Southern and Central Forests - IIB

Forest Range Types of Eastern North America

154. Progress of growth- Growth of coppice shoots of the black walnut introduced in the immediately preceding three slides five weeks thereafter. This tree was a member of mixed hardwood forest that developed on the floodplain of Modoc Creek in the western Springfield Plateau. Record drought and heat existed at time of photograph so growth was much less than would have been typical.

Black walnut produces an alleochemical, juglone (5-hydroxy-1,4-naphthalenedione) is an enzyme that inhibits respiration (Rice, 1984, p. 77) so as to be toxic to numerous plants (as well as insect herbivores) though not to all climax trees. For example, oak, maple, elm, and hackberry species as well as such shrubs including pawpaw poison ivy, grape, and Virginia creeper are tolerant of juglone (Ontario Ministry of Agriculture, Food, and Rural Affarirs, un dated). In other words, just about all the plant species growing in this bottomland forest. In this author's observation black walnut generally does not make palatable browse for white-tailed deer perhaps due to presencce of juglone although Burns and Honkala (1990) noted that deer ate buds of black walnut.led deer. On this same floodplain forest white-tailed deer browsed slightly on coppice shoots of sycamore and heavily stump sprouts of American elm and white ash.. Speaking of which, proceed to next and subsequent slides...

Modoc Creek floodplain, Ottawa County, Oklahoma. Early July; early summer.

155. Another tree that won't stay dead (and one with an additional handicap)- Stump sprouts in American elm that had been felled on floodplain of Modoc Creek in the preceding late summer-early autumn. Unlike coppicing black walnut featured immediated above, stump sprouts of elm were palatable to white-tailed deer which had severely browsed these shoots. Such defoliation could have had an adverse impact on tree regrowth or, alternatively, such browsing could have had a net beneficial effect on the tree by eliminating some of the competing shoots and--if regrowth of some shoots was left untouched--resulted in increased growth with elimination of apical dominance. At time of photograph every elm shoot had its apical bud removed by browsing deer, and this was early summer in a severe drought year. Deer-browsing can be detrimental to forest regeneration.

Some Virginia creeper and silky wildrye was growing by stump shoots.

Modoc Creek floodplain, Ottawa County, Oklahoma. Early July; early summer.

156. Two approaches to tree regeneration- Edge of a local clearcut patch of mixed hardwood bottomland forest that had been high-grade logged in late summer-early autumn of last year. Recovering forest vegetation included seedlings of American or white elm and both seedlings and stump sprouts of white ash demonstrating both sexual and asexual reproduction of tree species that were associate species of the climax forest. Two stumps of white ash were visible: one in left midground and one in background by standing bole (see also immediately below). American elm also had numerous stump sprouts as shown in the preceding photograph. Some browsing on a few elm seedlings by white-tailed deer was evident in this photograph. Some of these seedling had obviously germinated and emerged in the late autumn shortly after logging.

Other species in this "photoquadrant" including coppice shoots of spicebush, a dominant shrub of the understorey, that had been topkilled by winter temperatures as low as -20 degrees fahrenheit (an "Alberta clipper" combined with a blizzard in early February). Considerable cover of Virginia creeper was present. Dead buff-coplored ground cover was common chichweed, a naturalized Eurasian annual weed. There were also some plants of silky wildrye; these had not been grazed.

Modoc Creek floodplain, Ottawa County, Oklahoma. Early June; late spring.

157. More tree regereration- Interior of the local clearcut patch of mixed hardwood bottomland forest shown in the preceding photograph. First of these two photographs was a general view of recovering forest vegetation whereas second slide was a closer view of plants in right foreground of first slide. Inside this small area of forest denudation there were seedlings of trees that were either climax or seral species that persisted into the climax stage as well as herbaceous species. The latter included silky wildrye, James caric sedge, and Davis caric sedge. Tree seedlings in first slide were (left to right): western hackberry, northern red oak, and boxelder. The second slide featured the northern red oak and boxelder.

The taller seedlings had emerged and made some growth in late summer to early autumn of the previous year shortly after destructive high-grade logging by timber theives. (This land was owned by a neighbor who did not want his land logged, but woodcutters legally logging the adjoining property stole timber off of this absentee landowner.The theives also stole his metal Powder River gate. These bastards were bad to the bone.)

Modoc Creek floodplain, Ottawa County, Oklahoma. Late May; late spring.

158. More seedlings and sprouts later on- Seedlings of boxelder, northern red oak, and hackberry along with American bladdernut (left background) and resprouts from topkilled spicebush (left foreground) on a small local area that had been clearcut under high-grade logging late in the preceding year. Some of the larger seedlings (especially those of boxelder) appeared to have started growth shortly after logging in late summer or early autumn last year so as to be in about the middle of the second growing season. Other seedlings were of such small size as to indicate germination/emergence during the current (first-year) warm-growing season. Spicebush shoots also originated during the present spring-summer season following topkill from severe winter cold (temperatures at least -20 degrees fahrenheit) of a "blue norther" combined with a blizzard (early February).

A mixed hardwood bottomland forest in western Ozark (Springfield) Plateau.

Modoc Creek floodplain, Ottawa County, Oklahoma. Mid-July; early summer.

159. Daring to grow back (apparently at least the second time)- Initial growth of stump sprout or coppice shoot of white ash from trunk of a tree that had been felled the previous late summer-early autumn. This tree had two trunks that almost assuredly were themselves stump sprouts from logging approximately 60 years ago. (The author remembered from his boyhood when this forest had been partially logged and he had watched the forest "grow back".) These trunks were on the edge of a tract of mixed hardwood creek bottom forest on land owned by a neighbor that adjoined the part of this tract that had been contracted for logging. The railroad tie that served as a corner post was all that remained of the fence between the two properties, but it was conspicuous.

Most of the better timber--including every logged black walnut--was on the neighboring property. Woodcutters stole much of this higher-quality timber, seemingly as much as they thought they could "cover up" or get away with, including the log taken from the stump shown here. They left the other part of the white ash (outer trunk and crown) which was the same diameter and bole length as the one that was stolen. The logical conclusion drawn from investigation of this forest crime scene was that the uncut trunk was left as "cover" for the crime of timber theft. Such is the working of the criminal (and, in this instance, likely retarded) mind.

Note the Virginia creeper growing its way up the stump. Live stumps with the ability to coppice such as this one are known to foresters as stools (Helms, 1998).

Modoc Creek floodplain, Ottawa County, Oklahoma. Early June; late spring.

160. Five weeks of growth minus some- Five weeks after beginning growth from a stump of white ash a cluster of coppice shoots or stump sprouts had achieved this growth. Roughly half of the shoots had been browsed by resident white-tailed deer to the degree that most of the leader length of the browsed shoots had been removed. Deer-defoliated ash shoots were visible at base of stump and in greater detail in the next two-slide set.

Stump sprouts or coppice shoots are long shoots, shoots with a relatively large terminal (apical) bud with comparatively little branching or floral tissue that make rapid vegetative (vs. sexual) growth. Carboydrates stored in roots and, to some extent, in, cambium of shoots can be quickly converted and translocated as glucosethereby permitting tremendous vegetative growth of long shoots. Long shoots are sometimes called water shoots.

Modoc Creek floodplain, Ottawa County, Oklahoma. Mid-July; early summer.

161. One year later- Coppice sprouts of the same white ash stump that was shown in the two immediately preceding slide-caption sets. This set of stump sprouts, stump offshoots, or water shoots were midway through the second growing following felling by timber thieves. This was also the second consecutive year of Extreme to Extraordinary Drought (as measured on the Palmer Index).

The apical bud (apical meristem) had grown above reach of resident white-tailed deer which had removed most of browsed leaders the preceding spring-summer season. Now (currently) any browsing would have to be limited to lateral branches coming off of of the main coppice shoot, the apical meristematic tissue of which was now out of browsing range. This stump sprout--this clone or ramet of the original tree (the genet)--was no longer vulnerable to serious damage by browsing deer. Fast regrowth of coppice shoots is an example of natural selection and adaptation to defoliation.

Modoc Creek floodplain, Ottawa County, Oklahoma. Late July; early summer.

162. General view in the fourth year- Lower trunk of a white ash that had two trunks (ie. a forked-trunk tree) from which one trunk had been felled four years earlier by timber theives who left the outermost trunk in a vain (and stupid) attempt to hide their theft from the land-owner. There were two healthy, surviving coppice shoots or basal trunk sprouts from the stump of the removed trunk. These two coppice shoots were shown in the immediately preceding slides at two years post felling and in detail at four years post felling in the two immediately following slides.

Modoc Creek floodplain, Ottawa County, Oklahoma. Late July; early summer.

163. Four years later- Coppice sprouts of the same white ash stump that was shown in the six immediately preceding slide-caption sets. This set of stump sprouts, stump offshoots, or water shoots were midway through the fourth full growing following felling by timber thieves. This was also the fourth consecutive year of Extreme to Extraordinary Drought (as measured on the Palmer Index).

These two coppice shoots were only slightly larger than they had been two growing seasons earlier, but they were surviving (in contrast to coppice shoots of neighboring sycamore stumps which had died by the fourth and, in some cases, by the fifth year post felling). At this juncture, white ash reproduction (asexual) by coppice shoots appeared to be a viable form of reproduction and, at least partial, forest regeneration.

Modoc Creek floodplain, Ottawa County, Oklahoma. Late July; early summer.

164. Minus some- Degree of browsing on white ash coppice shoots by white-tailed deer. Two views of the white ash shoots introduced above. The first view showed the base of the live stump or stool (Helms, 1998) with adjacent forest vegetation which included silky wildrye, Short's caric sedge, horseweed or mare's-tail, Virginia creeper, trumpet creeper, poison ivy/oak, and several seedlings of American elm. This latter, one of the climax dominants of this mixed hardwood floodplain forest, reproduced by both asexual reproduction through stump coppicing and sexual regeneration as seen here.

The second view was a close-in view to give a detailed view of the pattern of browsing by white-tailed deer. Note from these two photographs and the one immediately above that some coppice shoots were not browsed (at least not at time of photographs). The possibility that defoliation of some shoots could result in more rapid growth of undefoliated shoots that could then rise beyond reach of browsing deer. Removal of some shoots and reduction in overall apical dominance could have resulted in elevation of remaining shoots out of reach of deer. Scientists must retain a degree of objectivity and consider as many possibilities as they can think of.

Modoc Creek floodplain, Ottawa County, Oklahoma. Mid-July; early summer.

165. Another white ash daring to grow back- Another stump resulting from timber theft on a mixed hardwood bottomland forest had coppiced and produced stump shoots of this size by late spring of the first full growing year after felling. This log was stolen in late summer-early autumn of the previous year and, unlike the white ash stump shoots of the preceding example, had initiated growth of these shoots soon after harvest (and quickly before the first fall frost).

This tree was growing farther inside the forest that was on adjoining land than the two-trunk ash seen above. Timber theives stood a better chance of stealing this log without detection than trunks that were standing closer to the edge of the forest.

Younger and/or smaller trees with generally thinner bark of more likely to produce stump sprouts than are older trees with thicker bark. Living stumps that can produce coppice shoots are known as stools (Helms, 1998).

Modoc Creek floodplain, Ottawa County, Oklahoma. Early June; late spring.

166. Five weeks of growth on this white ash- Five weeks following onset of coppice shoot growth of a white ash introduced in the immediately preceding slide, this much more shoot growth had taken place (and in a severe drought). This set of ash resprouts had largely escaped deer browsing. There were two young poles of western hackberry in left background and one pole-sized white in right background of first photograph. In the second slide here were some leaves of trumpet creeper visible in fromt of ash sprouts.

Modoc Creek floodplain, Ottawa County, Oklahoma. Mid-July; early summer.

167. Splintered wood, shattered dreams (but the dream lived on)- Hatchet-happy, don't-give-a-damn woodcutters carelessly felled two mid-sized white ash trees on top of this young black walnut (the would-have-been next lumber crop) breaking it completely off somewhere between ten to twelve feet above ground level. The young walnut (pole-sized; ready for sale as a veneer log in about a quarter century) "bravely" sent out coppice shoots from intercalary meristem.all along its trunk. The tree's "will to live" prevailed and, in spite of a blasted trunk, was in process of recovering. Obviously this tree will never produce a log of any consequence though it may develop cavities that provide shelter for wildlife.

This black walnut tree was stolen the same as the two white ash (stumps of both shown immediately above) that brought it down. Contrary to popular assumptions, timber theft is a major problem in areas where land-holdings are of small acreages. Forest robbery is not limited to land of large timber companies. In fact, "little men" landowners probably are more likely to be robbed of valuable timber (in some cases black walnut shade trees from front yards) because access to trees is more available and small land-owners less apt to be aware of the threat. (This is a parallel to cattle rustling from hobby stockmen often being more likely and commonplace than from large ranchmen who "know the ropes".) Local hill talk was that this wood was stolen to support a drug addiction.

Modoc Creek floodplain, Ottawa County, Oklahoma. Early June; late spring.

168. A recovering (and surviving?) sycamore- Coppicing of a sycamore felled the preceding late summer-early autumn (this same stump was shown in the preceding section dealing with the cut-and-run logging of this mixed hardwood bottomland forest. This stump was readily identifiable by the chunk of wood that remained attached to the stump surface when fellers either failed to made a large enough backcut and/or failed to properly place or follow through on the undercut. Also serving as a "landmark" was the hackberry sapling left uncut to immediate right of the sycamore.

This hackberry sapling was left uncut so that is canopy would serve as cover to hide (to the extent possible) from exterior of this forest tract the crime scene of this stolen sycamore. (Larger boxelder, hackberry, American elm, and chinquapin oak growing deeper inside the forest were senselessly felled by hatchet-happy fellers.) This sycamore was obviously growing on neighboring private property several feet from the fenceline. Woodcutters legally logging the adjoining part of this forest robbed the absentee landowner of his timber. This tree as well as the black walnut and white ash shown immediately above were all stolen from their absentee owner. Story in the woods was that the money from this timber theft was used by the theiving neighbor to support his drug addiction.

Modoc Creek floodplain, Ottawa County, Oklahoma. Early June; late spring.

169. Deer were at it again- Browsing by white-tailed deer on apical parts of stump (coppice) shoots of sycamore. This was a nice example of the phenomenon of apical dominance. Deer removed the apical (terminal) bud and upper part of this shoot whereupon the axillary bud below the removed (eaten) internode developed. Removal of the apical or terminal bud, which was the source of auxin that suppressed ("dominated") lower meristems (meristematic tissue), permitted development/differentiation of the subtending (lower or beneath) bud.

On the deer range of this mixed hardwood bottomland forest coppice shoots of sycamore were less palatable than coppice shoots of American elm andwhite ash, but more palatable than those of black walnut (described above).

Modoc Creek floodplain, Ottawa County, Oklahoma. Early June; late spring.

170. Sycamore stickin' to it- Growth of coppice shoots on felled sycamore five weeks after growth shown in two immediately preceding photographs. This was the same stump shown in the preceding section that described high-grading of this mixed hardwood bottomland forest. The sapling of western hackberry growing at immediate right of the stump in these (and above section) photographs served as a "landmark" as, for that matter, did the wood remaining on this stump that resulted from improper felling (either too small a backcut or incomplete uncercut). The hackberry sapling was left so that its canopy could serve as some cover of the crime scene when viewed from the exterior of the forest tract. (Other, and much larger hackberry, American elm, and boxelder trees growing in the interior of the tract were needlessly felled by the hatchet-happy theives.)

This stump had been left (the tree felled) in late summer or early autumn of the previous year and the stump sprouted almost immediately. Thus shoot growth seen in this photograph was that of a short, partial growing season and that through early summer of the first full growing season (ie. the "hind end" of one growing season and the "fore end" of a second growing season).

This sycamore stump constituted a stool, a live stump capable of growing shoots or coppicing (Helms, 1998).

Modoc Creek floodplain (tributary into Modoc Creek), Ottawa County, Oklahoma. Mid-July; early summer (Extreme Drought, Palmer Drought Index).

171. Sycamore stickin' to it the next year- Coppice shoots of the sycamore stool shown imediately above in mid-summer of the second full growing season after felling. These stump sprouts had attained this growth during two consecutive years of drought which at time of photograph as reached the highest stage of drought severity, Extraordinary. This photograph was taken from a greater camera distance and slightly different camera location from that of the photograph immediately above.

Actually, this tree had been felled (the log was stolen) in late summer or early autumn and had sprouted soon thereafter. Shoot development seen immediately above was in the first full growing season following logging so has to have growth made in one growing season and the "butt end" of another growing season (ie. a partial growing season).

The pole-sized bole at left margin was American elm. Large saplings and small poles in distant background were of hackberry and American elm, the ultimate co-dominants of the climax bottomland forest.Other plant species present included Canada wildrye and resprouts of sawed-off fox grape.

Modoc Creek floodplain (tributary into Modoc Creek), Ottawa County, Oklahoma. Late July; mid-summer (Estraordinary Drought, Palmer Index).

172. So much for conjecture- By fourth warm-growing seasons after logging of a middle-aged sycamore all of the coppice shoots ("stump aprouts"= "sucker sprouts") were dead except for one. One surviving shoot would be all it would take to replace the felled sycamore, and besides all of these "water shoots" or "sucker sprouts" could survive anyway. Sadly, however, look at the next, the fifth (and final) warm-growing season in the next slides ...

By the way, the shoot on right side of stump was hackberry, the number one climax dominant of this mixed hardwood bottomland forest. Two trees to left and rear of the stump were American elm and hackberry. Young seedlings in foreground were of pignut or bitternut hickory. All of these are climax tree species for this forest range site.

Bank of Modoc Creek, Ottawa County, Oklahoma. Late July.

173.

174. Sycamore that started a year later- Stool, a live stump capable of coppicing (Helms, 1998), of sycamore with young coppice shoots approximately midway through the second full growing season following felling by timber thieves. Unlike the sycamore stool shown in preceding photogrphs, this living stump did not send up new shoots (reprouts) the first year after felling, but instead "waited" until the second year post-felling. Actually the sycamore stool and shoots presented immediately above sprouted almost immediately following felling which took place in late summer or early autumn. Those coppice shoots had grown for a brief period of one summer/autumn season and throughout the entire first full growing season thereafter. In other words, those stump sprouts had grown parts of two years more than the ones here that had just recently developed. Reason(s) for later or delayed development of this stump was not known.

Immediate bank of Modoc Creek, Ottawa County, Oklahoma. Late June; early summer (Extreme Drought, Palmer Drought Index).

175. Sycamore sprouts did not make it- The sycamore stump introduced immediately above now shown in the fourth full warm-growing season post felling of the tree revealed that all stump sprouts (= stump suckers, coppice shoots, "sucker sprouts") were all dead except for one that was dying ("good as dead") and would be dead by end of this current growing season. Speculation that felling the adult tree resulting in stum shoots might extend life of the tree proved false. All the stump sprouts were good for was heralding that tree death would be prolonged and pathetic in the trees futile struggle for survival. Within four years of felling the last coppice shoots had died.

Coppicing was not a viable method for regeneration of sycamore on this bottomland forest site in the western Springfield Plateau. Even on the most favorable site for sycamore establishment and persistence, stump shoots succumed within four years of their initial growth. Timber theft had been responsible for loss of every adult and pole-sized subadult sycamore in this mixed hardwood bottomland forest. Worse, there was no sycamore regeneration through seedling establishment (sexual reproduction).

Sad ending (at least for foreseeable future) for this once diverse climax or late subclimax Ozark Plateau forest. This outcome continues to be the fate of most remaining forests in this area.

Bank of Modoc Creek, Ottawa County, Oklahoma. Late July.

176. Barely hanging on- In the fourth full warm-growing season post felling there was only one stump sprout or "water shoot" of this middle-aged sycamore still living on the bank of an Ozark Plateau stream. The log from this sycamore had been stolen and coppice shoots had sprouted from its stump in the second full warm-growing season after illegal harvest. It was noted that only one healthy, fast-growing shoot would be needed to replace the felled tree (which by the way was hollow and worthless as a saw log), but sadly even one surviving shoot was expecting too much. The next slides showed the fate by fifth (and final) growing season for stump sprouts...

Bank of Modoc Creek, Ottawa County, Oklahoma. Late July.

Do not live forever even at church- Stump sprouts or suckeres from the trunk of an immense sycamore that was at least 150 years old and that apparently died on natural causes. This sycamore was growing on the flood plain of Lost Creek (roughly 150 yards from the main stream channel). Age of the sycamore could be roughly estimated because it was a reasonably large tree growing on land that became the churchyard of the Presbyterian and, soon thereafter, Methodist Episcopal Church, South in Seneca, Missouri in the 1870s. Upper and outer parts of the sycamore starting dying in the second decade of the 21st century. By 2017 when these three slides were taken all its major limbs had fallen and the trunk had started to decompose and rot away (seee especially the third slide). Several coppice or adenntituous shoots, including several smaller ones higher up on the trunk (first and second slides) and a larger one from near bae of the trunk (third slide), developed and grew to sizes shown in these slides. The larger, basal sprout lived for five years until it died in 2023. This adventituous shoots appeared to arise from epicormic buds in intercallary meristem.

Though interesting and "promising" hope that these vegetative offshoots (at least the larger basal offshoot) would result in another sycamore tree of the same genotype ("a chip off he old block"), such was not to be the case. These vegetatively produced "daughter" shoots did not survive and grown into a replacement tree. Epocormic offshoots were only the rally bfore death, the last gasp before the last spark of life departed from the old tree.Things die even at church. Life after death "ain't" in the churchyard.

As was the case of stump sprouts in "sawed down" large, adult sycamore trees along Modoc Creek, coppice or other epicormic shoots of old sycamores sprout and shown considerable growth for only a few years before they die. This tree was about two miles away from (east of) sycamores along Modoc Creek.:

Sycamore regeneration does not occurr in old trees. Stump sprouts do not survive.

Seneca, Missouri, Newton County, Missouri. Late July.

178. Renewal in a bottomland forest- A western or common hackberry pole with small sapling of box elder at far left (left magin in first or vertical slide), stump sprouts or coppice shoots of white ash to immediate right of hackberry pole, seedling of white ash between box elder and hackberry pole, and seedling of sycamore near trunk base of hackberry pole (between this pole and white ash stump sprouts on a mixed hardwood bottomland forest disturbed by high grade logging five years prior to time of these slides. The amber-colored herbaceous understorey consisted mostly of Virginia wildrye.

Modoc Creek floodplain, Ottawa County, Oklahoma. Late July; mid-summer aspect. late spring. FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). Mesic Bottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

As if high-grade (and timber-theft) logging was not enough:

After having been in the same family for three generations a neighboring landowner bought the property on which the mixed hardwood forest that developed along Modoc Creek in the western Ozark (Springfield) Plateau and that had beeen high-graded by timber theives (as shown in detail immdiately above). The new owner sprigged a cultivar of sterile hybrid bermudagrass (Cynodon dactyledon) on part of the property, but the result was a poor stand of this introduced grass. Instead the "thin" stand of sprigged bermudagrass was invaded by naturalized annual ryegrass (Lolium multiflorum) which overwhelmed what was apparently a poorly adapted cultivar of bermudagrass. The cool-season annual ryegrass was absent from the field of bermudagrass in mid- to late summer, but from autumn through early summer ryegrass outcompeted the weakly established bermudagrass and appeared to prevent establishment of a stand of this introduced species.

The land owner could not kill out enough annual ryegrass with herbicide so he resorted to pasturing cattle on the ryegrass to graze it out in an effort to establish a stand of the non-competitive, "sickly" bermudagrass. The attempted biological control was only modestly successful (competition from ryegrass was apparaently not the only problem preventing establishment of this finicky cultivar), but cattle grazing was much more effective in reducing palatable woody species in the understorey of what remained of the mixed hardwood floodplain forest. It was shown below that native shrubs and seedlings of oak and hickory species in the forest understorey were much more palatable and preferred by cattle than any of the introduced grasses, both cool-season and warm-season, both planted and naturalized agronomic forages.

Further forest degradation was the result.

Selective browsing and regrowth- Pair of "nested photoplots" (second slide, close-up view of the immediate center foreground shown in first slide) showing selective browsing by beef cattle (cow/calf pairs) of shrubs in the woody layer of a mixed hardwood floodplain forest in the western Ozark (Springfield) Plateau. These two slides were taken less than one month after removal of cattle thus showing rapid regrowth shortly after excessively heavy defoliation (overuse of browse) of American bladdernut (Staphylea trifolia). Simultneously, there was zero use (not a single leaf touched) of spicebush (Lindera benzoin).

This was the scene after the first year of cattle browsing. As proven below, cattle were the only ungulate species that ever touched American bladdernut (just be patient; "don't git your panties in a wad").

The first or upper "photoplot" presented: 1) regrowth of American bladdernut (most of foreground) following overuse, 2) completenonuseof spicebush (larger leaves in foreground), and 3) unbrowsed American bladdernut (taller, darker-green leaves) in bckground. Cattle could not get through the dense shoots of American bladdernut--even after severe overuse reduced height of shoots to less than one-half of unbrowsed shoots--to broswe on the farther-back plants.

The second "photoplot" was the nested portion of the immediate foreground of the first "photoplot". Plants with larger, oblong leaves in the right foreground of this slide were spicebush which neither beef cattle nor white-tailed deer fed on.

These two slides presented American bladdernut in a partial opening in the bottomland forest. This immediate location or microhabitat received "full, direct sun" throughout most of the day light (diurnal) period.

Floodplain, Modoc Creek, Ottawa Count, Oklahoma. Late June; early estival aspect, regrowth of American bladdernut. FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). Mesic Bottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

More selective browsing by catle and woody regrowth- Regrowth of northern red oak (immediate foreground) and of Americn bladdernut (mid- and background) in the understorey of a mixed hardwood bottomland forest less than one month following removal of cow/calf pairs from this forest range. Upper portions of northern red oak and upper and outermost parts of American bladdernut had been eaten by cattle and subsequent regrowth of both woody species was paresented in this slide.

The author had run this bottomland forest range over a span of more than half century (since a grade-school boy) during which time white-tailed deer frequented the habitat. No cattle had grazed this unfenced property during that time. Your author cannot recall ever seeing any deer browsing of any northern red oak, chinquapin oak, bitternut or pignut hickory, or sycamore trees. He had observed removal of some buds of American elm and tips of box elder which would would have had to have been by deer.

Likewise, white-tailed deer never touched American bladdernut. (Slides of unbrowsed American bladdernut located less than 50 of the author's steps away from the heavily browsed bladdernut shown here that was fenced out from cattle yet accessable to white-tailed deer proved this statement unequivocally. These slides were presented below.)

Thus, any browsing of oaks, hickories, and American bladdernut had to be that of cow/calf pairs.

This photograph was taken at the edge of the bottomland forest that received sun light for much of the daylight hours.

Valiant effort of the part of a native American shrub- Excessive defoliation (overuse) of American bladdernut in the understorey of a mixed hardwood bottomland forest by cow/calf pairs less than one month after removal of cattle. Defoliation of shoots included trampling as much, if not more, than actual eating although removal of shoot portions by mouth was obvious. (As explained above and below, resident white-tailed deer never so much as touched American bladdernut in this floodplain forest.)

What appeared to be two local colonies of American bladdernut were presented in these two potographs. Both of these local stands had received more browsing than those presented in the two immediately preceding slide/caption units plus they were farther back in the forest that received less light than was the case for the preceding examples. The conspicuously less regrowth of the two stands shown here was a result of those two combined factors.

Even with that, American bladdernut showed a remarkable capacity to replace defoliated parts. This as the first year of animal browsing in the history of these shoots. Such rates of recovery seemed highly unlikely (probably impossible) after several bouts of excessive browsing. Extreme breakage of shoots--by both trampling and feeding--was conspicuous in both of these slides.

Furthermore, presence of green leaves was deceiving without description of the plant life seen in these two "photoplots". Green leaves in the first "photoplot" included those of spicebush, a native shrub that, on this range, was not touched by cattle or deer. Forbs were absent from this heavily browsed stand of American bladdernut. Many of the leaves in the second "photoplot" were those of giant ragweed (Ambrosia trifida) and the naturalized beefsteak plant (Perilla frutescens), a naturalized invasive exotic from the subcontinent of India. Both forbs are annual species that pioneer areas of disturbnce. Neither species can compete with perennials, and they werer absent from neighboring, unbrowsed, fenced-from-cattle stands of American bladdernut.

Any decent student with one course in Range Management "under his belt" can see where the direction of vegetation change in these two "photoplots". Plot 1 is "on target" to become a "solid stand" of spicebush while Plot 2 is on its way to becoming a weed patch. As pointed out above, this was the first time cattle had grazed this forest range in over a half century and white-tailed deer never touched American bladdernut.

Note on morphology: American bladdernut (Staphylea trifolia) forms colonies from a well-developed network of woody rhizomes ("rootstocks"). This author never encounteded American bladdernut in the Ozark Plateau Region as a single multi-shooted individual let alone as a single shoot (tree-form) individual. It always grows as a colony of shoots.

Another example of selective broswing, limited regrowth, and retrogression)- A local "spot" or micro-location that showed part of a (former) colony of American bladdernut in the understorey of a mixed hardwood bottomland forest less than one month after excessive browsing (both eating and trampling) by cow/calf pairs. Shoots of this obviously very palatable (to beef cattle not white-tailed deer) had been stripped of leaves to heights of three and a half feet. There was breakage of shoots due to both trampling and feeding.

Most of the green leaves in thee two views were those of giant ragweed and the exotic (from India) beefsteak plant. Both of these annual forbs are pioneer or colonizing species of disturbed habitats such as the heavily denuded forest understorey seen here.

This first-year disturbance furnished a textbook example of 1) selective and excessive browsing and 2) the onset of range retrogression. Retrogression as applied to range (as in range retrogression) is synonymous with range degeneration: "The process whereby the same area becomes sucessively occupied by different plant commnities of lower ecological order" (Kothman, 1974). Retrogression is the retrograde change on a given sere such that plant communities (vegetation) of consecutively lower successional order occupy that land. Retrogression is the opposite process (or the ecologically "reversible reaction") of succession, although the order of (or even the same) seral communities may not be in the exact order of change.

