Oak-Hickory Forests-II

The fundamental and practical distinction between coniferous and deciduous forests is useful (and was used herein), but precise, non-arbitrary "lines" are impossible when presenting and discussing forest range types in the eastern half of the continent. This is especially the case when climax or potential natural vegetation is used as the basis for forest types (ie. when cover types, or the more specific management cover types, are discussed as being more or less synonymous with permanent forest types). As discussed in detail below, the epic work of Lucy Braun (1950) is still the definitive basis for the ecological discussion and classification of those North American forests which extend from the Atlantic Coast to slightly beyond the Missouri and Mississippi River drainages. Braun (1950) included all the coniferous forests (forest types, regions, etc.)-- the generic "southeastern pine region"--as part of her one Deciduous Forest Formation. 

The forest range types included in the following section include coniferous, deciduous, and mixed coniferous-deciduous forests. This is confusing but unavoidable given the nature of the vegetation and the standard understanding (the Braun interpretation) of ecological relations and classification of  this forest vegetation. Most of the southeastern pine types presented are management cover types maintained silviculturally as more economically valuable coniferous forests rather than as the climax mixed hardwood-pine forest types. In other words, efforts were made to fit the Society of American Foresters (1980) cover types with the climax types of Braun (1950) and the potential natural vegetation units of Kuchler (1966).      

The major forest communities or forest zones of eastern North America are broad or wide in their spatial patterns unlike the narrow zonation characteristic of the forests of western North America. The “young” mountains of the western part of the continent are taller (in fact, still getting taller) and as a result have more elevation-based zonation of vegetation than do the geologically older and more eroded (lower) eastern mountains such as the Applachians or Ozarks. So too, are the soils of the Atlantic Coast more zonal (ie. major soil units are larger or broader in spational dimension like those of the vast continental interior whereas soils of the Rocky Mountains and the Pacific Slope ranges are more of the intrazonal spatial scale. See for illustration the national soil map of dominant soil orders and suborders (Soil Survey Staff, 1998).

 Vankat (1979, p. 137) wrote that relief within the eastern deciduous forest “is quite variable” yet earlier Vankat (1979, p. 41) had also correctly noted that “low hills “ were characteristic of much of this deciduous forest region. Again, contrast this with the extreme physiography of the Rockys or Sierra Nevada-Cascade Ranges.

Braun (1950, ps. 35-37) listed nine forest regions making up the Deciduous Forest Formation of eastern North America:

                1. Mixed Mesophytic Forest Region,

                2. Western Mesophytic Forest Region,

                3. Oak-Hickory Forest Region,

                4. Oak-Chestnut Forest Region,

                5. Oak Pine Forest Region,

                6. Southeastern Evergreen Forest Region,

                7. Beech-Maple Forest Region,

                8. Maple-Basswood Forest Region, and

                9. Eastern Hemlock-Eastern White Pine-Northern Hardwoods Region.

 Braun (1950, ps. 11-12) interpreted these same combinations of species as forest communities at the scale (both spatial, mostly, and, also, temporal) of climax association  from which, as quoted immediately above, Braun derived the names of forest regions. Braun (1950, ps. 11-12) distinguished between the association-abstract and the association-concrete, a distinction discussed in the review of the derivation of vegetation cover type from the concept of plant association. The Braun association is the association of F.E. Clements. Indeed the entire ecological paradigm on which Braun (1950, ps. 10-15) based her monographic treatment of the North American Deciduous Formation is Clementisan except allowance for and inclusion of edaphic and physiographic climaxes of Cowles, Tansley, etc.  Vankat (1979, ps. 137-150) and Delcourt and Delcourt in Barbour and Billings (2000, ps. 365-378) described eastern deciduous forest vegetation under the Braun (1950) associations of the Clementsian model.

It is important to bear in mind that the Braun associations can occur in more than the one forest region bearing the name of the association (eg. the Oak-Pine Association commonly occurs and the Maple-Basswood Association infrequently occurs in parts of the Oak-Hickory Forest Region).

Several of the species combinations that delineate deciduous forest regions and associations were also used as forest cover types by the Society of American Foresters (Eyre, 1980) as for example White Pine-Hemlock (SAF 22), White Pine-Northern Red Oak-Red Maple (SAF 20), Sugar Maple-Basswood (SAF 26), and Beech-Sugar Maple (SAF 60). The Society of American Foresters emphasized that it’s forest cover types were “based on existing tree cover” (… forest as they are today…”) and that some types may be climax while others are “transitory” (ie. seral stages leading to another climax).

