Southern and Central Forests - IB

Forest Range Types of Eastern North America

Willow Oak-Water Oak Type in a Mesic Hardwood Bottomland Forest

A second-growth though a seemingly climax-composition had developed on land of slightly higher elevation in a mixed hardwood bottomland forest on the old stream bed (a former meander) of Sabine River in northeast Texas. This was part of the western Pineywoods that a vegetational subdivision of the once-vast Eastern Deciduous Forest Region of eastern North America. In the section immediately below, an example of the Willow Oak-Water Oak-Diamondleaf (Laurel) Oak cover type (number 88) recognized by the Society of American Foresters (Eyre, 1980) was presented.

182. Still yet, more variation: another local forest community- This second-growth forest stand was one of many local communities comprised of various hardwood species that formed a mixed hardwood forest range that had developed on an old meander of the past stream channel of Sabine River in northeast Texas. This vegetation was mostly a group of willow oaks, hence the Willow Oak-Water Oak-Diamondleaf Oak forest cover type (see below).. Tree species represented here included five willow oaks beginning on the left, a green ash in center foreground with a Virginia creeper climbing it, two sweetgums to right of the green ash, and at farthest right (slightly behind) another willow oak.

Much of the green ground cover in foreground was peppervine, but there was also cover of trumpet creeper and broadleaf woodoats.

Old Sabine Bottom Wildlife Management Area, Smith County, Texas. Late June; early estival aspect. FRES 15 (Oak-Hickory Forest Ecosystem. K-91 (Oak-Hickory Forest). Forest cover type as described by the Society of American Foresters (Eyre, 1980) was Willow Oak-Water Oak-Diamondleaf oak (88). So varied that that there was not a "one-for-all" appropriate biotic community unit in Brown et al. (1998), much of this vegetation fit Mixed Hardwood Series 223.13 of Southeastern Swamp and Riparian Forests 223.1 of Warm Temperate Swamp and Riparian Forests 223 (Brown et al., 1998 p. 43). South Central Plains- Floodplains and Low Terraces Ecoregion 35b (Griffith et al., 2004).

183. Picturesque forest assemblage- Deep inside a second-growth, mixed hardwood bottomland forest that developed on an old meander of the upper Sabine River there was this forest stand co-dominated by water oak and willow (willowleaf) oak (sweetgum was the associate tree species) with a multi-layered understorey. Giant cane (native bamboo that in this forest was a semi-woody plant) formed an erratic or interrupted upper herbaceous layer, but there was a limited lower herbaceous layer. Instead there was a lower woody (shrub layer) comprised largely of liana species which also extended to the upper shrub layer when some of these woody vines climbed trees as, for example, Alabama supplejack on the trunk of the willow oak in the right foreground of the second slide. There was a fairly high density and shrub canopy of American hornbeam or blue beech in this forest stand.

The featured plant species in these two slides was dwarf palmetto (Sabal minor) that occupied the left foreground of the first slide and the center of the second slide. Together these two slides made up a "nested photo-plot" with the second photograph being a "sub-plot" of the first photograph or overall "photo-plot".

Sabine Bottom Wildlife Management Area, Smith County, Texas. Late June; early estival aspect. FRES 15 (Oak-Hickory Forest Ecosystem. K-91 (Oak-Hickory Forest). Forest cover type as described by the Society of American Foresters (Eyre, 1980) was Willow Oak-Water Oak-Diamondleaf oak (88). So varied that that there was not a "one-for-all" appropriate biotic community unit in Brown et al. (1998), much of this vegetation fit Mixed Hardwood Series 223.13 of Southeastern Swamp and Riparian Forests 223.1 of Warm Temperate Swamp and Riparian Forests 223 (Brown et al., 1998 p. 43). South Central Plains- Floodplains and Low Terraces Ecoregion 35b (Griffith et al., 2004).

184. Near ultimate Pineywoods forest and range- Structure and composition of a second-growth mixed hardwood bottomland forest in northeast Texas on an old meander of the upper Sabine River. A remarkable array of tree species was present in the first slide, which served as a general "photo-quadrant", while the second slide, which served as a "photographic sub-plot", featured tree species found on the right side of the first slide. In the first slide, the largest tree (left-most tree) was water oak playing host to a commensal poison ivy vine, willow oak was the right-most tree as were most of the trees in the middle with darker-colored trunks, swamp post oak (Quercus similis) was the second tree from the right, other lighter-colored trunks in the middle of the line-up were overcup oaks, the larger sapling in midground (immediately to left of the swamp post oak) was a water oak and the smaller, sapling-sized plants were eastern hop-hornbeam. There was a large seedling of sugarberry in front of the overcup oak in center midground of this first slide and--shown to better advantage--in left foreground of the second slide.

Forest vegetation in the second (vertical) slide was a "photographic sub-plot" of slightly less than the right half of the first slide or "photo-plot". These trees left to right were: overcup oak, water oak, swamp post oak (largest tree with knots on trunk), and willow oak. As noted in the preceding paragraph there was a large seedling of sugarberry in front of the overcup oak, left-most tree in left foreground.

Most of the understorey herbage (herbaceous layer) was that of broadleaf woodoats but there was some cover of immature (unidentifiable) caric-seddge. Besides tree-climbing poison ivy, trumpet creeper and roundleaf green-brier were common liana species.

185. Textbook example of Pineywoods range- A local herbage-rich community in a second-growth, mixed hardwood bottomland forest in the Pineywoods of northeast Texas. This forest had developed on an old meander of the Sabine River. Forest vegettion shown here was a local stand of willow oak with some water oak and a sapling (light trunk in dead-center) of green ash and a lush understorey of some species of caric sedge (Carex sp.), which this author could not identify at its pre-bloom stage, plus the ever-present broadleaf woodoats. The main shrub species were American hornbeam or blue beech and highbush or squaw huckleberry (Vaccinum Stamineum).

The extremely water-favorable soil environment made for a highly productive herbaceous understorey. Unfortunately for some uses, such floodplain habitats have their costs under certain conditions. The year after the slides in this section were taken, severe flooding drowned so many of the white-teailed deer fawns that there was no deer hunting permitted on this wildlife management area.

186. Local stand and a "mixed message" succession-wise- Local stand of forest vegetation that was part of a diverse, second-growth, mixed hardwood btottomland forest on the old stream channel of Sabine River in northeast Texas. Three slides at progressively shorter camera distance served as "double-sub-plotted photo-quadrant" of a local stand of willow oak with multiple vegetational layers including a prominent layer of shrubs and sapling trees and an herbaceous layer that was fairly consistent in its understorey coverage. The principal herbaceous species in this forest stand was Virginia wildrye (Elymus virginicus) although broadleaf woodoats was ever-present throughout this forest. There was also coverage of Panicum, Paspalum, and Carex species. These pre-bloom/pre-fruit graminoid species could not be identified based on vegetative features by your newly arrived author.

The lesson in forest succession provided by this set of three slides was built around presence of green ash (seen most conspicuously in center foreground of the second slide and in right foreground of the third slide). Green ash had unquestionably established after the willow oaks had made consoderable growth. Green ash had obviously germinated, emerged, and continued to survive in (under) the shade of willow oak. Whether green ash would come in successional time to replace the willow oaks or merely to persist along with them into the climax forest (or perhaps for both to remain only into some subclimax stage) remained to be seen. This could only be seen at a point in time beyond the temporal scale of this publication's continued growth and development.

In their nearly comprehensive coverage of silvics of North American tree species Burns and Honkala (1990) described willow oak as a subclimax species that was generally ranked as intolerant (intolerant of shade/competition). Green ash was inconsistent in its general tolerance (shade/competition) which ranged from intolerant to moderately tolerant (Burns and Honkala, 1990). Viewed from perspective of tolerance--which was not to rule out other variables (including chance) as being important--presence of green ash along with willow oak was a plausable outcome in this second-growth forest, the climax of which was not know with certainity.

Technical note and experienced advice: the first of these three slides was botched up by an Epson Perfection 700 scanner. Epson Far From Perfection over-exposed this slide. Even Adobe PhotoShop with "all its king's horses and all its king's men" could not restore a slide whose image at least approached perfection. Never invest in Epson products.

187. Dominant of moist woods- Trunk and lower branch with leaves of willow (sometimes, called willowleaf) oak (Quercus phellos) in a second-growth, mixed hardwood forest on an old meander of Sabine River in northeast Texas.

Willow oak is a member of the white oak subgenus, Leucobalanus.

Old Sabine Bottom Wildlife Management Area, Smith County, Texas. Late June.

188. Willowed oak leaves- Leaves of willow oak on the tree and branch introduced immediately above. In a second-growth, mixed hardwood bottomland forest on the old stream channel of the Sabine River in northeast Texas.

Old Sabine Bottom Wildlife Management Area, Smith County, Texas. Late June.

189. Bark dwellers on a branch in the bottomland- Two species of lichen growing on a dead branch that fell of of the willow oak featured in the two immediately preceding slide/caption sets. The flat, spreading lichen was a foliose lichen perhaps of the species known as common greenshield lichen (Flavoparmelia caperata). The fruticose lichen was a beard lichen of species Usnea endochrysea known by the common name of bushy beard lichen.

Old Sabine Bottom Wildlife Management Area, Smith County, Texas. Late June.

190. Beards on a branch in the bottom- Closer-in views of bushy beard lichen (Usnea endochrysea) growing on a dead branch of willow oak tht fell from the tree introduced above which was used as an example of willow oak. In the second slide there was also part of a foliose lichen, most likely common greenshield lichen (Flavoparmelia caperata).

These ectophytes (plant species that grow on the outside of other plants) were commensal species that grew on both live and dead willow oak. Commensalism is a symbiotic relationship in which one species (lichen in this case) lives off of another species, the host (willow oak in this instance), this latter of which gets nothing--good or bad--out of the relationship. f

This lesson only ends with commensalism. The more basic (if that be the most descriptive term) story of this living together (= symbiosis or symbiotic relationship) was the lichen itself. Lichen is actually two species: a fungus and an algae. These two diverse groups of plants grow in such intimate relationship that they literally become "one flesh" and are recognized as a species (one "fused" species actually made of two species).

The story then continues to herbivory as animals consume the lichen. In other words, lichen are the beginning of their own food chain.

Life on the range is remarkable. For that matter, all life anywhere is remarkable.

Old Sabine Bottom Wildlife Management Area, Smith County, Texas. Late June.

191. Another mixed message on forest succession- Local stand of vegetation in a second-growth mixed hardwood bottomland forest on an old meander of the upper Sabine River in northeast Texas. The "photo-dendrogram" of the forest community "sampled" here included young (still rapidly gorwing) yet sexually mature water and willow oaks showing for example a water oak (left margin) and two willow oaks (right midground and far-right background). Shrubs included poison ivy, which was "growing everywhere", and eastern hop-hormbeam (shorter shrubs in center midground). Broadleaf woodoats dominated the herbaceous understorey which formed a nearly complete layer in this forest range community. There were some Panicum and Paspalum species as well. Giant cane, the dominant grass over much of this forest, was not present in the local "sample" of forest plant community seen here.

The "real story" in this "photo-plot", however, was represented by a sapling of green ash (right midground). Willow and water oaks have generally been regarded as constituting climax or subclimax species in forests that form on alluvial sites. By contrast, green ash has been generally been regarded as a seral species that follows such pioneer species as cottonwood and willow, although green ash does sometimes persist into the climax forest as an "earlier entrant". In the definitive silvics treatment of North American trees, Burns and Honkala (1990) regarded willow oak and water oak as subclimax species and ranked them as intolerant of shade/competition. Burns and Honkala (1990) stated that green ash was inconsistent in its general tolerance (shade/competition) which "varies from intolerant to moderately tolerant to shade" though much of such work was based on more northern habitats. Regeneration of green ash in this bottomland forest currently dominated by willow oak and water oak presented something of a conflicting phenomenon, but based on shade/compoetition tolerance of these species it appeared that green ash was persisting along side the dominant subclimax oak species. This was consistent with large green ash that were approaching old-growth status on lower and more mesic habitats (see above, this section).

This local stand was an example of the Willow Oak-Water Oak-Diamondleaf (Laurel) Oak cover type (type number 88) recognized by the Society of American Foresters (Eyre, 1980). In describing this forest cover type, Shropshire in Eyre (1980, p, 63) concluded: "It probably represents a topographic/edaphic climax, but when it is heavily cut, species such as sugarberry, green ash, American elm, and red maple may capture the site, at least temporarily". This large tract of mixed hardwood bottomland forest (at least some parts of it) had been logged approximately a half century prior to time of these photographs.

Taken in total, presence of green ash alongside subclimax willow oak and water oak plus the tolerant shrub, eastrern hop-hornbeam (Wegner, 1984, p. 3) was not overly surprising.

192. Changes to be- A young adult (sexually mature) overcup oak (left) that was representative of one of the dominant to associate species of a second-growth mixed hardwood bottomand forest range appeared to be in the successional process of being replaced (succeeded) by sugarberry as represented by the sapling immediately to right of the overcup oak.

Overcup oak has a tolerance (general or shade/competition) rating of intermediate (Burns and Honkala, 1990) whereas Fowells (1965) rated sugarberry as tolerant of shade. On the other hand, for flooding tolerance Midleton (1999, p. 146) showed sugarberry as being Wealky Tolerant whereas overcup oak was rated as Highly Tolerant. On the somewhat higher ground (hence, drier or less wet soil) of the local habitat seen here, tolerance to flooding (= water tolerance) was less relevant or determinative than general (shade/competition) tolerance. Sugrberry is tolerant whereas overcup oak is only intermediate. It follows that in the grand scheme of forest succession --and the always specified, "other things being equal"-- sugarberry will replace overcup oak. Obviously other things are not always "being equal" hence factors such as, say, fire could "deflect" the normal path of succession (might modify the typical "trajectory" of community development on this forest sere; the ultimate or terminal vegetational state might be a pyric climax or an edaphic climax rather than a climatic climax).

Successional status of water oak and willow oak as subclimax species and co-existence (and regeneration) of green ash beside these oaks was treated in the immediately preceding slide/caption set. It was even more to be expected that the shade-tolerant sugarberry might replace its neighboring oak species (even the intermediately tolerant overcup oak).

Broadleaf woodoats dominated the herbaceous layer of this second-growth, mixed hardwood bottomland forest that was on the old stream channel of the upper Sabine River in northeast Texas. There were also some Panicum and Paspalum species as well as unidentified Carex species. The major shrub was American hornbeam or blue beech.

The habitat on which this local "photo-sample" of this second-growth, mixed hardwood bottomland forest developed was more mesic than that of the "photo-sample" of this same forest tract seen in the immediately preceding slide/caption set.

Old Sabine Bottom Wildlife Management Area, Smith County, Texas. Late June; early estival aspect. FRES 15 (Oak-Hickory Forest Ecosystem. K-91 (Oak-Hickory Forest). No one specific or single forest cover type described by the Society of American Foresters (Eyre, 1980). So varied that that there was not a "one-for-all" appropriate biotic community unit in Brown et al. (1998), much of this vegetation fit Mixed Hardwood Series 223.13 of Southeastern Swamp and Riparian Forests 223.1 of Warm Temperate Swamp and Riparian Forests 223 (Brown et al., 1998 p. 43). South Central Plains- Floodplains and Low Terraces Ecoregion 35b (Griffith et al., 2004).

Ending interesting note: that the mixed hardwood bottomland forest on the old channel of Sabine River is in a floodplain was demonstrated in graphic form the year after the above slides were taken. Spring rains were so heavy that this entire forest tract was flooded and under water for so long that all the white-tailed deer fawns drowned. There was no fawn crop in that year and the Old Sabine Wildlife Mangement Area was closed to deer hunting. Sometimes Mother Nature's rules seem harsh. The abiotic factor that is responsible for the great productivity--indeed the very life--of floodplain forests is the same abiotic factor that can bring death to these bottomland forest ecosystems. Students must always remember that death--the final end of all biotic life, the completion of the life cycle--is a part of the life of such forest ranges.

Location note: Presented in the chapter entitled Loblolly Pine Forest was a upland tract of loblolly pine (Pinus taeda) forest that developed on hillsides immediately above (contiguous with) the mixed hardwood bottomland forest on the old stream channel of Sabine River. This tract of second-growth loblolly pine was also part of Old Sabine Bottom Wildlife Management Area, Smith County, Texas.

Shortleaf Pine (Pinus echinata) Forests

Shortleaf pine, which is generally second only to loblolly pine (longleaf pine have been eliminated by man as a dominant species from much of its former range), occurs over much of upper southcentral North America (mid-South) especially on drier slopes and generally shallower, droughtier soils. Most of the pine and oak-pine forest of ancient mountains like the Ozarks, Ouchitas, and Kiamichis are those in which shortleaf pine is either a dominant or associate tree species. When Okies and Arkies speak of pineries they refer to shortleaf pine. Shortleaf pine is the State Tree of Arkansas.

193. Trunk of a mature shortleaf pine (Pinus echinata)- Red River County, Texas. July.

194. Branch of shortleaf pine- Shortleaf pine has the shortest needles of the major Pinus species in the Southern Pine Region. It was noted above that longer needles accumulate ice from storms more than short needles do. Shortleaf pine is therefore better adapted to more northern and western portions as in Arkansas and Oklahoma where ice storms not uncommonly inflict major damage to pines grown for shade and wood commodities.

Red River County, Texas. September.

194. Needless and cones of shortleaf pine- Relative length of leaves on shortleaf pine can be guaged by comparing the ones in this slide with those for longleaf, slash, or even ponderosa pines presented vriously in this publication. Cones of shortleaf pine are the smallest of the four major Pinus species often reaching only one and a half to two inches in length. Cones are frequently borne in a cluster of three at twig tips. Shortleaf pine is a prolific seed-producer beginning at relatively young ages. Fasicles bear two to three needles.

Red River County, Texas. September.

195. Shortleaf pine-oak-hickory forest- These two views are of a shortleaf pine-dominated forest with the physiogonomy and morphological form characteristic of old-growth (or approaching that stage). The author interpreted this state of forest vegetation development as, most likely, subclimax or perhaps even climax for this site. The crowns and general habit of the large pines had the appearance of very old trees. The understorey was of oaks (primarily the less mesic species like post, blackjack and black oak rather than the more mesic species like northern and southern red oaks). The hickory species could not be identified positively, but it was likely black hickory (Carya texana), a species adapted to drier sites and a local dominant in this area. Sweeetgum (Liquidambar styraciflua), one of the pioneer species on disturbed sites, was very common.

The oak, hickory, and sweetgum species appeared to have been suppressed by periodic fire. Perhaps this forest developed following a major fire and the fire-adapted shortleaf pine out-grew the oaks and hickories. The suppressed state of sweetgum would not be explained by the latter alternative.

There was regeneraation of shortleaf pine. It was an uneven-aged stand or population of P. echinata, but pine reproduction was less than that of the hardwood species.

Understorey shrubs included smooth sumac (Rhus glabra), trumpet creeper (Campsis radicans), Virginia creeper (Parthenocissus quinquefolia), redbud (Cercis canadensis), and poison oak (Toxicodendron radicans). There were two pronounced woody plant layers (two shrub understories). The herbaceous species comprised a single layer (excluding ground-surface species like mosses and lichens). This was the most species-diverse layer of vegetation. It included eastern gamagrass (Tripsacum dactyloides), longleaf wood oats (Uniola longispicata), beaked panicgrass (Panicum anceps), little bluestem (Andropogon scoparius), big bluestem (A. gerardii), and the composites, compassplant (Silphium integrifolium) and gayfeather (Liatris elegans).

Obviously this was an amazingly species-rich and structurally diverse plant community. It had for dominants climax species from both the tallgrass prairie and the oak-hickory-pine forest.

Ouachita National Forest, Scott County, Arkansas. July. Ouachita Mountains physiographic province. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Natural Vegetation Classification System of Arkansas for GAP Analysis Project, Natural Terrestrial Cover Unit 1.A.9.b.2 Evergreen Forest With Rounded Crowns Pinus echinata-Quercus marilandica-Quercus stellata. Oak-Pine Series of Brown et al. (1998). Ouachita Mountains- Fourche Mountains Ecoregion, 36d (Woods et al., 2004).

196. Shortleaf pine-oak-hickory forest near old-growth stage- Another view of the shortleaf pine-dominated forest with hardwood and herbaceous understories. Species were listed and the structure of this forest vegetation was discussed in the preceding caption. It will be emphasized again that the pine population was un-even aged, but recruitment of shortleaf pine was less than that of the oaks and hickories. The vegetation was obviously developing into a oak-hickory-shortleaf pine forest, but one with well-developed shrub and herbaceous layers composed of climax species.

The large old-growthlike shortleaf pine on the far right side of this photograph had numerous horizonally spreading limbs and bare spire-form of top which was typical of mature trees growing on savannas. This condition, taken in conjuction with the evidence of a tallgrass understorey (big bluestem, little bluestem, eastern gamagrass plus prairie composites), suggested that this vegetation might have been a shortleaf pine-tallgrass savanna. There were several other older individual pines in this and the preceding slides that had crowns made up of several major limbs which in turn branched diversely, a pattern or arrangement which indicated that these trees had developed in full light and not in a forest. This habit or morphological form indicated that these oldest trees lacked the apical dominance-development shape typical of trees growing up through an established forest. Some of these older trees had limbs below the terminal extension of the bole that were actually larger than this central apex. These trees had well-developed boles and would produce such fine logs as to self-start a Stihl, but the crown form and structure resembled that of wolf trees (trees whose big, spreading crowns occupy more canopy space than that of neighboring trees with larger, more valuable trunks).

Vegetation shown in these slides appeared to be a model example of the oak-hickory-shortleaf pine climax type at the subclimax stage which is most productive of high-quality conifer timber and forest range for wildlife and livestock.

Ouachita National Forest, Scott County, Arkansas. July. Ouachita Mountains physiographic province. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forests). SAF 76 (Shortleaf Pine Oak). Natural Vegetation Classification System of Arkansas for GAP Analysis Project, Natural Terrestrial Cover Unit 1A.9.b.2 Evergreen Forest With Rounded Crowns Pinus echinata-Quercus marilandica-Quercus stellata. Oak-Pine Series of Brown et al. (1998). Ouachita Mountains- Fourche Mountains Ecoregion, 36d (Woods et al., 2004).

197. Arkansawyer pineywoods- Second-growth shortleaf pine (in effect, a planttion) established as artificial regeneration after the native shortleaf pine--white oak-northern red oak-bitternut hickory forest had been logged. Artificial regeneration (also, artificial reproduction) refers to a stand of young trees established by human seeding or planting of seedlings or cuttings (Helms, 1998). In this forest tract in the Boston Mountains the U.S. Forest Service planted shortleaf pine and managed the forest for that single species with the result presented here: shortleaf pine plantation. The natural pine-oak forest was converted into a one-species forest (at least, more-or-less so; there was clearly a sole dominant tree species). The natural co-dominant oak and hickory component had been supressed by silvicultural practices so that only the natural co-dominant shortleaf pine pine held forest dominance. The hardwoods had reproduced and were growing well in this forest, but at this point (which was approaching harvest age-size for pine) oaks and bitternut hickory were not of size to share in dominance and, thus, in defining this forest community.

This tract of forest managed so as to have an unnaturally or atypical high cover and community dominance by one species, the economically most valuable species under current marketconditions, was an example of a forest plantation. The Society of American Foresters specified that a plantation was "a stand composed primarily of trees established by planting or artificial seeding" and with the specific notations that 1) plantations could have tree or understorey components from natural reproduction and 2) they could consist of single- or mixed spceies composition with either uniform or diverse structure, age classes, etc. (Helms, 1998). The example of shortleaf pine plantation shown here was a textbook example of those criteria.

In short, this was a man-modified forest. It was a forest with native species that had been manipulated to produce the forest crop species of choice while other species were managed for secondary considerations (eg. habitat of white-tail deer, a secondary crop). In this sense, this forest range community was transformed by management from a unit of potential natural vegetation (climax) into a politico-socio-economic unit of vegetation, an Arkansawyer forest.

Even though this was a forester's choice of forest, a forest modified by minds and hands of forestry professionals, at least one--a a glaring one at that--of these modifications was due to neglect or, perhaps more precisely, to human action that had "backfired". Exclusion of fire and/or, same thing with emphasis of inaction rather than action, lack of prescribed burning had allowed invasion of eastern red cedar (Juniperus virginiana).

The high relative canopy cover of sweet gum was typical of forests recovering from disturbances such as fire, blowdown, or, in this instance, timber harvest. Sweet gum is a seral tree species and one with a tolerance rating of Intolerant. It is, however, very effective in growth competitiveness on a wide range of soils and soil moisture conditions (Burns and , 1990). Sweet gum can readily come to dominate forests that were reestablished through artificial regeneration like planting as well as those that are redeveloping through secondary plant succession. This is simply part of the forest sere whether natural or aftificial reproduction. Sweet gum cover and density can be reduced substantially by grazing (even grazers like cattle are remarkably effective), herbicidal treatment, and prescribed burning. Such silvicultural practices may or may not be desirable and included as part of forest management.

Shrub species in this tract were listed in the next caption.

Ozark National Foraeest, Newton County, Arkansas. July, estival aspect. FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Pinus echinata Association (if recognized) in Oak-Pine Series of Brown et al (1998). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004).

198. Pine plantation in Boston Mountains- Interior of a shortleaf pine plantation showing composition and structure of a man-modified shortleaf pine-oak-hickory forest. Silvicultural practices had produced a forest in which one conifer species dominated the vegetation rather than the natural state in which three or four tree species (and both hardwoods and conifer) are dominants. This situation was explained in the immediately preceding caption.

