still an appropriate time to describe the postclimax climax vegetation of this forested range. This was the same tract of forest (along and above Modoc Creek) that was presented and discussed in the preceding slides of the Sugar Maple Forest section. (See again first photo-caption for comprehensive list of major plant species.). Plants featured in the autumn scene were sugar maple (big trees at extreme left and right margins, Big tree in center midground (midway up the bluff) still bearing green leaves was a northern red oak. Tree with green leaves at far left (left margin) was a chinquapin (chinkapin) oak. Trees with retained dead leaves were sugar maple.

Along Modoc Creek, Ottawa County, Oklahoma. Late November (autumnal aspect; leaf fall for most trees and shrubs except the oaks). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

235. Another instructive view of gone-past-autumn color- Vertical look at a sugar maple-dominated forest on a limestone bluff (above Modoc Creek) in the Springfield Plateau of the Ozark Highlands. Largest tree (center foreground; midway up the bluff) and the larger tree on top of the bluff (to left of largest tree) were sugar maples readily distinguished by their large, crooked limbs. Tree in front of and to left of largest tree (and still bearing green leaves) was a northern red oak. The several trees at far right mid- to background included sugar maple, basswood, and chinquapin oak. Most seedlings and saplings of understorey were sugar maple, the Very Tolerant, climax dominant tree species of this mesic-limestone forest (Nelson, 2005, ps. 122-125).

Attention was drawn to the large, flat rock in left-of-center foreground which was limestone.

236. Spring slope- "Photo-transect" on the north slope of the sugar maple-bitternut hickory in early spring. The spring flora of the mesic-limestone forest (Nelson, 2005, ps. 122-125), especially the lower herbaceous layers, that was described in this portion was one of the most unique or conspicuous features of this climax forest range vegetation. This early vernal aspect showed the temporal and spatial variation of this mesic Ozark Highlands forest. Species details were presented in the two immediately succeeding photographs.

The farmer has to "make hay while the sun shines", and so do most of the florest floor-dwelling plants in this sugar maple-dominated plant community. Once the leaves are fully developed on the maples and associated tree species, along with the shrubs, there is inadequate light for most shorter plant species (this includes almost of the herbaceous species). To survive, forest forbs (which dominate the herbaceous layers) must complete their annual cycle of life (= growth cycle) as early as spring temperatures permit their growth and reproduction.

Ottawa County, Oklahoma. April.

237. Details of spring slope- Two "photo-plots" of the vernal herbaceous understorey on a north slope, climax sugar maple-bitternut hickory forest that was introduced in the preceding slide. Species on the north slope of this forest floor included Jack-in-the-pulpit, wake robin, Solomon's seal, false Solomon's seal, wild geranium (Geranium maculatum), green dragon, trout lily, mayapple, Dutchman's breeches, and the fungus known simply as morel (Morchella rotunda..

The first and last of these forest range plant species were conspicuous. Students should try to find some more of the just-listed species in these two photographs. (Like a good matching question not all named species were present, and there could have been a species or two present in the photographs that was not listed. Good Luck.)

Ottawa County, Oklahoma. April.

Beginning and end of plant succession in the mesic-limestone, mixed hardwood forest of the Ozark Plateau. The following short section showed the beginning (pioneer stage) and ending (climax forest range vegetation) of a sere of mesic-limestone forest on a stream floodplain in the western Ozark (Springfield Plateau) Mountains. The sample of this range cover type was the same location and tract of forest treated immediately above. This was a continuation of the north-slope, limestone bluffs drainage of Modoc Creek, Ottawa County, Oklahoma.

238. Started on a rock bar- Two "photo-transects" of the exterior of a mesic-limestone forest that developed along a typical Ozark Plateau stream (Modoc Creek). These photographs were centered on the currently dry shallow creek (half on the stream bed and half on the stream bank) and extended upward taking in lower portions of a limestone bluff. The first furnished a distant view while the second of these two slides was a nearer view that showed details of streambank vegetation.

Most of this stream bank had been been scoured down to bare stream stones and gravel (chert and limestone) following removal of these creek bed materials in the foreground by county road crews (ie. land in the immediate foreground had been used as a borrow pit for roadbed material). This lowered land surface (bar pit) was refilled with more stones, gravel, sand, mud, and other alluvium the next time Modoc Creek flooded. These photographs were in taken in mid-spring two years after flood waters restored the creek bed and low stream bank (start of third warm-growing season following last flooding and stream channel restoration).

Vegetation that pioneered the fresh rubble of the stream channel included both annual and perennial, herbaceous and woody, and native and naturalized (or adventive) species. Sycamore seedlings (more obvious in foreground of the second of these photographs) from the dying adult sycamore (center midground in both photographs) had pioneered the disturbance site. This was the only tree species that had regenerated on this denuded area. There were seedlings of sugar maple in rocks along the creek bank, but these grew mostly under shade (to rear of the parent sycamore) and not on the disturbed part of the channel.. There were only a few individuals of annual pioneer forbs like giant ragweed (Ambrosia trifida). The most abundant (density and cover) annual pioneer was hairy crabgrass (Digitaria sanguinalis). The native festucoid grass, Virginia wildrye, was one of the climax grasses in this forest vegetation and it was the most grass even exceeding hairy crabgrass. Weedy perennial forbs were also better represented than annual species. The more common forbs included the Eurasian curly dock (Rumex crispus) and the native, colony forming Pennsylvania smartweed or pinkweed (Polygonum pensylvanicum) plus big blue strar (Amnosia tabernaemontana).. Woody vine on the dying sycamore was trumpet creeper (Campsis radicans). There were many seedlings of this linana.that had established on both bed and bank of the stream channel (near base of sycamore).

In exterior (physiogonomic) view of the mesic-limestone forest in the first of these slides the tallest tree (big crown in upper left corner) was northern red oak. The crown to left of the tall red oak was basswood. The next crown to the left was of chinquapin oak. All of the shorter trees and saplings (in front of the tall northern red oak and to right of the dying sycamore) with densely leaved crowns at outer edge of this stand were sugar maple. Details of this forest understorey, including grasses and forbs on the limestone bluffs was treated above.

Stream channel of Modoc Creek, Ottawa County, Oklahoma. May (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple-bitternut hickory forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

239. Later in the summer on the same rock bar- Same rock-gravel stream bed and bank of Modoc Creek described immediately above. These two photographs were taken two months later as follow-up "photo-plots" to the preceding two transect views of this pioneer stage of a climax sugar maple-bitternut hickory-white ash-northern red oak-chinquapin oak forest, a north-slope, mesic-limestone mixed hardwood forest in the Springfield Plateau portion of the Ozark Plateau Region. This seral vegetation was developing on newly laid-down, channel material (limestone and chert) in early summer of the third growing season following deposition.

The species composition was obviously the same as that of two months earlier (see above caption) except that some cool-season species (eg. curly dock, Virginia wildrye) had entered summer dormancy while warm-season species (giant ragweed, hairy crabgrass) were still growing toward full maturity and sexual reproduction. A single plant of brown-eyed Susan was blooming (at base of dying sycamore and not readily visible).

The most conspicuous feature of this range vegetation was growth of sycamore seedlings which were now the tallest plants on the gravel bank of Modoc Creek. The trumpet creeper on trunk of dying sycamore was in bloom, and its seedlings around its trellis had also grown considerably in the two months since first "photo-plots" were recorded.

Tree crown outline of the forest in background (second of these photographs) were from tallest crown at right margin moving leftward: northern red oak, basswood, chinquapin oak, and dying sycamore. Shorter trees in front (many were saplings) with crowns of especially dense leaves were all sugar maple.

240. Followup visit to the rock bar- Development of mesic-limestone forest vegetation on the same rock-gravel stream bed and bank of Modoc Creek introduced above except in the subsequent summer. after the above. This was still the pioneer (or, perhaps, second seral) stage on a sere of climax sugar maple-bitternut hickory-white ash-northern red oak-chinquapin oak forest. The sycamore had advanced to young sapling class, but this pioneer tree species was now joined visibly with seedlings and small saplings of American elm, white ash, and bitternut or pignut hickory. Elm had the apparent greatest cover of these three three species all of which are components of the climax forest.

The alluvium (stones, gravel, coarser sand, alluvial soil particles) that had been laid down three years earlier by flood waters still supported populations of "new land" pioneer annuals including giant ragweed and hairy crabgrass that had been so conspicuous one year earlier. In addition, common cocklebur (Xanthium strumarium) had joined the list of pioneer plants. Also noticed for the first time was bracted plantain (Plantago major), a naturalized Eurasian annual forb. It was on the more recently laid-down alluvial material in the stream bed itself. Curly dock was still present on the now three-year-old alluvium, but it was being overshaded (and, consequently, losing vigor, cover, and density. The population of Pennsylvania smartweed or pinkweed was (pretty much) stable while the perennial forb, big blue strar, had increased in cover and density. Apparently most of the many seedlings of trumpet creeper that had established last year were larger and thicker "than ever". Poison oak/ivy had noticably joined previous occupiants on the channel bed and bank.

Most telling succession-wise was migration of broadleaf woodoats down from higher up (older alluvium) on the shaded channel bank down onto the three-year-old alluvium where it joined Virginia wildrye, the other native, climax grass of this forest range site, that had established a year earlier. The introduced pasture grass, tall fescue (Festuca arundinacea) had also joined the cast of grasses, but it was moving in from the outer or sunlite edge whereas the two native grasses were dispersing and migrating in from the shady or forest side.

Stream channel of Modoc Creek, Ottawa County, Oklahoma. Early July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple-bitternut hickory forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

241. Climax of the mesic-limestone, mixed hardwood forest- Deep inside a bottomland (floodplain of Modoc Creek) in western Ozark [Springfield] Plateau) forest of white ash, bitternut hickory, sugar maple, sycamore, northern red oak, black walnut, and honey locust. The second=growth forest vegetation in this photograph developed on a meander of an Ozark stream. Large inward (leftward) leaning tree at far right was sycamore as was end of limb in upper right corner. The other left-leaning trunk that ran from lower right corner to top center was a white ash, the trunk of which was covered with Virginia creeper. Sapling in center foreground and seedlings in immediate foreground were all bitternut or pignut hickory (Carya cordiformis). The sapling above this lower layer of regenerated bitternut hickory with sunlit-bright leaves leaning from the extreme left margin rightward to center of photograph was also a bitternut hickory. The cluster of young well-lit young leaders and leaves were resprounts when the crown of this young hickory had been sheared "clean-off" by a large limb of the whte ash that was brought crashing down in an ice storm. The smaller (but now taller) sapling to left of the ice-topped hickory was northern red oak, the species that was second to bitternut hickory in reproduction. Twigs/leaves of black walnut in upper left corner.

White ash, bitternut hickory, and northern red oak are climax dominants of this forest site. Black walnut sycamore are pioneer (usually Intolerant species, especially walnut) species that live long enough to persist into the climax (at least subclimax) forest. There were no sugar maple on this creek overflow parcel of bottomland in the meander of Modoc Creek. Instead sugar maple was limited to the north slope of limestone bluffs above the creek where it was the dominant tree species.

Regeneration of bitternut hickory and northern red oak was so dense that there was almost no herbaceous growth other than scattered silky wildrye and broadleaf woodoats. Shrub cover were also very sparse and limited mostly to American bladdernut.

On a meander of Modoc Creek, Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple-bitternut hickory forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

A few of the characteristic and the unique species of the sugar maple-bitternut hickory forest were included below. Most of the species growing in the general oak-hickory forest (including sugar maple and bottomland forest cover types) that were included in Range Types of North America were presented in the chapter entitled Oak-Hickory Forest-I. The phyto-characters shown immediately below were placed here to whet the student's appetite for more range plants later on.

242. Leaves and fruit of sugar maple (Acer saccharum)- Details of leaves and nearly mature schizocarp of sugar maple. The fruit of maple has been interpreted as either a samaroid schizocarp having two winged mericarps or as two samaras joined together (Smith, 1977, p. 165). Schizocarp is a dry fruit with carpels separated from each other into single-seeded indehiscent segments called mericarps, which in Acer species are winged; samara is also a dry fruit and a winged one that is indehiscent with a single-seed (Smith, 1977, ps. 66, 307).

Ottawa County, Oklahoma. August. Stage of phenology: premature but late fruit stage (immediate or near-term maturity of fruit).

243. A real bundle- Cluster of schizocarps of sugar maple in mid-summer. This photograph presented better depth-of-field for abit more detail than that afforded immediately above. Ottawa County, Oklahoma. Late July.

244. Fall colors- Leaves of sugar maple in autumn coloration. This was near peak color (brightness; most reddish, yellowish, or orangish) for what is typical of sugar maple in the Ozark Highlands. Usually the colors of sugar maple in one of the most western (and marginal) extremities of the biological range of this mesic species are less brilliant and extreme than those in more eastern (especially northeastern) portions of this species' range.

Ottawa County, Oklahoma. October. Leaf fall was immanent.

245. Clusters of fruits in box elder (Acer negundo)- The fruit of maple species has been regarded as either a samaroid schizocarp consisting of two winged mericarps or as two samaras joined at their apexes. Throughout much of the Ozark Plateau Region box elder blooms and sets fruit earlier than sugar maple with which it is sometimes associated on bottomland forests. Box elder is a favored feed of beaver whereas this largest rodent in North American seems to avoid sugar maple.

Along Lost Creek, Ottawa County, Oklahoma. April.

246. American bladdernut (Staphyla trifolia) in understorey- One of the dominant shrubs in a sugar maple-bitternut hickory forest that developed on a north slope of a limestone bluff above Modoc Creek. American bladdernut was the sole shrub species of a lower shrub layer (vegetational stratum) on a lower elevational zone that extended from stream bank upslope to the point where there was a higher or upper zone consisting of one major shrub layer (dominated by eastern dogwood with eastern redbud as the associate shrub). There was also a taller shrub stratum in the lower zone (near the stream) that was made up almost largely of pawpaw. Pawpaw and bladdernut did not "mingle" or the layers they dominated overlap to any appreciable extent. Rather these two shrub species were largely segregated, although this was certainly not entirely the case.

American bladdernut is one of the more common shrubs in more mesic forests in the Ozark Plateau, especially along watercourses and bases of bluffs.

Bluffs above Modoc Creek, Ottawa County, Oklahoma. October.

247. New flowers and old fruit- A leader of American bladdernut retained one of last year's fruits while new inflorescences were in full-bloom during early spring along a north-facing bluff in the Springfield Plateau of the Ozark Hihglands. This was a dominant shrub in the climax sugar maple-bitternut hickory forest of which it was part.

Bluffs above Modoc Creek, Ottawa County, Oklahoma. April.

248. Flowers of American bladdernut- Flowers of bladdernut grown in clusters that hang down in a racmeme-like arrangement. These inflorescences often occur on tips of smaller branches off of the main limb of the shoot.

Bluffs above Modoc Creek, Ottawa County, Oklahoma. April.

249. Nuts! Bladdernuts- The fruit of American bladdernut is a inflated or bladder-like pod consisting of three subdivisions or compartments. Bladdernut is one of two other woody plant species found in the Ozark Highlands that are in the Celastraceae, staff-tree family. Burning -bush or eastern yahoo (Euonymus atropurpureus) and bittersweet (Celastrus scandens) are these other two species, both of which are distinctive or even unusual and niether of which is common.

Bluffs above Modoc Creek, Ottawa County, Oklahoma. October.

250. Rolling in brome- Local stand of hairy wood brome (Bromus purgans= B. pubescens) on an upland, black oak-dominated forest that had a surface wild fire in spring of the preceding spring (ie. this stand was in the second growing season following a a spring wild fire). This upland forest was just above the limestone bluffs along Modoc Creek which was the forest site of the a mesic-limestone mixed hardwood (sugar maple-bitternut hickory-white ash-basswood-northern red oak-chinquapin oak) forest that was featured in this section.

Hairy wood brome, which was shown as Canada brome in Barkworth et al., 2007, p. 220), is distributed sporadically throughout eastern North America ranging from central Manitoba across to and then south to Florida and westward to central Texas. B. pubescens has an interupted species (biological) range with local occurrence in in Arizona and in Colorado and Wyoming. Incidentially, the common name of Canada brome seemed to this author to be unwarrented and clearly inferior to hairy or hairy wood brome given the specifici epithet, "pubescens", rather than a commenerative one featuring Canada. The common habitat of hairy wood brome is forests and woodlands.

Ottawa County, Oklahoma. Late May; peaking standing crop of herbage, anthesis stages of phenology.

251. Beauty of a native brome- Characteristic nodding, spreading panicle (Barkworth et al., 2007, p. 220) of hairy wood or Canada brome. Anthesis to milk stage of phenology. Bluffs above Modoc Creek in western Ozark (Springfield) Plateau. Ottawa County, Oklahoma. Late May.

252. Hairy spikelets- Spikelets of hairy wood brome in anthesis. These units of the panicle were produced on plants growing in the brome stand on a burned-off (spring wild fire) upland forest dominated by black oak shown above.

Above bluffs of an Ozark Plateau stream (Modoc Creek)Ottawa County, Oklahoma. Late May; peaking standing crop of herbage, nthesis stages of phenology.

Sagging under a heavy yield- Panicle of hairy woodland brome

Ottawa County, Oklahoma. Mid-July; grain-ripe just before grain-shatter stage of phenology.

Full spikelets- Spikelets of hairy woodland brome on an Ozark Plateau upland, chert forest dominanted by black oak and with pignut or biternut hickory, as the associate tree species. Almost every floret had developed fully matured grain on this forest range that had been burnt by wildfire several years earlier.

Sexual reproduction of this native perennial had been very effective in establishing scattered populations of hairy woodland brome throughout the second-growth forest that had developed to an old-growth state climax chert forest.

Ottawa County, Oklahoma. Mid-July; grain-ripe just before grain-shatter stage of phenology.

Study in spikelets- Ripe spikelets of hairy woodland brome produed on a chert upland, black oak-dominated forest with bitternut or pignut hickory hickory as the associate tree species. The pubescent feature of this species, one scientific name of which incorporates this pbescence as Bromus pubescense, was visible in these macrolens views.

Ottawa County, Oklahoma. Mid-July; grain-ripe just before grain-shatter stage of phenology.

253. Shoot! Look at the shoots!- Culm details of shoots of hairy wood brome. Although organs of some shoots of this species are glabrous (Barkworth et al.(2007, ps. 220-221) they typically have a characteristic pubescence (hence the inclusion of "hairy" in common name and "pubescens" as specific epithet) as shown in these examples from the western Ozark Plateau. Another key characteristic of culms in this species is the prominently swollen nodes.

Bluffs above Modoc Creek, Ottawa County, Oklahoma. Late May (peak standing crop, anthesis phenological stage).

254. Woodland neighbor- Woodland bluegrass (Poa sylvestris) growing on and among limestone bluffs in western Springfield Plateau (of Ozark Plateau) in a sugar maple, bitternut hickory-white ash-basswood mesic-limestone, climax forest. Sugar maple seedlings were conspicuous. This photograph can be compared back to photographs (above) that featured the understorey of this remarkable forest range community.

Above Modoc Creek, Oklahoma. May.

255. Panicle in the bluff- Panicle of woodland bluegrass growing on limestone bluffs in western Ozark Plateau. Above Modoc Creek, Ottawa County, Oklahoma. May.

256. Woody spikelets- Details (as best these small ones can be shown in deep woods) of spikelets of woodland bluegrass. Bluffs above Modoc Creek, Ottawa County, Oklahoma. May, anthesis.

Organization Note: a black oak-bitternut hickory forest that had the species composition and structure of climax vegetation was shown and discussed below in the Use and Abuse of Oak-Hickory Forest portion of Oak-Hickory Forests-II. That forest was immediately back from bluff along Modoc Creek and contiguous with the preceding tract of forest that was dominated by sugar maple (with bitternut hickory, white ash, hackberry, northern red oak basswood, and American elm as associate species). The black oak-bitternut hickory forest had burnt in late March or early April just six to eight weeks before a series of photographs was taken recording forest response to the fire. It seemed more appropriate to deal with the recently burned black oak-bitternut hickory forest in the later portion of this chaper. The north slope, bluff forest described immediately above did not burn. This was an example of the relatively greater degree of isolation from disturbances afforded by more moist, north slopes.

The next series of photographs and their captions was of another sugar maple-dominated forest also along Modoc Creek in the Springfield Plateau section of the Ozark Highlands. This forest tract was an east slope forest located less than one mile downstream from the sugar maple-dominated forest just described. The east slope sugar maple forest had fewer species--except for ferns--overall. Most conspicuous was white ash (versus bitternut hickory) as the most frequent associate tree species followed by northern red oak and chinquapin oak. Basswood, bitternut hickory, and American elm were infrequent and did not approach associate species status even in local stands.

This east slope, limestone bluff, sugar maple forest--another variant of mesic-limestone forest (Nelson, 2005, ps. 122-125)-- was postclimax in the general Ozark Plateau oak-hickory forest. The sugar maple-dominated forest described in the following portion was contiguous with a mesic bottomland forest (Nelson, 2005, ps.147-150), except for the separation by stream corridor of Modoc Creek, that was described later in this section on Miscellaneous Forest Types.

Just outward and upward from where sugar maple was dominant:

Where the oaks and hickories took over- In contact with the sugar maple-basswood-white ash north-slope forest on creek bluffs that was described above there was a less mesic forest dominated by variously by black oak, northern red oak, and pignut or bitternut hickory with an herbaceous understorey of ticky tick-clover or pointed-leaf tick-trefoil and hairy woodland brome with various forbs, especially the mint-family member, heart-leaf skullcap (Scutellaria ovata). There were two lower woody layers to this slighter-higher elevation forest the upper one of which was characterized by flowering dogwood (Cornus florida), eastern redbud (Cercis canadensis), and serviceberry or shadbush (Amelanchier canadensis) with some small saplings of the dominant black oak, northern red oak, and bitternut hickory along with a few saplings of American elm and even sugar maple. A lower woody layer also had some sugar maple and American elm of seedling age/size as well as those of the dominant two dominant oaks and bitternut hickory.

Forest vegetation in the first of these two slides included one young adult of black oak (leftmost foreground tree; largest trunk in image), two young adult northern red oak (right-center tree and one to its right in background), a sapling of basswood or American linden (to immediate right of the center northern red oak), post oak (Quercus stellata; one background tree with burls on trunk) and bitternut or pignut hickory (rest of background trees). Seedling in lower right-hand corner was pignut or bitternut hickory. Other understorey species included those named in the preceding paragraph.

Forest plantspecies in the second slide featured a large, stately, adult Carya cordiformis, pignut or bitternut hickory. Adult background trees included more pignut hickory, black oak, northern red oak along with a few sugar maple and American elm. The lower forest layers were as described in the first paragraph of this caption, including various age and size classes of all these tree species along with sticky tick-trefoil, hairy woodland brome, and heart-leaf skullcap.

Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

257. East slope sugar maple forest- Another example or phase of sugar maple-dominated forest in the Ozark Plateau was also found along Modoc Creek. This forest tract had developed on an east slope and differed from the sugar maple-bitternut hickory forest type or subtype (variant) that developed on a north slope of Modoc Creek located only 3/4ths to one mile upstream from the forest vegetation described here. This sugar maple-dominated community was much less species-rich except that it had more ferns including maidenhair (Adianthum pedatum) and Christmas ferns (Polystichum acrostichoides). In this forest communitysugar maple was sole dominant with northern red oak and chinquapin oak (Quercus muehlembergia), and white ash being local associate species. Mature trees of these two oak species were generally larger than all but a few of the slower-growing sugar maples.

There were also larger, less abundant trees of sycamore, bitternut hickory, and black walnut. Most regeneration was of sugar maple with somewhat less reproduction in box elder and northern red oak with these two species swapping places depending on microhabitat. Co-dominant shrubs overall (and of two different layers)were American bladdernut (taller) and spicebush (shorter-growing). The tallest shrubs (those of the taller shrub and/or lower tree layer) were flowering dogwood (more abundant) and eastern redbud. Pawpaw was present, but was primarily on the adjacent (and other side of the creek hackberry-American elm-sycamore-eastern cottonwood bottomland forest). Lanceleaf buckthorn (Rhamnus lanceolata) grew in association with American bladdernut, and though usually overtopping the latter, R.lanceolata was uncommon. Almost no herbaceous species other than ferns. Occasional plants of Virginia or silky wild-ryes or beakgrain (Diarrhena americana) were present, but these-like woodreed grass (Cinna arundinacea)-were usually confined to rock or gravel bars away from the sugar maple forest. Grasses were much less common than on an adjacent hackberry-elm-box elder forest on the stream floodplain (described below).

The two wide-view photographs shown here presented species composition and structure of a postclimax sugar maple forest that developed on an east-facing limestone bluff along a typical stream in the Ozark Plateau. In the first of these photographs a large hackberry (largest trunk; dead center of photograph) and black walnut (tree to right of big hackberry) added some diversity to an otherwise solid stand of uneven-aged sugar maple. In the second of these slides an immense sugar maple (25 inch DBH) overlooks a nearly "pure" stand of its species (undoubtedly many of which were its own offspring). Even smaller adult trees at right were mostly sugar maple. Lanceleaf buckthorn (Rhamnus lanceolata), an exmple of which was tallest shrub in front (and slight to right) of large sugar maple, American bladdernut, flowering dogwood (conspicuous in right foreground), and American hazlenut (Corylus americana) were the primarily shrubs vbisible in the second photograph.

Along Modoc Creek, Ottawa County, Oklahoma. July (aestival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

258. Broad view of an Ozark Plateau bluff and its forest- A wide-angle "photo-transect" on an east slope of a limestone bluff along a typical stream in the western part of the Ozark Highlands on which a postclimax sugar maple-dominated forest (with white ash and, to lesser extent, northrn red oak were associate tree species) had developed. This was a mesic-limestone forest (Nelson, 2005, ps. 122-125) that in monoclimax theory (Clements, 1916) was postclimax to the regional (climatic) climax oak-hickory forest. This photograph introduced forest range vegetation that was viewed at increasingly closer-in scale in subsequent slides.The huge sugar maple at the right served as a "landmark" and featured topic in the subsequent slides. The smaller tree in left midground with the right angle-like trunk leaning sharply to the left was another sugar maple. The tree in upper left margin (upper right corner) was a northrn red oak.

Shrub species included American bladdernut (generally the dominant shrub), lanceleaf buckthorn, flowering dogwood, shadbush or, as it is also known, eastern serviceberry, and eastern redbud. There were only occasional herbaceous species that were not forbs, most of which were ferns, as most grasses and sedges grew closer to the creek. Along banks of the stream (Modoc Creek) there were occasional individuals of silky or Virginia wildryes, beakgrain, or giant woodreed. Most common ferns were Christmas fern and maidenhair fern.

Regeneration of climax dominant trees (mostly sugar maple, white ash, and northern red oak) comprised much of the lower layers of the understorey as the young of these trees ranged from seedlings to large saplings.

259. Postclimax and "pure"- An east slope of a limstone bluff above an Ozark Plateau stream provided the setting for a "nearly pure" stand of sugar maple. In the Clementsian monoclimax theory this was a consociation of Acer saccharum that was postclimax vegetation where the climatic =regional or zone) climax was oak-hickory forest, the Oak-Hickory Forest Region (an association) of Braun (1950, ps. 35, 162-191). The mighty sugar maple in center foreground was the sugar maple described as "huge" in the immediately preceding photograph. This large maple had a large (by standards of its species) lanceleaf buckthorn to the right and in front of it. Shrub in lower right corner was flowering dogwood. Shrub species in left foreground included American bladdernut, pawpaw, and flowering dogwood. Many of the shrubs higher up on the bluff were shadbush or, as is also known, eastern serviceberry. Almost all tree regeneration was sugar maple with traces of northern red oak and white ash.

Details of understorey of this immediate (local) site was given in the next slide and caption.

260. On the bank- On the bank of a typical stream in the Ozark Highlands there was a multi-layer understorey in a sugar maple-dominated forest on an east slope of a limestone bluff. Tree trunk in center midground was that of the mammoth sugar maple that served as the "landmark" in the immediately preceding slide. Shrub to right (and with its upper shoot growing to the right)was lanceleaf buckthorn. Most of the other shrub cover was that of American bladdernut. Ferns in edge of stream bank were Christmas fern, a major forb in this mesic-limestone, postclimax forest. Also present was a ground layer composed of luxuriant cover of moss, the species of which this bryophyte-ignorant author knew not.