Braun (1950, p. xiii) specified: “Some of the communities for which composition is given are readily referable to ‘forest cover types’ as defined by the Society of American Foresters”. She then added, “However, an attempt to classsify all communities as to ‘cover types’ would be artificial” and often impossible. Undoubtedly this was due to the differences in classification by Braun’s climax basis (with seral communities clearly specified) versus the existing or present-day forest communities basis of the SAF.

 The Society for Range Management (Shiflet, 1994, p. xi) also specified the criterion of “existing vegetation” and that some rangeland cover types are climax and others are seral. The author of this collection of photographs and descriptions repeatedly reminded readers of this situation, but specified that most of the rangeland and forest cover types included herein were climax vegetation. That criterion exist for forest range types of the Eastern Deciduous forest Formation with most photographs being of either old-growth or second-growth forest with climax species composition as described in the classic literature such as Braun (1950) or Shelford (1963, ps. 17-119).

The nine forest regions of Braun (1950, ps. 35-37) were retained with little modification as series in the fairly comprehensive system of vegetation (primarily, climax; secondly, disclimax or subclimax) used in A Classification of North American Biotic Communities by Brown et al. (1998). Their organization of the Eastern Deciduous Forest Formation was: Oak-Hickory Series, Oak-Chestnut series, Beech-Maple Series, Oak-Pine Series, Maple-Basswood Series, and Hemlock-White Pine-Mixed Hardwood Series within the Northeastern Deciduous Forest biotic community and Mixed Mesophytic Series and Pine Series within the Southeastern Deciduous and Evergreen Forest biotic community. The Brown et al. (1998) series were included below following SAF and/or SRM cover type designations.

Their like, their genre of comprehensive, panaramic, descriptive, first-hand accounts of vegetation on this grand scale, will not likely appear again before icicles hang in Hell. The contemporary research world is hung up on numbers, even generated or simulated (vs. real data) numbers often for numbers-sake alone, and especially numbers of publications. This has gone beyond Lord Kelvin’s admonition to “express it in numbers”,  (indeed Kelvin used actual numbers derived from physical experiments) to the point that quantity is everything and quality (always subsidary to quantity) itself is based on numbers. Not only is there little room for Descriptive Ecology, but there is hardly more for descriptive analysis of experiments and observations because the gold-standard of refereed publications has descended, has been perverted, to the quantitative entity of LPU (Lowest Publishable Unit). A natural length paper based on objectives of the study is split into as many LPUs as possible to extend the author’s bibliography. This procedure does not allow enough results to be included in any one paper to allow a discussion of  findings from a comprehensive perspective. Besides the experimental procedure (complete with lots of numbers and split-nine-ways-to-Sunday replications) is the most important part according to anonymous peer-reviewers.  

In an institutional culture where “Publish or Perish” has become prostituted to a realm of pot-boiler papers written from predictable-outcome, piss-ant projects the next generation of Brauns, Weavers, Shelfords are “dead meat” if they devote (ie. sacrifice) their careers to document for eternity the kind of knowledge their “takes a lifetime “ research produced. Such incredible work is left to not only the fully vested or tenured but the tenured full professor of independent financial means at career’s end (and then there is not enough time left to do the work). A key factor in the creative genius and amazing productivity of Frederic E.Clements was that he was able to spend most of his career working for the rich Carnegie Foundation which freed him from the routine of classroom teaching and daily chores of academia thereby enabling him the luxury of a self-proclaimed “escaped professor” (Brewer, 1988, p. 503).  Alternatively, the most lasting and useful research is the province of the academic martyr to whom pursuit of knowledge or satisfaction of curiosity are of higher utility than organizational rank and its financial renumeration.

 Thus the Ecology student is left with the classical works of those “giants in the earth” who reigned when knowledge was the domain of a more leisurely, honest, genteel, and collegial time and culture.

The scholar of biblical texts cannot read just the several English translations of the Holy Bible. He must also understand the native tongues of Hebrew, Arabic, or Greek in which Holy Writ was written. So too with the “scripture” of Ecology. And the language of vegetation, at least North American vegetation, is Clementsian. The serious student of vegetation must be knowledgable and conversant in this language given that so much of the all-encompassing vegetation literature was written predominately from the view of Clementsian Ecology (and vocabulary). These original, monographic works remain the basis, however distant, of current investigations or even classifications of vegetation. The basic ecological concepts in such natural resource fields as Range Management and Forestry remain Clementsian at root (eg. the Clementsian association is the basis of the forest and range cover types as used in North America).

Any who would refuse to familarize themselves with Clementsian Ecology because there are exceptions to and alternative models for some of its general, long temporal-large spatial scales traverse the terrain of ecological literature half blind. In their zeal to reform the basic vegetation paradigm to include, justifiably, the exceptions they end up “throwing the baby out with the bath water”.