Most of the understorey consisted of woody layers that were primarily different age classes of such climax dominants as white oak, northern red oak, black oak, bitternut or pignut hickory, mockernut hickory (Carya tomentosa) along with sweet gum, generally a pioneer tree species, and sassafras (Sassafras albidum), a large shrub or small tree species. Shrubs included flowering dogwood, eastern redbud, Carolina buckthorn (Rhamnus caroliniana), lowbush huckleberry (Vaccinum vacilians), poison oak, pison ivy, Virginian creeper, and blackberry (this latter probably of several species). Herbaceous species were generally absent (at least from a practical standpoint) which was undoubtedly due to the dense canopy produced by woody species.

This photograph included some of the larger trees of native hardwood dominants for comparison to the larger individuals of shortleaf pine. For example, the two northern red oak that appeared at this angle to be growing side-by-side in left foreground. The next succeeding slide showed large diameter of some of the pines.

199. Oaks, out; pines, in (or foresters' choice)- Interior of a shortleaf pine-oak forest, a plantation, produced by artificial regeneration. This forest had obviously been managed at (or near) time of planting pines resulting in their noticably greater size than almost all individuals of other tree species. This greater size (both height and diameter) was more than could be accounted for by growth. Sweet gum, the most widespread and abundant hardwood species, could easily have outgrown shortleaf pine on such sites as could stump sprouts from coppicing oaks and root-sprouting hickories.

Two of the conspicuous shrubs in foreground of this photograph were sassafras and Virginia creeper. Other shrub and tree species were listed in the preceding caption.

Viewers attention was directed to the large diameter of shortleaf pine (about time for logging) and in comparison to that of the much smaller hardwoods. A similar comparison was presented in the preceding slide.

Note on location of related forest range type: For purposes of comparison a white oak-northern red oak-bitternut hickory forest, that was more typical of forests in the Boston Mountains of the Ozark Plateau (closer to climax forest composition and structure), was located immediately opposite the shortleaf pine plantation just covered. These two tracts of forest were on the same forest site and both were parts of the Ozark National Forest. They were separated by the two-lane blacktopped state highway, Arkansas 21. That comparison tract was dealt with in the Oak-Hickory Forest (Part 1) chapter of Forest and Woodlands under the White Oak-Black Oak-Northern Red Oak Forest section.

200. Shortleaf pine (Pinus echinata)-oak pineywoods of eastern Oklahoma- Frequent fires have kept the oak-hickory-sweetgum component suppressed thus maintaining a nearly "pure pine type" in this second growth forest. (But note mostly hickory and oak sprouts in understory indicating recent fire suppression and succession toward the climatic climax of the region.) Herb layer is absent but flowering dogwood and redbud form an upper shrub layer while blackberry (Rubus spp.), gooseberry (Ribes spp.), and huckleberry (Vaccinium spp.) comprise a lower shrub layer. Old Military Road, Talimena State Park. LeFlore County, Oklahoma. July, estival aspect. FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem). Textbook example of K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Pinus echinata Association (if recognized) in Oak-Pine Series of Brown et al (1998). Ouachita Mountains-Western Ouachitas Ecoregion, 36e (Woods et al., 2005).

201. Shortleaf pine type with an open parklike understory dominated by big bluestem-Xeric south slope and a recent history of surface fires maintained this form of the white oak-shortleaf pine type as a pine-bluestem cmmunity. Benton County, Arkansas. June, early estival aspect. FRES No. 14 (Oak-Pine Forest Ecosystem) is the most accurate description of this type but FRES No.14 and FRES No.13 (Loblolly-Shortleaf Pine Ecosystem) both include the K-101 (Oak-Hickory-Pine Forest) unit. SAF 75 (Shortleaf Pine). Pinus echinata Association (if recognized) in Oak-Pine Series of Brown et al. (1998). Ozark Plateau- Dissected Springfield Plateau-Elk River Hills Ecoregion, 39b (Woods et al. 2004).

202. Virgin shortleaf pine-oak-hickory forest— One of the few remnants of old growth forest left in Texas is this shortleaf pine-white oak-chinkapin oak (Q. muhlenbergii)-shellbark hickory (Carya ovata)-pignut hickory (C. cordiformis) community seen here. There are several layers of vegetation including a second tree layer of young climax tree species and species like winged elm (Ulmus alata) and boisd'arc or Osage orange (Maclura pomifera) and a shrub layer of flowering dogwood, Arkansas traveler or pepperwood (Ampelopsisarborea), blackberry, gooseberry, and various wild grape vines. The prominent herb layer(s) include little bluestem, rosette panic grasses (Panicum spp.), slender- or longleaf wood oats, and scattered clumps of the native bamboo, giant cane (Arundinaria gigantea).

Lennox Woods (donated by Kirby Lumber Company to The Nature Conservancy), Red River County, Texas. May, vernal aspect. FRES No. 14 (Oak-Pine Ecosystem).FRES No.14 (Oak-Pine Forest Ecosystem). Classic example of K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Oak-Pine Series of Brown et al. (1998). South Central Plains- Tertiary Uplands Ecoregion, 35a (Griffith et al., 2004).

Following presentation of a virgin shortleaf pine-oak-hickory forest at old-growth state immediately above, a second-growth shortleaf pine-mixed hardwood forest in the central portion of Texas Pineywoods was presented in the following section. This forest rangecommunity was on the Davy Crockett National Forest where it was being managed as part of the restocking program of eastern wild turkey (Meleagris gallopavo silvestris) in the Pineywoods. Such forest range vegetation was an outstanding example of second-growth forests described by the Society of American Foresters (Eyre, 1980, ps. 7, 60) as the Shortleaf Pine-Oak forest cover type (SRM 76).

203. Physiogonomy and external architecture- Overall views of a second-growth, uplnd shortleaf pine-mixed hardwood stand from surface of forest floor to near top of forest canopy. Dominant woody vines in this particular local stand were Small's, lanceleaf, or coral greenbriar (Smilax smallii) and an unidentified grape (Vitis sp.?). Smooth sumac (Rhus glabra), American beautyberry (Callicarpa americana), and yaupon. Common grasses were longleaf woodoats, the dominant herbaceous species overall, purpletop (Tridens flavus), and beaked panicgrass (Panicum anceps). Forbs were extremely scarce so as to be of no notable relevance except to record presence of bracken fern (Pteridium aquilinum).

Davy Crockett National Forest, Houston County, Texas. October. FRES No. 14 (Oak-Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Pinus echinata Association (if recognized) in a Oak-Pine Series of Brown et al. (1998, p. 37) or, alternatively, Warm Temperate Forest and Woodland (123), Southeastern Deciduous and Evergreen Forest (123.1), Pine Series (123.12) of Brown et al. (1998, p. 38). South Central Plains-Southern Tertiary Uplands Ecoregion 35e (Griffith et al., 2004).

204. Internal structure and species composition- Two " photoplots" at progressively closer distance to range vegetation of a second-growth, upland shortleaf pine-mixed hardwood forest. The same shortleaf pine of relatively large size was in foreground of both slides. Dominant shrub overall as based on cover, density, and frequency was American beautyberry. Yaupon was a distant second based on these criteria. There were two shrub layers (upper and lower) in the forest range vegetation presented. The upper or higher shrub layer was very sporadic consisting of "here-and-there" isolated plants of flowering dogwood (left side of first slide), red maple (Acer rubrum) which, though a tree species, was represented as a shrub in this stand (left side of first slide), eastern redbud (right-center foreground of first slide and same plant at far-right in second slide). Common greenbriar (Smilax bona-nox) and an unidentified species of wild grape (Vitis sp?) formed a fairly continuous "botanical connection" from ground level to tree canopy.

205. Species composition of a second growth shortleaf pine-mixed hardwood stand with emphasis on shrub layers- In this forest range vegetation as shown in the first photograph hortleaf pines (in background) were joined by sweetgum (foremost tree at far right margin) and white oak (center midground) with an understorey dominated by the low shrub American beautyberry with yaupon as associate shrub species. An irregular or sporadic herbaceous layer consisted of longleaf woodoats (overwhelmingly the dominant herbaceous species) accompanied by splitbeard bluestem (Andropogon ternarius) as the associate herb.

Second and closer-in "photoplot" (second slide) presented this forest plant community from roughly the same camera point. Tree trunk in immediate foreground (left-center) was the sweetgum in far right margin of first slide. There was a sassafras seedling to right of this sweetgum. American beautyberry and yaupon (first and second, respectively, major shrubs in lower woody layer) were presented to better advantage than in the first of these two slides. Other ("also-ran") shrub species included common greenbriar; Small's, lanceleaf, or coral greenbriar, smooth sumac, winged sumac (Rhus copallinum), and an unindentified Vitis species. Red maple was present as large seedling and small sapling age classes so as to be part of this lower woody layer. Red maple was present as a small tree along margin of this forest tract (see below).

206. Synopsis views of vegetational layers- Interior of a second-growth shortleaf pine-mixed hardwoods forest showing the lower woody (shrub) layer. There were more shrub species and consequently greater diversity in understorey structure than in the local forest vegetation presented in the immediately preceding "photoplots". Hardwood species present in forest vegetation presented here included white oak, sweetgum, sassafras, and, especially, red maple. This latter species was present primarily as large seedlings to small saplings except on perimeter of forest where it grew to small tree size (shown bwelow).Here both smooth sumac and winged sumac were present at height intermediate between tall shrubs such as flowering dogwood and eastern redbud and lower shrubs like American beautyberry, the dominant shrub overall.

Shrub layers of the forest range vegetation in this local area was presented in the immediately succeeding pair of photographs at shorter focal length to better "sample" species composition and arrangement of this forest stand. Herbaceous species consisted mostly of grass species of which longleaf woodoats was "head-and-shoulders" above all others to rank as the dominant herb. Other important grass species included beaked panicgrass, rosette panicgrasses (Panicum species of the Dichanthelium section), purpletop, and splitbeard bluestem. Forbs were too limited to warrent remarks other than to note presence of an occasional plant of bracken fern, perhaps the most widely distributed plant species on Earth (greatest species range).

207. More emphasis on shrubs- Interior of a second-growth, upland shortleaf pine-mixed hardwoods forest presented so as to show greater detail of the shrub layer. This local area was the same "as introduced in " photoplots" represented in the immediately preceding pair of slides. More details of the lower woody and herbaceous layers (s) of this local stand were visible in these "photoquadrants". Dominance was a matter of extremely localized groups because some of the major species grew as clonal colonies. Smooth and winged sumac were most pronounced of these, but Small's or coral greenbriar (immediate center foreground in second of these slides) was also an obvious clonal plant spreading by "rootstocks" (woody rhizomes). Likewise, American beautyberry (the overall dominant shrub) grows in dense populations due perhaps to its typically abundant yields of fruits (immediate center foreground of second slide).

Other shrubs represented in these two photographs included poison ivy, an unindentified species of wild grape, and yaupon.Seedlings and small saplings of sweetgum and red maple were plentiful in this stand (they were especially noticable in the first slide). Herbaceous species were primarily grasses and included the dominant, longleaf woodoats, splitbeard bluestem, purpletop, beaked panicgrass, and rosette panicgrasses. the most conspicuous forb was bracken fern, but it was present as incidental, individual plants and did not form colonies or brakes.

208. On the outskirts- The outermost edge of an upland, second-growth shortleaf pine-mixed hardwoods forest featuring a large red maple (immediate center foreground). Most tree trunks were shortleaf pine except for an occasional white oak and sweetgum. Shrubs included American beautyberry, yaupon, and Small's or coral greenbriar. Herbaceous species were not dectible in this photograph taken at such distance as to show most of the nice specimen of red maple. (Details of lower layers of understorey were featured in the next two two-slide sets.)

209. Lower layers- Two "photoplots" of herbaceous and lower shrub layers of an upland, second-growth shortleaf pine-mixed hardwoods forest in the Texas Pineywoods. The first photograph presented beaked panicgrass and purple as local dominants of the herbaceous stratum (longleaf woodoats was the overall dominant herbaceous species; next set of slides). A species of rosette pancigrass was represented at far left. Yaupon, the overall associate shrub species, accounted for almost all cover in background of this first photograph. An unidentified species of wild grape was also present.

The second photograph included seedlings of red maple and sweetgum along with smooth sumac and American beautyberry at less-than-its-usul abundance for the lower woody layer. A plant of bracken fern, the only forb of much abundance, was also included for viewers' interest.

210. More detail of lower layers- Smaller "photoquadrants" of the lower woody and herbaceous layers of an upland, second-growth shortleaf pine-mixed hardwoods forest in the Texas Pineywoods. Longleaf woodoats was the star of this lineup and merited center-stage by nature of it being the dominant herbaceous on this forest range (across much of the Pineywoods Region for that matter). American beautyberry and yaupon with their characteristic leaves also were obvious in both photographs. Also present in the first photograph were beaked panicgrass and a rosette panicgrass as well as smooth sumac, all locally common species in this stand of forest vegetation.

211. Older pines, more hardwoods, and denser shade- Another stand of upland, second-growth shortleaf pine-mixed hardwoods forest that was adjacent to the stand featured in this section was made up of somewhat larger (resumedly older) shortleaf pines at conspicuously greater density and canopy closure. The downed pine was the victim of a hurricane six weeks prior to time of photograph. The tree closest to this laid-low pine (in front and at left of the trunk) was a sweetgum. Smaller tree with light gray bark in center midground was a Texas or black hickory (Carya texana) readily distinguishable to and handily identified by the photographer (not discernable in photographs) due to presence of 13 leaflets (the only hickory in this area to "sport" so many leaflets).

The only shrubs present with any remarkable cover were American beautyberry, dominant shrub of this forest plus flowering dofgwood and eastern redbud, those stewarts of the upper shrub layer across much of the eastern deciduous forest. All three of these shrub species provide mast for eastern wild turkey, the critical animal species for which this forest tract was being managed. Shade was too dense to permit much grass cover. Even the longleaf woodoats, the dominant herbaceous species, had been pretty much excluded from this "sylvan party".

212. Sunlite sample of structure- A small, local opening in a shortleaf pine-mixed hardwood second-growth forest afforded a view of forest structure and botanical composition of this foret range vegetation. In addition to the tall shortleaf pines there was a hardwood component of saweetgum, white oak, and black or Texas hickory in the tree layer. In addition there were two shrub layers: 1) taller layer of flowering dogwood and eastern redbud and 2) lower layer dominated by American beautyberry with yaupon as associate species of this stratum. There was an interrupted herbaceous layer dominated by longleaf woodoats.

213. Small's, lanceleaf, or coral greenbriar (Smilax smallii)- Common greenbriar (S. bona-nox) was comparatively more abundant than Small's greenbriar in the second-growth shortleaf pine-mixed hardwood forest featured in this section, but S. smallii was well-represented and afforded an opportunity to introduce viewers to another Smilax species.

Davy Crockett National Forest, Houston County, Texas. October; autumnal aspect.

Shortleaf Pine-Oak Forests

The shortleaf pine-oak range cover types originally covered extensive areas of the Boston and Ouachita Mountains as well as smaller portions of the central Ozark Plateau. Much of this was cleared for farming or as a result of cut-and-run logging practices (ie. cut-over land).Some of the cut-over land and a smaller proportion of the old fields went back to forest (at least of some sort). On much of this forest "go-back land" the various oak species regenerated better than the shortleaf pine, a successional development clearly exagerrated by cessation of fire which reduces hardwood cover substantially more than that of the highly fire-adapted shortleaf pine. In local areas there is still some of the shortleaf pine-hardwood (especially oak) forest, complete with natural understories (some better developed than others; some with more lower layers than others).

Local folk, particularily of older generations, knew these pine-hardwood forests by the colorful name of "pineries". In regions like the Ozarks (Ozark and Springfield Plateaus) and Boston-Ouachita Ranges (including the Kiamichi, Jack Fork, and San Bois Mountains) "pinery" was on par with "pineywoods" with both backwoods terms denoting the presence of hardwoods with pines (or vice versa) in contrast to forest of "pure pine". Examples of such pine-hardwood "pineries" werer included in this next section.

214. Pinery in the "Everlasting Hills of Oklahoma"- Exterior of a shortleaf pine-blackjack oak-post oak second-growth forest in the Elk River Hills of northeastern Oklahoma. Physiogonomy and structure of a typical second-growth shortleaf pine-oak forest known to locals as a "pinery". The "Everlasting Hills of Oklahoma" was a short, sweet song by the talented Sons of the Pioneers and used in the movie, Home in Oklahoma. Grasses in the foreground were broomsedge bluestem (Andropogon virginicus) and poverty oatgrass (Danthonia spicata). Herbaceous species in the understorey of this forest were limited to openings in the mostly closed canopy. The understorey consisted mostly of seedlings and saplings of the three tri-dominant tree species plus the huckleberry known variously as deerberry or squaw-huckleberry (Vaccinum stamineum).

Further details of this forest were given in the brief caption immediately below.

Delaware County, Oklahoma. Late June; early estival aspect. FRES No. 14 (Oak-Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Pinus echinata Association (if recognized) in a Oak-Pine Series of Brown et al. (1998, p. 37) or, alternatively, Warm Temperate Forest and Woodland (123), Southeastern Deciduous and Evergreen Forest (123.1), Pine Series (123.12) of Brown et al. (1998, p. 38). Ozark Highlands-Dissected Springfield Plateau-Elk River Hills ecoregion- 39b (Woods et al., 2005).

215. Pinery in the western Ozarks- Interior of shortleaf pine-blackjack oak-post oak second-growth forest in the Springfield Plateau of northeastern Oklahoma. Trees of the three dominant species were mostly small pole down to sapling and seedling. A local associate tree species ws red maple (Acer rubra). Trees of red maple were also small, mostly seedlings. The major shrub species was squaw-huckleberry or deerberry. Other shrub species included blackberry (Rubus spp.) and Virginia creeper (Parthenocissus quinquefolia). Absence of herbaceous species (except for occasional openings) was a function of closed forest canopy.

In this part of the Ozark Plateau the native shortleaf pine has been confined to shallow ridgetops where the native oaks and hickories cannot outcompete shortleaf pine. Lack of fire has been a major factor that has enabled native hardwoods to outcompete the also native shortleaf pine. The forest shown here was on a harsh site with a Clarksville stoney silt loam soil on roughly 10 to 20 percent slopes. This was an extremely shallow soil with cherty limestone rocks covering much of the land surface as well as extending down through the soil profile.

This was a cutover forest meaning that the virgin forest had been logged indiscriminately (they took everything merchantable; "slicked it off high and wide") with no provision (or even concern) for tree regeneration. The result was a "doghair" (excessively dense) stand of all tree species resulting in 1) stunted growth of trees and 2) limited understorey (due to dense shade from a closed tree canopy). Proper silviculture of this naturally regenerated forest would have included thinning.

A properly managed shortleaf pine-oak forest (and one with a proper understorey) and, in this instance, a climax forest community was shown next...

216. Introduction of a better example- The perimeter of a shortleaf pine-blackjack oak second-growth forest at edge of a big bluestem (Andropogon gerardii)-dominated tallgrass prairie in the Kiamichi Mountains of southeastern Oklahoma. Post oak was the associate (and, in local areas, the co-dominant) tree species in this forest. Othe locally important tree species included red maple and winged elm (Ulmus alata). Big bluestem was sole dominant (and, in many spots, the only) herbaceous species of this forest range vegetation. Other --though, again, minor--grass species included povery oatgrass, broomsedge bluestem, and little bluestem (Andropogon scoparius). At local scale there were patches of poverty oatgrass, which along with broomsedge bluestem, showed similarity of this shortleaf pine-oak forest with the one treated immediately above. The main shrub (and shrubs were mostly lacking in this climax forest vegetation) was winged sumac (Rhus copallina). There were very few forbs, but the prominent species was yellow wild-indigo (Baptisia bracteata).

This climax forest was, in effect, a tallgrass prairie (of mostly one species) with trees of shortleaf pine, blackjack oak, and (at lesser cover) post oak. From this perspective, this range plant community was "sort of" a savanna pattern except that tree crowns formed a nearly closed canopy so that this vegetation was forest and not even woodland let alone savanna. There was sexual reproduction of all three tree species. The understorey was a consociation of big bluestem with only a few other herbaceous species as given in the prededing paragraph. Trees had not reached their fully mature adult size, but the composition and basic structure of this climax forest was that of an old-growth forest. It was a climax forest that had yet to achieve the final form of an old-growth climax forest.

Choctaw Ranger District, Ouachita National Forest, LeFlore County, Oklahoma. Late June; early estival aspect. FRES No. 14 (Oak-Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Pinus echinata Association (if recognized) in a Oak-Pine Series of Brown et al. (1998, p. 37) or, alternatively, Warm Temperate Forest and Woodland (123), Southeastern Deciduous and Evergreen Forest (123.1), Pine Series (123.12) of Brown et al. (1998, p. 38). Ouachita Mountains- Fourche Mountains ecoregion 36d (Woods et al., 2005).

217. Ecological knife edge- The point of union of a tallgrass prairie dominated by big bluestem (foreground of both slides) and a shortleaf pine-blackjack oak-big bluestem forest in Kiamichi Mountains of southeastern Oklahoma. Post oak was the associate and, in local sreas, the co-dominant hardwood species. There were a few smaller trees (mostly saplings) of red maple and swinged elm (which natural or prescribed fire would have eliminated). Shrubs were very limited but the principal one was winged or shinning sumac. Inside this second growth-growth climax (composition-wise) forests there were some plants of poverty oatgrass along with some cover of broomsedge bluestem and little bluestem. Poverty oatgrass was not present out in the adjoining tallgrass prairie. The most common forb (in both forest and grassland) was yellow wild-indigo.

There was ample sexual reproduction of the three major tree species to maintain their continued presence (at the same relative composition) in this climax forest community.

218. Going to the inside (or standing on the outside looking in)-Outer edge of a climax shortleaf pine-blackjack oak-big bluestem second-growth forest in the Ouachita Mountains Region, specifically the Kiamichi Mountains of southeastern Oklahoma. Post oak was the associate tree species though it was locally the co-dominant hardwood tree species. Other trees present (mostly at sapling age/size classes) included red maple and winged elm. The major (and quite sparse in density) was winged or shining sumac. Density and cover of big bluestem was such that it was not only the dominant herbaceous species, but a defining dominant of this overall forests community. Other grasses in this forests range included pobverty oatgrass, broomsedge bluestem, and little bluestem.

This forest was the potential (climax) vegetation for this forest range site (based on species composition and general structure), but this forest range community was a younger second-growth forest that had not progresses to the ultimate old-growth state with numerous senescing trees and the with much dead and sowned timber.

219. From the inside- Interior of a second-growth shortleaf pine-blackjack oak-big bluestem forest in the Kiamichi Mountains of the overall Ouachita Mountain system. Post oak was the overall associate tree species, and being the co-dominant hardwood tree (with blackjack oak which was the overall co-cominant tree species with shortleaf pine). There were a few (and generally sparsely distributed) saplings of red maple and winged elm. For all practical purposes there was no shrub cover, although there were widely scattered plants (some with multiple shoots) pof winged sumac. Accompanying big bluestem (which comprised, in effect, a consociation of the herbaceous understorey) were poverty oatgrass, broomnsedge bluestem, and little bluestem. Most other grasses of tallgrass prairie in this area were not present.

This second-growtth forest was at climax state having the composition and basic structure of the terminal successional stage, but this forest vegetation lacked old-growth trees at the senescing/dying phase of their life cycles. As such there was not the downed, rotting tree component in this otherwise-climax forest range.

The second and third of this three-slide set featured leaves of shortleaf pine and blackjack oak growing side-by-side to represent the co-dominant trees, one conifer (gymnosperm), and one hardwood (angiosperm) species.

Students should nolte the well-developed and densely covered herbaceous layer made up largely of big bluestem.

220. Where trees were a little thicker- Interior of a shortleaf pine-blackjack oak-big bluestem second-growth forest in Ouachita Mountain Range (specifically, the Kiamichi Mountains) with a more limited herbaceous layer and more regeneration of blackjack oak. This was a different tract of forest than the tract featured in the immediately preceding four slide/caption sets. In the tract being "photo-sampled" in both of these slides large seedlings and small saplings of blackjack oak were featured prominentl. In the second slide a small sapling of shortleaf pine was growing to the rear and right of a large blackjack seedling (or very small sapling) indicating regeneration of the dominant conifer and dominant hardwood in this forest range. There were also a few small post oaks in this forest tract as in the tract from which the above four slide/caption sets were built. The major shrub species in the forest tract seen here was poison ivy and deerberry (Vaccinum stamineum).

There was a litter layer on the floor of this shortleaf pine-oak forest, but the duff layer was not so thick as to prevent growth of big bluestem as the dominant species of a sporadic herbaceous layer. The herbaceous layer was not nearly as well-developed or dense as in the forest tract feautured immediately above. This second-growth forest had the species composition of the potential natural (cvlimax) vegetation though without senescing old-growth trees.

The first or vertical slide was taken with temporary overcast sky while the second slide was taken as the sun once again gained the upper hand.

221. Where trees were not so thick- A second-growth (yet of climax species composition) shortleaf pine-blackjack oak-big bluestem forest in the the Kiamichi Mountains of the Ouachita Mountain Range in southeastern Oklahoma. In the two "photo-dendrographs" presented here the trees (most were small pole-size) were of a lower density and crown cover so that more sunlight reached the ground than in the slide-caption set presented immediately above. This forest range vegetation was in the same forest tract as in the preceding set, but there was both more grass (as in the second slide) and greater development of shrubs (as in the first slide). The most abundant shrub species in forest vegetation seen here were poison oak and deerberry or deerberry huckleberry.