261. A bluff full of beauties- An east-facing limestone bluff aligned along Modoc Creek in the western part (Springfield Plateau) of the Ozark Highlands supported a mesic forest dominated by sugar maple (with white ash and, secondly, northern red oak as associate tree speceis). This forest range was postclimax in the general oak-hickory forest of the Ozark Mountains.

This photograph featured species composition (especially of dominants and associates) and structure of this southern and western outlier of the sugar maple forest cover type. In this stand white ash was the clear associate to sugar maple, but with northern red oak a close "runner-up". Large tree at right midground (and midway up the bluff) was sugar maple. The large tree to the right of this sugar maple (along right margin of photograph) was northrn red oak. The adult tree closest to the big sugar maple (left of it with its upper bole in the crown by large left limb of the maple) was white ash as was the largest tree in this photograph which was largely obscured by two smaller hackberry trees. The largest tree, the white ash, had lost much of its crown which retained a large left-leaning limb.

Major shrub was American bladdernut, but flowering dogwood, American hazelnut, shadbush, and eastern redbud were present to locally common.

262. Sugar maple-white ash stand- Composite view of species composition and structure of a mesic-limestone (east slope; bluff habitat) with sugar maple the dominant and white ash the associate species. Biggest tree (right side) was sugar maple. Second largest tree ((left of center midground) was white ash. Most regeneration was sugar maple and white ash, including the larger saplings in baqckground. Hackberry at far left. There were two shrub layers: 1) taller layer dominated by flowering dogwood, lanceleaf buckthorn, and American hazelnut (the large shrub in front of the big sugar maple) and 2) lower layer almost exclusively American bladdernut but with considerable poison oak/ivy. Grape vines grew to top of canopy and formed a "unifying wooden thread" among the various layers of forest range vegetation.

Herbaceous layer(s) consisted primarily of Christmas and maidenhair ferns. Mosses made up a lush ground layer (the lowest layer of vascular plants).

The black circular "spot" in upper left was entrance to a small cave in the limestone bluffs. This is a common feature of the ancient Ozark Mountains.

263. Hole in the canopy- Example of gap or patch dynamics in a sugar maple-dominated east slope bluff forest in Ozark Plateau. A postclimax forest range in the general oak-hickory forest had extensive feeding by beaver (Castor canadensis) 17 years prior to this follow-up photographic session. Beaver killed a high percentage of the largest northern red oak (along with lesser browsing on American elm, box elder, sycamore), but fed on very little sugar maple. This gap was created 17 years earlier when beaver girdled three massive northern red oak (all were between two and a half and three feet DBH) which died in the spring following winter browsing a few months earlier (the three oaks never really leafed-out). The light-colored trunk in right background was the snag of the only one of the three northern red oaks that had not fallen.

Death of the three giant oaks left such a gap in the canopy that the stage was quickly set for the process of recovery of the forest vegetation through secondary plant succession. Such vegetation dynamics--to use the term made famous by F.E. Clements who adopted it from H.C. Cowles--is studied as patch or gap dynamics which is unique from the perspective of small spatial scalet as in, say, the larger spatial of a large forest clearcut, blowdown, fire, or old-fields as in cut-over lands.

Most tree regeneration was of sugar maple (including the sugar maple sapling in lower right foreground) and white ash. The large limb, which was shown at close range in the next (succceding) slide, was off of a large northern red oak (outside of camer range at left) brought down by a February icestorm. The falling red oak limb brought down canopies of two medium-sized black walnut trees (left midground). Beneficiaries of this tree damage were understorey plants, espceially herbaceous species the most common of which were maidenhair fern and the pioneer composite, giant ragweed (Ambrosia trifolia). American bladdernut was the most common shrub in the gap.

264. Ground level and ground zero- Lower and mid levels of forest range vegetation in a seventeen-year old gap in a sugar maple-dominated forest that was made when beaver girdled three massive northern red oak. In the 17 years following beaver-felling of old-growth trees the "dynamics of vegetation"--to borrow a title for selected writings of F.E. Clements (Allred and Clements, 1949)--had resulted in re-establishment of sugar maple and white ash, the two major (dominant) climax tree species for this forest site. Of course trees of these two climax domiants were still small, but the species composition had quickly progressed to that of the climax forest vegetation. The dominant shrub was American bladdernut which, while not as abundant as on the banks of the stream (Modoc Creek), still had appreciable cover and density.

The largest and one of the most common herbaceous species was giant ragweed. It was not known whether this pioneering annual composite was more plentiful due to (or even present only after) disturbance due to tree damage from the February icestorm .There was very occasional cover of Virginia wildrye, but in more shaded local habitats the most common forb was maidenhair fern.

The sugar maple sapling (right-of-center foreground), that became conspicuous within a couple or three years following death of three massive northern red oaks by beaver girdling, had in the interim become well-established and received limited beaver feeding (note blaze on lower trunk). Obviously beaver abandoned this feeding station quickly and before the sugar maple sustained enough feeding damage to be life-threatening, at least in the short-run. (Reduced performance or even death might be possible via disease entry through the debarking wound where beaver fed, however limited that was.)

265. Life among the fallern on the forest floor- A severe February icestorm brought down a large part of the crown of a northern red oak (which in a chain reaction brought down crowns of neighboring black walnut as shown in a preceding photograph). About six months later, as shown in this photograph, maidenhair fern and regenerated (seedling and/or root-sprouting) sugar maple were apparently benefiting from the ecological windfall caused by ice.

The following series of eight slides and six captions showed the initial stage (denudation) and subsequent early (pioneer) stages of the gap created by beaver browsing on (and quickly killing) three northern red oaks on an east slope (of a bluff) sugar maple-white ash climax forest that was described immediately above. This was the same forest gap or patch featured in the three immediately preceding photographs. Denudation (induced by defoliation, specifically bark feeding, by beaver) had been initiated 17 years before the time at which the three preceding photographs were taken. Several subsequently taken photographs revealed dynamics and development of forest range vegetation (forest recovery or revegetation) through secondary plant succession at periodic intervals. (The sylvan stage play ran backward to show viewers "the storey so far": forest succession on a Ozark Plateau bluff.).

266. A lingering death and the tragic end of their reign- The first act of the unfolding forest drama (tragedy or comedy was left up to viewers' discretion) that was revealed above began with the empty stomach and ever-growing incisors of beaver(s). Bark-feeding by beaver during winter months left three very large northern red oaks (each over two and a half foot DBH) girdled and soon-to-be-dead in an east slope, mesic-limestone, sugar maple-white ash climax forest. The tree in left background as well as the trunk in midground, and partially concealed by the foremost tree, were all northern red oaks and completely girdled. Death would "officially" wait until spring when, with rising sap, these once-magnificant oaks would fail to leaf-out. (As it turned out, the preformed, terminal buds of these oaks did not fully open before they fell off indicating that trees were dead.) None of the three trees sent up any stump sprouts. Lack of suckering (production of basal shoots) is the typical or normal physiological response of old hardwood trees. Death of the entire tree (not just top-kill) was "sure and swift" for all three mature "timber-ripe" oaks.

These three northern red oak were some of the largest trees in this east slope, limestone bluffs forest. Only a few sugar maple and white ash reached similar size. Trees of these latter two species were not browsed. This was a textbook example of feeding selectivity (= selective browsing). Locally these three oaks were (had been) "far and away" the largest trees that controlled the largest portion of the canopy (ie. had the greatest impact on how much and for how long light reached lower levels of the forest). Loss of the "masters" of the forest canopy set the stage for secondary plant succession on the bluffs above Modoc Creek.

Oange inner bark is characteristic of several of the red oak species (Erythrobalanus subgenus) , especially black oak, and it showed prominently on the three girdled northern red oaks featured here. "Reading sign" revealed that beaver gnawed higher (almost three feet above ground level) on the upslope side of the trunks showing these rodents (largest rodent species in North America) fed on their hind legs on the high side of trunks while "standing on all fours" on the downhill side.

Tthe author's 38-inch, hickory walking stick was placed lengthwise at base of the foremost (and largest) oak for scale.

Historically there have been high population densities of beaver along Modoc Creek and other streams in this local area. As a boy the photographer's younger brother trapped beaver for several years along these streams. He caught a few adult male beaver that exceeded 50 pounds in weight. These were (are) bank-beaver. The beaver do occasionally build high dams on streams causing localized flooding of bottomland forests, but as far back as local observers can remember beaver never built stick-and-mud lodges. Rather, long tunnels dug far back into creek banks served as beaver dens.

As was shown in this photograph adult beaver often fed at considerable distances from streams. This was never much more than half-way to top of bluffs (usually feeding was limited to approximately the lower one-third of bluff height). In other words, trees that grew high up on bluffs were safe from beaver browsing or, from a forester's perspective, beaver depredation.

Above Modoc Creek, Ottawa County, Oklahoma. February, 1990.

267. Standing tall, proud, and dead- On east-facing, limestone bluffs above Modoc Creek the tallest and largest-diameter (over 30 inches DBH) of three northern red oaks was quickly dying (after only partially leafing out) following bark-feeding by beaver during the immediately preceding winter months. Death of the three northern red oaks was complete in a month or month and a half following bud-opening. Most of the neighboring (surrounding) trees--all of which were considerably smaller and, presumedly, younger--were sugar maple and white ash plus a few black walnut.

Death of the largest--and locally dominant--northern red oaks created a forest gap or patch almost as rapidly as windthrow (blowdown) or lightening strike. Beaver were not only hungry and "eager" they were lethal. The stage was preparing for the second act of patch or gap dynamics.

Above Modoc Creek, Ottawa County, Oklahoma. August, 1991.

268. The new living among the old dead- Two local "photo-quadrants" showed details of and adjacent to the largest of three northern red oaks that died after being girdled by bark-feeding beaver over a period of roughly two to four months prior to an aborted effort by oaks to leaf-out in spring. The first fact to "leaf-out" from the "crime scene" at this photographer was that the large oak (three foot DBH) was largely hollow. The trunk of this otherwise healty northern red oak (prior to beaver strike) consisted almost solely of sapwood. This condition of "hollow-heart" (by which it is known among Ozark sawmill operators) probably had little, if any, influence on this tree because 1) heartwood is dead and provides no life-supporting function and 2) a hollow tree, like many hollow cylinderical things in nature (grass culm, bird quill) is almost as strong as trees with solid (non-hollow) trunks. Therefore, beaver did not bring down a tree that was dying (they killed a sound, healthy tree). On the other hand, beaver did not destroy a valuable lumber tree and cause economic loss to the firm or landowner. The huge tree was useless for lumber and was fit only for fuelwood (the trunk would not have made a saw log and loggers would have wasted their time felling the tree). In fact, beaver conducted their own version of "site preparation" by clearing the way (making resources available and conditions more favorable) for younger trees that could grow sound boles for future (and more valuable) wood products.

Let's here it for the beaver! Plus, the browse (bark) of the oaks contributed to the production of more beaver plew (pelts), another product of the forest. Why, it might even stir that younger brother to re-run his trap line (at least renew his subscription to Fur, Fish and Game).

The hollow-trunk northern red oak was an example of "over-ripe" timber known technically as overmaturity or overmature which when applied to individual trees refers to the condition of having reached the "...stage of development when it is declining in vigro and health and reaching the end of its natural life span" or "...one that has begun to lessen in commercial value because of size, age, decay, or other factors" (Helms, 1998).

It was possible (probably likely) that once it was dead, and with weakened and soon-rotting sapwood, the hollow oak crashed sooner than if it had been solid. In point of fact, one of the three girdled northern red oaks was still standing 17 years following its death (the snag standing conspicuously on the bluff in two of the photographs presented above). The solidness of this trunk was not determined by the author because cutting into the tree to determine its state of soundness could have caused it to fall when it might otherwise have continued to stand for decades. (Besides only a hollow-headed idiot would risk life and limb--catch the pun--to tote a double-bit up such a steep, slippery slope to cut a tree that was useless for wood. Now slinging a 35mm SLR Nikon for educational purposes--that is, to chop out ignorance--is a different grade of lumber.)

The other--and more relevant--fact from forest vegetation and succession standpoints was the tree species that were replacing the former "mighty oak". Saplings and seedlings of sugar maple began almost instantly to fill the gap created by the beaver. Obviously, maple saplings had been growing beneath this northern red oak prior to beaver "attack". Furthermore, sugar maple is rated as Very Tolerant (Wenger, 1984, p. 3) so that continued survival and growth of sugar maple was likely even with continued canopy cover of the potential still-living oak. Nonetheless, more resources and modified conditions (including increased light, space, and soil moisture) undoubtedly was of some benefit to sugar maple growth and regeneration. There were also several age-size classes of white ash (and of northern red oak) in the gap formed by death of the large northern red oak.

The tree behind the fallen oak was one of the other two northern red oaks killed--but still standing--by beaver.

Above Modoc Creek, Ottawa County, Oklahoma. September, 1991.

269. Down and out (and after only one "beaver strike")- The "long view" of the largest of three northern red oak killed by beaver girdling in the preceding winter. This hollow-trunk old patriarch fell across the channel of Modoc Creek creating a forest gap in which already active-growing sugar maple and white ash (even a few young northern red oak) were ready to utilize more available resources and different conditions (not least of which was more light, space, and soil water).

It would be humanly impossible to know all of the ways in which death and crash of this old-growth speciment affected life in this east slope climax forest, or of the creek below it or the bottomland forest (another forest cover type and separate tract of forest) separated from it by the stream. Even the trunk and crown of the oak across Modoc Creek could conceivably have some impact on some organisms. For instance, it could permit some animals to cross to other side of the stream. The potential barrier to movement, which though not exactly a grand canyon, had been breached. This was of no momemt to a coon, but it could be determinative to a biped (such as the hillbilly who took this photograph).

Above Modoc Creek, Ottawa County, Oklahoma. September, 1991.

270. Bridge to a new forest and return of the final order- Two views from opposite ends of the rotting log of a large northern red oak that was killed by bark-feeding beaver and fell in the first summer following its death in early spring. The tall tree in left foreground of first slide was a sugar maple. Much, probably most, of the tree regeneration was also of sugar maple though this was accompanied by that of white ash, northern red oak, bitternut hickory, hackberry, and black walnut. Sycamore, the dominant pioneering or colonizing tree species along stream channels and freshly scoured bottomlands, was not released or encouraged by loss of northern red oak.

Shrub species along this stream bank and the sides (slopes) of a limestone bluff above included American bladdernut, flowering dogwood, lanceleaf buckthorn, pawpaw, eastern redbud, and shadbush. These species were aligned along the slope from stream bank to top of bluff with bladdernut, lanceleaf buckthorn, and pawpaw typically restricted closer to water while shadbush grew highest up on the bluffs farthest from the stream. Flowering dogwood and eastern redbud generally grew all across this elevational gradient.

Understorey herbs ranged from colonizing species like annual giant ragweed and mare's tail (and where did that seed come from?) and the hugh tap-rooted, perennial pokeweed to maidenhair and Christmas ferns.There were a few individuals of the two grass species, woodreed and beakgrain. Mosses grew on rocks, logs, tree trunks, etc.

Above Modoc Creek, Ottawa County, Oklahoma. July,1997.

271. The young beneficiary (an immediate heir after assination)- A small pole of white ash was the largest tree above sapling size closest to the beaver-killed (and soon crashed) northern red oak described above on the east slope of a bluff above Modoc Creek that supported a climax sugar maple-white ash forest. Sugar maple (sole climax dominant), northern red oak, bitternut hickory, hackberry, and black walnut were generally the most abundant tree species and, not surprising, these species had the most regeneration in the gap created by death of three large northern red oaks. Of these, sugar maple was followed by white ash in general abundance (cover, density, etc.) as the major species to benefit from death of large, mature (actually, overmature) northern red oak. Not only were did sugar maple followed by white ash furnish the largest individual trees and greatest canopy cover, they were also the species which had greatest apparent (quite obvious) reproduction. Ergo, a sugar maple-dominated forest in which white ash was the associate species.

Above Modoc Creek, Ottawa County, Oklahoma. June, 1997.

Concluding observtion with regard to browsing by beaver- On this stream-bordering, bluff forest (an east slope form of mesic-limestone sugar maple forest) feeding by beaver had a major impact on existing forest composition and structure and on future forest development (revegetation). Beaver were a major biotic factor that at local scale were a major cause of denudation and subsequent changes in the forest plant community via secondary plant succession.

Beaver preferentially browsed northern red oak while largely leaving sugar maple and white ash untouched. Defoliation of some of the largest and oldest northern red oak by complete girdling resulted in quick death of some of those individual trees that had greatest control of canopy (forest) cover. Elimination of forest trees with some of the largest canopies produced forest gaps and patch dynamics. The main tree species to benefit from death of northern red oak by beaver and openings in the forest canopy created by such death were sugar maple and white ash, the climax dominant and associate of this forest which was a subtype of the sugar maple cover type, SAF 27 (Eyre, 1980, p.30). Some northern red oak--typically smaller and younger trees--was unbrowsed and remained in this forest as an important (though not an associate) species.

Northern red oak was most likely a subclimax tree species having some large, long-lived (and lucky!) trees that persisted into the climax forest vegetation dominated by sugar maple with white ash as its associate (at least, major associate) species. Selective browsing by beaver hastened dominance by and general increased cover of sugar maple and white ash.

Thus it appeared that sugar maple-dominated forest in the greater oak-hickory forest of the Ozark Highlands were partly a result of zootic influence and not just an edapho-topographic climax. Conversely, there were other tracts of sugar maple forests along this same stream (including a sugar-maple-bitternut hickory subtype of SAF 27) which supported no beaver populations and in which northern red oak was a major constituent but not an associate species.

While beaver ignored or even avoided sugar maple in these forests they readily consumed bark and harvested saplings and smaller branches of box elder, the other Acer species on this tract. Box elder was uncommon on the sugar maple-white ash forest on the east-facing bluff above Modoc Creek, but box elder was locally a major species on the adjacent bottomland forest (just across the channel of Modoc Creek) where it was readily and regularily fed on.

In contrast to browsing selectivity by beaver on a subclimax species (and avoidance of climax tree species) in this forest, a representative tract of SAF 27, beaver had a preference for and browsed selectively on climax American elm and hackberry on a bottomland forest, a representative of cover type SAF 93. These two tracts (and representatives of two forest range types) were contiguous except for separation by the narrow stream channel of Modoc Creek. On the bottomland forest the climax tree species were decreasers under beaver bvrowsing whereas on the adjoining bluff forest (and a different forest cover type) the climax tree species were avoided while beaver exhibited browsing selectivity for an important subclimax tree species which was a decreaser species..

Apparently there has been little evaluation or even recorded observations on the affect of beaver browsing on the development and compostion of forest range communities. Two interesting and extremely detailed studies on impacts of beaver on North American forests were included as chapters 18 (Donkor) and 19 (Smith) in the reference, Plant Disturbance Ecology, edited by Johnson and Miyanishi (2007, ps. 579-671).

Organization note: a climax, bottomland, mixed hardwood that developed on the floodplain of Modoc Creek and that was conterminous with the sugar maple-northern red oak-white ash-pignut hickory-white ash mesic forest on an east slope along bluffs above this creek was treated separately under Southern and Central forests-II.

A thrid series of slides and captions presented a third tract of sugar maple-dominated forest. This was also a third variant of form of the sugar maple forest cover type (SAF 27) in the Ozark Highlands which is the more westerly extent of the oak-hickory forest in general, the Oak-Hickory Forest Region of Barun (1950, ps. 35, 162-191). In monoclimax theory (Clements, 1916) sugar maple-dominated forests are postclimax vegetation in the climax oak-hickory forest region. Viewed from the polyclimax or climax pattern theories proposed by Tansley (1926) and Whittaker (1953), respectively, sugar maple forests are edaphic, topographic, etc. climaxes within the general or more widespread climatic climax (oak-hickory forest). By any of these interpretations sugar maple forests are the potential natural vegetation (Kuchler, 1964).

Just as sugar maple-dominated forests in the general oak-hickory forest are variants of this regional forest (sugar maple forests usually include an oak and/or hickory species as the associate) so too are there variants (variant forms) of the generic sugar maple forest. The first series of photographs and their captions in this sugar maple section of Oak-Hickory Forests-II was devoted to the sugar maple-bitternut hickory variant and the second series of photographs-captions dealt with the sugar maple-white ash variant which were subtype 4 and subtype 3, respectively, of the sugar maple forest cover type, SAF 27, (Eyre, 1980, p. 31). The current (third) series of slides dealt with a sugar maple-northern red oak variant. This subtype was not noted by the Society of American Foresters description (Eyre, 1980, p. 31). Nor was it described by Steyermark (1940) or Braun (1950). Likewise, the SAF description did not include the sugar maple-white oak subtype of Steyermark (1940) and Braun (1950).

It was somewhat surprising that Steyermark (1940) did not identify and describe the sugar maple-northern red oak variant because northern red oak is more mesic in its general habitat requirement than bitternut hickory and, even more so, than white oak. In moisture regimes of these tree species habitats, sugar maple is closer (more mesic) to northern red oak than to white oak or bittrnut hickory. In fact, among the three tracts of sugar maple forests described herein (western edge of Springfield Plateau of Ozark Highlands) northern red oak was the most consistently represented associate species. Nelson (1987, p. 28) listed the first three dominant plants of mesic-limestone forest as (in order): northern red oak, sugar maple, and white oak. In the first two tracts of sugar maple-dominated forest described above northern red oak was of lesser importance than bitternut hickory and white ash, but northern red oak was present at greater cover, density, frequency, etc. in all three tracts than the two associate species (other two species, if northern red oak was also an associate species).

The sugar maple-northern red oak was the least mesic and most exposed of the three forest tracts described herein. This third variant form os sugar maple-dominated forest was a north slope (and limestone) forest, but it was not near a stream and more upland than bottomland forest. It was about two miles (or less) from the second tract (east slope) of a sugar maple-dominated forest, and with white ash as overall associate species.

This third example of a sugar maple-dominated forest, or in this case co-dominated by northern red oak (Quercus rubra= Q. borealis), was on a steep north slope less than 2 miles from the two sugar maple forest stands along bluffs of Modoc Creek. The forest community featured next was a dry-mesic limestone forest (Nelson, 2005). This one was also in Ottawa County, Oklahoma; July, 2007). In strictest sense sugar maple was sole and defining dominant because most regeneration-especially in deeper shade-was of Acer saccharum and not Quercus rubra even though the latter was reproducing on some better-lite microsites (sometimes by established maples). Other species represented by occasional mature (and large) trees included bitternut hickory (Carya cordiformis) and white ash (in roughly equal proportions) along with lesser cover of basswood, all of which are climax species on this cover type and site. At upper (and drier) habitats in this forest chinquapin oak had both mature trees and regeneration. This species was regarded as a local climax member of this forest. Black walnut survived in more open spaces where it probably established in some gap and, having captured its share of canopy, survived to adulthood. At other spots in this forest black walnut succumbed to competition (was shaded out) by sugar maple and northern red oak. Major understorey (sub-canopy) shrubs included both eastern redbud and flowering dogwood throughoutalong with American bladdernut and pawpaw these latter two of which were limited to lower, more mesic habitats.

The lower ground shrub layer was comprised mostly of Virginia creeper (Parthenocissus quinquefolia), but like the common, well-distributed summer grape (Vitis aestivalis), this woody climber ascended into the canopy. Poison oak/poison ivy (Rhus toxicondendron) was present, but was less common than in drier forests. Dominant forbs were tick clovers (Desmodium nudiflorum and D. glutinosum). Forbs that were locally abundant (in a relative sense) included Jack-in-the pulpit, Bloodroot (Sanguinaria canadensis), and wild ginger (Asarum canadense). An uncommon (and uncommonly striking) forb was blue cardinal flower of giant lobelia (Lobelia siphilitica). The only graminoids were unidentified species of Carex (only non-fruiting specimens present).

These photographs were taken at abut 1330 hours Central Standard Time to get maximum direct light, but even then there was too much shade (too many shadows) to be able to catch key features such as bark. Ergo: not many good shots. Will try "next year" (wait until next summer) using with light cloud cover to get diffuse light.

272. Looking in on more sugar maple in the Ozarks- Exterior view of a north slope, mesic sugar maple-northern red oak forest at western extreme of the Ozark Plateau, and of the Oak-Hickory Forest Region (Barun, 1950, ps. 35, 162-191), in which this island of the sugar maple cover type (SAF 27) is postclimax (Clementsian monoclimax theory) or an edaphic or topographic climax (Tansley polyclimax and Whittaker climax pattern theories). The small pole-size trees were sugar maple. The tree species higher upslope (behind maples) included three oaks: chinquapin (chinkapin), northern red, and black oak with the latter more abundant on the drier, higher part of slope.

Shrub species included American bladdernut (the most abundant shrub in understorey shown in this photograph), pawpaw (second most common shrub here), flowering dogwood, and eastern redbud. The latter two shrub species were more abundant in the tract overall just not in the sample of the forest plant community presented here. Virginia creeper formed the bulk of a lower shrub laye,r but along with the less common summer grape, this woody climber ascended into the canopy layer.

Relatively little cover of herbaceous species, but the major forbs two species of ticktrefoil (Desmodium nudiflorum an D. glutinosum). Other forbs that were locally (and relatively) abundant in this forest vegetation were Jack-in-the-pulpit, mbloodroot (Sanguinaria canadensis), and wild ginger (Asarum canadense). An uncommmon (and uncommonly striking) forb wqas blue cardinal flower of giant lobeila (Lobelia siphilitica). The only graminoid observed by the author was an unidentifiable (pre-bloom) caric sedge (Carex sp.?).

Ottawa County, Oklahoma. July (aestival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

273. On a steep north slope- The mesic north slope (with the fertility of a limetone-derived soil) furnished an ideal environment for this sugar maple-northern red oak forest in the Ozark Highlands.Northern red oak was the associate species overall, but local stands were dominated by white ash or bitternut hickory. In this first of these two photographs the four trees in foreground were sugar maple. Trees to left and downslope were white ash. In the second (vertical) photograph four sugar maples were shown at closer distance, but the more critical feature of this view was the understorey which included a diverse array od species including Virginia creeper, tick trefoils, a caric sedge, and seedlings to small saplings of sugar maple, bitternut hickory, and northern red oak. Regeneration of these tree species comprised most of the low and middle shrub layers in these two "photo-plots". Overall sugar maple, the tree with the high tolerance rating of Very Tolerant (Wenger, 1984, p.3), was the primarily reproducing tree species followed closely by bitternut hickory (eg. large seedling in front of and to left of the foremnost sugar maple).

In this forest tract as a whole most reproduction was sugar maple and not northern red oak, white ash, or bitternut hickory. On some local sites (microsites) most of the seedlings and sapling stocking was of associate tree species. Examples of such stocking of younger age-size classes was shown in these and some of the following slides.

There was very little herbaceous growth in this tract (including the "photo-samples" presented here) based on density and relative cover.

274. In the shade of their parents- A large sugar maple--by standards of this forest site in the Ozark Plateau--"looked down" on its progeny (and that of other sugar maples) as this climax dominant tree successfully reproduced in the mature soil and final stage of development of this forest sere. In this climax forest (potential natural vegetation) tree regenertation was primarily of the same species as the dominant trees (ie. same species of trees in the understorey layers as tree species in the canopy). Plant succession had stopped. The sere had reached its final "destination", climax stage. The final plant community for this natural vegetation--termination of plant succession on this forest site (this naturally occurring habitat of north slope, steep, limestone hillside in the Ozark Highlands)-- was complete until some disturbance (eg. fire, icestorm, disease or insect outbreak, logging, blowdown) reset vegetation on this sere back to a earlier state of vegetational development (plant succession).

This was not, however, a "perfect match" of canopy and regeneration age-size classes in lower layers of this range vegetation. This was clearly a stand of sugar maple with all larger (adult) trees, including the one in foreground and those to its left in midground, as well as many larger seedlings and small saplings in foreground being maples. Also in foreground (and growing side-by-side with sugar maples seedlings) were seedlings of bitternut hickory, northern red oak, and chinquapin oak (these three hardwood species were discernable in lower right corner). It was uncertain as to which, if any, seedlings of these latter three species (all of approximate Intermediate tolerance) would grown to adulthood in shade of the mature sugar maples. It was a "good bet" that seedlings of the Very Tolerant sugar maple (Wenger, 1984, p. 3) would survive in the shade of their parent species.