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

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

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

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

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

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

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

Dominant grass overall was silky wldrye (Elymus villous) though locally broadleaf woodoats (Uniola latifolia) was common to dominant. Naturalized Kentucky bluegrass (Poa pratensis) was also common at local scale.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Ottawa County, Oklahoma. April.

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

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

Ottawa County, Oklahoma. April.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Along Lost Creek, Ottawa County, Oklahoma. April.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Above Modoc Creek, Oklahoma. May.

A sticky stand- Large colony of pointed-leaf or cluster-leaf ticktrefoil, or also known as sticky tick-trefoil or sticky tick-clover, (Desmodium glutinosum) on a black oak-pignut forest that was burnt three yearsearlier by a moderate-intensity, surface wild fire. This species of papilionaceous legume was one of the most important local dominants on that upland chert oak-hickory forest which was contiguous with the relict tract of sugar maple forest featured above. In fact, D. glutinosum was the overall dominant herbaceous of the black oak-pignut forest range.

Ottawa County, Oklahoma. Late June.

A showy and nutritious dominant- Cluster-leaf ticktrefoil, pointed-leaf tick-trefoil, or sticky tick-clover as the dominant plant species of the herbaceous layer of a black oak=pignut or bitternut hickory upland, chert forest in the western Springfield Plateau. This relict tract of oak-hickory forest had been burned three years earlier by a moderate-intensity, surface wild fire. Many shrubs and trees up to small sapling size were top-killed by this spring wild fire. One of the main beneficiaries of this pyric disturbance was cluster-leaf tick-trefoil.

Ottawa County, Oklahoma. Late June; phenological stages ranging from opening-bloom to maturing fruit.

About the size of a tick (a full one)- Inmflorescence of pointed-leaf tick-clover or cluster-leaf tick-trefoil comprised of numerous, small, papilionaceous flowers arranged on opposite sides of the central axis. This one was growing on one of many plants on the floor of a black oak-pignut hickory forest subjected to a moderate-intensity, surface, wild fire three years earlier.

Ottawa County, Oklahoma. Late June.

Forest legumes with a peanut-flavor- Sexual shoot of sticky tick-clover or pointed-leaf tick-trefoil with several ripening legumes along with some young flowers ( ie. various stages of phenology on the same central stalk of the inflorescence. Distinctively shaped legumes are unique in their own right, but the even more unique aspect of theser fruits is their pronounced peanut flavor. The adjective "tick" attached to trefoil or clover is derived from the readily adhering feature of these legumes due to minute pubescence that permits fruits to be easily attached to animals, including man, and thereby be dispersed by them (zoochory in formal parlance). Hunters, photographers, and all manner of biped woods-walkers pick off the "botanical ticks" and, unlike the case with actual arthropods, put these in their mouths and upon biting down discover tha they just eat a tiny peanut. Arachis species (ie. peanuts or goobers) and those of Desmodium are in the same Leguminosae tribe, Hedyscreae. Ain't Botany interesting?

Ottawa County, Oklahoma. Late June.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

This is one of three Lactuca species known generically as "wild lettuce" that became well-established in the Ozark Highlands. All these species of "wild lettuce" do best in newly and frequently disturbed ground where they commonly thrive. In so doing they provide both forage from their foliage and small concentrates from their tiny achenes.

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

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

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

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

Ottawa County, Oklahoma. Early July.

Another naturalized commoner- Common sow thistle (Sonchos oleraceus) on an oak-hickory-tallgrass savanna about three-fourths of a mile from this clearcut. These plants (two plants in first slide; upper shoot with inflorescence of right plant in sescond slide) were on the edge of the same black oak-dominated forest in photographs of prickly lettuce immediately above. It is common for several species of Lactuca and Sonchos to grow side-by-side (or nearly so) in the Ozark Highlands, especially on harsher habitats such as those of shallow, rocky soils. The ridgetop of this clearcut forest was just such an environment (in contrast to the deeper and moister soil of the shaded north slope below this high point of the forest.

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

Ottawa County, Oklahoma. Mid-July.