Now for the bottomland- the six immediately preceding slide/caption sets were of shortleaf pine-blackjack oak-big bluestem forest on two tracts on an upland site. Now follow two sets of slide/caption units that showed forest range vegetation in a bottomland site. This bottomland forest community was on one tract that adjoined the last tract of upland forest covered above. All three tracts were second-growth forest that had the species composition and structure of the climax state (potential natural vegetation) except that senescing adult trees (and thus broken and fallen crowns, rotting limbs, and standing snags) were lacking.

222. Pines without the oaks- Exterior of a second-growth forest dominated by shortleaf pine on a bottomland site. Trees in this tract were fully mature and at or nearing their approximate full size (ie. they were in their "prime of adult life". This bottomland tract in the Kiamichi Mountains in southeastern Oklahoma adjoined the upland forest tract treated in the immediately preceding two slide/caption sets.

In this bottomland forest there were no hardwood tree species to speak of other some seedlings to small saplings of red maple and winged elm at outer edge of this lowland forest, the tree part of which was an even-aged stand of full-maturity shortleaf pine. Also at outer margin of this forest there were colonies of winged or shinning sumac which had been the principal shrub species on an adjoining tract of shortleaf pine-blackjack oak-big bluestem forest on an upland site that had a more open canopy. Also on the perimeter of this bottomland shortleaf pine forest the naturalized (frequently weedy) serecia lespedeza (Lespedeza sericea= L. cuneata) had grown into local populations.

Within this even-aged, lowland shortleaf pine forest there was almost no (essentially zero) regeneration of pine. Shrub species were extremely limited in this second-growth forest that was approaching the old-growth state of development. The dominant shrub was poison ivy which was distributed throughout this forest community. In ocal areas poison ivy had developed into a nearly complete covering of the forest floor. There was an herbaceous layer of eradic or sporadic cover that was made up of silky wildrye (Elymus villosus) and, in confined wet spots, common threesquare or chairmaker's bulrush (Scirpus pungens) was present as robust plants.

223. Shortleaf pine on a bottomland site- A tract of second-growth lowland forest with the tree component comprised of even-aged shortleaf pine. These pines that were nearing their "prime of life" (just before onset of senescent stage) with the single-species, even-aged stand approaching the old-growth state of development. There was, however, no regeneration of shortleaf pine (at least for all practical purposes). In a few small openings under the nearly closed forest canopy a few large seedlings of red maple and winged elm. No winged sumac was under the forest canopy although this shurb species grew prolificly on the margin of this forest and sparsely throughout an adjoining upland tract of shortleaf pine-blackjack oak-big bluestem.

In this even-aged, lowland shortleaf pine forest there was a two-layer, well-developed understorey consisting of 1) a shrub layer and 2) a sporadic herbaceous layer. The dominant shrub species of this bottomland forest was poison ivy which grew nearly everywhere and, in certain local areas, had developed into nearly complete coverage of the ground surface. The dominant species of the broken or interrupted herbaceous layer was the large-growing, big-spiked silky wildrye. In confined wet spots there were some robust plants of chairmaker's or common threesquare bulrush. This latter, grasslike plant which is characteristic of marshes indicated that this lowland shortleaf pine forest was almost a wetland or wetland forest. This was an atypical habitat for shortleaf pine which is less mesic in its requirements than loblolly pine (Pinus taeda), this latter species of which is not native to this area (Burns and Honkala, 1990). Unexpected in this mesic habitat were local populations of one or the several taxonomic variety of black-eyed susan (Rudbeckia hirta) which made this composite the major (and about the only) forb of this bottomland singe tree- species forest.

The first two (the vertical) slides of this three slide-set were obviously of the same image (the same "photo-dendrograph"), but the first image was under a full-sun sky whereas the second image was taken a few moments later under an overcast sky as a large cumulus cloud came between the sun and forest vegetation. Sometimes overcast images show forest vegetation to an advantage as there is less shade or sunlite and shaded spots intermingled. Other times, full-sun shots show forest plant communities to better advantage. In this instance, viewers have their choice. The third (horizontal) slide was of the same location as the two vertical shots.

224. Seedtree regeneration method for shortleaf pine- A shortleaf pine-post oak about two and a half years following harvest using the seedtree method of reproduction. Seedtree cutting is an even-aged method of forest regenertion whereby all trees (shortleaf pine in this example) are cut other than a reduced number that are left at wide spacings for the purpose of seed production and thereby production of the next crop of trees. In turn the seed trees are cmmonly harvested upon establishment of the next tree (shortleaf pine) crop (Helms, 1998, p. 151). The example seen here was of the shortleaf pine-oak-hickory forest cover type in the Ouachita Mountains Region (specifically in the Kiamichi Mountains) of southeastern Oklahoma.

In this example the apparent main result of seedtree cutting was release of existing trees and sexual reproduction oak and hickory rather than establishment of the next crop of shortleaf pine. Yes, there were some seedlings of shortleaf pine, but the main outcome was establishment of two main hardwood species: 1) post oak which, with both numerous seedlings and released sapling to small pole-sized trees, became co-dominant with shortleaf pine and 2) mockernut hickory (Carya tomentosa) which became the associate tree species in this seedtree cutting.

Seedtree regeneration also resulted in increase of herbaceous cover in the understorey. Grasses dominated the herbaceous understorey of this seedtree method with poverty oatgrass, silky wildrye, and forked or cypress rosette panicgrass (Panicum dichotomum) being the major Gramineae species. The major forb species in this newly released uynderstorey was pokeberry or pokeweed (Phytolacca americana) which was growing everywhere.

The first (horizontal) slide presented an exterior view of this shortleaf pine-post oak pine forest to show physiogonomy and overall structure while the second (vertical) slide was an interior view to present details of structure and species composition, including numerous seedlings of post oak and mockernut hickory in the predominant herbaceous understorey.

225. Seedtree method in shortleaf pine- A second-growth shortleaf pine-post oak upland forest in the Kiamichi Mountains) of southeastern Oklahoma roughly two and a half years following harvest. The seed tree method of regeneration is a silvicultural approach to even-aged management. Most trees of harvestable age/size are cut yet leaving some widely spaced trees of the harvest cohort as seed-producers to establish the next even-aged stand from the new cohort seedlings. Later, the seed trees can be harvested.

In the example of the forest tract seen here and in the immediately preceding slide/caption unit the main apparent outcome was not so much establishment of the next crop of shortleaf pine (although there was some of this) was release of existing trees of post oak and, at lesser cover, mockernut hickory along with establishment of seedlings of these two hardwood treee species. Seedtree cutting also resulted in a tremendous increase of pokeberry or poke-salad along with greater cover and biomass of poverty oatgrass, silky wildrye, and forked or cypress rosette panicgrass.

The first (horizontal) of these two slides showed co-dominant post oak as both a small pole and large seedlings in foreground to shortleaf pine left as seed trees in mid- and background. The second (vertical) slide presented the bole of a shortleaf pine along far right margin with a seedling of post oak to its left in center foreground. These two images portrayed the co-dominant post oak and revealed that post oak and mockernut hickory had benefitted more than shortleaf pine, at least up to this time post-harvest of the pine crop.

The seed tree method in this forest tract had restored more of the climax (potential natural) vegetation of the pine-oak-hickory forest rather than insuring the next crop of shortleaf pine. Plus, from a rangeman's perspective the seedtree method amounted to tree-thinning and commensurate increase in the herbaceous understorey, the range forage crop.

Side note: just after your author had taken these photographs and returned to his rig, a carload of tourists stopped asking if this photographer had gotten any pictures of the bear. Apparently an adult black bear (Ursus americana) had crossed the road just below this seed tree forest tract and ran up throught it. The bear went over the mountain and this photographer did not see the bruin, but maybe its likeness is somewhere in the cover of vegetation in one of these images. Or, maybe not. Either way viewers can use their imagination and understand how this forest manipulation provided increased cover for black bear habitat.

226. Sylvan line-up- Crowns of southern red oak (Quercus falcata), shortleaf pine, white oak (Q. alba), and black tupelo or blackgum (Nyssa sylvatica), left to right, respectively in a relict forest tract of these four species plus sweetgum (Liquidambar styraciflua), red maple, and winged elm in the Kiamichi Mountains (a range in the Ouachita Mountain mountain series) in southeastern Oklahoma. There was a very limited understorey in this forest, most seedlings and saplings of the dominant hardwood species with fewere of shortleaf pine. The limited herbaceous component was dminated by narrowleaf woodoats (Uniola sessifolia).

LeFlore County, Oklahoma. Late June; early estival aspect. FRES No. 14 (Oak-Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Pinus echinata Association (if recognized) in a Oak-Pine Series of Brown et al. (1998, p. 37) or, alternatively, Warm Temperate Forest and Woodland (123), Southeastern Deciduous and Evergreen Forest (123.1), Pine Series (123.12) of Brown et al. (1998, p. 38). Ouachita Mountains- Fourche Mountains ecoregion 36d (Woods et al., 2005).

227. Heavy hittters of the shortleaf pine-mixed oak type- These two slides were closer-in views of the crown canopy introduced and described in the immediately preceding slide. In the first of these two slides tree crowns were--left to right, respectively--shortleaf pine, white oak, and southern red oak. In the second (vertical) slide the upper trunk and lower crown of a white oak was on the left and of a shortleaf pine on the right. These trees were part of a mixed hardwood-shortleaf pine forest in which sweetgum, blackgum or black tupelo, red aple, and winged elm accompanied the three major species (shortleaf pine, white oak, and southern red oak). There was not a well-developed understorey in this forest of dense shade. Narrowleaf woodoats was the major herbaceous species. There were some seedlings and saplings of all tree species, but the dominant oaks were most plentiful.

228. Scarred heavy hitters- Interior of a shortleaf pine-mixed hardwood (mostly oak) forest in the Ouchita Mountains (actually the Kimichi Mountains, a smaller or sub-range within the general Ouachita Range) in southeastern Oklahoma. The two slides in this set comprise a "nested photoplot" with the second slide being a "sub-plot" of the first or general "plot". This second (horizontal) photograph was taken from a different angle to show the three dominant tree species (tri-dominants) of this climax forest community: shortleaf pine, white oak, and southern red oak. Southern red oak was represented by the two-trunked (forked just above ground level) tree in left-center foreground in the first slide and at left foreground in the second slide. White oak was in left midground (to immediate left of the forked-trunk southern red oak) in the first slide and in center midground of the second slide. Shortleaf pine was behind and to immediate right of the forked-trunk southern red oak in the first (vertical) slide and conspicuous in right foreground in the second (horizontal) slide.

While the two dominant hardwood species displayed prominent fire scars (visible in white oak only in the second image), the more fire-tolerant shortleaf pine proudly bore only its beautifully fire-burnished bark. The shortleaf pine and shortleaf pine-oak forest cover types (SAF 75 and SAF 76, respectively, Eyre, 19890) are fire types that evolved under natural fire as a determinative atmospheric phenomenon. Fire is essential in maintenance of these two timber types.

It was not known to this author if the fires in this forest were wild fires or prescribed fires set by the US Forest Service. The Forest Service (one of the finest conservation organizations on Earth) could certainly use the credit for prescribed buring after a century of it's carried-to-excess fire-suppression policy, it 's enamorment with the fire-preventing Smoky Bear (a sweetheart of its own invention).

There was a surprisingly limited understorey in parts of this climax composition, seccond-growth shortleaf pine-mixed oak forest. In other parts (as in background of both these views) the understorey had both a shrub and an herbaceous layer. In these two views, lowbush blueberry or lowbush huckleberry (Vaccinum vacillans), a major low-growing shrub in this forest range, stood between the southern red oak and shortleaf pine in the second slide and to right of the southern red oak and to left front of the shortleaf pine in the first slide. Narrowleaf woodoats was generally the dominant herbaceous species of the understorey.

Ouchita National Forest, LeFlore County, Oklahoma. Late June; early estival aspect. FRES No. 14 (Oak-Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Pinus echinata Association (if recognized) in a Oak-Pine Series of Brown et al. (1998, p. 37) or, alternatively, Warm Temperate Forest and Woodland (123), Southeastern Deciduous and Evergreen Forest (123.1), Pine Series (123.12) of Brown et al. (1998, p. 38). Ouachita Mountains- Fourche Mountains ecoregion 36d (Woods et al., 2005).

229. Densely southern- Loweeree limb of southern red oak (Quercus falcata) in an upland shortleaf pine-mixed hardwood second-growth forest in the Kimichi Mountains (a smaller range within the Ouachita Mountain chain) in southeastern Oklahoma. Southrn red oak is adapted to slightly less mesic habitts than its cousin, northern red oak (Q. rubra).

Ouachita national Forest, LeFlore County, Oklahoma. Late June.

230. Southern leaves- Dense leaves on lower leaders (woody branches) of southern red oak in the the Kimichi Mountains (one of several smaller mountain ranges within the Ouachita Mountain chain) in southeastern Oklahoma (first slide). An example of the cluster arrangement of southern red oak (second slide). Southern red oak is in the red (or, sometimes, black) oak subgenus, Erythrobalanus. Species in the red (black) oak group are characterized by extension of veins or ribs beyond leaf margins and by a period of two years (two growing seasons) for acorns to reach maturity. Acorns in the white oak group (subgenus Leucobalanus) mature in one year (one growing season).

Ouachita national Forest, LeFlore County, Oklahoma. Late June.

231. An example of the shortleaf pine-northern red oak forest- Exterior view of an upland shortleaf pine-northern red oak climax forest showing its physiogonomy and gross structure. While northern red oak was co-dominant with shortleaf pine there were also trees of post oak, mockernut hickory, and red maple. Major shrub species included eastern hop-hornbeam (Ostrya virginiana), poison ivy, and cat green-brier (Smilax glauca), and muscadine grape (Vitis rotundifolia). The understorey of this second-growth forest varied between two traacts that comprised it (see next paragraph), but generally lower layers of forest vegetation consisted of taller shrubs, lower shrubs, seedlings and saplings of the tree species, and widely (and sparsely) scattered herbaceous species, expecially narowleaf woodoats (Uniola sessifolia).

This forest tract consisted of three adjoining management units: 1) two stands of young trees of both shortleaf pine (these were barely half-grown) and oak species and 2) one stand of mature, starting-to-senesce shortleaf pine and younger trees of the oak species. Both stands of this tract had evidence of relatively low-intensity surface fires (perhaps prescribed fire).

This second-growth forest was an example of the shortleaf pine-northern red oak subtype or variant of the general shortleaf pine-oak forest cover type (Eyre, 1980).

Note on forest foliage- The slidesof this forest tract (these three forest stands) were taken immediately after a moderate intensity rain shower that created the effect of glistening leaves and shoots (bark). Nice effect for a fine forest.

232. Interior of a stand of young trees- Deep inside a stand of half-grown to nearly mature shortleaf pine and northern red oak in the Kimichi Mountains (of the general Ouachita Mountain Range or chain) in southeastern Oklahoma. This tract had obvious evidence of comparatively low intensity surface fire (natural or prescribed fire was not known to the photographer). The understorey was primarily of woody plant species except for a few sparsely populated plants of narrowleaf woodoats and, even more sparse in cover and density, big bluestem. These two grasses were the classic decreaser species for this forest range habitat.

The firest (horizontal) slide presented the general structure and species composition of this second-growth forest that was co-dominated by young trees of shortleaf pine and northern red oak and with a varied understorey that included cat green-brier (Smilax glauca), poison ivy, and eastern hop-hornbeam (Ostrya virginiana). There were a large number of mockernut hickory seedlings at the lower understorey level as shown in the first slide. The second (vertical) slide also presented the structure of this forest emphasizing the locally well-developed understorey and featuring characteristic leaves of eastern hop-hornbeam in lower left corner opposite from the lichen-covered trunk of a young northern red oak (right margin).

Although the trees of this forest tract were young, the species composition of this forest range was that of the climax vegetation.

233. One for the shade- Shoots with characteristic leaves of eastern hop-hornbeam (Ostrya virginiana) one of the most shade-tolerant shrubs in the eastern deciduous forest of eastern North America (Wenger, 1984) and a major understorey woody species in a second-growth (though climax) shortleaf pine-northern red oak in Kimichi Mountainsin southeastern Oklahoma. There were also some leaves of cat green-brier in top-center portion of this slide. This foliage was in the understorey of the forest tract seen in the immediately preceding slide/caption unit.

234. Inside another stand of young 'uns- Another stand of the shortleaf pine-northern red oak subtype of the general shortleaf pine-oak forest cover type that had developed in a tract of this type within Kimichi Mountains (part of the overall Ouachita Mountain Range or chain) in southeastern Oklahoma.. This stand was conterminous with the stand presented immediately above other than being on the opposite side of a road. This stand had relatively less cover and density of northern red oak and ccomparatively more cover of post oak, this latter of which was the associate species to the co-dominants. In the views shown here big bluestem had almost as much cover as narrowleaf woodoats. The major species of the lower woody portion of the understorey was cat green-brier. There were also a lot of plants of poison ivy as well as several plants of muscadine grape.

235. Where the trees were older- The third of three stands of a shortleaf pine-northern red oak forest in one forest tract in the Kimichi Mountains (of the general Ouachita Mountain Rangeor chain ) in southeastern Oklahoma. These three slides showed the interior of a second-growth shortleaf pine-northern red oak climax forest, a subunit or variant of the "coverall" shortleaf pine-oak forest cover type (Eyre, 1980), with adult pine trees approaching what was probably their maximum possible (their genetic potential) size and northern red oak at roughly one-third to one-half of their ultimate adult size. It appeared that northern red oak established after colonization of shortleaf pine. Northern red oaks were identifiable in these three slides by the dark-grey bark with long, shallow furrows. Pine bark was obvious except in the third slide where shortleaf pine was in left foreground and northern red oak was in right midground.

Shrubs in this second-growth (and climax species composition) forest included eastern hop-hornbeam, cat green-brier, poison ivy, and muscadine grape. There was almost no herbaceous growth (cover) in the understorey of this nearly closed-canopy forest.

In the first and third of these three slides there many seedlings of mockernut hickory near the ground level. In the second slide there was a small sapling of mockernut hickory in the lower left extending up the left margin while there were leaves of this same sapling that extended across to the upper right corner. There were conspicuous shade leaves of mockernut hickory in the lower left corner and immediate foreground in the third slide. Edge of a crown of a blackgum or black tupelo sapling filled the right border of this third slide. Considerable cover of eastern hop-hornbeam was in center fore- and midground of this third slide also.

236. Co-dominant and their forest friends- Northern red oak (left) and shortleaf pine (right) were co-dominants of an upland shortleaf pine-northrn red oak subtype of the shortleaf pine-oak forest cover type (Eyre, 1980) in the Kimichi Mountains (local range in the of Ouachita Mountain chain) in southeastern Oklahoma. There were a few smaller trees (seedlings to small saplings) of red maple. There was a well-developed woody understorey (generally of two layers) in this second-growth (climax vegetation as to species composition) forest. Shrub species included eastern hop-hornbeam, Virginia creeper, poison oak, cat gren-brier, and muscadine grape. This woody understorey pretty much prevented development of herbaceous species other than occasional plants of narrowleaf woodoats.

237. An "understanding" at the edge- Example of lower herbaceous layer of a shortleaf pine-northern red oak second-growth forest in the Kimichi Mountains (part of the of Ouachita Mountain chain) in southeastern Oklahoma. The "curly" looking foliage-appearing material that occupied the greatest cover in the first slide and that dominated the center of the second slide was Dixie reindeer moss (Cladonia subtenius), a species of fruticose lichen with a habit or growth morphology comprised of hollow "limbs" or stalks. The grass in these two "photo-plots" (the second slide being a "sub-plot" of the overall "plot" or the first slide) was big bluestem (Andropogon gerardii). There was some species of moss and an unknown graminoid.

238. Another version of shortleaf pine-oak forest- Exterior view of an upland shortleaf pine-mixed oak climax forest showing its physiogonomy and gross structure. This was a remarkabley diverse forest community with a nearly unbelievable array of oak and other hardwood species. Oak species present included white, northern red, southern red, black (Quercus velutina), and post oaks. Other major hardwood tree species included mockernut hickory, red maple, blackgum or black tupelo, winged elm, and, sweetgum with this latter early seral species generally at lesser cover and, more commonly, confined to the outer perimeter of this climax forest. Major shrub species included eastern hop-hornbeam, poison ivy, both cat green-brier and roundleaf green-brier (Smilax rotundifolia), and muscadine grape. Herbaceous cover was limited, but there were scattered plants of narrowleaf woodoats.

This forest community was in the Caddo Mountains which are part of the larger or general Ouchita Mountains of southwestern Arkansas.

Ouchita National Forest, Polk County, Arkansas. Mid-July; early estival aspect. FRES No. 14 (Oak-Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Pinus echinata Association (if recognized) in a Oak-Pine Series of Brown et al. (1998, p. 37) or, alternatively, Warm Temperate Forest and Woodland (123), Southeastern Deciduous and Evergreen Forest (123.1), Pine Series (123.12) of Brown et al. (1998, p. 38). Ouachita Mountains- Fourche Mountains ecoregion 36d (Woods et al., 2004).

239. Shortleaf pine and a lot of hardwood friends- Interior of an upland shortleaf pine-mixed hardwood second-growth (climax species composition) forest in the Caddo Mountains (a range within the ( general Ouchita Mountain chain) of southwestern Arkansas. Shortleaf pine was visible in each of these three images as was blackgum or black tupelo (Nyssa sylvatica), a highly shade/competition tolerant hardwood of the dogwood family (Cornaceae) or, alternatively, its own family, Nyssaceae. Cat green-brier (Smilax glauca) and, to a lesser cover, roundleaf green-brier were widespread in this forest commmunity. Muscatine grape was also common though less so than cat green-brier. Poison ivy was a species that occurred as both a lower bush-like shrub as well as a tree-climbing woody vine. These three species added a liana or woody vine component to the two shrub layers of this forest. Tree-like shrubs such as eastern hop-hornbeam made up a sporadic or erradic upper shrub layer. There seedlings as well as saplings of hardwood tree species including white oak, northern red oak, black oak (Quercus velutina), red maple, and mockernut hickory (Carya tomentosa). Seedlings of mockernut hickory frequently grew in local clusters, all except one of two of which would eventually succumb due to shading or some other form of competition. Herbaceous plants were limited mostly to narrowleaf woodoats which is a decreaser grass species and commonly the herbaceous dominant in shortleaf pine forests in all (at least most) of its varaints and subtypes.

Blackgum or black tupelo was conspicuous as several saplings in the first slide and as a large sapling or small pole in the right foreground (opposite a shortleaf pine to its left) in the third slide. The large sapling or small pole in left foreground of the secondslide was a white oak. There wer several plants of eastern hop-hornbeam visible in each of these three slides. Plants of narrowleaf woodoats were most common in local patches such as one in center midground of the second slide.

240. Big old pines and their understudies- Four old-growth specimens of shortleaf pine with associated understorey species were the subjects of these two views of species composition and structure of a climax upland shortleaf pine-mixed hardwood forest in the Caddo Mountains (part of the Ouachita Mountain chain) in southwestern Arkansas. These four shortleaf pines were at the ultimate state of adulthood just before onset of old-age senescence. They could tell some woodsy tales (note the large fire scar on right lower trunk of the pine in right midground of the first (vertical) slide. The sapling between the two old-growth shortleaf pine in this first slide was a white oak. Behind (to rear) of this white oak were two young trees of sweetgum. The bush in left-center foreground of the first slide was eastern hop-hornbeam. Eastern hop-hornbeam was well-represented in the second (horizontal) slide by two small saplings in center foreground and left midground to immediate right of the left shortleaf pine trunk. Poison ivy was also represented in the second slide (immediate foreground).

A leaf layer covered the soil surface in this climax forest range, buth this did not appear to have hampered development of hardwoods (either shrub or tree species). There was not, however, much regeneration of shortleaf pine, seedlings of which were 'few and far between". Overall, even mature trees of shortleaf pine were less plentiful than plants of more tolerant woody species like blackgum (= black tupelo) and eastern hop-hornbeam or even than white oak which is less tolerant than the former two hardwood species (Wenger, 1984).

241. Side-by-side differences- The concept of tolerance (Wenger, 1984) was graphically (and beautifully) shown in this two views deep inside an upland shortleaf pine-mixed hardwood forest in the Caddo Mountains (a range within the Ouachita Mountain chain) in southwestern Arkansas. Shortleaf pine, the climax dominant (and only) conifer of this forest cover type, was represented in the first slide in center midground (to immediate left of the foremost trunk) and at left margin in the second slide. Black gum or black tupelo was present in the first slide as the foremost trunk (a large sapling) and in the second slide as a small pole and sapling, respectively, to right of the shortleaf pine. A white oak seedling was present in the lower right corner of the first slide. Shoots of poison ivy were present in foreground of both slides.

The complex phenomenon of tolerance (the general, relative or comparative adaptation of forest tree and shrub species to competition; especially for light, hence general adaptation to shade) can be seen readily by students in this narrow array of native woody plants. Wenger (1984, ps. 2-3) presented accepted tolerance ratings (categories) of vrious forest species. Shortleaf pine was rated as Intolerant whereas eastern hop-hornbeam (which grew throughout the understorey of this forest stand) was Very Tolerant and the tupelo species ranked as Intermediate, though not conclusively so. Burns and Honkala (1990) described black tupelo (= blackgum) as "tolerant of shade", but they noted that black tupelo seldom attains dominance in the upper forest canopy (forest crown layer) and instead typically occcupies an "intermediate crown position". In summarizing findings related to tolerance of mockernut hickory, Burns and Hankala (1990) reporting classifiction ratings from Tolerant to Intolerant. Thus it appears that tolerance of mockernut hickory, which is widespread in the Pineywoods Region, is forest-site specific. The author of Range Types of North America found mockernut hickory to respond to shade as Tolerant or, at least, Intermediate in the western Pineywoods.