The term overbrowsing--like overgrazing in reference to herbaceous plants--is prolongued overuse leading to deteriorated range as defined and shown by downward (retrograde) changes in composition, structure, etc. in the range plant community to lower stages of vegetation on the sere (the natural potential plant commnity for that particular area of ground, including soil, relief, exposure, etc.). Overbrowsing typically requires several grazing seasons (or years) of overutilization (the meaning of prolongued overuse) before there are noticeable shifts (relative changes) in species composition in the range vegetation.

In the instances shown here, however, the climax (and extremely palatable to cattle) American bladdernut was already being replaced by a pioneer annual composite and an invasive exotic. In this instance, overuse and overbrowsing were one and the same. The native woody understorey of this mixed hardwood bottomland forest was being replaced (instaneously in plant succession time scale) by annual weeds. Overstocking (having more animals on the range than the natural vegetation can sustain; range resource exploitation) with beef cattle had started to destroy the woody component of this forest range in one short grazing season.

No cows, all sorts of deer, no browsing of bladdernut- Two views of a one colony of American bladdernut growing in an area of a mixed hardwood bottomland forest range from which cattle had been excluded for over a half century yet easily accessible to resident white-tailed deer. As the author specified above, in "running these woods" over a span close to 60 years, this author never found any browsing of American bladdernut by deer.

The colony presented here was only 50 to 60 of the author's steps away from the overbrowsed (by beef cattle) colonies of American bladder shown in the immediately preceding four slide/caption units. All of these slides were taken within a quarter to half hour of each other. Complete absence of deer feeding onbladdernut in the area from which cattle were excluded and the heavy feeding and trampling by cattle in the portion of this bottomland forest accessible to both ruminant species was unequivocal, self-evident evidence that defoliation was solely by cow/calf pairs.

This case provided a textbook example of animal species-specific preference (and consequent selectivity) for specific plant species. It is also an example that there are exceptions to "thumb rules" such as "cattle prefer and eat grass; deer prefer and eat shrubs".

The tree trunk at right margin of the second slide was that of a "prime adult" northern red oak.

Browsed off, regrew; others untouched- "Photoquadrant" of the lower woody layer in a mixed hardwood bottomland forest range in the Springfield Plateau after being browsed by cow/calf pairs throughout the spring season. Seedling of chinquapin oak that was browsed by beef cattle to the level at which two uppermost branches originated and subsequently in less than one month's time had made this much regrowth. Leafy shoots behind the chinquapin oak seedling were those of spicebush, none of which had received any use by cattle or white-tailed deer.

Another example of grazing selectivity and an eample of tree seedling regeneration.

It was remarked above that this photographer never found any evidence of deer browsing on any oak or hickory species in this bottomland forest over a span of over half century. Deer feeding in this mixed hardwood forest wwas confined to white ash, eastern cottonwood, and boxelder among trees and mast (mostly acorns), forbs, caric-sedges, and, infrequently, grasses like Elymus species. Cattle had not been stocked on this unfenced property for 50 or 60 years. The slides shown here were taken following he first grazing season by cattle.

Based on these observations plus cow tracks, dung, and cattle hair that was rubbed-off on tree trunks, it was obvious that browsing on oaks, hickories and shrubs like Americn bladdernut was solely by cattle.

Moving upstream: An ephemeral stream that was a tributary of Modoc Creek entered immediately upstream from the bottomland (floodplain) forest that was just described. In summary that forest which was developing on the greater floodplain of Modoc Creek was progressing through processes of plant succession from a subclimax sycamore-eastern cottonwood- northern red oak-Shummard oak-dominated community to a climax western hackberry-American elm-white ash-bitternut hickory forest.

Another tract ("branch" or "arm") of that bottomland forest had developed along the tributary of the "wet weather" drainage. This forest range community was contiguous with the one just presented. It bordered on the other side of the old fiield shown and described immediately above. The upstream bottomland forest range was slightly above the floodplain of Modoc Creek though, again, conterminous with it. Forest vegetation on the Modoc Creek tributary had developed on a less mesic forest site with shallower and, almost assuredly, less fertile soil.

Many of the same plant species were common to both floodplain and tributory forest range; however, there were conspicuous and ecologically meaningful differences between the two tracts of bottomland forest including those in forest structure or architecture, age class structure of trees, density and cover of trees, and herbaceous layer (in addition to those in plant species composition).

The most important and practical difference (at least from standpoint of this photographic presentation) was in light conditions, both intensity and duration. This was due to greater density of smaller and generally less mature trees in the tributary forest the result of which--from much of spring through to leaf fall during mid-autumn--in more complete and denser canopy cover. This vegetational structure excluded most light from the lower forest layers and permitted much shorter periods of light when photosynthetically active radiation could penetrate the upper forest cover. As such, photographing of this forest vegetation--especially the herbaceous layer-- and details of tree trunks could only or, at least, best be achieved during winter. Even then the angle of winter sunrise and sunset was so much farther to the south that there was never a period of time when direct light reached more northerly exposures. This resulted in dimlite appearances combined with shadows from tree trunks as the bane of this winter forest photographer. Viewers will have to suffer along with him at some shots.

179. Seral and climax- Second-growth mixed hardwood forest on tributary of Modoc Creek (western border of Ozark Plateau) immediately above creek's floodplain with western hackberry (the climax dominant) and American elm, slippery elm, pignut or bitternut hickory, mockernut hickory, and white ash (climax associates or, locally, co-dominants in various combinations with western hackberry) succeeding black walnut and honey locust. Chinquapin oak, boxelder, and red mulberry were also present at meaningful porportions of relative cover. The successional status of these latter three species was unclear.

The major shrub was buckbrush or coralberry on land having less tree density and canopy cover and fox or frost grape where tree density and crown cover were greater.Common greenbriar, poison ivy, and trumpet creeper were present sporadically. Other than local habitats where buckbrush was dominant, the preeminent part of the understorey of this tributary forest was the herbaceous layer which consisted almost exclusively of Canada or nodding wildrye. This was strikingly different from the more mesic forest on the Modoc Creek floodplain which supported a more diverse understorey (see above) including an herbaceous layer dominated by silky and Virginia wildrye.

The first of these two photographs was a second-growth bottomland forest dominated by pignut or bitternut hickory (two foremost trees in center to right foreground, the clump of three trunks in center background on bank of the ephemeral stream channel) and hackberry (largest tree that was also on stream bank [at left margin of midground], large sapling in front of the clump of pignut hickory in background). Both pignut hickory and western hackberry were climax species. There were also a few trees of mockernut hickory (Carya tomentosa= ) that were indistinguishable in background. There was regeneration of both Carya species represented by various age classes from large seedling through sapling and pole to mature tree. Dead shoots from last year and green shoots of the current cool-growing season of Canada or nodding wildrye (both visible in foreground) indicated the sole dominant of the herbaceous layer of the understorey.

In the second photograph young hackberry (foremost pole-sized tree in left foreground; sapling in right midground; "sliver" of a trunk at left margin) were succeeding two log-sized, mature black walnut (diagonally opposite each other in midground) while white ash (three large trees in right background), pignut or bitternut hickory (foremost tree situated between the two walnuts) and boxelder (tree to left of the pignut hickory in far background) persisted (probably as climax) into this subclimax state forest. Canada wildrye was present (and generally conspicuous) as both last year's dead and current cool-growing seson shoots.

Ottawa County, Oklahoma. Early January (hibernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). Mesic Bottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

180. Seral forest on a tributary of an Ozark creek- A second-growth mixed hardwood forest dominated by black walnut and honey locust had been successfully invaded by hackberry, slippery or red elm, and American or white elm which were succeeding the seral black walnut and honey locust. Fox grape (conspicuous speicmen climbing black walnut in right foreground of first photograph) and buckbrush or coralberry (inconspicuous in these photographs) were the major shrubs. Herbaceous layer was the predominant layer of this forest understorey. It was dominated almost exclusively by Canada or nodding wildrye. Winter annual grasses, especially Japanese chess (Bromus japonicus), was present, but stood no chance at widespread invasion given dominance by vigerous plants of the the robust native, Canada wildrye. Winter forbs were absent. Dead shoots of elephantfoot (Elephantopus carolinianus), the major warm-season forb, were present. Elephantfoot is a native perennial composite.

In the first photograph the three large trees on the right bank of this ephemeral channel were black walnut. The sapling on the opposite (left) bank was an American or white elm. Large tree at right margin in midground, sapling appearing to left of this tree and right of the three walnuts, and the largest tree in this scene (orn right bank behind the black walnuts and to front of a rotting snag ) were western hackberry which is the number climax dominant and the species with greatest recruitment (= regeneration). A large boxelder was near center of channel in distant background. Dead shoots of a dense colony of Canada wildrye were visible on the left channel bank in midground.

The second photograph presented a textbook example of forest succession with one seral stage replacing another and displayed dramatically by two seral species of aging trees being succeeded by two climax species with saplings. Black walnut, an Intolerant (Wenger, 1984, p. 2) seral species, represented by the two adult comparatively large trees in right foreground and one behind them, and honey locust, an Intolerant (Wenger, 1984, p. 3) seral species, represented by the two largest trees in center midground were being succeeded by western hackberry (foremost tree, a sapling, at far right; several small trees in badkground including the one behind the fox grape in center), American elm (sapling directly in front of second or center black walnut), and slippery or red elm (sapling behind foremost hackberry sapling and in front of foremost black walnut). Shoots, both dead and live, of Canada or nodding wildrye (the dominant herbaceous species) were conspicuous. Greater detail of this vegetation at local scale was presented in photograhs below.

181. Being replaced (more-or-less) while othr stay- Vegetation dynamics in a bottomland mixed hardwood forest on an ephemeral drainage in the Ozark Plateau. Dramatic example of elm replacing in successional order black walnut while younger trees of black walnut along with older trees of white ash, chinquapin oak, and western hackberry remain. Through all this turnover among tree species Canada or nodding wildrye (a climax decreaser) remained the overwhelming dominant of the herbaceous layer which was the predominant layer of the forest understorey.

The first photograph presented a broader view of a "photographic sample" of a local stand in which largest (and, by inference, oldest) trees were black walnut as represented here by the largest tree (left midground). There had been regeneration of the Intolerant (Wenger, 1984, p. 2) black walnut in the past as shown by trees of intermediate size such as foremost tree in immediate right foreground, but recent regeneration (recruitment) was of slippery or red elm as represented by the two saplings in front of big black walnut (left foreground), American or white elm, and western hackberry. Chinquapin oak (tree with light-colored bark to immediate right of big black walnut) and white ash such as the one opposite the chinquapin oak (right-center midground) persisted into what was developing into the climax forest vegetation. There were three large white ash in left background beyond the large black walnut. Big tree in center of distant background was an immense boxelder. Conspicuous shoots of Canada wildrye (several with clearly defined spikes such as those at base of big black walnut) proclaimed this species as the dominant of the herbaceous layer which was the most prominent part of the forest understorey.

Second photograph was a "nested photoplot" within the larger "photosample" jsut described. The two saplings of red or slippery elm in front of the large, mature black walnut that was "escorted" by a small log-size chinquapin oak. Three nice mature (log-size) white ash as well as the white ash opposite the chinquapin oak represented this climax tree species. Small pole-size tree behind and to immediate of the white ash in right foreground was American elm. The very large boxelder in center of distant background was also still shown. Various plants of Canada wildrye, which was the herbaceous dominant, were obvious (due largely to the prominence of last year's dead shoots).

These two photographs provided a textbook example of sucession and transition from one species to others of higher successional order. Black walnut, an Intolerant (Wenger, 1984, p, 2) and hence seral species (on this bottomland sere) was slowly being replaced through tree reproduction by red or slippery and American or white elm which, though somewhat variable, are generally Imtermediate in tolerance (Wenger, 1984, ps. 2-3). Likewise, white ash was also listed as listed by Wenger (1984, ps. 2-3) as Intermediate. For this forest cover type (SAF 93: ugarberry-American Elm- Green Ash) the Intermediate species comprise a variant form of the climax eolm-ash forest of Kuchler (1964, p. 101) and as recognized in Garrison et al. (1977, ps. 15-16). This forest range vegetation was in process of achieving species composition and structure of the terminal forest vegetation on this bottomland site .

182. A lesson in plant succession- Close-up "photoplot" of the large black walnut and one of two slippery elm saplings introduced in the two immediately preceding photographs. This is "a picture worth a thousand words". Intolerant black walnut being replaced by Intermediate tolerance elm (Wenger, 1984, ps. 2-3) as elaborated in the preceding caption. No further comment deemed necessary here. Canada or nodding wildrye was presented in clariity and promience appropriate for its presence as the overall dominant of the understorey. This natiive, perennial, festucoid grass was sole dominant of the herbaceous layer which was the most consistent and prominent stratum of this forest's understorey.

183. Another stand, a different seral dominant, the same successional story- Honey locust, as represented by two "really big" and senescing trees, was being replaced by younger trees of western hackberry and American and red elms on this sere of a bottomland forest site just above the floodplain of Modoc Creek near the western border of the Ozark Plateau. The first slide was at distance to give some idea as to physiogonomy as well showing internal structure and species composition of this form of a mesic bottomland forest in the ecotone between eastern deciduous forest and the vegetational mosaic of tallgrass prairie and tallgrass-hardwood savannah immediately to the west. Second slide presented sufficient detail for description of species composition, structure (arrangement) of forest vegetation, and progression of plant succession.

The two largest trees (center foreground and standing diagonal to one another) were comparatively ancient (over mature) honey locust with dead and dying limbs (complete with woodpecker nesting cavities) in their crowns and just enough thorns to proclaim their species identy. These "long-past-their-prime" senescing trees still looked out on younger honey locust trees (just beyond camera range) proving that their species' moment in the "successional sun" (span of time on this sere) was not completely over, but it was clear that their floristic reign as local dominant was past as honey locust was being replaced by both American or white elm and slippery or red elm which were dsstined to share climax dominance with western hackberry, the overall forest dominant.

American elm was represented by the largest tree and one small pole or large sapling to right-rear of the two honey locusts in both slides while slippery elm claimed the other large sapling or small pole by the big American elm in the first slide. Western hackberry was present as the young adult tree behind and immediate left of foremost honey locust (both slides), young adult tree to left of big American elm (second slide), and as small sapling (immediate left foreground of second slide). The aray of age classes of the Intermediate in tolerance elms and hackberry and Intolerant honey locust (Wenger, 1984, ps. 2-3) bespoke the inevitable replacement of honey locust by the hackberry-elm climax. This inexorable successional pathway, this progression along the sere, depends on absence of disturbance(s) severe enough that western hackberry and elm species are not halted in their advance so that honey locust (or species even more seral) reclaim their transitory throne.

A plant of fox grape provided a continuous "thread" from lowest (ground) layer to highest layer (foret canopy) of this forest plant community. Here, as throughout the rest of this forest range vegetation, Canada or nodding wildrye dominated (essentially, exclusively so) the herbaceous layer which was the prominent one of the understorey. Buckbrush or coralberry was also present, but its leafless, short woody shoots (foreground of both slides as in front of foremost honey locust trunk) did not adequately proclaim it to be the most widespread shrub on this forest range.

184. An opposite view of the stand just described- From the opposite end of the honey locust-dominated seral stand presented in the preceding two-slide set and caption the laina of fox or frost grape along with the short, woody shoots of buckbrush or coralberry (absent leaves and most fruit) presented the shrub layers of this bottomland forest range. Canada wildrye was present of course, but less conspicuously featured than in preceding slides.

Successional replacement of Intolerant honey locust by Intermediate tolerance (and climax on this forest site) western hackberry American elm, and slippery elm was again the central theme of this photograph. Hackberry was represented by the large sapling or small pole to the forward and left of the big honey locust. Both Ulmus species were well-represented by saplings and small pole-sized trees to left of and behind the honey locust. There were also older, larger elm trees in distant background.

This same local forest range vegetation was shown again below and from the same photo-point at a slightly farther distance in its advanced vernal aspect (late spring). And the old, slowly dying honey locusts were still alive.

All this forest vegetation was on the banks of an ephemeral stream that was a tributary to Modoc Creek, the floodplain forest vegetation of which was described in detail above.

and

185. Calf pen trees- Numerous shoots of chinquapin oak that developed as stump (epicormic) sprouts from the previous tree on a tributary of Modoc Creek in the western Ozark Plateau. The first two photographs (side-by-side) are of the same set of young adult clones, the second (right slide) being a closer view than that shown in the first. The third photograph was of a second set of epicormic shoots from a second previous tree. Agent of defoliation that removed the preceding tree trunks was unknown. There were no remains of stumps or logs. Based on surmise and fragmentary history of land use it seemed most likely that the former trees had been cut for firewood, but this was not known with certainity. It remained irrelevant because the original (at least preceding trunks or shoots) of both chinquapin oaks had been topkilled and the two genetic individuals resprouted. Burns and Honkala (1990) cited work which concluded that chinquapin oak readily resprouts resulting in shoots that are usually younger than their root systems. Resprouts (resgrown shoots) from sawed stumps are common.

Such epicormic or water shoots are clonal organisms. These coppices or clones are ramets of the original genet (the genotype or unique genetic individual). Such coppicing can result in shoots developing into trees that originated from trees of genetypes that appeared several "generations" (cycles of shoot cutting and regrowing) earlier.

Burns and Honkala,(1990) also explained that chinquapin oak is shade-intolerant, but young trees are less so than older ones. This means that regeneration of chinquapin oak is disproportionately greater than than indicated by tolerance of established individuals. Hence, chinquapin oak can reproduce asexually from stumps and, by relying to some extxent on reserve food stored in roots, grow into mature trees under some degree--perhaps substantial--of shading. These same authors interpreted chinquapin oak as climax on soils derived from limestone parent material even when these are droughty natured soils. Even on mesic forest sites chinquapin oak is "subclimax to climax" and present as part of the climax forest vegetation (Burns and Honkala, 1990).

Shoots originating from previous trunks (phenomenon of coppicing) can grow into mature trees as large or larger than the original or, at least., preceding shoots (trees). Guargantian clumps of resprouted coast redwood (Sequoia sempervirens) have clonal shoots which grow to such immense size that they are known reverently as "cathedral trees" (examples presented in the chapter, Coast Redwood Forests, under Woodlands and Forests). This Okie author has also been to the National Catheral in our Nation's Capital, but he feels more at home in the smaller Oakdale churchhouse. Having had experience with both beef and dairy cattle in the Ozark Region this photographer felt it appropriate to dub these clonal clumps of chinquapin oak "calf pen trees". Eat your evergreen, redwooded hearts out prune-pickers.

Ottawa County, Oklahoma. On a tributary of Modoc Creek. Early January; dormancy phenological stage.

186. Spring perspective on calf pen trees- The stump sprouts of chinquapin oak described immediately above in late spring. The herbaceous layer consisted of Canada . wildrye and silky wildrye; James caric sedge (Carex jamesii); Virginia creeper; and seedlings of hackberry, American elm, and bitternut hickory the climax dominants for this forest range type.

Ottawa County, Oklahoma. On a tributary of Modoc Creek. Early June (vernal aspect).

187. Stand of pignut or bitternut hickory (Carya cordiformis)- Local stand of pignut hickory on bank of an ephemeral stream that was a tributary into Modoc Creek in the western Ozark (Springfield) Plateau. All trees in photographs were pignut or bitternut hickory showing reproduction of this species as represented by various age classes. Downed branches resulted from an ice storm in late autumn of the previous year. Most plants in understorey were of buckbrush or coralberry and Canada wildrye and/or silky wildrye.

The interlacing (anastomising) network of bark yet with a rather smooth appearance, especially, on upper trunk and limbs was characteristic of Carya cordiformis. Positive identificaation of these trees (trunks) was easy and foolproof given plentiful specimens of lower, terminal buds which in C. cordiformis are a giveaway sulphur-color. This is the only species of hickory to have this coloration.

Some specimens of mockernut hickory (Carya tomentosa) also grew on the banks of this ephemeral drainage. These trees were just beyond depth-of-field in these photographs, but they were presented later (below) in this section. These two Carya species are readily distinguished by features of winter twigs, especially buds, as was described immediately above as well as below.

Ottawa County, Oklahoma. Late December; winter dormancy.

188. Two more trunks- Another example of trunks and bark pattern in pignut or bitternut hickory. These two trunks were in background of second slide shown immediately above. All were growing on the bank of an ephemeral stream serving as a tributary of Modoc Creek in the western Ozark Highlands. Bitternut hickory is regarded as a climax species in several communities of bottomland and/or north slope mesic forests in the Ozark Region, including, as described in Eyre (1980), the hackberry-elm-ash cover type (SAF 93) shown in this section as well as sugar maple-basswood cover type (SAF 26) and white oak-black oak-northern red oak cover type (SAF 52) all of which were found were found either adjacent to this forest or within two-thirds of a mile upstream from it.

Ottawa County, Oklahoma. Late December; winter dormancy stage of phenology.

189. Bitter buds- Three examples of bitternut or pignut hickory twigs (C. cordiformis), including sulphur-colored buds, in winter phase. This is the only species of Carya (at least in the Ozarks and, probably, North America) that has buds with the color of sulfur or bright buckskin (Kurz, 2003, p. 80). Buds of this species also have a "pinched" or "squeezed" appearance often with an upswept orientation. Buds of C. cordiformis are considerably smaller than those of C. tomentosa with which it sometimes grows in close proximity (as on this stream bank) and, while generally of the same relative size as those of C. texana, easily distinguished from this smaller-growing hickory by its one-of-a-kind sulfur-colored buds.

These examples were growing on the example trees in the stand on the Modoc Creek tributary presented above.

Ottawa County, Oklahoma. Late December; winter dormancy.

190. Bunch of bitter nuts- Nuts of bitternut or pignut hickory gathered beneath the trees (and their buds) introduced above growing on the bank of an ephemeral stream draining into Modoc Creek.

Ottawa County, Oklahoma. Late December; ripe fruit stage, but they live up to their name (and the fact that humans would just as soon their hogs fattened on them).

191. Bitter fruits above Modoc Creek- Leaves and hickory nuts of pignut or bitternut hickory on a young tree on a steep slope directly above the channel of a typical stream in the Ozark Plateau. Ottawa County, Oklahoma, fruit-filling stage of phenology.

192. Instructive look- Leaves, characteristic young fruits, and the positive-identifying, sulphur-colored buds of pignut or bitternut hickory. Sutures of the husks or hulls of the immature nuts of Carya cordiformis frequently portrude outward like miniture "wings". This tree was producing fruit along with next year's leaves, catkins, and twigs as primordial tissues as its vote that life should go on. Bear on ole tree. Same organs as introduced in the two immediately preceding slides.

Ottawa County, Oklahoma, fruit-filling stage of phenology.

193. Trunk of mockernut or white hickory (Carya tomentosa)- Specimen of mockernut growing on bank of an ephemeral stream that drained into Modoc Creek in western Ozzrk Plateau. This hickory species was much less abundant than pignut or bitternut hickory which was a minor dominant on a western hackberry-elm-white ash tract of forest that was part of a larger bottomland forest that included sycamore, eastern cottonwood, boxelder, chinquapin oak, and black walnut along with the more consistently present climax dominants.

Bark of mockernut hickory is noticably smaller and more scalelike in general appearance than that of pignut or bitternut hickory with which it was closely associated in this bottomland forest. The only foolproof distinguishing difference, at least in late autumn through early spring prior to leafing-out are twigs. Both buds and lenticels differ drastically between mockernut and pignut hickories. Proceed to next two sets of slides and then compare back to pingnut twigs shown above.

Ottawa County, Oklahoma. Late December; winter dormancy.

194. Mockernut twigs- Winter twigs of mockernut or white hickory showing the distinctive lenticels and "tear drop" winter buds, especially the larger terminal buds. Besides being much larger than buds of associated pignut or bitternut hickory and black, Ozark, or Texas hickory (Carya texana) mockernut buds have a unique pattern of longitudinal bud scales as shown in the next three-slide set.

Ottawa County, Oklahoma. Late December; winter dormancy.

195. Detail of mockernut twigs- Greater detail of winter buds of mockernutor white hickory. The distinctive elongated lenticels and the longitudinally arranged, prominent bud scales handily distinguish Carya tomentosa from the smaller and less distinctive buds of C. texana two species of which often grow side-by-side on upland forest and savannah range sites with relatively shallow soils in the Ozark Plateau. Mockernut hickory also grows beside pignut or bitternut hickory on bottomland forest sites that have deep, rich alluvial soils (as shown in this section devoted to floodplain and tributary bottomland forests that was also in the Ozark Plateau). In such bottomland forest ranges pignut hickory is quickly distinguished from mockernut (or any other hickory) by its sulfur-colored, "squeezed-in-the-middle" buds (see again above).

Ottawa County, Oklahoma. Late December; winter dormancy.

Note on location: twigs of Ozark or black hickory (Carya texana) were included in the chapter, Tallgrass Savanna (under Grasslands) where this was a major (often, dominant) tree species along with mockernut hickory (C. tomentosa). Twigs of C. texana were not included in this section on bottomland because it is not a species of such environments.

196. Mockernuts- A "passel" of hickory nuts, all of mockernut hickory, that had naturally washed into this group following recent heavy rains. This view presented an example of the sometimes "super yields" of hickory fruit possible in "good years". There are, however, many more years when little or even no hickory nuts aae produced. The author did not find any fruit produced by the trees that yielded this bounty for five (5) years prior to this picturesque crop. Buds on these mockernut hickory trees were--as is typical of Carya species in general--an every year source of feed for wildlife ranging from white-tail deer to squirrels. So eat up and savor the moment ye 'beasts of the forest" (including hillbillies because mockernuts are relatively sweet in bitter contrast to those of pignut hickory).

Ottawa County, Oklahoma. Late December; ripe fruit stage.

197. Mockernut hickory in summer- leaves and young hickory nuts of Carya tomentosa growing in an oak-hickory forest on western ozark Plateau. Ottawa County, Oklahoma. Early July, fruit-filling stage.

198. Bidding winter and successional farewell- Final view of species composition during early winter of a bottomland forest in the western Ozark Plateau in which western hackberry, the "number one" climax dominant tree species (two leftmost trunks: big trunk in foreground and smaller trunk to left and behind it) was successionally replacing black walnut (four diagonically aligned trunks from right foreground extending to left midground). Most of the smaller (indistinguishable) trees in background were American and slippery elm (return to previous slides in this series for details concerning these two Ulmus species).

The most consistent understorey dominant was Canada wildrye. It was the only herbaceous species in the understorey of any consequence, Japanese brome or chess being present only as rare, isolated individuals. Buckbrush and fox or frost grape were the major shrubs, and of two sporadic layers. Canada wildrye, by constrast, was omnipresent comprising a nearly universal layer in this bottomland forest that developed along an ephemeral stream draining into Modoc Creek, an Ozark stream.

199. Carrying the banner- Canada or nodding wildrye was--as shown and described throughout this section--the dominant and basically the sole species of the hrbaceous layer (the predominant stratum of the understorey)--of a second-growth bottomland forest currently dominated by black walnut, honey locust, and white ash but progressing along its sere to a western hackberry-elm (American and slippery)-white ash climax.

Canada wildrye was present as both dormant shoots from last year and new (and very young) shoots of the current cool-growing season. Livestock were never present on this forest range, but white-tail deer as well as cottontail rabbits were abundant throughout and at all seasons. Squirrels, coons, and 'possums also called this tributary forest home, but of course fed only on fruits and buds.

One of the interesting differences between this bottomland forest range and the floodplain forest range (treated above), with which it was contiguous and immediately unstream from, was presence as local dominants and, sometimes, associates of silky and Virginia wildrye (Elymus villosa, E. virginica) downstream on deeper alluvial soil versus nodding wildrye (E. canadensis) on higher, shallower, less mesic soil. Also, the herbaceous layer (s) of the floodplain forest was (were) much more diverse in species and structure (see again above).

Canada wildrye was presented at level (the photographic scale) of local population in these two slides and at levels of individual plant and individual tiller immediately below.

Ottawa County, Oklahoma. Late December; all (at least, most) basal leaves stage (four to seven-inch length).

200. A real cool grass- The cool-season, perennial, native grass, Canada or nodding wildrye, was the dominant--indeed, sole species of any consequence--in the herbaceous layer of a bottomland forest on a tributary of Modoc Creek in the Ozark Plateau. Canada wildrye was present in almost every forest community photograph and description presented in this section (above). This species was presented "up close" at scale or level of individual, cespitose plant in these two slides. Canada wildrye was shown at the organizational level (scale) of local population in the immediately preceding set of slides and at level of individual tiller in the succeeding two-slide set.

Both current, cool-season shoots and dead, last year's shoots were present on this forest range.

Ottawa County, Oklahoma. Late December; all (at least, most) basal leaves stage (four to seven- inch length).

201. New growth in winter- Tillers of Canada or nodding wildrye in early winter in the herbaceous layer of a bottomland forest (ephemeral drainage into Modoc Creek, western Ozark Plateau) beneath black walnut and honey locust (Intolerant species) with western hackberry, American elm, and slippery elm in process of achieving domiance. These two slides presented Canada wildrye at photographic scale (plant organizational level) of individual shoot. Scales (organizational levels) of local population and individual plant were presented immediately above as two two-slide sets.

Canada wildrye is a strictly cespitose (bunchgrass) species meaning that the only shoots produced by this grass are tillers. These vertical (intravaginated) shoots do, however, grow in a somewhat decumbant form just below the soil surface as they grow away from the base of last year's shoots. That morphology was presented in these two photographs.

Both current, cool-season shoots and dead, last year's shoots were present on this forest range.

Ottawa County, Oklahoma. Late December; all (at least, most) basal leaves stage (four to seven-inch length).

The following series of slides of the ephemeral tributary (wet weather branch) of Modoc Creek presented forest range vegetation in late summer (early to middle of June). In addition to leaves on trees to show another feature of pioneer, dominant, and associate woody species the understorey herbaceous plants were also shown (as best they could be given dense shade) thereby supplying details of major grasses and grasslike plants as well as both structure and species composition of the forest plant community.

The first four slide/caption sets showed the lower end of this tributary just as it entered the greater floodplain of Modoc Creek.Subsequent slides gave views of this forest range farther upstream to present the same vegetation shown above in hibernal aspect in its vernal aspect.