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

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

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

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

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

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

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

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

Thus this transect includes SAF cover types 61 (River Birch-Sycamore, variant form) or variant of 94 (Sycamore-Sweetgum-American Elm), 26 (Sugar Maple-Basswood), and 52 ((White Oak-Black Oak-Northern Red Oak), the latter is indistinguishable in background. Ottawa County, Oklahoma (bank and bluffs of Lost Creek). March, early vernal aspect. Mesic bottomland forest according to Missouri Natural Areas Committee (1987). FRES No. 15 (Oak-Hickory Forest). No obvious Kuchler unit. Brown et al (1998) units of Mixed Hardwood Series in Southeastern Swamp and Riparian Forest biotic community. Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

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

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

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

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

The following slides and captions described a creek bottom hardwood forest in the Springfield Plateau section of the Ozark Highlands (Mountains) from perspectives of: 1) forest range and 2) plant succession or forest development (dynamics of a forest community). This forest range type was an example of the Society of American Foresters (Eyre, 1980, p. 65) cover type 93 (Sugarberry-American Elm- Green Ash). It was an old-growth forest, but in spite of some hugh trees in the forest the vegetation was at subclimax stage developing into the forest community that is climax for this forest site. Trees of the climax forest community were still young and much smaller than some of the immense individuals of subclimax species. This tract of forest had been undisturbed for decades and it was questionable if any woodcutting had ever been done in this forest other than that associated with clearing of a narrow fenceline along one side of the property line that had been done at least 60 years prior to time of photographs.

This forest range community was described--to partial degree or some extent-- by Nelson (1987, p. 52; 2005, p. 148) as mesic bottomland forest, but as shown below even when general descriptions were provided for the Ozark Plateau or Ozark Border there were inconsistencies between those and the vegetation that developed on this undisturbed tract of bottomland forest.

This forest developed in the flood plain of Modoc Creek between the old creek channel and the current channel. Forest range vegetation varied with distance from current and former creek channels. The dynamic development of forest vegetation was of such time scale as to a study in "still life", but this ecological drama in the forest stage presented one undeniable successional fact:: The Old Order Passeth Away...

52. The billboard cast of a bottomland forest- General view of a subclimax bottomland forest dominated by the aged or senior actors of sycamore and eastern cottonwood (Populus deltoides var. deltoides) with a supporting arboreal cast including hackberry or western hackberry (Celtis occidentalis var. canina), American elm, and pignut or bitternut hickory, slippery or red elm, chinkapin (chinquapin) oak, black cherry, black walnut, box elder (Acer negundo), red mulberry (Morus rubra), northern red oak, and Shumard oak (Quercus shumardii var. shumardii). The largest trees were individuals of sycamore, eastern cottonwood, and chinquapin oak though some trees of northrn red oak, Shumard oak and box elder were almost as large. There were more species of trees than of shrubs and the more common herbaceous species.

The most abundant tall shrub was pawpaw (Asimina triloba) which formed local groves or colonies from estensive rootstocks. Hazlenut (Corylus americana) was a taller shrub that was also present though at much lower cover and density. A lower shrub layer was dominated overall by spicebush (Lindera benzoin), but closer to the current creek channel American bladdernut (Staphylea trifolia) was dominant with dense colonies at local scale. Large woody vines of grape extended from ground to tops of canopies of the tallest trees. Grapes were of two species: summer grape (Vitis aestivalis) and fox or frost (V. vulpina= V. cordifolia).

The upper herbaceous layer was dominated by colonies of Virginia wildrye and silkly wildrye (local consociations). These cool-season grasses had green--though small--basal shoots throughout autumn and winter and became dormant by late spring or early summer. Other major herbaceous species were tall nettle (Urtica dioica var. procera), slender nettle (U. gracilis= U. dioica . var. gracilis), and wood nettle (Laportea canadensis), both members of the nettle family (Urticaceae) followed by purple or green-stemed Joe Pye weed (Eupatorium purpureum) which grew in groups of widely speced individuals, and lopseed. These forbs persisted throughout the warm-growing season. Another common and colony-forming forb (though one having shorter growing season) was Virginia waterleaf (Hydrophylloum virginianum) and bigleaf waterleaf (H. canadense). Thes spring-blooming forbs dominated the lowest level of the herbaceous vegetation layer in this bottomland forest range.

The species composition of this largely undisturbed (ie. direct human impact had been minimal to non-existent) bottomland forest was meaningful different from the currently most-apt descriptions of natural forest vegetation (Nelson, 1987 and Nelson, 2005 for mesic bottomland forest) that corresponded to the creek floodplain forest described here. Nelson (1987, p. 52) included western hackberry and bitternut hickory as dominant species and Shumard oak as a characteristic species. Nelson (2005, p. 148) omitted hackberry, but did list both American and slippery or red elm along with black walnut as dominant species. Neither Nelson (1987) nor Nelson (2005) listed chinquapin oak, box elder, or black cherry as even being present in mesic (or dry-mesic) bottomland forests in the Ozark Plateau or Ozark Border. Neither sugar maple nor white oak, dominant tree species according to Nelson (1987, 2005), were present in this creek bottom forest although sugar maple dominated an east-facing on the other side of Modoc Creek.