Tolerance ratings (Wenger, 1984, ps. 2-3) of other important (at least locally important ) tree species in this forest were: red maple, Tolerant; white oak, Intermediate, and sweetgum, Intolerant. This shortleaf pine-mixed hardwood forest had developed a more open forest canopy (it was not a completely closed canopy forest) such that there was a wide array of tolerance ratings among a diverse group of woody plants distributed in one adult tree layer and two shrub or, more accurately, woody layers (one of which had sapling- and pole-sizes and the other, seedlings).

This wide variation in tolerance ratings was exhibited even more in the next two-slide/caption set…

242. Pronounced differences- Another two views in the interior of an upland shortleaf pine-mixed hardwood forest in the Caddo Mountains of southwestrn Arkansas taught the concept of woody plant tolerance tolerance as applied to Forest Ecology. In the first slide there were two mature, adult shortleaf pines, a species with a tolerance rating (Wenger, 1984, ps. 2-3) of Intolerant, along with a number of young saplings and poles (both classes having various sizes) of blackgum or black tupelo, a hardwood species with a tolerance rating of Intermediate (Wenger, 1984, ps. 2-3) and seedlings of red maple (eg. lower left corner of first slide), a species that Wenger (1984, ps. 2-3) reported as Tolerant. The sapling in the immediate center foreground and the small pole to right of the right shortleaf pine in the first slide were both blackgum.

In the second slide the crooked ltrunk in left foreground was eastern hop-hornbeam, shrub species with a tolerance rating of Very Tolerant (Wenger, 1984, ps. 2-3), and the biggest trunk and the sapling to its left in right midground were of the Intermediate blackgum. The lower-growing (short) broadleaved plants on the forest floor in the second were seedlings of mockernut hickory, red maple, black oak, and white oak along with poison ivy, cat green-brier, roundleaf green-brier, and muscadine grape. There was some herbaceous cover of narrowleaf woodoats, a decreaser and the dominant herb of this upland forest.

Detailed treatment of the lower layers of vegetation on and near the forest floor were presented in the next two slide/caption units.

243. Give me shade, lots of shade or, alternatively, sun, lots of sun- Two views of a small pole of blackgum or black tupelo in the understorey of an upland shortleaf pine-mixed hardwood forest in the Caddo Mountains of southwest Arkansas. The first slide presented an overall view of the lower trunk and the entire length of a lower leader (woody branch) of blackgumwhile the second slide showed more details of bark and the simple, alternate leaves of this common hardwood tree species that is widespread throughout much of the Southeast Pine Belt and, for that matter, the vast Deciduous Forest Region of Eastern North America (Burns and Hankala, 1990).

Black tupelo is a hardwood tree species that is extremely versatile in its light requirements. Blackgum has generally been regarded as having a tolerance rating of Intermediate (Wenger, 1984, ps. 2-3) although Burns and Hankala (1990) concluded that black tupelo ranges from Intermediate to Tolerant depending on forest site and local habitat.

Black tupelo is typically second only to sweetgum as a pioneer on freshly clearcut loblolly pine plantations yet it also thrives in the understorey of shortleaf pine and shortleaf pine-hardwood forest cover types. as, for instance, in the western Arkansas-eastern Oklahoma portions of the Pineywoods Region such as in the Ouachita Mountains area featured in this section.

Ouachita National Forest, Polk County, Arkansas. Mid-July.

Versatile leaves- Outer portion of a young leader (woody branch) of black tupelo showing the alternate arrangement of oblong simple leaves along this stem. Recommended refrerences for blackgum included Sargent (1933, ps. 780-781), Vines (1963, ps. 801-802), Harlow et al. (1979, 428-429), Burns and Hankala (1990), and Samuelson and Hogan (2003, ps. 148-149). As described in these references (especially Sargent, 1933) leaves of black tupelo are quite varied as, for that matter, is the bark (see immediately below).

Ouachita National Forest, Polk County, Arkansas. Mid-July.

Barked in shade (this time)- Part of the trunk (with a short lower leader and leaves) of a small (young?) blackgum or black tupelo in the understorey of an upland shortleaf pine-mixed hardwood forest in the Caddo Mountains of southwest Arkansas. Blackgum is adapted to a wide variety of habitats over a large species range that cover much of the huge (formerly huge) Eastern Deciduous Forest which include much of the Pineywoods Complex such as that which was treated in this section.

The bark of blackgum is remarkable varied or diverse, seemingly becoming more smooth with age and increases in trunk size. (Refer to the various references cited in the immediately preceding caption.)

Ouachita National Forest, Polk County, Arkansas. Mid-July.

Shade tolerance in spades- Small sapling-sized eastern hop-hornbeam (Ostrya virginiana) growing in the woody intermediate layer of a shortleaf pine forest in the ancient Ouachita Mountains (specifically the Caddo Mountains which is a relatively small range within the Ouachita Mountain chain) in southwestern Arkansas. No native hardwood species has a greater tolerance rating than eastern hop-hornbeam which scored in the highest tolerance rating group designated as Very Tolerant (Wenger (1984, ps. 2-3). The species (biological) range for eastern hop-hornbeam extends from the Maritime Provinces like Nova Scotia westward through Quebec to parts of the Great Plains as in eastern Wyoming and as far south as the Pineywoods of Deep East Texas. Eastern hop-hornbeam is native to every state and province bordering the Mississippi River and eastward to the Atlantic Ocean.

A good reference for eastern hop-hornbeam included Ritz (2012) and the ever-ready Vines (1963, p. 145). Eastern hop-hornbeam was regarded as a tree species by Sargent (1933, ps.202-204), Harlow et al. (1979, p. 280), and the U.S. Forest Service and, as such, appeared in the agency's compendium on tree silvics (Burns and B. H. Honkala, 1990). Eastern hop-hornbeam is typically a smaller, shorter tree reaching heights of 20 to 30 feet, but some individuals reach up to 60 feet (Sargent, 1933, p. 203).

Ouachita National Forest, Polk County, Arkansas.

Let it shade- Branch with typical leaves of eastern hop-hornbeam growing in a shortleaf pine-mixed hardwood forest in the Ouachita Mountains of southwest Arkansas. Eastern hop-hornbeam falls in the highest tolerance rating of any North American woody species: Very Tolerant (Wenger (1984, ps. 2-3). This species grew along with blackgum or black tupelo in the same mid-level of the woody layer of this Pineywoods forest.

Ouachita National Forest, Polk County, Arkansas.

Starting at the bottom; reaching for the top- The lower woody layer of an upland shortleaf pine-mixed hardwood forest in the Cddo Mountains of southwestern Arkansas. In the first of these two slides plants were mostly seedlings of red maple and mockernut hickory with substantial cover of poison ivy. In the second slide most plant cover was also of seedlings, in this case of tree species , black oak and red maple, and of the shrub eastern hop-hornbeam. Also in the part of the forest floor seen in this second slide there was shrub cover of poison ivy and muscadine grape along with narrowleaf woodoats.

Numerous fallen branches on the forest was indicative of older, maturing (senescing) trees, primarily of shortleaf pine. There was considerably less regeneration of pine than of red maple, mockernut hickory, and the oak species. Nonetheless, pine regeneration appeared to be adequate to maintain shortleaf pine in a shared dominance status with the vrious hardwood species which suggested that this upland forest had the species composition--though not the structture--of a climax forest.

The surface of this forest soil was completely covered by a fairly thin mulch of leaves (mostly of hardwood species), but this leaf litter did not hinder germination, emergence, and development of shortleaf pine or hardwood tree and shrub species. Any impact on growth and development of herbaceous species was not known though, again, there was locally (patchy or erradic) heavy cover of narrowleaf woodoats, the dominant decreaser grass species of shortleaf pine types throughout much of the southeastern Pineywoods Region.

Still on the bottom and headed for the top- Two closer-in views of the lower woody layer on or near the floor of an upland shortleaf pine-mixed hardwood forest in the Caddo Mountains (a range in the Ouachita Mountain chain) in southwestern Arkansas. Plants in the first slide were mostly seedlings of black oak (left) and northern red oak (right) along with seedlings of eastern hop-hornbeam as well as cover of adult plants of cat green-brier, roundleaf green-brier, and muscadine grape (Vitis rotundifolia). There were also a few seedlings of winged elm (Ulmus alata) and blackgum or black tupelo. These plants were growing amid downed branches of senescing shortleaf pine.

Plants in the second slide were muscadine grape with lesser cover of cat green-brier, poison ivy and (as represented by a few leaves in immediate left-center foreground) mockernut hickory (Carya tomentosa). There was also a high proportion of the cover in poison ivy.

Not actually the cat's meow- Cat green-brier (Smilax glauca), a major, widespread liana (woody vine), along with some eastern hop-hornbeam (Ostyra virginiana) in an upland shortleaf pine-mixed hardwood forest in the Caddo Mountains (a range within the Ouachita Mountain chain) in southwestern Arkansas.

Ouachita National Forest, Polk County, Arkansas. Mid-July.

Still not the cat's meow- Roundleaf green-brier (Smilax rotundifolia) as part of the shrub component of an upland shortleaf pine-mixed hardwood forest in the Caddo Mountains (a range within the Ouachita Mountain chain) in southwestern Arkansas.

Ouachita National Forest, Polk County, Arkansas. Mid-July.

The cat's something- Shoots of roundleaf green-brier which is one of several Smilax species found in the western part of the Pineywoods Complex. In much of this large geographic area cat green-brier is more common than roundleaf green-brier, but these two often grow in close proximity to each other. First of these two slides was in a loblolly pine (Pinus taeda)-bald cypress (Taxodium distichum) bottomland forest in the Red River Valley of southwestern Arkansas. The example in the second (vertical) slide was also in a loblolly pine-dominated forest on an upland forest range site. This second slide showed the wide variation in roundleaf green-brier.

First slide: Sevier County, Arkansas. Mid-July. Second slide: Old Sabine Bottom Wildlife Management Area, Smith County, Texas. Late June.

Sampling another tract- Still yet another tract of an upland shortleaf pine-mixed hardwood forest in the Caddo Mountains (a small range in the Ouachita Mountain chain) in southwestern Arkansas provided an example of the shortleaf pine-oak forest cover type (Eyre, 1980). In addition to the trunk of a mature shortleaf pine in left foreground there were trees of white oak (right-center midground) and northern red oak (right foreground to midground) along with saplings and small poles of black oak, red maple, winged elm, black tupelo, and mockernut hickory.

Shrub species included the understorey woody dominant, eastern hop-hornbeam, plus the woody vine species of poison ivy, cat green-brier, roundleaf green-brier, and muscadine grape. Herbaceous species were limited, but as in similar tracts of the shortleaf pine-oak cover type, narrowleaf woodoats (a decreaser grass) was the dominant herb of the understorey.

Nested plots of another tract- Interior of an upland shortleaf pine-mixed hardwood forest in the Caddo Mountains of southwestern Arkansas showing its structure and species composition. Shortleaf pine was featured prominently as the two foremost trunks in the first slide and by the trunk at left margin in the sedond slide. Close observation revealed that the pine trunk at the left margin (immediate left foreground) in the first slide was naught but a broken snag. Likewise there were several broken branches of shortleaf pine lying on the ground or even driven into the ground in the first slide. Some of hese trees were actually old-growth shortleaf pine that had died and were falling to the ground. Even though these dead and rotting pines were rather small they had reached adult age and size for this shallow, dry, upland forest site. There was enough shortleaf pine reproduction to maintain the proportionate cover of this conifer in the present climax forest.

These two slides comprised a "nested photo-plolt" with the second slice being a closer-in view--a "sub-plot"--of part of the bigger spatial-scale, first slide which was the overall "photo-plot". The tree with the forked upper trunk (base of tree crown) in right background of the first slide and right midground of the second slide was a white oak. The dark trunk in raight margin of first slide and right foreground of second slide was a mockernut hickory.

Most of the lower (understorey) cover was of poison ivy, cat green-brier, roundleaf green-brier, and muscadine grape, but there were some tree seedlings (some baby crown cover) mostly of white oak, mockernut hickory, northern red oak, and red maple. There was not much herbaceous cover,and most of this was that of narrowleaf woodoats.

Rich mix in this tract (inside and out)- Interior (first slide) and outer edge (second slide) of an upland shortleaf pine-mixed hardwood forest in the Caddo Mountain range of southwestern Arkansas. The trunks in left immediate foreground of both slides were of shortleaf pine. There was a second adult shortleaf pine in center midground of the second slide. The tree seedling in center of immediate ground in the first (horizontal) slide was sweetgum (Liquidambar styraciflua), a species that on the forest tolerance scale was regarded as Intolerant (Wenger, 1984, p.2; Burns and Hankala, 1990). Other tree species in the first slide included white oak (right midground), red maple, and blackgum or black tupelo. Much of the remaining foliage was of poison ivy, cat green-brier, roundleaf green-brier, muscadine grape, and eastern hop-hornbeam along with seedlings and saplings of hardwood tree species (and precious little cover of shortleaf pine seedlings).

In the second slide, which was at the outer margin of this forest, besides the two prominent, adult, shortleaf pine trees (do not overlook the second one in center midground), there was a blackgum or black tupelo (large sapling or small pole) at right in the foreground, a small pole or very large sapling of northern red oak framed by the two shortleaf pines (and to their rear), an eastern hop-hornbeam to immediate left of the blackgum as well as poison ivy, cat green-brier, muscadine grape and, in lower right corner) smooth sumac (Rhus glabra). There were thus two interrupted or "broken" shrub layers: 1) taller upright shrubs like eastern hop-hornbeam and smooth sumac and 2) lower trailing shrubs (poison ivy, cat green-brier, muscadine grape) with tree seedlings. The vine climbing the trunk of the foremost shortleaf pine was cat green-brier.

There was so much cover of woody plants (trees at various age/size classes and various forms of shrubs) that there was not much by way of herbaceous growth or cover (it was "shaded out").

Hardwood stand in another tract- Local stand of hardwood tree species in an upland shortleaf pine-mixed hardwood in the Caddo Mountains of southwestern Arkansas. Tree species in this "photo-dendrograph" included mockernut hickory (crooked sapling in left foreground), northern red oak (to right of crooked mockernut hickory sapling), white oak (to right of northern red oak), and rd maple (leaves of lower crown in upper right). There was a lot of cover of cat green-brier, less cover of roundleaf green-brier, and muscadine grape plus some cover of eastern hop-hornbeam.

In this local stand the shade was so dense and constantly present that there was essentially no herbaceous cover.

At the outside edge- At the outermost edge of a shortleaf pine-mixed hardwood forest (barely under the drip line of tree canopy) vegetation seen in these two (and the two immediately following) slides had developed into a low layer of plant life. Cover in this layer of vegetation consisting of lichen, grass, seedlings of various hardwood tree species, and seedlings of shortleaf pine. Dixie reindeer moss (Cladonia subtenius) a cup lichen in family Cladoniaceae, was spread over a considerable portion of the soil surface and made up a high proportion of total plant cover. Seedlings of shortleaf pine alsomade up sizable proportions of plant cover, especially in the first slide (which served as a "photo-quadrant"), but hardwood tree seedlings (mostly of white oak and red maple) also comprised as much if not more of total planc cover (as seen in the second "photo-quadrant"). There were also seedlings of blackgum or black tupelo and, as throughout the forest community, of the ubiquitous cat green-brier.

Leaves of shortleaf pine and the hardwood species covered much of the ground surface. Pine needles were distinguishable in the second "photo-quadrant", but these needles and tree seedlings were shown at closer camera distance in the two slides following the next two slides…

Reindeer in Dixie?- Details of Dixie reindeer moss (Cladonia subtenius), a in the understorey (lowest hrbaceous layer) of a shortleaf pine-mixed hardwood forest in Fourche Mountains within the larger or overall Ouachita Mountains System. These two images were of thalluses on one of the plants shown in the immediately preceding slide (second image in the preceding slide/caption set). The various Clatonia (club lichen group) are fruticose lichens in their own family Cladoniaceae.

Lichen species are actually an amazing mutualistic symbiosis of one species of an alga and one species of a fungus that have grown so integrated with each other that the various alga-fungus "organisms" or relationships are interpreted as single species. The fundamental study of lichens is the discipline known as (what else) Lichenology.

Clatonia species are important range forage plants in diets of certain animal species, especially native caribou and the domesticated caribou called "reindeer". For this reason, Clatonia species are known generally by the common name of "reindeer moss". One of the more widespread and locally abundant kinds of reindeer moss in southern North America is appropriately designated as Dixie reindeer moss (although , of course, is not a moss at all).

Ouchita National Forest, Polk County, Arkansas. Mid-July.

More views of the outside edge- Just under (or slightly beyond) the dripline of tree crowns of an upland shortleaf pine-mixed hardwood an assembledge of plants had developed as represented by these two slides (two "photo-plots"). The first slide was a closer-up view of the forest range vegetation introduced in the immediately two-slide/caption unit. In the center of the first "photo-quadrant" Dixie reindeer moss (a fruticose lichen with hollow stalks ) was "escorted" by a seedling of white oak to its left and a seedling of shortleaf pine to its right. Needles of crowns of shortleaf pine seedlings were also in the lower left and upper right corner of this slide. Shed pine needles covered much of the ground surface not covered by lichen or tree seedlings.

The second slide featured a plant of big bluestem, frequently the dominant herbaceous species in shortleaf pine-dominated forests in the Pineywoods Region, with a big seedling of sweetgum behind it and a smaller sweetgum seedling to fore of big bluestem. Dixie reindeer moss was also present in this vertically oriented "photo-plot".

Grew back- A seedling of shortleaf pine that grew at the outer edge of a shortleaf pine-mixed hardwood forest (in the Caddo Mountains within the Ouachita Mountains chain) also happened to be within the mowed zone of a highway right-of-way. The unfortunate little pine had its upper shoot shredded off. Not to worry, shortleaf pine is one of the few Pinus species that has the ability to regrow. In this seedling the tiny branch at the next lower node/internode unit (the one immediatly below the defoliated apical bud) became "activated", grew out, and was already growing its apical or terminal bud. Regrowth capacity (ability to grow back after shoot removal or tfollowing defoliation) is an obvious and prominent adaptation to fire. Students see in these two views evolution or natural selection in action.

Miscellaneous Forest Types

151. Old growth white oak-shellbark hickory-shortleaf pine community-A bottomland site but on this sandy soil species composition is more typical of upland and mesic sites. Composite shot of the climatic or regional climax of northern portions of Texas Pineywoods. Same species composition as in previous slide. Lennox Woods, The Nature Conservancy, Red River County, Texas. May, vernal aspect. FRES No. 14 (Oak-Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Oak-Pine Series of Brown et al. (1998). South Central Plains- Tertiary Uplands Ecoregion, 35a (Griffith et al., 2004).

152. Climax bottomland White Oak-Shagbark Hickory-Shortleaf Pine Forest- The more mesic bottomlands of this forest cover type are of the oak-hickory affiliation with very little pine present. This massive old-growth white oak stands as evidence of what even the more western reaches of the Pineywoods can produce. The hat between the flutes of the trunk is a standard 4 inch brim-size so it is about a foot end-to-end. The oak is over 1 yard Diameter Breast Height. Countless thousands of white oaks such as this were logged from Texas� virgin forests for railroad ties and building timbers to help build a young nation, but many, probably most in many forests were felled for cooperage (mostly to make staves for whiskey barrels). Such is the dual nature of man. The grass understory is made up of scattered, depauperate shoots of the native bamboo (Arundinaria gigantea), longleaf uniola (Uniola sessiliflora= Chasmanthium sessiliflorum) along with Canada wildrye and various species of Panicum and Paspalum. It is meaningful from a range perspective how much herbaceous and woody understory there is in this old-growth forest, and how much feed there will be if stocking rates are kept very low or super-conservative. The Nature Conservancy Lennox Woods, Red River County, Texas. Vernal aspect, May.

The four slides presented immediately below were taken of an unusually mesic form of climax oak-hickory forest in the Boston Mountains section of the Ozark Plateau. The photographs were in the location specifically identified by Braun (Braun, 1950, ps. 170-172) as being an outlier or island of the Mixed Mesophytic Association (Braun, 1950, p.11) of the Mixed Mesophytic Forest Association but found in the Western Mixed Mesophytic Forest Association (Braun, 1950, p. 35). Braun (1950, p. 170-172) concluded that this specific forest vegetation was typical of that in the Cumberland and Allegheny Plateaus. Braun's conclusions were based on species composition, specifically of key species like beech (Table 33, p. 172) and local dominance into the climax by species like sweetgum. Beech was largely extirpated from this locale, but the combination of species mentioned by Braun including Kentucky coffeetree (Gymnocladus dioicus), American or white elm (Ulmus americana), and chinquapin oak (Quercus muhlenbergii) along with the typical sassafras, persimmon, and flowering dogwood as shrubs or understorey trees distinguished this as a unique community.

Ecologically significant by their absence were post and blackjack oaks, and even black oak (Q. velutina), this latter the dominant species and key species over much of the Ozark Plateau. Commonness of black locust (Robinia pseudoacacia), classified as Very Intolerant, along with Intolerant species like sweetgum and Kentucky coffeetree were also indicators of a "choice blend" of the oak-hickory "brand". This was further verified by presence of northern red oak (Q. borealis= Q. rubra), southern red oak (Q. falcata), shagbark hickory (Carya ovata), and bitternut hickory (C. cordiformis), one of the more tolerant hickories

Deemed by the author of substantial indicator value was commonness of wild hydrangea (Hydrangea arborescens), an understorey shrub limited to the most moist habitats such as seeps, springs, and north slopes. Relative abundance of this species and of hop hornbeam (Ostrya virginiana) along with the more typical poison oak and ivy, Virginia creeper, and pawpaw (yet nearly complete absence of herbaceous species) indicated an understorey that also varied from the typical Ozark Mountains oak-hickory forest.

The conclusion reached by Braun (1950, p. 172) was :"These isolated mixed mesophytic communities are related to past forest migrations. Their preservation here, in a region whose physiographic history is similar to that of the Cumberland Plateau, is significant."

This was an example of the point made by Braun (1950, p. 34) that each of the climax associations which characterize a specific forest region also occur in other forest regions characterized by, and thus named after, another climax forest association. This illustrated the dual nature of a Clementsian association: it was both an abstraction (abstract concept) and an actual climax plant community depending on both 1) the context in which association was applied and 2) the precise spatial and temporal location of the vegetation.

The specific forest vegetation shown in this three-slide series illustrated a forest outlier, "an area of forest separated from the main occurrence of its type generally because of some local variation in ecological conditions or past migration of vegetation associated with major climatic changes" (Helms, 1998). Braun (1950, p. 172) specified that this forest outlier was largely a product of "past forest migrations".

The following three photographs were taken on the upper terraces of the Mulberry River south of the community of Cass in Franklin County, Arkansas on a moderately steep northeast slope. July.

The closest reference for native plant communities in Arkansas is that of neighboring Missouri (two counties north of the vegetation shown in this series) by Nelson (1987) who named and described forest natural communities as to either upland or bottomland forests. These two general groups were then divided on edaphic features such as depth, soil moisture and parent material. From this base the white oak-red oak-hickory forest introduced below would be either Mesic Forest (Nelson, 1987, p. 21) or Mesic Sandstone Forest based on the geologic aspect of the sandstone-capped Boston Mountains and absence of shortleaf pine (Pinus echinata) found on Dry-Mesic Forest (Nelson, 1987, ps. 37-38.

According to the elaborate (and confusing, to this author) Natural Vegetation Classification System of Arkansas for GAP Analysis Project the Natural Terrestrial Cover of this forest was:1.B.3.a.6 Quercus alba-Carya tomentosa-C. ovata listed under Temperate Lowland and Submontane Broad-leaved Cold-Deciduous Forest. Ahh, right. The U.S. Forest Service Forest Type and Management Type Code designation was White Oak-Red Oak-Hickory and 53 for Type Name and Code, respectively. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest).

Society of American Foresters general designation was SAF 52 (White Oak-Black Oak-Northern Red Oak) (Eyre, 1980), BUT this was much less accurate than the SAF 1954 designation of White Oak-Red Oak-Hickory. The SAF (Eyre, 1980, p. 42) explained that "as hickories seldom make up more than 10 percent of the stocking, they have been dropped from the type name and black oak, a more common component, has been added". This was a true statement if applied at a landscape or regional scale (ie. across the Ozark Mountains where this type is climax according to the SAF and where black oak is a common dominant), but it most certainly is not a true statement if applied at the stand scale. The stand scale was used in the current publication of photographs and descriptions because stands-- and not landscapes or larger units-- are all that can be photographed with any detail to show vegetation. As shown below, hickories were often not only the obvious dominant but the most tolerant species and those accounting for most regeneration. As such, the SAF number was used below with the specification that hickory was co-dominant. Furthermore, as noted seven paragraphs above, black oak was not common on this Boston Mountains location but instead was generally absent from this more mesic area whose forest vegetation was an island of the Western Mesophytic Forest Association.