Two species of ticktrefoil were the dominant herbaceous plants (eg. a plant at prebloom stage to left of the center of sugar maple trunk in immediate foreground). Other herbaceous species were not identifiable in this photograph.

275. Layers and niches in climax sugar maple forest- Beneath a large, forked-trunk sugar maples saplings of bitternut hickory (left of the sugar maple) along with pawpaw (eg. shrub with light-colored, crooked trunk to the right and slightly behind the maple), eastern redbud (not discernable in this slide), flowering dogwood (not able to be detected in this photograph), and American bladdernut (shrubs of intermediate-height in lower right foreground) comprised much of the understorey of this climax sugar maple-northern red oak forest. There were actually up to three layers of woody plants in the understorey of this north (and very sttep) slope in the western perimeter of the Ozark Highlands. Pawpaw, flowering dogwood, and eastern redbud made up a high (the taller of two below-canopy) shrub layers while American bladdernut with seedlings and small saplings and ground-level Virginia creeper comprised the lower of these two shrub layers (those that did not extend to the canopy). Summer grape and some (lesser cover) poison oak/ivy and a few woody vines of Virginia creeper climbed into the canopy so as to extend the shrub layer into tops of tree crowns and form a third shrub layer in this forest plant community.

Herbaceous species were sparse in this denser part of the understorey and limited mostly to two species of ticktrefoil.

276. "They have their exits and their entrances" (Shakespeare, As You Like It, II.7)- In a north slope, climax sugar maple-dominated forest it was curtains for a black walnut, but the snag of the old tree was still serving a function in the forest ecosystem. A large summer grape used this snag of a long-dead black walnut as a Nature-provided trellis. The large bole behind and to left of the snag and that of the big bole behind and to right of snag were northern red oak each with a straight, standing log of clear lumber on the stump. Northern red oak produces some of the finest, clearest, straight-grained wood available for durable, attractive lumber that is commonly used in church furniture. This is a most appropriate use of this renewable natural resource especially in the Ozark Highlands section of the Bible Belt where hillfolk praise God and good timber. Of course there are heathens everywhere, but fortunately the owner of this timber land appreciated the aesthetic value afforded by the beauty of natural forest vegetation that blessed his property.

This tract of climax, sugar maple-northern red oak forest was safe (at least for the foreseeable future) to perpetuate the species composition and structure of this potential natural vegetation, and re-develop that forest range vegetation should natural disturbances restart plant succession on this north slope, limestone sere.

Most tree regeneration on this climax Ozark forest was sugar maple (including saplings in the background), but there was also some reproduction of bitternut hickory, white ash, and, of course, northern red oak. Herbaceous species in this vertical "photo-transect" were mostly ticktrefoils.

277. The players at play's end- Species composition of a climax sugar maple-dominated forest was furnished by this photograph of a north slope, mesic-limestone forest in the western Ozark Highlands. All visible trees were sugar maple except the tree with single (left center midground) and largest trunk (slkightly right of center background)both of which were northern red oak. In this tract of forest a high proportion of sugar maple had multiple trunks. Boles of the same tree were different sizes (hence of apparaently different ages). These were secondary shoots (basal trunk sprouts) which demonstrated the importance of vegetative (asexual) reproduction of this species (especially in this specific forest).

The part of this forest's understorey was unusually sparse compared to most of the rest of its vegetation. Ticktrefoils and regeneration of sugar maple, northern red oak, and bitternut hickory comprised most of the lower strata.

278. Entrance of some new members of the cast- The next generation of sugar maple and northern red oak assured perpetuation of this climax, north slope, Ozark Hghlands forest range vegetation. Individuals of sugar maple, the dominant species of this forest (and the forest type it was representative of), included the larger tree at right foreground and the two-trunked, mature tree in center background as well as the three or four saplings with light-colored bark (left foreground back to background). Individuals of northern red oak, associate species, were the large pole-sized tree at left margin and larger tree in right-of-center background. Many of the larger seedlings and smaller saplings were bitternut hickory which were interspersed with those of sugar maple and northern red oak.

A considerable number of flowering dogwood and eastern redbud were present, but these were in the background of this photographic view. A high percentage of the ground level layer of vegetation was Virginia creeper, which in this tract of forest range, tended to be more of the ground cover form than the vertical, tree climbing form.

279. The last set of players in the play of a sugar maple forest- Two photographs in the interior of a north slope, mesic-limestone forest in the western portion of the Ozark Plateau provided strong ecological evidence that northern red oak, the general associate tree species of this tract of forest, was subclimax to the climax sugar maple. Both of these slides featured a local stand of uneven-aged northern red oak, including some seedlings of this associate species. Most tree regeneration, however, was that of the Very Tolerant (Wenger, 1984, p. 3) sugar maple. While all the large, mature and pole-sized trees were northern red oak almost all of the seedlings and saplings (up to near pole-size) were sugar maple.

Northern red oak, bitternut hickory, and white ash were represented in seedling and small sapling classes, but in such small (probably closer to trace) proportions that sugar maple was undeniably going to dominate almost all of the canopy of this forest in the not-too-distant future. These three species are generally relatively long-lived (on mesic, north slopes) and grow into large trees (comparatively speaking for trees in the Ozark Highlands) so that mature trees of these hardwoods persist into the climax sugar maple-dominated forest type. Any of these three tree species plus, on a more restricted basis, basswood are local associates of sugar maple. The latter has much slower growth rates on forest sites (even the most mesic and favorable of them) in the Ozark Plateau, but it exceeds all other trees in forest tolerance so as to regenerate in the forests it dominates, and at rates of reproduction that so "out-distance" its associate species that sugar maple is the last of the cast on the forest stage before curtains close. Sugar maple "steals the show" in the final act as it becomes the sole dominant of the climax forest.

Shrubs in the understorey of this northern red oak stand included eastern redbud, flowering dogwood, and summer grape, the latter of which extended up through all layers of this climax forest. The most common, abundant herbaceous plants were Desmodium species.

Starting over (chapter one): One year (almost to the week) after photographing this third tract of sugar maple-northern red oak-dominated forest (one of old-growth status on a steep north slope with mesic, limestone-derived soil) the forest was clearcut by independent (gypo) loggers. Nothing was done on the land or with the forest vegetation except to remove logs (for pallats, including the sugar maple though a log of black walnut might have found other uses). Slash was not flat cut to enhance rotting but merely left after bucking up what logs were wanted. Some limbs of crowns protruded several feet in the air. Several larger logs were hollow and where left as they had fallen. There did appear to have been any fuel wood recovery although some portions of loaded logs might have eventually found their way into fire wood.

Logging operations were completed in August and there was some plant growth prior to first killing frost in late October. Some cool-season species germinated (mostly annuals and biennials) or sprouted (mostly trees and shrubs; some perennial grasses) that autumn and/or early winter. In late May and early June (late spring), the beginning of first full growing season for warm-season species (including all tree and shrub species), a series of photographs was taken of the clearcut forest. Several of these slides were shared below. They presented the essence vegetation development, the process forest community recovery through secondary plant succession. This was an example of the action of "dynamic vegetation" as described by Henry Chandler Colwes and Frederic Edwards Clements.

To reiterate specifics basic to forest range type and potential natural (climax) vegetation it was again noted that this forest was at western edge of the Springfield section of the Ozark Plateau, which was part of the Oak-Hickory Forest Region (Barun, 1950, ps. 35, 162-191).

280. Slicked off and starting over (or What hath man wrought?)- Clearcut old-growth sugar maple-northern red oak-dominated forest (with pignut or bitternut hickory, white ash, and basswood as associates). The north slope, limestone soil, mesic, mixed hardwood forest at old-growth state explored above was clearcut (just plain ole highgrading for pallet wood) in July and August of the year after the above photographs were taken. (God's guidance that your hillbilly professor got there a mere year before the white trash with their chain saws.) In these first two and a series of slides below the recovering (redeveloping) forest range vegetation was described and analyzed at season of late spring (late May to early June) of the first full-growing season (first spring) after logging (about nine months post-logging). There had been approximately six to eight weeks immediately after logging before the first killing frost and then one autumn and winter before this series of photographs was taken.

These first two wide-angle "phototransects" gave overall views of the logged forest from a ridgeline (top of the north slope) vantage point. Finer details were presented in subsequent "photoquadrants" at closer camera range.

Atop the ridgeline (hilltop) the species most visible on this new clearcut included mostly pioneering annual composites such as the horseweed or mare's tail, giant ragweed, and daisy or whitetop fleabane (Erigeron strigosus) as well as naturalized (from Eurasia) common sow thistle (Sonchus oleraceus) and prickly lettuce (Lactuca serriola= L. scariola). Japanese brome or soft chess (Bromus japonicus), a naturalized, cool-season, Eurasian annual grass, was also abundant. Japanesse brome was visible as several straw-colored, dead, small bunches (ie. it had completed its life cycle, and produced copius grain for next year's crop sunlight permitting). Another annual forb with considerable cover was common bedstraw (Gallium aparine). Numerous plants of poke or pokeweed (Phytolacca americana), a native perennial forb, were also present. Pokeweed was the first forb species to successfully invade the clearcut forest. Plants of pokeweed appeared within a few weeks following clearcutting and made normal-size growth in the remaining time of the first late summer-autumn following logging. There were a few saplings of redbud that survived the "woody massacre" (first of these two slides) and a fortunate-indeed small sapling of pignut or bitternut hickory (second photograph). Blackberry (Rubus sp.) and a seedling of smooth sumac (Rhus glabra) were other shrubs present on the ridgeline (second photograph).

The shallower soil of and the more exposed (hotter, drier) land surface of the hilltop was more-or-less an old field environment, the classic cutover forest or bare ground point of denudation on this sere. This was in contrast to the greater area of land that was on the north slope that provided a "successional medium" which produced stump-sprouting and seedling establishment of the climax forest tree species along with release of native decreaser grasses in addition to the annual colonizing species that dominated the comparatively area of the hilltop or ridgeline.

Only one of four oak stumps on the hilltop had coppiced (second slide).

Ottawa County, Oklahoma. Late May (late spring). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

281. New life amid manmade devestation- Two views of the north slope sugar maple-northern red oak (dominants)-bitternut hickory-white ash-basswood (associates) mesic forest that had been clearcut about nine months prior to time of these two photographs. These shots were from the base of the north slope extending up to the hilltop (ridgeline) so as to provide a summary or overall view of the clearcut forest and recovering forest vegetation about six to eight weeks after the last killing frost. Species in the first of these two slides included sugar maple, both suckers (stump sprouts; coppice shoots) and seedlings of which were present only two months into the first full-growing season after logging. Stump-sprouting basswood or American linden (lkeft foreground). Coppicing might have begun in the last six to eight weeks right after logging and before the first killing autumn frost. Shrub species present included American hazlenut (Corylus americana) that was beside the basswood, greenbriar (Smilax tamnoides), and fox grape (Vitis vulpinia), sassafras (Sassafras albidum), and redbud, this lattter was the overall most abundant shrub on the clearcut. Herbaceous species ranged from native and introduced annuals like giant ragweed and common sow thistle to the tall-growing (left side) figwort or carpenter's square (Scrophularia marilandica).

In second slide there many resprout shoots (from stump) of sugar maple plus a few of northern red oak. Redbud, most common shrub, was again conspicuous. In this second "phototransect" the herbaceous component was dominated by the native perennial woodland brome (eg. lower right and center foreground). Hairy wood brome is the dominant decreaser grass on north slope, mesic, limestone-soil forests, especially those in high successional status. The plants of woodland brome had been present prior to clearcutting as confirmed in a pre-logging photographic session by the author one year prior to clearcutting. Also present--as in left midground--were a few plants of tall fescue (Festuca arundinacea), an introduced, cool-season, perennial that is managed as an agronomic pasture grass and that has naturalized throughout this region. Tall fescue had undoubtedly established over the entire time period from end of logging until the present (a complete autumn through winter to late spring). There were a number of plants of pokeweed, a native perennial that was the first forb to successfully invade the clearcut (within weeks after logging). The most abundant forbs, however, were annual, colonizing, composite species: common sow thistle (naturalized Eeurasian weed) and mare's tail or horseweed (native) the standard pioneer of old-fields and cutover forests throughout this region..

Saplings in the background (both slides) were of northern red oak, bitternut hickory, and American elm along with redbud.

Ottawa County, Oklahoma. First slide, early June (late spring); second slide, late May (late spring). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

282. All manner of new life- Two views midway up the steep north slope of a clearcut mesic, limestone-soil sugar maple-northern red oak forest in the western Ozark Plateau only two months into the first full-growing season after careless (as in "don't give a damn") logging. In the first of these two "photoplots" there was a copoicing American linden or basswood (right foreground) and two stumps of northern red oak neither of which produced sprouts (adventituous shoots or suckers). There was, however, a new seedling (not more than two-months-old) midway between these stumps. Most new phytomass (plant biomass or "weight") was of horseweed or mare's tail and common sow thistle, native and naturalized annual composites, respectively. There was also much common dandelion (Taraxacum officinale), another naturalized, annual, Eurasian composite. There was also fair representation of hairy wood brome, the dominant decreaser grass for this forest site. The small adult tree or large pole in center background was a northern red oak that somehow miraculously survived the onslaught. (This clearcutting was strictly harvest for saw logs, and for pallets only, such that many pre-log size trees were not cut. This was a logging not a forest-clearing operation. Nonetheless, most trees of any size with only a few exceptions like the small adult of northern red oak, were destroyed by careless logging. Most of this by felling saw timber-trees on top of the next crop and breaking them off or over.Loggers were careless, apathic, and hatchet-happy.) There were a number of seedlings of pignut or bittternut hickory surrounding the remaining young northern red oak. There was a single seedling of eastern red cedar (Juniperus virginiana) that was clearly visible in front of the young adult northern red oak. This young eastern red cedar had to have been present prior to clearcutting. The plant in lower left corner was leather-flower (Clematis pitcheri).

The second "photoplot" featured a stump sprouted American elm amid much pokeberry, a perennial and the first forb to successfully invade the clearcut having grown to full size within two months after logging. There was also much hairy wood brome as in the previous "photoplot". Other common forbs were whitetop or daisy fleabane and mare's tail or horssweed, both native annuals. Another forb was golden Alexanders (Zizia aurea). Most of the remaining saplings in background of this second photograph were sugar maple. There were fewer northern red oak. Saplings of both species had been battered (including broken crowns) by careless logging; time will tell how they progress.

283. Diversity of life on a clearcut- Two more "photoquadrants" roughly midway up a north slope of a limestone-derived soil from which a mesic sugar maple-northern red oak (two dominants)-basswood-bitternut hickory-white ash forest of old-growth state been clearcut about nine months previously. Present forest vegetation had developed from about six to eight weeks before first killing frost after logging through winter until late spring (three weeks before summer solstice). The first photograph was in a local spot where there were numerous stump sprouts and seedlings of American elm and much hairy wood brome, the dominant climax grass for this forest range site. Woodland bromegrass was a peak standing crop with maturing grain; in fact, some plants were senescing and entering dormancy. Redbud which was the dominant shrub across the clearcut was conspicuous in the foreground. Ther was also quite a bit of pokeweed or pokeberry, the first forb to successfully invade the clearcut, having established in the two months remaining of summer after logging. The most abundant forb in the first full-growing season after timber harvest was mare's tail or horseweed. There was also much giant ragweed. Both of these annual native composites are pioneering species. The yellow composite in background was the perennial composite, wingstem (Verbesina helianthoides).

The second photograph centered on stump of a young northern red oak that had not commiced (stump sprouted). It was surrounded and being overwhelmed by fox grape and some tick-clover or tick-trefoil (Desmodium sp.). There was much of the dominant decreaser grass, hairy wood brome (almost everywhere). In addition there were several nice (and quite large) plants of woodland bluegrass (Poa slyvestris) which was readily distinguished by its cespitose habit, tall height, and dormant state as seen by flaxen, amber-colored straw. A specimen of woodland bluegrass was to upper right of the oak stump. Woodland bluegrass was the climax associate grass species for this range site. The tallest forb (eg. to left of stump) was a goldenrod (Solidago sp.) The white-flowered composite was daisy or whitetop fleabane. There were a lot of American elm seedlings to rear of this second "photoquadrant". Redbud, the major shrub, was also common.

Ottawa County, Oklahoma. Early June (late spring). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

284. Up from the stumps- Coppicing of sugar maple two months into the first full growing season following high-grade clearcutting of a sugar maple-northern red oak dominated forest on a north slope, limestone-derived soil. Coppicing is the foresters' term for stump sprouting Two different sugar maple stumps in these two photographs. In the first slide there was also stump sprouting of American elm along with elm seedlings (ie. both asexual ans sexual reproduction of American elm). Also sprouting (from rhizomes) was the shrub, smooth sumac (Rhus glabra) which was very obvious at left of this first slide. Two annual native forbs were present in this first "photoplot": bedstraw and daisy or whitetop fleabane. This sugar maple stump had a profussion of stump sprouts.

The second slide presented a second coppicing sugar maple stump with less prolific sprouting, but with an asexual shoot having mrvelously red-pigmented leaves (autumn-like leaves in late spring). In this second "photoplot" there were seedlings of pignut or bitternut hickory. Seedlings of the native pioneer species, mare's tail or horseweed (Conyza canadensis), were all around the sugar maple stump (eg. a big one at upper right of stump). A second annual composite--though a naturalized Eurasian one--was common sow thistle. Wild ginger (Asarum canadensis) which is a native, climax, perennial forb of mesic north and east slopes in the Ozarks was thriving with its new-found habitat of full light. A second, native, perennial forb--though a much more common one and one that thrives best in "full sun"-- was the mint, wild bergamot (Monarda fistulosa).

In Silvics of North America Burns and Honkala (1990) reported the following findings: 1) sugar maple does have asexual reproduction via stump sprouting (ie. coppicing), 2) percentage of stumps that coppice decrease with increase in tree size, stand density, and years since cutting 3) average number of sprouts per stump decreased with years following cutting, 4) coppicing is more prolific in northern parts of the species range, and 5) coppicing is less in sugar maple than associated hardwoods in the southern part of its biological range.

It appeared that, all factors considered, the degree (proportion or percentage) of coppicing bysugar maple and other dominant tree species on this clearcut was fortunately very high. It was underscored that there was abundant regeneration--axexual and/or sexual--of the climax dominants. Students of forest succession should note well this phenomenon. There certainly was an abundance of colonizing or pioneer plants (a number of plants of several weedy annuals, including the naturaalized aliens), but of native climax trees there was also abundant regeneration axexually (= regrowth of the same genetic individual) and sexually (seedlings of new genotypes). In regards the latter, on to the next slide, please.

285. Up from seeds- Seedlings of sugar maple that were becoming established on a mesic, north slope forest dominated by sugar maple and northrn red oak that had progressed to old-growth state and was then clearcut. Logging amounted to nothing but high-grading by a bunch of local hicks (hillbilly was much too fine a term for this scalawags) and yet by the second month of the first full growing season after clearcutting seedlings of the climax dominant tree species had become established. it was possible that these sedlings had sprouted immediately following clearcutting in late summer of the previous year. However, there was only a period of six to eight weeks after clearcutting before the first killing frost that killed any remaining leaves of warm-season species (this included all woody species on this forest).

Burns and Honkala (1990) reported that sugar maple seedlings are quite sensitive to mositure-stress near the soil surface because they have "a shallow, fibrous root system that lies between the litter-mineral soil interfaces". On this clearcut there had been much wetter than average moisture conditions in both O and A horizons of the soil due to heavy continuous rains in late summer-autumn immediately after logging through winter and spring of the following year. Such an abnormally wet edaphic environment favored establishment of seedling (and probably also stump sprouts) of sugar maple. Pignut hickory and American elm may not be as sensitive to dry soil habitats as the more mesic sugar maple, but abundant precipitation on rocky, well-drained soils could not have hurt these climax dominants either. With increaed light root systems of sugar maple grow down through the organic matter layer and into mineral soil which permits shoot growth and favors plant establishment (Burns and Honkala, 1990). Thess little sugar maples were "off and running".

It was possible that the limb of a felled tree that lay to the west (upper portion of this photograph) of these seedling provided a key factor of shade from late afternoon sun that otherwise might have tried out litter and soil spelling doom to these delicate seedlings. This was an example of microclimate or a microsite habitat.

Conspicuous other species in this "photoplot" included climbing wild buckwheat (Polygonum scandens), wake robbin (Trillium sessile), wild ginger, an unknown perennial composite, and a lichen on the downed limb. This was a combination of climax forbs (wake robbin and wild ginger) and weedy ones (climbing wild buckwheat).

Reestablishment of existing trees (same genetic individuals) had also taken place by coppicing as shown in the two immediately preceding slides.

286. Basswood from the base- Basswood or American linden coppicing from a stump following clearcut logging (exploitative highgrading) of an old-growth north slope forest in the Springfield section of the Ozark Plateau. Sugr maple and northern red oak were co-dominants of this mesic, limestone forest, but climax associate tree species included pignut or bitternut hickory, white ash, basswood, black walnut, and chinquapin (chinkapin) oak.

Examples of coppicing (stum sprouting) in basswood or American linden were presented above at which juncture this form of asexual reproduction was explained in context of this species which is one of the more readily coppicing trees in Ozark forests.

Other plant species shown in this "photoquadrant" included several forbs. Among these broadleaf herbs were the native annual composites, giant ragweed and horseweed or mare's tail, and the naturalized Eurasian biennial common or bull thistle (Cirsium vulgare) all three of which aree pioneer or colonizing species. Shrub species visible in this photograph included redbud, the most common ro abundanrt shrub, one of the wild grapes (Vitis sp.), and Virginia creeper. Grasses and grasslike plants were uncommon on the ground of this photograph or otherwise covered up by broadleaf species, forb and/or woody species.

Oh, but you (like me) wanted grass? OK, next slide please.

287 Released and growing - Hairy woodland or woodland brome (Bromus purgans) growing robustly on the north slope of a clearcut sugar maple-northern red oak-pignut or bitternut hickory-white ash-basswood mesic climax forest. Woodland brome was at its peak standing crop (soft-dough grain) stage in the first spring following highgrade logging. B. purgans is the only native perennial bromegrass in this region. This festucoid grass is restricted to the most favorable (especially most mesic; north slopes) habitats of hardwood forest ranges. Woodland brome is the dominant decreaser grass for this north slope, limestone soil forest site. The major (most important) climax grass benefitted from clercutting, at least initially.

(Woodland brome was treated above in this chapter under the first of three sugar maple forests discussed.)

Accompanying woodland brome in this "phytograph" were seedlings and/or stump sprouts of American elm and pignut or bitternut hickory, redbud (sprouts or already established seedlings released by loss of forest canopy) and Virginia creeper. Redbud was the most common shrub (had most foliar cover) on the clearcut. Some species of wild grape and poison oak/ivy were also common on this clearcut though not visible in this photograph.

Students should note again in this "photoquadrant" (as in several of those above) that clearcutting this climax sugar maple-northern red oak forest created an initial forest range community that was a "blend" of plant species ranging from typical or standare colonizing species (including naturalized aliens) to sprouted clones of pre-existing climax trees as well as their seedlings (new genotypes) along with decreaser grasses. This forest was not going to have to regeneration from the bare ground stage of the sere.

Important ecological lesson: forest plant community in the first growing season after clearcutting was a composite of herbaceous pionering or colonizing species (both annual and biennial; native and naturalized exotic), climax herbaceous species (including perennial cool-season grasses), shrubs (both seral and climax), and all species of climax trees (dominant and associate species; both sexual and asexual in orgin). In other words, on this mesic, north slope Ozark forest site, clearcutting an old-growth sugar maple-northern red oak-pignut hickory-white ash-basswood forest in summer did not return the sere back to earlier (let alone initial) stages of vegetation development. Clearcutting was not the same as forest clearing. Even though harvest for saw timber was an isidious form of highgrading with zero concern for forest regeneration this mismanagement, this abuse of the forest resource, was not the equivalent of an old-field or typical cutover forest.

Secondary succession was a combination of old-field succession plus regrowth of existing trees and shrubs by coppicing (asexual reproduction), establishment of new seedlings (new genotypes) of climax trees (dominants and assocites), and invigoration of climax (decreaser) grasses and forbs.

288. 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.

289. 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.

290. 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.

291. 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.

292. 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.

293. 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.

294. 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 a regrouping or composite shot of the "business end" of prickly lettuce:

295. Inflorescence of a naturalized import- Blooming heads and unopened floral buds (first slide) and clusters of achenes (second slide) of prickly lettuce (Lactuca serriola= L. scariola). These were on plants on an oak-hickory-tallgrass savanna about three/fourths of mile (as the crow flies) from the clearcut sugar maple-northern red oak-pignut hickory-white ash-basswood forest shown in the preceding photographs. One or two branches of one entire inflorescence (flower cluster) on a prickly lettuce were displayed.

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 "new ground" of the clearcut forest, especially on the ridgetop, was ideal habitat for this pioneering annual of the Liguliflorae, the subfamily of Compositae whose members secrete a "milky" latex in their sap in contrast to those of subfamily Tubuliflorae.

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.

296. 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.

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".

Ottawa County, Oklahoma. Early July.

297. Another naturalized commoner- Common sow thistle (Sonchos oleraceus) on an oak-hickory-tallgrass savanna about three-fourths of a mile from this clearcut. 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. The ridgetop of this clearcut forest was just such an environment (in contrast to the deeper and moister soil of the shaded north slope below this high point of the forest.

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.

298. 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.

299. Alien pioneer- Flannel mullein (Verbascum thapsus) is a Eurasian, biennial range forb of the Scrophulariaceae, the snapdragon family, that has completely adapted (=naturalized) to disturbed habitats in North America. Flannel mullein is one of the more widely distributed weeds on its adopted continent. That is, weed by definition (ie. an alien not grown under domestication, at least not purposely) but not weed by function or in an ecological sense. By functioning as a pioneer or colonizing species of drstically disturbed land this cool-season biennial--albeit it an invasive--is often a highly beneficial range and forest plant. Its broad-leafed, overwintering, basal rosettes covering more-or-less bare land serve as protection against accelerated soil erosion. Upon decay, rotting organic matter from flannel mullein serves in the role Clements describead as reaction (Wever and Clements, 1938, ps.234-247 passim) or what others have since labeled facilitation.(Connell and Slatyer, 1977)

Plants of flannel mullein presented here were growing on the upper slope of a two-year old barrow ditch along a county road. These members of an exotic species were preparing the way for range plant species of higher succcessional order. In immediate time scale they were retaining some soil from being washed away from its natural resting place and covering a freshly paved road surface. Such plants were even more beneficial on the forest clearcut described here.

As a secondary benefit this naturalized range forb and weed of tilled soil is a wildflower that always struck this author is quite attractive. Flannel mullein is afterall a tall "snapdragon of sorts", and one that plants itself. Furthermore, when drought comes early this biennial weed will still flower and add some beauty to a "burnt-up" country.

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

300. Flower stalk and flower- Part of the often much-branched flowering shoot of flannel mullein (first slide) and some individual flowers (second slide) of this anturalized Eurasian member of the snapdragon or figwort family. If a nurseryman sold a clueless city slicker the seed of this plant the urbanite would, quite rightly, view it as a garden flower. Yes, a large, rank-growing one, but a desirable (at least, acceptable) gardenflower nonetheless. If a farmer somehow missed a plant of flannel mullein until it reached this stage he would almost assuredly hoe it out cussing it as he did so. Now who has the more impartial perspective.

There is often a common bond of sorts between the rangeman and/or grazier and the "little ole lady in tennis shoes" who enjoys "pretty wildflowers". Certainly there is also a bond between farmer and stockman. They may be one and the same. There has also been an antagonism between the herdsman and sod-busting plowman, one going back Cain and Abel.