Furthermore, both versions of Nelson (1987, 2005) listed numerous forbs as herbaceous species while largely ignoring grasses. On the forest range described here two species of wildrye overwhelming dominated much of the herbaceous layer of this forest. Also, Nelson (1987, 2005) did not list purple or green-stemed Joe Pye which was one of the dominant forbs of this bottomland forest. Virginia waterleaf was listed (Nelson, 1987, 2005), but this was a minor species--both spatially and temporally--compared to the wildryes, Joe Pye weed, and tall nettle.

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

Critical qualifying observation: the fungus (Ophiostoma novo-ulmi)-caused Dutch elm disease spread by Elm bark beetles (Hylurgopinus rufipes and Scolytus multistriatus) had catastrophic impacts on both American and slippery elms in the area of this climax hackberry-American elm bottomland forest. Elm phloem necrosis is another common disease of these two elm species in this same area. This disease is caused by a mycoplasma-like organism ( a kind of virus) for which the whitebanded elm leafhopper (Scaphoideus luteolus) is the vector. Both of these diseases commonly kill many elm trees throughout this region of the western portion of the eastern deciduous forest formation.

In fact, it is often difficult for other than forest pathologists or other trained specialists to tell whether a given elm died of Dutch elm disease or elm phloem necrosis. For unknown reasons neither of these two widespread diseases that are common to the Ozark Highlands (and that have destroyed millions of elms) was not a factor--at least, not enough of a factor-- to eliminate American elm and or the less common slippery elm from the forest described below. This is not to say that either or both of these diseases were absent, but only that they did not prevent American elm from becoming the co-dominant of this climax forested range plant community.

53. The stage and introduction of the principal actors- An old (apparently the prior) channel--bed and banks--of Modoc Creek was evident on this bottomland (floodplain of Modoc Creek) forest range. Growing along the banks of the earlier channel were immense individuals of sycamore, eastern cottonwood, northern red oak, and Shummard oak. At greater distance from the old channel banks (as well as from current channel) there were chinquapin oak, box elder, and some(fewer) white ash (Fraxinus americana). Hackberry (also designated as western hackberry; Celtis occidentalis) and American elm plus some box elder and bitternut hickory were the primary species of younger trees (including those with on-going regeneration). These were growing by the old (prior or former) stream banks and outward from them (= landward from stream) as well as in between both old and current stream channels. Red mulberry was present as a smaller tree in the taller shrub/sapling layer along with American bladdernut and American hazelnut.

Sycamore and eastern cottonwood (typically pioneer species or colonizers) unquestionably had the oldest (and largest) trees, but it would be erroneous to equate all differences in size to differences in age as this would ignore different rates of growth (or, same thing, assume equal growth rates) among tree species. Obviously such was (is) not the case. For example, eastern cottonwood is one of the fastest-growing hardwood species in North America (Burns and Honkala, 1990, p. 530). Regardless, it was self-evident that sycamore and eastern cottonwood had become established on bare gravel or soil on banks of the old channel and subsequently persisted into the subclimax (or early climax) stage of this bottomland forest.

Large specimens of the two grape species grew up to tops of tallest sycamores and cottonwoods. Notable by their absence in this plant community were other species of woody vines such as Virginia creeper, trumpet creeper, and (more-or-less) poison oak/ivy, There were some individuals of bullbriar or catbriar (Smilax bona-nox). Other shrubs included pawpaw (the dominant and colony forming, taller shrub), spicebush (overall major shrub) and American bladdernut, the two species that costituted a middle shrub layer, and buckbrush or coralberry which formed the sporadic or discontinuous lowest shrub layer.

Throughout most of the growing portion of the year--both cool- and warm-seasons--there was a single herbaceous layer extending to a height of three to four feet. This was dominated during the cool-season portion of the growing season by local--typically separate--colonies of Virginia and silky wildryes. Elymus species were in dormancy during most of the warm- growing season (typically dormant by late spring or early summer). Dominants of the herbaceous layer during summer and sutumn were three forbs (all of which grew in colonies or local consociations): tall nettle, wood nettle, and green-stemed or purple Joe Pye weed. Virginia waterleaf and a few incidental forbs made up a lower herbaceous layer in late winter to mid-spring.

Various species of fuungi and lichen grew on downed logs and limbs and on rotting leaves on the soil surface.

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

54. Curtain rise as the cast rises from the old channel- Vertical view down a former channel of Modoc Creek along which pioneer sycamore and eastern cottonwood persisted into the late subclimax or early climax stage of vegetation that developed into ths moist bottomland forest. At some later stage(s) following the pioneer plant community northern red oak and Shummard oak became established along the sere of this forest vegetation. These trees also persisted into the current subclimax or climax range plant community. Large woody shoots of summer grape and fox or frost grape extended into the crowns of these trees, especially sycamore and eastern cottonwood.