202. Up the crick- Climax mixed hardwood bottomland forest on an ephemeral tributary at its entry into the greater floodplain of Modoc Creek in the western Ozark Plateau. The first of these two slides showed relatively young (pole-size) trees of hackberry and American or white elm (foreground) which are the climax co-dominants of this bottomland range vegetation. The largest tree (as seen here) in slightly left-of-center midground was hackberry. Clump of trees in background were all pignut or bitternut hickory which essentially had reached peak adult size (at their prime maturity). Behind the pignut was a mockernut hickory. Pignut or bitternut hickory was the major associate though it held dominance in some some local microsites. The second slide showed the large hackberry and the clump of pibnut hickory at closer camera range. These pignut and mockernut hickories were shown above in winter aspect and were treated again (below) in vernal aspect.

The herbaceous understorey was dominated by silky wildrye, Canada wildrye, James caric sedge, and Virginia creeper which climbed to the mid-canopy of some trees so as to be in several layers of vegetation. Also in the lower layer of the understorey were numerous seedlings of hackberry, American and, though fewer, slippery or red elm, and pignut hickory.

Tributary of Modoc Creek, Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). Mesic Bottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

203. Farther down the tributary- Where an ephemeral tributary formed a local delta at its narrrow mouth upon entry unto the floodplain of Modoc Creek a climax forest of young (pole-size) hackberry and American or white elm were growing to adulthood with a luxurant herbaceous understorey of both silky and Canada wildrye, James caric sedge, and Virginia creeper. The woody shoots of Virginia creeper were climbing tree trunks and ascending into mid-canopy. There were scattered plants of buckbrush or coralberry (as eeen in lower right corner of first photograph). The foremost sapling (center foreground) in both of these photographs was American elm. The two seedlings in front of this elm sapling (in second slide only) were one Americn elm and one pignut hickory. There were also several even-smaller seedlings of pignut hickory in the second slide. Trees behind and to the left of this elm were three hackberry, one American elm (far right), and a large log-size chinquapin oak (largest trunk in group at right). .The largest trees in left distant background of first slide and left midground of second slide were black walnut (tree at right with remnants of dead lower limb; larger of the two) and chinquapin oak (immediate left of black walnut). These latter two trees were either pioneers or early seral entrants into this forest.

Students should pay particular attention to the fact that this is a climax forest as to species composition (of both dominance by the climax dominant tree species an of climax herbaceous species), but structure of the climax forest had not been reached yet because hackberry and American elm were not at old-growth state of maturity. In fact, the climax dominants were still saplings to young poles.

So class, how could this be? Simple: it was a second-growth forest that had developed (undergone secondary plant succession) to the climax species composition yet with trees still in sub-adult age/size classes. Herbaceous perennials have only annual aboveground structures (shoots) so that climax species composition and climax structure of the herbaceous layer(s) were the same. Meanwhile long-lived seral tree species like black walnut and black walnut persisted in the climax stage, at least the onset of climax when trees of climax species were still young. Now, see how easy that was. You just have to learn to read vegetation.

204. Oldsters and youngsters- Old-age trees of seral species and young trees of the climax species with an understorey of climax herbaceous and shrub species on a bank of an ephemeral tributary of an Ozark stream. This was a closer-in view of the old black walnut with its rotten lower limb and a chinquapin oak to the immediate left of the walnut that were introduced in the immediately preceding photographs. The large sapling or small pole in right foreground was a hackberry. Pole-size trees behing this hackberry (midground) were both hackberry and American elm. Understorey was mostly an herbaceous layer consisting of Canada and silky wildrye (both species) and James caric sedge. Virginia creeper joined these graminoids, but it also ascended into the cnopy layer. Two other shrubs were buckbrush or coralberry and Missouri gooseberry (Ribes missouriensis).

205. A confusing state of successional affairs- Local assemblage of plant species of different succcessional states in a climax mixed hardwood bottomland forest on a bank of an ephemeral tributary into Modoc Creek at western edge of Ozark Plateau. The three largest trees were hackberry, the number one climax dominant. The tree immediately behind them (and barely visible) was American elm. The tree behind and to the right of this elm (far background) was a black walnut. The two straight and barely tapering trunks at far left midground were white ash. The sapling (foremost trunk, in front and to left of the big hackberries was a chinquapin oak. Leaves at far right and upper right corner were those of a pole-size northern red oak. The herbaceous layer was primarily Canada wildrye and some silky wildrye with lesser amounts of James caric sedge. Virginia creeper and buckbrush or coral berry were the two most common shrubs. Missouri gooseberry ranked a distant third. Forbs were not represented.

This array of range plant species encompassed a considerable successional span as well as presenting broad botanical diversity from species and family perspectives. Black walnut is this general area (over several forest sites) is pioneer to early seral speceis whereas hackberry and elm are climax dominants. White ash and chinquapin oak more-or-less fall in between these community extremes on the sere. Burns and Honkala (1990) interpreted white ash as a species of "intermediate as well as early stages of natural plant succession". White ash pioneers disturbed, yet fertile, sites (such as cutover bottomland forest like the one described here) and it can survive through on-going forest succession because white ash seedlings and young trees are shade-tolerant. White ash becomes less tolerant with advancing age (hence, increasing size) so that it "is classed overall as intolerant" (Burns and Honkala, 1990). With these silvics, white ash persist into later seral or even climax states.Under some forest conditions white ash is more shade-tolerant than northern red oak (Burns and Honkala, 1990). Consistent with these features was the fact that there were no seedling and very few young white ash in any part of this floodplain forest. Taken in total, the presence of white ash in the vegetation of this bottomland forest in early age climax (species composition but not vegetation structure of climax) made "successional sense".

Chinquapin oak is another somewhat "ambiguous" tree species with regard to successional status, but it bears a remarkable similarity to white ash in its silvics. Chinkapin oak is shade-intolerant yet, like white ash, it tolerates shady environments better at younger ages (though it may grow slowly under such conditions) until it eventually developes less tolerance of shade and general competition with advancing age (Burns and Honkala, 1990). This explained presence of older chinquapin oaks but not younger trees of sapling classes in this forest.

206. Afterall, this is part of the Oak-Hickory Forest Region- Young adult to mature adult trees of American elm, pignut or bitternut hickory, mockernut hickory, and white ash on the bank of an ephemeral stream that drained into a Modoc Creek, a typical perennial Ozark stream. Trees in this "lineup" in both photographs (from left to right; somewhat front to rear) were: American elm (first trunk), pignut hickory (three trunks--only two of which are visible--to immediate right of the elm), mockernut hickory (the siccessive four trunks after pignut hickories), and white ash (two right- and rearmost trunks). In the first of these two slides the three trunks (two smaller than the other) at far left were American elm while the large seedling or small sapling to right of the two ash (far-right midground) was pignut hickory. In the second slide the nice pole-size tree in right midground was also pignut hickory.

Braun (1950, p. 34) was careful to explain that although her forest regions where named for the normal or most widespread climax association (an application of the Clementsian concept of association) in each region, there were several climaxes that developed in regions and that taken altogether these (and even seral communities) characterized a specific regions. In addition to the climax association there were edaphic, topographic, etc. climaxes. (Braun [1950, ps. 33-38] actually merged or united the Clementsian monoclimax and Tanslian polyclimax in her classifiction of forests of eastern North America.) By way of example, the Oak-Hickory Forest Region was the center of development of the Oak-Hickory Association so this region was named after or designated by that association. Braun (1950, p. 35). Within the Prairie Peninsula Section of the Oak-Hickory Region Braun (1950, p. 189) recognized and described bottomland forests. Some of the tree species that she listed were walnut, ash elm, hackberry, and honey locust with sycamore, American elm, and cottonwood in lower habitats and the riparian zone. In a paragraph of that page Braun (1950. p.189) described the mixed hardwood bottomland forest that was treated here.

Dominance of the canopy layer by two species of hickory certaily indicated that this overall climax hackberry-American elm forest was indeed within the Oak-Hickory Forest Region.

207. Again, this is part of the Oak-Hickory Forest Region- At a different point of the bank of a tributary (upstream and a less mesic environment) into Modoc Creek another local forest community included old trees of pioneer and seral species being joined with younger (and smaller) trees of climax species. These two photographs showed the same forest range vegetation and the same plants with a closer-in camera distance in the second slide.The foremost tree (left margin) was an aging chinquapin oak with lower dead and rotting limbs. To the right of this chinkapin oak, generally an Intolerant (hence pioneer and early seral) species, was a sapling of American elm, climax co-dominant of this forest community and for this forest site. In center to center-left background were two over-mature honey locusts with much dead material in their crowns (ie. the old-timers were more dead than alive though still very much a part of this forest). To the left of the left honey locust was an American elm sapling. To the right of these honey locust was another large tree which was a box elder. The two large trees with dark-colored trees to right of the box elder (at extreme right margin in the second slide) were black walnut. This is another pioneer tree species that lives long enough to persist into the climax forest in the same manner that sycamore and eastern cottonwood did lower on the floodplain of Modoc Creek (see above discussions). Saplings in the center of these two "photoplots" were a mixture of hackberry and elm. Some of the latter keyed to slippery or red elm so the author interpreted this as a hackberry-elm--and probably more precisely as a hackberry-elm-hickory climax.

The understorey of this bottomland forest was primarily an herbaceous layer comprised of Canada wildrye, silky wildrye, and James caric sedge. Shrubs included Virginia creeper, poison ivy (= poison oak). buckbrush or coralberry, Missouri gooseberry, and, in the views shown here, fox or frost grape that had "clumb to the top of the honey locust". These two honey locust served as the focal point for the very next slide-caption as well as some of those presented above.

The unknown component in the ecological equation of this forest: One of the most profound, detrimental, (and downright sad) impacts of white man on the eastern deciduous forest, including (again, sadly) the Ozark Plateau was inadvertant introduction of Dutch Elm Disease. As a lad this author watched one giant and stately American and slippery elm after the other succumb to this beetle-borne fungal disease 40 to 50 years prior to the period over which these photographs were taken. Elm phloem necrosis in another disease common throughout the Ozark Region that kills elms, many of them huge beloved shade trees as well as those of field and forest. Phloem necrosis appears to be less prevalent (and only slightly slower to kill elms) in the Ozarks. For whatever reasons, American or white and slippery or red elms in Ozark forests are much less apt to be stricken by either of these dreaded tree diseases. This is a personal observation on the author's part and without any quantiative data to back up his assertion. It is still a fact.

The reason for this phenomenon was unknown to this observer, but it was the case that American and slippery elms that stood by themselves in yards, pastures, roadsides, field edges (ie. isolated trees, trees not in forests) were taken out at a much faster rate and at a higher percentage of total trees killed than those that were intermixed with other tree species. For example, isolated elms in yards, fencerows, and pastures within a quarter- to half-mile radius of elms in the bottomland forest described here were killed a half century ago while slightly smaller though--also old--elms in the floodplain of Modoc Creek were still alive.

Perhaps trees in forest were less likely to be found and used for feeding by American elm-bark beetles (Hylurgopinus rufipes), the principal vector of Dutch elm disease. Or, maybe, beetles feed only on larger trees so that when elm seedlings, saplings, and poles in the Modoc Creek forest reach adult size they will also be killed.

Whatever the explanation, the final outcome of these two dreaded elm diseases on forests in which American and slippery elms are major species remains unknown. Also unknown is what impact substantial loss of elm species would have on the ultimate climax (or, disclimax) forest. Would it become a hackberry-pignut hickory or a hackberry-pignut hickory-box elder forest? Perhaps it will remain a hackberry-elm climax, but of elms that never reach adult dimensions due to beetle feeding and transmission of Dutch Elm Disease. Even in some of the areas within the Ozark Plateau that were hardest-hit by elm diseases there are enough sexually mature American and slippery elms to replinish these species. There has not been enough time passed for these young trees to reach their mature sizes and, in point of fact, many of these are killed by diseases, especially Dutch Elm Disease, by the time trees are roughly "half-grown". Again, however, most of these are isolated trees not those in dense forests. At this time such observations continue to raise the question as to whether elms can remain undetected long enough to reach monster size, or if this can occur only in forests.

208. Climax newcommers join a pioneer veteran (some from both recent and earlier times)- Farther upstream on the same bank of the ephemeral branch of Modoc Creek the overall habitat and local habitats (microhabitats) were noticeably drier than the environments farther downstream and at the mouth of the branch that was presented above. On the xeric (less mesic) environments tree reproduction was proportionately greater for pignut hickory than for hackberry and American elm. Regeneration of northern red oak also was apparently more upstream than on downstream banks of this ephemeral barnch. Herbaceous dominants were the same as downstream: Canada wildrye, silky wildrye, and James caric sedge. The most abundant shrub in this scene was buckbrush or coralberry. There was also appreciable cover of Virginia creeper, but on ths "photosample" of vegetation the latter was the associate species. There were even a few large plants (in background) of Missouri gooseberry.

Most conspicuous in this local assembly of mixed hardwood bottomland forest were two senescing honey locusts with fox grape climbing into their dying crowns. These two aging, "over-ripe" honey locusts and the fox grape were shown in photographs above during winter (hibernal aspect). One of those slides was taken from this same photo-point (at a slightly closer distance) and two other slides showed the honey locusts and adjacent vegetation from the opposite direction. Honey locust is a pioneer tree species on disturbed land of this and similar range sites such as old fields ("go-back ground"). Wenger (1984, ps. 2-3) gave a tolerance rating for honey locust of Intolerant indicting from landowners in this area know all too well: honey locust invades--often, aggressively--abandoned fields, cutover land, and overgrazed pastures. These two old-growth trees had obviously established themselves on this second-growth forest shortly after the former forest had been logged which had been at least three-quarters of a century earlier. Braun (1950, p. 189) listed honey locust as a tree species characteristic of floodplain and bottomland forest in the Oak-Hickory Region to which this forest vegetaion belonged.

The trunk of the live tree to immediate left of the honey locusts was hackberry while the trunk behind this hackberry was that of an American elm that had died and fallen over on the honey locusts. The two trees at far right (smaller pole-size at left; larger yound adult at right margin) were American elm. Sapling at left margin was American elm (northern red oak in front of it).

Seedlings in the foreground were pignut hickory center, immediate foreground), American elm (right foreground), and northern red oak (left foreground). in that relative order of apparent cover and density.

A study of the understorey-The following sequence of slides and their captions dealt with the grazable-browseable layers of a climax mixed hardwood bottomland forest in the western Ozark (Spriungfield) Plateau that developed on an ephemeral stream functioning as a tributary into the main channel of a perennial stream, Modoc Creek. The understorey was treated with emphasis because this publication was devoted to range types and, in particular, the feed resoures of such natural pastures. The lower layers of this portion of the bottomland forest was somewhat different from that of the forest range vegetation that developed on the floodplain of Modoc Creek which was at slightly lower elevations.

209. Turf by trunks- Floor of a climax mixed hardwood bottomland forest on an ephemeral branch of a perennial stream in western Ozark Plateau. Canada and/or silky wildrye with James' caric sedge at bases of a black walnut (left) and chinquapin oak (right) filled this "photoquadrant" to show the herbaceous dominants and two associate tree species of what was apparently the potential natural range vegetation for this mesic forest site. These two tree species were most likely pioneers (certainly early seral colonizers) that due to long life spans persisted into the early climax ( a forest with species composition, but not old-growth structure of the climax).

The herbaceous species as well as those of woody vines were adapted to the shady forest floor and sun flecks that were a key environmental factor. (Note sun flecks in this photograph.) Some Elymus plants keyed to E. canadensis while other individuals keyed to E. villosus, plus still yet other plants appeared to be E.canadensis X E. villosus hybrids. However it was, both of these Elymus species are also adapted to growing and reproducing in full sun as, for example, beside dominant tallgrass species such as big bluestem (Andropogon gerardii) on nearly adjacent tallgrass prairies .

By contrast James' caric sedge appeared to be restricted to the densely shaded florest floor of well-drained soils on this upstream tributary whereas Davis' caric sedge was found only on the bank of the lower floodplain and bank of the channel of the main stream. Neither of these Carex species was ever found within the general area of the western Ozarks except in densely shaded microhabitats. C. jamesii and C. davisii were never found growing together, but instead were segregated according to location, soil drainage, and even presence of certain tree species. For example sycamore and C. davisii often grew in association (close proximity) whereas C. jamesii was never seen to grow beneath (or even near) sycamore, a tree which grew only on the floodplain of this forest.

From the perspective of the most practical management this range plant community was an edaphic and/or topographic climax viewed in Tansley's polyclimax theory and, probably, Whittaker's climax pattern theory. From monoclimax interpretation this forest vegetation was postclimax within the Oak-Hickory Association of Weaver and Clements (1938, 514-516). Braun (1950, p. 189) regarded bottomland forests like this as being wthin the Prairie Peninsula Section of the Oak-Hickory Region.

210. Lower-layer dwellers all together- The two lower layers (herbaceous and woody) of a climax mixed hardwood bottomland forest immediately above the floodplain of an Ozark Plateau perennial stream. The herbaceous zone was comprised of graminoids (Canada wildrye, silky wildrye, James' caric sedge) and no forbs. The lowest woody layer consisted of shoot bases of Virginia creeper along with seedlings of pignut hickory (locally most abundant), American elm, and hackberry. The latter two of these tree species were dominants and pignut or bitternut hickory was the associate of the canopy layer of this climax forest. On this forest range site these three trees were generally of Tolerant (or higher) tolerance ranking. On this higher-elevation portion of the bottomland forest Virginia creeper was the overall most common shrub with highest apparent cover among the lianas which also included (at other locations in this forest) fox grape, poison ivy, and trumpet creeper.

In this "photoplot" (and the one shown in the next slide) Virginia creeper was the only shrub species associated with seedlings of climax trees and the climax herbaceous species. Like the other woody vines Virginia creeper extended through all layers of the forest vegetation. It was, however, the only shrub species at maturity to still have its lower leaves intermixed with those of herbaceous species. For example, adult plants of fox grape and poison ivy had their lowest leaves many feet (say, about 10 or 12 or more) above ground level, often beginning just below the lower limbs of mature trees.

211. Grasslike plant and sort-of-a shrub in the lower layer- Several cespitose plants of James' caric sedge and basal leaves of Virginia creeper made up all of the lower layer of vascular plants in a climax hackberry-American elm bottomland forest. (Pignut hickory was the local associate to even dominant tree species at this forest location.) This species combination--usually with Canada wildrye and/or silky wildrye as herbaceous co-dominants--was characteristic of the lowest vegetational layer on an ephemeral stream branch immediately above the floodplain of an Ozark Plateau perennial stream.

This "photoplot" preented a closer-in, more detailed view than the "photoplot" shown immediately above which, at greater camera distance, showded more area and more range plant species (including the two wildryes). The herbaceous layer of forest vegetation presented here featured presence of lower leaves of Virginia creeper as a companion species to herbaceous plants. Even mature plants of Virginia creeper had their basal leaves intermixed with grasses and caric sedge whereas lowermost leaves of other lianas like fox grape and poison oak where high up on trunks of trees typically beginning just below lower levels of tree crowns.

212. What the future seems to be- The future composition of a mixed hardwood bottomland forest just above the immediate floodplain of a perennial stream in the Springfield Plateau was suggested by seedlings of tree species represented in this lowest layer of vascular plants. Here along with James' caric sedge, Canada wildrye, and silky wildrye (climax herbaceous species) were seedlings of pignut or bitternut hickory, hackberry, chinquapin oak, and American elm (in that relative order based on apparent cover and density). Chinquapin oak has usually been interpreted as Intolerant and thereby a pioneer or early seral tree species (Burns and Honkala, 1990). It was discussed above that presence of sexual reproduction in chinquapin oak added an element of ambiguity to the pattern of plant succession and forest development on this forest site. The other three tree species were viewed by Burns and Honkala (1990)as as being about Intermediate in tolerance rating with a degree of uncertainity. More specific evaluations of hackberry (and sugarberry) and elm species on bottomland forests (Diamond et al. 1987; Diamond 1993; Hoagland (2000) ) strongly suggest that these are the climax dominants of floodplain forest types. Thus, lowest cover and number of American elm seedlings was also inconsistent with the basic pattern of succession throughout the overall entirity of this bottomland forest.

Viewers should bear in mind that to get a photograph with tree seedlings large enough to be recognized the area photographed had to be quite small in size and restricted (limited) in location. Length (longest dimension) of this "photoquadrant" was less than four feet: not a large enough size to fortoituously find the exact composite mix of range plants for the average of the forest. Such samples that varied slightly from the arithmetic average were still common and within the realm of being representative. In other words students, do not jump to the conclusion that the individualistic view of the plant community carefully and skillfully proposed by Henry Gleason (1917, 1926) was verified by this roughly four foot by two and a half foot plot. Neither is this to imply that the Gleasonian view of plant communities is without some merit or that chance occurrence (current presence) of parent plants is not determinative (or largely so) in initial stages or final outcome of forest composition. Quite the contrary: it was in this instance. Presence of specific, climax tree species at specific locations appeared to be the major variable that determined which species dominated (= made up most of) the tree seedling component layer. Go to the next two sets of slides and captions for more of this compelling finding.

213. Pignuts in all layers (or indicators of climax)- Portion of a climax mixed hardwood bottomland forest on which pignut hickory dominated all tree layers from seedling through sapling to pole, and, ultimally, adult trees. The two large trunks with a fallen branch between them were mature pignut hickory. (The two large trunks at far right margin in the second photograph were white ash.) Given that pignut hickory was the local dominant tree species with the largest adult trees on this microsite and the species that produces the largest fruit it would seem logical to expect most younger trees to also be pignut hickory. This was not necessarily the situation. Ecology is biological not logical. The light-weight, wind-borne samaras of American and slippery elm were readily dispersed to "'every nook and cranny" of this forest range. The heavier yet also small fruit of hackberry, a drupe, was also dispersed widely. Even the bulky, heavy nuts of pignut hickory were not wholly dependent on gravity for dispersion. This bottomland forest abounded in squirrels, white-tailed deer, and eastern wild turkey all of which eat and disperse hickory nuts. Finally, even though this ephemeral stream bank was slightly higher than the floodplain of the main stream channel, local seasonal flooding permitted water to be yet another agent of seed dispersal. In final analysis, then, immediate proximity of adult pignut hickories to seedlings, saplings, and poles of this same species was not necessarily the determinate (at least not the only) variable in recruitment of pigunut hickory into the climax forest vegetation.

Dominant herbaceous species were graminoids: silky wildrye, Canada wildrye, and James' caric sedge. There were no forbs to speak of. Shrubs included Virginia creeper, the local dominant shrub, poison oak or poison ivy, buckbrush or coralberry, Missouri gooseberry. This forest range community was on the greater bank of an ephemeral stream that served as a branch to a perennial stream such that this range vegetation differed slightly from that on the lower floodplain of this stream system.

214. All together, but not by themselves- Details of a climax mixed hardwood bottomland forest on which pignut or bitternut hickory was the dominant tree species by having greatest cover and density of trees from seedling- to sapling- and up through pole-size classes. In the overall forest covered in this section pignut hickory was the associate while hackberry and American elm were dominants. In "photoplots" seen immediately above American elm exhibited considerably lower sexual regeneration. In this current "photoplot" American elm "got back in the game" of future climax forest by having the second greatest rate of sexual reproduction.

Other important forest plant species included Canada wildrye, silky wildrye, and James' caric sedge as dominant herbaceous species with prominence of particular species varying by microsite. The dominant shrub was Virginia creeper, a sort-of (relatively small) liana or woody vine.

215. Dominant grasslike plant on less mesic habitats- On the prominent bank of a tributary into Modoc Creek the bottomland forest had a different species composition (though similar structure) than that of the lower bottomland forest that developed on the primary floodplain of this Ozark stream. On this higher forest site Carex jamesii replaced Davis' caric sedge (Carex davisii), the major Carex species on the floodplain forest (shown below).

Ottawa County, Oklahoma. Mid-June.

216. James caric sedge- Several plants of Carex jamesii at full shoot development stage (peak standing crop) on the upper bank of a tributary into Modoc Creek in the Ozark (Springfield) Plateau. This sedge species was co-dominant with silky and/or Canada wildrye (see discussion of Elymus sp(p). above). Ottawa County, Oklahoma. Mid-June.

217. A college try- At least an attempt was made to show details (including miniscule sexual shoot) of James caric sedge, the dominant grasslike plant on bank of a tributary into Modoc Creek. Ottawa County, Oklahoma. Mid-June.

218. Tiny but ready to go- Current season's sexual shoot of James caric sedge that just senesced at peak standing crop and on-coming shoot dormancy. This shoot was lying on a shed leaf of sycamore. (Nice effect I thought.) Ottawa County, Oklahoma. Mid-June.

219. Grasslike floor dweller- Davis' caric sedge (Carex davisii) was the most common grasslike plant in the herbaceous layer of a western hackberry-American elm-sycamore-boxelder bottomland forest that developed in the floodplain of bottom of Modoc Creek in the western Ozark Plateau. The major herbaceous species were silky and Virginia wildrye and there were numerous forbs (many covered above) immediately. This was one of the few grasslike plants on this forest range.It was obviously a cool-season species being at this degree of vegetative development in early winter.

Ottawa County, Oklahoma. Christmas Day.

220. Doing it mostly in the shade- Vegetative and sexually reproductive shoots of Davis' caric sedge on the floor of a mixed haedwood (western hackberry-American elm-sycamore-boxelder) bottomland forest that developed along the floodplain of Modoc Creek in the Ozark Plateau. This is a cool-season species that--as shown in the immediately preceding slide--makes much of its vegetative growth during autumn through early winter. This pattern of growth permits Davis' sedge to survive (even thrive) under conditions of dense shade. This grasslike plant must, nonetheless, make some growth and undergo sexual reproduction during spring under heavy shade of a nearly completely closed forest canopy.

Ottawa County, Oklahoma. Early June, soft-dough stage.

221. Fruits on floor of forest- Detail of clusters of fruit (spikelets) of Davis' caric sedge, including individual units of fruit. The perigynium (plural is perigynia) is the thin sac (actually a sheath) surrounding the gynoecium and, as fruit matures, the achene in Carex species (Smith, 1977, p. 303). Perigynia and achenes are inside of the spikeletes in the fruit clusters.

This plant was in almost constant shade once tree leaves fully developed. Light adequate enough for this pictures was extremely short-lived. These photographs were a gift from God and fruits of photographic patience, but a long wait in the woods is itself a form of Almighty grace.

Ottawa County, Oklahoma. Early June, soft-dough stage.

Two that were closer to the water: Within the mixed hardwood bottomland forest (hackberry-elm climax) that developed on the floodplain of Modoc Creek there two other Carex species: 1) narrowleaf or eastern narrowleaf (C. amphibola var. amphibola= C. grisea) and 2) Short's caric sedge (C. shortiana). These two species grew much closer to the stream bank such that their habitat was wetter (ie. hydric or, at least, very mesic) than that of Jame's and Davis' cric sedges discussed immediately above.

222. In a puddle- Narrowleaf or eastern narrowleaf caric sedge (C. amphibola var. amphibola= C. grisea) introduced with shoot portion of a whole plant of (first photograph) and sexual shoots (second photograph) growing on outer bank of Modoc Creek as a part of a bottomland mixed hardwood forest. Narrowleaf caric sedge was one of three Carex species found in the herbaceous layer of this Ozark Plateau forest.

Some authors interpreted C. amphibola and C.grisea as synonymous (Steyermark (1963, ps. 356-359; Correll and Correll, 1975, 554-555; McGregor et al., 1986, p. 1069; Diggs et al. 1999, p. 1122; Kaul et. al. 2006, p. 391). Steyermark (1963, ps. 356-359), the authority covering this local area, and Fernald (1950, ps. 358-359) regarded C. amphibola and C. oligorarpa as two distinct--though very similar--species.

Ottawa County, Oklahoma. Late May.

223. Business end of a sometime water-lover-Distal or apical portions of sexual shoots (first slide) and general view of the infloresecnce (consisting of separate male and female spikes) situated within two terminal leaves of a sexual shoot of narrowleaf caric sedge. Greater detail of spikes were presented in the next succeeding slide set.

The specific epithet, amphibola, apparently refers to "equivocal" or "ambiguous" which on the floodplain forest in which these examples grew was appropriate given that some plants were on microhabitats that were considerably wetter (eg. these shown here) than others.

Ottawa County, Oklahoma. Late May; early fruit stage (immediate-past anthesis) of phenology.

224. "Male and female created he them" (Genesis 5:2)- Staminate and pistillate spikes, the inflorescences, (above and below, respectively; first two slides) and female spike with ripening achenes, the fruit type, (third slide) of narrowleaf caric sedge. Spikelets of narrowleaf sedge are androgynous, the arrangement in which staminate (male0 florets are above the lower pistillate (female) florets; in other words female florets subtend the male floral units (Steyermark, 1963, ps. 311, 357; McGregor et al., 1986, p. 1069).These reproductive structures were on plants growing on the flooded outer bank of ModocCreek, a typical stream in the western Ozark Plateau.

Explanation of third photograph: this was a nearly perfect shot, except for two water-drop resembling ghosts in upper left corner, but Epson Perfection Scanner never could scan this shot correctly for whatever reason. Readers were warned throughout this publication to never purchase any Epson products. They are inferior and the company will provide next to no help in assisting buyers of theri products. Inferior product; inferior business people.

Ottawa County, Oklahoma. Late May; early fruit stage (immediate-past anthesis) of phenology.

225. Not short on size- Short's caric sedge (Carex shortiana) is a species characterized by relatively large cespitose plants with "stout" shoots (Correll and Correll, 1975, p. 541) and thick rhizomes (Fernald, 1950, ps. 349-350). Individuals of Short's sedge are impressive in regards size and shape of plants, leaves and spikes (as shown in these two slides and several immediately below).

The particular specimen seen here was growing on the outer bank of Modoc Creek which had covered it first terrace as a recent flood. The sycamore branch against which shoots of this plant had lodged was debris from that flooding. Further evidence of flood water was deposition of sediment on leaves of the plant.

Short's caric sedge is a widespread species ranging from Ontario south and west to Oklahoma (Steyermark, 1963, p. 366).

Ottawa County, Oklahoma. Late May.

226. "The long and the short of it" (William Shakespear)- Leaves and several sexual shoots (first photograph) and entire inflorescence with several spikes (second photograph) of Short's caric sedge on outer bank of Modoc Creek following recent flooding. Short's caric sedge produced plants of largest individual size with largest, widest leaves of any of the four Carex sedges in the understorey of this Ozark Plateau mixed-hardwood floodplain forest. It was also the least common of the four Carex species, being restricted to habitats close to the stream and subject to overflow.