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

Six years after clearcutting (chapter two): the following section presented and described the steep north slope mixed hardwood forest in the western Ozark (Springfield) Plateau. Six years prior to images presented in this section an old-growth forest dominated by sugar maple and northern red oak (with pignut or bitternut hickory, white ash, and basswood as associates) had been cut over leaving no standing trees. This logging /firewood harvest was a "clearcut", but not a sound forest management clearcut as in even-age silviculture. Logging was simply a textbook example of "cut-and-run", "take-everything-and-don't-look-back", "sleek-her-off" forest cutting that left a textbook case of cutover forest.

This case of forest plunder set the stage for forest regeneration through secondary plant (forest succession) succession. The earliest stages of this forest recover (reforestation) via secondary succession (including the major pioneer plant species) was presented immediately above.

The following section treated the seral stage beyond the pioneer or initial (= first seral) stage. It was shown above that a relatively high proportion of climax tree species--individual trees of climax species--coppiced or sprouted from stumps of felled trees. Also on this cutover forest in (at) the pioneer stage or first seral stage there was successful recruitment or establishment of seedlings of the climax tree species. Thus, there was a remarkably high cover (density, general presence)--through both sexual and asexual reproduction--of the potential natural (= climax) dominants.

Also shown immediately above, however, were the typical to-be-expected pioneer annual species--both native and introduced naturalized; grasses and forbs--along with shrubs such as grape as well as pioneer hardwood trees including, as shown below, sycamore, chinquapin oak (Quercus muhlenbergia), and black cherry (Prunus serotina).

The section immediately following presented details of the second seral stage of the recovering sugar maple-northern red oak forest. Presented in the following section were some of the stump sprouts of coppiced trees along with growing--in fact, rapidly growing--saplings that had been seedlings of pioneer trees in the above section.

Now watch natural reforestation as secondary plant succession progresses back toward the ultimate climax of a north slope, Ozark Plateau sugar maple-northern red oak forest.

301. Early stage of recovery- Representative sample of the second seral stage of a clearcut sugar maple-northern red oak forest (bitternut hickory, white ash, black walnut, and basswood were associate tree species) six years following a "cut-and-run" logging operation for pallet lumber and fuel wood. Box elder (Acer negundo) and chinquapin oak were minor tree species.

Tallest saplings (and only six year-old trees) in this "phytograph" were sycamore, a common pioneer tree that often persist into the climax forest. Presence of sycamore as the most abundant tree species in this second seral stage forest was noteworthy because sycamore had been absent from the climax forest. Saplings seen here (at least five with distinctive crowns) had all grown from seed. Seed of sycamore is "light and fluffy" so as to be readily carried by wind, but this was unusual habitat for sycamore in this general area. The more common environment for sycamore is bottomland forest, especially in the riparian zone. There were also seedlings and saplings of sassafras (Sassafras albidum).

Major shrub species included Rhus copallinum known variously as winged sumac, shining sumac dwarf sumac, and flameleaf sumac. Eastern or common redbud redbud (Cercis canadensis), which can grow to size of a small tree. Lianas (woody vines) including an unidentified grape (Vitis sp.), 'possum grape (Amelopsis cordata), and Virginia creeper (Parthenocissus quinquefolia) were well represented. There were occasional (and small) patches of blackberry (Rubus spp.), but not nearly as much cover of "briars" as on cutover forest in this general locality. The more mesic environment of this north slope could have permitted more rapid development of cover by tree species that would have out-competed blackberry than on more widespread drier habitats.

Principal forbs ranged from the pioneering perennial commonly called pokeweed or pokeberry (Phytolacca americana) to the introduced and now naturalized forage legume, sericea lespedeza (Lespedeza cuneata). Grass species (not discernable in this view) were given in subsequent slide/caption sets.

This assemblage of plant species formed a nearly impenetrable "thicket". Photographs seen in this section were taken at midday in late July so as to have maximum sunlight for views of the lower layers of seral forest vegetation. Temperature was in low 90s Fahrenheit with approximately 60% relative humidity (and no wind movement inside the "thicket". Percent slope was roughly 60%. This photographer was"wringing wet" with sweat, ripped shirt, and torn pants, but the Nikon FM was safe and the to-be-shared images procured. It was deemed unnecessary to photograph progression of forest succession annually.

302. A young recovering forest- Re-establishment of tree and shrub species on a north slope Ozark (Springfield) Plateau six years after an old growth forest dominated by sugr maple and northern red oak had been coutover (a high grade form of clearcut) and left to regenerate (or not ) on its own. These view were from the base of the steep north slope. Trees were at the small sapling stage whereas shrubs such as lianas, including wild grape, 'possum grape, and Virginia creeper had progressed further in their life cycle, but no woody plants even appproached adulthood. Tallest plants were saplings of slippery or red elm (eg. first and second slide). Other tree species included basswood, box elder, black cherry, chinquapin oak, and basswood or American linden.

The shrub to small tree species, common or eastern redbud was well represented. There was even greater shrub cover of the rhizomatous, colony forming sumac known under various common names including winged sumac, flameleaf sumac, dwarf sumac, and shining sumac. Flowering dogwood (Cornus florida) was much less common, but contributed considerable cover locally.

Two grass species growing in small open patches (eg. foreground, second slide) were both introduced and naturalized perennials: the cool-season tall fescue (Festuca arundinacea) and the warm-season Johnsongrass (Sorghum halapense). There was also some cover of the naturalized Eurasian annual, Japanese chess (Bromus japonicus).The major forbs were the naturalized Eurasian annual wild lettuce (see numerous slides in preceding section), serecia lespedeza, and an unidentified goldenrod (Solidago sp.).

303. From the stump (and otherwise)- Coppice shoots (stump sprouts) of chinquapin oak along with 'possum grape, sessileleaf tickclover or sessileleaf tick trefoil (Desmodium sessilfolium) six years after cut-and-run logging of an Ozark Plateau, north slope, old-growth sugar maple-northern red oak forest (white ash, pignut hickory, black walnut, basswood were some of the other--sometimes associate--hardwood tree species). Every tree in this climax forest had been felled so the "clearcut" operation was not a form of forest regeneration, but instead abusive, explotiative overharvest.

A good number--though fairly small percentage--of stumps of adult trees had coppice sprouts. Sexual reproduction was more important in establishment of hardwood tree species, but asexual reproduction was important as illlustrated by this young chinquapin oak resprout shoot. Coppice regeneration in other tree species was presented in subsequent slides.

304. Early recovery up closer- Interior of a cutover north slope sugar maple-northern red oak forest (bitternut hickory, white ash, black walnut, basswood were associate tree species) in the western Ozark Plateau (Springfield Plateau portion) six years after the old-growth forest had been completely deforested (an improper form of "clearcut").

Vegetation in the first of these two slides was primarily that of the rhizomatous, colony forming shrub going under the common names of winged sumac, flameleaf sumac, shining sumac or dwarf sumac. Plant species in the second slide had developed on and around a large pile of slash. Plant species included 'possum grape (on the slash, foreground), wild lettuce, flameleaf or winged sumac, black cherry, and a young plant of some grape species (growing all around the pile of slash).

305. Mostly pioneer or early seral tree species- Three "photoquadrants" in the deep interior of a recovering (naturally reforesting) cutover former old-growth sugar maple-northern red oak forest on a steep north slope in the western Ozark Plateau (Springfield Plateau portion) six years after all trees had been felled. Other tree species in the old-growth forest had included bitternut hickory, white ash, black walnut, basswood, and chinquapin oak.

The first "photoquadrant" gave a more general (= a broader or larger area) view inside the thicket-like forest plant community at the second seral stage of revegetation. In this view of secondary plant succession six years post logging, the most abundant tree species (one with greatest cover and stem density) was sassafra and the second most plentiful tree species was black cherry. In this field of view the most common shrub was winged or shining sumac while the second most plentiful shrub was Virginia creeper. Soil surface was covered by leaves of these (and other) hardwood species.

The second photograph of the dense, almost-impassable interior of cutover forest vegetation featured a large seedling of bitternut hickory, a climax and associate species in the former old-growth forest, in the foreground and seedlings of sassafras in the background. Sassafras was the most abundant tree species in this second seral-stage forest. Seeds of sassafras had to have been part of the forest soil seed bank for decades because there were no plants of any age/size class (neither trees, saplings, nor seedlings) of sassafras in the old-growth forest. The seedling of bitternut hickory also had to have grown from the soil seed bank, but the nut from which it grew could have been much younger than seeds of sassafras because bitternut hickory was a major associate member of the climax forest at time of logging.

Note that the soil surface was covered completely by leaves of hardwood (tree and shrub) species. Pieces of slash of varying sizes and degrees of decay were also present on the land surface throughout the cutover forest. Examples of slash (which further increased difficulty of movement through the almost impassable seral forest vegetation) were in immediate foreground of the second slide.

The third photograph also showed seedlings (and forest slash) of the seral hardwood species. The two larger stems at left were of black cherry while the the stem in immediate right-center foreground was flowering dogwood. The shade-tolerant flowering dogwood had been a member of the former old-growth forest being a local dominant in the lower woody layer of that climax forest. The stem to right and slightly behind the flowering dogwood seedling was a seedling of bitternut or, as less frequently called, pignut hickory. Bitternut hickory is a shade- tolerant climax tree species that in the former old-growth forest had been an associate to the climax co-dominants, sugar maple and northern red oak. Most other stems in this third (and closer-in) view were those of sassafras which, again, was the most abundant (had greatest cover and stem density) tree species in the revegetating forest.

Leaves of shrubs and trees also covered the soil surface in this third "photoplot" forming a young O horizon of this north-slope forest soil. Note also in this third view the presence of forest slash which, in this case, was the stump of a small tree of bitternut or pignut hickory. That stump showed clearly the cutover ("took it all"; "slicked it off high, wide, and handsome") pattern of complete deforestation of this old-growth stand. Even saplings growing away from mature trees were cut off maliciously and either left to rot where they fell, drug into rough piles of slash, or had some of the lower portions of trunks (they were almost all sapwood) taken off presumedly for firewood as these trunks were far too small to be used as pallet lumber.

All-in-all this logging had beeen one of the most abusive forest operations that one could imagine. It was as if fellers had a hatred for trees and were out to kill as many of them as possible. That is not to say that such thoughts were in the minds of loggers (this rangeman is not a psychologist or psychiatrist), but the results on the land clearly showed that those who "slicked off" this forest had no love or respect for the very resources that provided them their sustenance. Ecological white trash.

306. Some climax tree babies- Three examples of sexual and asexual reproduction of seral and climax tree species in a cutover sugar maple-northern red oak old-growth forest six years following removal of all adult trees, poles, and saplings from this north-slope, Springfield Plateau (far-western Ozarks) hardwood forest.

The first photograph was of a seedling of bitternut or, at times (and confusingly), called pignut hickory (Carya cordiformis) and a cespitose plant of beaked panicgrass (Panicum anceps). Some Virginia creeper and, in far background, winged or shining sumac.

In the second slide presented a seedling of bittrenut or, sometimes, pignut hickory (center) flanked by seedlings of sassafras (left and right). The large, rotting stump was (had been) in the second slide was unidentifiable on-site, but its high degree of decomposition after just six years showed that it could not be a stump of an oak, hickory, or black walnut but was more likely the soft wood of basswood or white ash. There were no stump sprouts from this readily deccomposing stump. Considerable cover of young Virginia creeper grew throughout the recovering forest febetation shown in the second "photoquadrant".

The third slide featured two large stump sprouts (= coppice shoots) of northern red oak. To left of these resprouts there were small seedlings of black cherry and to the right of these asexual small sapling-sized shoots there were seedlings of bitternut hickory. Virginia creeper and small plants of blackberry were common in seral vegetation shown in the third slide.

As shown in this slide-caption set as well as several of those above there were numerous seedlings of bitternut hickory, one of the major associate tree species in this climax sugar maple-northern red oak forest, established thoroughout this seral forest vegetation. Burns and Honkala (1990) explained thaat although bitternut hickory was often interpreted as an Intolerant species, its tolerance was variable. The current author observed quite the opposite response of Carya cordiformis instead noting that it was quite tolerant of shade and competition and readily reseeded even in climax forests--on both upland and bottomland forest sites--nthroughout the far-western Ozark (Springfield) Plateau.

Note: It was possible that some of the apparent seedlings of bitternut hickory were actually root suckers, asexual or vegetative shoots that often arise prolifically from the relatively dense root system of bitternut hickory (Burns and Honkala, 1990). By the way, bitternut or pignut hickory is classified as a "pecan hickory" rather than a "true hickory" and its lighter, less denser, weaker wood is less valuable for tool handles and firewood (Burns and Honkala, 1990).

Savanna of Oak-Hickory Forest and Tallgrass Prairie

The tallgrass prairie-hardwood forest savanna or transition was given a chapter of its own in this publication entitled, Tallgrass Savvanna, plus a section on Prairie Peninsulac in the chapter, Tallgrass Prairie (Interior). One photograph and caption was included at this point to highlight locations for these important deciduous forest and eastern prairie grasslands.

307. Oak-hickory forest and savanna- This is the widespread transition zone or intermediate community type between actual oak-hickory tallgrass prairie understory savanna and the western edge of the oak-hickory forest proper.Post oak and black hickory dominate the tree layer while black and red oak are associated species. Understory is primarily nodding or Canada wildrye and purpletop (Tridens flavus). Virginia creeper covers much of the understory and climbs into tree tops. Blackberry, buckbrush, Mayapple, and black-eyed susan dominate various microsites depending on cover of tree canopy. Chert savanna-chert forest composite type with fire determining which form prevails.

Ottawa County, Oklahoma.June, late vernal aspect. FRES No. 15 (Oak-Hickory Ecosystem). K-73 (Mosaic of Bluestem Prairie [K-66] and Oak-Hickory Forest [K-91]). SAF 52 (White Oak-Black Oak-Northern Red Oak); SRM 801 (Dry Savanna). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

Dominant, Associate, Widespread, and Just Plain Interesting Plants
of the Central and Southern Forests

308. Composite view of Oak-Hickory Forest- Sample of species richness and biological diverstiy of eastern deciduous forest in central and southern regions. Tree on left is pignut or bitternut hickory, the forked trunks with burl on left trunk is black cherry, a young pignut hickory to its immediate right, and the two larger white-barked trunks in background visible between the hickory and cherry are post oaks. The barely detectable short shrub dominating the understory is buckbrush which in this winter season has shed its fruit. Hibernal aspect . Ottawa County, Oklahoma. Hibernal aspect, Late December. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 40 (Post Oak-Blackjack Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

309. Cherries in the woods- Leaves and ripening fruit of black cherry (Prunus serotina). The fruit type of Prunus species is pome. Black cherry is the source of commercial cherry wood, one of the most beautiful of all North Americn hardwoods. Black cherry is not a domiinant or even associate tree species in climax oak-hickory forests, but in absence of fire it increases quickly and can overwhelm the climax dominant and associate oak and hickory species.

Ottawa County, Oklahoma. Late June.

310. Fruit of buckbrush or coralberry (Symphoricarpos orbiculatus)- This fleshy fruit is widely held to be valuable for bobwhite quail, squirrels, and rabbits. It is perhaps the most dominant species of the lower shrub layer (taller shrubs like dogwood, persimmon, sassafras, and redbud being local dominants of the taller shrub layer). Ottawa County, Oklahoma. Early December (and fruits may be shed within the month or persist on twigs in dried form until spring depending on the year).

311. Flowers of flowering dogwood (Cornus florida)- This is probably the most widespread understorey shrub or small tree in the oak-hickory forest forms and, therefore, is typically the dominant of the lower woody plant canopy. In the spring flowering dogwood (often accompanied by redbud) turns the dark, drab-colored, bare woods into a colorful botanical banner announcing start of another growing season. This species with it's delightful spring and, as shown immediately below, fall display is the State Tree of Missouri. Ozark Plateau. Newton County, Missouri. April.

312. Leaves and mature fruit of flowering dogwood in fall coloration- It is an age-old argument as to whether the spring or autumn is the more colorful in the deciduous forests of eastern North America. The argument cannot be resolved by admiring the flowering dogwood which contributes to the beauty of the timberlands in both seasons. The fruit is a major food source for wild turkey (Meleagris gallopavo) and bobwhite quail (Colinus virginanus), especially in the south as in the Ozark Mountains, while the twigs are important browse for white-tailed deer (Odocoileus virginianus). Indians had several uses for this shrub (eg. dyes, herbal remedies).The hard, tough, tight-grained wood has many non-construction uses.

Newton County, Missouri. October.

313. Shadbush or eastern or downy serviceberry (Amelanchier arborea)- Another shrub of the oak-hickory-- and general eastern deciduous-- forest is downy serviceberry. The common name of shadbush comers from the hill folk who noted that blooming of this species often coincided with the spring runs of shad (Alosa spp.). Downy serviceberry is often confused with flowering dogwood because the flowering periods of these two species often overlap and that of one species may precede or lag behind the other depending on conditions in any one spring. (Dogwood gets all the credit by the way.)

Serviceberry is one of the many members of the rose family which is the single most important family of range browse plants in North America. Amelanchier species are in the Pomoideae, the pome fruits subfamily, of Rosaceae. Shadbush undoubtedly provides some browse and the fruits are eaten by birds and furbearers, but it is not common enough to be a major feed plant.

The bluffs of Modoc Creek, Ottawa County, Oklahoma. April.

314. Downy serviceberry in full bloom- In the opinion of your author when this species is flowering it is often the most striking and showy shrub in the oak-hickory forest. Much of the enjoyment of the spring blooming in the hills is due to serviceberry and not flowering dogwood, but a human population of predominantly city dudes does not know the difference and just calls everything other than redbud a "dogwood". Ottawa County, Oklahoma. April.

315. Flowering shoot tip of downy serviceberry- The inflorescence and two newly emergent leaves of shadbush or eastern serviceberry. Ottawa County, Oklahoma. April.

316. Goatsbeard or Goat's Beard (Aruncus dioicus var. pubescens= A. pubescens)- Goatsbeard is another striking member of the Rosaceae. It is far from common in the oak-hickory forest, but when it is encountered it causes the most hill-hardened hillbilly to pause and "take a gander". Aruncus literally means "goat's beard" and according to various manuals this name can be traced to Pliny and later applications to Eurasian species of this genus. This North American shrub has a range from the extreme northeastern part of the continent south to Alabama and west to Oklahoma.

Springfield Plateau section of Ozark Plateau; base of limestone hill in Newton County, Missouri. June.

317. Inflorescence of Goat's Beard- Goats's Beard is a dioecious, but the flower clusters are similar for both sexes. Newton County, Missouri. June.

318. Catkins (male flowers) and mature fall fruit of common hazelnut (Corylus americana)- Another (but usually sparsely populated) shrub of the oak-hickory forest is the hazelnut. This nut was highly prized by the early American backwoodsmen, the proper historic name for the first wave of frontiersmen who moved westward beyond the Fall Line into the dark, forbidding, and mysterious deciduous forest of eastern North America. For millinia prior to the first footprint of man (God only knows what shade of skin or species of Homo it was) hazelnut was feed for wildlife ranging from wild turkey to furbearer to native ruminant.

Male flowers of this monecious species are borne in catkins that are some of the first flowers in the late winter or early spring in the oak-hickory forest. The specimen shown here is from an individual of the variety C. americana var. indehiscens (formerly interpreted as C. cornuta) delineated as a separate taxon by the united fruit bracts that make the fruit to appear unopened. Fruits appear singularly up to as many as four in a cluster like the one pictured here.

The family to which this little nutbearer belongs is another source of controversy. Some treat it as a member of the birch family (Betulaceae) with separate tribes of Betuleae and Coryleae while other workers place it in it's own hazel family (Corylaceae).

On second terrace of Modoc Creek in Ozark Plateau. Ottawa County, Oklahoma. Catkins, February; fruit, October.

319. Pawpaw (Asimina triloba)- Pawpaw or custard apple is another (though less common) shrub of the eastern deciduous forest, in particluar the oak-hickory forest. As is the case for many of the understorey plants, especially forbs, pawpaw reproduces asexually by suckering from extensive creeping rootstocks. In fact, this is usually the main mode of reproduction there being springs when pawpaw does not bloom (and fruit-set is much less frequent than flowering). On bottomland forests pawpaw often forms estensive colonies with broad, shiny leaves. Ottawa County, Oklahoma. August.

320. Immature fruits of pawpaw- When pawpaw does bloom and set fruit, which is less frequent than non-fruiting years, it produces this backwoods delicacy for 'possums (Didelphis virginiana), coons (Pyocyon lotor), and coon- and 'possum-hunting hillbillies. The green color of the thin fruit skin indicated that this fruit was still immature. Ripe fruit turns yellow and finally brown when over-ripe. Ottawa County, Oklahoma. August.

321. Spicebush (Lindera bezoin) leader- Twig of spicebush with staminate flower clusters. Spicebush is dioecious, usually with male flowers more conspicuous than the females. This shrub species is usually restricted to the fertile, moist conditions of forest bottomlands. The common name was probably derived from the use of the dried and ground fruit as a substitute for allspice during the American Revolution. Later fruits , leaves, and twigs were used by backswoodsmen to brew a fragrant tea, a practice almost certain to have been adopted from the Indians. Spicebush is in the laurel family (Lauraceae) which includes that finest of all North American teas, sassafras (Sassafras albidum). That member of the family was presented under the tallgrass savanna (Grasslands).

Western part of Ozark Plateau. Flood plain of Modoc Creek, Ottawa County, Oklahoma. March.

322. Spicebush in summer- Twig, leaves, and fruit (mature and immature) of spicebush. Fruit is ripe for gathering and powdering to make a unique seasoning relished by Ozark hillfolk. This shrub seldom attains a height exceeding three or four feet and is usually limited to the understorey of bottomland deciduous forests. It is a member of the climax plant community.

Terrace of Modoc Creek, Ottawa County, Oklahoma. July.

323. Wild hydrangaea (Hydrangea arborescens)- This (and the fine specimens shown in the next set of two slides) grew on a moist north slope by a wet-weather srping. Wild hydrangea is not only one the most showy of the understorey woody species in the eastern deiduous forests, but also one of the most mesic shrubs of this formation with occurrence of this member of the Saxifragaceae limited to moist or even wet habitats. Newton County, Missouri. June.

324. More beauty- Details of wild hydrangea inflorescence. The larger, conspicuous, three to four-petaled flowers on the periphrey of the flower cluster are sterile (without sex organs). These appear to serve as attractants to insect pollinators. Newton County, Missouri. June.

325. Lowbush huckleberry (Vaccinium vacillans)- Ericaceae Vaccinioideae Flag Springs State Park, McDonald County, Missouri. June.

326. Highbush huckleberry, squaw huckleberry, or deer berry (Vaccinium stamineum). Ottawa County, Oklahoma. June.

327. Poverty oatgrass (Danthonia spicata)- This cool-season bunchgrass often forms rather extensive carpet-like stands made up of these green clumps in the understorey of oak-hickory forests. This occurs mostly on the drier habitats like the steeper of south slopes and the more shallow, stoney, and acidic soils where the larger grasses (eg. the bluestems, panicgrasses, beakgrain, and woodreed grass), forbs, and shrubs like blackberry (Rubus spp.) are absent or stunted. Poverty oatgrass is also more common on "edges" between deciduous forest and tallgrass prairie where the more drought- tolerant trees predominate. For example, in the habitat shown here oak leaves in the background belonged to blackjack (Quercus marilandica). Ottawa County, Oklahoma. June.

328. Poverty oatgrass at more advanced maturity- Several cespitose plants of poverty oatgrass on the same precise microsite as shown in the immediately preceding slide (first of these two slides) and of a single plant of poverty oatgrass (second slide) just a short distance removed from those in the first photograph. These shoots were photographed in another year and at more advanced maturity (phenological age) from those presented immediately above.

The rocky (chert limestone) nature of the microhabitat of these examples was representative of the environment on which plants of this species excell. It was shown above, however, that big bluestem will sometimes grow beside plants of poverty oatgrass such that these two species are range "pals" on oak-hickory/tallgrass savannas.

Ottawa County, Oklahoma. Early July; grain at maturity just prior to its shatter stage.

329. Rock-poor or rock-rich?- Spikelets of poverty oatgrass at grain-shatter phenological stage. Number of florets per spikelet ranges from three to seven, rarely up to nine in poverty oatgrass (Steyermark, 1963, p. 142; Diggs et al., 1999, p. 1260).

The rock on which these spikelets were laid was common chert, the erosion-resistant remains of Mississippi limestone that formed much of the parent material of the Ozark (Springfield) Plateau.

Ottawa County, Oklahoma. Late May. (Maturity of caryopses varies considerably in poverty oatgrass even within a local population. These spikelets were at grain-shatter stageat this time whereas othr spikelets in the same local population had not reached grain-ripe/grain-shatter stage six weeks later.)

330. Not impoverished- Local dense stand (population) of poverty oatgrass in the western Ozark (Springfield) Plateau. This example of a single-species stand (or colony) at peak standing crop was a particularily--though not at all unusually-productive sward growing on a extremely rocky yet deep soil on the perimeter of a mixed oak-black or Texas hickory second-growth forest.

Agrostologists cannot make up their minds (and that is not unusual either) as to proper taxonomic placement of Danthonia species. This genus has been placed in its own tribe, Danthonieae, within either subfamily Arundinoideae (Gould, 1975, ps. 56-58; Gould and Shaw, 1983, ps. 329-331) or Eragrostoideae (=Chorideae) (Diggs et al, 1999, 1259-1260).

Ottawa County, Oklahoma. Late May; grain at mid-maturity stage.

331. Panicle of stout woodreed or woodreed grass- This festucoid grass (tribe Aveneae) is a major provider of nutritious forage in the understories of carefully managed (ie.conservatively stocked) oak-hickory forests. It is a major species only on a local basis because years of overgrazing have greatly reduced it's abundance and even occurrence. Woodreed is adapted to diverse sites within the oak-hickory forest range type thriving on both bottomland and stream bank habitats as well as shallow upland soils. Bank of Modoc Creek, Ottawa County, Oklahoma, September.

332. Stout woodreed is a perennial grass that has an obvious prolonged growing season and pronounced allocation of resources. In the Ozark Plateau flowering occurs in late summer or early fall (August to October), but new basal shoots appear soon after completion of the annual life cycle (death) of current-season shoots. Thus there are usually green shoots in some phenological stage yearlong, but the growth and development of these shoots is extremely slow. In this photograph young shoots (tillers: intravaginated shoots) of two to four inches in length are growing among last year's tillers which reached their maximum mature height of three to four feet. Shallow savanna range site (a shallow upland soil overlying a solid layer of chert). Ottawa County, Oklahoma, December.

333. Close-up of the woodreed tillers of the previous slide- Allocation of resources so as to maintain some live tillers yearlong is likely an adaptation to the dark shade existing from spring through autum due to dense canopy of the forest during this frost-free period.(mid-April to mid-October in the Ozarks). Woodreed requires a full year under these conditions of limited light to fix enough carbon to complete the annual life cycle and store root reserves to initiate next year's growth. Any plant living in habitats with such a low light intensity and a protracted regime of photosynthesis and growth will obviously be quite vulnerable to defoliation and require careful grazing management.

Not only is conservative stocking important for regeneration of shrubs and lumber trees but it is also essential to prevent overgrazing of grass species growing in the dimlite, stressful understory. Given the shallowness of soils on some sites in deciduous forest range in conjunction with the commonness of periodic summer drought it is likely that grasses like stout woodreed and beakgrain suffer from the stress of water shortage. This would be �true in spades� for these summer-flowering festucoid grasses which must compete with the more heat- and drought- tolerant dominant panicoid grasses like big bluestem and beaked panicgrass (Panicum anceps). Combinations of dense shade, shallow soils, drought, competition, etc. make for a harsh environment even in humid precipitation zones. Excerbation of this condition with overgrazing or improper season of use is one reason why deciduous forest ranges have become deteriorated.