153. Mesic white oak-red oak-shagbark hickory forest- A stand of shagbark hickory within the specific mesic form or community indicated. Tolerance of shagbark hickory-- as for all Carya species-- is apparently open to debate. Harlow et al. (1979, p. 251) rated shagbark hickory as "moderately tolerant" while Burns and Honkala (1990, p. 222) regarded it as "intermediate". Both authorities agreed that shagbark hickory produces a deep, rapidly penetrating taproot and that younger trees of the species respout prolificly. Shagbark hickory is a minor component of six forest cover types recognized by the Society of American Foresters (Eyre, 1980) and probably of others including the more mesic Beech-Sugar Maple Type (Burns and Honkala (1990, p. 221). Local dominance by shagbark hickory throughout this specific oak-hickory forest community in the Boston Mountains was one indication that this forest vegetation was a geologic-determined remnant or relict of the more eastern and mesic Western Mixed Mesophytic Forest Association as discussed immediately above.

The trunk with the bleached-color bark in left background was one of many of the red or black oak species (subgenus Erythrobalanus) killed by an outbreak of the red oak borer (Enaphalodes rufulus). The center and foremost tree was red mulberry (Morus rubra) that, while not a rare species in the Ozarks, in combination with the other species of this community was yet another indicator of the botanical diversity and uniqueness of this specific vegetation.

Understorey species were strictly woody and included flowering dogwood, sassafras, persimmon, wild hydrangea, poison ivy, smooth sumac, and Virginia creeper. Black locust as a small tree was present just to the right of the photograph. Interestingly, and ecologically significant, was the fact that the most common tree species that was regenerating in the understorey was shagbark hickory. This indicated that this species was indeed relatively tolerant. (See also the slide below of a white oak stand in this forest-- Ozark National Forest, Franklin County, Arkansas-- where regeneration beneath large, mature Q. alba was shagbark hickory.)

Boston Mountains of Ozark Plateau. Ozark National Forest, Franklin County, Arkansas (near Cass). July. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak), but with hickory and not black oak. Oak-Hickory Series of Brown et al. (1998). Boston Mountains- Upper Boston Mountains Ecoregion, 38a (Woods et al., 2004).

154. Mesic white oak-red oak-shagbark hickory forest- Large and, thus, quite old shagbark hickories (two center, obvious trees) and white oak (two trees at far left) grew alongside an also-old sweetgum (leaves visible on far left, background trees behind and to left of hickories) for an unusual combination of species in the Ozark Plateau (Boston Mountains section). Understorey species were all woody plants with Virginia creeper dominant on the ground and with poison ivy growing up every tree trunk of any size. Shrubs throughout the community of this and other photo-plots in this series included persimmon, pawpaw, sassafras, flowering dogwood, and hop hornbeam. Wild hydrangea was common indicating the mesic nature of the general habitat. There was considerable sexual reproduction by shagbark hickory.

Locally the red oak borer had destroyed many trees in the Erythrobalanus subgenus. Oaks in this group included both northern red oak (Quercus rubra= Q. borealis= combinations of both epithets) and southern red oak (Q. falcata). Q. velutina was conspicuous by its absence in this forest community as were post and blackjack oaks, but chinquapin oak was present in small numbers and cover in localized spots.

In general, white oak was-- as is typical-- relatively more abundant on less mesic sites like south slopes while the various red/black oak species were more common on the more mesic sites, but there were many examples where all were "fully integrated" and grew side-by-side.

155. Mesic white oak-red oak-hickory forest in Ozark Plateau- Here is a "sample" of the Mixed Western Mesophytic Forest Association "lost" a "fur piece" from it's Cumberland Plateau region here in the Boston Mountains section of the Ozark Plateau. It is a remarkably species-rich community in a small "plot". The half of a trunk on far left is of shagbark hickory. The four trees to the right of it and in center background were white oak. The largest tree on the right was bitternut hickory (C. cordiformis), often regarded as intermediate in tolerance and more tolerant than it's associates (Harlow et al. (9179, p. 263; Burns and Honkala, 1990, p. 194). It will be seen that there were several lower small branches coming directly off the trunk of this large tree (leaves on these and interlacing furrows on bark made identification possible) suggesting relative tolerance in a dense forest.

Understorey species included Virginia creeper all over the ground and poison ivy growing to tops of large trees. All the usual understorey shrub/small tree species of this area grew on or close to this photo-spot, including smooth sumac, persimmon, flowering dogwood, and wild hydrangea. Hop hornbeam was least common. None of the early spring forest forbs, like mayapple for example, were visible. Grasses and grasslike plants were absent.

Boston Mountains of Ozark Plateau. Ozark National Forest, Franklin county, Arkansas (near Cass). July. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak), but with hickory and not black oak. Oak-Hickory Series of Brown et al. (1998). Boston Mountains- Upper Boston Mountains Ecoregion, 38a (Woods et al., 2004).

156. White oak stand representing the white oak phase of mesic white oak-red oak-hickory forest- Here all the large trees were white oak. The largest tree was approaching size of old-growth and was ripe for harvest. Regeneration was almost exclusively hickory, mostly shagbark and some bitternut. This hickory reproduction dominated the understorey and excluded most of the shrubs and small trees of the lower woody layers. For understorey species see captions for three slides of mesic white oak-red oak-hickory in this same forest (near community of Cass in Franklin County, Arkansas) shown above. Ecological implications of this were unknown, but in this local area the Carya species appeared to be tolerant enough to regenerate in what was obviously a dense forest and crowded understorey.

Boston Mountains of Ozark Plateau. Ozark National Forest, Franklin County, Arkansas (near Cass). July. FRES No. 15 (Oak-Hickory Froest Ecosysstem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak), but with hickory and not black oak. Oak-Hickory Series of Brown et al. (1998). Boston Mountains- Upper Boston Mountains Ecoregion, 38a (Woods et al., 2004).

157. Shortleaf pine-oak forest with a canebrake understorey, a pine-oak-giant cane savanna- This was a most unique composite vegetation. It was an oak-pine forest with tree and shrub layers that were diverse in their own rights to which was "added" a canebrake of giant or southern cane, the native bamboo (Arundinaria gigantea subsp. gigantea). There was no layer of vegetation (at least of macroscopic vascular plants) beneath the canebrake which is the typical condition for canebrakes of this species. Transeau et al. (1940, p. 762) specified that canebrakes were "often invaded" by evergreen oaks. As described in the next paragraph evergreen species in this community were pines and the oak species were not "live oaks". This community was similar, however, to that described in a standard Botany text widely used decades ago (ie. old textbook knowledge). Prior to settlement by European man, giant cane formed vast "brakes" or colonies (as in endless miles of bamboo) along water courses ranging in size from major rivers to creeks throughout southeastern North America. These seemingly infinite canebrakes occurred as far north as the Ohio River and westward to central Texas. Giant cane also grew (and still grows) as isolated plants of one or a few shoots up to small colonies of several hundred square feet as part of the understorey vegetation in oak-hickory and pine-hardwood forests. An exclusive (a homogenous or "pure") stand of uninterrupted canebrake understorey beneath an adult tree canopy was-- in this vegetation hunter's experience-- quite rare. The species composition, structure, and physiogonomy of this forest-woody grass vegetation was likely that of the climax. This was relict vegetation. Even though it was not virgin forest this was interpreted as a climax community tht lacked the component of very old trees of the species present. It was one of the most unique assemblages of native plants encountered by your photographer-author.

Shortleaf pine was the dominant tree, but there were a few individuals of loblolly pine (or perhaps of the widespread natural hybrid of these two species). Other trees included white oak, southern red oak, water oak, and sweet gum in about that order. There was a shrub layer composed of small individuals of persimmon and sassafras as well as smooth sumac. These shrubs and/or short trees grew in the canebrake, but were shorter than the taller shoots of the bamboo except at the edge of the canebrake where there was light of greater intensity and frequency. Pensacola bahiagrass (Paspalum notatum var. saurae) and hurrahgrass (P. pubescens= P. muhlenbergii= P. setaceum var. muhlenbergii) also grew on these extreme edges where they got adequate light for survival.

The development of spreading, horizonal branches on all trees, including the dominant pines that are usually strongly apically dominant, indicated that these trees had developed on a savanna which was exactly what this community appeared to be: an oak-pine-giant cane savanna. The soil was moist enough to support water oak and the scarce loblolly pine (or a hybrid thereof), but soil water conditions were not periodically ponded let alone swampish.

The grassland expression of Arundinaria gigantea canebrake (exclusive, homogenous stands or colonies of giant cane with widely scattered or infrequent trees) was covered in the Miscellaneous portion of the Grasslands section.

Miller County, Arkansas. July. West Gulf Coastal Plain portion of the Coastal Plain physiographic province. Pineywoods vegetation, part of the Southern Pine Region. According to U.S. Forest Service Ecological Divisions (Forest Service, Final Environmental Impact Statement, 1999) this was in the Southeastern Mixed Forest Province (231), Mid Coastal Plains, Westen Section (231E), South Central Arkansas Subsection (231Ea). FRES No. 14 (Oak-Pine Forest Ecosystem). K-102 (Southern Mixed Forest). SAF 76 (Shortleaf Pine-Oak, variant of). No Kuchler or SRM units for canebrake. South Central Plains - Tertiary Uplands Ecoregion, 35a (Woods et al. 2004).

158. Shortleaf pine-mixed oak-giant cane savanna or a shortleaf pine-mixed oak forest with a giant cane understorey- A more interior view of the composite pineywoods vegetation displayed in the preceding slide. Interpretation of this vegetation was subject to many subjective factors including disciplinary background or professional allegiance. It could be viewed as a mixed oak-southern pine forest with an understorey comprised exclusively of a "brake" of giant cane, the one species of native North American bamboo. Alternatively this could be viewed as a combination of diverse pineywoods forest and giant cane or bamboo grassland. Beyond any debate these two "elements" (components) were present. Also beyond debate is the ecological fact that both of these vegetational elements do occur (they formerly occurred on massive scales in the virgin vegetation) in their own forms separate and distinct from each other as potential natural vegetation. It was a different form of canebrake; it was a unique developmental expression of oak-pine forest growing as it were "out of" or in the midst of a canebrake.

The characteristic leaves of southern red oak appeared very distinctive along both the left and right borders of this photograph. Smooth sumac and shrublike or scrubby trees of persimmon and sassafras were overwhelmed in a "sea" of native bamboo.

Canebrake of Arundinaria gigantea as a grassland form devoid of other than occasional trees was presented and discussed in some detail in the Miscellaneous portion of the Grasslands section herein.

Miller County, Arkansas. July. West Gulf Coastal Plain unit of Coastal Plain physiographic province. Gorest Service ecological divisions given in preceding caption. FRES No. 14 (Oak Pine Forest Ecosystem). K-102 (Southern Mixed Forest). SAF 76 (Shortleaf Pine-Oak, variant of). No Kuchler or SRM unit for canebrake. South Central Plains - Tertiary Uplands Ecoregion, 35a (Woods et al. 2004).

159. Bottomland (floodplain) gallery oak-hickory forest- A "finger" of the eastern deciduous forest projects into the climatic or regional climax tallgrass paririe here in the Cherokee Prairie in the Osage Plains division of the Central Lowlands physiographic province. This gallery forest community is classified by the Missouri Natural Areas Committee (1987) as wet-mesic bottomland forest. It is dominated by pin oak (Quercus palustris) represented here by the largest tree with the light-colored trunk (center). Associated species also visible include: western hackberry (Celtis occidentalis), black cherry (Prunus serotina), persimmon (Diospyros virginiana), bois d'arc, red mulberry (Morus rubra), and silver maple (Acer saccharinum). Dominant shrub is Missouri gooseberry (Ribes missouriense). Herb layer is absent.

Missouri State Prairie Park, Barton County, Missouri. June, early estival aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). One riparian form or part of K-73 (Mosaic of Bluestem Prairie [K-66] and Oak-Hickory Forest [K-91]). Variant of SAF 65 (Pin Oak-Sweetgum). Central Irregular Plains- Cherokee Plains Ecoregion, 40d (Chapman et al., 2002).

160. Oak-hickory forest- Landscape scale view in central Ozark Mountains of a white oak-black hickory (Carya texana)-black oak forest that is the mesophytic or climatic climax of this western-most extension of the deciduous forest proper of eastern North America. Shortleaf pine is an associate that is locally dominant. Post oak and scarlet oak (Q. coccinea) are also common upperstory associates. Flowering dogwood, persimmon, sumac (Rhus spp.), summer grape (Vitis aestivalis), and lowbush huckleberry (Vaccinum vacillans) dominate the shrub layer. The herb layer is composed of prairie grasses and forbs of the tallgrass prairie to the west. In addition to typical prairie species, a major legume component is present including tick clovers (Desmodium rotundifolium, D. nudiflorum), wild indigo (Baptisia leucophaea), and native Lespedeza spp.

Classified as dry-chert forest by the Missouri Natural Areas Committee (1987). Christian County, Missouri. July, estival aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). Classic example of K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- White River Hills Ecoregion, 39c (Chapman et al., 2002).

161. Dormancy afforded a good "look-see"- With leaves on the ground instead of in the canopy an all-in-one view was provided of structure, arrangement, species composition, and lumber crop of a climax black oak-northern red oak forest full of mature trees. Black or Ozark pignut hickory and white or mockernut hickory (Carya texana and C. tometosa, respectively) were associate tree species. There were some post oaks, but this was clearly a forest site for the black oak species. There was also mentionable cover of black cherry, which probably indicated infrequent surface fires in this stand. Grasses were few in understorey, but dominant was broomsedge bluestem. There had been a history of cattle grazing in this stand, but it was generally light (mostly breechy ones looking for better pasture and finding worse than they left). Canopy was too dense for much herbaceous understorey other than for that of early season species like Mayapple (Podophyllum peltatum).

Many of the larger trees in this stand were over-ripe and dying or even dead. There were several snags. However most regeneration was hickory. Ottawa County, Oklahoma. FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak).

Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

162. Now a summer view- With leaves back up in the canopy a vastly different perspective was afforded of the same climax black oak-northern red oak of mature trees as introduced immediately above. There was ample regeneration of hardwoods, but hickory predominated.

Ottawa County, Oklahoma. FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak).

Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

163. Interior of the climax black oak-northern red oak forest presented in the immediately preceding two sets of two slides each. Abundant reproduction of hickory so that this stand was becoming a hickory phase or variant of the black oak cover type.

Ottawa County, Oklahoma. FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak).

Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

Forest remains: harvest of the black and northern red oaks-black and mockernut hickories forest- The tract of upland black and northern red oaks-black and white or mockernut hickories forest shown immediately above was highgrade-logged and basically clearcut in winter (and on the sly) 15 years after the preceding photographs were taken. The cutover land was re-photographed in July of the second spring-summer growing season post-harvest. Redevelopment of forest vegetation on this improperly logged, degraded tract was a combination of typical old-field plant succession beginning with pioneer or colonizing species including giant ragweed (Ambrosia trifida), horseweed or mare's tail (Conyza canadensis), hairy crabgrass (Digitaria sanginualis), and the naturalized annual, beefsteak plant (Perilla frutescens) along with climax dominants that were regenerating both sexually and asexually. Most asexual reproduction was in black oak (and some northern red oak) by coppicing from mid-size stumps. The other three species besides climax dominants that had appreciable cover were black cherry (Prunus serotina) and sycamore (Platanus occidentalis). Trees with noticeably less cover were , Kentucky coffee tree (Gymnocladus dioica), American elm (Ulmus americana), black walnut (Juglans nigra), and eastern red cedar (Juniperus virginiana). Surprisingly, the typical pioneer tree species, sycamore had less cover than would be expected judging from recovering vegetation on other severely disturbed sites in the immediate vicinity. Other tree species commonly found as pioneer or early seral plants included sassafras (Sassafras albidum) and persimmon (Diospyros virginiana). These two species were also less abundant than in the typical situation observed for disturbance in this vicinity. This cutover land had not been denuded to the extreme old-field state.

Most regeneration of dominant climax trees was of the hickory species present as saplings before logging (described and ahown above). Most reproduction of oak (especially black oak) at this point in forest recovery was by stump-sprouting. There were few oak or hickory seedlings at this stage of secondary succession.Most seedlings were sycamore, black cherry, American elm, and sassafras. Some smaller though sexually mature trees that were left standing in the clearcut as well as uncut adult trees on the perimeter of the forest (see below) served as seed trees. Otherwise, re-establishment of oaks and hickories depended on coppicing and/or the soil seed bank.

Shrubs were very important in this recovering vegetation. Far and away the most important shrubs were blackberry (probably
including several Rubus species) that formed immense thickets that were summarily invaded by this author adequately equipped with buckets at berry picking time. Another important shrub (though less productive of tasty fruit) was smooth sumac (Rhus glabra). Fox grape (Vitis vulpina) and summer grape (V. aestivalis) had spread quickly on the cutover land. Eastern redbud (Celtis canadensis) and flowering dogwood (Cornus florida) that had already been growing in this forest remained minus minimal mechanical damage.

This remarkably diverse recovery vegetation also included numerous species of forbs, both native and naturalized. The latter included the biennial, flannel mullein (Verbascum thapsus), and introduced legumes, both perennials such as white clover (Trifolium repens) and red clover (T. pretense) and annuals like Korean or Japanese lespedeza (Lespedeza striata). Other forbs included numerous natives like biennial evening primrose (Oenothera biennis); pokeweed (Phytolacca americana); and various composites, especially giant ragweed (Ambrosia trifida), common ragweed (A. artemisiifolia), Carolina elephant foot (Elephantopus carolinianus), Baldwin ironweed (Vernonia baldwinii), black-eyed susan (Rudbeckia hirta), Canada wild lettuce (Lactuca canadensis), and wingstem crown-beard (Verbesina alternifolia). The pioneering composite, horseweed or mare's tail was widespread and locally dominant although generally not in dominating proportions that might have been expected (and, probably, present in the first season following harvest). There was also some immature smartweed or knotweed plants (Polygonum sp.) that could not be identified as to species along with toothed spurge (Euphorbia dentata). Pokeweed was the most widespread and overall most important forb, but giant ragweed was a close second forming dense stands from which almost all plants of other species were excluded.

The only common grass was hairy crabgrass. The perennial grasses were represented almost exclusively by broomsedge bluestem (Andropogon virginicus). Poverty oatgrass (Danthonia spicata) already present in the understorey persisted by larger stumps where it was not "overrun by pioneer species. There was also an occasional plant of purpletop (Tridens flavesus) next to shelter (eg. uncut tree).There were also some plants of green and/or yellow foxtail (Setaria viridis, S. glauca, respectively) which could not be distinguished at vegetative stages present at time of photographs. Likewise there were incidental plants of Carex and Cyperus species.

White-tailed deer where the only species of large vertebrate that had access to this was black oak-northern red oak forest which had not been grazed/browsed by cattle or hogs for several decades.

In the climax oak-hickory forest that approached old-growth state there were very few plants available for grazing and/or browsing in the understorey. These were limited to leaves and buds on regenerating oak and hickory species, flowering dogwood, poison oak/ivy, and incidental plants (trace amounts in absolute and relative cover) of blackberry, poverty oakgrass, and sedge.

This tract of black oak-dominated upland Ozark Plateau forest was clearly transitory forest range. Once (after) canopy cover reached almost 100% almost no light that could penetrate throught the dense foliage to reach the ground (soil) surface. After leaves were fully grown in the forest canopy each spring light could only reach to the level delineated by lower leaves on shrubs like flowering dogwood and saplings of oak, hickory, black cherry, etc. In this climax oak-hickory forest with mature ("over-ripe") trees spring forbs like mayapple (Podophyllum peltatum) were not present except in natural forest gaps and at forest edges.

Organization Note: a black oak-pignut or mockernut climax forest that served as permanent forest range was presented and described in the Use and Abuse of Oak-Hickory Forests section of Oak-Hickory Forests-II.

Harvest of this forest was an example that socioeconomic factors often override biological ones, and that many if not most human endeavors (not excluding silvicultural operations) involve human emotions including greed and jealosy. Logging of this oak-hickory forest was a case of timber theft. A thieving son snuck in and stole this standing timber from his aged father who for romatic and aesthetic reasons wanted the forest with its many mature trees left as it was. The conniving son left uncut the trees around the edges of this forest to hide what he was doing inside the tract. The timber-buyer left some of the smaller trees inside the perimeter that he felt were not worth felling (probably to the chagrin of the greedy son who worked up all the slash to sell as fuelwood). Not only was this dysfunctional family relations, but it was also improper forest practices: cut-and-run, sloppy, (and illegal) logging of the worst form, the kind that gives the forest products industry a bad reputation. Thief of "free grass" and "timber for the taking" was standard fare in early history of use and abuse of forest and rangeland resources. The important thing for rangemen and foresters to bear in mind is that these resources are, to large degree, renewable (even with improper harvest methods and other forms of abuse).

The following series of slides was of the black and northern red oak-black and/or Texas and mockernut hickories upland forest in the second growing season following highgrade logging that was, in effect, land-clearing.

164. Missing trees, stolen timber- The second-growth black oak-dominated (northern red oak, associate) forest shown and described here was secretly clearcut with a few smaller trees left around the perimeter so that a son could hide the timber theft from his aged father. The crime scene and second-year successional vegetation was presented in two photographs that gave a general view of the cleared forest. The wasted sound logs in the first of these slides added resource waste and abuse to the weed patch effect. In both slides remaining trees (a young northern red oak in the first; black and northern red oaks, Texas or Ozark and mockernut hickories in the second) "looked on" cutover land supporting a pioneer stage of recovering forest vegetation dominated by such colonizing species as the annual composite, giant ragweed, along with common ragweed, hairy crabgrass and such disturbance-loving perennials as pokeweed, black-eyed susan, Baldwin ironweed, wingstem crown-beard , Carolina elephant's foot. Foreign weeds that benefitted from this disturbance included flannel mullein and beefstake plant. Common shrubs included several species of blackberry, summer and fox grape, and smooth sumac.

This stage of secondary plant succession on a black oak-dominated Ozark Highlands upland forest was about half-way into the second warm-growning season following clearcutting of a second-growth forest that approached old-growth status.

Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak).

Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

165. Into the weeds (and the ticks and chiggers)- Two close-in views of seral vegetation on a forest site that two winters or one an a half warm-growing seasons before had been a climax black oak-northern red oak (dominant and associate species, respectively) upland forest that was harvested by clearcutting (in effect a land-clearing operation). The most abundant herbaceous species were giant and common ragweeds. Other composite forbs included black-eyed susan, common horseweed or mare's tail, an Carolina elephant's foot. Pokeweed and flannel mullein represented native perennial and naturalized Eurasian biennial forbs, respectively. Passionflower (Passiflora incarnata), a perennial herbaceous vine or twining forb, was conspicuous along right margin (about half-way up) in the first slide. The only grass with cover and density worthy of note in the seral range vegetation presented in these two photographs was hairy crabgrass.

Shrubs included blackberry, smooth sumac, and summer and fox grape. Tree species present as pre-existing (present before logging), stump sprouts, or seedlings included black oak, northern red oak, post oak, Texas or Ozark hickory, mockernut hickory, black cherry, American elm, red elm, sassafras, and persimmon. Young trees (either too small to make a saw log and/or serving as a cover to conceal the crime sceene from outside the clearcut) included all of the oak and hickory species as well as a few black cherry.

Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak).

Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

166. Edge effect, a benefit of forest harvest- Two more general views of a clearcut in a black oak-dominated climax forest in the Springfield Plateau. These two photographs high-lighted the effect of edge, the union of standing trees at perimeter of the now clearcut forest and seral range vegetation developing on the clearcut through secondary plant successtion midway through the second warm-growing season post-harvest. Some of the more abundant and conspicuous herbaceous species on this clearing included native forbs common to disturbed habitats: smartweed, Baldwin ironweed, pokeweed, giant ragweed, common ragweed. Most of these were of little or, at least, very limited forage value. The forb of most forage value was red clover, a naturalized, Eurasian, perennial legume (barely visible in the first photograph as pink clusters). How such abundant cover of this valuable forage plant developed by the second growing following logging remained a mystery. Farmers in this local vicinity have overseeded permanent pastures to red clover.

Clearcutting had converted a climax forest with limited understorey (most of that regenerating hardwoods, especially hickory species) into a cutover pasture or range that, though a far cry from standards of high-quality tame pasture, provided valuable forage plants for livestock and wildlife including an introduced, perennial legume; a palatable, annual grass, and forbs of diverse palatability). This was transitory forest range.

The second of these slides featured the numerous woody species that invaded the clearing the first growing season following clearcutting. Most conspicuous were two-year-old seedlings of sycamore (center of photograph). There were stump sprouts and seedlings of black oak, Ozark and mockernut hickories and black cherry. Most of the latter were seedlings or pre-established saplings. Prominent forbs in successional range vegetation shown in the second slide included Baldwin ironweed, giant ragweed, and common ragweed. Young vines of summer and fox grape trailed and wound their way around newly established large patches of blackberry out of which grew black cherry, sassafras, and elm (both American and red).

In this arrangement of seral vegetation there were edges within edges:. edges where blackberry patches met weed (eg. colonizing composites, pokeweed, flannel mullein) patches within the edge that was the perimeter of the climax black oak-red northern oak forest. This was shown in more detail in the next set of two slides.

In Nature any disturbance--no matter how traumatic or disturbing of existing species, communities, ecosystems, landscapes, etc--is a boon to some other species, communities, ecosystems, and so on. Clearcutting the climax, near old-growth black oak-northern red oak forest that occupied this site was an extreme perturbation, a drastic alternation of that forest ecosystem, that had dire consequences on many living things ranging from old trees and their dependent fungi species, squirrels, pileated woodpeckers (Dryocopus pileatus), and humans due to dissolutionous family relations between a father and a theiving son. That action, that extreme ecological disturbance, also created greately improved habitat for white-tailed deer and northern bobwhite (Colinus virginianus).