Viewers could see in the last two of these three slides pistillate spikes filled with ripening fruit (achenes) atop staminate spikes that had senesced. The inflorescences with their large female spikes subtended by male spikes (now in the "spent-state" in these photographs) were typical of Short's cric sedge.

In this bottomland forest, plants of Short's caric sedge did not consistently reach the impressive size shown here. Neither were such high yields of spikelets (ie. such well-filled spikes) typical. (Photographers often have to take advantage of unusually fruitful crops and atypically large or vigerous plants in order to show students what plant species and range landscapes look like "in person", that is their "in-the-field" appearance as seen by those fortunate enough to be on the land and observe live plants in their natural environments).

Ottawa County, Oklahoma. Late May; mid-to ripening-fruit stage of phenology.

227. Not short on fruit- Sequence of photographs showing distribution (first slide), general morphology (second slide), and details of spikes (third slide) of Short's caric sedge growing on outer bank of a stream (Modoc Creek) in the western Ozark Plateau. These photographs were of the same plant that was introduced in the two immediately preceding slide-caption sets.

The spikes seen here bore extremely heavy loads of spikelets in an abnormally wet spring. Mud on various organs was sediment left by a recent (and relatively) small flash flood.

Ottawa County, Oklahoma. Late May; mid-to ripening-fruit stage of phenology.

Farther up the crick- the following set of slides furnished another example of a second-growth hackberry-dominated bottomland forest that was upstream from the forest tract presented and described provided above. This forest tract treated below included "sylvan scenes" a few weeks following damage from a low-intensity (EF-1) tornado that were just outside and adjacent to the tornado-affected area..

A simple forest community- Interior of a second-growth hackberry forest (American elm was associate tree speies) with a highly productive herbaceous understorey that was almost exclusively silky wildrye. Although hackberry trees in this second-growth forest were mostly of the small pole size/age class (decades away from mature stage) this young forest was of the physiogonomy, structure, and composition of the climax forest. In other words, this forest range community was a second-growth climax forest of young trees and a fully developed, mature understorey.

Throughout extent of this second-growth forest there were scattered trees of American elm, sycamore, white ash, black walnut, pignut or bitternut hickory, chinquapin oak, and basswood or American linden.

In the Daubenmire (1968) habitat system this was a hackberry-silky wildrye (Celtis occidentalis-Elymus villosus) habitat type. This forest had developed in the western Ozark Plateau or, as it is sometimes designated, Ozark Highlands.

Floodplain of Modoc Creek, Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). Mesic Bottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

Dominant woosy and herbaceous plant- Interior of a second-growth mixed hardwood bottomland forest with a pole-sized (left) and adult tree (right) of hackberry with an undetstorey of silky wildrye representing the two dominants of this Celtis occidentalis-Elymus villosus habitat type of the Daubenmire (1968) model. The partially exposed upper trunk to adjacent right of pole-size hackberry was a young or immature tree of black walnut tree.

This climax (perhaps late or advanced subclimax) forest was in the western Ozark (Springfield) Plateau.

Earlier arrival that persisted- Local stand of white ash in a hackberry-dominated mixed hardwood bottomland forest. White ash is a pioneer species and a dominant of earlier seral stages with individual trees becoming increasingly less tolerant as they age and reach maturitysuch that the species has an overall tolerance rating of Intolerant (Burns and ). Seedlings (which can survive in threee percent full-light environments) and young trees of white ash are shade-tolerant (Burns and , 1998).

Given these silvic characteristics it was obvious that the large white ash seen here had established and grown to more mature stages when this second-growth climax forest of primarily young trees had been more open. As forest succession progressed, Tolerant species like the climax dominant, hackberry along with pignut or bitternut hickory, American elm, and basswood or American linden (mostly associate tree species) established leaving white ash--like the Intolerant sycamore (see below)--to persist into the climax, but without regeneration (at least not very much) of the next generation.

In the two views of this local stand of white ash there were just a few ash seedlings (with their survival yet to be known) whereas the tree component of the lower woody layer consisted mostly of seedlings and small saplings of hackberry and pignut hickory with a few chinquapin oaks of these age/size classes. The shrub component of this lower woody layer consisted mostly of spicebush (Lindera benzoin).

The herbaceous layer of the understorey (which ws not well-developed in this local area) was silky wildrye with a few individuals of Virginia wildrye.

Now more "sylvan scenes" of the the year thereafter (second full growing season after an EF-1 twister), but in parts of this second-growth floodplain forest that were around the perimeter of the tornado-damaged part of the forest which in the Daubenmire (1968) scheme was a hackberry-silky wildrye (Celtis occidentalis-Elymus villosus) habitat type...

Inside views- Two "photo-transectss" of the interior of a second-growth, climax-composition mixed hardwood forest that developed on the bottomland of a small stream in the Ozark (Springfield) Plateau. In the Daubenmire (1968) scheme of vegetatiion classification this was a hackberry-silky wildrye habitat type. There were also plants of Canada wildrye and broadleaf woodoats the ovrall associate grass species, but dominance varied locally. In these slides the yellow-colored (straw) was mostly of Canada wildrye or, about as often here, silky wildrye while broadleaf oats was featured in the first of these two slides. Trees, saplings and, seedlings of hackberry, American elm, and pignut or bitternut hickory were present in forest range vegetation shown in both photographs. Trees in first slide also included white ash, black walnut, sycamore, northern red oak, honey locust (Gleditsia triacanthos), basswood or American linden (Tilia americana), and chinquapin oak. Trees in the second slide were mostly hackberry (bigger trunks), American elm (mostly saplings and seedlings), and pignut hickory, but chinquapin oak and black walnut werre represented..

Major shrubs in both photographic scenes were spicebush fox grape, and trumpet creeper. Poison ivy and Virginia creeper were present, but not as widespread as in most forests in this area. Main forb was frostweed or white crownbeard (Verbesina virginica) or pokeweed (pokeberry) depending on local spot..

Floodplain of Modoc Creek, Ottawa County, Oklahoma. Mid-June (late vernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). Mesic Bottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

Interior composition I- Vertical and horizontal "photplot" (photographic dendrogram) of the same spot inside a second-growth, climax-composition mixed hardwood forest that developed on thefloodplain of a small stream in the Ozark (Springfield) Plateau. Two larger trunks and attending, sapling at at right were hackberry, the dominant tree. Small pole at right margin was Ameican elm.Tree in right background was sycamore. Yellow grass straw was that of both Canada and ssilky wikdrye. Shrubs were not present in visible range of these two photographs.

. Interior composition II- Horizontal and vertical "photplots" (photographic dendrogram) of a second-growth, climax-composition mixed hardwood forest that developed on the bottomland of a small stream in the Ozark (Springfield) Plateau. Forest dominat was hackberry (all foremost trees). The still-green grass om fpregrpund of slides was broadleaf woodoats while the straw (dead yellow shoots) was that of both Canada and sikly wildrye. Tree species scattered variously throughout this forest tract included American elm and pignut hickory, the two associate tree species, chinquapin,oak, sycamore, northern red oak, white ash, basswood or American linden, black walnut, and honey locust. Shrubs included trumpet creeper, fox grape, spicebush, poisonivy, and Virginia creeper. Widespread forbs included white crownbeard or frsotweed and pokeberry.

Interior composition III- Two horizontal " photplots" (photographic dendrograms) of a second-growth, climax-composition mixed hardwood forest that developed on the bottomland of a small stream in the Ozark (Springfield) Plateau. In the first (upper) slide trees were pignut or bitternut hickory (left margin, foreground) and hackberry (right foreground)). Other tree species scattered throughout this forest tract included: chinquapin oak, white ash, northern red oak, basswood or American linden, black walnut, and honey locust. The most common shrubs included trumpet creeper, fox grape, spicebush, poison ivy, and Virginia creeper..Mmajor forbs were white crownbeard or frsotweed and pokeberry.

Major tree, shrub, and grass- Scarred (cause unknown; maybe fire) trunk of aging hackberry, dominant tree species; poison ivy or poison oak, widespread (but only locally dominant) woody vine; and Canada wildrye (vying with silky wildrye and broadleaf woodoas as dominant grass) in a local microsite in a second-growth, climax-composition mixed hardwood forest on the bottomland of a small stream in the Ozark (Springfield) Plateau. American or white elm and pignut or bitternut hickory were associate tree species to hackberry. Chinquapin oak, white ash, northern red oak, black walnut, basswood or American linden, and honey locust were tree species well-represented in this second-growth forest. Spicebush and fox grape weere major shrubs. .

Right after the tornado- Below is a section of this second-growth, climax-composition, mixed hardwood forest along a small stream in the Ozark Mountins (Springfield Platteau), a hackberry-silky wildrye (Celtis occidentalis-Elymus villosus) habitat type, showing damage from an EF-1 tornado that dropped down sporadically throughout the forest a few weeks before these slides were taken.

Took off their tops- Crowns of two young adult sycamore trees (left background) and crown of one American elm (right mid-ground) blown off (twisted and torn away) by an EF-1 tornado in a second-growth mixed hardwood bottomland forest in the western Ozark (Springfield) Plateau.

Most of the lower woody layer consisted of saplings of the climax dominant, hackberry and associate species, pignut or bitternt hickory and American elm. Spicebush made up the shrub compnent of this lower woody layer. Young trees in this understorey certainly will benefit from the removal of adult tree crowns by the tornado as will the dominant herb, silky wildrye. Spicebush is so shade-tolerant that increased sunlight would likely have little or no impact on this shrub species.

This was a textbook example of a forest gap with the "stage set" for the phenomenon of vegetation dynamics known as patch dynamics or gap dynamics.

Blowed-over: pulled 'em up; took 'em down- Three views of a local stand of young adult sycamores trees within which three trees were uprooted from water-saturated soil by winds of an EF-1 tornado. These three trees all faced the same direction and, unlike neighboring trees that were twisted off at heights ranging from mid-trunk to beginning of crowns, these were laid over with root systems exposed. This pattern of blowdown indicated that it was possible (perhaps highly likely) that straight-line--rather than cyclonic--winds had been responsible for downing these three sycamores. Straight-lines could have preceded, followed or even accompanied cyclonic (counter-clockwise circular) winds.

This classic wind-created forest gap set the stage for "release" of understorey plant species and consequent changes in the local forest community, a process known as patch or gap dynamics.

This forest range was in the western Springfield Plateau.

Stright-line or cyclonic? Two adult hackberry trees in a second-growth, climax-composition mixed hardwood bottomland forest that were blown over and uprootd from saturated soil during a spring thunderstorm. Hackberry was the dole dominant of this climax (or, perhpas, late subclimax) stage forest populated by all age/size classes of hackberry and, also, pignut hickory and American elm, the principal associate tree species.

Silky wildrye was the dominant herbaceous species of this Ozark (Springfield) Plateau floodplain forest that would be a hackberry-silky wldrye haitat type by the Daubenmire (1968) system. Other tree species in this climax (or, perhaps, late subclimax) forest included white ash, black walnut, chinquapin oak, and basswood or American linden. The dominant shrub was spicebush and there were some plants of fox grape (Vitis vulpina).

For the "florensic ecologist" investigating this natural disturbance "crime scene" (or, maybe, the insurance adjustor surveying the "damage") the obvious question was whether these two trees were blow down by straight-line or circular (cyclonic) winds. Water-saturated soils from recent heavy rains explained (at least partly explained) the uprooted condition of these adult trees, but their having been blown over in the same direction and blow over intact rather than twister or broken off suggested high-speed straight winds. This conclusion was bolstered by neighboring trees that had been twister or torn off at lower levels of tree rowns or, even, midway up on tree trunks.

It was possible that straight-line winds could have accompanied the cyclonic winds of the small EF-1 tornado. Conversely--or also possible--straight winds preceded or followed in the wake of the little twister. Well, no the tornado was not little from the perspective of the mangled trees, but compared to the EF-5 tornado that had razed Joplin, Missouri only 15 miles way five years earlier it was a little twister. It does not take a major disturbance to set in motion what F.E. Clements called the dynamics of vegetation.

Straight or circular winds, these adult hackberries were laid over and down either way. Likewise, whatever category or rating of wind or extent of wind damage, hackberry was the predominant tree species that was present as young trees to replace the toppled adult hackberry trees (probably progeny of the uprooted adult trees). In essence, the forest gap and subsequent gap or patch dynamics was going to be that of age and size replacement of the climax tree species and not that of plant species replacement. Furthermore, the dominant climax herbaceous species, silky wildrye was still present and might even benefit from increased light.

Forest dynamics set into motion (pun intended) by damaging winds was not going to be plant succession, but rather population dynamics. This was a case of Population Ecology not Community Ecology. Interesting, hugh?

Sylvan orphan; took one generation, left another- As a follow-up or closer view of the last point about wind-created gaps or patchs and, subsequent patch dynamics being about changes in an existing population and an already present forest community, take a close look at this local gap. An adult hackberry tree was blown over and uprooted from water-saturated floodplain soil by either cyclonic or straight winds during a spring thunderstorm. Immediately in front (from this perspective) of the uprooted adult hackberry was a large sapling or small pole of--what else--hackberry. Furthermore, most of the smaller saplings and seedlings were also hackberry.

The thunderstorm that took out an adult hackberry and created this forest gap merely replaced one age/size class of a tree species with another (or others) age/size class of the same tree species. In this case the tree species was the dominant of a climax (or, at least, advanced subclimax) floodplain forest.

Plus, silky wildrye, the dominant herbaceous species of this hackberry-silky wildrye habitat type was also poised to either benefit from increased light or be unaffected (silky wildrye also thrives in the full sun of tallgrass prairie in this area of the Springfield Plateau).

"The more things changethe more they stay the same".

Twisted it off- Large adult hackberry with its trunk ripped off about twelve feet above ground level by circular winds of an EF-1 tornado that touched down in a second-growth, mixed hardwood botomland forest during a spring thunderstorm. Most of the trees that were left standing were hackberry, the sole dominant of this climax-composition forest. Hackberries ranged in size from seedlings, saplings, small poles to one adult that was almost as large as the snapped off tree (see very next slide below).

Other tree species that remained after the twister passed included pignut or bitternt hickory, white ash, American elm, chinquapin oak, sycamore, basswood, and black walnut, but there were more hackberry of all age/size classes than of all these other species combined. This was more so in the area seen here than throughout most of the forest due to greater dispersion of more progeny surrounding the two adult--as well as several younger, smaller--parent trees.

Major shrubs included spicebush and fox grape whereas the dominant herbaceous species was silky wildrye. These forest plants of the lower layers sustained no wind damage although one grape vine lost part of its 'arboreal trellis". The ccondition of greater sunlight in this newly creatd forest gap was likely to be of short duration as smaller trees around the snapped off hackberry were "released" and their crowns would "in short order" fill in this gap in the forest canopy.

Wind is a selective browser- Close-in view of a tornado-topped hackberry in a second-growth, climax-composition mixed hardwood forest that developed on the bottomland of a small stream in the Ozark (Springfield) Plateau. The shattered trunk on the right margin was a lower portion of the large torn-off trunk featured in the two immediately preceding slides as seen from a different camera angle. The tree in the left-center midground was another hackberry that was almost as large as the twisted-off hackberry.

The cyclonic winds of a small EF-1 tornado shattered the trunk of one hackberry and left completely untouched an almost-as-big hackberry less than 20 human steps away. All agents of defoliation (animals, disease organisms, fire, frost, ice, and wind) are (or, at least, can be) selective. Tornadoes are famous (infamous) for their wierd patterns of destruction. There have been numerous instances in which tornadoes have demolished a house yet left something untouched (eg. a cupboard with one teacup remaining while every other piece of china is gone). That was the situation here--though not quite so exacting in scale--where two trees of the same species and nearly the same size standing reasonably close together had one demolished while the other was unscathed.

The next year after the tornado- Immediately below are slides taken the second growing season after an EF-1 tornado "hopped skipped, and jumped" through a second-growth mixed hardwood forest that developed on the floodplain of a small stream in the Ozark (Springfield) Plateau.

Broke and spared- A scene inside a a second-growth mixed hardwood floodplain forest in the western Ozark (Springfield) Plateau in the second year (and growing season) after an EF-1 tornado moved sporadically through the woods breaking off limbs and trunks and uprooting large trees. The snapped trunk (snag) in right bckground was hackberry. Small pole at left margin was Anrerican elm. Most tree seedlings and your saplings were hackberry, the dominant tree. Two trees in left center were hackberry and American elm. here were also pignut hickories of all age classes in this second-growth forest that was of climax species composition.Grass inforeground was broadleaf woodoats (Uniola latifolia). Pokeberry (Phytolacca americana) accompanied it.

Gap edge- Two slightly different views of a forest gap created by blowdown from an EF-1 tornado a year (and growing season) previous to these photographs in a second-growth, climax composition mixed hardwood forest on a floodplain of a small stream n the Springfield Plateau. .The three trunks of large trees were middle-age, mid-size sycamore. Sapling at left in first slide was hackberry, the overall dominant tree of this bottomland forest. A small pole of American or white elm was aalong right margin of second slide. Major shrubs were trumpet creeper and fox grape. Grasses were Canada wildrye and broadleaf woodoats. Inaddition to saplings and seedlings of hackberry and American elm there also all age classes of pignut or bitternut hicklry and chinaquapin oak present.

This forest on the move- Below is a section showing secondary plant succession on an old field adjoing the second-growth, climax-composition mixed hardwood forest described immediately above. This little bottomland field, in the floodplain of Modoc Creek, was used for various field crops over a span of at least a half century before it was abandoned and became "go-back ground". This rangeman hauled "pea and millett" (actual species unknown) hay off this field as a kid. The sequence of secondary (old-field) succession for the climax vegetation of this bottomland site was altered by continued--though sporadic-- hay-making. Whatever plants grew on the old-field were put up as hay in little rectangular 65 pound bales. After about 40 years the old-field or "go-back" plant community had progressed to a herbaceous community dominated by eastern gamagrass (Tripsacum dactyloides) and rosinweed (Silphium integrifolium) and with various other primarily prairie plants including blackeyed susan (Rudbeckia hirta) and purpletop (Tridens flavus) along with nturalized tall fescue (Festuca arundinacea). Over time, seedlings of black walnut appeared and grew from one up to two or, maybe, three years of age depending on intervals between hay-making. Eventually a few seedlings of northern red oak began to appear followed by white ash, hackberry, and American elm.from the adjoining mixed hardwood forest. First, however, there were were seedlings of upland trees including redbud (Cercis canadensis), post oak (Quercus stellata), blackjack oak (Q. marilandica), and eastern persimmon ( Diospyros virginiana). Where fruit and seeds (or maybe vegetative parts like washed out roots) of these plant came from is known but to God (and He might be scratching His head).

Early stages of tree invasion were shown in these two slide-caption portions followed by more recent stages of tree invasion of the old field, abandone farm field. These two slide-caption sets were first included in the chapter herein entitled, Tallgrass Prairie (Interior) - ID. They precede the next slide-caption sets taken a few years later showing extent of tree invasion and conversion to an early tree-growth forest

The edge of the contiguous second-growth,.climax composition mixed hardwood was clearly visible in the background or along side margins in each of these slides.

Grassland or Forest?- That is the concluding question for this change in grassland community development at this point in vegetation dynamics. Students are seeing the essence in dramatic, graphic form of what Frederic Clements termed "dynamic plant ecology". Is this dynamics on-going plant succession? Or is it a textbook example that grasslands in humid and subhumid zones can only persist with some continual defoliating agent such as natural fire or, in this immediate instance, mowing or hay harvest?

Reign change- By the third year of not mowing or harvesting for hay it was becoming obvious that the grassland vegetation dominated by eastern gamagrass which had established on an old field was, in fact, clearly on the vegetational way to becoming a bottomland forest. It was not known whether this dynamics in range plant community was natural secondary succession or, alternatively, woody invasion of climax grassland in absence of tree control (whether defoliation was natural by fire or grazing/browsing or human action such as mowing or chemical treatment).

For whatever reason(s) eastrn gamagrass mysteriously died out simultaneously with encroachment of tree and shrub species. Some of these woody species such as black walnut and northern red oak, were present in a closely adjoining climax bottomland forest of hackberry (Celtis occidentalis), American elm, and white ash, but most woody plants were species of upland forest or oak-hicklory forest-tallgrass savanna and not present in the adjoining floodplain forest. These species included redbud, post oak, blackjack oak, and eastern persimmon Shrub species included smooth sumac (Rhus glabra) and blackberry (Rubus spp.)..

A phenomenon that heightened the intrigue of the mysterious disappearance of eastern gamagrass was persistence of the exotic (an introduced forage species) tall fescue which had died out in a field located just across a section line road during recent years of Extreme Drought. Native climax (decreaser) forbs, including rosin weed (Silphium integrifolium), were still thriving on this old field. There were also plants of the naturalized forage legume sericea lespedeza (Lespedeza cuneata).

On the floodplain of Modoc Creek, Ottawa County, Oklahoma. Mid-July.

Recovered grassland lost- An old field that had "gone back" to an eastern gamagrass-dominated bottomland grassland and served as a hay meadow for over two decades was being invaded by native species of trees and shrubs three years after hay harvest ceased (three years after cesssation of mowing and hay production).

Woody plant species visible in this heratbreak "photo-plot" of Paradise Lost included redbud, eastern persimmon, black walnut, post oak, blackjack oak, and northern red oak. Black walnut and eastern red oak were the only species growing in an adjoining climax bottomland forest (upper right corner of slide). All other woody species were found on neighboring uplands of oak-hickory forest, tallgrass prairie, of savannas (ecotones) of these two climax plant communities. Shrubs included smooth sumac and blackberry.

There was still some cover of tall fescue which had "volunteered" (invaded without human propagation) in this old field even before colonization by the climax dominant, esstern gamagrass. In a hay field just across a county road from this old field tall fescue had been planted some 40 years earlier and that stand of tall fescue had mostly died out after several years of Extreme Drought. Why, in absence of hay harvest (or, at least, mowing/shredding) the native eastern gamagrass died out (for undeternined reasons) and the introduced coo-season tall fescue persisted remained what Paul called a "great mystery" (Ephesians 5:32) .Unfortunately from a rangeman's perspective it was a sad mystery.

This old-field was contiguous to a a second-growth, climax-composition mixed hardwood forest of hackberry, American elm, pignut or bitternut hickory, sycamore, northern red oak, white ash, basswood, honey locust trees. The herbaceous understorey of this adjoining forest was silky wildrye, Canada wildrye, and broadleaf woodoats none of which were present in the (apparently) secondary succession community.

On the floodplain of Modoc Creek, Ottawa County, Oklahoma. Mid-July.

Three years later (six years post-mowing)- Six years after annual haying operations stopped hardwwood sprouts were bigger and thicker on an old-field immediately adjoining a second-growth, climax-composition mixed hardwood forest. This old crop field had been abandoned over 40 years before and through secondary plant succession with no human management input other than one annual hay- mowing had returned to a gassland dominated by easstern gamagrass and rosinweed. When annual hay-making operations ceased hardwood tree species (mainly redbud, persimmon, post oak and blackjack oak with some black walnut) with the shrubs, ssmooth sumac and blackberry invaded with established woody plants continuing to grow while more seedlings continued to increasse. At this successional stage seedlings of honey locust, American elm, and hackberry became obvious. Silky wildrye, Canada wildrye, and broadleaf woodoats which dominated the herbaceous understorey of the adjoining second-growth forest could not be found in the range plant community at this stage of apparent secondary plant succession.

It was emphasized that this appeared to be secondary plant succession, but what could not be explained as consistent with secondary succession was why eatern gamagrass and rosinweed, which are climax prairie plants, were in the process of being "crowded out", "shaded out" and being replaced by woody plants both climax forest tree species like hackberry American elm, and pignut hickory and typical colonizing tree and shrub species such as redbud, persimmon, sumac and blackberry.

"I know of no basis for a climatic grassland climax, but only of a fire grass “climax” for soils pern&ting deep rooting" (Sauer, 1950). Could fire, either natural-occcuring (lightening-ignited) fire or Indian-set, have been a factor in maintenance of prairies in this south and western part of the Prairie Peninsula (Transeau, 1935)? Had this old-field been a natural local prairie (which was why it had been tilled to begin with) amid what certainly appeared to be a a young second-growth climax floodplain forest? Both the second-growth forest and the recovered eastern gamagrass grassland appeared to be climax vegetation.

The cliamx decreaser species, eastern gamagrass, including grain-bearng inflorescences was visible in the foreground and midground of both of these photographs.The two naturalized introduced forage species, tall fescue and sericea lespedeza were also present (and abundant in case of tall fescur) and neither of these exotics had been planted in this old field. This further complicted interpretation of plant dynamics on this range.

On the floodplain of Modoc Creek, Ottawa County, Oklahoma. Mid-July.

After six years of "no mow"-On an old-field abandoned from farming 40 years earlier and retired from annual mowing for hay six yearsago seedlings and saplings of redbud, common persimmon, blackjack oak, post oak, black walnut, northern red oak continued to invade and grow while smooth sumac and blackberry patches increased in areaas. These first woody colonizers or pioneers were now joined with seedlings and small saplings of honey locust, American elm, and hackberry. These latter two tree species were associate and dominant trees, respectively, on an adjoining second-growth, climax-composition mixed hardwoody floodplain forest.

At this point (successional stage?) the only two tree species that had not invaded were sycamore and basswood. Basswood was a minor tree species, but sycamore was a local or spot dominant that was the first major tree species to colonize bare soil, such as gravel beds along streams, in this habitat. Eastern cottonwood (Populus deltoides) was also a first-tree pioneer on these sites, but cotttonwood was much less common (lower density and frequency) than sycamore. Neither sycamore or eastern cottonwood invaded this old-field-- at any stage.

Some of the eastern gamagrass, climax prairie grass (a decreaser), was visible in immediate foreground of both of these slides, but so also were plants of sericea lespedeza and, especially, tall fescue which were naturalized exotic forage species neither of which had been planted in this old field. Presence (and abundance in instance of tall fescue) of these naturalized forages, both of which are frequently weedy, made interpretation of vegetational dynamics.

On the floodplain of Modoc Creek, Ottawa County, Oklahoma. Mid-July.

Arrival of climax (for forest) trees- Now even climas forest trees had joined the woody invasion of this old field just six years after the last annual mowing for hay. Here, one of the few surviving plants of eastern gamagrass in full anthesis (anthers not visible at this size) was featured in the foreground in front of an approximate five or six year-old black walnut. The straw of sexual shoots of tall fescue were conspicuous. Tree species pressent at this stage of woody plant invasion included black waalnut, redbud, persimmon, blackjack oak, post oak, northern red oak, honey locust, pignut hickory, American elm, pignut hickory, and hackberry. Shrubs such as smooth sumac and blackberry are thicker than ever.

There were no trees of sycamore in this ol field. In this area of the Prairie Peninsula in the Springfield Plateau sycamore is a pioneer tree species that persist into the climax stage.Apparently seedlings of sycamore could not establish in the well-developed plant community at this (successional?) stage. Likewise, at this stage of vegetation dynamics there were no plants of silky wildrye, Canada or nodding wildrye, or broadleaf woodoats which were all dominant species in the herbaceous understorey of the adjoining second-growth, coimax-composition mixed hardwood forest.

Important specification: the foregoing discussion of bottomland forests represented Society of American Foresters (Eyre, 1980) cover types 93 (sugarberry-American elm-green ash) and/or 94 (sycamore-sweetgum-American elm) or some combination thereof. While these dominance types are unquestionably bottomland forests some--if not all--of these species can also be dominats on some highly favorable upland sites as well as on local or isolated habitats. For example, in the same area (within a half mile radius) of the Modoc Creek floodplain forest just discussed hackberry, American elm, slippery elm, white ash, and even sycamore grow to large size in fencerows, field corners, and other protected small areas. With spread of the dreaded Dutch Elm Disease large elm trees are not as abundant as prior to this epidemic. Nonetheless, trees of largest size in such protected and favorable haitats in the Ozark Plateau and eastern Osage Plains are typically those just named along with black cherry (Prunus serotina). Loss of natural and aboriginal fires has unquestionably been a major factor is this vegetational pattern.

Regardless, there are tracts--usually small and local in spatial scale--of forest or prairie woodland where hackberry, elm, and even sycamore on upland forest sites form the climatic (or, perhaps,edaphic) climax vegetation. At least such forest vegetation is climatic climax if one disregards-- momentarily, arbitrarily and for purposes of argument--natural fire which is, of course, a function of climatic (ie. long-term atmospheric phenomenon).

A short treatment of such forest range vegetation was presented immediately below. In this example honey locust, the pioneering or colonizing tree species persist throughout the sere until it is replaced by hackberry and American elm.as the climax dominants. Units of climax vegetation were not obvious so such units were not provided. It was deemed that the pattern of forest succession was the lessor for this short session.

228. Had their moment in the sun- Snags of honey locust that lived their lives long enough to see their kind being replaced by climax American elm (at least if and until Dutch Elm Disease claims them). This example of tree species succession was along a spring that fed into the tributary of Modoc Creek that was just described. The understorey included 1) an herbaceous layer dominated by silky wildrye with broomsedge bluestem, wild bergamot or purple beebalm (Monarda fistulosa), giant ragweed (Ambrosia trifolia), and common evening-primrose (Oenothera biennis) and 2) a shrub layer that consisted of buckbrush or coralberry (Symphoricarpos orbiculatus) and poison ivy/oak. In addition there was a lower tree layer (that was still at height of tall shrub) comprised of hackberry seedlings and small saplings which would--if not disturbed by man--grow into mature trees to co-dominant with American elm.

This local area had been dozed and graded with a county road backsloping and fenceline clearing operation about 60 years prior to time of photographs. It was on this "new land" (cleared ground along the fenceline) that the seedlings of the pioneer species, honey locust, had colonized. They had lived out their normal comparatively short life cycle. There were some neighboring, adult honey locust trees (just to the right of these two photographs) growing along this same old fenceline that were larger (though probably no older). They were still in their "natural prime" and preented in the immediately following slide.

Ottawa County, Oklahoma. Late December; winter dormancy.