Furthermore (and contrary to popular misconception) these deciduous woods ranges were the last open range (the true meaning of open to the public, a grazing commons, and not just absence of fences) in the United States. The actual Public Domain ranges of the Intermountain West were closed and came under some scientific regulation by the Grazing Service (later, Bureau of Land Management) with attempted enforcement of the Taylor Grazing Act in the 1930s. By contrast, state laws officially closing the range (passage of mandatory "fence-in" laws) in Missouri, Arkansas, Oklahoma, and Texas were not passed until the 1950s. Free-ranging, acorn-eating, razorback, rooter hogs could be found sleeping and rooting on county courthouse squares through the first half of the 1950s throughout portions of the Ozark, Boston, Quachita Mountains in Oklahoma, Missouri, and Arkansas and in the Big Thicket of Texas. The account for the latter can be found in Sitton (1995, ps. 194-273 passim). Rowley (1985, ps.239-241) noted that control of livestock grazing on National Forest "…in the South [Region 8] remained an unfinished business in the early 1960s…", and that in "… the 1950s range continued to be the southern region's 'bastard child''. Closing the southern ranges had always been a battle in intself (King, 1982) because (and this is another scientific, economic fact that is contrary to popular, and incorrect, opinion) livestock raising was far more important economically and culturally than it was credited with. This was true even for the antebellum South when the planter class was at its zenith (McDonald and McWhiney, 1975). It is no wonder why professionally trained foresters still harbor strong prejudices against even proper (sustainable) livestock grazinig in the eastern deciduous forests.

Location Note: more detailed coverage of woodreed grass was offered in Range Types of North America in the chapter, Central and Southern Forests-II.

334. Beakgrain ( Diarrhena americana)- The understory of oak-hickory forest and its ecotone or leading edge with tallgrass prairie supports species of grasses which are not commonly discussed in standard range plant references (even though they often produce appreciable amounts of palatable, nutritious forage). Beakgrain is one of these species which was included here as an example of a situation commonly confronting the range practitioner (especially one just starting his career). A relatively obscure species which is given but short-shrift in handbooks or range guides is nonetheless of local or occasional importance. Rangemen must learn these important "locals" much like they must learn the locally powerful politicians.

335. Spikelets in a panicle of beakgrain- This member of the Festucoideae is in its own tribe (Diarrheneae). Floodplain of Lost Creek, Ottawa County, Oklahoma, July.

336. 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.

337. 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.

338. 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.

339. 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.

340. 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.

Most of the herbaceous understorey species in the various eastern deciduous forest cover types are forbs. Forb is a term used by foresters, rangemen, and wildlifers in reference to any herbs (=. herbaceous plants) that are not grasses or grasslike plants. In other words, forb refers to all soft-stemed dicots and to any monocots that have conspicuous petals. Forb is not a precise botanical term , but rather one used by professionals in the natural resource management fields. Range and forest plants are either 1) woody or 2) herbaceous. The woody plants are either trees or shrubs, the distinction between which is not always obvious. The herbaceous species (herbs) are either grasses, grass-like plants, or forbs. (Together these are generally the five "kinds"-- as in categories or groups not species-- of vascular range and forest plants.)

Some of the more common and conspicuous forbs of the eastern deciduous forest communities were included immediately below. All of these were growing in the Springfield Plateau section of the general Ozark Plateau or Ozark Mountains.

341. Jack-in-the-pulpit or Indian turnip (Arisaema atrorubens)- This unique and picturesque forest forb is in the arum family. It has one of the more expansive species ranges of any understorey species in the eastern deciduous forest formation. Jack-in-the-pulpit is not always plentiful where it grows, but it does indicate a general lack of abnormal disturbance and botanically diverse forest community when present. This is a very interesting species. Indians made much use of it. It does cause some animal poisoning. Of most interest perhaps are the basic botanical aspects of this monocotyledonous wild flower. The inflorescence is a spadix in which "Preacher Jack" is the fleshy spike of imbeded flowers "who" is inside of an enveloping bract known as a spath. Individual plants are either monoecious or dioecious and, reportedly, individual plants can change the sex of flower from one year to the next. Jack-in-the-pulpit also reproduces vegetatively from creeping rootstocks. Plants perform the most "kinky sex". This is but one of countless interesting things about Range Management and Forestry.

In bottomland forest of Modoc Creek, Ottawa County, Oklahoma. April.

342. Green dragon (A. dracontium)- This Arisaema species is another woodland forb which is also enjoyed as a wild flower like it's "cousin" jack-in-the-pulpit. These two species sometimes occur together in rich, moist forest soils. Green dragon spreads by creeping rootstocks, a form of asexual reproduction that is apparently quite efficient in the dim-lite forest floor of the deciduous forest. Arisaema species begin growth and flower early in the spring before the deciduous trees leaf-out and form their light-excluding canopy.

First terrace of Modoc Creek, Ottawa County, Oklahoma. May.

343. Detail view of green dragon- Leaves and the spadix and spath of the "flower" of green dragon were presented here. Modoc Creek, Ottawa County, Oklahoma. May.

344. Deer's tongue, adder's tongue, or dog-tooth violet (Erythronium albidum)- This is a forest lily that emerges early in the spring through the deep layer of shed oak and hickory leaves on the forest floor. It then promptly flowers before the overstorey trees grow their annual canopy that affectively blocks most light from reaching the ground. Hill-tromping hillbillies enjoy this delightful forb as a harbinger of spring. Ottawa County, Oklahoma. March.

345. Trout lily (Erythronium americanum)- This even showier species is more restricted in its environments in the Ozark Plateau oak-hickory forest than is E. albidum. The trout lily featured here was growing on a bench on the north slope of limestone bluffs that supported a sugar maple-bitternut hickory (Acer saccaharum-Carya cordiformis) mesic-limestone forest (Oak-Hickory Forest- II).

On bluffs above Modoc Creek, Ottawa County, Oklahoma.April.

346. Wake robin (Trillium sessile)- This is another member of the Liliaceae that emerges early in the spring when it can get enough light to complete it's annual cycle of life. Wake robbin grows best and to it's largest size on rich, moist soil. Ottawa County, Oklahoma. April.

347. Walking up on wake robbin- Two succcessively closer views of T. sessile in the western Ozark Highlands. These fine specimens were growing on the boggy soil of a green ash (Fraxinus pennsylvanica)-sycamore (Platanus occidentalis) wet bottomland forest (see Oak-Hickory Forest-II).

348. May apple or mandrake (Podophyllum peltatum)- May apple is probably the most widespread and, as it grows in sizeable colonies from extensive roostocks and sports large, spreading, uniquely shaped leaves, the most conspicuous forb of oak-hickory forests. The fruit is edible, but the foliage does not appear to be grazed by vertebrates. This mandrake should not be confused with the other species of that name that is native to the Mediterranean Region (the mandrake of the Holy Bible). This forb is often the locally dominant species of the upper herbaceous layer of the multi-layered deciduous forest. Ottawa County, Oklahoma. May.

349. Berry on the bottom- A ripe fruit of may apple still attached to the crotch or point of branching in this species was presented in the first of these three photographs. The second and third slides showed the fruit cut open with seeds remaining attached and then with seed cluster removed and set aside, respectively. May apple fruit is a berry. Though edible, this berry is pretty much tasteless (unless one is extremely hungry-- as in starving). A bona fide hillbilly would eat his specimen. The one who gave you these shots fulfilled this hill culture requirement.

Ottawa County, Oklahoma. Early July.

350. Dutchman's Breeches (Dicentra cucullaria)- The range of this forest forb extends from Quebec westward to Oregon and Washington and south to Missouri and Oklahoma. As seen from this photograph it "does it's thing" early in the spring before the trees leaf out and exclude light from the forest floor. Even then, as also clear from this slide, light is limited by trunks and larger limbs.

This species was discussed in Notes on Western Range Forbs (Dayton, 1960, p. 220-222) where it was noted that the bulbs are poisonous to cattle when they are pulled from the soil and eaten along with the shoot. Burrows and Tyrl (2001, ps.706-709) provided a recent synthesis. They noted that Dicentra species contain several isoquinoline alkaloids that affect the nervous system and cause trembling and stggering, but they added that the plants were so uncommon as pose no serious problem, and besides animals usually completely recovered.

Your author noted that this is another application of the Cardinal Principles of Range Management, Proper Season of Use in this case. When forbs like the ones shown in this section are growing there is very little valuable forage in the oak-hickory forest because what grasses and valuable forbs (eg. legumes) do grow have not produced feed. Ergo: stay the heck off the oak-hickory forest ranges at this time. Perhaps of even more importance is the fact that twigs and buds are very susceptible to browsing injury at this season and growth such that regeneration of hardwood species is easily adversely affected. The wood products from these oak-hickory forests are a greater source of revenue than is the 10-30 acres per AUM valuation. It does not require a post doc to figure this out folks.

Base of limestone bluff along Modoc Creek in Springfield Plateau section of Ozark Plateau; Ottawa County, Oklahoma. March.

351. Detail of Dutchman's Breeches- The source of the colorful common name for this range forb can be seen in this close-up shot of Dicentra cucullaria. Limestone bluff along Modoc Creek, Ottawa County, Oklahoma. March.

352. Bloodroot (Sanguinaria canadensis)- Here is yet another picturesque range forb from the early spring floor of the oak-hickory forest. And it comes with another colorful common name. This one derived from the red- or orange-colored exudate from the rhizomes that was used by backwoodsmen as a dye to add a little color to their drab garb of buckskins and hunting shirts and gingham or flour sack dresses.

Bloodroot is a flowering neighbor of it's pantalooned "cousin" just presented. Both bloodroot and Dutchman's breeches bloom at about the same time and in close proximity to one another. Bloodroot seems to grow slightly higher on hillsides, especially those adjoining streams, than does Dutchman's breeches. Yes, these two species are "kissin' cousins" of a sort. Some taxonomists placed Dicentra species in the Fumariaceae (fumatory family) while others interpreted Dicentra as belonging to the subfamily Fumarioideae in the Papaveraceae (poppy family). Bloodroot has commonly been interpreted as a member of the Papaveroideae subfamily of Papaveraceae.

On a limestone bluff alongside Lost Creek in the Springfield Plateau section of the Ozark Plateau physiographic provinece. Ottawa County, Oklahoma. March.

353. One shoot (from a rhizome) of bloodroot at peak bloom and accompanied by a detailed view of it's inflorescence- Take note of the fruit immediately above the leaf. Fruit of bloodroot is a one-locular (locule= cavity of, in this case, the ovary) capsule. This fruit is very similar, obviously, to the capsule of poppy.

Bloodroot is one of the first native forbs to flower in the spring on the leaf-covered floor of oak-hickory forests. To be able to live through another hard winter and share the joy of the bloodroot in bloom is one of Nature's blessings to the hill-billy.

Bluff above Lost Creek, Ottawa County, Oklahoma. March.

354. Some other seasoning to go with oak and hickory- Wild ginger (Asarum canadense) grows farom New Brunswick westeward to Minnesota and south through Kansas and Oklahoma to Arkansas and Alabama thereby encompassing most of the eastern deciduous forest formation in its range. Its best habitat is rich wooded slopes, especially calcareous soils associated with bluffs like the ones above Lost Creek in the western part of the Ozark Plateau on which this lovely plant was growing.

The common name of wild ginger indicated the use that Indians and white frontiersman made of the rhizomes (either fresh or dried) as a seasoning and substitute for the real thing (Steyermark, 1963, p. 572).

On east-facing bluffs above Lost Creek, Ottawa County, Oklahoma. April.

355. In rocks and earth tones- Wild ginger growing amid rocks on the east side of limestone bluffs above Lost Creek in the western part of the Ozark Plateau (Springfield Plateau section). The purplish-brown color of flowers of this species blend in with fallen leaves and moist soil almost as if camoflage was the objective. Wild ginger propagates asexually from long rhizomes that give off a pungent aroma (Fernald, 1950, p. 562) which as explained in the preceding caption were used as a seasoning.

On east-facing bluffs above Lost Creek, Ottawa County, Oklahoma. April, full-bloom stage.

Bur on the bottom- A young plant of bur-cucumber (Siccyos angulatus) growing in the understorey of a mesic forest of sycamore, box elder, white ash, and northern red oak. This particular was in immediate pre-bloom phenological stage. Herbaceous neighbors included hop caric sedge (Carex lupulina), rice cutgrass (Leersia oryzoides), cardinal flower (Lobelia cardinalis), and spotted jewelweed (Impatiens camssis) .Bur-cucumber is an intereesting member of the cucumber,pumpkin, or gourd family (Cucurbitaceae) that grows (mostly) in bottomland forests,especially those in floodplains of small streams.

Floodplain of Modoc Creek, Ottawa County, Oklahoma. .Early August.

356. Solomon's seal (Polygonatum commutatum) - This and the next species are members of the asparagus tribe of the lilly family (Liliaceae) are found in the vast deciduous forest region of eastern North America. The pictured specimens of these two species were growing in the oak-hickory forest association in the Ozark Plateau. They were conspicuous plants and though of no commercial value often elicit favorable responses from people visiting the "woods", in particular native plant and wild flower buffs. Young spring shoots of Polygonatum and Smilacina species were eaten by Indians and backwoodsmen. The range of P. communatum is from New England westward to Oklahoma. Ottawa County, Oklahoma. May.

357. Flowers and leaves of Solomon's seal- The bell-shaped flowers and rounded to broadly acute leaves were obvious between views in these two photographs. Polygonatum species are strongly rhizomatous. The origin of the common name was from the seal-resembling scars on the rhizomes. Ottawa County, Oklahoma. May.

358. Synopsis views- More summary images of shoots of Solomon's seal. Thee plants were growing on a transition zone between tallgrass pariaire and oak-hickory forest in the western part of the Ozark (Springfield) Plateau. Shoots of these plants, including a super-abundant crop of flowes, were the result of a record wet spring (fololowing a record cold winter).

Solomon's seal has been treated consistently as a member of the asparagus subfamily (Asparagoideae) of the Liliaceae (Dayton, 1960, 23-24; Smith, 1977, p.256). Parallel veins of the Monocotyledoneae were obvious in these photographs.

The author never witnessed directly any grazing of Solomon's seal, but the best examples of this species (both specimens in these two and the next three slides as well as in the immediately preceding two slides) were in fence rows just outside of (on the other side of fences) extremely heavily grazed pastures. The inescapable, self-evident conclusion was that beef cattle had grazed out shoots of Solomon's seal that otherwise would have spread via robust rhizomes into the two adjoining pastures. It was more likely that plants (or shoots) of Solomon's seal had existed in these pasture until they had been killed out by heavy cattle-grazing with only remnants persisting in outer fence rows (from which cattle had been excluded). It is amazing what plant species will be eaten by grass-preferring Bos taurus and B. indicus.

Ottawa County, Oklahoma. Late May; peak stage of blooming.

359. Musta' been a record- An extremely abundant crop of inflorescences on plants of Solomon's seal at edge of an oak-hickory forest-tallgrass prairie transition zone. After the coldest temperatures ever recorded for this area in February (in excess of -20 degrees Fahrenheit) and one of the wettest springs in history these monocotyledonous forbs produced this remarkable yield of flowers. With a recognition of the rarity and raw beauty of this pehnomenon the Nikon FM was brought into action for some heavy "photo-artillery shelling". This action in combination with the ground-clutching posture of the infantry trained photographer and the green color-capturing quality of Fujichrome 100F produced textbook images for those who appreciate God's own wildflowers and picturesque range forbs.

Solomon's seal is first thought of as a forest plant, but in the Springfield portion of the Ozark Plateau this forb is equally at home in dense grassland, both tallgrass prairies and even thick patches of Johnsongrass (Sorghum halapense) which will not have not reached great heights by the time Solomon's seal bears flowers.

Ottawa County, Oklahoma. Late May.

360. Cluster of woodland beauties- Example of the inflorescence (consisting of numerous flowers) arising from the leaf axil in a plant of Solomon's seal in an open oak-hickory forest in the western edge of the Ozark Highlands. The drooping (pendulant) feature of the clustered flowers is characteristic of this forb. Kaul et al. (2006, ps. 562-563) provided an especially good description of one of the more distinctive herbaceous forest plants in the Ozark Plateau.

Ottawa County, Oklahoma. Late May.

361. False Solomon's seal or false spikenard (Smilacina racemosa)- This forest forb was growing on a moist north slope on top of a bluff in an oak-hickory forest of which sugar maple (Acer saccharum) was the dominant. The understorey woody (= shrub) layer was dominated by flowering dogwood a trunk of which was pictured along the right margin of the photograph. Forbs such as this have little or no feed value although Dayton (1960, p. 23) reported that deer eat the berries of Smilacina species. Their main practical vlaue in Forestry and Range Management is as biotic diversity. Professionals in these natural resource fields are frequently called upon to provide names for conspicuous plant species and questioning laymen are always impressed when rangemen and foresters can spout back the name. This is more so the case for those plants that have little economic value because it shows that professional resource managers know even the minor species, those that are not major lumber or forage and browse plants. Above Modoc Creek, Ottawa County, Oklahoma. May.

362. Inflorescence of false Solomon's seal or Solomon's plumes- This bright flower cluster was on a specimen growing above Modoc Creek in Ottawa County, Oklahoma. May.

363. Plant on a Pineywoods pond- Grassleaf arrowhead (Sagittaria graminea) on a local ponded habitat in natural forest clearing in the Big Thicket of Texas Pineywoods. There are several Saggitaria species in North America. These are in the monocotyledon family, Alismaceae (arrowhead family). Big Thicket National Monument, Hardin County, Texas, April.

364. Flowers and young fruits on a Pineywoods pond- Inflorescence on the specimen of grassleaf arrowhead introduced in the preceding slide. This provided a very good example of an indeterminate inflorescence, one that matures "top-down and outside-in" (flowering begins at apices of inflorescence, or on the upper- and outer-most floral tips, al-most and progresses downward and inward). Big Thicket National Monument, Hardin County, Texas, April.

365. Common pitcherplant or yellow trumpets (Sarracenia alata)- Habit and general appearance of common pitcherplant (also shown as pitcher plant) one of the more unique and distinctive of moist forests in the southeastern portion of the North American deciduous forest formation. This specimen featured was at the edge of a colony of its species on highly acid soil in the Big Thicket part of the Texas-Louisiana Pineywoods.

Big Thicket National Monument, Hardin County, Texas, April.

366. Pitchers in the Pineywoods- Pitcherplants are these largest of Earth's carnivorous plants and are most readily distinguished by "modified leaves that form hollow, water-containing vessels that are adapted to trapping and digesting animal prey"(McPherson, 2007, p.3). These modified, cylinderical leaves that usually form complete enclosures open only at the top are referred to as "pitchers". Hence, the most commonly used common generic name.

The pitcherplant featured here is the species most common in western portions of the southeastern region of the North American deciduous forest formation. It is also typically a generally large and showy species (even when it is out of season for its light lemon-colored corolla). Sarracenia alata is a highly variable species, especially given it rather limited species range. Definitive description of this species (and other species of the pitcher plant family native to the Americas) was McPherson (2007, ps. 195-203). As of this writing, the definitive work on carnivorous plants on ranges and forests of North America is that of Schnell (2002).

Big Thicket National Monument, Hardin County, Texas, April.

367. Fruit pitcher- Shoot apices of the yellow trumpets pitcherplant bearing the leaf-enclosed fruit. The fruit has been interpreted as a loculicidal capsule, which is that dehisces due to or through openings in the locules between partitioning tissue (Smith, 1977, ps. 122, 300). Pitcher plants are in the family Sarraceniaceae.

Big Thicket National Monument, Hardin County, Texas, April.

368. Plant eats bugs- Portion of lower pitcher of yellow trumpets pitcherplant revealing a partly digested insect being used as a source of nitrogen for a plant highly pted to impoverished soils.

Big Thicket National Monument, Hardin County, Texas, April.

Pterophyte is the term for spore-bearing vascular plants. These are "intermediate" (in evolutionary development, taxonomy, etc.) between the traditional units of Byrophyta and the Spermatophyta (seed-bearing plants, usually a taxon at the division level). Traditionally the pterophyte taxonomic level (division) has been called Pteridophyta. All three of these traditional units or taxa were included in the Embryophyta (subkingdom of plants in which the zygote develops into a multicellular embryo while enclosed within the female sex organ or within the embryo sac). In the traditional taxonomic hierarchy the vascular plants (those possessing xylem and phloem including both Pteridophyta and Spermatophyta) were distinguished from the Bryophyta by being placed in the Tracheophyta (the taxon, variously a division or superdivision, of vascular plants). In other words, Tracheophyta minus Pteridophyta leaves Spermatophyta (Gymnospermae and Angiospermae). The Pteridophyta included the club mosses (Lycopocineae), horse-tails or scouring rushes (Equisetineae), and the ferns (Filicineae). Some taxonomic schemes had a taxon designated Pteropsida that grouped (largely on basis of complex, relatively large leaves) the ferns, gymnosperms, and angiosperms.

Spore-bearing plants from the thallophytes through and including the pteriophytes have been called cryptogams (pteridophyts are vascular crypotgams). Plants have thus traditionally been grouped taxonomically in various arrangements depending on what features were of concern for different purposes. In effect, this is a form of taxonomic bilingualism. It is desirable, even essential, in instances where several features are of interest simultaneously. This is often the case when studying vegetation and, even more so, when applying this study to production agriculture. Those individuals are uninformed who view terms and taxa like the ones just discussed as meaningless or useless because they are archaic or, even, obsolete (two different things). If these individuals arrogantly persist in viewing such words as meaningless these folks are stupid (incapable of learning).

There are several taxa of pteridophytes in range and forest vegetation. Pteridophytes are especially common in the various deciduous forest cover types. A few were selected for inclusion in this section devoted to the oak-hickory forest.

369. Colony of smooth scouring rush (Equisetum laevigatum)- Equisetum is, as obvious from the prefix, the genus of pteridophytes given the common name of horsetail. Many of the Equisetum species do not have the namesake horsetail arrangement of primitive leaves encircyling the shoot. Those Equisteum species whose leaves are reduced to rudiments were given the common name of scouring rush, a common name which appeared to aptly describe even those having a "horsetail". In fact, given that most Equisetum species in many locales are "bald" tails, scouring rush is often the more apt common name (genus designation notwithstanding).

Lack of well-developed leaves that have a typical leaf form was a criterion used in classifying ferns as more advanced and closer to spermatophytes than are the club mosses and scouring rushes (ie. ferns are the least primitive of the vascular cryptogams).

Within the oak-hickory forest many of the vascular cryptogams grow best on the more moist habitats.This colony of smooth rush was growing along a high bank of Lost Creek flowing through the extreme western edge of the Ozark Plateau section of the oak-hickory forest. Ottawa County, Oklahoma. July.

370. Shoots of smooth scouring rush- The texture of the shoot and a spore-bearing strobilus of smooth scouring rush were shown in this photograph. Strobilus (plural, strobolii) in this usage refers to a group or unit of sporophylls (including their sporangia) more or less densely encirclying the central sexual axis (the shoot apex in this genus). Sporophylls are modified leaves or leaf-like organs that bear spore-containing sproangia (singluar, sporangium; the case in which spores are formed and stored before release).

Specimen from the stand shown immediately above. Bank of Lost Creek, Ottawa County, Oklahoma. July.

371. Walking fern (Camptosorus rhizophyllus= Asplenium rhizophyllum)- This unique (and rare) fern was growing on a moss-covered limestone bluff above a creek in the western oak-hickory forest of the Ozark Plateau. The common name of this species comes from it's form of asexual reproduction. The pointed tip of the frond (the leaf of a fern) often roots and produces a new daughter unit (a module or ramet) which, upon complete development, can repeat this pattern of propagation. This phenomenon was "going hog-wild" in the specimen shown here. Walking fern also reproduces sexually as do other ferns by producing and releasing spores from sori (singular, sorus; the clusters of sproangia in ferns) on the undersides of their fronds.

Fern was growing amidst or a "carpet" of the gametophytic generation of a "true moss" (see slides below).

On a limestone bluff above Modoc Creek, Ottawa County, Oklahoma. March.

372. Sori on the underside of a walking fern frond- Close-up of the vertically oriented frond of the walking fern seen in the preceding slide (right side of plant). Limestone bluff above Modoc Creek, Ottawa County, Oklahoma. March.

373. Colony of rattlesnake fern (Botrychium virginianum)- A high proportion of forest pteridophytes require relative moist (= mesic) soils (ie. they are at the "higher end" of mesphytes). One exception is rattlesnake fern which grows on soils having average quantities of soil water. The general common name for Botychinum species is grape fern, but B. virginanum is usually called rattlesnake fern. This colony of rattlesnake fern was thriving on a microsite that had a deep layer of rotting oak and hickory leaves in a second-growth oak-hickory forest along the extreme western edge of the Ozark Plateau.

Newton County, Missouri. May.

374. Shoot of rattlesnake fern- An example of the large and intricately patterned leaves and the sori-bearing sporophyll of rattlesnake fern were "captured" in this slide. The sporophyll is the spore-bearing leaf of the pteridophytes. Sori (plural of sorus) are the "fruit structures" (typically borne on undersides of leaves) which bear numbers (usually large numbers of) spores. Newton County, Missouri. May.

375. Christmas fern (Polystichum acrostichoides)- This fern allegedly got it's common name from the fact that it is characteristically green (and often producing new fronds) in the middle of winter (ie. evergreen fronds). Christmas fern prefers moist (but typically well-drained) habitats especially along banks of shaded, flowing streams. This specimen was one of numerous plants of this species growing along the bank of Modoc Creek in a bottomland forest of sycamore, sugar maple, and box elder (Acer negundo), but here beneath huge black oaks. Beaver (Castor canadensis) had cut off many of the fronds and carried them into their lodges which were burrows in the creek bank (ie. bank beaver). What use beaver made of them was not be determined.

Ottawa County, Oklahoma. January.

376. Put-together morphology- Details of frond of Christmas fern shown in a three-slide sequence from general morphology of frond blade (first slide)to details of ets or secondary leaves (second and third slides) of the frond. Leaflets are alsso known as pinnae (pinna, singluar). Students were referred to Diggs et al. 91999, p. 178) and standard Botany texts such as Weoer et al. (1982, ps. 586-589) for explanations of morphological structure of fern fronds, rhizomes, and roots.

Christmas fern is aptly named (in the northern hemisphere) as it thrives in winter when it gets the greatest amount of sunlight with leaves shed from trees and shrubs.

Bank of Modoc Creek, Ottawa county, Oklahoma. Late December.

377. Maidenhair fern (Adiantum pedatum)- This is perhaps one of the most graceful of all forest forbs. This was one of several individuals of this species that grew in close proximity to the Christmas fern shown in the preceding slide. Maidenhair ferns grew several feet higher up (on a bluff) from the Christmas ferns on the bank of a slow-moving stream in the far-western Ozark Plateau. Unlike the evergreen Christmas fern, maidenhair ferns were dormant and died back to the ground surface in winter. The maidenhair ferns shown in this slide were were growing in late summer in dense shade and on a moist east slope. They received less than three hours of direct sunlight during the longest days and had to survive on that and/or what diffuse light "filtered" through the leafy overstorey of black oak and sugar maple.

On a limestone bluff above Modoc Creek, Ottawa County, Oklahoma. August.

Bryophytes are those range and forest nonvascular plants of the general groups of mosses, hornworts, and liverworts. (Recall that "plants" was used herein as in the older or more traditional usage of the two kingdoms of organisms that the author viewed as more practical for discussion of vegetation.). More specifically bryophytes are members of the Bryophyta (a division of plants in the Plantae); in short, they are nonvascular spore-producing plants. Bryophytes have alternation of generations (gametophyte and sporophyte), but in contrast to the vascular plants the dominant and conspicuous generation of bryophytes (ie. the so-called plant) is the gametophyte (gametophytic generation). While bryophytes lack differentiation into true roots they have root-like structures called rhizoids which anchor the gametophytes to their substrate (bryophyte rhizoids do not function in absorption).

Bryophytes, like thallophytes (thallus-- the type of plant body that is undifferentiated into root and shoot-- plants such as algae, bacteria, and fungi), are primarily important in Range Management and Forestry as reducers or decomposers and therefore in processes like soil formation and plant succession (ie. the Clementsian process of reaction now more commonly known as facilitation). This was discussed above immediately before presentation of fungus species.

Raven et al. (1992, ps. 298-316) provided an excellent introductory discussion of the Bryophyta. Shaw and Goffinet (2000) wrote a comprehensive and the recent classic text on the Bryophyta for the "really serious" student.