The two smaller (younger) foremost trees (left and right bank of previous creek channel) were hackberry. The tall shrub to right of the left bank hackberry was American bladdernut. Large tree in center midground was a dead sycamore. A pole-sized hackberry was growing to left of this sycamore. Large tree in background was eastern cottonwood.

Shrubs in this view of the bottomland forest were spicebush and buckbrush or coralberry.

Straw in the herbaceous layer of the understorey was of recently gone dormant Virginia and silky wildryes. Green leaves in the herbaceous layer were those of purple or green-stemed Joe Pye weed, tall nettle, and wood neetle.

Some of the lower green leaves were those of seedlings and small saplings of western hackberry and American elm (the climax tree species with most sexual reproduction), bitternut hickory (a less abundant climax tree species), and, least of all, box elder.

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

55. Oldsters along the old channel: the old order, and the new- A previous stream channel of Modoc Creek was still the home to pioneer sycamore and eastern cottonwood (large trees in background) as well as northern red oak (eg. second left-leaning tree trunk on right bank) and Shummard oak, but this old order was giving way in the progression of plant succession to hackberry and America elm (eg. the nice, big, foremost left-leaning tree trunk on right bank), and bitternut hickory. The left of center tree with gray bark in far foreground or near midground was chinquapin (chinkapin) oak. The successional status of chinquapin oak was unknown, but its presence suggested a status of subclimax to climax on this forest range site. Chinquapin oak was regarded as Intolerant and yet to be subclimax to climax on mesic, limestone-origin soils (Burns and Honkala, 1990, p. 699), en edaphic condition met on this site. Large woody vines of summer and fox grape traced across the old stream channel into crowns of the tallest trees.

The straw was that of colonies of Virginia wildrye and silky wildrye that were dormant at this point in early summer. Most shrubs were spicebush, coralberry or buckbrush, and American bladdernut. No forbs were visible in this forest range scene.

This bottomland hardwood forest range was in the western Ozark Highlands (Mountains) just a few miles from eastern edge of the Cherokee Prairie of the Central Lowlands physiographic province. Species like eastern cottonwood, American elm, chinquapin oak, and the wildryes were botanical proof of the affinity of these two distinct yet contiguous floristic regions.

The bottomland habitat on which this forest had developed was formed when the perennially flooding Modoc Creek underwent a large enough flood and other "just right" conditions to form another channel which left the former creek channel high and dry. At that time (when the new--the latest or more recent--channel of Modoc Creek was carved in its floodplain), the old channel became just a geologic reminder of the dyamic nature of streams. The former riparian habitat (a kind of wetland) along the banks of Modoc Creek was changed into a slightly elevated, mesic habitat that ultimately became nothing but a rise with a "haired-over" (vegetated), meandering, dry ditch down the middle. The pioneering sycamore and eastern cottonwood and the somewhat later invading oaks (northern red, Shummard, chinquapin) continued to grow on banks and bed of the previous channel of Modoc Creek. But the growing and, eventually, large trees of these early seral species did reproduce (sexually or asexually). They did not replace their own kind in the shade they cast (ie. they did not replace themselves in the habitat they modified). The individual plants of these species had modified their environment (affected soil, air currents in the forest, God only knows what else) so that the changed habitat was "improved" (made more amenagle) for the next stage on this sere which, in this case, was the climax.

This was the phenomenon of reaction in the Clementsian model of plant succession or what was later labeled the facilitation model by (Connell and Slatyer, 1977). The other relevant phenomenon of the Clementsian paradigm was competition. The pioneering and persistent (through longevity) sycamores and cottonwoods--large though they were and commanding the canopy as they did-- still, and simply put, could not compete effectively with western hackberry, American elm, bitternut hickory, box elder or, even, black cherry. The latter were the new--and presumedly final order--on the sere of this floodplain forest range. The former (subclimax) dominants were the old order that, having had their successional moment in the sun, surrendered the "ground" they had modified (reaction in action) to the final cast of characters at termination of this sylvan play. These as-the-curtain-falls actors were the dominants of the potential natural vegetation, the climax forest that bore their names as that of the forest cover type.

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

56. Views down the channel: "The Times They Are A'changin'"- The title line from the old protest song of Bob Dylan (1964) summed up the dynamics of this forest range vegetation. These two photographs provided a farther and a closer view of species composition and structure of a mesic bottomland forest in the western portion of the Ozark Highlands by focusing on the forest community that had developed on the old (previous) channel of Modoc Creek. Both the long and short focal perspectives provided a classic example of change in forest plant species with progression of plant succession. This was an example of the Clementsian model of dynamic vegetation.