229. Armed and unarmed- Lower trunks of two adult honey locust growing in a fencerow along a spring that drained into the Modoc Creek tributary treated above. The tree on the left was a thorned genotype (G. triacanthos var. triacanthos) whereas the tree on the right was a thornless honey locust (G. triacanthos var. inermis) The traits of thornless and thorned in honey locust were noted by numerous workers over the years. It appears that these genetic characters are not straightforward Mendelian traits. Santamour and McArdie (1983) remarked that the thornless feature is a sexually inherited trait and that recognizing thornless honey locust as a . distinctive taxonomic entity was of questionable legitimacy. Given drought tolerance, rapid growth (at least over course of a relatively short lifespan), and picturesque morphology of honey locust plant breeders have developed numerous cultivars of the native woody legume.

These trees and associated range vegetation (next paragraph) developed over course of roughly 60 years following backsloping by county owned and operated equipment. These two honey locusts pioneered the scraped land surface and over a half century expired before hackberry and American elm became widely established. Neighboring honey locust trees of this same approximate age (immediately to the left of this vertical camera range and shown in the two preceding slides) had already died. The author readily established age of these trees because as a school boy he walked along the county road bordering this fence row within three or four years of backsloping and new fence construction. (The photographer has continued to walk this road periodically since that time and has watched the redevelopment (recovery) of range vegetation over course of his 60 plus years lifetime.)

The liana (woody vine) growing between the two trees and upward on the throned trunk was possum grape (The lower woody shoots were buckbrush or coralberry. The several conspicuous small saplings or large seedlings were hackberry, the ultimate tree species that will become the climax co-dominant (with American elm)--unless of course man and his taxpayer-subsidized bulldozers and roadgraders backslope this fencerow again. This camera view did not include the herbaceous layer of the understorey. That layer of the range plant community was descriped in the immediately preceding caption.

Ottawa County, Oklahoma. Late December; winter dormancy.

230. An older patch and the same story- An isolated patch of woodland of approximately two acres in winter aspect that had developed in an old crop field ("go-back land"), the virgin vegettion of which had a big bluestem-Indiangrass tallgrass prairie here in the far western edge of the Ozark (Springfield) Plateau. The origin of this woody plant community was unknown to the author (or any living human), but the author rented and ran beef cattle on this land 45 years prior to time of this and the next five photographs (this and the next three slide-caption sets). Over the more than half century that the author was aware of this spot of Earth the vegetation consisted of a few honey locust and some black locust (Robinia psuedoacacia) trees within a population of broomsedge bluestem (Andropogon virginicus) and briars (Rubus sp.) the latter of which never produced a decent rop of blackberries. That plant community was at the edge of abandoned farmland (mostly dent corn and winter wheat). Broomsedge and briars comprised the ubiquitous vegetation on such "go-back ground" across much of this Ozark-Cherokee Prairie Region. (A decent place to hunt rabbits was about the most complimentary comment one could offer for such plant life.)

For whatever reason the strongly rhizomatous black locust gradually died. Their passing was followed about a quarter cnetury later by death of the larger honey locusts (the largest of which was featured shortly below). All the dead trees--starkly conspicuous by their lost bark--seen here were honey locust. The live trees that were their "replacements" were all hackberry (smaller leafeless trees of sapling to pole size) except for prominent (needle-bearing) seedlings of eastern red cedar (Juniperus virginiana) that were growing to right and/or behind the angiosperms.

There was very limited herbaceous growth it being overwhelmed by the lower woody (shrub) layer that consisted almost exclusively of buckbrush or coral except for a few plants of the dreadfully invasive, exotic multiflora rose (Rosa multiflora).

Ottawa County, Oklahoma. Late December; winter dormancy.

231. From honey locust to hackberry- On a local patch of woodland surrounded by recovering vegetation of an old field ("go-back ground") pioneering seedlings of honey locust that grew into adult trees were being replaced by hackberry, a climax dominant tree species on such forest sites. There were no other tree species on this sere the tree species sequence of which was honey locust then hackberry. Likewise, the understorey was alomst exclusively buckbrush or coralberry which established on the seral understorey of broomsedge bluestem and blackberry briars. For all practical purposes there were no plants of herbaceous species except for a few individual patches of Japanese chess or Japanese brome (Bromus japonicus) and these were so rare as to be of no consequence.

Hackberry plants were all larger saplings to small poles. These slides were taken in the winter aspect so that smaller saplings and seedlings of hackberry were not conspicuous. Nonetheless, most plants of Celtis occidentalis were saplings and poles. An example of pole-size hackberry was the foremost tree in the second of these two slides.

There were two dead honey locusts in far mid-ground of the first photograph, one consisting of a single trunk (inches DBH) and the other comprised of four trunks. The latter was the only honey locust in the second photograph. This four-trunk honey locust tree was featured in two slides below. The one-trunk honey locust was included in the next one-slide and caption set. Next slide, please.

Ottawa County, Oklahoma. Late December; winter dormancy.

232. No doubt as to the next regime- Hackberry or western hackberry ranging in size from small saplings to poles formed the exclusive tree species in an isolated woodland situated within an old field as seen in winter aspect. One large (and "very dead") honey locust tree (right midground) with a DBH 25 inches of served as the sole remaining evidence (in this photograph) of the previous dominant woody species in this local stand. The surrounding "go-back land" had been cropped for decades following breaking the virgin sod of a tallgrass prairie dominated by big bluestem and Indiangrass. Broomsedge bluestem and blackberry comprised the understorey beneath the living honey locust. Gradually under the shade of the honey locust and a few smaller and younger hackberry trees buckbrush or coralberry succeeded the broomsedge and blackberry until this low shrub formed the exclusive underlayer of this recovering plant community.

Distinctive knobby bark of hackberry was evident in the foremost trunk at far left. Hackberry appeared to be on its way to becoming the climax sole dominant of this isolated tract or woodland or forest.

Ottawa County, Oklahoma. Late December; winter dormancy.

233. Impressive even in death-Large dead honey locust consisting of four boles with DBHs of 17, 17, 16, and 15 inches. The photographer had rented this land and ran cattle on it for 15 years beginning about 45 years prior to time of photographs. At that time this tree was growing at the edge of an old field ("go-back ground") "carved out of" a former tallgrass prairie. Following cropping (dent corn and winter wheat) for decades the farmland was abandoned and supported a primary forage crop of Johnsongrass (Sorghum halapense). Just beyond the former farmland honey locust and some black locust formed a woodland with an understorey made up of broomsedge bluestem and blackberry.

It appeared as if the plant community of honey locust, black locust, broomsedge, and blackberry had been part of the cropland that had been abandoned before that larger part of the field which supported Johnsongrass (ie the woodland was the outer edge of the farmland) and had been undergoing secondary succession for some years prior to abandonment of the rest of the field.

Preceding photographs showed on-going replacement of large and dead honey locust by hackberry. Several of these large sapling to pole-sized hackberries were visible in both of these shots. The second slide provided a closer-in perspective of the four trunks of the dead honey locust. In this second slide the smaller trunk in the "gunsight" view created by these four boles (and to the rear of them) was a hackberry.

There was a low shrub layer made up exclusively of buckbrush or coralberry that formed the understorey of this range plant community. Herbaceous plants were for all practical purposes absent.

Visible (barey) in background of the second slide were several large sedlings of eastern red cedar. Cessation of natural prairie fires had permitted establishment of this sole coniferous species. For that matter, loss of the historic fire regime was quite likely responsible for establishment of this woodland or forest community to begin with. Tillage of land was one factor responsible for disappearance of fire as a natural climatic phenomenon and disturbance that retarded development of woody vegetation on this former tallgrass prairie in the extrme western edge of the Ozark (Springfield) Plateau.

Ottawa County, Oklahoma. Late December; winter dormancy.

234. American elm (Ulmus americana) in full-fruit- Twigs of American elm bearing bountiful fruit. The fruit type in elms is a samara which is a winged achene or, more specifically, a dry, indehiscent, single-seeded fruit fitted with a prominent wing (Smith, 1977, ps. 66, 90, 307). Most elm species have the season of their sexual reproduction "backassards" compared to most other angiosperms. Both American and slippery elm bear fruit before leafing out in the spring.

Ottawa County, Oklahoma. March.

235. Comparison of two major elm species- Distinction between American or, sometimes, white elm and slippery or red elm-- especially in young trees--is difficult (and not always correct even by "experts"). This can be a perplexing and frustrating problem in forests where both species are found (often trunk-by-trunk) such as on the bottomland forest discussed here and the sugar maple-bitternut hickory foresst covered previously. There is one brief period, however, when identification and separation of these two species is "a chinch". When these two species are in fruit (for ever so brief a time) they can be distinguished instantly by the distinctively different features of their samaras.

The first of these two photographs was a comparison of twigs heavily loaded with samars. In this first slide the twig of red or slippery elm was on top while that of American or white elm was on the bottom. These were typical spring twigs with the classic differences between these two species quite evident. The samaras of slippery elm are noticeably larger and the wings are "wavy" or "crinkled". By contrast samaras of American elm are smaller with more-less "flat" wings. The buds of red or slippery elm are covered with orangish-red (dull rust-colored) short hairs and at time of opening these reddish buds are much smaller (often less than half the size) of American elm which often have an elongated appearance (visible here).

The second photograph was a comparison of samaras of these two species. The smaller samara on the left was American elm and the larger samara with the "wavy" wing on the right was that of slippery or red elm.

Ottawa County, Oklahoma. March.

236. Slippery elm sample- Three-year-old seedling of slippery or red elm just entering its third growing season. Gives an indication of growth rate possible under favorable growing conditions (start of third wet spring-summer growing season). Characteristic leaves presented in second shot. Ottawa County, Oklahoma. July.

237. Leaves and fruit of the big oak in these bottoms- Chinquapin oak (Quercus muhlenbergia) was the only oak of consequence in the bottomland forest that developed on the floodplain of Modoc Creek. An example of leaves and ripe acorns was included to represent this tree species. Ottawa County, Oklahoma. October; fruit-shedding stage.

238. King shrub- Portions of leaders with immature fruit (drupe is fruit type) of spicebush (Lindera benzoin) which was clearly the dominant shrub of a climax mixed haradwood bottomland forest that developed along a typical Ozark (Springfield) Plateau stream. The author has observed that there is almost no feeding by animals on leaves or twigs of spicebush (even by often abundant white-tailed deer), but many animals (both mird and mammal) readily consume the ripe drupes.

Spicebush is in the sassafras or laurel family (Lauraceae). Good sources for discussion of spicebush include Vines (1963, 293-294; Stephens (1973, ps. 186-187), Kurtz (1997, ps. 142-143) and Nesom (2003). This shrub is dioecious with separate male and female plants being the typical case. Most reproduction is asexual or vegetative with thickets or colonies forming clonally (Nesom, 2003).

Spicebush got its common name from the fact that during the Revolutionary War, and subsequently (and probably even before), the drupe fruit was used for a form of allspice (Steyermark, 1963, p. 718; Kurz, 1997, p. 142). Even plant leaves have a fragrant odor, especially after a nice summer rain.

Floodplain of Modoc Creek, Ottawa County, Oklahoma. Late July; immature (though nearly full-sized) fruit phenological stage.

Bladdery pods on the bottom- American bladdernut (Stpahylea trifolia) was one of the more picturesque shrubs in the mixed hardwood bottomland forest that developed on the floodplain of Modoc Creek in the western Ozark (Springfield) Plateau. American bladdernut was covered immediately below.

239. Easy identification- Leaves and fruits of American bladdernut (Stpahylea trifolia) in a large colony on a mixed hardwood bottomland forest in the Springfield Plateau. Basis of the specific epithet, trfolia is the prominent and conspicuous trifoliate leaf. The fruit of American bladdernut is an inflated, papery, sutured, three-champered capsule with one to four seeds per locule (Vines, 1963, p. 667; McGregor, 1986, p. 567; Gleason and Cronquist, 1991, p. 350).

Staphylea species, of which there is only one in the immense Eastern Deciduous Forest Region, are in their own family, Staphyleaceae. This is an eye-catching forest shrub. On this bottomland forest range, from which cattle were excluded, white-tailed deer were the main range animal. The author never saw any browsing on leaders of American bladdernut. The major utilitrian function of this shrub to man was very effective watershed protection and soil conservation. The vast network of woody rootstocks (rhizomes) of American bladdernut colonies made for a "natural mesh" that provided an all but indestructable and nearly 100 percent effective barrier against rushing floodwaters.

Floodplain of Modoc Creek, Ottawa county, Oklahoma. Late May; immature fruit stage of phenology.

240. Bladders on the crick bottom- Capsules of American bladdernut in a large colony growing on the deep, rich, alluvial soil of a mixed hardwood bottomland forest in the western Ozark Plateau. Steyermark (1963, p. 1011) described the fruit as becoming "inflated and bladder-like at maturity " and "making a popping sound when crushed between the fingers". In this author's experience this latter effect is much more the case when the capsules are extremely dry and mature as in late autumn.

Floodplain of Modoc Creek, Ottawa county, Oklahoma. Late May; immature fruit stage of phenology.

241. Foxy foliage- New shoots (first slide) and characteristic leaves (second slide) of shoots of fox, frost, or chicken grape (Vitis vulpina) that regrew in the first spring-summer season following cutting by amateurish and hatchet-happy so-called "loggers" (described in agonishing detail above). Typical leaf patter, especially of leaf margins.

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

242. Woody climber in the bottoms- Fox, frost, or chicken grape in the western hackberry-elm (mostly American; some slippery)-white ash- boxelder (with persistent sycamore and eastern cottonwood) bottomland forest in the floodplain of Modoc Creek. This liana was brought to ground by the ice storm treated above which facilitated its visual reproduction here. This grape was growing in the forest gap described previously in various stages of growth and recovery Terminal portions of this woody shoot subsequently began the plant's ascent back into the forest canopy (but not before it left a contribution to educational of the next generation of rangemen, foresters, and wildlifers.

Frost or fox (the literal interpretation of vulpina) grape is one of the more common Vitis species in the Ozark Plateau as well as on this particular bottonlamd forest in the Springfield Plateau portion of the Ozark Region.

Floodplain of Modoc Creek, Ottawa County, Oklahoma. May; peak bloom phenological stage.

243. Foxy flowers- Leaves and flower cluster of the plant of fox grape in bottomland hardwood forest that was shown in the preceding photograph. Details of the inflorescence in second photograph. This woods-walking author has observed blooming fox grape over the decades, but only seldom found fruit produced. Sexual reproduction by any of the wild grape species is not particularily successful, at least not in the western Ozark Plateau. Return trips by the photographer to this location showed a complete "crop failure" of this particular flower cluster on this grape vine. All was not lost however: the sexual scene was reproduced and preserved for continuing education of viewers and would-be hillbillies.

Floodplain of Modoc Creek, Ottawa County, Oklahoma. May; peak bloom phenological stage.

Full foxy view- Lower portion of a smaller (younger?) plant of fox grape with young leaves and inflorescences in full-flower growing up in trees on the Springfield Plateau of the Ozark Uplift in northeastern Oklahoma.

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

More foxy views- Leaves and inflorescences of fox grape growing in the western Ozark Uplift (Springfield Plateau) in northeastern Oklahoma. Neighboring trees included young trees of hackberry and American elm plus senescing trees of honey locust (Gleditsia triacanthos) and adult blackjack oak (Quercus marilandica). The dominant understorey herbaceous species was silky wildrye (Elymus villosa).

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

New foxy leaves- New leaves of fox grape ranging in size and age with some "baby" leaves less than 1/5 adult size to mature (though still yound) adult dimensions. Fox grape was the most common Vitis species "hereabouts" (on a mixed hardwood bottomland forest).

Ottawa County, Oklahoma. Late May.

Foxy flowers- Clusters of flowers of fox grape growing up in hackberry, American elm, honey locust, and blackjack oak in the Springfield Plateau portion of the Ozark Highlands in northeastern Oklahoma. The first and second slides showed flowers while the third photograph presented tiny grapes in a very early stage of fruit development.

Ottawa County, Oklahoma. Late May, first and second slide; mid-June, third slide.

When it does bear fruit- Heavy crop of grapes in a large sprawling plant of fox grape growing in a sycamore tree in the Springfield Plateau portion of the Ozark Highlands in northeastern Oklahoma. Crops like this are quite infrequent. This was the most abundant fruit-set in fox grape ever seen by this photographer. Obviosuly, sporadic or infrequ;ent episodic fruit production would be adequate for sexual reproduction of fox grape. From seed viability studies of native plants it would be axiomatic that seed of native Vitis species would remain viable for years if not decades From the standpoint of providing food (feed) for animals, including humans, atypical, unusual high yields (or just any sort of yield) would be a challenge. Just another example of the hard-scramble life of Stone Age American Indians (and of European frontiersmen).In fact, infrequent crops in plants such as grapes would be a challenge for wildlife. Little wonder that corn (Zea mays) was central to human survival in much of North America, and that some species of wildlife thrived on corn grain.

Ottawa County, Oklahoma. Above Modoc Creek. Late July, ripening fruit stage.

Fruit of the vine- A single cluster of fox grapes. This was one cluster or bunch among the many such bunches presented in the immediately preceding slides. Fruit type of Vitis species is a berry. Note that within a cluster of individual grapes (berries) were in various stages of development and maturity.

Ottawa County, Oklahoma. Above Modoc Creek. Late July, ripening fruit stage.

Another vital species- Young shoot and leaves of summer grape (Vitis aestivalis) growing in the Prairie Peninsula of the western Ozark Uplift (Springfield Plateau). Summer and fox grape both grew in a mixed hardwood bottomland forest that developed on the floodplain of Modoc Creek. The overall dominant of this bottomland forest was hackberry (Celtis occidentalis). Leaves of hackberry seedlings were visible in the upper right corner of the first slide and along the left quarter of the second slide.

Taxonomic difficulty- Positive identification of the Vitis species is quite problematic given variation in leaf feaures, especially lobes nd margins, on even a short section of shoot. The five lobes on leaves of V. aetivalis is typically characteristic. Leaves presented in these photographs were pretty much "textbook". Best reference for the grape species of the Ozark Region found by this author was Kurtz (2004, ps. 364-375)

Ottawa County, Oklahoma. Late May.

Now for major forbs, but first:

Observational note: There are comparatively few publications devoted to forest forbs (per se) in the eastern deciduous forests. This contrasted to treatment devoted forbs in the Western Range Region. The serious student is left to rumage through "wild flower" field guides and related lay publications. Fortunately some of these are very good. Two such books that were especially relevant for forests of the Ozark Plateau Region were Denison (1989) and Kurz (1999).

244. Joe Pye weed on the bottomland- Portion of a local colongy of sweet, purple, purple-jointed, or green-stemed Joe Pye weed (Eupatorium purpureum), a composite of tribe Eupatorieae, growing on a bottomland Ozark Plateau forest. This forest on the floodplain of Modoc Creek had the species composition of a climax forest being dominated by hackberry or western hackberry and American elm (with some slippery elm) along with old-growth individuals of sycamore and eastern cottonwood that had pioneered this forest sere. This large composite was a local dominant during much of the warm-growing season.

Obviously, sweet or green-stemed Joe Pye weed is adapted--in fact, thrives in--shade. This species was a local dominant of the taller of two layers of herbaceous vegetation. The lower layer was mostly limited to early spring (before trees become fully leafed-out), but purple-jointed Joe Pye weed and three species of nettle (shown below) formed local, tall layers of herbaceous plants. All of these forb species formed local colonies from which other species were excluded. Apparently, microsite or "luck" (original or initial presence of plant species) was determinative as to which forbs grew where in this climax bottomland forest.

Eupatorium species are known variously as thoroughwort, boneset, Joe Pye weed, and, even for select species, wild ageratum.

Ottawa County, Oklahoma. July, most shoots were at full-bloom stage of phenology.

245. One Joe Pye weed- A single plant of sweet, purple-jointed, or green-stemed Joe Pye weed standing on outskirts of a colony of this species presented morphological and other key features of this composite species that is well-adapted to the "deep, dark woods" of bottomland forests.

Pole-sized tree in left background was hackberry, one of two climax dominant tree species. Liana was that of summer grape.

Ottawa County, Oklahoma. July, full bloom phenological stage.

246. Ready, set, flower- Two stages in sexual reproduction in green-stemed, purple, or purple-jointed Joe Pye weed. First photograph was of immediate pre-bloom with the infloresecence "fixing" to emerge. Second photograph was the full-bloom stage of the forest floor-dwelling composite. These plants were growing as part of a colony in the dark understorey of an Ozark bottomland forest of climax species composition in which western hackberry and American elm (some slippery elm) were the natural dominant tree species. There were several large (old-growth) trees of sycamore, eastern cottonwood, and chinquapin oak which had been pioneers (at least members of early seral stages if not colonizers) on this land when it was along the bank of an Ozark Plateau stream (Modoc Creek). Over time Modoc Creek changed its course so that the land occupied by most of this forest became farther removed from the previous channel and part of the creek's greater floodplain.

This is a beautiful forb that would appear to have great value as an ornamental, horticultural wild flower for those who enjoy landscaping with native plant species.It is obviously well-adapted to "deep shade".

Ottawa County, Oklahoma. July.

247. Ornate cluster- The showy inflorescence of sweet, purple-jointed, or green-stemed Joe Pye weed in understorey of a mesic bottomland forest with the species composition of a climax hackberry and American (some slippery) elm-dominated range plant community. There were several old-growth specimens of sycamore, eastern cottonwood, and chinquapin oak that still commanded much of the canopy of this forest. These species had colonized this sere (or arrived early in the seral sequence) and persisted into the climax forest through their long life and "good fortune".

Ottawa County, Oklahoma. July.

248. Blue mist along the creek- Blue boneset or mist-flower (Eupatorium coelestinum) growing on the bank of a typical stream in the Ozark Plateau. Blue boneset is particularily well-adapted to quickly colonizing and stablilzing riparian habitats and similar moist environments because it has an extensive system of rhizomes coupled with plentiful seed (achene) production. This specimen was growing on a creek bank comprised primarily of rocks (but wet rocks from stream outflow).

Lost Creek, Ottawa County, Oklahoma. September; peak bloom.

249. Set on blue bones- Upper shoots (first slide) and inflorescences (second slide) of blue boneset or mist-flower in the Ozark (Springfield) Plateau. This is another one of the Eupatorium species that prefers wet to fairly mesic microsites. It is, however, extremely drought-tolerant even on moist upland habitats. The specimen shown here was growing on a rocky upland microhabitat after autumn rains broke a record summer-long drought. Although this microenvironment was tremendously less moist than the specimen shown in the immediately preceding and succeeding slides the blue boneset plant (and it had numerous companions) featured here grew under moderate shade of a black walnut tree and thicket of black raspberry (Rubus occidentalis). It is the combination of all factors impacting an organism that comprises the ecological niche and local home (environment) of that living individual.

Purchase wrning: These were two more images that were improperly scanned by an Epson 700 Perfection scanner. These were very crisp and properly colored slides (Provial 100F film) before the Epson scanner miraculously marred them. These two scanner-ravashed images were included as a warning to would-be purchasers of scanners: never buy Epson products. Not only is the equipment inferior, but company are arrogant and will not assist customers. Sorry equipment to match quality of company management.

Ottawa County, Oklahoma. late October; full-bloom stage.

250. "Now blue ain't the word for the way that I feel"- Ray Price (compliments of song writers Ralph Mooney and Charles Seals) did not have this sort of blue in his "yearning mind" but "crazy arms" of the photographer did capture this example of blue boneset or mist-flower growing on a wet stony bank of a typical creek in the Ozark Highlands. There is a wide array of habitats on which the various Eupatorium species grow ranging from shallow soils on dry prairies to moist or, even, wet bottomlands. E. coelestinum is one of the more moisture-requiring of these species.

Species in tribe Eupatorieae have only tubular or disk flowers.

Lost Creek, Ottawa County, Oklahoma. September; peak bloom.

251. Virginia waterleaf (Hydrophyllum virginianum)- In early spring the herbaceous vegetation of the climax, hackberry-American elm-sycamore bottomland forest typically consists of a single layer the height of which seldom extends far above the forest floor. In the mesic bottomland forest that developed along Modoc Creek this early vernal herbaceous layer was dominated by two species of Hydrophyllum which is the namesake species of the Hydrophyllaceae or waterleaf family. The more common of these species was H. virginianum commonly known as Virginia waterleaf.

This author could not detect any sign of herbivory on either of these species other than nector and/or pollen feeding by bees. It followed that waterleaf species were locally valuable bee plants.

Ottawa County, Oklahoma. April.

252. Virginia waterleaf at closer and close range- Virginia waterleaf with the only animal species seen by this author to interact with waterleaf being a few insects, including the common bumplebee (Bombus americanus).

Ottawa County, Oklahoma. First photograph taken in April; seocnd photograph taken in May.

253. Bigleaf waterleaf (Hydrophyllum canadense)- This was the larger (and less common) of two Hydrophyllum species which together comprised the vast bulk and cover of the early vernal herbaceous layer in the climax, mesic, bottomland forest (Nelson, 2005, ps. 147-150) that was a hackberry-American elm-sycamore cover type, an Ozark Plateau variant of SAF 93 (Eyre, 1980, p. 65).

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

254. Its own family- Lopseed (Pharyma leptostachya) "down deep" on the floor of a mixed-hardwood bottomland forest. This species was associated with Virginia waterleaf, bigleaf waterleaf, purple-jointed Joe Pye weed, Davis' caric sedge (Carex davisii), Canada wildrye, and silky wildrye in the herbaceous layer of this botanically rich forest range.

Pharma is the sole genus of the lopseed family (Phrymaceae), and P. leptostachya is the single species of this genus in North America (Fernald, 1950, p. 1313). This truly is a unique plant that added all the more to the biological diversity of this forest type.

Floodplain of Modoc Creek, Ottawa County, Oklahoma. Late July (Exceptional Drought and severely drought-stressed plant).

Explanation for another run at "ye old college try": The Epson "Perfection" (not) 700 scanner made such a mess of the two preceding slides that four more were presented trying to "do justice" to the unique forest forb, lopseed.

255. A unique though distinguishing species- Large plant of American lopseed on floor of climax mixed hardwood (western hackberry-American elm-sycamore) bottomland forest in western Ozark (Springfield) Plateau. Diggs et al. (1999, ps. 880-881) described this species has having a "disjunct distribution pattern" in Asia from India to Japan and the deciduous forest of eastern North America.

Lopseed is a perennial forest forb that is well adapted to deeply shaded growing conditions and that prefers mesic habitats such as moist deep alluvial soils.This plant was suffering through the second consecutive year of Extreme to Extraordinary Drought (Palmer Index). This photograph was taken after the author waited over an hour for a brief period of sunlight forest floor.

Floodplain of Modoc Creek, Ottawa County, Oklahoma. Late June; still-in-bloom stage.

256. Emphasis on the inflorescence- Upper part of sexual shoot with its spike-like inflorescence (Diggs et al., 1999, ps. 880-881) at maximum development of American lopseed on floor of a climax western hackberry-American elm-sycamore bottomland forest. Viewers should note the deeply shaded environment in which this individual was growing. The author never did find a period of time when there was any more sunlight that could penetrate the forest canopy. This plant, like the ones shown above, was severely water-stressed being in the second consecutive year of Extreme to Extraordinary Drought (Palmer Scale).

Floodplain of Modoc Creek, Ottawa County, Oklahoma. Late June; still-in-bloom stage.

257. Another view (or attempted view)- Another plant of lopseed growing on the floor of a mixed hardwood bottomland forest along a typical heavily human-impacted stream in the western Ozark (Springfield) Plateau. It proved to be extremely difficult (and frustrting) to get even "a half-way decent" shot of lopseed given that it grows in deep shade; in this instance with less than three-quarters of a hour of direct light (a shaft of sunlight) that permitted acceptable photographs using natural light conditions.

Loppseed has to be even more tolerant of low-light conditions as the tolerant dominant tree species such as western hackberry and American elm. This forest forb was not abundant to provide any feed "to speak of" for larger range animals such as the resident herd of white-tailed deer living in this forest.

Floodplain, Modoc Creek, Ottawa County, Oklahoma. Late July; mid-fruit maturation stage.

258. A hard-won sequence- Series of views of lopseed going from late bloom (the last bloom of the cluster) to mid-maturity phenology growing on the floor of a climax hackberry-American elm-sycamore floodplain forest that developed along a degraded stream in the western ozark (Springfield) Plateau. Pharma letostachyia is a unique species being the solitary member of the lopseed family (Phrymaceae) found in the ancient, worn-to-their-roots Ozark Highlands, but it is a fairly widespread occurring in every county in Missouri (Steyermark, 1963, p. 1380).

Interesting enough, a search of the literature did not reveal any detailed tratments of P. letostachyia in its "own right" beyond a treatment of this speies morphology which remarked on its sexual and asexual reproduction (Holm, 1913).

Although this species has no feed value as a member of the forest range plant community and is not limited in distribution so as to be endemic it is an indicator plant of rich, moist forest habitats. In addition Phrymaceae is a monotypic family (Diggs et al., 1999, p. 880). Inclusion of lopseed certainly added a touch of variety to this treatment of a mixed hardwood bottomland forest range. (Or so it was hoped given the great difficulty and patience required to deliver these images from a dark summer forest to viewers.)

Floodplain, Modoc Creek, Ottawa County, Oklahoma. Late July; late-bloom stage (first slide) to mid-fruit maturation stage (second and third slides).

259. Monotypic flowers- Portion of inflorescence of American lopseed, the sole species in genus Phryma. This was the right branch on the spike-like inflorescence presented in the immediately preceding slide. The fruit of lopseed is an achene (or achene-like fruit) that is enclosed within its calyces (Diggs et al (1999, p. 881).

Floodplain of Modoc Creek, Ottawa County, Oklahoma. Late June; still-in-bloom stage.

260. A different sort among a common sort- Virginia knotweed or jumpseed (Polygonum virginiianum= Tovara virginiana) on the deeply shaded floor of a mixed hardwood bottomland forest in the western Ozark Plateau.As treated by Steyermark (1963, ps. 582-598) there are 24 Polgonum species in Missouri, the state that bordered on this county in Oklahoma (a state for which there is no published flora or manual). Styermark (1963, ps. 596-597) put Virginia knotweed in the Polygonum section of Tovara, which was formerly the genus Tovara that included T. virginiana. Small (1933, p. 454) used Tovara virginiana for this species. Gleason and Cronquist (1991, p. 138) showed both Tovara and the even earlier genus, Antenoron as previous genera for this species.