One of numerous reasons why populations of bobwhite quail have declined throughout much of the Ozark Plateau Region is that their prime habitat of old-fields, recovering cleared forests, and small farm fields had revegetated back (had secondary plant succession progress closer toward) the pre-white man forest. Advanced seral, subclimax, and climax stage oak-hickory forests provide marginal to poor habitat for bobwhite. "Setting back" forest vegetation to pioneer and other early seral stages was (is) of immense benefit to seral species like bobwhite quail and white-tailed deer. When this climax forest (with its high proportion of rotten trees that were "overripe" for quality hardwood lumber) was logged pileated woodpeckers that lived off of insects that thrived on decaying wood as well as cavity nesting mammals including squirrels, 'possums (Didelphis virginiana), and coons (Procyon lotor) came out "loosers", at least in the short run. Critical parts of their habitat were eliminated. Perhaps this was the situation for wild turkey (Meleagris gallopavo) and raptors like great horned owls (Bubo virginianus) and red-tailed hawks (Buteo jamaciensis), all species that commonly lived in the black oak-northern red oak forest with its old age, over-mature trees. However, turkey and coon soon benefitted from increased production of blackberries, pokeberries, etc. Reduction of canopy cover facilitated predation by raptors although owls are still going to have to find dense tree canopy to escape their arch enemy, the annoying, sleep-depriving crow (Corvus brachyrhynchos). The 'possum will miss her favorite hollow tree, but see what a crop of persimmons will be produced for her descendants in a few years.

The edge where uncut, timber theft-blocking trees meet and merge with the new weed and brush patch provided a new corridor as well as ample cover for new feed sources for various species of animals and new growing environments for plants. Use of natural resources always producers "winners"and "loosers". Wise use to conservation prophets like Gifford Pinchot consisted of finding that "blend" which, in the spirit of Jeremy Bentham's utilitarianism, "provided the greatest good to the greatest number for the longest period of time". As was shown in later descriptions and discussions of clearcutting this climax black oak-forest, it was quite likely that in the "long run" Pinchot's standard for wise use was fulfilled on this land, in spite of the villainous crime committed by a treasonous son against the rest of his family.

Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak).

Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

167. A closer look at edges and species make-up of seral vegetation- Two photographs showing seral forest range vegetation about half-way through its second warm-growing season growing adjacent to adult black and northern red oak trees that were left uncut to hid from view a forest clearcut of a climax black oak-dominated upland forest in the western Ozark Plateau. The plant communites that developed along the uncut forest perimeter occured spatially as "rows" (narrow zones) of woody plants (both tree saplings and shrubs) while there were larger patches of "weedy" colonizing species interior to the "belts" of wody vegetation.

In the first photograph a narrow belt of adult black and northern red oaks was in the background while the foreground was an area dominated by giant and common ragweed. The bulk of the recovering forest vegetation (midground of photograph) was a blackberry patch with numerous saplings of black cherry, American and red elms, and sassafras (less persimmon) along with fox and summer grapes. The second photograph also showed a "weed patch" that was almost exclusively giant ragweed with some common ragweed and mare's tail orhorseweed (foreground) contiguous with a zone or "natural row" of blackberry patch with tree saplings and shrubs the major ones of which were smooth sumac, the two grape speceis, black cherry, American elm with some climax hardwoods (more hickories than black or red oaks).

Such strips of seral woody vegetation are almost impenetrable to humans as the blackberry picking photographer attested. This is bobwhite habitat par excellence. Even pointers experience some problems getting through such brushy barriers.

Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak).

Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

168. Sere colonizers, both pioneer newcomers and climax holdovers- Two close-up "photo-plots" of seral range vegetation on a climax black oak-dominated upland forest site in wesstern Ozark Highlands that had been clearcut two winters previously. Recovering forest vegetation in these photographs was about half-way through the second warm-growing season after logging. The plant community of this pioneer or initial seral stage was a combination of colonizing species (giant ragweed, common ragweed, mare's tail or horseweed, hairy crabgrass, and beefsteak plant) native perennial composite forbs (Baldwin ironweed and Carolina elephant's foot), biennial forbs (flannel mullein and biennial evening primrose), native shrubs (blackberry, smooth sumac, summer and fox grape, Virginia creeper), colonizing trees or generally less tolerant tree species (sycamore, black cherry, sassafras, American and red elm), and seedlings and stump srpouts of climax trees (black and red oaks, Ozark or Texas and mockernut hickories). An amazing array of plant species in small local habitats.

Seral range vegetation shown in the first slide consisted of a weed patch dominated by giant ragweed with Canada wild lettuce, biennial evening primerose, and greeen and/or yellow foxtail mixed with woody species ranging from black oak and hickory seedlings and stump sprouts to summer grape that had developed in front of an edge of woody vegetation made up of taller shrubs and young tree saplings. Shrubs included blackberry, smooth sumac, fox and summer grapes while saplings ranged from black oaks and two hickory species to American and red elms, black cherry, and sassafras. This was another example of the edge where different plant communities meet producing the edge effect that is so important for certain species of wildlife. Northern bobwhite quail was a species of great local importance that benefits immensely from this kind of seral range vegetation..

The second slide was of seral forest range vegetation dominated by forbs and seedlings and stump sprouts mostly of black oak and the two hickory species. Conspicuous forbs included giant ragweed, horseweed or mare's tail (many of which had upper parts of shoots grazed off by white-tailed deer), Canada wild lettuce (also grazed by deer though less so than mare's tail), toothed spurge, and biennial evening primrose. Red clover was also present but less conspicuous in spite of its showy inflorescneces than the taller-growing "weedy" colonizers. Tha annual grasses, airy crabgrass and green or yellow foxtail, were the representatives of the Gramineae in this photograph There was considerable cover of young Virginia creeper. The tree trunk in center midground was a black oak. Sprouts surrounding this black oak were mostly Ozark and mockernut hickories. Also in this slide was a hollow butt portion (roughly one ana a half foot diameter) of a black oak log that was later used for fuelwood.

Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak).

Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

169. Forgiving oaks- Four examples of stump sprouting or coppicing in black oak on a black oak-northern red oak upland forest in the western Ozark Highlands (Springfield Plateau section) midway through the second growing season for hardwood species following clearcutting. Coppice was defined by the Society of American Foresters (Helms, 1998) as: "1. the production of new stems from the stump or roots 2. to cut the main stem (particularly of broadleaved species) at the base or to injure the roots to stimulate the production of new shoots for regneration 3. a plant derived by coppicing 4. any shoot arising from an adenvtitious or dormant bud near the base of a woody plant that has been cut back". All four meanings were appropriate for the morphological/physiological phenomenon presented in the four black oak stumps and their second-season suckers. Helms (1998) also provided: "stool- a living stump (capable of) producing sprouts".

Coppice shoots or stools (and related forms of vegetative reproduction including water shoots, suckers, and sprouts) are, of course, clones, ramets, modules or modular units of the parent plant, genet, entire vegetative plant, respectively. A simplified explanation is that stump sprouts are natural grafts or naturally formed scions arising from the stock (the stump or root).

Forest regeneration by coppicing is a major silvicultural system (or component treatment thereof) for certain hardwood species. Black oak has not generally or typically been viewed as a coppice species or black oak lumber as a coppice crop to the extent as for some other species (eg. black cherry), but on the clearcut described in the above captions a high proportion of the felled black oak did coppice (sprout back from living stumps). In fact, this is the usual regeneration response of black oak, at least of the younger trees. Stump sprouting in black oak was described in Silvics of Trees of the United States (Fowells, 1965, p. 560; Burns and Honkala, 1990, Vol. 2.p. 747). The latter reported that roughly 95% of black oak regeneration in clearcutting was by sprouts from either stumps or "advance reprodution" (new sprouts from dormant buds near juincture of shoot and root).

An interesting inverse relationship exist between stump size and successful stump sprouting. Shoots (sprouts= stools) from black oaks that were larger at harves grow faster than those of smaller cut trees; however, stump sprouting was inversely related to stump size, tree age, and forest site with larger stumps and those of older trees (often "same difference") having reduced sprout perduction. In other words stumps of bigger trees (ie. larger diameter stumps) and, hence, usually older trees are less apt to sprout, but if they do sprout these suckers (stump shoots) grow faster (have more rapid grow rates) than suckers from smaller stumps. This is a common--if not nearly universal--physiological response in hardwood species.

Black oak stump-sprouts less readily than norther red oak which, as on this oak-hickory forest, is commonly associated with black oak. Fowels (1965, p. 591) cited research that reported over two-thirds of black oak reproduction in the Missouri Ozarks was "of sprout origin". Nonetheless, coppicing is the best possible means to replace 1) "original" (genetically identical) trees and shrubs and 2) climax woody species. In this regard, sawing logs amounts to the same thing as mowing shoots of perennial forage species for hay. With properly harvest the species composition of the tree component of the clearcut forest is essentially the same as it was before logging.

In the instance of the upland Ozark Highlands black oak-dominated forest that served as the example for this lesson there was a cruel irony to the clearcutting operation in which a theiving son stole timber from his trusting father. The criminal act of forest harvest actually benefitted the black oak-northern red oak forest. The felled trees were still of an age and/or size that a high proportion of them coppiced (stump-sprouted) resulting in regeneration of both 1) the climax tree species and 2) reproduction of the same genetic individuals (ie. restoration of the identical harvested trees). Obviously the shoots (trunks= boles) of the mature (actually, over-mature) trees will not be restored because they were hauled of to the sawmill for pallats and framing lumber, but genetically these identical trees will regrow (grow back) so as to be be replaced as they were before forest harvest. There will be the same original root systems and basal trunks, the stumps, but morphologically new trunks.

Based on canopy cover the proportion of black and northern red oaks in the recovering (seral) forest was less than in the climax forest, but climax tree species (and the same individual trees) were in the seral forest vegetation from the initial (pioneer) stage of plant succession (forest restoration). With asexual reproduction (coppicing) climax trees were present in the initial plant community following clearcutting rather than entering the forest sere later in the sequence of secondary plant succession as would be necessary if climax tree establishment depended solely of sexual reproduction (production of seedlings). Net result will be (barring other or continued severe forest disturbances) a more rapid return to the terminal stage of plant succession (ie. a shorter time interval to replace the climax black oak-northern red oak forest).

Ironically, if the near old-growth black and red oaks (many of them already "overripe" with trunks partly hollow with heart rot) had remained standing for more years most of them would have been too old to coppice. Thus regeneration of the climax oak species almost assuredly would have been much lower and slower. Sexual reproduction (seedlings from acorns) of black and red oak is much less likely (much lower probability of successful tree establishment) than is asexual reproduction by coppicing. Furthermore, recall (from photographs and descriptions of the unlogged, nearly old-growth forest) that most tree reproduction in this climax black oak-dominated forest was of the associates, Ozark hickory (a small tree at maturity) and, secondly, mockernut hickory. In this forest of mature oak trees and in absence of natural disturbances like windthrow (blowdown) and fire the climax oak species were, through a combination of natural death due to old age and lower rates of regeneration, being replaced by replaced by hickory species.

Clearcutting this black oak-dominated forest resulted in 1) regeneration of the climax oaks along with that of the associate hickory species, 2) increased plant biodiversity due to a combination of regeneration of climax species and colonization by pioneer ("weedy") species, 3) increased forage and browse production for livestock and deer, 4) improved habitat for bobwhite quail, and 5) caused loss of habitat for plant and animal species dependant on climax forest vegetation.

170. The black oak cover type- Interior view of dry chert upland forest dominated by black oak with post oak, northern red oak, blackjack oak, and black hickory as associates. There are two obvious shrub layers: 1) an upper one dominated by flowering dogwood (State Tree of Missouri; conspicuous here), redbud (State Tree of Oklahoma), and shadbush= eastern serviceberry (Amelanchier arborea) and 2) a lower one dominated by buckbrush= coral berry (Symphoricarposorbiculatus) and blackberry. Wild grape and Virginia creeper (Parthenocissus quinquefolia) form an aboreal shrub layer while poison ivy (Rhus radicans = Toxicodendron radicans) occurs in both shrub layers ranging from lianas extending to tops of trees to non-climbing thickets.The herb layer is usually limited to early spring species that complete their annual cycle before greening of the forest canopy. Mayapple (Podophyllum peltatum) is the conspicuous dominant herb. Head of hollow on a chert upland. April, early vernal aspect.

Ottawa County, Oklahoma. FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak).

Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

IMPORTANT: As a general rule, browsing animals find deciduous shrubs and trees considerably more palatable than coniferous ones with the general response being that deciduous forests are much more prone to suffer damage, especially retarded regeneration, by overbrowsing than are coniferous forests. Understandably, foresters are reluctant to recommend (typically adamantly oppose) stocking livestock on hardwood forest types.Swine with their incessant rooting and feeding on mast are the livestock species that cause the most damage to these remarkably fragile range types.Proper livestock stocking rates on hardwood range are those described generically as “conservative”. These are forests that are usually most valuable for watershed and whose main crop or commodity is wood.HARDWOOD FORESTS ARE NOT "STOMP LOTS"!

171. Dormant but healthy- An Ozark Plateau upland forest (south slope) co-dominated by black oak and post oak that had received no livestock grazing for decades supported various age classes of trees. Other major trees included northern red oak, black hickory, black walnut, Kentucky coffeetree (Gymnocladus dioicus) and black cherry (Prunus serotina). Typical understorey shrubs included redbud, flowering dogwood, and woody vines such as various species of greenbriar and grape along with Virginia creeper and poison ivy (oak). This forest stand was so dense and had a nearly closed canopy so as to exclude development of an herbaceous understorey other than for early growing season species like mayapple.

In most precise terms, the potential natural vegetation of this tree-dominated plant community was more woodland than forest per se. Climax vegetation would most likely consist of a more open or incomplete canopy cover (ie. tree crowns would not be interlocking). Nelson (1987, 2005) made a rational, well-written distinction between forest and woodland vegetation of the Ozark Plateau. The stand of black and post oak-dominated vegetation described here and immediately below were Dry-Mesic Chert Woodland (Nelson, 2005, ps. 190-193).

Otawa County, Oklahoma. January. An upland forest of mixed oak and hickory species, but given overall dominance of this and adjoining forest stands the Society of American Foresters (Eyre, 1980) cover type that most closely fit this forest vegetation was Black Oak (SRM 110). Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Chapman et al., 2002).

172. Also dormant but not healthy- An Ozark Plateau upland forest (south slope) co-dominated by black oak and post oak that had been grazed by beef cattle for decades. This stand (if that term could be used loosely for comparitive purposes) was about 150 yards down a county road from the stand shown in the preceding slide. In addition to mature black oak, post oak, and, fewer, northern red oak (some of each species were on the ridge crest in background) there was a pole-size black walnut. There were also numerous and very conspicuous seedlings to small saplings (say, two to eight years in age) of eastern red cedar, an eastern juniper (Juniperus virginiana). There was zero regeneration of hardwoods of any species including the strongly smelling, usually unpalatable black walnut.

This stand was a degraded Dry-Mesic Chert Woodland (Nelson, 2005, ps. 190-193) with potential natural vegetation for this forest site being an open or sporadic (vs. closed or complete) canopy of an actual forest having interlocking crowns. Foresters and rangemen would still management this as a stand of hardwood trees capable of producing high-quality oak, hickory, black walnut lumber as as having a grazable understorey for light stocking of livestock and habitat for wildlife including white-tailed deer, bobwhite quail, and squirrels.

Ottawa County, Oklahoma. January.

173. Older hardwoods and younger cedars (or Where are the young hardwoods?)- The same "stand" of Ozark Plateau upland forest (south slope) co-dominated by black oak and post oak as presented in the immdeiately preceding slide. Large, two-trunked tree in center foreground was northern red oak. Extreme overrgrazing/overbrowsing for unknown decades (probably half a century or longer) had prevented regeneration of hardwood species. This management of a hardwood stand as a "stomp lot" had taken place over such a long time that even pole-size trees were absent from these "woods".

Overgrazing/overbrowsing by cattle had not only been responsible for failure of hardwood reproduction, but this poor forestry (and animal husbandry) practice had also prevented periodic surface fires in what would otherwise have been an oak-hickory-walnut forest. Removal of essentially all herbage and prevention of production of fine woody material pre-empted light forest burning (ie. there was no fuel). Meanwhile birds that had eaten the fleshy seeds of eastern red cedar on rocky north slopes and bluffs above a nearby creek perched in and defeacted cedar seeds from the mature oak and black walnut trees. This avian behavior resulted in establishment of young cedar seedlings and saplings in the understory. In absence of fuel for light surface fires eastern red cedar was becoming established as the new forest cover type (SAF 46, Eastern Redcedar). Cattle will not eat eastern red cedar even inside corrals (or "cowpens" as such enclosures are called by many Ozark hillfolk). Barring disease these eastern junipers are safe-- at least until they become so large and close together (adequate canopy cover) that an accidental fire can spread almost instaneously through their crowns (ie. a crown fire, which is what any self-respecting rangeman would be hoping for in this degraded forest range site).

This is horrid mismanagement of resources resulted in anthropogenic vegetation that was textbook case of grazing disclimax. In fact, this stand fit perfectly the description of the Eastern Redcedar forest cover type (SAF 46) by the Society of American Foresters (Eyre, 1980, p.50-51).

Ottawa County, Oklahoma. January.

174. "In the Good Ole Summertime" but still "sick"- Sleek cattle and green leaves do not change the fact that this should-be or one-time forest is a degraded plant community (again, notheing but a "stomp lot"). What should be an oak-hickory forest with miscellaneous hardwoods such as black walnut, black cherry, and Kentucky coffeetree become a degraded pasture of mostly Eurasian annual grasses and a few mature trees of climax species from the previous forest stand. Cattle through overgrazing and overbrowsing prevented regeneration of the hardwood trees and also precluded light surface fires that would have killed these non-sprouting junipers and benefitted former climax understorey grasses.

Yes, it is true that a climax or late seral stage oak-hickory forest like the stand that was about 150 yards down the road from this "mess" (and that was used as the control plot to present this lesson) would have little grazable/browsable understorey. A comparison of that late successional stage of forest vegetation with the "cow pasture" shown here would suggest to the neophyte that there is more "cow feed" on this degraded former forest. That is not true, not the case at all. The near-climax forest of the control plot is ready for logging. Following harvest of oak, hickory, and walnut logs, native grasses (big bluestem, Indiangrass, and purpletop are the main ones) and numerous shrubs (including blackberry, sumac, buckbrush, and wild plum) as well as regenerated hardwoods (mostly seedlings with some stump sprouts) will soon become re-established and provide range forage and browse for livestock and wildlife (notably white-tailed deer and bobwhite quail). Most importanlty from a forest perspective is the fact that the wood crop (hardwood logs are the agricultural commodity) is a source of revenue along with cattle (feeder calves and cull cows) and wildlife (either as recreational products or a sources of income from egress fees).

Over the longterm, a properly managed oak-hickory forest will generate more revenue and produce more resources and commodities than this degraded barnyard with shade trees. Even the latter will eventually die to be replaced by juniper which will be a fire hazard by that time. Of course, agricultural producers are the world's greatest and most sustained optimists. In that spirit one can always hope that the eastern red cedar will escape crown fires and live long enough to grow into lumber that will fetch a fair price and that can be made into fragrant cedar chests to sell to tourists in flea markets.

If this owner wanted nothing but pasture for these cows and calves the proper farming practice would be to plant this land to introduced (agronomic) pasture grasses like bermudagrass (Cynodon dactylodon) or tall fescue (Festuca arundinacea), both of which are well-adapted to these shallow rocky Ozark hills. The owner could then properly manage this tame pasture for economical production of beef cattle. Instead and as it was this landowner had nothing but "bragging rights" to running some cows and a lot less income than if he had wisely managed his forest, range, and livestock resources.

This joker had not done justice to the revered title of "hillbilly" (just plain "hick" about covered it).

Ottawa County, Oklahoma.

175. Some respectible-size hardwoods on a rock pile- A climax oak-hickory forest with black oak (two foremost trees on the left; two left foreground trees) the most common tree species along with black hickory and bitternut hickory (Carya cordiformis) and even one blackjack oak (rightmost larger tree). The large tree with the high scar (tree behind foremost tree on left) was a bitternut hickory with a DBH of 24 inches. Not much herbaceous understorey but big bluestem and broomsedge were main species. Flowering dogwood (left margin; just coming into bloom) was major species of the upper shrub layer. A second or lower shrub layer consisted of buckbrush, blackberry, and Virginia creeper, this latter of which covered much of the ground surface and also reached up into tree crowns so as to be in both shrub layers. Grape (right foreground) also extended in both shrub layers.

Very marginal land (Land Capability Unit #8).

April, early vernal aspect. Ottawa County, Oklahoma. FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak).

Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

White Oak (Quercus alba) Forests

White oak is one of the most widely distributed Quercus species in North America. It is also a widespread dominant or associate species being a major and defining member of several of the climax forest regions of Braun (1950, ps., 35-38): Western Mesophytic, Oak-Hickory, Oak-Chestnut, and Oak-Pine. Forest cover types in which white oak was co-dominant, especially with a conifer (eg. white oak-shortleaf pine, white oak-loblolly pine) or was only an associate species, were treated separately from this short section which was devoted only to cover types White Oak (SAF 53) and White Oak-Black Oak--Northern Red Oak (SAF 52).

176. The white oak cover type- As stated by the Society of American Foresters (1980), the white oak forest cover type is "pure". In classic Clementsian terms this primarily a consociation (certainly in the photo-plot presented here). Seen here is a stand of vigerous young white oaks on a moist north slope in the Missouri Ozarks. The dominant herb is the widespread composite, black-eyed susan (Rudbeckia hirta). Also visible is the unique natural spiderwort hybrid (Tradescantiaozarkana X T. ernestiana). The main shrub growing amidst the oaks is flowering dogwood.

Roaring River State Park, Barry County, Missouri. May, vernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). White Oak form of K-91 (Oak-Hickory Forest). SAF 53 (White Oak). Quercus alba Association (if recognized as such) in Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Dry-Mesic Chert Forest (Nelson 2005, ps. 125-130). Ozark Highlands- White River Hills Ecoregion, 39c (Chapman et al., 2002).

177. Members of an upland white oak-dominated forest- A species-rich upland forest community had developed on this upland Ozark Highlands location. At some local sites white oak formed a consociation. At other local sites white oak was joined by shagbark hickory (Carya ovata) and sugar maple (Acer saccharum) as associates. Other tree species included black oak, northern red oak, post oak (of course as nearly always present), western hackberry, sycamore, and honey locust (in that approximate order). Flowering dogwood comprised most of a lower woody layer other regenerating young trees of above listed species. An herbaceous layer at this particular location included shooting star or American cowslip (Dodecatheon meadia) and Christmas fern (Polystichum acrostichoides) along with the hybrid spiderwort specified in the preceding caption.

Roaring River State Park, Barry County, Missouri. May, vernal aspect.FRES No. 15 (Oak-Hickory Forest Ecosystem). White Oak form of K-91 (Oak-Hickory Forest). SAF 53 (White Oak). Quercus alba Association (if recognized as such) in Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Dry-Mesic Chert Forest (Nelson 2005, ps. 125-130). Ozark Highlands- White River Hills Ecoregion, 39c (Chapman et al., 2002).

178. Deep in "them thar hills"- White oak-dominated upland forest in Ozark Mountains.Associates of white oak were (locally or at local site scale) shagbark hickory , sugar maple, black oak, and the ever-present post oak. Flowering dogwood was present throughout as the principal shrub species though it was not in bloom during this mid-spring season. Redbud was also present, but it was much less common than flowering dogwood. In these two "photo-plots" herbaceous were sparse and limited mostly to the hybrid spiderwort noted above.

Roaring River State Park, Barry County, Missouri. May, vernal aspect.FRES No. 15 (Oak-Hickory Forest Ecosystem). White Oak form of K-91 (Oak-Hickory Forest). SAF 53 (White Oak). Quercus alba Association (if recognized as such) in Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Dry-Mesic Chert Forest (Nelson 2005, ps. 125-130). Ozark Highlands- White River Hills Ecoregion, 39c (Chapman et al., 2002).

179. Scond-growth white oak et al. forest with amazing species diverstiy- At head of a hollow deep in the Ozark Plateau white oak dominated a north slope of a Dry-Mesic Chert Forest (Nelson, 2005, ps. 125-130). Associate tree species were of the red or black oak group (Erythrobalanus subgenus): black oak and northern red oak. Hickory was common as was (though less so) black walnut. Sugar maple was also present in more mesic microsites. Dominant large woody understory species (large or tall shrub layer) was flowering dogwood. Understorey shrubs of a second or lower layer included wild hydrangia, blackberry, poison oak, grape, Virginia creeper, and lowbush huckleberry (Vaccinium vacillans). The most abundant grass was hairy, downy, or silky wildrye (Elymus villosus) while the dominant forb was pokeweed (Phytolaca americana). Several species of tickclover (Desmodium spp.) were also widely distributed.

Flag Springs State Park, McDonald County, Missouri. June, late vernal aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Most plausible Society of American Foreters (Eyre, 1980) cover type was SAF 52 (White Oak-Black Oak-Northern Red Oak) and this was predominant cover type of this locality: white oak was not as much as obvious dominant and defining species as was case for SAF 53 (White Oak) which this closely resembled. Oak-Hickory Series of Brown et al. (1998). Dry-Mesic Chert Forest (Nelson 2005, ps. 125-130). Ozark Highlands- Elk River Hills Ecoregion, 39b (Chapman et al., 2002).