The largest tree (right center midground) was an old-growth specimen of northern red oak. The large tree to right of this northern red oak, the upper trunk of which leaned right, was an old-growth Shummard oak. Incidentially both of these oak species are generic red oaks (Erythrobalanus subgenus of Quercus). The two trunks opposite of the old-growth northern red oak (left side of creek channel) were sycamore. So much for the pioneer sycamore and later seral red oaks, these stalwarts of the old order. The new order--the forest climax--which, in the grand scheme of forest development (plant succession; the dynamics of vegetation) was ousting the Old Arboreal Guard, was that of western hackberry (the major climax dominant), American elm, and bitternut hickory.

Box elder and, to less extent, black cherry were also well-represented by younger trees of large pole-size but, like chinquapin oak in the preceding photograph, the successional status of these two species was unknown, although they appeared to be subclimax or climax. Black cherry was classed as Intolerant (Wenger, 1990, p. 3), yet it "grows very fast" (Burns and Honkala, 1990, p. 599). Perhaps rapid growth was an adaptation that enabled black cherry to survive in this bottomland forest. Furthrmore, in this author's observation, black cherry propagates readily in the understorey of oak-hickory and bottomland forests in the western Ozark Plateau and, as was shown below in the Use and Abuse portion of this chapter below, factors such as fire and browsing prevented greater density and cover of black cherry in these forest ranges. A tolerance rating of Intolerant did not seem appropriate for black cherry on these Ozark forest types (including this largely undisturbed, creek bottom forest).

In the first of these two photographs there were two western hackberry trees growing in fromt of the sycamores (left midground) while a straight-trunked black cherry grew in right foreground. In the second (closer-in or short view) photograph only one of the two foremost hackberry trees was visible. In both photographs the hackberry growing to left and slightly behind the old-growth northern red oak was visible. Hackberry was clearly in the successional ascendency.

Most shrubs in these photographs were spicebush (eg. center shrub--and in center of old creek channel--in second of these photographs) with buckbrush or coralberry being the second most abundant in these views of this forest vegatation (eg. foreground of first slide by black cherry).

The straw was that of recently gone-dormant Virginia and silky wildryes.

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

57. The action scene: changing of the guard- In a bottomland (floodplain) forest along a stream (Modoc Creek) in the Ozark Highlands sycamore and eastern cottonwood had pioneered this former creek channel (now esentially the first stream terrace) and were being replaced by the climax dominants which were hackberry, American elm, and bitternut hickory (often a north-slope dominant with sugar maple). Box elder and chinquapin oak appeared to be other climax (at least, subclimax) tree species on this site along with climax spicebush, American bladdernut, pawpaw, Virginia and silky wildryes, purple Joe Pye weed, tall nettle, and wood nettle. Successional status of black cherry on this creek bottom forest site was not known. On drier upland habitats buckbrush or coralberry was locally common. On moister drainages into the current stream channel American bladdernut was a local dominant shrub.

Two photographs presented a long view (first slide) and a shorter or closer view (second slide) of this floodplain forest vegetation. In the center foreground (both slides) was a pioneer (ultimately a large and very old) sycamore that had died two years prior to time of photographs (a two-year old snag). The two-year sycamore snag died without replacement progeny was being replaced instead by: western hackberry (young sapling immediately to right of the bark-exfoliating snag; also, tree at right margin of foreground in the first slide only), bitternut hickory (tree immediately to left of snag; also leaves of sapling in left corner foreground), and American elm (tree to left of the bitternut hickory that was immediately to left of snag) along with fewer and more scattered box elder. The shrub in front of and slightly to left side of sycamore snag was spicebush, the dominant shrub in this creek bottom forest. Behind the sycamore snag with its flaking bark and the young hackberry growing immediately to right of the dead sycamore (center background of both slides) was a pioneer tree of eastern cottonwood. Straw in foreground extending back to midground consisted of colonies of Virginia and silky wildryes, the former somewhat more abundant.

Long-lived sycamore and eastern cottonwood persisted into the subclimax or early comix forest much Douglas-fir foes on certain forest cover types. Sycamore and cottonwood were not regenerating on this bottomland site. Tolerance ratings of sycamore, eastern cottonwood, American elm, and box elder by the Society of American Foresters (Wenger, 1984, ps.2-3) were Intermediate, Very Intolerant, Intermediate, and Tolerant, respectively though with some uncertainty for sycamore and American elm.

Like the preceding slides this was a two-photograph sequence that provided both a long and short view of mesic bottomland forest range with an emphasis on dynamics of vegetation with the progression of forest development (ie. community and species changes with progression of plant succession).