Virginia smartweed is uniquely different from other Polygonum species in the region of the eastern deciduous forest. Gleason and Cronquist (1991, p. 138) gave the biological range for this species as being from New Hampshire west to Nebraska and south to Florida and Texas (ie. almost all of the Eastern Deciduous Forest Formation). I

Virginia knotweed (or smartweed, the other generic common name for the Polygonum species) differes from others in this genus by having an elongated and narrow inflorescence with fruit having "hooked or recurved beaks" (Steyermark, 1963, p. 582). This unusual feature among Polygonum species aside, Virginia smartweed is a common, widely occurring species being found in every small county throughout Missouri (Steyermark, 1963, p.) 596).

Virginia knotweed grew in association with American lopseed on a mixed hardwood bottomland forest (which is why both were included 'beside each other" at this juncture in coverage of this forest range cover type).

Floodplain of Modoc Creek, Ottawa County, Oklahoma. First slide: mid-June at pre-bloom stage. Second slide: late June; late-flowering/early fruit stage of phenology.

261. Hooked flowers and fruit- Inflorescence with both flowers and fruit (lenticular or three-sided achene; Smith, 1977, p. 112) of Virginia smartweed on the shaded floor of a mixed hardwood bottomland climax (or late subclimax) forest. These two views were of the upper potion of the inflorescence on the plant shown in the second slide of the preceding slide-caption set (the immediately preceding photograph).

Small (1933, p. 454) explained that the common name of jump-seed was derived from "the surprising distance the ripe achenes fly when pressed under the thmb nail".

Floodplain of Modoc Creek, Ottawa County, Oklahoma. Late June; late-flowering/early fruit stage of phenology.

262. Two more views of a different sort- One plant of Virginia knotweed or jumpseed seen from the side (first slide) and top-down (second slide) in the floor of a maixed hardwood bottomland foresst in the western Ozark (Springfield) Plateau. Associated plants in these "photoplots" included the naturalized Asian mint hereabouts called "beefsteak plant" (Perilla frutescens) and Davis' caric sedge (Carex davisii). The latter of these was treated earlier (above) in this chapter.

Comparison of the plant in these two images with those plants presented immediately above revealed differences in phenological advance from one year to the next. Plants in above slides had reaached early (immature) fruit stage one month earlier (late-June) that this plant (late-July) that had not even reached pre-bloom stage. The plant in these two slides was photographed in nearly the exact spot of forest floor though two years later than those shown above. The plant shown in these two slides was growing in a spring/summr growing season that had been one of the coolest in weather records. Both years were drought years.

Note evidence of insect feeding on leaves of this particular plant. It was not the best of years for this Virginia knotweed.

Floodplain of Modoc Creek, Ottawa County, Oklahoma. Late July; pre-bloom stage of phenology.

263. Rose family forb- White avens (Geum canadense) at base of half-century old black walnut on an Ozark Plateau botomland forest. Detail of leaf and arrangement of flowers on ssexual shoot of white avens (second slide). This is an interesting forest forb in that it is commonplace in shaded understories that vary from virgin or old-growth forest to disturbed areas in yards and gardens. Kurz (199, p.48) commented that leaves of white avens are grazed by white-tailed deer while seeds provide a feed source for forest birds.

Rosaceae, the rose family, is generally more important and better known for being the single most important family of browse plants, but the family also boast of a few notewoerthy range forbs. White avens is one of these.

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

264. Tiny flower on large forb- The individual inflorescence of white avens as well as the entire group of these flowers is small compared to size of its stem and shaply leaves.

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

There were three species of the Urticaceae, nettle family, that at local scale were dominants of the summer herbaceous layer of the understorey of this hackberry-elm bottomland forest. Individuals of all three species have stinging hairs that canse a burning and/or itching sensation when they come into contact with human skin. As these species often form colonies of considerable size their presence in the woods presents problems to humans stupid enough to go into forest wearing short sleeves, short pants, and other inappropriate clothing. All three species are monoecious with separate male and female flower clusters near apex of their shoots. It is doubtful that any of these three species provided forage to grazing animals. These three nettle species were included because they were native species that were local dominants of this forest range type.

265. Colony of wood nettle (Laportea canadensis)- Portion of an extensive colony of wood nettle in the hackberry-American elm-sycamore bottomland forest described here. Shoots of these plants ranged in height from two to almost four feet in height. Wood nettle was a local dominant of the taller portion of the herbaceous understorey.

Ottawa County, Oklahoma. June.

266. Close, but don't get too close- Details of shoots, including inflorescence, of wood nettle. This is the most skin-irritating member of the nettle family to grow on the climax hackberry-American elm bottomland in the Springfield Plateau of the Ozark Highlands region. The smaller inflorescences at tip of shoots were staminate clusters while the lower and larger floral clusters were the pistillate inflorescence.

Pubescence of this species include stinging or poison hairs that can be painfully irritating to careless woods walkers. It is, however, an attractive and interesting forest forb as well as a locally dominant species of the understorey.

Ottawa County, Oklahoma. First of these two photographs taken in September; second photograph taken in August.

267. Prime females and withered males- Details of the past-its-prime, staminate flower cluster (top; tip of shoot) and the height-of- sexual-reproduction, pistillate inflorescence (rest of shoot) in wood nettle. This was one of many shoots in the colony of this species presented above.

Apparently there are no vertebrate feeders on this mechanically injurious forest range plant.of phenology t

Ottawa County, Oklahoma. September; peak bloom and early fruit-set stages of phenology.

268. Shoot at fruit time- Upper portion of a shoot of wood nettle with young fruit developing in the pistillate inflorescence. This plant (shoot) was one of many in the colony presented above.

Ottawa County, Oklahoma. September; ripening fruit.

269. Slender nettle (Urtica gracilis= Urtica dioica ssp. gracilis )- A second species in the nettle family that formed extensive colonies and was a local dominant forest range forb in the climax hackberry-American elm-sycamore bottomland forest described herein.

Ottawa County, Oklahoma. July.

270. Shoot of slender nettle- Detail of one of the shoots in the colony of Urtica dioica ssp. gracilis shown above that was growing on the climax bottomland forest on floodplain of Modoc Creek.

Ottawa County, Oklahoma. September.

271. The third Urticaceae species in bottomland forest- The third species of nettle in the hackberry-American elm-sycamore floodplain forest of Modoc Creek was tall nettle (Urtica procera= U. dioica var. or ssp. procera). This is the more common form of U. dioica. It is found along most streams in the Ozark Mountains. It grows to twice the height of U. dioica ssp. gracilis, at least in the forests described herein. The plants presented here and in the next photograph were growing along Lost Creek, a stream sharing a confluence with Modoc Creek. Individuals of tall nettle along Modoc Creek were more shaded and did not provide photographs that were as clear and detailed as those shown here.

Ottawa County, Oklahoma. November.

272. Stings by a stream- Upper portion of a shoot of tall nettle that was growing beside Lost Creek. According to Steyermark (1963. p. 567) young shoots of this species makes up one of several native forbs used as a pot-herb or what hillbillies know simply as "greens" (ie. Ozarker spinach). It thus shares the same pot as pokeweed, wild lettuce, dock, and other forbs.

Ottawa County, Oklahoma. Novermber.

273. Flowers on a stinging shoot- Pistillate inflorescences at leaf axils (union of shoot and leaf petiole) of tall nettle. Details of an individual shoot of Urtica procera= U. dioica var. or ssp. procera. Along Lost Creek, Ottawa County, Oklahoma. October, early bloom stage.

274. Two forbs of the wet woods- Allegheny or square-stemed monkeyflower (Mimulus ringens), on the left, and Cardinal flower (Lobelia cardinalis), on the right, growing on the coarse gravel, rich alluvium of a small stream flowing through a mixed hardwood bottomland forest in the western Ozark Plateau. Monkeyflower was treated immediately below while cardinal flower was cover farther below under an example of an ash-elm-hackberry that developed as a spring-fed wetland forest.

Bank of Modoc Creek, Ottawa County, Oklahoma. Late July; peak-bloom phenological stage in both species.

Arguably, the showiest of all- Cardinal flower (Lobelia cardinalis) is commonly regarded as the (certainly, one of the) most showy, picturesque, flashy, etc. of all the woodland forbs of the eastern deciduous forest. Cardinal flower thrives best in wet, particularily heavy wet clay, soils such as shown in the first of these two slides which was in an artesian spring site. Cardinal flower can also do well in very rocky soils--so long as it stays wet--as shown in the second of these two slides which was taken on the banks of Modoc Creek, a sream in the far-western Ozark Plateau. These plants were almost four foot tall. Amazing plants. The author had to share the treat.

Ottawa County, Oklahoma. Early August.

Ain't none purtier- Upper shoot of cardinal flower showeing leaves and inflorescence (first slide) and several individaual flowers within this inflorescence (second slide). that grrew on the banks of Modoc Creek, an Ozark Plateau stream. This lovely secimen was growing in a forest of sycamore, white ash, hackberry, box elder, and northern red oak.

Ottawa County, Oklahoma. Early August.

275. Monkeying around the crick- Several plants of Allegheny or square-stemed monkeyflower (Mimulus ringens) growing along the banks of a typical Ozark stream in the western Springfield Plateau. "Typical" in this usage implies severe degradation by human action. It was not known whether past abortive efforts at re-routing the stream channel and destructive logging, accompanying by increased rates of flashflooding and severe soil erosion, had impacts on this species of hydrophyte. The nearest neighboring species was cardinal flower (Lobelia cardinalis), a climax species.

Bank of Modoc Creek, Ottawa County, Oklahoma. Late July; peak-bloom phenological stage.

276. All monkeyed up- Closer-in views of plants of Allegheny monkeyflower that were introduced immediately above. This is a hydrophytic species (Gleason and Cronquist, 1991, p. 469) that was growing along a drastically altered (more-or-less destroyed for the foreseeable future in human time scale) stream in the Ozark Highlands (Springfield Plateau). Forage value of this species was unknown, but there was no use made (no defoliation) of it, and a resident herd of white-tailed deer frequented this location and made use of grasses and grasslike plants in close proximity to these plants.

Bank of Modoc Creek, Ottawa County, Oklahoma. Late July; peak-bloom phenological stage.

277. What comes from monkeying around- Details of flowers and leaves of Allegheny monkeyflower growing on gravel bed of an Ozark Plateau stream that effectively had been rendered dysfunctional by human action. Stems of this hydrophyte are four-angled and, even, slightly winged (Gleason and Cronquist, 1991, p. 469).

Bank of Modoc Creek, Ottawa County, Oklahoma. Late July; peak-bloom phenological 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.

278. Renewal in the riparian zone- A short--and representative--streatch of riparian vegetation along the bank of a small Ozark Plateau stream during mid-summer. A seedling of sycamore (one of the very few of its kind along this reach of the stream) of two or three years age was growing with with rice cutgrass (Leersia oxyzoides) and ditch stonecrop (Penthorum sedoides).

This portion of the stream bank had been drastically impacted by logging of both destructive (high-grading) and illegal (timber theft) action. Logging equipment had scrapped most of the riparian vegetation from this part of the stream bank, but this permitted germination of the seed of sycamore while rice cutgrass ha come back from rootcrowns in the favorable habitat of permanently wet, stoney soil of the riparian zone. Ditch stonecrop is well-adapted to disturbance, especially in such moisture-favorable environments. Cardinal flower (Lobelia cardinalis), which was growing in several large populations along this stream bank prior to logging had not fared so well: it was obliterated--totally eliminated--by soil-displacement of the timber theft activities.

Modoc Creek, Ottawa County, Oklahoma. Late July; mid-estival aspect.

279. It can cut (sort of)- A colony of rice cutgrass (Leersia oxyzoides) growing on the immediate bank (and actually in the stream itself) of a stream in the western Ozark (Springfield) Plateau. This local population of the aquatic grass species was in the stand of riparian range vegetation presented in the immediately preceding slide. This colony of rice cutgrass was growing with ditch stonecrop.

Some of the shoots of rice cutgrass seen here were starting to turn a brown color as they were dying at end (completion) of the annual growth cycle of plants in this population of this species, a perennial oryzoid grass.

Bank of Modoc Creek, Ottawa County, Oklahoma. Late July; grain-ripe stage.

280. It cut a fine figure- Composite view of leaves and part of the panicle of rice cutgrass, the dominant species of riparian range vegetation at edge of a mixed hardwood bottomland forest in the Springfield Plateau. Rice cutgrass is one of only three grasses (all Leersia species) of oryzoid grasses (those in subfamily, Oryzoideae; tribe Oryzeae) growing naturally in this region (Steyermark, 1963). One (L. virginica, white grass) of the other two Leersia species was also found in this mixed hardwood bottomland forest and treated above.

Bank of Modoc Creek, Ottawa County, Oklahoma. Late July; grain-ripe stage.

281. Ozark rice (kind of)- Panicle of rice cutgrass (first slide) and closer view of part of that panicle (second slide) growing as the dominant of riparian vegetation at edge of a mixed hardwood bottomland forest range in the western Ozark (Springfield) Plateau.

Bank of Modoc Creek, Ottawa County, Oklahoma. Late July; grain-ripe stage.

Another example below: More slides of another specimen of rice cutgrass were presented below (under Ash-Elm-Hackberry Forests).

Organization reminder: Grasses and grasslike plants (specifically, Carex species) growing on this bottomland forest were presented above (between sections devoted to the actual floodplain and a tributary of Modoc Creek).

282. Growing in a ditch- Ditch stonecrop (Penthorum sedoides) growing in a stream typical (anymore) of the Ozark Plateau. These plants were at the edge of a stand of riparian vegetation along with rice cutgrass. This annual aquatic (at least, semi-aquatic) forb (Fassett, 1957, p. 239) is relatively widely distributed as, for example, across Missouri (Steyermark, 1963, ps. 772, 774).

Palatability of this aquatic forb was not known to this author, but there had been no feeding on these plants by white-tailed deer, the only grazing ungulate on this bottomland forest range. Likeswise, sucessional status of ditch stonecrop was unknow, but it is a perennial whch suggested that at least this aquatic plant (Fassett, 1957, p. 239) was not a weedy pioneer or colonizing forb species.

Modoc Creek, Ottawa County, Oklahoma. Late July; peak-bloom phenologyical stage.

283. Starting a crop in the ditch- Apices of shoots (first slide) and detailed view of a single shoot apex (second slide) of ditch stonecrop growing in a stream in the Springfield Plateau. This local population was part of the riparian range vegetation dominataed by rice cutgrasss that was introduced above.

Modoc Creek, Ottawa County, Oklahoma. Late July; peak-bloom phenologyical stage.

284. To whom do we belong?- Flowers and apical leaves of ditch stonecrop. As indicated by both common name and the specific epithet of sedoides, this typically aquatic species has been interpreted as being in either the stonecrop family (Crassicaceae), as for example by McGregor et al. (1986, p. 356-357) and Fassett (1957, p. 239), or, more frequently, in the related saxifrage family (Saxifragaceae) as treated by Fernald (1950, p. 736), Steyermark, 1963, p. 773-774), and Smith (1977, p. 147). Fernald (1950, p. 736) treated ditch stonecrop as being in Penthoroideae, a sumfamily of Saxifragaceae. In the most recent (as of this writing) treatment, Haines (2011, p.703) placed this species in its own family, Penthoraceae.

Modoc Creek, Ottawa County, Oklahoma. Late July; peak-bloom phenologyical stage.

285. Shorter stage star- Fruiting body of an earthstar (Geastrum fimbriatum) on floor of the hackberry-American elm-sycamore floodplain forest of Modoc Creek. This member of the Geastradaceae is in the order Lycoperdales which includes the larger and generally more conspicuous puffballs. The central pollen sac had begun to dry and shrivel up indicating that opening and relaease of spores was only a few days away.

Ottawa County, Oklahoma. June.

286. Rotter on a root- Black-staining polypore (Meripilus giganteus= M. sumstiner= Grifola giganteus), member of Polyporaceae, growing on a major surface root of an American elm that died of elm phloem necrosis or elm yellows. Causive agent is a phytoplasma, bacteria that are obligate parasites which attack phloem. Elm yellows is transmitted by insect vectors including the whitebanded elm leafhopper (Scaphoideus luteolus ), the meadow spittlebug (Philaenus spumarius) and the introduced Eurasian species of leafhopper (Allygus atomarius). In this instance, elm phloem necrosis had been transmitted by removal of a large lower limb by a contaminated chainsaw. The specimen of black-staining polypore did not seem to care about the source of the pathen that provided it with a fitting food source. This was the fifth year following death of the once-stately elm.

Ottawa County, Oklahoma. July.

287. Rotter in another year- Fruiting bodies (caprophores) of the same genetic plant (genotype) of black-staining polypore as shown above. This and the next four images (the two immediately succeeding two-slide/caption sets) were of caprophores two years after the image immediately above was taken. The caprophores shown here and in the next two pairs of slides were produced seven yers following death of an American elm that was killed by elm phloem necrosis (elm yellows) spread by a contaminated chainsaw.

Ottawa County, Oklahoma. Early July.

288. Bigger rotter in another year- Two images of a caprophore (so-called fruiting body) of black-staining polypore that grew on the rotting lateral surface root of a large, dead American elm root killed by elm phloem necrosis. The first of these two slides was obviously taken in full sun while the second was taken in shade. a The second image of the fruiting body was taken from the opposite side of the first image as well s being at closer camera distance.

This caprophore was 18 inches across.

Ottawa County, Oklahoma. Early July.

289. Rotter skin- Details of a portion of the fruiting body (caprophore) of black-staining polypore that was shown in the immdiately preceding pair of images. The first of these two slides presented the adaxial (upper or dorsal) surface (left) and the abaxial (lower or ventral) surface (right). The second image presented the adaxial surface at closer camera distance and with greater detail.

The Polyporaceae (polypore family) is in the Basidomycetes, a former taxonomic classs (Class level in classification or systematics hierarchy) that is now regarded as being in the division, Basidiomycotina. The name comes from basidiospore which is a sexual spore produced by and borne on an anatomical structure (typically a single cell) called a basidium.

Ottawa County, Oklahoma. Early July.

290. Predator on the same root- A quick eyescan farther down this rotting root takes the class from decomposer to invertebrate consumer. A large, and gravid Carolina wolf spider (Hogna carolinensis) was well-camoflaged on the decaying root of an American elm that died of elm phloem necrosis while this colorfully adorned arthropod "laid-low" in wait for unsuspecting prey, themselves finding food and shelter in their little part of the world within a hackberry-elm bottomland forest in the Ozark Plateau.

Life amid death: the state of all life. Another short lesson in Range Ecology.

Ottawa County, Oklahoma. July.

291. Oysters on a dead tree- Caprophores, fleshy fruiting bodies (spore-bearing parts or organs of fungi), of elm oyster fungus (Agaricus ulmarius= Pleurcrotus ulmaria= Hypsizygus ulmarius) on the snag of a young adult American or white elm that died of elm phloem necurosis. This disease, to which American or white elm is extremely susceptible, is caused by a "mycoplasma-like organism" that is transmitted by a the insect vector, whitebanded elm leafhopper (Scaphoideus luteolus). In the instance of this formerly very healthy American elm, died of elm phloem necurosis when the disease-causing organism was spread by clueless, careless, etc. workers using contaminated chainsaws for tree-pruning at edge of a powerline corridor.

Actual cause of tree death was largely irrelevant to this lesson which was to present a native fungal detrivore (any organism that feeds on detritis or dead organic matter) that represented decomposers or reducers, members of the biotic component of ecosystems whose role or function is to decompose or rot dead material.

The elm oyster mushroom possibly has medicinal value being suspected of possessing compounds that serve as antioxidants as well as having anti-tumor and anti-inflammatory properties.

Elm oyster is in the Basidiomycetes (the taxonomic division of filamentous fungi that are distinguished as having basidia, sexual organs producing basidiospores which are specialized sexual spores), order Agricales and family Tricholomataceaee, a "catchall" group of gilled mushrooms producing pale-colored spores.

The elm oyster mushroom is technically speaking edible (though apparently not particularily tasty to those so inclined to eat such fungal "fruits of the woods").

Ottawa County, Oklahoma. Late December.

292. Oysters up closer- Top view (first slide) and bottom view (second slide) of caprophores of elm oyster mushroom growing on a rotting snag of a young American elm that died from elm phloem necrosis.

Gilled mushrooms are in the fungus order, Agaricales.

Ottawa County, Oklahoma. Late December.

293. Oysters dried out- "Spent" caprophores of elm oyster mushroom on the dead trunk of an American elm that died of elm phloen necrosis. These organs were photographed at the same time as the live ones shown in the two immediately preceding slide/caption sets. In fact, this group of caprophores can be seen on the left margin of the elm trunk as shown in the first slide that introduced this fungus.

The elm oyster toadstool is a common and widely distributed fungus. The species was included here to show students the extent of plant life in and the vegetational structure of a mixed hardwood bottomland forest.

Ottawa County, Oklahoma. Late December.

Ash-Elm-Hackberry Forests- Example From a Spring-Fed Wetland In the Ozark Plateau

The next series of slides and captions that began immediately below covered another form of bottomland forest that had developed in close proximity to that described in the last (preceding) slide series. Sycamore and pawpaw were key or indicator species (tree and shrub, respectively) in both forest range types, but the current forest vegetation was that of a wet bottomland (versus the mesic bottomland of the preceding forest) habitat.

294. White ash (Fraxinus americana)-sycamore (Platanus occidentalis) wet bottomland forest- A natural spring in this bottomland at the western edge of the Ozark Plateau supported a unique combination of mesophytic and hydrophytic species. The dominant tree was white ash based on both relative forest canopy cover, number of mature trees, and regeneration of young trees. Sycamore was conspicuous by the white, exfoliating bark of large tree trunks, but there were no trees of immature age classes (ie. no reproduction). From perspective of tree size and apparent age, sycamore was interpreted as semi- co-dominant to ash (an ecological-successional status between that of a dominant and an associate species). Black oak (Quercus velutina) and northern red oak (Q. rubra) were obvious associate (tree with retained dead leaves in right background); hackberry (Celtis occidentalis), boxelder (Acer negundo), and red or slippery elm (Ulmus rubra) were also present in several age classes with regeneration of hackberry and boxelder prouonced. Larger elms had succomed to Dutch elm disease (Ceratocystis ulmi). Black walnut (Juglans nigra) and chinkapin oak (Q. muehlenbergii) grew on better drained soils at outer edges of this community.

The lower shrub layer was dominated by buckbrush or coralberry which occurred throughout the understorey. Pawpaw and gray dogwood (Cornus racemosa) grew at restricted, local scale. Woody climbers, which grew abundantly on trunks and high into the crowns of almost all the larger trees, consisted of trumpet creeper (Campsis radicans), Virginia creeper, and poison ivy.

Herbaceous species were covered in succeeding photographs. The green patches of understorey visible in this slide were composed of two sedge species (Carex lurida and C. lupulina).

The author observed use of this vegetation-- specifically that immediately adjacent to the spring-- in autumn by migrating woodcock or timber doodle (Philohela minor). There were many probe holes (earthworms were most likely the sought prey) in the mud of this habitat. All-in-all, a most unique combination of biotic range community.

Ottawa County, Oklahoma. Hiebernal aspect, December. One of numerous forms or variants of wet bottomland forest based on classification of natural communities by Missouri Natural Areas Committee (1987). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). Combination of SAF 93 (Sugarberry-American elm-Green Ash ) and SAF 94 (Sycamore-Sweetgum-American Elm). Too small for Kuchler units. Mixed Hardwood Series in Southeastern Swamp and Riparian Forest biotic community of Brown et al. (1998). Wet Bottomland Forest (Nelson, 2005, ps.153-157). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

295. White ash-sycamore wet bottomland forest in summer verdure- Same vegetation as seen in winter aspect in the preceding slide. The dominant herb averaged over entire herbaceous layer was cyber or lurid sedge (Carex lurida). It was frequently "accompanied" by hop sedge (C. lupulina). The dominant forb in the vernal aspect was wake robbin (Trillium sessile); dominant forb in estival aspect was spotted jewelweed (Impatiens capensis) and halbertleaf mallow (Hibiscus militaris). Other forbs present were cardinal flower (Lobelia cardinalis) and swamp milkweed (Asclepias incarnata). Dominant grass in and along the spring was rice cutgrass (Leersia oryzoides) while woodreed grass (see slides above) occurred farther from the aquatic habitat and scattered among the trees. The floating green plant on the water surface was lesser duckweed (Lemna minor).

Ottawa County, Oklahoma. Estival aspect, July. Designation of wet bottomland forest was determined from classification by the Missouri Natural Areas Committee (1987). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). Combination of SAF 93 (Sugarberry-American Elm-Green Ash) and SAF 94 (Sycamore-Sweetgum-American Elm). No Kuchler unit at this small scale. Mixed Hardwood Series in Southeastern Swamp and Riparian Forest biotic community of Brown et al. (1998). Wet Bottomland Forest (Nelson, 2005, ps.153-157). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

296. White ash-sycamore wet bottomland forest- The rice cutgrass and sedge understorey (in and immediately adjacent to the spring feeding this forest range community) was shown here at peak standing crop. The saplings were all green ash. Regeneration of green ash established this as the dominant species of this vegetation. Green ash was rated as Intermediate in tolerance and Moderate in flood tolerance; sycamore had these same ratings (Wenger, 1984, ps. 3, 7). Pawpaw is visible in far left foreground. Trunk of mature tree is green ash with 32 inch DBH (shown immediately below).

Ottawa County, Oklahoma. Estival aspect, July. FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). Combination SAF 93 (Sugarberry-American Elm-Green Ash) X SAF 94 (Sycamore-Sweetgum-American Elm). Mixed Hardwood Series in Southeastern Swamp and Riparian Forest biotic community of Brown et al. (1998).Wet Bottomland Forest (Nelson, 2005, ps.153-157). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

Wet intrior- Inside a mesic to wet white ash-sycamore-northern red oak that developed around an artesian spring in the western Springfield Plateau of the Ozark Highlands. The herbaceous understorey featured here consisted of hop caric sedge (Carex lupulina), rice cutgrass (Leersia oxyzoides), sessile-flowered wake robin (Trllium sessile), spotted jewel weed or spotted touch-me-not (Impatiens campsis), cardinal flower (Lobelia cardinalis), Virginia smartweed or Virginia knotweed (Polygonum virginianum), winged monkey flower (Mimulus alatus), bur-cucumber (Sicyos angulatus), ditch stonecrop (Pentharum sedoides), sweet anise (Osmorhiza longistylis), and moon-seed (Menispermum canadense).

Shrubs included pawpaw (Asimina triloba), trumpet creeper (Campsis radicans), and ;poison ivy (Rhus radicans= R. toxicodendron=Toxicodendron radicans=).

Ottawa County, Oklahoma. Estival aspect, Early August. FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). Combination SAF 93 (Sugarberry-American Elm-Green Ash) X SAF 94 (Sycamore-Sweetgum-American Elm). Mixed Hardwood Series in Southeastern Swamp and Riparian Forest biotic community of Brown et al. (1998).Wet Bottomland Forest (Nelson, 2005, ps.153-157). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

Wet understorey- Closer-in views of the understorey of a white ash-sycamore-northern red oak wet or, at least, mesic forest that developed around an artesian spring in the western Springfield Plateau. Herbaceous species were mostly forbs such as sessile-flowered wake robin, spotted jewel weed or spotted touch-me-not, cardinal flower, Virginia smartweed or Virginia knotweed, winged monkey flower, bur-cucumber, ditch stonecrop, sweet anise or sweet cisely, and moon-seed. Hop caric sedge was the principal grasslike plant while rice cutgrass and were the grass species.

Sweet anise and spotted jewelweed were feature in the third slide.

297. White ash (Fraxinus americana)- The trunk of this mature white ash is the one shown in the preceding slide. DBH: 32 inches. The grasslike herb at base of trunk was lurid or cyber sedge. Ottawa County, Oklahoma. December.

298. Leaves of white ash- Ottawa County, Okahoma. July.

299. Bumper crop of samaras- Leaves and heavy yield of fruit in a fairly young white ash graowing on banks of Modoc Creek, Detail of leaves and samaras on a typical twig. Ottawa County, Oklahoma. June.

300. Once in a while- White ash does not bear fruit regularly. More often than not trees of this species do not produce samaras in the western Ozark Plateau. The reason for this relatively rare sexual event was not known. It was almost assuredly not due to water stress given that trees such as the specimen "sampled" here grow on stream banks and even on inundated habitats such as around around springs. Once in a while, however, its a "Katy bar the door", "bang-up" year with whoppin' yields of ash fruit. Evidence of such a phenomenon was offered here.

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

301. Samara study- Cluster of samaras of white ash laid across lichen-covered, fallen branches of the parent plant. The samara fruit type is big-winged achene. A more detailed definition is a dry, indehiscent, one-seeded, winged fruit (Smith, 1977, p. 307)..

302. Another year, another example- Leader and leaf with clustr of samaras (first slide) and details of cluster of samaras of this unit (second slide) of white ash that grew along a spring in the western Springfield Plateau section of the Ozark Plateau.Ottawa County, Oklahoma.

Identification note: Identification features and published morphological characteristics of leaves and fruit that distinguish (supposedly) between white ash and green ash are often conflicting and generally confusing. For instance many, if not most, authorities (eg. Harlow et al., 1979; Elias, 1980), described leaflets of white ash as having margins varying from entire to serrate or toothed. There remains considerable variation and apparent descreptancies in descriptions and identifying features of samaras both green and white ash when different flora and texts were consulted. Atlas of the Flora of the Great Plains (McGregor et al., 1977, p. 298) and the USDA Plant Profile (www..usda.gov/java/profile?symbol=FRPE) did not show green ash as occurring in Ottawa County, Oklahoma which was location of photographs of ash presented above. Given, however, that maps showed green ash occurring in all counties contiguous with small Ottawa County, Oklahoma it seemed likely that green ash does grow there. Steyermark (1960, p. 1178), the definitive flora closest to Ottawa county, Oklahoma, distinguished between samaras of green and white ash on basis being winged more than 1/3 vs. no more than 1/3 and having 2-4 vs. more than 4 ridges on portions of the wings covering seeds. Based on these features and plant distribution maps provided by thedefinitive authorities (at time of this writing) ash trees and component parts (leaves, samaras, twigs) presented above were white and not green ash.