180. White oak-black oak-noethern red oak forest- White oak clearly dominated this north slope of a western Ozark Plateau forest, but it was not cover or density domiance to the degree that white oak dominated the north slope shown in the preceding set of slides (Roaring River State Park, White River Hills Ecoregion). Development and species diversity of two prominent woody layers below canopy layer was greater and different from that of a more overwhelming dominance by white oak as described for the immediately preceding Ozark Plateau forest.. This north slope and forest hollow was substantially more mesic and had less light exposure than the preceding forest.

Flowering dogwood was the dominant of the taller shrub layer. A lower and more diverse shrub layer included wild hydrangia, blackberry, poison oak, grape, Virginia creeper, and lowbush huckleberry. The herbaceous layer(s) was not as diverse as the white oak-dominated north slope forest described above, but the grass component (mostly downy, silky, or hairy wildrye) was much more productive and of far greater foliar cover on this mixed oak Ozark forest. Most common forb was pokeweed (which is more commonly a forest forb on locally disturbed areas).

181. White oak-shortleaf pine forest- White oak is frequently the dominant oak in the shortleaf pine-oak type (SAF 76), especially on more mesic forest sites. By definition and description dominated more by pine than by oak (even with co-cominance). The forest shown here, and included under the White Oak section of this chaper, was clearly dominated by white oak with shortleaf pine ranging from being lesser of two co-cominants to the main associate species. In local stands of this forest in which white oak and shortleaf pine were co-dominant post oak was associate tree species. A list of shrub species in the forest community presented here was a long one. Flowering dogwood, sassafras, persimmon, grape, smooth sumac (Rhus glabra), shining or winged sumac (R. copallina), blackberry, redbud, and poison oak were good "for starters". Indiangrass, little bluestem, and Canada wildrye were dominant grasses in that order. The Eurasian orchardgrass (Dactylis glomerata) was also present.

Hobbs Wildlife Management Area, Benton County, Arkansas.October, autumnal aspect. FRES No. 14 (Oak-Pine Forest and Woodland Ecosystem). K-101 (Oak-Hickory-Pine Forest). White oak-dominated variant of SAF 76 (Shortleaf Pine-Oak). Oak-Pine Series 122.14, Northeastern Deciduous Forest 122.1 of Brown et al., 1998, p. 37). Ozark Highlands- Dissected Springfield Plateau-Elk River Hills 39b (Woods et al., 2004).

182. Views of an Ozark "pinnery"- Hillfolk in the Ozarks traditionally refered to hardwood (most commonly oak-hickory) forest with pines (usually scattered individuals rather in groups) as a "pinnery". "Setting the woods afire" (often for the expoused purpose of "killing them *#&%** ticks" by white hillbillies was a lesson well-learned from the Indians and such flaming rituals of spring undoubted gave some competitive advantge to the more fire-adapted conifers. The sundown autumn scenes shown here from the western Ozark Highlands accurately represented a typical "pinnery". There were enough adult shortleaf pines and they were reproducing adequately to add a "pine flavoring" to the white oak-dominated form or phase of the Ozark oak-hickory forest.

As the sun sets there's just time to do the chores, eat a leisurely supper, and then load up the hounds to spend an evening listening to the mountain music as Black and Tans, Blueticks, and Redbones inform us of their progress in pursuit of coon or fox. Bring plenty of crackers and sardines, boys. It'll be a fine fall night in the pinnery.

183. A well-formed Ozarker- Mature white oaks growing in the open develop magnificant crowns formed by large limbs that branch and rebranch repeatedly. Such white oaks are priceless shade trees rather than forest or timber trees that form large to massive boles which are free of limbs for distances sometimes in excess of 50 feet and that are prized for their yields of high- grade white oak lumber. The grand specimen paraded here grew on a fertile upland site in the graveyard of a rural church where it stood in stark testimony of the sorts of trees that can be produced in the western Ozark Plateau.

McDonald County, Missouri. June.

Bur Oak (Quercus macrocarpa) Forest

Bur oak as a forest cover type (SAF 42; Eyre, 1980, ps. 39-40, 236) is the most northward and westward of the eastern and central continental tree oaks. Bur oak comprises the only oak forest cover type in more northern portions of the Central Lowlands and Great Plains physiographic provinces. Bur oak is extremely tolerant of drought and fire and forms forests, groves and savannahs within (and within) the central grasslands of North America from the eastern edge of the Prairie Peninsula (Transeau, 1935) through to the Black Hills (Eyre, 1980, p. 39). Bur oak is to this vast region what post oak and blackjack oak are in the south (eg. Cross Timbers). These two forest range types meet near northern reaaches of the Flint Hills Region.

Presented in this section was an example of the bur oak forest cover type (SAF 42) found on a north slope and outward to a ridge top in the Nebraska Sandhills. This was about as dry (least mesic) an environment as bur oak is adapted to, at least to the extent of forming forest communities. This example was a forest and not a woodland or savanna although at its outer edges it did form small groves with tree density and understorey more suggestive, or even typical, of woodland. This contiguous bur oak-dominated community (both forest and grove phases, or two communities if so interpreted) was the climax (ptential natural) range vegetation on this sere except where frequent, close, mechanical mowing had converted the native herbaceous understorey to a manmade one made up of Eurasian perennial grass species adapted to such intensity and frequency of defoliation. Examples of both native and naturalized herbaceous layers were presented following a brief section that introduced the foliage and fruit features of bur oak.

This relict stand of bur oak forest had developed on what was at this time the Valentine, Nebraska city park (and almost any city park says it all). This forest range vegetation occupied both upland, including and especially a mesic north-slope, and bottomland site. Vegetation on all but the north-slope was highly modified by yard-mowing, manicure-it urban man (even in this over-whelmingly rural region). The natural herbaceous understorey of the upland phase was Canada wildrye (Elymus canadensis) except on north-slope upland forest range where the dominant herb was—as on the bottomland phase—long-beak(ed) or Sprengel’s sedge (Carex sprengelii). It was likely that there was also some bristleleaf or ebony sedge (C. eburnea) was growing in association with long-beak sedge, but C. eburnea could not be positively identified given absence of inflorescences. Under frequent, close mowing Kentucky bluegrass (Poa pratensis) dominated the lowland understorey. With less frequent mowing smooth brome (Bromus inermis) dominated both bottomland and upland phases.

Both of these introduced Eurasian grasses have naturalized widely and under a farmer (vs. forester or rangeman) frame of mind, and management consistent therewith, the native grasses and sedges were crowded out by aggressive, highly competitive agronomic forages. The timeless story since Cain and Abel. The farmer (cornhusker in the Cornhusker State) again won out over the pastoralist.

184. Big tree on the prairie- Beautiful and massive specimen of bur oak that made its home in the floodplain of the Bosque River in the Western Cross Timbers of northcentral Texas. Bur oak is obviously a minor member--especially when compared to the dominant post and blackjack oaks--of the scattered forest and woodland communities in southern portions of the tallgrass prairie region. In more northernly parts of this vegetational realm bur oaks replace blackjack and even post oak as the dominant oak. In these northern parts of the Prairie Peninsula and tallgrass-true prairie region bur oak would be second--if that--only to eastern cottonwood as a dominant tree both on grassland and isolated forest communities such as gallary forests. Bur oak is the most widely distributed of all the oak species that have the role of hardwood dominant across the North American central prairies (McGregor et al., 1977, ps. 39-41). Post oak and blackjack oak barely extend northward to Iowa whereas bur oak extends to Ontario.

Bur oak forms forests, woodlands, and groves (distinguished by tree density, extent of canopy cover, and size of stand resulting in varying degrees of understorey development) in more northern areas of North America as in the Nebraska Sandhills, an example of which was used for photographs and corresponding descriptions and explanations in this section.

Hamilton County, Texas. July.

185. Bur(r) or mossy cup oak (Quercus macrocarpa)- Young leaves and catkins of the dominant oak of the northern tallgrass, true, and mixed prairies northward from central Kansas. Bur oak can be interpreted as the northern ecological equivalent of post and blackjack oaks as the aboreal dominant of the grassland-deciduous forest ecotone (Vankat, 1979, p. 221). This species produces the largest acorns of any oak in North America and it is the oak of the famous oak groves and savannas of the northern grasslands (Peattie, 1938). Seedlings rapidly send their tap roots deep into the fertile prairie soils and become quickly established after germination (Weaver, 1968, ps. 135-139). This genetic adaptation to drought combined with the species’ thick fire resistant bark (Allen, 1967, p. 15) make it admirably suited to drought- and fire-prone prairies. It’s range extends far south of it’s region of dominance into central Texas where it sometimes dominates bottomland savannas.

Hamilton County, Texas, April.

186. Leaves and fruit of buroak- Burr oak (either one or two "rs" are used) is regarded as having the largest acorn of any Quercus species in North America. Like other white oak species buroak bears fruit every year (versus a two-year cycle) in the red or black oaks. The combination of large acorns and production each year makes this species one of the most valuable producers of mast in the eastern deciduous forest (ie. the eastern part of North America).

Burroak is also known as mossy-cup oak, both common names in reference to the conspicuous tapered tips of the scales of the cup which present a fringe-like appearance. The geographic range of. Q. macrocarpa extends from eastern Canada (New Brunswick and Quebec) to Texas.

Hamilton County, Texas. September.

187. Unusual leaf arrangement in bur oak- Bur oak has a unique arrangement of leaves along its leaders. Leaves grow in dense clusters or whorls at intervals along young branches (leaders) with one of these clusters (complete with the current year's acorns) on each of several internodes extending back toward the trunk until older internodes cease to produce such clusters. Two such leaf and acorn clusters on one leader were shown in this photograph. These were obviously the two youngest internodes which showed the last two year's growth of this shoot (one internode being produced each growing season).

Erath County, Texas. Late September; approaching fruit-ripe stage of phenology.

188. Clusters of leaves (and acorns) in bur oak- Two clusters or whorls of leaves and acorns in bur oak. The first photograph presented a whorl from a lengthwise view whereas the second photograph presented more of an oblique and semiend-on view of the second whorl. Bur oak concentrates its leaves in large clusters infrequently along its branches (one cluster per internode) whereas most oak species produce individual leaves and some secondary (usually short) shoots off of young branches (= leaders or shoots) at more frequent intervals along their internodes.

Erath County, Texas. Late September; approaching fruit-ripe stage of phenology.

189. Clustered at the end- End-on views of two different clusters or whorls (at progressively closer distance) of leaves in bur oak. This is a unique and eye-catching arrange of foliage and fruit in a Quercus species especially well-adapted to drought and fire. Bur oaks grow at variuos densities ranging from isolated individual trees to small groves to dense forests (see rest of section immediately below). Function of this unique arrangement (pattern) of leaves in bur oak was unknown to this author.

Erath County, Texas. Late September; approaching fruit-ripe stage of phenology.

Technical Note: The following sequence of photographs of the Bur Oak forest cover type (SAF 42) was taken in early summer (late-June). Two aspects were presented and described: 1) north-slope and 2) hilltop. These represented two distinct aspect-based bur oak forest communities even though both were: 1) dominated by bur oak, 2) the same cover (dominance) type, and 3) upland forest. Both aspect-determined bur oak communities were photographed at the same time (about 0730 to 100 hours Central Standard Time). This time of day was most advantageous for pphotographing the north-slope forest which, having a general northeast orientation (hillside alignment), received most light in the understorey (maximum land surface area exposed to full-sun lighting; greatest understorey coverage of greatest light intensity) in early to mid-morning. Obviously time of daylight hours (excepting very early morning or late evening) was largely irrelevant for photographing vegetation of hilltop forest.

Relatively high density of adult bur oaks on the north-slope forest coupled with dense individual crowns of this large-leafed species resulted in one of the darkest understorey habitats ever seen by this photographer. Most of the understorey area (square footage, yardage, meterage, etc.) of the north-slope forest range received somewhere between half to three-fourths of an hour of direct light daily. During the remainder of daylight time plants--especially herbaceous species--were in relatively dense shade and obtained only indirect light. Herbaceous species in bur oak forest have to be some of the best adapted and most extreme examples of sciophytes (skiophyes) or sciaphlic (skiaphilic) plants, "shade-loving" or plants that have evolved to shade (shady habitats), in the eastern deciduous forest.

Viewers should realize that the shady (generally dark; poorly lite) images presented below, while of relatively poor viewing quality (compared to range types in other biomes), "caught" exactly normal light conditions as they existed, and at their brightest (time of day in which most light penetrated forest crown canopy to reach ground level). No artificial lighting was used. In other words, intensity and quality of light seen in photographs were those that the viewer himself would have experienced, the images that would have traveled through his cones, rods, and optic nerve, had he moved through this forest environment.One can but marvel at the adaptation of the herbaceous plants in bur oak forest range.

190. Sylvan scene in Nebraska Sandhills (Are you kiddin"?)- Send this as a postcard from the Nebraska Sandhills and those who deal in sterotypes will asert, "You're kidding me". Nope, the real thing.. Bur oak Sandhill forest on a north slope in early summer. Most of the trees in these two "dendrographic" photographs were bur oak (including the largest tree and the pole-size sampling in left front of the "big 'un"), but white or American elm (Ulmus americana) and green ash (Fraxinus pennsylvanica) were associates that were present (as in background of these two slides). (Incidentially the commerative specific epitthet pennsylvanica means literally "Penn's woods" [ in honor of the Quaker founder, William Penn] ; " sylva from the Latin and Greek roots is in reference to "wood" or "forest").

This was an old-growth stand of bur oak with the climax herbaceous species still dominating the understorey. This was clearly virgin range vegetation.

The graceful herbaceous understorey in these two "photo-plots" was almost exclusively made up of long-beak or Sprengel's sedge (Carex sprengelii). Canada wildrye (Elymus canadensis) was the associate herbaceous species overall though very little of this grass was present in forest range vegetation presented here. Eastern red cedar (Juniperus virginiana) had in absence of fire begun to encroach as an invader into this sylvan Eden.

Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Juchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001).

191. Appreciative of what it receives- Close-up "photo-quadrant" of local sward of long-beak or Sprengel's sedge presented in the first photograph of bur oak forest treated in this series. This is the very fleeting maximum intensity of light that these shoots will receive.Short duration of light from early morning rays on a north slope in Nebraska Sandhills.

Cherry County, Nebraska. Late June.

192. Forest and sunflecks- Nebraska Sandhills bur oak forest and shade-adapted herbaceous species. Light conditions of early morning, the major part of day during which light could reach ground level of this north-slope forest, provided a textbook illustration of the sunfleck phenomenon. In dense-canopy plant communities like forest (and even grassland swards) sunlight penetrates through foliage at varying lengths of time. The shortest of these light periods or bouts (time lengths or durations of light at whatever light intensity) which can last only a few seconds up to several minutes are known as sunflecks (Smith et al., 1989; Chazdon and Pearcy, 1991). The converse of sunflecks are shadeflecks which are shade fluctuations due to cloud cover between shaded spots and the sun.

In vegetation, such as that of a bur oak forest featured here, wind movement of leaves, travel of Earth relative to sun, and any cloud cover are responsible for the ever-changing state of light that penetrates to lower strata of the range plant community. The general condition or phenomenon of changing or dynamic light as to duration and related intensity or "fluctuation in irradiance" as it was described by Chazadon and Pearcy (1991, p. 760) is an all-important abiotic factor in growth and survival of plants in lower layers of range vegetation-- the understorey of this bur oak forest. Light dynamics interacts with other factors (abiotic and biotic) such as defoliation, precipitation, and, as in this north-slope forest, aspect. Dense shade produced by the large leaves and close interlocking crowns of bur oak on the aspect most shielded from sunlight resulted in severe conditions of light deficiency related stress for understorey plants in this bur oak forest.

Range plants in lower layers of this north-slope bur oak-dominated forest community were sciophytes (sciophytes), "shade-loving" species of the highest adaptation. There were a few, though extremely patchy or irregular, layers of woody plants below the forest canopy. These vegetational strata were made up primarily of regenerating bur oak (seedlings, saplings, and poles), American elm, green ash, and, least of all, eastern red cedar. This photographer did not see any shrub species. The herbaceous layer (which was also the ground layer) of this forest range was dominated mostly by long-beak or Sprengel's sedge with Virginia wildrye the associate species. The major--and about the only--forb was anise root (Osmorhiza longistylis).

This virgin range vegetation was a fine representation of old-growth bur oak forest (even if there were too many shadows to show it to best advantage).

193. A study in shadows- Closer-in photograph of the two big bur oaks and associated herbaceous layer introduced in the last slide (second slide of the preceding pair of photographs). This will be about all--and clearly the most (greatest intensity and longest duration of)--light that the understorey of this north slope bur oak forest will receive, and this was close to longest day of the year. Early morning and late evening were the major (about the only) times that sunlight could fall on the ground at this extreme north aspect.

The phenomenon of sunflecks as explained in the preceding caption was obvious.

Understorey was herbaceous, except for basal sprouts of bur oak (see below), and consisted primarily of long-beak sedge with some Canada wildrye. The only forb of note was anise root (Osmorhiza longistylis). This was virgin vegetation in structure and composition and in all layers; a good representation of climax bur oak forest range.

194. About as bright as it ever gets- The slant of earlier morning light on this north slope is just about as much photosynthetically active radiation as the understorey of this north-slope, Nebraska Sandhills bur oak forest ever gets.

These paired photographs presented an example of light dynamics in the understorey layer of this densely shaded forest range. The second of these two photographs was at a closer distance to trees, but the major difference in pattern of light and shade on the forest floor was due to changing radiation that occurred with passage of time. These photo-dynamics resulted from 1) variation due to the diurnal (day-and-night) cycle of light and 2) annual cycle due to travel of Earth on its yearly orbit around the sun. Sunflecks lasting only a few minutes or, perhaps more commonly, mere seconds (or fractions of seconds) are largely due to wind movement of leaves in the forest canopy. This rapid photo-dynamics--very short-term spots of alternating sun and shade--could not be captured in photographs or sequences photographs, but the prevalence of light dynamics, hence the concept of sunflecks, was represented photographically.

Trees and saplings in these slides were all bur oak. The herbaceous layer was mostly long-beak sedge with some Canada wildrye and, even less, anise root. Old-growth bur oak forest serving as a reminder of what a virgin forested range was like in a most unlikely grassland region.

195. Shade-adapted range plants- This two-slide set of a north-slope, Nebraska Sandhills bur oak forest featured the 1) herbaceous understorey dominated by long-beak sedge with Canada wildrye as associate species and anise root as principal (essentially sole) forb and 2) base of a mature bur oak with stump sprouts and a sapling. The sapling was at such distance (about a foot and a half) away from the trunk of the adult tree that it was apparently a separate tree derived from an acorn (most likely the adult bur oak it was closest to). Thus this bur oak sapling appeared to be a sexual offspring of the adult oak, a separate and unique genetic individual produced by an adult bur oak through fruit. It was possible that this sapling grew from an acorn produced by a neighboring bur oak and transported next to the big oak shown here by a combination of gravity, wind, and steep slope or by a food-storing squirrel. There was some probability that this sapling was actually a basal sprout, secondary shoot, that arose from the adult tree, but this seemed not to be the case.

By contrast, the small stump sprouts to the right of the sapling were asexual offspring from the large bur oak. In this case the new oak shoots (stump sprouts or stump suckers which are secondary shoots) were clones (clonal progeny) of the parent tree. These were genetically identical to the big bur oak from which they arose. Stump sprouts were secondary shoots of the existing bur oak tree the same as new limbs, branches, and buds. Such clonal shoots are often called offshoots. This asexual reproduction is vegetative reproduction (synonyms), human horticultural forms of include grafting and budding. More (and more clear) examples of suckering from healthy, uninjured, mature bur oak was shown below near end of this section.

Almost all tree species are capable of both sexual and asexual reproduction, but sexual regeneration is less successful in forest of dense shade unless the species are relative in tolerance (Tolerant or Very Tolerant). Vegetative regeneration is not limited (at least not nearly as much so) because the parent tree can translocate photosynthate and other nutrients to its own shoots (clonal organs). By contrast, sexually produced progeny are their "own plants" and must be able to survive under the shade of their parents at age classes ranging from seedling through sapling to pole-size. Most of these individual smaller trees--at least in species having lower tolerance--die. An example of this reality was the stick (at left of trunk and sapling) which was the remains of a small sapling tht could not survive in the shade of (compete with) the aduld bur oak.

Burns and Honkala (1990) reported that bur oak has usually be interpreted as being Intermediate in tolerance rating although certain observers viewed bur oak as more tolerant than other climax oaks such as northern red and white oak. In the Prairie Peninsula Region however bur oak stands have been invaded by other oaks including white oak, and bitternut hickory (Burns and Honkala, 1990). This was far to the east and on more mesic habitats than in the Nebraska Sandhills where the more mesophytic oaks and hickories are absent. These same authors specified that in forests to the east and north bur oak died from suppression (due largely to shading?) after reaching sapling size. That same response was frequent in the north-slope Sandhills bur oak forest described here.

196. Dominants of a deep, dark lower layer- Local sunlit patch of the herbaceous layer in a north-slope bur oak forest in Nebraska Sandhills. Plant species seen here were the climax herbaceous species that dominates the understorey of a sandhills bur oak forest..Long-beak or Sprengel's caric sedge (Carex sprengelii) was the dominant--almost exclusive--species with Canada wildrye the associate species. Anise root was the only forb of consequence in the herbaceous layer. These were reported as characteristic (indicator) herbaceous species in bur-oak dominated range vegetation (Barker and Whitman, 1989, ps.17-18).

The first of these two photographs was of a single-species stand of long-beak sedge. Second photograph included a shoot of Canada wildrye accompanying the long-beak sedge.

197. Contaminated understorey- Sward of herbaceous layer of a north-slope bur oak forest in Nebraska Sandhills consisting of long-beak or Sprengel's sedge and of naturalized Kentucky bluegrass (Poa pratensis). Kentucky bluegrass and smooth bromegrass (Bromus inermis) are two widespread Eurasian, perennial festucoid grasses introduced by the white man that have natrualized widely in North America. These two introduced grasses are commonly used throughout the greater Northern Great Plains Region (as well as areas of adjoining regions) for grazing and, in case of smooth brome, for both pasture and hay. They continue to have ample opportunities for naturalization and establishment across a vast acreage, especially on disturbed sites in humid through semiarid zones, so they are properly regarded as naturalized range plants..

Continued heavy mowing up to edge of the north-slope, old-growth bur oak forest described heein enabled both Kentucky bluegrass (as shown here) and smooth brome (covered below) to invade outer parts of the native herbaceous understorey of this bur oak stand. This invasion allowed displacment, to varying extents, of native graminoid species like long-beak sedge and Canada wildrye. These two agronomic, Eurasian grasses were able to outcompete natives only where there was continued disturbance by repeated, heavy mowing over specific locations or of adjacent areas from which the exotics could spread for short distances into unmowed native sward. The sward shown here was an example of the latter situation.

Cherry County, Nebraska. Late June, peak flowering stage of both long-beak sedge and Kentucky bluegrass.

198. Long-beak or Sprengel's caric sedge (Carex sprengelii)- Three photographs showing progressively greater detail of sexual shoots on the dominant herbaceous species in the understorey of a north-slope Nebraska Sandhills bur oak forest.

Cherry County, Nebraska. Late June, peak flowering stage just after anthesis.

199. Broadleafed Lone Ranger- Anise root (Osmorhiza longistylis) in the herbaceous layer of the understorey of a north-slope bur oak forest in Nebraska Sandhills. This was only forb that this photographer could find in this climax forest range. It is a characteristic species; generally a dominant and indicator species in the bur oak-dominated forests of this region (Barker and Whitman, 1989, p.18). Its graminoid neighbors were long-beak caric sedge, the dominant, and Canada wildrye, the associate species.

Cherry County, Nebraska. Late June, late pre-bloom phenological stage.

200. Invasion at the edge- Lower and outer edge of a north-slope bur oak forest in Nebraska Sandhills with an individual eastern red cedar prominently and happily growing in a mid-layer of vegetation. This immediate local was just above Minnechaduza Creek. This accounted for the lush stand of herbaceous riparian vegetation in the foreground. which "crept in" into this "photo-plot" useed to show size of the cedar relative to typical size of adult bur oak (to immediate right of eastern red cedar). Human suppression of fire had permitted eastern red cedar to establish sporadically both along edge and in interior of the bur oak-dominated forest.

Most of the herbaceous riparian vegetation was the introduced Eurasian grass, smooth brome which invaded the bur oak understorey under frequent, heavy mowing. Also present with much cover and density, though lacking height of the bromegrass,was another Eurasian grass, Kentucky bluegrass. Other riparian species were native grasslike plants (Cyperaceae) that were still only in vegetative stages and unidentifiable by this worker. These had, however, been overwhelmed by the introduced grasses. The only native plants that could persist under cover of the agronomic grasses (which were competitive with native herbaceous species only under heavy mowing) were bur oak, green ash, American elm, and eastern red cedar. And mowing wiped out all plants of these tree species except saplings and adult trees.

201. Bur oak stand at the edge and changed by man- Part of the perimeter of the north-slope Sandhills bur oak forest adjacent to a field of smooth bromegrass. Extension of close (short-height) mowing, over a number of years, into the edge of the bur oak forest had either directly killed out long-beak sedge, Canada wildrye, and anise root and/or permitted invasion of smooth brome and Kentucky bluegrass that "choked out" (outcompeted, overshadded, etc.) these natives.