378. Gametophye of a "true moss" (order of Bryales)- "Moss" is one of the most confusing and most misleading names in all of Botany. "Moss" is applied to every sort of "plant" from reindeer moss (a lichen eaten by caribou and reindeer) to Spanish moss (a monocotyledonous epiphyte often eaten by native and domestic ruminants). Obviously such "mosses" are range plants by definition because they are native plants that serve as feed for grazing/browsing animals. Absent this axiomatic and self-evident definition, these and other "plants" growing on range are still range plants even if they function in less conspicuous roles such as decomposition, soil formation, nutrient cycling, and plant succession. True mossses provided such examples of these roles or functions in forest and range ecosystems.

"True mosses" has been the designation traditionally used for the one (of three) class in the division of Bryophyta know as Musci. In some of the more recent works (eg. Raven et al., 1992, p. 308) Bryophyta division still consist of three classes one of which is that of the true mosses but known as Bryidae (instead of Musci). The Musci or Bryidae bryophytes are partially saprophytic being dependent on decaying organic mattter. The specimen photographed here (the species of which was not identified ) was attached to a piece of chert that had a deep facet which had acumulated rotting oak leaves. Oak-hickory forest in Ozark Plateau. Ottawa County, Oklahoma. January.

379. Gametophytic and sporophytic generations of a species of true moss (Funaria hygrometrica), Musci- In these two macrolense shots the leafy gameotphyte is visible at the base of the moss while the mature sporophytes are very prominent as the apex of the moss body. These shown here were female sporophytes known as archegonia (singlular is archegonium) or archegonial heads which consist of a capsule (= sporangium; plural is sporangia) that are borne on a seta (stalk). The first of these two slides presented a view from the top to show the overall appearance of moss in the sporophytic stage.

The second slide was a side-view of the moss which showed more clearly both the leafy gametophytic generation and the stalked capsule of the sporophytic generation. The covering of the capsule is the calyptra the sharp, pointed tip of which is the operculum. This is the lid of the capsule which when shed opens up the capsule, the inside of which contains spores which are released upon this opening. While these are not vascular plants and are not differentiated into roots, stems, and leaves there is some differentiation of tissues. This occurs in the stalk as well as in the leafy gametophyte.

At the base of this sectional sample of moss are the rhizoids which are the root-like structures (more like root-hairs actually) of the gametophytic generation that function only to anchor or hold the plants (absorption of water and mineral nutrients occurs directly through the gametophyte).

Oak-hickory forest in Ozark Plateau. Ottawa County, Oklahoma. March.

380. Colony of juniper hair cap moss (Polytrichum juniperinum)- This well-developed stand was on the floor of a post oak flats site of the oak-hickory forest in the Ozark Plateau. Post oak flatwoods is a very mesic form of the oak-dominated deciduous forest. It is not a swamp or any form of wetland, but it is poorly drained flatland of primarily clay soils. In the Ozark Mountains and adjoining savanna of the Prairie Peninsula post oak is usually the only species that can survive the poor drainage of this site (hence the name of post oak flats). The high moisture condition of the soil and shade from post oaks provided a microhabitat (microsite) satisfactory for juniper hair moss.

Newton County, Missouri. April.

381. Close-up of juniper hiar cap moss- This is the gametophyte (gametophytic generation) of this species. Newton County, Missouri. April.

The relatively high primary productivity of the oak-hickory forest results in heavy accumulations of detritis ("yields" of necromass) that are rich substrata for many kinds of reducers (= decomposers) among forest and range plants. (The author acknowledged the value and legitimacy of the five kingdom taxonomic system, but herein used "plants" in the traditional botanical usage for simplicity, utility, and practical application.) Botanical organisms functioning as reducers extend from bacteria, algae, fungi, lichens, and bryophytes to vascular plants. A sample of these was included below beginning with the fungi. Not all fungi are saprophytes. Some fungi are saprohytic, of course, but others form mycorrhiza or the mycorrhizal symbiosis with roots of vascular plants while others are parasitic (on vascular plants, insects, or even othr fungi). Some fungus species are a combination of saprophyte and parasite (examples were presented below). Likewise, not all saprophytes are fungi or bryophytes. An example of a saprophytic vascular plant (a dicotyledon) was presented below following presentation of several fungal species.

Fungi are one of the three major groups of organisms traditionally regarded as plants (or plantlike taxa) that lack vascular tissue (ie. simplistically defined as nonvascular, undifferentiated lower plants). These "plants" (plantlike organisms) were historically regarded as thallus plants or thallophytes, members of the Thallophyta (usually listed as a subkingdom). Thallophytes are those organisms having plant bodies not differentiated into roots and shoots, lacking vascular tissue, and having gametes enclosed only by a cell wall such that their zygotes do not develop into embryos while inside the female sex organ (Wilson et al., 1971, ps. 447-451). With acceptance of the five kingdom classification system of organisms proposed by such stellar scientists as Robert H. Whittaker the older or traditional two kingdom scheme fell into disuse. While this development was generally an improvement (especially for the teaching of Biology to beginners) some of the taxonomic groupings (eg. thallophyte) still make sense especially for applied use in Agriculture. The author of this web publication periodically resorted to groups like the Thallophyta for practical use (ie. ease of teaching cencepts in conservation, agricultural production, etc.).

Thallophytes and, as discussed below, bryophytes are essential to development of vegetation and soils. These lower plants typically form one or more layers of vegetation. Also, they are usually pioneer species. These primitive plants are among the first organisms to grow on the raw parent material of a sere such that they improve the habitat for higher plants of later seral communities. This was what Clements termed reaction, one of the processes in development of vegetation (Clements, 1916a, 79-96 passim; Weaver and Clements, 1938, ps. 234-241). Years later this process was termed facilitation (Connell and Slatyer, 1977) and in time became known as the facilitation model of plant succession (Begon et al., 1990, ps. 632-633, 635, 641-642). Clements' specific view (almost as much philosophical as theoretical) was that "[e]ach climax formation had its individual or ontogenetic development"... such that "it shows a phylogenetic development from a preceding climax or community". "If the phylogeny of the community comprises the same general process as that of the species, it should be recapitulated by the ontogeny as seen in the sere" (Clements, 1916a, ps. 344-345).

An assumed phylogeny beginning with lower plants such as algae, liverworts and mosses, lichens, etc. (as determined from the fossil record) implied-- at least to Clements-- that development of vegetation on current seres began with these thallophytes and bryophytes. This recapitulation component of Clements' grand theory of plant succession was probably the least understood part of his complicated model of vegetation development, but undoubtedly all students of plant succession have observed that some of the earliest species to pioneer an area (especially a prisere, a fresh or newly created bare area of parent material or the area having to undergo primary succession) are the primitive plant forms. Whatever processes and paths of development are involved, the thallophytes and bryophytes do facilitate development of range and forest plant communities by the processes of plant succession. It was for this reason that a few of these species were included in this publication on range and forest cover types.

There are numerous ouytstanding field guides to the fungi (ie. mushrooms) of North America including Krieger (1967), Orr and Orr (1979), Lincoff (1981), Pacioni (1981), McKnight and McKnight (1987), Metzler and Metzler (1992), and the massive Arora (1986). Mycology is the study of fungi. There are numerous outstanding texts and references for the fungi. Pritchard and Bradt (1984) was recommended for readability yet thoroughness. Carlile and Watkinson (1994) was a comprehensive text that covered basic biology, including ecology, evolution, genetics, etc., but for overall, concise reference (especially for beginners) the basic Botany textbook of Raven et al. (1992, ps. 208-243) seemed easiest to use.

382. Morel or yellow morel (Morchella esculenta= M. rotunda)- This fruiting (=fruit) body (reproductive structure; in higher fungi the fruit body is sometimes designated the carpophore) of this fungus is delicious and a delicacy to hillbillies who seek it out in the leafy forest floors of such ancient mountains as the Appalachians and Ozarks. Morchella species are ascomycetes (Ascomycotina is the largest subdivision of true or fleshy fungi) within the Discomycetes class. There are several species of morels across North America but the species presented here is the most common one in the oak-hickory forest of the Ozark Mountains.

Banks of Modoc Creek, Ottawa County, Oklahoma. April.

383. Ink cap (Coprinus radians)- On this rotting log covered with decomposing oak leaves is the complete body of a fungus in the group, basidiomycetes (Basidiomycotina is a subdivision of the higher, fleshy, or true fungi), more specifically the Hymenomycetes class therein. The species shown here is one of the best or standard textbook examples of the true fungi, those species that produce fleshy fruit-bodies know generically as mushrooms and/or toadstools. The two general parts of a mushroom are 1) the filaments of hyphae which form a network known collectively as the mycelium or spawn and 2) the fruiting body, often called the carpophore, which bears the reproductive structures that produce spores. The fruit-body is the obvious, often conspicuous, part of the true or higher fungi (ie. the "mushroom") which forms from the mycelium. The latter part of the true fungi are not differentiated into roots and shoots and are typically not seen by human eyes. The only part of fungi usually visible (again, to humans) is the fruit-body, the mushroom. If some of the mycelium (hyphae filaments) remain attached to the carpophore when someone picks or knocks over the mushroom he would naturally think of the mycelium as the "roots" which is obviously incorrect.

C. radians is unusual in having bright orange hyphae threads and, when these are accompanied by the carpophore (as shown here), students have the rare opportunity to see the whole "plant" (entire fungus body). In this slide there is one live or "fresh" and one dead (withered) carpophore. Some fungal species have a carpophore or mushroom that includes a prominent "cap" as its hymenium, the spore-bearing surface of the Ascoymetes and Basidiomycetes. In C. radians the spore-bearing underside of the cap is dark-colored hence the common name of ink cap (next slide). Obviously this fungus is a saprophyte. It was shown to offer students a classic example of a saprophyte or saprophytic "plant" which is one of the major categories within the reducer or decomposer group of organisms in forest and range ecosystems.

First terrace of Modoc Creek, Ottawa County, Oklahoma. April.

384. Detail of the carpophore of ink cap and surrounded by the mycelium- The underside of the cap of Coprinus radians showing the gills that bear the spores of this species being supported or held by the stipe, the stalk or "stem" of the carpophore that bears the hymenium (in this species the latter developes as a cap). The substrate was a rotting oak log.

Bank of Modoc Creek, Ottawa County, Oklahoma. April.

385. Close-up of the hyphae filaments comprising the mycelium of ink cap- This is the vegetative (non-fruit-body portion) of a fleshy or true fungus growing on a decaying oak log. An individual filament is a hypha (hyphae is the plural). The many hyphae form the mycelium which in this species is a brilliant orange. It was photographed in Kodochrome (hence without color-enhancement). Modoc Creek, Ottawa County, Oklahoma. April.

386. Wood ear or tree ear (Auriculularia auricula) on dead elm (Ulmus americana) limb- In the first slide, almost indistinguishable on this dead limb, are dry fruit-bodies (accompanied by equally dried-out lichens) of a species of the Hymenomycetes (class) of the Basidomycetes (subdivision) known by the colorful (and most appropriate) common name of wood ear.

In the second slide, which had been taken 24 hours earlier when the fruit-bodies (the "mushrooms") were conspicuous, wood ear carpophores (and accompanying lichens) had swollen from a recent rain.

This saprophytic species is but one of many fungi that quickly decompose the vast detritis or necromass (dead organic matter that was previously biomass) produced in the oak-hickory forest. This species of Hymenomycetes was growing in the "dead of winter", a common season for appearance of many carpophores due to abundance of water in detritis as a result of low evaporation rates.

Ottawa County, Oklahoma. December.

387. Wood or tree ear (Auriculularia auricula) capophores fully hydrated- Close-up shot of wood ear immediately after a warm winter rain. Ottawa County, Oklahoma. December.

388. Wood ear fruit-body drying out- Appearance of the member of Hymenomycetes known as wood ear about six or seven hours after cessation of winter rain. The capophore was quickly returning to the more common dehydrated state as shown above. Ottawa County, Oklahoma. December.

389. Tooth fungus, old man's baerd fungus, or lion's mane fungus (Hydnum erinaceus= Hericium erinaceus) growing on dead wood of a fire scar- An old fire scar of black oak (Quercus velutina) in an oak-hickory forest of the Ozark Plateau was the substrate for this primarily saprophytic fungus. This fungal species (and the next one in this line-up) can be viewed as partly parasitic because it frequently grows on wounds of living deciduous trees as well as on totally dead wood. Like the preceding species this fungus is in the Hymenomycetes class of the Basidiomycetes subdivision. (As in the case for all groups of organisms there are different taxonomic schemes or arrangements and names for the hierarchial levels such that the class level may be a subclass in the hierarchy of another author). The fruit-body of H. erinaceus is edible when young, but it takes a long to cook the tough tissue.

Newton County, Missouri. December.

390. Fruit-body of a tooth or comb fungus (H. erinaceus)- Detail of the carpophore on the fire scar of the preceding slide. The fruiting body of members of the Basidiomycetes has been designated the basidiocarp The basidocarp bears the basidia (singlular is basidium) which are the individual microscopic, spore-producing cells. H. erinaceus is a saprophyte, but it is also somewhat parasitic hastening the rotting of live wood adjacent to dead wood that is the main substrate.

Oak-hickory in Ozark Plateau. Newton County, Missouri. December.

391. Basidiocarps of oak conk or cracked cap polypore (Fomes robustus= Phellinus robustus) on an ancient black oak- The trunk of this old black oak that had survived for decades on a rockpile of a site "sported" several carpophores (fruiting bodies which are basidocarps in this class of fungi) of this member of the Hymenomycetes (in the Basidiomycetes class). Oak conk is a shelf or cork fungus within the group known as polypores and/or bracket fungi. Polypore refers to the many pores (more precisely, spore-producing tubes) on the underside of the cap (carpophore, specifically the basidiocarp).The polypores or shelf mushrooms (conks are but one category of these bracket fungi) are in the general group viewed as true or fleshy fungi.

Oak-hickory forest in Ozark Plateau. Ottawa County, Oklahoma. April.

392. Oak conk- The basidiocarp of a woody shelf fungus showing the underside with the layer of tissue from which the spores are released from the numerous spore-producing tubes. The typical cracked cap of the polypores (family: Polyporaceae) was also clear in this photograph. This was one of numerous carpophores on the trunk of the old black oak shown in the preceding slide.The shelf- or bracket-like basidiocarps of this and related species are just as woody as the substrata they grow on.

Polypores have generally been viewed as more saprophytic than parasitic, but most species function in both capaacities. Arora (1986, p. 549) pointed out that the mushrooms of this group (Polyporaceae and related genera) "are absolutely indispensable to the forests of this world" in their function as decomposers. These are the major "wood-rotting fungi" causing 90% of the rotten-wood damage to standing timber in North America. This fact could be viewed as indicating a major forest pathology crisis, and at one time foresters who interpreted Forestry as mere tree farming (and thus sought to eliminate all "enemies of the forest") may have held such views. That was before application of the ecosystem concept brought back a more balanced perspective or maybe just some old-fashioned woods wisdom in new terms for original principles. Today's foresters and rangemen appreciate the essential role of decomposers in reduction of detritis (plus the ecological role of detritis itself), cycling and availability of nutrients, and provision of food for consumers (both vertebrates and invertebrates). Arora (1986, p. 549) stated the case for the polypore fungi and put the role of decomposition in proper perspective. "Without them there would be no logging industy in the first place: every cut stump, felled log, and lopped-off limb would be indefinitely on the forest floor, the woods would quickly become impenetrable, and the new trees would have neither room nor nutrients to grow".

Oak-hickory forest in Ozark Plateau. Ottawa County, Oklahoma. April.

393. Sulphur shelf, chicken mushroom, or chicken of the woods (Polyporus sulphureus= Grifola sulphurea= Laetiporus sulphureus)- This colorful member of the Polyporaceae is edible (when young) and so distinctive that it is not likely to be confused or misidentified). Sulphur shelf is also both saprophytic and parasitic. Krieger (1967, p. 157) labeled the chicken mushroom as one of "the wood-destroying fungi" and "a most persistent enemy of coniferous as well as deciduous trees". He listed oaks, locusts, maples, alders, walnuts, pines, hemlocks, and spruces among it's host species. The fine specimen shown here was growing on an old fire scar at the base of a black oak in the Ozark Plateau.

Ottawa County, Oklahoma. October.

394. Pinesap (Monotropa hypopithys)- This saprophyte is a vascular plant included by various treatments in such families as wintergreen (Pyrolaceae), Indian pipe (Monotropaceae), or as a separate subfamily (Monotropoideae) of the heath (Ericaceae) family. Color alone (absence of chlorophyll) indicated that this dicotylendous species cannot carry out photosynthesis and must be either parasitic or saprophytic. It is the latter by means of forming mutualism with mycorrhiza on its roots. This specimen was growing on rotting litter on the floor of a black-oak dominated oak-hickory forest in the Ozark Plateau. The dead stems from the previous growing season were photographic evidence of the perennial length of this species' life cycle.

Ottawa County, Oklahoma. June.

395. Pinesap- Close-up photograph of the vascular saprophyte presented in the preceding slide. This specimen had less pubescence than is typical for this species. It was growing on the organic matter of rotting oak and hickory leaves in the western part of the Ozark Plateau. Ottawa County, Oklahoma. June.

396. Interior of an upland mixed oak (white, red, post, black)-hickory forest that has extremely lush and diverse shrub and herb layers. Understory plants include tick clover, blackberry, grape vine, Virginia creeper, poison oak (Rhus toxicodendron= Toxicodendron toxicarium), bluestem, and panic grasses. Second growth forest with excellent regeneration of climax tree species, especially hickories. Note that this excellent botanical diversity and forest regeneration is occurring under light or conservative grazing/browsing by cattle (Herefords are visible in center).

Adair County, Oklahoma. May, late vernal aspect. FRES No. 15 (Oak-Hickory Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Dry-mesic chert forest. Oak-Hickory Series of Brown et al. (1998).

397. Uneven-aged management in upland oak-hickory forest- Selective cutting (= selective felling) in an Ozark Plateau chert forest dominated by black oak which was harvested for veneer to be used in making church furniture. Trees were felled in winter and this is the scene three months after logging. Most harvested trees were 70-90 years old. Note total absence of any soil disturbance or damage to young trees. Logs were carried— not drug — out by a rubber-tired skidder. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998).

Selective cutting like clearcutting has both proper and improper applications. The benefits of proper selection-cutting (= selection felling) to maintain forest with all, or most all, age classes with minimal adverse impacts on the forest resouces is forest practice par excellence. Unfortunately, all too often selective cutting is not selection cutting (= the selection method) but simply amounts to high-grading, "the removal of the most commercially valuable trees (high-grade trees), often leaving a residual stand composed of trees of poor condition or species composition" Selective cutting is "a cutting that removes only a portion of trees in a stand… it is a general term that should not be confused with cutting done in accordance with the selection method" (Helms, 1998). Selection method (= selection felling) was defined under slides of the Sierran Mixed Conifer Type. Even-aged and uneven-aged management or regeneration methods are only one part of silviculture which in turn is but one component of forest management.

"Forest Management- the practical application of biological, physical, quantitative,
managerial, economic, social, and policy principles to the regeneration,
management, utilization and conservation of forests to meet specified goals
and objectives while maintaining the productivity of the forest" (Helms, 1998).

Although the selective harvest seen here was not conducted according to a forest harvest plan (including harvest scheduling) or the guidelines of scientific forest management, the intensity and frequency of cutting was so low as to have minimal, if any, detrimental effects on the forest. Acorn production is so great as to overwhelm feed demands of wildlife and assure natural regeneration by sexual reproduction. Plus there is asexual or vegetative reproduction by:

398. Stump sprouting (= coppicing) of young black oak- These are stump sprouts at the near end of the first growing season post-logging on the black oak-dominated chert upland oak-hickory in the Ozark Highlands seen in the preceding slide. Stump sprouts like these from the root collar or basal trunk arise from dormant buds whereas stool sprouts arise from adventitious buds between bark and wood. The latter are short-lived and of no value for regeneration (Kramer and Kozlowski, 1979, ps. 61, 150-151). Stump sprouting among haradwoods varies due to many factors including species, age of felled tree, and season of felling. Generally, younger trees sprout better than older ones because the bark is thicker in older trees and the dormant bud may not be able to emerge through it.In addition, older trees are more likely to have connections between bud and pith interrupted (Smith, 1986, p. 471). Sprouting is most abundant and vigerous when trees are cut during dormacy (Kramer and Kozlowski, 1979, p. 272). Sprouts that arise at the basal part of a stump are superior to those growing from the top of a stump because they are stronger and less apt to break off (Smith, 1986, p. 472). Cutting low stumps like the ones seen in these two slides is a trademark of good forest harvest practice. It is the mark of sloth and sloppiness to leave high stumps with trees of this size and straight boles with little butt swell.

Ottawa County, Oklahoma. August. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

399. Healthy low stump sprouts of a young black oak- Regeneration of certain hardwood species like oak by coppicing is a sound regeneration method. Combined with uneven-aged management it provides a minimum of impact on the forest ecosystem and wildlife habitat. There are times when clearcutting or even-aged management is also beneficial. It may result in faster regeneration, greater wood yield and improve habitat for deer through greater browse production. It can also release climax tallgrass species and produce more forage for livestock and wildlife as shown in the next slide.

Ottawa County, Oklahoma. Estival aspect, September. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

400. Release of big bluestem the first summer following selective cutting in an oak-hickory forest- This is a fine colony of big bluestem (appropriately called "timber grass" by local hillbillies) at anthesis at the end of the first growing seasonfollowing the selective harvest (eight months post-logging) seen in the last three slides. This stand of the native decreaser grass was released from growth-limiting shade along a skid trail. Other such examples of release occurred throughout this chert upland black oak-dominated forest in the Ozark Plateau. It is a textbook example of transitory forest range (as soon as the crowns of oak and hickory trees fill in the openings the grass and forbs will again be suppressed). This demonstrates how Forestry and Range Manageament often go hand-in-glove.

Organization Note: More examples of forestry practices (including harvest and regeneration) in the oak-hickory types were presented in the chapter, Oak-Hickory Forests-II (under the section, Use and Abuse of Oak-Hickory Forests).

Bottomland Forests

401. Bottomland hardwood forest on north slope in Ozark Plateau— Extremely species-diverse community shown here in early spring. Species include sycamore, extreme right foreground; western hackberry, foremost trunk; black walnut (Juglans nigra), two trees immediately behind preceding foreground trees; chinkapin oak, grey trunk just behind walnuts; sugar maple (Acer saccharum), two black trunks forming V-shape in background and one behind and partly concealed by the sycamore; basswood or American linden (Tilia americana), on upslope to left of maples; and white ash (Fraxinus americana), left-center midslope. Redbud and flowering dogwood are in the shrub layer but the dominant shrub is spicebush (Lindera benzoin) which comprises a lower shrub layer.The herbaceous layer, which is clearly visible in this early spring stage, includes bloodroot (Sanguinaria canadensis), trout lily or yellow dog-tooth violet (Erythronium americanum), broadleaf waterleaf (Hydrophyllum canadense), rue anemone (Anemonella thalictroides), false rue anemone (Isopyrum biternatum), and scattered colonies of the ever-present Mayapple. Following completion of their annual growing season these species are replaced in their location by great Indian plantain (Cacalia muhlenbergii) which then dominates a tall herb layer through mid-summer. This is a unique transect view going from the first terrace of a mid-size creek to the top of limestone bluffs. The upper-most forest is dominated by black and red oaks as shown in preceding slides. Downslope from the upland oak forest is a mid-slope sugar maple-basswood-chinkapin oak zone. The bottomland or floodplain forest is the sycamore-hackberry-black walnut-white ash forest. Thus this transect includes SAF cover types 61 (River Birch-Sycamore, variant form) or 94 (Sycamore-Sweetgum-American Elm, variant of), 26 (Sugar Maple-Basswood), and 52 (White Oak-Black Oak-Northern Red Oak), the latter is indistinguishable in background.

Ottawa County, Oklahoma (bank and bluffs of Lost Creek). March, early vernal aspect. Mesic bottomland forest according to Missouri Natural Areas Committee (1987). FRES No. 15 (Oak-Hickory Forest). No obvious Kuchler unit. Mixed Hardwood Series in Southeastern Swamp and Riparian Forest biotic community of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

402. Methuselah blooming- One of the few remaining live branches on the more-dead-than-alive, ancient black walnut introduced immediately above, and it was in early stages of sexual reproduction. With backcrop of an azure-blue sky (right after a late norther) this sexual leader might well have been this ancient tree's last attempt at fruit production. These catkins of the monoecious black walnut were all males (closer views immeidately below).

South bank of Bosque River, Erath County, Texas. Mid-April; blooming of black walnut.

403. Methuselah details- View of several catkins and a few new leaves (first slide) and close-up of one of these catkins (second slide) on the leader of the ancient black walnut presented above. Black walnut flowers are monoecious; these long, hanging catkins were males. All flowers on this leader were male.

South bank of Bosque River, Erath County, Texas. Mid-April; blooming of black walnut.

404. Bearing fruit in the deep woods- Branch of black walnut with fruit at approximate stage of mid-maturity and with the compound leaves characteristic of this marvelous nut-bearing North American hardwood. Black walnut is the single most valuable hardwood lumber species in the United States. The quality wood of this species is unsurpassed for particular kinds of furniture uses and, even more for its beauty, strength, durability, and light-weight features for gunstocks. Black walnut is equally prized for the eating quality of its nuts the flavor of which to palates of many consumers exceeds the strength and delightful aftertaste of all other North America nuts with exception of the now-extinct American chestnut.

Springfield Plateau of the Ozarks, Ottawa County, Oklahoma. July.

405. Green or unripe fruit of black walnut- Even the odor of walnut hulls is distinctive. To those hillbillies whose boyhoods included the autumn ritual of walut-picking this was a never-to-beforgotten, pleasant aroma. Even the persistent stain from the hulls was a much milder form of distraction than wounds left from ticks, chiggers, black berry briars, and other summertime perils. Somehow, no other nut so flavors ice cream as the strong, woodsy taste of black walnut (especially when accompanied with pleasant memories of cool fall days spent beneath native walnuts in the eastern deciduous woods).

Interesting note: the difference in color between foliage and fruit covering of these two slides was due to differences in film (not the usual culprit, an Epson Perfection 600 printer). The darker green color in the lower slide was the more nearly correct tone of green. It was slide film of what else but the "gold standard"; Kodachrome 64. Nothing reproduced color of plants like Kodachrome.

Springfield Plateau of the Ozarks, Ottawa County, Oklahoma. July.

406. Start of a black walnut- Young seedling of black walnut that was growing in the Springfield Plateau of northeastern Oklahoma. Seedlings such as this specimen grow very rapidly due to considerable storage of nutrients in the richly flavorable (to humans and squirrels) and extremely hard-shelled nut.

Ottawa County, Oklahoma. Mid-June; early seedling stage of pnehology.

407. How it starts in the soil- Three views at progrsssively closer camera distance of the basal part of a recently germinated black walnut seedling (the one presented in the immediately preceding photograph) in the western Ozark (Springfield) Plateau orf northeastern Oklahoma. This is near the far western edge of the oak-hickory forest (as distinguished from the Cross Timbers form oak-hickory-tallgrass savanna).

The "meat" of the delicious black walnut fruit is primarily the two cotyledons which serve as the stored nutrients for the seedling during germination and emergence.

Ottawa County, Oklahoma. Mid-June; early seedling stage of pnehology.

That chapter also has other bottomland forest types or forms as well.

Pecan Bottoms- Example of Southern Bottomland Forests

The following section treated bottomland forests dominated by pecan (Carya illinonensis= Hickoria pecan) to furnish an example of a specific form or subtype of either Sugarberry-American Elm-Green Ash (SAF 93) or Sycamore-Sweetgum-American Elm (SAF 94). The latter forest cover type (SAF 94) had previously been designated as Sycamore-Pecan-American Elm (Erye, 1980, ps. 65-66).