303. Ice ain't nice- Severe damage from an ice storm to trees in a green ash-sycamore wet bottomland forest in the Ozark Plateau. A late autumn precipitation event resulted when warm, moist air from the Gulf of Mexico rose over cold, dry Arctic air (a Norther) resulting in the "winter mix" of, in order, rain, freezing rain, sleet, and snow that accumulated as ice to depths of one to four inches. The heavy accumulations of ice added so much additional weight to trees, power lines, roofs, etc. that numerous of these objects crashed to the ground. Such precipitation events, known as ice storms, can cause some of the most incredible damage in Nature to trees. Locally, extreme damage to trees and shrubs may rival that from tornadoes and hurricanes. Area of impact is frequently larger than that from tornadoes. The ice storm responsible for the damage shown here and in subsequent slides cut a swath about three hundred miles long and a one hundred and fifty miles wide.

The wet bottomlad forest shown here was the green ash-sycamore community introduced above that developed around a natural spring. Most of the downed crowns and individual limbs at this site were from sycamore, the locally dominant tree, but Shummard oak, black oak, and green ash also suffered major damage.

304. The double-edged sword of water- A hollow and nearly-dead-but-still-standing sycamore was toppled by heavy ice accumulations resulting from late autumn ice storm caused by the atmospheric phenomenon of overriding (warm, moist air rising above cold, dry, heavy air resulting in mixed precipitation of rain, freezing rain, sleet, and snow). This old sycamore and its cohorts, including its side-by-side "sister" tree, became established and survived as facultative phreatophytes along a natural spring in the western edge of the Ozark (= Springfield) Plateau. The ice-downed sycamore was the left sycamore (leftmost, conspicuous, white-barked trunk in left midground) shown in the first slide above that introduced this local green ash-sycamore wet bottomland forest.

Ponded water from the spring was visible in the left midground of the first of these two slides. Ironically it was liquid water that permitted establishment and continued survival of sycamore and it was solid (frozen water in form of ice) that finally ended the life of the old, toppled sycamore. Details of the hollow trunk and its downfall were covered in the immediately following set of photographs and caption.

From perspective of plant succession and development of this wet bottomland forest the most obvious change in this forest community was the on-going replacement of sycamore by western hackberry. The characteristic netted or deeply furrowed bark of the latter species was distinctive on the trunk of the pole-size tree in left foreground of the first photograph and in the three centermost saplings (included the leftmost one on which the sycamore had fallen and bent). The sapling on the right of the first slide and in left midground of second slide was a sapling of Shummard oak (the same sapling in both photographs incidentially). These species dynamics were already in play several years before the old sycamore fell. They will certainly be accelerated thereafter. This pattern of species replacement (the succession of plant species) was taking place at the edge of the wet bottomland forest. In closer proximity to the pool of spring water sycamore and green ash were still the species having greatest recruitment and, thus, were an edaphic climax (in polyclimax theory), postclimax (in monoclimax theory), or a climax that developed along a gradient in which soil moisture was determinative (in climax pattern theory).

The lack of sycamore reproduction and, instead, reproduction of western hackberry was consistent with changes in the hackberry-American elm-bitternut hickory floodplain forest along Modoc Creek described in detail above. These two forest range communities were about 350 yards apart.

305. Hollow-grounded- Details of the hollow and almost-dead-though-still-standing sycamore shown above that was felled by heavy accumulation of ice in late autumn ice storm. Hollow trees (trees with hollow trunks), contrary to the erroneous popular view, are usually about as strong as trees with solid trunks. Not quite, but close. This is the same basic physics as the fact that under certain kinds of strain a steel or iron pipe is quite strong compared to a solid piece of steel or iron of the same metal and diameter. This is why shafts of bird feathers, the hollow culms of many grasses (think bamboo), and hollow airplane wings (used as fuel tanks with different degrees of fullness) are very strong, especially when compared to their weight. In the same manner hollow trees can stand much strain and stress. In the case of this sycamore the added weight of accumulated ice (about two inches) and accompanied with gusty winds was more than the trunk that was 9/10ths or more hollow could bear up under. In this same ice storm there were large, completely intact trees (including sycamore, American elm, and hackberry) that also crashed. It was no wonder that this over-ripe oldster ended its life cycle thusly.

There was a small, but very deep fire scar on the base of the fallen sycamore that appeared to the point of entry of decay agents (fungal mycelium were present) that permitted decomposition of the dead wood (heartwood) inside the trunk. The lower end of this fire scar was visible in the second of these photographs (at extreme left margin of the photograph about 3/8ths of the vertical distance from bottom of slide). Referral to any freshman Botany text will remind viewers that the layers tissue of the tree trunk moving from interior to exterior were pith (very little of that); xylem of heartwood; followed consecutively by living xylem, cambium, and live phloem; and, finally, bark. It was the heartwood, which in a tree of this age comprised vast bulk of the trunk, that was susceptible to decay by microorganisms and fungii. In fact, in this tree there was remarkably little sapwood and bark to keep this old fellow upright. It was a testament to the strength of hollow trees that this more-dead-than-alive, rotten sycamore stood as long as it did and that it took such extreme stress to bring it down.

Even down this sycamore remained a critical component in the forest range ecosystem of which it was part for decades. As this trunk continues to rot it will release nutrients stored in wood to enter the biogeochemical cycles of carbon, nitrogen, etc. It hollow will undoubtedly serve as a den for coon, possum, feral house cat, etc.

Ottawa County, Oklahoma. December.

Life cycle- To "go the way of all the earth" (1 Kings 2:2 KJV).Like all living things trees start life, grow up, grow old, decay, and die (and, ideally, reproduce in this endles cycle of life). The following four sets of slides (in addition to the three sets of slides immediately above) documented the death of late maturity, senescing and decaing trees of sycamore and northern red oakk in wet-mesic bottomtland forest that developed around an artesianspring in the westeernOzark Plateau physiographic province.

Down it came-. A late-life (in relative terms, ancient and dying) northern red oak was an uprooted victim in a wet bottomland whte ash-sycamore-northern red oak forest that developed around an artisian spring in the westernSpringfield Plateau of the Ozark Highlandss. The straight-trunked oak was about two foot diameter breast height and was completely hollow exceept for a narrow ring of sapwood. The tree had obviously been hollow for several decades, but showed no signs other than for some dead uppermost branches, a sign of aging and senescence. Otherwise the tree appeared to still be prime lumber from outer appearances. Older roots in the peg root system of oaks this age were likely also hollow and weaker than in a younger tree. The old forest denizen was on the way out when it came down. It was not known if this uprooting was windfall or whether it just finally succumed to gravity with hollow upper roots,a in the always-wet soil.

This example of treefall provided an example of a tree cradle, the "pit" or cavity left in the soilThe life cycle of this tree had ended, but it left a cradle for it's own (or for other tree's) progeny. The herbaceous plant growing around the tree cradle were mostly forbs lincluding sessile-flowered wake robin, spotted jewel weed or spotted touch-me-not, cardinal flower, Virginia smartweed or Virginia knotweed, winged monkey flower, bur-cucumber, ditch stonecrop, sweet anise or sweet cisely, and moon-seed, but there was also rice cutgrass, hop caric-sedge, and . The plants in the foreground of the third slide were those of a colony of spotted touch-me-not, The second most abundant forb was Virgina knotweed.

The only tree regeneration (by sexual reproduction) was of white ash.

Over it went- Another very old, "over-mature" northern red oak came down in the same wet bottomland forest of white ash, scamore, and northern red oak that was shown above. This northern red oak had a forked trunk very near the soil surface. Like the northern red oak presented immediately above in a three-slide set, this trunk was almost completely hollow and even without very much sapwood. This was obviously "death of natural causes" and probably without any contributing physical force other than gravity although wind cannot be ruled out. If wind was a factor in the fall it was windthrow of breakage and not uprooting as was the case immediately above. Either way one of the two trunk forks had completed its life cycle.

Neighboring plants were the same as listed above, but visible in foreground of the second slide were the woody vine, trumpet creeper and shrub, buckbrush (Symphorocarpus orbicularis) along with Virginia smartweed, spotted jewelweed, and sweet anise or sweet cisely. In immediate background were large seedling/small saplings of white ash.

There was no regeneration of northern red oak or of sycamore in this forest.

Waiting to go- Trunk base of an "ancient" (senescing; "more dead than alive") sycamore with its hollow trunk rotted clear through to the outside. Like the northrn red oak discussed immediately above this was a two-trunked tree. It was only a matter of time until this sycamore joined its two neighboring northern red oaks that only a few feet away had already succumbed to old age death and crashed to the soil that had supported them for decades.

It is a common observation that most of the old sycamores growing along water courses are mostly hollow(Steyermark, 1963, p.).790). Hollows of all trees can serve as dens for cavity nesting animals like coons (Procyon lotor) and ''pssums (Didelphis virginiana).

Plants growing in front of the trunk were young shoots of trumpet creeper.

Ottawa County, Oklahoma. Late July.

306. Sycamore (Platanus occidentalis)- Distinctive bark, branching pattern, and habit of sycamore. This massive old-growth specimen of sycamore was growing along the flood plain of Modoc Creek (Ottawa County, Oklahoma) and was over 7 feet DBH (where the branch-like sprout originated). Authorities regard the sycamore as one of the hardwood (angiosperm) species that attains largest mature size (especially in diameter) in North America.

Sycamore is interpreted as a pioneer species that persist into the climax forest, often as ancient behemenths like this splendon specimen. Such trees often live for decades as a member of climax riparian vegetation where they function as facultative phreatophytes. Even at some distance from streams sycamores like the one presented here get a continuous water supply from the water course by lateral flow. April.

307. Crown and upper trunk of intermediate-age sycamore. The exfoliating, mottled mature bark against a background of grayish immature bark is a distinctive pattern to those familar with the eastern deciduous forest. Newton County, Missouri. December.

308. Leaves and fruit of sycamore- Ottawa County, Oklahoma. July.

309. Sexy sycamore- Leader of sycamore bearing a male and feamae inflorescence. More detailed (close-up) views of these were presented immediately below. Lampasas County, Texas. April.

310. Male equipment- Staminate flower clusters (heads)of sycamore consist of a myriad of stamen mingled in with tiny scales. Lampasas County, Texas. April.

311. Female equipment- Pistillate flower clusters (heads)of sycamore are comprised of numerous ovaries mixed up with minute scales. Lampasas County, Texas. April.

312. Mature fruit of sycamore- Fruit of sycamore is a dry aggregate of achenes. The second of these slides showed numerous of these achenes close up. The native hardwood with the largest trunk diameter and second overall in size begins it growth as a tree from this small dry fruits. Another of God's miracles in the woods.

313. Part of sycamore's secret- In contrast to detrminate growth of many other--probably most-- hardwoods (eg. oaks, hickories, walnut) and conifers with preformed buds at shoot (leader) tips, sycamore and some other angiosperms like sweetgum and willows have indeterminate growth. Whereas extension of the branch in determinate growth is in one seasonlong bout of growth (at least as growing conditions are favorable and at rates consistent with growth favorability) indeterminate growth occurs in phases or "flushes". These flushed are obvious when new leaves appear periodically throughout course of the growing season

Examples of new leaves at tips of sycamore can often be seen throughout summer months. The three photographs provided examples of such recurrent or flushes of growth at ends of lower branches in sycamore growing on banks of Modoc Creek, Ottawa County, Oklahoma. June.

314. Saprophytic teeth on a sycamore- Fruiting body of bearded tooth fungus (Hericium erinaceus). This outstanding specimen of this picturesque species somehow made itself "to home" in the tall crotch of this more-dead-than-alive old sycamore. How the spore found its way to this habitat is anybody's guess. H. erinaceus is not common, but it is found far more frequently in oak-hickory forest to the east than this one along the San Saba River in an ecotone between the eastern Edwards Plateau and the southern tip of the Cross Timbers and Prairies vegetational area of central Texas. This seemed an appropriate place to include this saprophyte on a sycamore while that tree species was being featured.The taxonomic hierarchy of H. erinaceus going from class down to family was Basidomycetes, Hymenomycetes, Aphyllophales, Hydnaceae.

Lampasas County, Texas. January, just post-maturity and beginning to dry up.

315. Next year's saprophytic toothed fruit- Fruiting body of the same bearded tooth fungus as introduced in the preceding photograph. This spore-bearing structure appeared ten months following the one shown immediately above. This fruit body grew out of the opposite side of the sycamore snag. It was photographed at a slightly earlier stage of maturity and from more of a ventral (bottom-up) view than its predecessor.

Lampasas County, Texas. November; peak maturity.

316. Gray dogwood (Cornus racemosa)- Whole plant of gray dogwood growing along the streambank of the spring shown in this series of photographs of wet bottomland forest. This is a many stemmed, sprawling riparian shrub. Ottawa County, Oklahoma. July.

317. Leaves of gray dogwood- Leaves on plant in preceding slide. Ottawa County, Oklahoma. July.

318. Colony of shallow or lurid caric sedge (Carex lurida) in understorey of white ash-sycamore wet bottomland forest- Lurid or shallow caric sedge is the species most abundant here, but there were some "strays" of hop caric sedge(C. lupulina) present. Ottawa County, Oklahoma. April.

319. Shallow or lurid caric sedge (Carex lurida)- Staminate (upper, tassel-like) and pistillate inflorescences of lurid caric sedge. In understorey of white ash-sycamore wet bottomland forest. Ottawa County, June.

320. Successful sex- Mature flower clusters of shallow or lurid caric sedge with achenes falling from pistillate inflorescence stalks.There were staminte flower clusters above the pistillate inflorescences in both of these examples. These sexual shoots had senesced (died with completion of sexual reproduction) and though they were dead, yet they were alive as they bore live fruit or, more precisely, viable seed inside of fruit.

Shallow caric sedge is a strongly rhizomatous (hence, colonal) species with most reproduction being asexual through production of more clonal shoots each of which is a modular unit of the same genotype. By contrast each achene (see immediately below) is a new geontype, a new organism of this same species produced through genetic recombination. Many of the shoots in this colony of shallow or lurid caric sedge had successfully reproduceed both sexually and asexually. The process of evolution fitted this species for survival in its wetland habitat.

Ottawa County, Oklahoma. Late July-fruit (achene) shedding stage.

321. Cluster of dry fruit on drying mud- Cluster of ripe or mature achenes (fruit type of Carex species) of shallow or lurid caric sedge. Central stalk of the pistillate inflorescence-- a spike--that held female spikelets (flowers) was serving as point of attachment for achenes which were being shed at fruit-shatter stage of phenology. Each of the fertile (viable) achenes on this spike is a potential new plant, a genetically unique individual of C. lurida. It holds the possibility that it is more fit 9better adapted) to its wetland environment than its parent. That is, assuming that the process of evolution is still in process.

Ottawa County, Oklahoma. Late July-fruit (achene) shedding stage.

322. Hop caric sedge (Carex lupulina)- Flowering shoot of hop sedge: male inflorescence (upper) and female inflorescences (below). Growing in understorey of white ash-sycamore wet bottomland forest. Ottawa County, Oklahoma. April.

323. Rice cutgrass (Leersia oxyzoides)- Small colony of rice cutgrass on an alluvial bar along an Ozark Plateau creek. Cutgrass is a hydrophytic grass that usually grows on gravel bars or along banks of streams but in spots that receive direct sunlight throughout most of the day. Cutgrass derived it's common name from the fact that the rough leaves can cause mild abrasions on bare skin.

There are two Leersia species native to the Ozark Plateau Region. Whitegrass (L. virginica) is the other species. A few isolated plants of both of these Leersia species were growing on the floodplain of an Ozark stream in the Springfield Plateau portion of the Ozark Plateau physiographic province. Whitegrass is a mesophytic species and grew farthr up on the first terrace of the stream whereas rice cutgrass grew cirectly in the channel and in the immediate riparian zone of the stream.

Location note: Given the occurrence (different habitats) of the two Leersia species, the hydrophytic L. oxyzoides, rice cutgrass, was positioned here whereas the mesiphytic L. virginica, whitegrass, was presented above in this same chapter with the mesic mixed hardwood bottomland forest. Likewise more slides of another specimen of rice cutgrass that was also growing on the bank 9actually in the stream) of Modoc Creek were presented above (under Bottomland Forests).

Modoc Creek, Ottawa County, Oklahoma. September.

Cutting (more like scrapping) shoots- Shoots of rice cutgrass that were growing on an ephemeral artesian spring that fed into Modoc Creek, an Ozark (Springfield) Plateu stream. This bunch was growing with common or broadleaf arrowhead (Sagiittaria latifolia). Yes, the stiff pubescence on the mrgins of leaves (mostly) and along the culm will pull against/along human skin and can leave minor abrasions or scratches; hence the common name of rice cutgrass.

Ottawa County, Oklahoma, feeder artesian spring into Modoc Creek, Mid-July, immediate pre-bloom phenological stage.

Cutting characteristics- The "swollen"-appearing. node adjoining internodes, and leaf axil (first slide) and details of leaves (second slide) of rice cutgrass that was growing with broadleaf arrowhead in an artesian spring that fed into Modoc Creek in the Springfield Plateau. The "swollen" node and its pubescece (prominent in both slides) is described as a pulvinus (a swollen-like structure umore commonly found at bases of leaves and inflorescences (Gould, 1975, p. 635). essually found

Ottawa County, Oklahoma, feeder artesian spring into Modoc Creek, Mid-July.

Ripening- Upper sexual shoots (featuring panicles) of rice cutgrass that was growing on an artesiaan spring (usually ephemeral) that drained into an Ozark streaam in the western Springfield Plateau.

Ottawa County, Oklahoma, feeder artesian spring into Modoc Creek, Late July, grain-ripe stage of maturity.

324. Spikelets of rice cutgrass- Modoc Creek, Ottawa County, Oklahoma. September.

524. Two colonies of broadleafed (broad-leaf) arrowhead (Sagittaria latifolia)- A wetland or, on some habitats, an aquatic plant that is widespread on wet prairie. It is another monocotyledon. First (upper) slide, Cherokee County, Kansas; second slide, Ottawa County, Oklahoma. July.

525A. Broadleaf arrowhead on tallgrass prairie- Details of leaves and inflorescence on broadleaf arrowhead. Due to presence of a conspicuous corolla the arrowhead species have traditionally been interpreted as forbs, but given that they are monocots a case could be made for regarding them as grasslike plants. Slough along Indian Creek, Newton County, Missouri. August.

Nymphaceae (water lily family) includes some of the most spectacular of all aquatic plants both from standpoints of size of plants, beauty of their flowers, and unique fruit structures. Strange as it may seem to the prairie neophyte some of these aquatic forbs are indigenous to North American grasslands and associated wetlands. A beautous sample was presented to enlighten the initiates of Range Management.

This small--though somewhat sensational--sample of aquatic prairie forbs once again presented a lesson in "lumpers" versus "splitters", this time at the taxonomic level of family. Older interpretations included the two Nelumbo species in Nymphaceae More recent treatments view this two-species genus as constituting a family all to themselves, Nelumbonaceae (Cronquist, 1981, ps. 107-111; Takhtajan, 1997, ps. 77-78, 83-84).

Further details of common or broadleaf arrowhead:

525B. Foliage and flower- Leaf and flower of broadleaf arrowhead growing in a spring in the western edge of Springfield Plateau of Ozark Highlands.

Ottawa County, Oklahoma. Mid-July..

525C. Inflorescence and single floer of common or broadleaf arrowhead growing in a spring in the far-westerrn Springfield Plateau.

Ottawa County, Oklahoma. Mid-July..

525D. Wetland fruit- Fruit of broadleaf arrowhead produced in a spring in the karst features of the far-western Springfield Plateau of the Ozark Highlands.

Ottawa County, Oklahoma. Mid-July..

Sweet and ciceley- Flowering plant of sweet ciseley or sweet anise (Osmorhiza longistylis) growing in an artesian spring in western Springfield Plateau of the Ozark Highland (Ozarl {;ateau [hysiographic province) This range forb was in the underrstorey of a white ash-sycamore-northern red oak wet bottomland forest. The overall shoot portion of the plant was presentted in the first slide wehreas the inflorescence and upper shoot were featured in the seond slide.

This specimen was not a robust plant, but it did well to srvive in the dense shade of this forest and the dense understorey of numerous competitive plants, including its nearest neighbor spotted jewelweed or spotted touch-me-not (Impatiens capensis).

Ottawa county, Oklahoma. Late July, early bloom phenological stage.

325. Halbertleaf rose mallow and cardinal flower as local dominant forbs in white ash-sycamore wet bottomland forest- This spring-fed unique form of the eastern deciduous forest (combination SAF 93 X SAF 94 forest cover types) was botanically diverse in both species and structure. The herbaceous layer was a mosaic of microsites composed here-and-there variously of tall, robust forbs; hydrophytic sedges; or grasses of varying water requirements. In this view two of the most mesic (and strinkingly photogenic) forbs native to the deciduous forest of eastern North America grew as pals in the rich mud of a bottomland forest.

This forest community was at the extreme western edge of the Ozark Plateau where it contacts the Cherokee Prairie of the Central Lowlands physiographic province. Forbs like cardinal flower are more typical of "wild flowers" found in the Applachians or Catskills. Wetland vegetation was an interesting assemblage of species usually not growing together. They also occur more commonly on habitats different from the environment seen here.

Ottawa County, Oklahoma. Late estival-early autumnal aspect, September.

326. Cardinal flower (Lobelia cardinalis)- Cardinal flower growing on the source of a spring within a white ash-sycamore bottomland forest in the western Ozark (Springfield) Plateau. This member of the the Lobelioideae (Lobelia subfamily) of the Campulaceae is unquestionably one of the showest, most brillantly colored forbs in the Eastern Deciduous Forest Super-region of North America. Several slide-caption sets were included herein to more fully cover this conspicuous forest range forb.

Other forbs in this photograph included an unidentified (at pre-bloom stage) species of knotweed or smartweed (Polygonum sp.) and beefsteak plant (Perilla frutescens), a naturalized native of the subcontinent of India.that has longbeen widespread throughout the Ozark Plateau.

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

327. A robust forb- This large specimen (over a yard in height) of cardinal flower or cardinal plant growing at the bank edge of the spring that was the basis for the wet bottomland eastern hardwood forest presented in this series of photographs. Cyber and hop sedge were visible in the background as was the trunk of the large green ash shown previously.

Ottawa County, Oklahoma. Late estival-early autumnal aspect, September.

328. Too purty not to show more- Details of shoots of cardinal flower or cardinal plant growing on a previously wet bank along a spring within a white ash-sycamore bottomland forest in the western Ozark (Springfield) Plateau. Extreme Drought (Palmer Severity Scale) for the second summer in a row induced flowering in these plants a full month earlier than is typical for this area. Such extreme water stress had not diminished the beauty of this striking forest forb.

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

329. More yet- Upper shoot (first slide) and overall view of young inflorescence (second slide) of cardinal flower growing on moist bank of headwaters of a spring within a white ash-sycamore bottomland forest in the western Springfield Plateau. Cardinal flower plants presented in this and the immediately preceding two-slide set were flowering fully one month earlier than typical for this locality due to plant an stress and advanced season and phenological stage due to Extreme Drought (the second summer in a row of such precipitation shortage) even though the root zone of these plants had been comparatively moist until the spring ran dry a few weeks earlier (prior to time of photograph).

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

330. Inflorescence of cardinal flower- Understorey of green ash-sycamore wet bottomland forest. Ottawa County, Oklahoma. September.

331. Defending (?) the dominant forb- Wheel bugs or southern assin bugs (Arilus cristatus) lurking in the immediate pre-loom inflorescences of two plants of cardinal flower. In the first photograph there is one wheel bug in both (each) of these immature inflorescences. Can ya'll find both bugs? To help you those with digitally blinded eyes the second photograph was included. This is the inflorescence on the right of the first photograph.

Wheel bugs are predators par excellence. They will feed on about any invertebrate they feel big or hungry enough to engage. In the case shown here, wheel bugs were waiting to ambush visitors to flowers of cardinal plant. Undoubtedly some of these visitors would be pollinators. Students should recall or be informed that pollination is, in many cases, a form of mutualism. The question follows, is the wheel bug benefitting cardinal plant? (See next slide-caption set for another view.)

Warning: only those skilled in handling these creatures (or being knowledably instructed in that skill) should attempt to catch these insects which can inflict a painful puncture wound with their proboscis-fitted piercing/sucking mouthparts. Another lesson: wheel bugs are true bugs. They are in order Hemiptera, those insects with piercing/sucking mouthparts. (Again, in this larger bug such equipment is can be effective against Homo sapiens).

Ottawa County, Oklahoma. Late July.

332. Biological control in a wetland forest- A wheel bug or southern assin bug "sucking the juice" out of a Japanese beetle (Popillia japonica). This exotic coleopteran (Coleoptera is the beetle order) is a major pest of many plant species (both native and introduced), especially woody plants and forbs. This hapless invader's luck ran out while it was feeding on sawtooth sunflower (Helianthus grosseserratus) growing on the perimeter of the forested wetland being described in this section.

Details could not be shown with clarity in this photograph which had to taken at an aperature of 2.8 at 1/15 second in the dim light of this forest wetland. In addition the assin bug was not keen for publicity and several times threatened "flight or fight". These insects are cantankerous characters.The tenured full professor taking the photogaph empathized with the insect, the one killing invasive vermin.

In this instance the wheel bug was doing the native range vegetation a good turn. Perhaps this predaceous bug is a mixed blessing to range plants (see immediately above), but then again most blessings are mixed with something or somebody getting the short end of the bug's proboscis. Take salvation as an example. Just a born-again thought.

Ottawa County, Oklahoma. Late June.

333. Halbertleaf rose mallow (Hibiscus militaris)- The flowering shoot of this native forb portrayed the staminal column that immediately marked it a member of the Malvaceae (mallow family). This remarkable family includes not only the greenhouse or indoor Hibiscus species but such yard beauties as Rose of Sharon (H. syriacus) and hollyhock (Althaea rosea) as well as okra or gumbo (Hibiscus esulentus), the beloved southern garden vegetable, and the world's most important fiber crop, the Gossypium species. Those familar with King Cotton will immediately note the close resemblance of the unopened flower buds on the rose mallow shown here with the cotton "square".

The descriptive name of halbertleaf is in reference to the halbert, a combination spear or pike and battle-axe used by soliders in the Fifteenth and Sixteenth Centuries. Hence, the Latin militaris. This high seed-yielding prairie and forest forb should be a favorite for those landscaping with native plants in humid regions.

Ottawa County, Oklahoma. August.

334. Colony of lesser duckweed (Lemna minor)- The common name of this "nifty" little monocotyledon is in reference to the palatable forage that it affords waterfowl. In regions where precipitation is sufficient to maintain streams and ponded water Lemna species often cover water surfaces so as to superficially resemble algae. L. minor accounts for about 90% of the Lemna species consumed by waterfowl (Martin et al., 1951, p. 448). Steyermark (1963, p. 389) reported that it was eaten by muskrat (Ondatra zibethicus) and other aquatic animals. The current author observed lesser duckweed being grazed by the red-eared slider turtle (Chrysemys scripta).

Lemna species are submerged to partly floating plants. Permanent spring, Ottawa County, Oklahoma. April.

335. Close-up of lesser duckweed- Several adult-sized plants of lesser duckweed were placed on this sycamore leaf for photographing and to portray the relative size of this tiny monocot. Distinction of individual plants, each of which has only one root, is difficult without magnification. These minute aquatic plants are monocotyledons but the sporophytic generation consist of leafless plants whose bodies are reduced to a thallus ( a plant body that is not differentiated into leaf and stem).

Permanent spring, Ottawa County, Oklahoma. April.

336. Lesser duckweed bearing fruit- Individual plants of the Lemna species are perennial and reproduction is typically vegetative. Duckweeds are monoecious with much-reduced flowers. The tiny fruit (seen here as numerous smaller, green, ovoid parts) is a utricle (an indehiscent, one-seeded, bladdery fruit).

Bottomland Forest- Example on an Ephemeral Stream in Ozark Plateau

The following example showed a forest co-dominated with both hackberry or western hackberry (Celtis occidentalis) and sugarberry (C. laevigata) along with American elm (Ulmus americana) with an understory almost exclusively of broad-leaf wood oats (Uniola latifolia= Chasmanthium latifolium). Details of the two Celtis species and broad-leaf wood oats was provided.

Part of this forest had suffered severe damage due to windthrow by straight-line winds. Uprooting of trees by combined action of wind and saturated soil from recent heavy rains furnished an example of secondary plant succession in the "tree cradles" formed by tree blowdown (uprooting).

337. Bottomland forest of western hackberry, sugarberry (Celtis laevigata), and American elm (Ulmus americana) with an understory almost exclusively of broad-leaf wood oats (Uniola latifolia= Chasmanthium latifolium) with scattered solidary plants of the shade-tolerant composite, elephant's foot (Elepohantopus carolinianus). Beautiful plant community and an example of permanent deciduous forest range in contrast to the typical transitory feature of these dense hardwood forest types. Ottawa County, Oklahoma. September, late estival aspect. No obvious FRES or Kuchler designations as this is a smaller unit. Neither was this relatively unique community described by Braun (1950) though it corresponds to general creek and river bottom vegetation in the Ozark Mountains. SAF Cover Type 93 (Sugarberry-American Elm-Green Ash). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

338. Side-by-side seedlings- Hackberry or western hackberry and sugarberry frequently grow in close proximity to each other. This can simplify or complicate identification in the field, especially when there is only one species present and even more so with large trees whose leaves are considerable distance from the observer. When Celtis occidentalis and C. laevigata grow next to each other leaves can be compared and differences between the two species are more obvious. Such was the instance presented here and in subsequent slides.

Larger (two- to three-year-old) seedlings of hackberry and sugarberry germinated and continued to grow side-by-side on a tree cradle created when several adult trees were uprooted from saturated soil by straight line winds. This tree cradle example was in the western Ozark (Springfield) Plateau provided a textbook comparison of seedlings of hackberry (on the left) and sugarberry (one seedlinmg on the right).

(Images and descriptions of several tree cradles where presented below. Creation or formation of tree cradles happens by one form of the phenomenon known as wind throw which can occur when high straight-line winds and moist soil coincide resulting in uprooting of trees, especially those with large, leafy crowns that catch more wind).

Ottawa County, Oklahoma. July.

339. Side-by-side Celtis- Closer views of leaves of the two seedlings introduced above. In both photographs hackberry (Celtis occidentalis) was on left and sugarberry (C. laevigata) on the right. Hackberry leaves typically have serrated margins while those of sugarberry are entire. Otherwise, no need for words when there are comparison photographs like these.