202. Advance guard of bur oak-dominated vegetation being invaded by enemy legions- Two views of a bur oak forest with herbaceous layer(s) that were highly man-modified. This local small stand of bur oak stood in stark contrast to the virgin climax bur oak forest featured above. These two photographs showed the outer- and lowermost perimeter of the bur oak community covered in this section. The adult bur oaks and a few patches of the natural understorey beneath mature trees had persisted (where oak trunks were too close together to permit entry and operation of rotary mowers [=shreaders]) along this perimenter. Generally, however, continued close mowing of the range vegetation had done two things: 1) prevented regeneration of bur oak by cutting off all bur oak seedlings and 2) largely converted the natural herbaceous understorey to an anthropogenic or manmade (=artificial) layer of smooth brome with local stands of Kentucky bluegrass.

Change in species composition of the herbaceous layer(s), replacement of native species with naturalized Eurasian species, was due to either direct killing of long-beak sedge, Canada wildrye, and anise root by defoliation or indirectly by reducing competitiveness of these range species and/or favoring the taller-growing smooth bromegrass along with denser-growing, more cutting-toleraant Kentucky bluegrass. It was explained above that both of these introduced, agronomic forage grasses have naturalized throughout much of the North America range country. Where mowing disturbance (= abnormal defoliation regimen) occurred throughout much of the growing season the understorey was converted from an herbaceous layer of native grasslike, grass, and forb species to one of naturalized agronomic grasses (ie. conversion from native to naturalized range plants). Also, the lower woody layer(s) was totally eliminated because mowing prevented regeneration of all tree species.

Beyond doubt or debate, continued close mowing as had been practiced on this forest range for years will eventually eliminate the bur oak forest (as soon as the already old trees die off) and convert the plant community into a manmade grassland of smooth brome and Kentucky bluegrass, a farm field of domestic forages.

203. Artificial understorey- Continued close mowing by roatary shredders killed out--directly or indirectly--the climax herbaceous species (long-beak sedge, Canada wildrye, anise root) of a bur oak forest understorey and converted the sward to a single-species layer of smooth brome.A small remnant of smooth bromegrass remained uncut where this beautiful trunk of an old-growth specimen of bur oak prevented mowing (an inexperienced or lazy shredder operator did not get close enough to the oak).

The other glaring outcome of mowing was absence of regeneration of bur oak, dominant tree species, and green ash, the associate species. If present mowing practices continue (and they obviously had been in effect for a number of years) there will eventually be no bur oak forest left. Instead, the plant community will be a monoculture of smooth brome (with local spots of Kentucky bluegrass). All native vegetation will be wiped out (ie. the forest range will be no more and a hay field will have replaced it).

204. Genuine-article understorey- Two trunks of large bur oak prevented close mowing of the native herbaceous understorey comprised of long-beak or Sprengel's caric sedge, the dominant, and Canada wildrye, the associate species. Survival of these climax species permitted some of the composition and structure of a bur oak forest range to persist. This part of the old-growth bur oak forest that had developed on a north slope in the Nebraska Sandhills was mowed with less frequency than some of the other forest understorey with the result that the climax species composition of the sward (herbaceaous layer) persisted. Tree regeneration (bur oak, green ash, American elm) was still completely prevented as any seedling was mowed off where density of adult trees and saplings did not prevent entry of rotary mowers.

205. Nebraska Sandhills bur oak grove- A bur oak grove, the outermost edge of which was shown here, had developed on a sunlit ridge conterminous with the north-slope bur oak forest described immediately above. Both phases--open, fullsun grove and dense, deeply shaded forest--were climax range vegetation and represented the extremes physiographic and structural manisfestations of the bur oak forest cover type. The greatest differences were in the understorey vegetation, as to vegetational structure and species composition of herbaceous and woody (shrub and smalltree) layers.

The bur oak grove phase (or community, if the two extremes were to be viewed as two different range plant communities) had no species of woody plants except bur oak (in contrast to American elm, green ash, and eastern red cedar in the north-slope forest). Also absent from the bur oak grove on the drier and more sunlit ridge was long-beak cric sedge which made up most of the herbaceous biomass on the north-slope forest. Instead, Canada wildrye, associate on the north-slope forest, was the overall dominant of the herbaceous layer in the bur oak grove. Giant ragweed (Ambrosia trifida) was a locally dominant herbaceous species and the most abundant forb in the oak grove. Anise root, the only forb, on the north-slope forest was absent in the grove vegetation.

This bur oak grove was on the border of a field of the Eurasian perennial grass, smooth brome, which, as shown above, had invaded the understorey of a north-slope bur oak forest under repeaded close, mechanical mowing when the native long-beak sedge, Canada wildrye, and anise root were killed out by such extreme (intense and frequent) defoliation. Where trunks of bur oak were too close together to permit intrusion of this bloody equipment and overmowing the native, cool-season Canaada wildrye dominated the understorey other than in microsites where giant ragweed and bur oak stump sprouts held this honor.

Physiogonomy and structure of this Sandhills bur oak grove was presented in this two-slide episode. Details of the herbaceous layer of this grove was presented in the next two-slide episode.

206. Edge of a bur oak grove- Lower zone of tree trunks and herbaceous understorey of a Nebraska Sandhills bur oak grove. This stand of climax vegetation was conterminous with the north-slope bur oak forest featured earlier. In contrast to the deeply shaded understorey of the bur oak forest in which long-beak caric sedge was dominant and Canada wildrye was the associate species, the herbaceous layer of the grove phase of the bur oak cover type was dominated by Canada wildrye except in small areas (microhabitats) where giant ragweed held supremancy or where basal sprouts of bur oak overtopped the dominant cool-season, festucoide grass.

Canada wildrye was featured in both of these photographs with the local sward in left foreground of first slide shown at close range in the second slide. Some of the dead (light brown or buckskin-colored) shoots of Canada wildrye included some of the previous year's growth (slightly more faded) as well as some of those of the current growing season. Almost all of the current year's shoots of Canada wildrye were still in the boot. It was not known why a few of the current growing season's shoots had already matured, died and gone into dormancy. Did make for a nice contrasting picture with Canada wildrye very conspicuous.

Giant ragweed, a warm-season annual composite of mature size that lives up to its common name, was still in early growth and thus not shown in proportion to its yield of biomass at peak standing crop. This herbaceous understorey was the vernal society even though it was summer and bur oak were representative of the estival aspect.

207. Parents, progeny, and playmates- Sprouting (suckering), the production of secondary shoots, from base of trunks in non-injured adult bur oak trees. Bur oak in both of these "photo-plots" were on a ridge at edge of a bur oak grove where light could penetrate from all directions throughout most of the day. This was a different condition of radiation than that of the north-slope bur oak forest described at beginning of this section. Differences in understories of the bur oak grove considered here and the bur oak forest described previously were explained in the immediately preceding caption.

It was strikingly evident that full-grown (adult) bur oak produced basal sprouts in both deep shade and full sunlight (forest and grove stands, respectively). This was an understandable characteristic when it was born in mind that these secondary shoots were being supported by the parent shoot (ie. sprouts were a photosynthate sink from the parent tree that was the a nutrient (food) source for the basal suckers. (Sprouting frrom stumps and basal trunks of bur oak was considered in more detail in the next succeeding set of photographs.)

What was substantially different in understories of bur oak forest versus grove was in herbaceous species and tree species other than bur oak (see again preceding caption). Mature sexual shoots of Canada wildrye, the overall dominant herbaceous species in the bur oak grove, with conspicuous spikes were present in the first of these two p;hotographs. Also prominent in this first slide was the pioneering, annual composite, giant ragweed. The second slide presented a fullsun shot of a basal sprout of bur oak at edge of the grove where frequent mowing had killed out the native Canada wildrye and its replacement with the introduced Eurasian Kentucky bluegrass.

208. Living stumps- Two stumps of bur oak with prolific and vigorous sprouting or suckering. These trees were at edge of a bur oak-dominated forest in the Nebraska Sandhills. Stump sprouting is an adaptation to fire, a key abiotic factor of grasslands. Bur oak is one of the most fire-adapted Quercus species in North America with sprouting from stumps or injured tree trunks one of its most important means of regeneration. In fact, bur oak sprouts prolifically even from old trees and those not subjected to injury. This was shown in several preceding photographs for bur oak growing in deep shade and fullsun environments.

Bur oak has long been recognized for its "thick fire-resistant bark" (Burns and Honkala, 1990), but prolific sprouting (suckering) is an accompanying adaptation to recurrent fire that was evolved in the generally fire-prone habitat of this species biological range. Burns and Honkala (1990) explained that prolific sprouting was common following burning or cutting of bur oak only up to pole-size with such secondary shoots being of poor quality except those of seedlings. This latter conclusion was confusing to say the least given that shoots of seedlings are not sprouts (secondary shoots) at all but rather primary shoots derived from acorns. These same authors noted that larger trees (mature or full-grown age class?) also produced basal sprouts but that vigor and quality of these sprouts had not been evaluated relative to age, size, etc. of parent trees.

The two examples shown here were pictorial evidence that seemed to contradict findings reported in Burns and Honkala (1990). The profuse (and plenty vigorous) stump sprouts shown off here were from mid-size trees (a foot or more in basal diameter). This was considerably smaller than the full-grown bur oaks presented in the immediately preceding set of two photographs, and these appeared vigorous enough too.

Cherry County, Nebraska. Late June.

209. Fire ran through the bur oaks- At the outer edge of a grove form of a bur oak forest along a small stream through a hot fire with high flames had burnt just two to three prior to time of photograph. This forest range was in the Dissected Till Plains of the Central Lowlands physiographic province of northeast Kansas which is about the southern limit of forest dominated by bur oak. Thee two slides presented an outside-the-forest view to show physiogonomy and structure of the plant community.

The lush growth of the herbaceous layer was dominated by seedlings of giant ragweed (Ambrosia trifida) with scattered plants of Canada wildrye, Canada wood nettle (Laportea canadensis), and pokeberry or pokeweed (Phytolacca americana). Giant ragweed is a pioneering annual; in fact, it is the most abundant pioneer or colonizing forb in this area. The hot fire had been a disturbance of such severity as to create an ideal habitat for invasion by giant ragweed which "eclipsed" the other herbaceous species which were perennials. These perennials had been present years before the burn, but they, too, appeared to have benefitted from the fire jucging by growth of other plants of these species in a few isolated spots that did not burn. This phenomenon was explained in captions below.

The saplings in foreground of these (and subsequent) photographs were all hackberry and American elm. Almost every one of these saplings (and seedlings that were too small to show up in the slides) were top-killed by the heat of the obviously hot fire which burned with intensity adequate to partially kill lower branches of bur oak. This latter feature of the fire was shown and described in greater detail below. In this region most climax bottomland forests are dominated by various combinations of hackberry, American elm, slippery elm, and green ash (the Tolerant climax species) with old trees of such Intolerant, colonizing species as eastern cottonwood, sycamore, black walnut, and honey locust persisting into climax as associates. This is Society of American Foresters (Eyre, 1980) cover types 93 and 94. These mixed hardwood forests are the climatic climax. By contrast, bur oak forests, Society of American Foresters (Eyre, 1980) cover type 42, are a pyric climax. Differences in alluvial soils of these floodplain forest are not so different as to account for differences in forest range types.

In the absence of periodic fire the bur oak forest would eventually proceed through plant succession to a hackberry-American elm climatic climax. That is, it would be the potential natural vegetation along streams in this climatic regime as interpreted from perspective of polyclimax and climax pattern theories. Or, what amounts to the same thing, bottomland forests dominated by hackberry and American elm would be the edaphic climax, the natural termination of such stream bottom seres. Either way, it is only periodic hot fires that maintain bur oak, which is Intermediate in tolerance (Burns and Honkala, 1990), as climax along stream bottoms (ie. bur oak is a fire type, a pyric climax). In monoclimax theory all such forests are postclimax to the climatic (= zonal or regional) climax of tallgrass prairie.

Individuals of bur oak would undoubted persist into a hackberry-American elm climax as bur oaks are relatively long-lived. Burns and Honkala, (1990) reported that some bur oaks bear fruit up to ages of 400 years, the longest of any Quercus species in North America.

Progression through plant succession from a seral bur oak-dominated forest to a hackberry or hackberry-elm climax forest was the pattern of vegetation development predicted for gallery or floodplain forests in northeast Kansas by Bellah and Hulbert (1974) and Abrams (1986). Abrams (1985) concluded that fire frequency (mean fire interval) bur oak-chinquapin oak gallery forests in the Flint Hills ranged from about eleven to twenty years with an average "somewhere between that range" but with a historic interval of two to three years. Clearly the historic frequency of two to three years would have maintained the oak forests and prevented progression to hackberry or hackberry-elm. It is likely that even an average fire return interval of roughly fifteen years in contemporary time would maintain the bur oak-chinquapin oak forest. Again, bur oak forest (SAF 42) is a fire type or pyric climax.

Spring Creek, Marshall County, Kansas. Mid- June, vernal aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Kuchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Western Corn Belt Plains, Loess and Glacial Drift Plains Ecoregion 47i (Chapman et al., 2001).

210. Late spring after fire-Apperaance of a bur oak grove two to three months following a hot fire that effectively top-killed seedlings and saplings of hackberry and American elm that had established beneath the bur oak canopy. Fire intensity was adequate to create a disturbance that favored rapid establishment of a dense population of giant ragweed, the most common and prolific pioneering plant in this region of the Dissected Till Plains of northeast Kansas. Rapidly growing seedlings of giant ragweed, an annual composite, accounted for the greatest herbaceous cover, but there was also an abundance of Canada wildrye and pokeweed (though widely scattered with most cover being local) along with local exclusive colonies of Canada wood nettle (see photograph and caption below). At outer margin (just under forest canopy) there were large individuals of what appeared to be--without much familarity of early growth shoots--giant sumpweed (Iva xanthifolia).

The first of these two photographs was a closer-in view of the bur oak at center-left in the second photograph of the preceding two-slide set. The trunk of this tree had split almost to ground level yet was still standing and healthy. Heartwood of a tree is dead tissue anyway so as long as the tree was not weakened to point of crashing and did not have heart-rot it remained as healthy is if the trunk was intact. The second of the photographs of this caption presented a summary view showing several top-killed saplings of hackberry and American elm. The rightmost bur oak (center-right) sported an old fire scar and ws in every way healthy from all outward appearances. This fire-scarred tree was featured below.

211. Old and the new after a fire- Close-growing and very old bur oaks (first slide) along with seedlings of giant ragweed and this season's shoots of Canada wood neetle, pokeweed, and Canada wildrye as an herbaceous layer plus some saplings of hackberry and American elm (see especially second slide) comprised the range vegetation of a bur oak forest two to three months following a hot surface fire. In the background there were several pole-sized trees and saplings of red mulberry (Morus rubra) that formed an erratic lower tree layer. One of these saplings was shown in center midground of the second slide. The sapling in center foreground of this second photograph was one of the few young hackberries that was not top-killed by the fire. Other saplings of hackberry and American elm that were not topkilled outright had already started to "peel bark" and appearedlikely to die. The hackberry sapling featured here defied the odds and appeared ready--at this juncture anyway--to take its place ultimately as an adult in the grove of bur oak.

It was explained in the immediately preceding caption that bur oak is a fire type forest. In absence of periodic fire bur oak--other than as a few persistent old-age trees--would be replaced by hackberry and American elm which would be the climax dominants on this bottomland sere. The latter two are Tolerant species whereas bur oak is Intermediate in tolerance rank (Burns and Honkala, 1990). Of course, fire is as much as a part of climate (or a consequence of climate if one prefers) as drought, wind storms, and floods. More precisely, lightening is as much part of the atmosphee as precipitation and temperature and fires from lightening with fuel made possible by climate are major factors responsible for maintenance of bur oak forests within the surrounding zonal vegetation of tallgrass parairie.

Various studies such as those by Kucera (1960) and Bragg and Hulbert (1976) showed that woody plant communities, including forests, expanded in the tallgrass prairie region since European settlement (due in large part to fire suppression). Abrams (1985) estimated that the mean fire interval for bur oak-chinquapin dominated gallery forest in the Flint Hills of Kansas was two to three years and roughly eleven to twenty years with intervention by white settlers and contemporary man. Abrams (1986) documented this phenomenon and predicted that hackberry, elm, and redbud (Cercis canadensis) would gradually replace bur oak and chinquapin oak--at least in absence or reduced frequency of fire--as the climax forests for bottomland forest sites in the Flint Hills and Dissected Till Plains in northeast Kansas. Abrams (1992) issued a later report based on his earlier findings and concluded that for some areas of Kansas the presettlement vegetation of tallgrass prairie had become either a bur oak-hackberry overstorey with a hackberry-elm understorey or a chinquapin oak overstorey with a elm-redbud-chinquapin oak understorey. This analysis struck the current author as an interim report because it seemed likely to him that the ultimate state would be a climax hackberry-elm forest.

212. Just another encounter with fire- Outer edge of a bur oak grove two to three months following a surface fire that burnt through this bottomland forest. Judging from fire scars and general understanding of bur oak forest and the natural envioroment of this area it was obvious that this was the most recent of many such fires in these old bur oaks. The fire had been hot enough to create a disturbance so severe that giant ragweed, the most abundant pioneer species in this area, had already established populations that pretty much excluded other herbaceous species except the climax perennials, Canada wildrye and Canada woodnettle (wood nettle). Even these shade-tolerant and already present perennial herbs appeared to have benefitted greatly form the fire. Without this fire a dense layer of bur oak leaves would functioned as mulch and largely smothered most herbaceous plants. Such a layer of large bur oak leaves (= mulch) was present in several small spots that did not burn. Giant ragweed was absent from these and Canada wildrye plants were smaller than those growing on land that had burned.

Note that the lowermost branches of bur oak were largely leafless, but with enough leaves to prove that they were still alive (barely) and that heat from the last flames injured (appeared to have more-or-less killed) these smaller, lower limbs. This was an example of fire-pruning, but such defoliation did not necessarily kill these organs. It was likely that new shoots would arise from intercalary meristem on these lower branches.

The sapling to right of foremost bur oak was hackberry. It leafed-out following the blaze, but was nonetheless severely injured by the heat from had to be an unusually hot fire. This one of the few hackberry saplings that was not totally top-killed and it had entire strips the total height of the shoot that already had peeling bark. All American elm sapling in the understorey were top-killed. Abrams (1985) studied fire frequency within gallery forests of northeast Kansas (the Fllint Hills) and found a natural fire frequency of two to three years and an extended mean fire interval of roughly fifteen years during time of occupation by European man. Either the natural or anthropogenic fire frequency would probably maintain bur oak groves and prevent development of hackberry-American elm forest on the forest range site featured here.

The fire scar on the foremost bur oak was an old wound that was already present and in process of healing before the recent blaze. This "honor scar" was presented and described further in the next slide.

213. New beauty in an scar- An old fire scar on an ancient bur oak that was renewed by a surface fire only two to three months prior to time of photograph. The tree had started started healing the wound caused from previous fire(s) and the most recent burn did not harm the new bark tht was slosly growing over the scar. This was a direct view of the scar that was shown from the side in the foremost bur oak featured in the immediately preceding two slides. Forbs growing in front of the oak were young seedlings of giant ragweed, the herbaceous species that most benefitted from the recent fire.

This is the "price" ("rent" so to speak) to be paid by the fire-adapted bur oak for living in a pyric habitat in which less fire-adapted and more tolerant hardwoods such as hackberry and elms cannot survive. In absence of periodic fire hackberry, green ash, American elm, and perhaps boxelder would eventually replace bur oak through plant succession. These more tolerant species are the climatic climax whereas bur oak is a pyric climax (ie. bur oak forest is, in effect, a fire type in the overall environment of the Dissected Till Plains or, sometimes, Glacial Drift Plains of northeastern Kansas).

Spring Creek, Marshall County, Kansas. Mid- June.

214. After the latest in a long history- Inside a bur oak grove through which a hot surface fire burned two or three months before. This bottomland forest range had developed along a small stream at edge of a tallgrass prairie in the Dissected Drift Plains of northeastern Kansas. There were several "featured attractions" in these two photogrphs. The most prominent of these was the large fire scar on one bur oak trunk that extended from the ground to over 15 feet in height. this was an old scar from previous fire(s) that was "freshened up" by the most recent burn.

Another, though much smaller, fire scar on another bur oak tunk was presented and discussed in the immediately preceding photograph and caption. The current author interpreted bur oak forests in the Dissected Till Plains as a fire-climax type with bottomland forests on this forest site in absence of recurrent fire developing through plant succession to a climax hackberry-American elm forest.

Effects from fire wounds and scars on bur oak are probably not know with any certainity. Entry of disease pathogens (eg. various forms of fungal rot) and insects through tree wounds, including those caused by fire, is always a possibility. One common suspicion--though perhaps without scientific proof--is that fire (heat or charring of wood) has something of a cauterizing effect so that disease entry is less likely with fire wounds/scars than with similar wounds (eg. those ranging from ax blazes to debarking by falling trees). The charred (charcoal) surface of a fire wound, even a fresh one) is a different (drier, perhaps chemically) environment than the moist, peeled-surface wound left by mechanical injury.

What is certain from even brief scanning of the literature is that bur oak fire scars as records of fire regimes, especially fire frequency, have been analyzed in considerably more detail than the impacts of fire damage on tree health. It is also certain that fire in bur oak-dominated and influenced communities (forests, open groves, savannahs, even prairies) is a natural part of the habitat of this range vegetation. Furthermore, it follows that bur oak is adapted to fire and, was the species not so adapted to and tolerant of fire to the degree it is, this species would not be a member of these range plant communities. It is axiomatic that the overall impact of fire on bur oak is minimally adversive, neutral at least, and almost assuredly positive for survival in range vegetation of which it is an important member.

Peterson and Reich (2001) concluded that on bur oak was a fire-resister, a designation given by Rowe (1983) to shade-intolerant tree species that suffer little or no damage from low-severity fres. Peterson and Reich (2001) explained that "[f]ire rarely killed mature bur oaks, even those in the smaller size classes ...". They (Peterson and Reich, 2001) noted further that even saplings of bur oak grew corky bark of such thickness as to protect the cambium from most fires. Burns and Honkala (1990) cited a volume of literature attesting to the thick fire-resistant bark of bur oak, which along with its general drought-tolerance, accounted for presnece of bur oak on xeric sites as well as mesic ones where bur oak was an associate on sugar maple-American beech forests.

Another "featured attraction" in this pair of photographs, especially the second one, was the death (or near death) of shoots of hackberry and American elm saplings standing in stark contrast to the survival of adult trees of bur oak even with fire scars where sizable portions of their trunks were removed. This is a photographic lesson showing that it is primarily periodic fire (with browsing obviously less important) that maintains bur oak groves and forests which would otherwise develop into hackberry-elm forest with only persistent and senescing adult bur oaks (at status of associate species at most). Bur oak forests and woodland as pyric climax (a climax fire type) was discussed in above photo captions (complete with citation of relevant literature) so that further discussion was not deemed necessary or desirable in this current caption. This was an opportune point to acknowledge that in more xeric environments (eg. those to the west the Dissected Till Plains such as the Smoky Hills or Sandhills in subhumid to semiarid precipitation zones) it is probable that limited soil moisture, especially in drought, rather than fire is the primary variable responsible for maintenance of bur oak and restriction of Tolerant tree species like hackberry, elm, box elder, etc.

Also featured here was presence of red mulberry (lower branch with large leaves extending downward from upper-left corner of first photograph). Red mulbery was usually observed to be a smaller tree of the second woody (lower tree) layer, but as one of the--if not the single most--consistent tree species in bottomland forests in humid and subhumid zones extending from the Cross Timbers and Central Prairies in Texas, Ozark Mountains in Missouri and Oklahoma, and through to the Great Plains in Nebraska. In southern mixed hardwood forests, such as those in northcentral Texas, red mulberry frequently grew to relatively large size (eg. 16-18 inch DBH) with straight boles. Red mulberry was found as a consistent member of bottomland forests were dominance varied from that by eastern cottonwood, sycamore, pecan, bur oak, elm, green ash, and hackberry or sugarberry and where soil texture ranged from primarily sandy to predominantly clay. In progressing northward, dominance by members of Ulmaceae goes from exclusively sugarberry (Celtis laevigata) as in north Texas to side-by-side co-dominance of sugarberry and hackberry (C. occidentalis) in northern Oklahoma to exclusively hackberry in northern Kansas and Nebraska. Red mulberry grew with an array of dominant tree species. Again this was usually, though not always, as an understorey or lower-height tree species.

Species in the herbaceous layer shown in these two photographs included giant ragweed, Canad wood nettle, Canada wildrye, pokeweed, and giant sumpweed in that order based on estimated relative cover. Giant ragweed, an annual composite, was the number one pioneer species of denuded land in this area and far-and-away the dominant of the herbaceous layer except where there were local colonies of Canada woodnettle farther in the interior of this bur oak forest. Canada wildrye, a festucoid grass of the barley or wheat tribe (Hordeae or Tritaceae) that responds to disturbance as a decreasere, was able to "hold its own" against the rapid-growing and rank seedlings of giant ragweed. Giant sumpweed was present as sparse though very conspicuous individuals.

215. Sting after the burn- Local colony of Canada wood nettle or, sometimes, stinging nettle (Laportea canadensis) in the interior of a bur oak forest in late spring after a fire in late winter or early spring only a few months earlier. This species is widely distributed in moist forest with a species range in North America from Saskachian east to the Maritime Provinces and south to Florida and west to eastern third of Kansas. This member of the spurge family (Euphorbiaceae) packs quite a sting, condiderably more so according to Stephens (1980, p. 21) than the stinging nettles (Urtica species). Wear long pants and move through it gingerly. The author has negotiated many of a patch of Canada wood nettle and never gotten more than a tiny tingle by showing it the respect it deserves.

The wood nettle was accompanied by several plants of giant ragweed.

Marshall County, Kansas. Mid-June.

[ Home ]