More detailed coverage of these forest range cover types was included with chapters, Miscellaneous Forests and Southern and Central Forests-II

408. Bottomland forest (actually more of a woodland physiogonomy as crowns overlap but slightly) of pecan (Carya illinoinenisis) with an understory devoid of shrub layers and with the herb layer dominated by Canada wildrye with frostweed (Verbesina virginica) an important associate. Indiangrass and eastern gamagrass add a tallgrass element. In the upper two post oaks (left center) entered this otherwise single-species stand or pecan consociation. In the second photograph two large pecans (only their lower trunks showing) shaded a carpet of the two cool-season perennial native grasses, Canada wildrye Texas wintergrass, while warm-season perennials like little bluestem, Indiangrass, and perennial dropseeds "waited in the wings" for their time on the shaded stage. This tree-dominated vegetation was an irregular gallery forest along the South Llano River. The pecan is the stately State Tree of Texas and the community shown here is known as "pecan bottoms" by locals who frequent it come nut season. Another beautiful permanent deciduous forest range type.

Kimble County, Texas. June, estival aspect. An "island" or isolated part of FRES No. 15 (Oak-Hickory Forest Ecosystem) and K-91 (Oak-Hickory Forest). No SAF designation readily fit this range vegetation. This forest range appeared similar to the Cottonwood and Live oak Bottomland Types (SAF 63 and SAF 89), but most likely a Variant SAF 94 (Sycamore-Sweetgum-American Elm) that had the former SAF designation of Sycamore-Pecan-American Elm. Edwards Plateau- Edwards Plateau Woodland Ecoregion, 30a (Griffith eat al., 2004).

409. Texas-size- Though it was hard to show with all the shade, several widely scattered giant pecans formed a pecan bottom much like those that existed in Texas river bottoms before European man presumptuously assumed that he could improve things. For big trees like our "centerfold hero" here stocking rate has be relatively low. Prof. Benton Storey (Texas A&M Universty) felt that production of big pecan trees and high nut yields were best obtained with only one tree per acre. Pecan bottoms would thus be savannahs or, at most, woodlands because their crowns would not contact each other at such stocking.

Studded T steel post serving as stakes for pecan seedlings and saplings and the top of one post by the pecan-picker's "centerfold" provided scale for size comparison. This part of the understorey of this river bottom woodland had been mowed (perhaps in anticipation of nut season).

410. East Texas pecan bottoms- Bottomland mixed forest in the Texas Pineywoods dominated by pecan with hackberry and water oak as associates. Foremost tree at left margin was a large water oak with a single bole of high-quality timber. The large tree at right-of-center with numerous major limbs and scaley, brown bark was pecan as were most of the trees (pole-size) in background. The smallest of three trunks in foreground (in front of and aligned along left edge of pecan was sugarberry (Celtis laevigata). Flowering dogwood (center green shrub) formed an interupted upper shrub layer. The invasive alien Japanese honeysuckle (Lonicera japonica), seen here as the green plant in left to center foreground, comprised a spotty though dense lower shrub layer. The exotic and now widely naturalized Japanese honeysuckle (a horticultural escape "gone wild") is a major noxious plant problem in forest understories, especially in pecan bottoms. There were only a few grasses (panicgrasses were most obvious), grasslike plants (limited to Carex spp.), or forbs in the understorey of this bottomland forest. The alien L. janpoica appeared to have crowded out herbaceous plants.Species of green briar and grape along with rattan vine or Alabama supplejacck formed a "jungle" of vines among the smaller pecans in bacdground. Local mowing at edge of this forest near a campground had apparently effectively controlled woody vines in this sample of lowland Pineywoods forest.

Davy Crockett National Forest, Houston County, Texas. March, vernal aspect (post dogwood blooming stage ). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). Variant SAF 94 (Sycamore-Sweetgum-American Elm) that had former SAF designation of Sycamore-Pecan-American Elm. Closest biotic community designation of Brown et al. (1998, ps. 37, 38) was Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1, but there should have been a Oak-Hickory Series for Southeastern Deciduous and Evergreen Forest 123.. South Central Plains- Southern Tertiary Uplands Ecoregion, 35e (Griffith et al., 2004).

411. Another example of a pecan bottom- This example of bottomland forest dominated by pecan was along the small slow-moving Alarm Creek on extremely fertile alluvium. The four big trunks in right foreground and center midground were pecan. Other trees included both post and blackjack oaks. Hackberry was also well-distributed. Greenbriar was the dominant shrub. There was a well-developd herbaceous layer of Canada wildrye, little bluestem, and sand lovergrass (Eragrostis trichoides). Small isolated patches of giant cane (Arundinaria gigantea) grew along banks of the creek.

Erath County, Texas. April. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-75 (Cross Timbers). Variant SAF 94 (Sycamore-Sweetgum-American Elm) that had former SAF designation of Sycamore-Pecan-American Elm. (Griffith et al., 2004). Closest biotic community designation of Brown et al. (1998, ps. 37, 38) was Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1, or more descriptively perhaps, a transition of this Oak-Hickory Series and Bluestem "Tall-grass" Series, 142.11, Plains Grassland 142.1 Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

412. Young pecans and tallgrasses in summer- Part of the floodplain of the Bosque River in the Western Cross Timbers supported this stand of "half-growed" young pecans with bur oaks of the approximate same age cohort as associate tree species. Understorey was locally dominated by Canada wildrye, especially prominent in the second of these photographs where that cool-season member of the Hordeae or Tritaceae tribe was taller than the top wire of the fence enclosing this nice sample of "pecan bottoms". The associate herbaceous species varied locally from such species as the native and-should-have-been-one-of-the- climax -dominants Indiangrass to naturalized Johnsongrass. Other grasses included big, little, and silver bluestems; Texas wintergrass, several perennial dropseeds, sideoats grama (Bouteloua curtipendula) tumble windmillgrass (Chloris verticillata), and tumblegrass (Schedonnardus paniculatus). Forbs were varied but, as to be expected, composites "ruled" with prominent and common species including western ragweed (Ambrosia psilostachya), Baldwin ironweed (Vernonia baldwinii), iand frostweed or white crownbeard (Verbesina virginica). The major shrub was common greenbriar or catbriar (Smilax rotundifolia). Other shrubs were trumpet creeper (Campsis radicans) and poison oak.

Hamilton County, Texas. June. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-75 (Cross Timbers). Variant SAF 94 (Sycamore-Sweetgum-American Elm) that had former SAF designation of Sycamore-Pecan-American Elm. (Griffith et al., 2004). Closest biotic community designation of Brown et al. (1998, ps. 37, 38) was Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1, or more descriptively perhaps, a transition of this Oak-Hickory Series and Bluestem "Tall-grass" Series, 142.11, Plains Grassland 142.1 Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

413. Pecan timber- The forest form of pecan bottoms was represented by this stand of midddle-aged trees on the first terrace of a small creek floodplain in the patchwork of range plant communities where there is an intermixing of Western Cross Timbers and Grand Prairie in northcentral Texas. All trees were pecan whose tall straight trunks supported such woody vines as mustang grape, fiddleleaf or fringed greenbriar, and Alabama supplejack. Dominant understorey shrub was Carolina buckthorn. Most woody ground cover was the invasive, alien Japanese honeysuckle (Lonicera japonica), a treadful, horrid, etc. (adjectives fall short of vile description) woody weed. Major grasses included Canada wildrye, purpletop, Johnsongrass, and Texas wintergrass in that order. Major forbs were Baldwin ironweed (Vernonia baldwinii), and Indian plantain (Cacalia plantaginea).

Erath County, Texas. April, early to mid-vernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-75 (Cross Timbers). Variant SAF 94 (Sycamore-Sweetgum-American Elm) that had former SAF designation of Sycamore-Pecan-American Elm. (Griffith et al., 2004). Closest biotic community designation of Brown et al. (1998, ps. 37, 38) was Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1, or more descriptively perhaps, a transition of this Oak-Hickory Series and Bluestem "Tall-grass" Series, 142.11, Plains Grassland 142.1 Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

414. Bottomland pecan forest- Higher stocking of pecan trees on bottomland produces a forest in contrast to the savanna oar open woodland form of this range type as shown above. All trees in these two photographs were pecan. Dense stocking resulted in tall, relatively straight boles in marked contradistinction from the widely spreading crowns of pecans with more spacing among trees. Pecans with stocking rates represented by forest vegetation presented in these two photographs produce wood and not fruit. This is the forest community of a lumberman not an orchardman, and the form of vegetation that develops in absence of fire.

There was a well-developed woody vine layer that extended from ground level to tops of forest canopy which was comprised of mustang grape, rattanvine or Alabama supplejack, and fiddleleaf greenbriar. Carolina buckthorn grew as a smaller tree or larger shrub in the lower woody layer. The herbaceous layers included tallgrass species like Canada wildrye, purpletop, and the naturalized Johnsongrass along with composite forbs the two most common of which were Baldwin ironweed and Indian plantain. Much of the lower layer was composed of the invsive (and apparantely naturalized) Japanese honeysuckle.

415. Woody understorey of pecan bottom- From ground level and extending upward just shy of tree branching the range vegetation of a bottomland pecan forest was comprised of three woody vine species (mustang grape, fiddleleaf greenbriar, and Alabama supplejack) and the shrub or small tree, Carolina buckthorn. The wsidespread exotic invader, Japanese honeysuckle, was absent from this photoplot. The herbaceous portion of this forest vegetation visible in the background was presented in the next photograph.

416. Herbaceous layer (s) of a bottomland pecan forest- In addition to the woody vines of mustang grape, fiddleleaf greenbrier, and Alabama subblejack or rattanvine in right foreground and background of this photoquadrant important forbs of the herbaceous component included Indian plantain (left corner of foreground) and scattered smaller, apparently shade-stunted Baldwin ironweed. Many of the small green shoots with forb-like leaves were new sprouts of greenbrier and supplejack.

417. Pecan in winter- Examples of winter twigs, with emphasis on terminal bud, of pecan. The last photograph was a more detailed (closer-up) view of that shown in the third photograph. Upper bank above Bosque River, Erath County, Texas. January; winter dormancy.

418. Young leaves and catkins of pecan- A leader or shoot (first slide) and details of leaves and catkins (second slide) of pecan in West Cross Timbers of Texas. Erath County, Texas. March, pre-anthesis stage.

419. Fruit in the forest- Terminal end of a short shoot of native (hard-shell) pecan in the Grand Prairie of northcentral Texas. End of a short shoot on a pecan in the small bottomland forest range site along Richardson's Creek.

Hunewell Ranch, Tarleton State University. Erath County, Texas. October; nearly fruit-ripe stage.

Eastern Cottonwood (Populis deltoides)-dominated bottomland forests

420. Floodplain forest of eastern cottonwood in the old bed of the Republican River— Seen here is a wetland forest site and a wetland range site (background and foreground, respectively). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). Cottonwood form ofK-89 (Northern Flood-Plain Forest).The forest site is an eastern cottonwood consociation (SAF Forest Cover Type 63) with an herbaceous understory of Indiangrass. The range site also has Indiangrass as the dominant herb, but Canada wild rye and prairie sandreed are important associated species. Dundy County, Nebraska. Populus deltoides Association (if and when such is recognized), Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). October, autumnal aspect. Dundy County, Nebraska.

421. Interior of a floodplain or wetland forest on the former bed of the Republican River— View inside the eastern cottonwood forest of the preceding slide, but showing local importance of scattered green ash (Fraxinus pennsylvanica). This vegetation is about 60 years old having begun development when the river flooded and changed course. The climax tree species were already established at time of that disturbance, but this perturbation removed most of the woody plants except the larger cottonwoods and ash allowing migration and establishment of prairie grasses as well as sedges and rushes more typical of the numerous marshes in the area.

Dundy County, Nebraska. October, autumnal aspect. FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem).Cottonwood form of K-89 (Northern Flood-Plain Forest). SAF 63 (Cottonwood). Populus deltoides Association (if and when such is recognized), Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). October, autumnal aspect.

422. Semiarid gallery forest- Exterior view of a riparian zone forest of Rio Grande cottonwood (Populus deltoides ssp. wislizenii), netleaf hackberry (Celtis reticulata), Mexican walnut (Juglans major), and lanceleaf sumac (Rhus lanceolata) At outer (least mesic) edge of this gallery forest some of the more moisture-loving grasses formed a thick sward. The large cespitose grass was Wright's sacaton (Sporobolus wrightii) the dominant understorey species; Canada wildrye was the major associate species. The most common forb was horehound (Marrubium vulgare), a naturalized Eurasian weed.

This range vegetation was in the Trans-Pecos Basin and Range vegetational area, the west Texas portion of the Chihuhuan Desert Region. It was in Chihuhuan Deserts 24 (ecoregion level III), Chihuhuan Montane Woodlands 24d (ecoregion level IV) based on Griffith et al. (2004). It was an example of mesic forest in a semiarid climate. FRES No 17 (Elm-Ash-Cottonwood Forest Ecosystem). No aqppropriate Kuchler unit for southern and.or southwestern cottonwood gallery forest. SAF 63 (Cottonwood). Populus deltoides Association (if and when such is recognized), Cottonwoow-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Trans-Pecos Basin & Range Vegetational Area: Chihuhuan Deserts- Chihuhuan Montane Woodlands Ecoregion 24d (Griffith et al., 2004).

June, summer aspect. Calamity Creek, Jeff Davis County, Texas.

423. Great Plains floodplain forest- Eastern cottonwood and various species of willow formed a species-rich forest complete with prolific regeneration and a prominent herbaceous understorey. In these two slides the dominant herbaceous plant at local level ws American licorice (Glycyrrhiza lepidota), but grasses, sedges (Carex spp.), spike rush (Elocharis spp.) rushes (Juncus spp), and cattails (Typha domingensis) were the major understorey plants throughout most of the vegetation. Canada wildrye and bottomland switchgrass were most common with Johnsongrass (Sorghum halepense) the associate. Even the annual Eurasian grass, cheatgrass (Bromus tectorum) was common locally.

FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). There is no Kuchler unit describing southern or southwestern cottonwood-dominated riparain forests. SAF 63 (Cottonwood). Populus deltoides Association (if and when such is recognized), Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Rolling Red Plains Vegetational Area: Central Great Plains- Red Prairie Ecoregion 27h (Griffith et al., 2004).

June, summer aspect. Floodplain, North Fork of Red River. Wheeler County, Texas.

Members of the Salicaceae, the cottonwood-willow family, are generally dioecious; that is having the characteristic or phenomenon of unisexual genotypes (of separate male and female plants). In the section below examples of male and female catkins on eastern cottonwood were presented. These unisexual (imperfect) flowers were borne on two separate (genetically distinct) trees that were located less than one adult cottonwood crown apart.

424. Gold and blue- Rich amber-colored staminate catkins of eastern cottonwood against a backdrop of a rare (for Texas) azure blue sky demonstrated colorfully and at progressively closer camera distance the male-flower cluster of this iconic tree of the tallgrass prairie.

Several spring-swollen lateral buds were visible on the leader presented in the first two of these three slides.

This example came from the Grand Prairie-Western Cross Timbers vegetational area of northcentral Texas.

Erath County,Texas. Late March.

425. Paler gold with the same blue- Pale amber-colored pistillate catkins of eastern cottonwood against a backdrop of a rare (for Texas) azure blue sky demonstrated colorfully and at progressively closer camera distance the female-flower cluster of this iconic tree of the tallgrass prairie.

The female tree that bore these lady catkins was less than felling distance (less than a felled trees distance) from the gentleman catkins seen immediately above. In addition, the second image shown here sported a large and swollen terminal or apical preformed bud.

Erath County,Texas. Late March.

426. Flowers of both sexes- Female (pistillate) catkins were placed above and below a male (staminate) catkin of eastern cottonwood that was growing in the Grand Prairie-Western Cross Timbers vegetational area of northcentral Texas. These catkins were some of those presented in the two immediately preceding slide-caption sets.

Erath County,Texas. Late March.

427. Parts of a foliaceous, voluptous female- Young, full-grown leaves and a pendulous cluster of immature capsules of eastern cottonwood (Populus deltoides sp. deltoides) in the Western Cross Timbers of northcentral Texas. Populus species are dioecious so this was obviously a female tree.

The cobalt-blue sky was result of a late-season norther that took out drab, high-humidity, polluted air and replaced it with fresh, clean Arctic air.

Silvics of eastern cottonwood was provided in Burns and Honkala (1990).

.Erath County, Texas. Mid-April.

428. Spring-bedecked female parts- Young, fully developed leaves and a cluster of immature capsules of eastern cottonwood in the West Cross Timbers of northcentral Texas. There are any number of fine references for this State Tree of Kansas including the following which were given in contest of taxonomy and morphology of eastern cottonwood.

Systematics of Populus species (even within P. deltoides) has and continues to be a matter of confusion and conflict with any number of taxonomic treatments ranging from those of Coulter (1891-1894, ps. 419-420) and Sargent (1933, ps. 132-137 passim) through Vines (1960, ps. 89-92), Steyermark (1963, ps. 507-508), Correll and Johnston (1979, p.455), Great Plains Flora Association (1986, ps. 278-279), Diggs et al. (1999, ps. 975-976), and Allred and Ivey (2012, p. 529-530) for the general region herein involved. It seemed to this rather independent author that the treatment by Diggs et al. (1999, ps. 975-976) was most accurate and appropriate, perhaps primarily because it was the closest flora for this immediate area.

Diggs et al. (1999, ps. 975-976) elevated to two subspecies of P. deltoides some taxa (species, varieties, forms) formerly used for eastern cottonwood, plains cottonwood, and perhaps others such as Sargent's cottonwood by previously published work. In their dichotomy of these subspecies, subspecies deltoides had 10 to 20 coarse teeth on each side of the deltoid leaf whereas subspecies monilifera had only five to 10 coarse teeth along each. Viewers can handily see in various of the views presented in this short treatment that this example was clearly P. deltoides subsp. deltoides.

Great Plains Flora Association (1986, p. 279) lumped all Populus deltoides for the Great Plains as being P. deltoides subsp. monilifera with a note that it and P. deltoides subsp. deltoides intergraded with each other along eastern margins of the Great Plains. These authors (Great Plains Flora Association, 1986, p. 2) extended their version of the "Great Plains" (presumedly so that they could include all of Kansas and northward states) into the Central Lowlands physiographic province. This implied that there would be no specimens of P. deltoides subsp. deltoides in Kansas, western Missouri and so on (Great Plains Flora Association, 1986, ps. 2, 279). This would certainly to be erroneous, but again these authorities distinguished between numbers of teeth on leaf margins as between early leaves and late leaves with many more margin teeth on late leaves.

Either way, treatment by Great Plains Flora Association (1985, p. 279) clearly conflicted with treatment by Allred and Ivey (2012, ps. 529-530) who broke P. deltoides into four varieties including P. deltoides var. deltoides which entered ("scarcely") northeastern New Mexco. This is, of course, west of Kansas, Nebraska, Missouri, etc.

Burns and Honkala (1990)--official US Forest Service version--treated eastern cottonwood (P. deltoides) as P. deltoides var. deltoides for "eastern cottonwood (typical)" and P. deltoides var. occidentalis for "plains cottonwood".

Confused, yet?

.Erath County, Texas. Mid-April.

429. How're they hanging?- Immature many-seeded capsules of eastern cottonwood. The pendulous position of the fruit cluster in the first slide is the natural or undisturbed habit whereas the "wrapped" or "coiled" position in the second slide was the result of wind action. The Populus species are not only wind-pollinated and with wind-dispersed seeds, but they also subject to numerous other "avenues", "venues", or "expressions" of wind action

.Erath County, Texas. Mid-April.

430. Cotton candy on a tree- About one month later (after the bearing of unisexual flowers) some of the pistillate (female) catkins presented above bore ripened ovaries that were now capsules which were dehiscing and releasing seeds equipped with their namessake "cottonwood" parachute-like pilose hairs. This morphological featurewas described by Smith (1977, ps. 128, 292) as comose meaning to have a coma which is a tuft of hairs.

Erath County, Texas. Late April.

431. Foliage and fruit of prairie silvan queen- First of these two photographs was young shoots (two small seedlings or small saplings) of eastern cottonwood. They had been flattened by running water from recent spring (June) rains. (They straightened up.) Second photograph was the typical deltoid leaf with ripe fruit ( a capsule) of eastern cottonwood. Cottonwood capsules are described as comose, an adjective referring to having a coma (a tuft of hairs). Cottonwood derived its common name from the phenomenon of conspicuous, usually copious (and inconvenient) shedding of the comose ("cottony") fruit. All members of the Salicaceae are dioecious so that a tree is male or female and thus produces only staminate or only pistillate flowers. Of course only female plants produce fruit so that occurrence of the "messy fluff" comes only from the ladies. (Some range men find girls troublesome while others enjoy the mess.) Female catkins from this same plant were shown in the next photograph. Erath County, Texas. May.

432. Wild or American licorice (Glycyrrhiza lepidota)- Wild licorice is one of the more common legumes along watercourses throughout the Great Plains. It occurs in both the riparian zone and at outer (drier) margins of rivers, creeks, and floodplains. July, full bloom. Custer State Park, Custer County, South Dakota.

433. Licoriced sticks- Shoots with ripening legumes of wild or American licorice in the floodplain of the Platte River. The two closer-in (more detailed) shots when compared to similar photographs of other range species of Leguminosae served as examples of the great diversity in shapes, sizes, pubescence, etc. of legumes.

Adams County, Colorado. Early July; advanced immature stage of fruit development.

434. Healthy regeneration of cottonwood and willow- One of the major problems in structure and sustainability of riparian and floodplain forests is reproduction under concentrated livestock grazing. This is especially so in arid and semiarid regions where livestock, especially cattle, tend to "hang in on water". The forest shown here was grazed periodically at light utilization (low degrees of use). There was extremely vigerous reproduction, both sexual and axexual, of both cottonwood and willow species. Generally members of the Salicaceae sprout vigerously particularily under moist soil conditions as on this subirrigated floodplain in the rolling red prairies. Successful reproduction of the salicaceous shrubs is a good indicator of proper management of the grazing resource and range ecosystem.

Correct identifiction of willows (Salix spp.) to the correct species is a challenging task. It is often impossible to obtain positive identification without the catkins. There appeared to two and, probably, three species in this bottomland forest. This most likely "mixture" being some combination of black willow (S. nigra), peachleaf williw (S. amygdaloides), coyote willow (S. exigua), or sandbar willow (S. interior).

FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). No Kuchler unit for southern or southwestern cottonwood and/or willow riparian forest. SAF 63 (Cottonwood). Populus deltoides association (if and when such is recognized), Cottonwood- Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Rolling Red Plains Vegetational Area: Central Great Plains- Red Prairie Ecoregion 27h (Griffith et al., 2004).

June, summer aspect. Flood plain, North Fork of Red River. Wheeler County, Texas.

435. Excellent regeneration of eastern cottonwood and willow species on a Great Plains bottomland forest- This is a close-up view of outstanding production of the "next generation" of eastern cottonwood (foreground) and willow (background) on a subirrigated river bottom in the Great Plains. Mixed prairie vegetation surrounded this wetland ecosystem.Trees were functioning as phreatophytes.

FRES No.17 (Elm-Ash-Cottonwood Forest Ecosystem) .No Kuchler unit for southern or southwestern cottonwood and/or willow riparian forest. SAF 63 (Cottonwood) Populus deltoids association (if and when such is recognized), Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Rolling Red Plains Vegetational Area: Central Great Plains- Red Prairie Ecoregion 27h (Griffith et al., 2005).

June, summer aspect. Flood plain, North Fork of Red River. Wheeler County, Texas.

436. Understorey of a cottonwood-willow bottomland forest- A lower woody layer of regenerating cottonwood and willow, an herbaceous layer of grasses and grass-like plants, and a multi-storied liana layer of greenbriar (Smilax sp.) and grape (Vitis sp.) added to the mature tree layer such that this range plant community formed a true forest structure on a wide expanse of river floodplain. Understorey graminoids included bottomland switchgrass, Canada wildrye, Johnsongrass, sedges, spike rushes, rushes, and cattail. This was another of the widespread and naturalized Johnsongrass. The most common forb was wild or American licorice.

Proper range management with emphasis on light and periodic grazing/browsing (vs. the widespread practice of continuous heavy grazing) permitted perpetuation of this forest. This was an outstanding example of good stewardship and sustainability of range resources.

FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). No Kuchler unit for southern and southwestern cottonwood and/or willow gallery forest. SAF 63 (Cottonwood). Populus deltoides association (if and when such is recognized), Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Rolling Red Plains Vegetational Area: Central Great Plains- Red Prairie Ecoregion 27h (Griffith et al., 2004).

June, summer aspect. Flood plain, North Fork of Red River. Wheeler County, Texas.

437. Plains cottonwood bottomland forest in the far-western plains- Introduced in this photograph and treated further in subsequent slides was a forest range that developed along an *intermittant stream or, perhaps, interrupted stream (East Bijou Creek) in the Colordo Piedmont on which the dominant, the only, tree species was plains cottonwood (Populus deltoides ssp. monilifera). Plains cottonwood is another subspecies of eastern cottonwood native to the semiarid zone This forest range had been grazed by cattle for decades and was still in Good range condition class so as to approach the state of potential natural (climax) vegetation. There was limited reproduction of cottonwood, but this appeared adequate to maintain the uneven-aged population that dominated and defined this range plant community. Besides the upper woody (crown or canopy) layer there was a lower woody layer--though it was sporadic or "spotty" in its pattern of distribution-- made up of saplings and young poles of plains cottonwood. (This structural feature was shown in sebsequent slides.) A few seedlings of cottonwood were present in the herbaceous understorey. There were three herbaceous layers all of whch were dominated by grass species. These three were tallgrass, midgrass, and shortgrass layers so the understorey was mixed prairie. In fact this range vegetation could be visualized or even described as mixed prairie savanna with plains cottonwood as the woody element. Cottonwoods were dense enough, at least in numerous microhabitats where there were local populations (or, perhaps, individual stands would be a more apt term) that crowns typically interlocked. Thus, forest was the proper designation. Still, the overall or general vegetation could be conceived of as a savanna or, at least, as having a savanna-like physiogonomy, and a wetland savanna (or savanna wetland) at that.

Portions of this range plant community actually comprised a gallery forest along the channel of this seasonally and spatially intermittant stream. Such a dispersion pattern and structure was shown in this photograph and the next photgraph where East Bijou Creek actually held water. Along other short reaches of this stream the creek channel was dry with only moister soil to distinguish the more-or-less interrupted riparian zone from the environment of the farther (outer) channel banks and adjacent floodplain.

The herbaceous vegetation that was actually in the stream channel and extending outward on the nearest (lowr) banks was made up of prairie cordgrass (Spartina pectinata), bottomland ecotype(s) of switchgrass, and Canada wildrye (in that order of cover and biomass or standing crop).

It should be noted that in addition to the light degree of use of tallgrass species there were a lot of cottonwood leaves and lower branches within browsing reach of cattle. This forest range was not being abused by the rancher who used. A well-earned tip of the hat and public praise to the faithful steward of this land. If the author had known he name of this commendable range manager he would have printed it.

A note on stream terminology- The author was "only pasing through" this country and, as he did reside here, could not determine the nature of East Bijou Creek and use the most accurate designation. Specifically, the photographer could not distinguish whether it was an intermittent or intermediate stream.

*Intermittent stream- "A stream or reach of a stream tht flows only at certain times of the year, as when it receives water from springs or from some surface source....A stream that does not flow continuously, as when water losses from evaporation or seepage exceed the available streamflow...Synonyms: temporaray stream, seasonal stream" and

*Interrupeted stream- "A stream that contains perennial reaches with intervening intermittent or emphemeral reaches, or a stream that contains intermittent reaches with intervening ephemeral reaches" (Wilson and Moore, 1998).

East Bijou Creek, Elbert County, Colorado. Mid-July (estival aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). No Kuchler unit for southern and southwestern cottonwood forest. No SRM. SAF 63 (Cottonwood). Populus deltoides association (if and when such is recognized), Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Southwestern Tablelands-Piedmont Plains and Tablelands Ecosystem, 26e (Chapman et al., 2006).

438. A little farther downstream- At another short reach with surface water of the channel of an intermediate or interupted stream (East Bijou Creek) along which a forest of plains cottonwood with a n herbaceous understorey had developed. In the reach of the creek viewed here the stream chaannel was smaller (narrower and shallower) so that water persisted for shorter periods than in the reach shown immediately above.

Range vegetation along this segment of East Bijou Creek was more of a mixture of aquatic and also of less mesic grass species than in the reach with a bigger channel and greater riparian zone surface seen above. In this stream segment major range plants were Torrey's rush (Juncus torreyi) and western wheatgrass as well as Canada wildrye and prairie cordgrass which were relatively more abundant around the larger area of water shown in the preceding slide. Cheatgrass or downy bromegrass (Bromus tectorum) was present in minor down to trace amounts.