Fernald (1950, p. 553) suspected that hackberry and sugarberry hybridize.

Ottawa County, Oklahoma. July.

340. Two hackberry seedlings/sprouts- A seedling (left) and a sprout from a three-year-old sapling (right) of hackberry or western hackberry in the western Ozark Plateau. The sprout (secondary shoot) came from a sapling that had been broken-off by a roatary shredder. This slide provided a comparison of growth between a first-season seedling and a first-season stump sprout. It also showed variation in leaves of this species. Toothed (serate) leaf margins was a key identifiction feature.

Ottawa County, Oklahoma. July.

341. Leafy samples #1- Examples of leaves of hackberry (Celtis occidentalis). These organs were on seedlings growing in a large hole called a tree cradle formed by adult trees being urooted by high winds and wet soil. These tree cradle-hackberry seedlings were introduced above (and the tree cradles wereshown and described below).

Ottawa County, Oklahoma. July.

342. Study of sugarberry- Trunk and basal leaves of sugarberry (Celtis laevigata) in the western Ozark Plateau. This tree was growing in the same fencerow as sugarberry seedlings introduced above. Bark of sugarberry or smooth hackberry is not always as corky or knobby as this specimen, but this was a fairly typical example of this species.

Ottawa County, Oklahoma. July.

343. Study of hackberry- Trunk of hackberry (Celtis occidentalis) in the western Ozark Plateau. This young tree was growing on the floodplain of Modoc Creek. The scaly bark was typical of this species and can be contrasted to the smoother bark yet with knobs (knobby bark) of sugarberry shown immediately above.

Ottawa County, Oklahoma.

344. Leafy samples #2- Examples of leaves of sugarberry (Celtis laevigata). From the cradle tree specimen introduced above.

Ottawa County, Oklahoma. July.

345. Hacked leaders- Terminal (distal) ends of branches of hackberry. This hackberry was growing on the floodplain of Modoc Creek in the Springfield Plateau of northeastern Oklahlma.

Ottawa County, Oklahoma. July.

346. Foliage and fruit- The fruit (drupe) and leaves of hackberry growing in a bottomland forest that developed along Modoc Creek in the far-western Springfield Plateau of northeastern Oklahoma. The broad-based leaves with "saw-tooth" serrations along their margins provide simple (and key) identifying features of this climax dominant of floodplain forests. These still-green drupes were immature. They turn a dark brown to almost-black color (varying with conditions) when ripe.

Hackberry is a widely distributed species with a biological range extending from New England and Quebec to Idaho and Florida to Oklahoma and New Mexico(Fernald, 1950, p. 553).

Ottawa County, Oklahoma. July.

347. Hacked fruit- Drupes, drupe is a fruit type characterized as a fleshy or pulpy fruit with th inner portion of the pericarp... " being "hard or stony". (Fernald, 1950, p.1574), of common or western hackberry. A drupe is commonly known as a stone fruit; examples include peaches and plums. The green drupes seen here were immature. Ripe hackberry fruit is a dark red, brown to almost black.

In this author's observation a high proportion of hackberry drupes are fertile. "Zillions" of hackberry seedlings can be found scattered around adult trees. In this context, hackberry is often very invasive of grasslands and fields that are in the general area of hackberry trees.

Ottawa County, Oklahoma. July.

348. Leafy samples #3 (for real confusion)- Examples of leaves of hackberry. These leaves were produced by a young adult tree growing on the floodplain of Modoc Creek, a typical Ozark Plateau stream. The broader leaf with visible srations along its margins is an identifying feature that distinguishes hackberry from sugarberry, especially when the two species grow close together (as was shown above).

Ottawa County, Oklahoma. July.

349. Catchy branching pattern in sugarberry- A combination of alternate branches coming off of opposite sides of a leader is a frequent branching arrangement in Celtis species, especially C. laevigata. Perhaps this pattern permits large numbers of leaves with a minimal degree of shading so as to facilitate photosynthesis in forest canopies. Would make a eye-catching pattern for neck ties, particularily for dressed-up foresters.

Erath County, Texas. September; fruit-ripe stage.

350. Shiny leaves and sweet fruit (well, sort of)- Leaves and fruit of sugarberry shown in photographs that featured leaf position and density along with a better-than-typical crop of fruit. Fruit type in Celtis species is a drupe. Fernald (1950, p. 553) remarked that these sweet-tasting drupes were basis of the name Celtis as used by Pliny for what he termed lotus. Anyway, wildlife of various species consume this fruit and undoubtedly contribute to dispersal of Celtis species.

Erath County, Texas. September; fruit-ripe stage.

351. Leaves of red (though green)- Leaves on a branch of slippery or red elm (Ulmus rubra) . These were shade leaves of a large seedling or small sapling.

Ottawa County, Oklahoma. Early June.

352. Spikelets of broadleaf wood oats (Uniola latifolia)- The inflorescence of broadleaf wood oats is one of the most attractive of North American Gramineae. In extensive stands (like the one seen immediately above) wood oats creates a stikingly beautiful aspect. Uniola (= Chasmanthium) species have some of the most laterally compressed spikelets of any grass.

Ottawa County, Oklahoma. August.

353. Heavy yield on wet ground- Shoots of vigerous broadleaf woodoats plants growing on wet prairie in the western Ozark (Springfield) Plateau showed the high-yield grain crop of which this species is capable under optimum growing conditions. This images also presented the relative broad leaf blades from which this species got its common name.

The more common habitat for broadleaf woodoats is the moist deep soil of floodplains extending down to the streamside riparian zone. Dense stands of broadleaf woodoats are a characteristic of lightly grazed ranges of bottomland forests, especially those dominated by such hardwood tree species as hackberry/sugarberry, American elm, sycamore, box elder, and chinquapin oak.

Ottawa County, Oklahoma. Late July.

354. Tipping over flat- Upper parts of sexual shoots of broadleaf woodoats bearing panicles bent over with heavy yields of grain in pronounced laterally compressed spikelets. These plants were growing on wet prairie in the Springfield Plateau of northeastern Oklahoma. Abundant spikelets compressed laterally to the "unth degree" and broadly expanded leaf blades make broadleaf woodoats one of the more readily identifiable grasses on habitats varying from bottomland forests to moist grasslands.

Ottawa County, Oklahoma. Late July.

355. About as flat as they come- Laterally compressed spikelets of broadleaf woodoats as seen against early evening light. Uniola species provide some of the best examples of lateral compression of spikelets of any grasses in North America. These organs were produced on one of the many plants growing on wet prairie sshown above.

Ottawa County, Oklahoma. Late July.

Windthrow or blowdown in a lowland forest of hackberry or western hackberry, sugarberry or southern hackberry, honey locust, chinquapin oak, and American or white elm provided an opportunity to study secondary plant succession at local scale through the phenomenon of forest gap or patch dynamics. Progression of revegetation, vegetation development by secondary succession, on a sere of climax lowland deciduous forest, SAF Cover Type 93 (Sugarberry-American Elm-Green Ash), in the western edge of the Ozark (Springfield) Plateau was preented in the following section of this chapter. Straight line winds of over 70 (probably closer to 80) miles per hour on Easter Sunday (15 April 2001) in conjuction with wet--though not saturated--soil resulted in uprooting of numerous mature trees of the species just listed. This blowdown occurred along the perimeter of a climax second-growth forest on slopes just above an intermittant stream. Deeper inside the forest there was minimal windthrow. This was a typical pattern resulting from the wind blockage affect of outermost trees on edges of the forest community.

A series of sequential photographs over a period of years following blowdown of trees on the perimeter of this lowland hackberry-sugarberry-elm forest documented early recovery of forest range vegetation in the cavities and immediately surrounding areas of denudation caused by high winds and wet soil conditions. Such forest gaps with large holes or craters left by uprooting of mature trees provided initial bare ground for the process of plant community development (forest revegetation by secondary plant succcession) at local scale, known as patch dynamics. Foresters affectionately (by standards of stoic woodsmen) refer to such soil cavities as "tree cradles" due to the frequent germination and subsequent establishment of trees in the hole and/or surrounding area of tree fall. Cavities thus created provide bare soil and more mesic land (edaphic-microtopographic) habitat for tree seedlings or root suckers.

As the following photographs showed development of vegetation (secondary plant succession) pm disturbance microsites caused by uprooting and related creation of bare soil (wind as "Nature's bulldozer") is not simply a matter of tree seedling pioneering the denuded microhabitat. Yes, this is sometimes the case as for example when seedlings of sycamore, eastern cottonwood, willow, sweetgum, and other hardwoods sprout up almost immediately on newly cut stream bnks, sand bars, roadcuts, or clearcuts. Douglas-fir (Pseudotsuga menziesii) and coast redwood (Sequoia sempervirens) are examples of western conifers that typically have the same pioneering or early seral stage establishment following creation of bare soil environments by disturbance. Coast redwood also sends up sprouts (vegetative reproduction).

Classic studies of secondary succession on old fields or go-back land have shown unequivocally that such rapid establishment of late seral tree species (some persisting into climax) and/or climax tree species is not the typical pattern of vegetation development on the seres of many range and forest sites. The photographic study presented below showed the same phenomenon, the same pattern, of old field succession commencing with annual, pioneer ("weedy") species which were then followed in succession by short-lived, perennial, herbaceous species over course of several growing seasons before seedlings of tree species vegetated the "tree cradle". There was a "successional twist", however, as some root and stump sprouts from unearthed trees quickly emerged and grew alongside the herbaceous pioneers that sprouted from the soil seed bank and quickly populated "tree cradles" .

Local microsites of denudation caused by uprooted trees were designated as "tree clods" (for the mass of soil, rock, tree roots) and their "holes" or "craters" (cavities left in the land surface by removal of "tree clods") were followed over spans of time beginning with initial plant communities in the first growing season starting after tree uprooting. A single "tree clod" typically consisted of two or more mature trees that had been uprooted together as a unit. A few "tree clods" were made up of a single large tree whereas other "tree clods" and their "holes" were made up of as many as five trees that were uprooted en mass.

In this photographic treatment two examples of uprooted tree units ("tree clods") caused by the Easter Sunday blowdown were presented to show recovery of plants on these local spots or patches of disturbance. The first of these examples was of one large "tree clod" consisting of five individual mature tress. The second example included two somewhat smaller "tree clods" each of which was comprised of a single mature tree and adjacent immature trees (saplings and/or poles) These two examples of "tree clods" and their "root holes" was followed over a sequence of years to provide photographic documentation of secondary plant succession on (in and around) each denuded microsite (forest gap).

356. Broad view of windthrow- Uprooted trees (straight line winds of 70-80mph combined with wet soil) viewed as "tree clods" at outermost edge of a lowland forest of hackberry, sugarberry, hony locust, chinquapin oak, and American elm. A total of four "tree clods" and "craters" created by uplifted earth were visible in this wide-angle photograph. The first of these "tree clods" was caused by uprooting of a large chinquapin oak which broke off crowns of hackberry and two honey locust. The second (and largest) "clod" was created by uprooting of a large hackberry and very large black walnut which coame up together as a unit and which brought down a sugarberry, American elm and honey locust. The third "tree clod" consisted of a large honey locust, two young American elm, and two young (pole-size) hackberry or sugarberry (the two could not be distinguished without leaves). The fourth "tree clod" was comprised of a massive honey locust that brought down (uprooted with it) two smaller hackberry or sugarberry. "Tree clods" consisted of both: 1) direct victims or causalities (larger, mature trees initially uprooted through storm strain, combination of straight line winds and wet soil) and 2) indirect or secondary victims or causalities (smaller trees that survived storm stress of wind and wet soil but were brought down, and their roots partially up, by crashing larger trees uprooted by storm action).

The large trees that were toppled by direct results of storm strain (high winds and wet soil) were almost entirely uprooted or else the major roots had been broken in the crash. As such, the main shoot of all large trees (chinquapin oak, honey locust, black walnut, sugarberry, and hackberry) had died by late spring. Some of these large trees sent up root suckers so that the actual genetic tree did not die completely and instead persisted as clonal offspring (asexual or vegetative reproduction), but these were relatively weak and lacking in vigor. In subsequent growing seasons some of these sprouts died. Some of the smaller trees that were brought down by uprooted windthrown trees were not completely uprooted with the result that they leaved albeit in the horizontal plane.

The second and massive "clod" (directly uprooted large hackberry and even bigger black walnut with secondary causalities of Ameican elm, sugarberry or hackberry [whichever], and honey locust) served as one example described below. The third "clod" (directly uprooted honey locust and secondary causalities of elm and hackberry or sugarberry) and fourth "tree clod" (directly uprooted huge honey locust and two young or pole-sized hackberry as secondary causalities) were in close porximity and together served as the second example to be described.

Photographs were taken periodically over a span of years in either late June or early July to present late spring or early summer aspects of successional range vegetation. Given that denuded microhabitats were created in mid April, these photographs represented the number of growing seasons (mid-way through) since denudation.

Ottawa County, Oklahoma. June, 2001.

357. First growing season of first blowdown example- "Tree clod" and "crater", the second of four such forest gaps of "clods and craters" presented in the preceding photograph, caused when a very large black walnut and large hackberry were uprooted by straight line winds and wet soil. Plant species growing on this denuded microenvironment included such pioneers as giant ragweed (Ambrosia trifida) and hairy crabgrass (Digitaria sanguinalis) as well as surviving, long-established perennials such as silky wildrye (Elymus villosus). There was some mare'stail or horseweed (Conyza canadensis= Erigerion cnaadensis), but much less of this species than of giant ragweed (which was somewht surprising given that these two annual composites are frequently co-dominant in early years of old-field succession). There were also abundant root sprouts from the uprooted trees, but these were generally lacking in vigor so that most were dead by the second growing season. Most plants of poison ivy (Rhus toxicodendron= Toxicodendron radicans) and fox or frost grape (Vitis vulpina) survived blowdown and even benefitted from increased light.

Ottawa County, Oklahoma. June, 2001.

358. Fifth growing season of first blowdown example- Successional vegetation in fifth growing season on a "tree clod and crater" caused by uprooting of a very large black walnut and large hackberry. Species of pioneer annuals--including giant ragweed, marestail or horseweed, and hairy crabgrass--were still common and dominant in bottom and sides of craters as in first growing season following windthrow. There was abundant cover of the Eurasian annual grass, Japanese chess or brome (Bromus japonicus) which had many straw-colored, dead or dying shoots at time of photograph (July). This grass had not been a prominent component in the first warm-growing season following blowdown (15 April, 2001) because it is a cool season species. The most abundant grass was silky wildrye (several spikes visible in foreground of second slide), the native, cool-seaaon perennial decreaser which had been present in understorey of the climax forest and survived on "tree clods" even in the first growing season following uprooting of trees. Another relative common and abundant annual that had been present since the first cool-growing season following denudation was the naturalized Eurasian forb, hedge parsley (Torilis japonica).

Seedlings of American elm, hackberry, and sugarberry were now well-established. Sprouts of honey locust origining from roots of huge, uprooted trees were obviously larger. An example of one of these vigerous honey locust root sprouts was in lower right corner of first photograph. Note: it was very difficult to determine whether the young trees of hackberry, sugarberry, and elm were from seed or root sprouts, but based on plants in the first couple of growing seasons following blowdown it was determined that they were, in fact, seedlings and not resprouts off of roots of downed adult trees (ie. sexual not asexual reproduction had been responsible for regeneration of climax tree species. It was the opposite situation for honey locust in which regenerated trees were root sprouts of huge uprooted trees.

These climax tree species did not dominate the denuded site on basis of density, cover, biomass, etc., but they were becoming more abundant. The biennial (and native) tall thistle (Cirsium altissimum) had become locally common (eg. lower left foreground of second photograph). Also, perennial forbs had begun to occupy the denuded site. This included an unidentified (not yet flowering) goldenrod (Solidago sp.) like the plant in the right corner of the second photograph. A common (and perennial) forb was climbing false buckwheat (Polygonum scandens). This species was conspicuous in the foregroung of both photographs, especially the second.

Ottawa County, Oklahoma. July, 2005.

359. Eighth growing season of first blowdown example- Recovering vegetation in eighth growing season on a "tree clod and crater" caused by uprooting of a very large black walnut and large hackberry. Almost all foliar cover was that from seedlings of the climax tree species, the adults of which had been uprooted eight growing seasons ago. The two dominant species of seedlings were hackberry and sugarberry (with leaves clearly distinguishable and distinct). Seedlings of American elm were smaller and less common making this the associate species. In spite of vigerous root sprouts (resprouts) early on in the denuded areas, honey locust had become a distant "runner-up" among the climax tree species.

Almost all forbs were gone from the sere at this stage of secondary plant succession. There were some scattered individuals of wingstem (Verbesina helianthoides). There were also individuals of silky wildrye which had survived windthrow. These were being overshadowed by seedlings and sprouts of climax tree species, but wildrye plants survived because they are a cool-season perennial species and make much of their growth during winter and early spring when hardwood tree species are bare (devoid of leaves).

Ottawa County, Oklahoma. July, 2008.

360. Detail in eighth growing season of first blowdown example- Successional stage of developing range vegetation on a denuded microsite, a "tree clod and crater", caused by uprooting of a very large black walnut and large hackberry. This was appearance and composition of the successional plant community about midway through the eighth warm-growing season following blowdown. Almost all plant cover was that of seedlings and sprouts of the climax tree species of which hackberry and sugarberry were the dominants (followed by American elm and, as in distant fourth-place, honey locust).

Note that the species composition--not structure or physiogonomy--was now that of the original climax hackberry-sugarberry-elm-honey locust lowland forest. In only eight years following denudation by windthrow, and old-field succession that began with weedy annuals like giant ragweed, marestail or horseweed, and hairy crabgrass, the original species composition of the forest had been restored. The rest of time taken for recovery of this natural forest range vegstoration on the sere will be maturing of established individual trees of the climax dominant species.

In descriptive vegetational analysis (at least to this point on the revegetating sere), succession of patch dynamics created by tree blowdown was amazingly rapid with restoration of dominance by climax tree species completed by the large seeding stage (age class) of tree growth, and in eight years following initial denudation.

Ottawa County, Oklahoma. July, 2008.

361. First growing season of second blowdown example- Appearance of two "tree clods" and "craters", the third and fourth of four such forest gaps of "clods and craters" that were presented in the introductory wide-angle photograph given above, about midway through the first warm-growing season after being caused by tree blowdown. The third "clod and crater" consisted of a large honey locust that was uprooted and took with it as it crashed two young American elm and two young (pole-size) hackberry or sugarberry (these two species could not be distinguished without leaves). The fourth 'clod and crater" was formed when an uprooted huge honey locust crashed taking with it to earth two young or pole-sized hackberry.

The most conspicuous species to show benefit from denudation was giant ragweed and, later in the summer, hairy crabgrass. There was some horseweed or marestail, but much less than giant ragweed. As was the case for the first "tree clod and crater" example, liana species like fox grape and poison ivy survived and beneffited (at least initially) from increased sunlight. There were root sprouts from the uprooted, large, adult trees. These were generally weak and lacked vigor typical of sprouts (especially, it seems, those that are brush) except for those of honey locust.

Ottawa County, Oklahoma. June 2001.

362. First growing season (close-in view) of second blowdown example- Detailed view of the "tree clod" on the :right as shown in the two immediately preceding photographs. These two "clods" and their "craters" served as the second example of old-field (secondary plant) succession on a local denuded microsite caused by windthrow (straight line winds up to 80mph and wet soil). This "tree clod" was caused by uprooting of a huge honey locust, the crashing of which caused uprooting (secondary impact) of two pole-sized hackberry. Many plants of giiant ragweed were the immediate successional "beneficiaries" of this devestation and denudation that left bare soil and a hole to temporarily pond and provide supplemental water to "lucky" plants. Giant ragweed grew so thick in this colony of seedlings that even hairy crabgrass was largely excluded.

Existing shoots of uprooted trees, including the huge honey locust and two hackberry, were killed. These trees temporarily leafed out at the appropriate time in spring following toppling (which was on 15 April, 2001), but leaves soon died and fell off before achieving any appreciable size. The big honey locust did send up vigerous and persistent root sprouts. Honey locust was the only tree species to successfully reproduce asexually (vegetatively).

Ottawa County, Oklahoma. June 2001.

363. Fifth growing season of second blowdown example- Appearance of recovering (successional) range vegetation about mid-way through the fifth growing season on two "tree clods" and their "craters" caused by uprooting of a one large to huge honey locust which on crashing ripped up or broke off smaller (sapling to pole-size) American elm and hackberry. These were the third and fourth "tree clods" of four such uprooted clumps of trees. They were the same "clods" shown above at the first growing season after uprooting. (More thorough descriptions of tree species that made up "tree clods" were provided above when "clods" described for the first growing season following blowdown.)

At this stage of secondary succession on denuded areas seedlings and root sprouts of the climax trees were well-established. Most of these new shoots of American elm, hackberry, and sugarberry were seedlings, the initial resprouts off of roots of mature trees having died by end of second warm-growing season. Sprouts of the immense trees of honey locust were vigerous and rapidly growing. Fox or frost grape was the major woody vine growing in association with seedlings and sprouts. Fox grape was presented in greater detail in photographs below.

Also present as robust and vigerous plants were pioneer annuals both cool- and warm-season species. In this fifth spring-summer period following denudation the dominant (and native), annual pioneer species remained giant ragweed (the same as in the first growing season following blowdown) with hairy crabgrass also common. Perennial herbaceous species included pokeweed (Phytolacca americana) which was the largest, most rank-growing, and dominant of this category of plant. Pokesalad was obvious in interior of denuded spaces in both of these photographs. Poke was featured in close-in, detailed slides shown and described below. Another herbaceous perennial (represented by scattered individuals) was goldenrod, the plants of which were not blooming and thus could not be identified as to species. Climbing false buckwheat was another perennial forb and olne that was abundant, especially around perimeter and near "craters". Another abaundant, perennial forb was giant yellow hyssop (Agastache nepetoides) which was presented as scattered individuals in the second of these two photographs. Also still present at this stage was Japanese chess or Japanese brome (a Eurasian, cool-season, annual grass which was visible as brown- or amber-colored straw in foreground of second slide) along with silky wildrye which was the major native grass (a cool-season perennial and climax decreaser species).Another Eurasian annual that was common on the more-or-less still bare patches of soil was hedge parsley. This species had been common since the first cool-growing season following uprooting of trees.

Observation: much ado is often about biodiversity, especially species diversity, in regards to plant succession and stages of vegetation development. Some ecologists have argued that plant diversity is greater in this or that stage, often arguing in favor of climax or late (advanced) seral stages. On tree-lowdown denuded spaces (forest gaps and, thus, patch dynamics) in climax hackberry-sugarberry-elm-honey locust lowland forest in the western Ozark Plateau, biodiversity as viewed by species number and plant forms was greatest--far and away so--in mid-sere, and clearly not at stage(s) in which plant species composition is typical of climax. More will be shown of this successional fact in photographs and captions taken of denuded spaces in the eighth warm-growing season following denudation (see below).

Ottawa County, Oklahoma. July, 2005.

364. Fifth growing season (close-in view) of second blowdown example- More detailed perspective of successional, forest range vegetation on a "tree clod" and adjoining "tree crater" created when huge honey locust trees were uprooted by high straight line winds (70-80mph) eight warm-growing seasons ago (prior to time of these photographs). The denuded microhabitat (forest gap) was in a climax hackberry-sugarberry-American elm-honey locust lowland forest in the Springfield Plateau (western partion of Ozark Plateau Region). This was the "tree clod" on the right (see above photographs) in second blowdown example being used in this section (see again first warm-growing season after tree blowdown). This was the fourth of four uprooted tree clumps described herein. It was created with the uprooting of a huge honey locust, the unearthing of which tore out (ripped out) as a secondary impact two pole-sized hackberry.

The first of these two photographs showed climbing false buckwheat growing on face (bottomside) of "tree clod". Pokeweed, the dominant herbaceous species growing on the "clod-crater" complex, was also featured prominently in this slide. Giant ragweed, the dominant annual pioneer species that was still abundant five years after denudation, was conspicuous in upper right margin. A spindly plant of marestail or horseweed (slightly right-of-center in foreground) represented another native, annual pioneer of the Compositae. A tall (almost 10 feet in height) root sprout of honey locust that had been present since the first growing season following blowdown (spring, summer, autumn after 15 April, 2001) of the large honey locust was in upper left portion of this slide. Several large seedlings or small saplings (all apparently from seed) of hackberry, sugarberry, and American elm were also present, especially in background, as were shoots of fox grape.

The second of these photographs was of the "crater", the hole created when soil and other earth materials were lifted out by the uprooted trees. The large, robust shoots of pokeweed that dominated (overwhelmed was more like it) much of the denuded area were featured in this slide. Also visible in this slide was a vigerous plant of silky wildrye (the consist dominant grass of blowdown-denuded habitats since their creation), Japanese chess or Japanese brome, marestail or horseweed, giant ragweed, poison ivy, and gray-headed coneflower (Ratibida pinnata), perennial composite. Pokeweed, a large perennial with an immense and fleshy taproot, is capable of living for years. Pokeberry is readily "shaded-out" by growing trees. The prime habitat for pokeweed is disturbed ground a few years after initial creation of the denuded spot. Ash from burning large accumulations of wood, bulldozed piles of brush, cleared road sides, and fencerows are ideal environments for pokeweed, especially if there are perches for birds that recently feasted on pokeberry fruit.

Ottawa County, Oklahoma. July, 2005.

365. Eighth growing season of second blowdown example- Late seral (advanced successional stage) forest range vegetation roughly mid-way into the eighth growing season on a "tree clod and crater" caused by uprooting and cavity creation by straight line winds (70-80mph) and wet soil in early spring. Local sites of denudation (both of these "clod-crater" microsites) were caused when a single massive honey locust was uprooted and, in secondary consequences of the crashing honey locust, smaller hackberry and/or sugarberry and American elm were ripped up.

Development of forest vegetation had achieved species composition of the climax hackberry-sugarberry-American elm-honey locust community. The plant community on "tree clods and craters" was dominated overwhelmingly at this stage by small saplings of hackberry, sugarberry, American elm, and as feaured in these two slides, black walnut. Almost all of the young trees of these species were from seed and not root suckers (ie. seedlings and not sprouts of uprooted trees) Young trees of honey locust were, in contrast, primarily root sprouts off of severed roots that remained in the soil when adult shoots (combination of trunks nad crowns) of large honey locust were uprooted. These honey locust saplings of root origin had been abundant and conspicuous by the fourth and fifth spring-summer growing seasons (see slides and captions above, especially the first example of blowdown). At this stage, eighth tree-growing season following windthrow, Celtis and Ulmus species had grown so thick that their larger leaves "blocked the view" of honey locust with their smaller, finner leaves. On these two denuded microsites black walnut seedlings and saplings had benefitted especially from the "craters" that temporarily ponded extra water.

At this stage of secondary plant sucession the terminal (climax) forest community had been reached from standpoint of dominant species composition, but not from criteria of physiogonomy, architecture (structure), physical arrangement, or, perhaps, total number of climax species. No, ecosystem structure and function had not been restored. That point of ecosystem restoration would require presence of mature (adult-size or fully grown) trees of the dominant species along with species associated with them as, for example, such lianas as grape, poison ivy, and trumpet creeper.

Gone from this plant community with its species composition comprised almost entirely of small trees of the climax dominant tree species were almost all annuals including giant ragweed, the dominant pioneer and later seral stage herbaceous species, as well as perennials of advanced stages like pokeweed. At edge or outer perimeter of this "tree clod and crater" there was one locally common, annual weed which wass shown in details of close-in views presented in the nest two slides immediately below.

Ottawa County, Oklahoma. July, 2008.

366. Eighth growing season (close-in view) of second blowdown example- Detailed views of the second (rightmost) "tree clod and crater" of the two "clods" that served as the second example of secondary plant succession on local sites (microsites) of denudation caused by uprooted trees resulting from high straight line winds (70-80mph) and wet soil eight growing seasons ago. This "clod and crater" resulted from uprooting of a huge honey locust and two pole-sized hackberry.

At this stage of forest recovery (vegetation development by old-field succesion on a lowland forest sere) the denuded site was almost completely populated by hackberry and sugarberry with some American elm and honey locust as associate species. Annuals, biennials, and herbaceous perennials had been "overtopped and overrun" by the climax tree species. For example, the large plants of pokeweed present from about the third to sixth year of secondary succession (recall from above) had either died out or survived as depauperate specimens (presumedly due to competition for light, growing space, soil water, or whatever from young saplings of climax dominants).

In other words, species composition was restored to that of the climax hackberry-sugarberry-American elm-honey locust forest. Physiogonomy, structure, and function of the climax forest was far from restored. These features of the climax stage of forest range vegetation would not be restored, the forest would not be fully recovered (ie. the fully functional climax forest ecosystem will not be achieved or reacquired) until these large seedlings and small saplings reach adulthood and mature size. On basis of species makeup as determined by relative cover or biomass the botanical composition of climax had been reached. Decades would be required for complete recovery of the forest ecosystem and full development of forest vegetation.

The phenomenon shown in this section devoted to forest gaps and patch dynamics was that the time span (temporal aspect or element) of secondary plant (old-field) succession from initial denudation back to the climax stage based on botanical composition of the plant community, the vegertation, was remarkably (indeed, almost miraculously) short or brief. Number of growing seasons was very few (only seven or eight) in the progression of vegetation development from bare soil to dominance of the disturbed site by climax tree dominants. Development of vegetation, progression along the forest sere, from denudation to termination of successional stages (advance of seral stages culminating in the climax community based on plant species) is rapid on the small spaces laid bare by uprooted individual large trees and/or small numbers of affilitated smaller trees. Secondary succession is rapid on the small microhabitats that are characteristic of forest gaps. Patch dynamics--secondary plant succession on small gaps (micro-spatial scale) created by local disturbances (tree blowdown in these examples)--is considerably faster than on larger areas such as old (abanodoned) farm fields or cutover foresst land. There is an interaction of temporal and spatial affects: temporal aspects operate (proceed) faster at smaller spatial scale (ie. vegetation development by secondary succession takes place more rapidly when areas of disturbance are smaller). Recovery of late seral and climax vegertation depends on size of disturbance (how much land, the area, was denuded) as well as time and plant-growing conditions related to time.

Ottawa County, Oklahoma. July, 2008.

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