Both smaller (presumedly younger) and full-sized adult trees of plains cottonwood were present in this portion of the floodplain forest.

Only range vegetation of the general riparizn zone was described in this and the preceding photograph. Composition and structure of the floodplain away from the immediate riparian zone was treated in the slides and their captions that followed immediately below.

439. Along a dry reach of the interupted stream- Range vegetation of a bottomland forest dominated by plains cottonwood along East Bijou Creek, an interrupted or intermittent (whichever it was) stream. This view across across (at a rough right angle to) the stream channel rather than a headon or inline view as in the two preceding photographs. Different age/size classes of trees were present showing this to be an uneven-aged population of plains cottonwood. There were relatively few cottonwood seedlings, but their number appeared adequate (though barely) to maintain the only tree species in this plant community.

Density and spatial arrangement (dispersion pattern) of cottonwood varied from closed canopy forest along the riparian zone to such wide (sparse) spacing of trees as to constitute a savanna on the outskirts of this bottomland range plant community. Cottonwoods grew (occurred) as individual mature adult trees or small groups of two or three trunks up small groves of eight to ten trees. Groves characteristically included pole-sized as well as fully grown trees. These larger groups were closer to the stream (generally in the riparian zone) and tree density declined with distance from the creek until the savanna spacing at outer border of the bottomland community. Dead wood at base of the adult cottonwood across the creek was mostly debris washed in and lodged against the trunk although a minor amount of this wood pile was branches from the crown of the tree that served as a barrier to water-borne material on this floodplain.

Although it was not obvious in this wide view there were three layers comprising the herbaceous understorey: tallgrass, midgrass, and shortgrass that were dominated by prairie sandreed, western wheatgrass, and buffalograss, respectively. Downy brome or cheatgrass was present (as it alway is) but it was minor on this range that was under superb management (see shortly below).

Forbs present included wild licorice, the most common and important; prairie coneflower or Mexican hat (Ratibida columnifera); showy milkweed (Asclepias speciosa); annual sunflower (Helianthus annus); four o'clock (Mirabilis linearis); prairie sagewort (Artemisia frigida); western ragweed (Ambrosia psilostachya) and kochia (Kochia scoparia). There were sparsely scattered plants of plains pricklypear (Opuntia polycantha).

Range condition class was Good. Cattle had most likely grazed this range for decades, certainly over the last several years to the present time as proven by dung piles of varying ages. In all these photographs there were cottonwood leaves and leaders within reach of cattle indicating that there had not been heavy browsing of of this species which has relatively high palatability to cattle. (High preference by cattle for Populus deltoides and the relative ease with which this palatable species is browsed out was described in detail below.) Presence of this palatable browse and fairly high abundance (author's subjective judgement based on cover, density, vigor, etc.) of prairie sandreed and prairie cordgrass along the creek was undeniable evidence that this range had not been abused (overgrazing/overbrowsing had not taken place) in the recent history of use by livestock. Accolades to the range manager.

East Bijou Creek, Elbert County, Colorado. Mid-July (estival.aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). No Kuchler unit for southern and southwestern cottonwood forest. No SRM. SAF 63 (Cottonwood). Populus deltoides association (if and when such is recognized), Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Southwestern Tablelands-Piedmont Plains and Tablelands Ecosystem, 26e (Chapman et al., 2006).

440. On the higher and outer bank of the interrupted stream- View looking up the channel and along the bank of East Bijou Creek, an interrupted or intermittent (whichever it was) stream, in the semiarid Colorado Piedmont. From the physiogonomic and structural perspectives, an upper woody layer of plains cottonwood dominated this bottomland range vegetation. Tree density varied from that of a gallery forest in the riparian zones which remained wetter later into the summer to a savanna at outer margins of the floodplain plant community. This range community was, in essence, a mixed prairie with a tree element that varied greatly in tree density and dispersion pattern.

From perspective of annual biomass production (standing crop productivity) the dominant plant species of this range ecosystem was almost assuredly prairie sandreed. Western wheatgrass was the overall community associate. Switchgrass held this honor within the riparain zone in which the dominant herbaceous species was prairie cordgrass. Thus it was that a tallgrass species was the herbaceous dominant of both the wetland (riparian zone) and the greater floodplain.

Dominance of the greater floodplain (beginning at top of stream bank) by prairie sandreed and the close association of western wheatgrass, the major midgrass species, was depicted in this photograph. Buffalograss and blue grama were not conspicuously featured, but they were certainly well-represented. Mesic tallgrass species (notably prairie cordgrass and switchgrass) that dominated the lower stream channel were absent. Forb species included wild licorice, generally the most important; prairie coneflower of Mexican hat, showy milkweed, annual sunflower, kochia, prairie sagewort, and narrowleaf four o'clock. There were sparsely scattered plants of plains pricklypear, but nothing approaching density and cover to form even an interrupted shrub layer.

441. A savanna form on the outer boundary- At outside margins of plains cottonwood-dominated bottomland range vegetation the plant community was actually a savanna or, at best from a crown canopy criterion, a woodland rather than a forest. This lower stand density or wider tree spacing was in contrast to the gallery forest that developed in the riparian zone of this interrupted stream as was shown above.

Downed limbs immediately beneath cottonwood boles were a combination of those that fell from directly overhead off of senescing adult trees as well as those that had been carried by flood water to rest against the trunks. Such debris offered protection for the all-too-scant number of cottonwood seedlings available to replace their aging elders.

The understorey consisted of three distinct herbaceous layers of tallgrass, midgrass, and (to a lesser degree) shortgrass species, the dominants of whch were, respectively, prairie sandreed, western wheatgrass, and buffalograss (blue grama was less abundant having lower relative cover than buffalograss). Forbs included wild licorice, the most common and important; prairie coneflower or Mexican hat; showy milkweed, narrowleaf four o'clock, prairie sagewort, annual sunflower, and kochia.

442. Village meeting- Local assemblage of range plants on plains cottonwood-dominated bottomland range on floodplain of an interrupted or, possibly, an intermittent stream in the semiarid zone. Dominant plant species was prairie sandreed. Other range plant species included prairie sagewort, western ragweed, Mexican hat or prairie coneflower, and wild licorice. The second (vertical) slide was a closer-in, more detailed view of the left foreground of the first slide. This second photograph presented a better view of shoots of praiirie sandreed.

443. At base- Basal portion of shoot of prairie sandreed with parts of sheath and blade of the four lowermost leaves. More details were presented in the next subsequent slide. This shoot was growing on East Bijou Creek, an intermittent or interrupted (whichever one), in the Colorado Plateau, the physiographic unit just west of the High Plains or Llano Estacho (Staked Plains) of the Great Plains province.

Elbert County, Colorado. Mid-July.

444. Unit of a reed- Phytomer--fundamental, repeating unit of a grass shoot including upper half of a node, internode, and lower half of next highest node complete with leaf--of prairie sandreed (first slide) and details of culm, sheath, and colar of this phytomer (second slide). Floodplain of East Bijou Creek, an interrupted or, maybe, an intermittent stream in the semiarid tableland of the Colorado Piedmont.

Elbert County, Colorado. Mid-July.

445. Atop the reed of a reed- The panicle--the entire structure--of prairie sandreed (first slide), upper or distal portion of this panicle (second slide), and one branch of this panicle at onset of anthesis (third slide).

East Bijou Creek, Elbert County, Colorado. Mid-July; initiation of anthesis.

446. Unfulfilled florets- Spikelets of prairie sandreed showing florets at pre-anthesis stage. There is only one floret per spikelet in this species and the upper glume is considerably longer than the lower (Shaw, 2008, p.90). Both of these features were presented in these two photographs.

East Bijou Creek, Elbert County, Colorado. Mid-July; initiation of anthesis.

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

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

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

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

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

449. Hanging lightly- Drooping leaders (woody shoots) with pendulous clusters of fruit on eastern cottonwood (Populus deltoides subsp. monilifera). The delta-shaped leaves, obvious basis of the specific epithet,.is a key identifying feture of this species. Vines (1960, p. 91) described leaf shape of P. deltoides as "broadly deltoid, margin crenate-serrate, apex abruptly acute or acuminate, base truncate to heart-shaped or abruptly cunete".

Larimer County, Colorado. Mid-June (late spring), seed-shatter stage.

450. Lightly but effectively- Fruit (first slide) and individual seeds (second slide) of eastern cottonwood. Eastern cottonwood "wrote the book" on anemochory or aerochory, dispersion of plant disseminules or propagules by wind. Many prairie plants evolved (natural selection, survival of the fittest) anemochory as a means of effectively and efficiently dispersing pollen, but cottonwood went a step farther and has one of the best adaptations to dispersal of its fruit and seed via aerochory of any woody plant in North America. The liguliferous composites like thistles and dandelions have nothing on Populus species.

Eastern cottonwood is a dioecious species. Its fruit is a capsule, numerous of which are borne in racemes (first slide) and each of which consist of 10 to 30 seeds (second slide) with each of these being equipped with pubescence in the form of numerous, long, cottony fibers (Vines, 1960, p. 91). It is the cottony pubescence that permits cottonwood seeds to become air-borne so as to float and finally settle by gravity into every available "nook and cranny".

Larimer County, Colorado. Mid-June (late spring), seed-shatter stage.

451. Palatable resprouts- Root sprouts (suckers) from a felled, senescing eastern cottonwood on a prairie in the western Ozark Plateau. The three sets of root shoots presented here (first and second photographs) were two-year-old suckers or heterophyllous shoots produced from lateral roots (third photograph) of a relatively old, "half-rotten" eastern cottonwood after it was felled for fuel wood. The shoots, some of which in these photographs were over eight feet in height, had been completely defoliated (100% of leaves removed) by commercial beef cattle (cows and calves of mixed breeding) in both the first and second years of shoot development. The small leaves present on shoots at time of photographs were current-season's regrowth (early part of current--second--warm growing season).

Several (a high proprotion) of cottonwood root suckers were broken off by browsing cattle so that this extreme or severe degree of use included both eating of leaves and upper twigs of current growing season as well as complete breaking off of many root suckers that were the production of two growing seasons. Several of the broken shoots and branches were visible in some of the photographs presented here and immediately below. Cattle defoliation in the first growing season was heavy, but not total as in the second season of growth. In the second growing season many of the first and current year's root shoots were broken off at their point of origin on the lateral roots. This was done by cattle who "rode down" the root suckers in order to reach the uppermost leaves and terminal buds (ie. cattle ate all leaves and much of the upper parts of shoots, terminal twigs, plus broke off entire shoots by the browsing technique of "riding down").

"Riding down" is the feeding behavior whereby browsing animals reach up as high as they can on plants and then commence walking forward so that forage or browse plants bend down beneath the animals' weight and pass progressively beneath the feeding animals' brisket and belly until the animals have fed as far as the plant will bend to within reach of the animals' mouth. In the process, some of the brittle shoots and branches are broken off while the more limber shoots spring back-- minus leaves, fruit, buds, twigs, etc. that were consumed by the animals. In this feeding pattern animals can remove plant material at much greater heights than they could otherwise reach by merely streatching their necks and extending their tongues.

Herbaceous plant species growing adjacent to and beneath eastern cottonwood shoots was mostly the introduced pasture grass, tall fescue (Festuca arundinaceae), which has naturalized across most of the greater Ozark Region and had self-seeded onto this former tallgrass prairie. The second-most common herbaceous species on this local habitat was Japanese chess or Japanese brome (Bromus japonicus), a naturalized Eurasian annual grass. It appeared that none of the herbage of these two non-native grasses had been eaten by cattle although considerable quantities of biomass had been trampled. Cattle seemingly had quite a browsing bout on this spot as quantities of dung were left. There was no evidence of browsing by deer, rodents or rabbits.

Total herbivory (=herbivorous action) that did and would continue to influence regeneration of eastern cottonwood on this pasture included at least four components: 1) complete removal of leaves from cottonwood shoots, 2) total elimination of some cottonwood shoots by breakage at ground level (ie. total loss of some two-year old trees or two growing seasons of plant regeneration), 3) feeding selectivity that severely injured or setback cottonwood while at the same time having much less defoliation impact on annual and perennial grasses (both non-natives), and 4) deposition of dung with importation of more potential plant nutrients than was recycled or otherwise generated on this local environment.

Question to the "village idiot" (so that he can be "above average"): "will grasses or cottonwood derive most benefit for plant growth and/or survival from the combined influences of cattle feeding?"

Eastern cottonwood (both eastern and plains varieties), like most other members of the Salicaceae, readily coppices or resprouts following major injury or defoliation. Fowells (1965, ps. 516, 521) and Burns and Honkala (1990, vol. 2, ps. 533, 539) described vegetative reproduction in eastern cottonwood. Eastern cottonwood sprouts vigerously from both stumps and roots. In the pattern of most sprouting species, vegetative (asexual) reproduction declines with increaseing age of trees while vegetative propagation, both natural and atrtificial, is vigerous in young plants. Fowells (1965, p. 516) and Burns and Honkala (1990, vol. 2, ps. 533) cited research which found that eastern cottonwood (P. deltoides var. deltoides) stump-sprouted up to ages of 25 years.

Heterophyllous shoots (root sprouts or suckers) shown here were from a much older tree. Over half of the trunk (beyond the center growth rings) of this eastern cottonwood had rotted away so that it was impossible to accurately determine tree age. (Rotting appeared to have begun around an old fire scar.) Recollections by the author and his conversations with elderly observers firmly established age of this felled tree in excess of 80 years. Root suckering began in early spring following sawing of the old tree in the preceding winter.

On various occasions this author has observed resprouting by various kinds of cottonwood (Populus spp.) in larger and older tindividual trees following sawing, bulldozing, f lood breakage, etc. Obviously resprouting in cottonwood is not limited to trees that are younger than a quarter century.

It was also obvious that regeneration of eastern cottonwood depends upon protection from overbrowsing. It was possible that some of the herbivory described above benefitted (or had potential to benefit) eastern cottonwood. Dunging provided extra plant nutrients, especially nitrogen, available to surviving cottonwood shoots. Dung might also benefit grass proportionately more than cottonwood so that increased competition from grass resulting from soil fertilization was detrimental to cottonwood. It was conceivable that breakage of some cottonwood root sprouts (suckers) had the effect of beneficial prunning which would enable surviving shoots to grow larger faster and become too large for cattle to "ride down" and thereby escape most further browsing impacts (other than continued dunging if and when cattle rested in shade of growing cottonwoods).

Overall, however, and up to the point of time recorded here, even the village idiot made 100% of this pop quiz by explaining (in simple talk) that the browsing shown here was excessive and could result in death of resprouts and ultimate loss of eastern cottonwood from this pasture.

Ottawa County, Oklahoma. June.

452. Overutilization of palatable eastern cottonwood by cattle- Excessive (extreme or severe) defoliation of two-ear old root suckers of eastern cottonwood on a prairie in the Ozark Plateau. Beef cattle (crossbred cows and calves) removed all leaves and many shoot tips (including developing buds) from these heterophyllous shoots by early to mid-spring of the second growing season. This necessitated production and development of a new set of leaves (the leaves present in these and preceding photographs). Numerous branches were broken from shoots that survived "rding down" browsing by cattle, and lots of the two-year old shoots (root suckers) did not survive the aggressive feeding by cattle.

This was an example of overuse which, if continued, might well result in killing and loss of eastern cottonwood from this pasture. These slides demonstrated the importance of protection of young eastern cottonwoods that are extremely palatable (even to animal species that are grazers more than browsers) from excessive browsing. Regeneration of eastern or plains cottonwood such as that shown above in a plains cottonwood-willow bottomland forest would not have been possible with continued overuse at the severity shown here and in the three immediately preceding slides. If degree(s) of use showh here continued for just a few more years (number unknown; would vary depending on many factors) the eastern cottonwood will die out on this pasture. The old felled cottonwood tree described in the preceding caption will be dead afterall.

The main lesson from these photographs was the concept of proper degree of use (the first and most important of the Four Cardinal Principles of Range Management), often expressed as Proper Use Factors. Secondary lessons from these slides involved range animal behavior, including both the phenomenon of "riding down" and feeding selectivity. A degree of defoliation approaching "anililation" of eastern cottonwood coupled with non-eating of tall fescue and Japanese brome (though with some trampling and covering with dung of these grasses) provided a "textbook example" of feeding selectivity, the threeway interaction of animal preferences for certain plant species, relative palatability of different plant species, and availability of herbage and browse of certain plant species.

Ottawa County, Oklahoma. June.

453. It'll do its part and its best, but eastern cottonwood cannot rise from the dead indefinitely- Close-up view of one of the severely browsed shoots that suckered roots of a felled eastern cottonwood described above. All leaves are regrowth that replaced earlier full-grown leaves that were eaten by browsing beef cattle. These were some shoots that survived extreme degrees of browsing. Many shoots did not survive. (This exterme defoliation might be expected from overpopulations of browsers like deer, but this was from just "plain ole cows".)

If the rangeman takes care of the feed resource it will take care of him. If this cattleman wanted natural replacement of a nice shade tree a drastic change of management was in order. A quicky, makeshift exclosure of wovn wire would protect these young shoots until they could grown beyond reach of the cattle. If this cattleman had the goal of elimination of eastern cottonwood from this pasture he had a good start toward that objective. In other words, if eastern cottonwood was regarded as a brush species (a woody noxious plant) and a single-species stand of tall fescue was the desired result then cattle browsing appeared to be a very effective method of biological control to reduce (perhaps totally eliminate) this woody species from this grazing (or haying) unit of land. As it happened such was not the case. This cattle-owner was simply running cattle for tax purposes and social status (ie."dodging taxes" and "bragging rights").

This photographic case study was used to illustrate how excessive degree of use and/or general imporper animal browsing can prevent or greatly reduce regeneration of eastern cottonwood. Degree of use on eastern cottonwood would almost certainly have been much less if cattle-grazing had been restricted to late autumn through winter-early spring when cottonwood was dormant and nutritive value and palatability of the naturalized cool-season grasses was higher.

Ottawa County, Oklahoma. June.

454. Standing in river water- Mature Rio Grande cottonwood trees were flooded by water from the Rio Grande River that flowed a short distance from this location. River water rose above the ground level via subsurface flow. Ponded water was visible because manicuring with power tools for a picnic area (in a national park) eliminated the understorey of what was (would have been) a bottomland or gallery forest in the Chihuhuan Desert. The herbaceous layer in this radically modified vegetation consisted almost exclusively of introduced perennial grasses including bermudagrass (Cynodon dactylon), King Ranch bluestem (Andropoogon ischaemum= Bothriochloa ischaemum), and Dallisgrass (Paspalum dilatatum). Natural vegetation of diverse structure, including a shrub layer below the tree layer and the invasive Eurasian salt cedar (Tamarix chinensis), was present in the background.

FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). No Kuchler unit for southern or southwestern cottonwood and/or gallery forest.. SAF 63 (Cottonwood). Populus deltoides association (if and when such is recognized), Cottonwood-Willow Series in Southwestern Riparian Deciduous Forest botic community of Brown et al. (1998). Trans-Pecos Basin & Range Vegetational Area: Chihuhuan Deserts- Low Mountains & Bajadas Ecoregion 24c (Griffith et al. 2004).

June, summer aspect. Floodplain of Rio Grande (water mostly from Rio Concho), Big Bend National Park, Brewster County, Texas.

Some "Odd Lots" Forests

455. Red oak-linden (basswood) mesic forest— This community was described by Weaver (1965, p. 14-17) as the most mesophytic forest in Nebraska. It extends from draws to north slopes and is one of the best-developed forest types at the edge of the vast grasslands of the continental interior. Ironwood or hop hornbeam (Ostrya virginiana) is the dominant species of the shrub layer, but Virginia creeper and wild grape vines climb trees to give an aboreal shrub component. The Missouri Natural Areas Committee (1987) described a similar community as mesic upland forest. The Society of American Foresters (1980) did not list or describe this type, but Braun (1950, p. 234) interpreted it as comparable to the red oak-sugar maple-basswood community in Pennsylvania and in the Mississippi Valley section of the oak-hickory region in Nebraska.Westveld (1939, pgs. 46, 51, 67-72, 77, 86, 114-117, 267, 268) recognized the red oak-basswood-white ash forest type for the New England white pine and central hardwood regions as well as for the mid-Atlanic oak region. Thus it seemed likely that the red oak-basswood mesophytic forest which occurs at the western-most edge of the immense eastern deciduous forest formations and the eastern origins of the vast interior grasslands is a variant of SAF 26 (Sugar Maple-Basswood) which was described as grading into other cover types throughout the deciduous forest zone.No obvious FRES or Kuchler designation, but basswood is a dominant of K-90 (Maple-Basswood Forest) to which it bears some taxonomic kin based on the Braun interpretation. Western Corn Belt Plains- Nebraska-Kansas Loess Hills Ecoregion, 47h (Chapman et al., 2001). Schramm State Park, Sarpy Cass County, Nebraska.August, estival aspect.

456. Floodplain (gallery) forest of Wichita River- Exterior view of a forest community composed of discrete zones of vegetation along the terrace of a meandering river in the . This was a gallery forest that lined the bank and terrace of a river bordering on the mixed-tallgrass prairie transition. Immediately adjacent to the river was a vegetation zone consisting of green ash as a tree layer with an understorey of Canada or nodding wildrye. Also in the tree layer were occasional individuals of eastern cottonwood, hackberry or sugarberry (Celtis laevigata var.reticulata), pecan, and very rarely, black willow (Salix nigra). The herbaceous understorey was "pure" or "solid" wildrye with essentially no other species present so as to constitute a consociation. Green ash approached consociation status. The vegetation zone outer (farther from the river bank) from the green ash-nodding wildrye zone was eastern cottonwood with cottonwood leaves forming a mulch or litter layer devoid of plant species except the lianas, poison oak and trumpet creeper (Campsis radicans). Both of these woody climbers reached tops of cottonwoods. Poison oak was present as both climber and "regular form" of shrub. Outward from the cottonwood zone was one dominated by black willow. The fourth or outermost (from the river bank) vegetation zone was composed of the arboreal (and almost exclusive) dominant, western soapberry (Sapindus saponaria var. drummondii= S. drummondii) with an herbaceous understorey dominated by western wheatgrass (Agropyron smithii) with purpletop (Tridens flavus) as the associate. Buffalograss (Buchloe dactyloides) and Texas bluegrass (Poa arachnifera) were present also, the former being an occasional "spot" (microsite) dominant.

Lucy Park, Wichita County, Texas. October. Estival aspect. FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). Combination of SAF 63 (Eastern Cottonwood) and green ash-dominated variant of SAF 93 (Sugarberry-American Elm-Green Ash). Loamy Bottomland range site. Variant of Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Central Great Plains- Broken Red Plains Ecoregion, 27i (Griffith et al., 2004).

457. Deciduous riparian forest on terrace of Wichita River- Two interior views of the floodplain or gallery forest shown from the exterior in the preceeding photograph. The arboreal layer of this bottomland community was dominated by green ash with scattered trees of eastern cottonwood, sugarberry, and pecan making up trace amounts of the canopy cover. The herbaceous understorey layer was a consociation of Canada wildrye. There were no arborescent shrubs or woody climbers in this vegetation zone which bordered the river. Reason(s) for lack of adult trees and presence of even-age structure of immature trees could not be determined. There were no stumps, trunks, or other remains of previously live trees. Numerous sprouts from young stumps browsed by beaver (Castor canadensis) were present (see below).-

Lucy Park, Wichita County, Texas. October. Estival aspect.FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). Green ash variant of SAF 93 (Sugarberry-American Elm-Green Ash). Loamy Bottomland range site. Variant of Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Central Great Plains- Broken Red Plains Ecoregion, 27i (Griffith et al., 2004).

458. Leader of green ash- Leaves and fruit along an internode of green ash. Length of internode was approximately 20 inches which indicated potential for annual increment of green ash on a bottomland site in a subhumid preacipitation zone. Lucy Park, Wichita County, Texas. October. Estival aspect.

459. Leaves and fruit of green ash- Fruit type is a samara (winged achene). Lucy Park, Wichita County, Texas.October.

460. Upper reaches of a deciduous bottomland (gallery) forest along Wichita River- Western soapberry dominated this forest community growing on the upper bench or reach of the river terrace. This was the uppermost zone of the riverine forest that was comprised of green ash, eastern cottonwood, hackberry, and pecan adjacent to the meandering river. At this greater elevation and less mesic soil western soapberry was the only tree species. Shrubs were extremely scarse with infrequent individuals of greenbriar (Smilax bona-nox) and skunkbush sumac (Rhus trilobata= R. aromatica). Note prolific regeneration of western soapberry.

Lucy Park, Wichita County, Texas. October. Estival aspect.Perimeter of FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosytem). Given the western soapberry consociation and absence of other tree species in this local vegetational zone Kuchler and SAF units were deemed inappropriate; this vegetation was the edge of such units designated above for the bottomland forest. Loamy Bottomland range site. Central Great Plains- Broken Red Plains Ecoregion, 27i (Griffith et al., 2004).

461. Western soapberry zone of bottomland deciduous forest- Plant community of a gallery forest along the meandering Wichita River in an area transitional between mixed and tallgrass prairie in the Rolling Red Plains. Except for an incidental individual shrub like greenbriar or skunkbush sumac soapberry was the only woody species present. The understorey was primarily western wheatgrass, purpletop, little bluestem, and the naturalized Johnsongrass (Sorghum halepense). Buffalograss and hairy tridens (Tridens pilosus= Erioneuron pilosum) were locally abundant and added a shortgrass component so as to form a two-layer herbaceous understorey of a patchwork appearance.

Lucy Park, Wichita County, Texas. October. Estival aspect. Perimeter of FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). Kuchler and SAF units were not descriptive but were given above for the more mesic and typical plant communities of this range cover type. Central Great Plains- Broken Red Plains Ecoregion, 27i (Griffith et al., 2004).

462. Western soapberry (Sapindus saponaria var. drummondii= S. drummondii)- Young plants of western soapberry in early fall (October, but still estival aspect) in fruit. Lucy Park, Wichita County, Texas.

463. Leader of western soapberry- Compound leaves and large inflorescence of western soapberry. This specimen was thriving in the western Ozark Plateau. July. Newton County, Missouri.

464. Infloresscence of western soapberry- An immense flower cluster of western soapberry on a tree that grew on the easternmost extension of this species' range. July. Newton County, Missouri.

465. Leaves and fruit of western soapberry- Lucy Park (just above Wichita River), Wichita County, Texas.

466. White Christmas in the Ozarks- Snowfall (especially of major accumulation like a half foot or more) is rare in the oak-hickory forests of the Ozark Mountains. In fact, it is not unusual for there to be winters without snow and with winter precipitation being limited to rain or the treacherous "winter mix" of rain, freezing rain, sleet, and snow (ice storms). The "winter wonderland" of wet snow and an accumulation of 10 inches shown in these photographs demonstrated the ever-changing weather of the Ozark Region and the necessity of some winter adaptation for it's species. Besides it was "kinda purty".

Post oak and black hickory were dominant tree species with northern red oak and black cherry the major associates of the canopy layer. Understorey was dominated by buckbrush and poison oak; Virginia creeper formed a uniform intermediate layer extending from ground level to tops of trees while wild grapes (Vitis spp.) were local dominants of the intermediate layer. In early spring the herbaceous layer was usually composed exclusively of Mayapple (Podophyllum peltatum) colonies. All plant life had settled in for a short winter nap in this scene which had all but entirely melted within 24 hours of these photographs.

Ottawa County, Oklahoma. Christmas Day, 2002.

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Southern and Central Forests - IC

Forest Range Types of Eastern North America

Sugar Maple-Dominated Forests

Savanna of Oak-Hickory Forest and Tallgrass Prairie

Dominant, Associate, Widespread, and Just Plain Interesting Plants of the Central and Southern Forests

Bottomland Forests

Pecan Bottoms- Example of Southern Bottomland Forests

Eastern Cottonwood (Populis deltoides)-dominated bottomland forests

Some "Odd Lots" Forests

Dominant, Associate, Widespread, and Just Plain Interesting Plants
of the Central and Southern Forests