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

58. Close in scene of the leading characters-Third photograph in a series with the same point of focus to provide consecutively greater detail on the featured plant, a two-year-dead sycamore. "Old as a big tree" (or "as big as an old tree") was not old (or big) enough enough to save this old sycamore from the ultimate end of all things mortal or, more meaningful from the perspective of vegetation, from the inevitable changes as a sere progresses through plant succession to the climax, the final or terminal plant community for that forest or range site. This large sycamore, which was introduced in the immediately preceding two-slide set, had pioneered the fresh, denuded sere on the bank of a new-cut channel of Modoc Creek decades, a century, (or however long) ago. Since the creation of that channel, Modoc Creek flooded again and formed another channel resulting in abandonment and isolation of the previous stream channel. The by-now established sycamore and eastern cottonwoods (probably along with northern red, Shumards, and some chinquapin oaks) continued to grow on the land of the old channel which remained as a terrace of the Modoc Creek floodplain.

As geomorphic and successional time passed the pioneering sycamore and eastern cottonwood and the somewhat-later invading oaks (northern red, Shummard, chinquapin) continued to grow on banks and bed of the previous channel.The growing and, eventually large, trees of these early seral species did not reproduce (sexually or asexually), at least not at rates sufficient to be competitive with more reproductively effective tree species. The pioneering and seral tree species did not replace their own kind in the shade they cast. Instead these trees modified their habitat so that other (later-appearing) species of trees out-reproduced the early colonizing and other seral tree species. Older and bigger individual sycamores, eastern cottonwoods, and red oaks persisted in the forest, but they did not replace themselves in the habitat they modified. At least, they did not replace their own species as prolifically as did the tree species that were newer to the sere. The successionally more advanced species produced more progeny than the earlier seral species Successionally advanced species were more competitive because they produced (and continued to produce) more offspring so that as old trees of sycamore, eastern cottonwood, northern red oak, and Shumard oak died the remaining members of their respective species made up ever-smaller proportions of the forest vegetation, especially in the light-controlling canopy. With these disproportionate rates of regeneration over a long enough span of time hackberry, American elm, bitternut hickory, and box elder eventually replaced (or largely so) sycamore, eastern cottonwood, and the red oaks.

This on-going struggle for control of the bottomland Ozark forest--and for survival of species on the sere of this floodplain forest site--was poignantly visible in this third photograph that featured the sycamore snag and successor species. In addition to the previously shown hackberry sapling (immediate right of snag), bitternut hickory (left of snag), and American elm (left of bitternut hickory) there were more western hackberry (two at far left midground and more behind and to right of them) and bitternut hickory (eg. left leaning tree behind and to left of sycamore snag). Several of the trees farther back were box elder. Again, shrub in front and slightly to left of snag was spicebush, the dominant shrub species. Also shown again was one of the pioneering eastern cottonwoods (right background).

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

59. A view from the opposite side of the stage: enter the new order- The featured two-year-old sycamore snag, the "landmark" of this series, was shown from the other end of the former channel of Modoc Creek to present other species in "successional ascendency" on this sere of an Ozark Plateau bottomland forest. Besides the previously referenced sapling of western hackberry (left front of snag) and faded straw of Virginia and silky wildrye, this view featured a box elder (crooked trunk in left foreground), a Tolerant species that was ascending into the climax forest vegetation. Readily visible leaves of hackberry (upper left) and bitternut hickory (blurred; upper center) along with flaking bark on the sycamore snag emphasized the Clementsian dynamics of this forest vegetation. Forbs in left midground (to left of snag) were a colony of purple or green-stemed Joe Pye weed.

This was an opportune place to re-emphasize that this was--based on local folk knowledge and biological-physical evidence (eg. absence of stumps and slash)--a humanly undisturbed forest. It was an out-of-the-way, isolated tract of forest protected by its owner and a limestone bluff.

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

60. Easier to see the story- Absence of leaves on trees and shrubs in winter vegetation made it easier to "read ecology" of the western hackberry-American elm-sycamore-boxelder bottomland forest described above. The previous (old) channel of Modoc Creek was shown to good advantage revealing development of a floodplain hardwood forest. Sycamore like the large, old-growth individual in right foreground and eastern cottonwood, an example of which was the big tree in center background, had pioneered banks and bed of the previous and channel of Modoc Creek after this Ozark Plateau stream flooded and cut a new channel in the distant past. This geologic event left the now-abandoned channel to undergo primary plant succession beginning with colonization that included sycamore and eastern cottonwood. The individuals of these species along with some boxelder, which pioneered the raw site (along with typical annual forbs and grasses), persisted into the climax bottomland forest that at termination of plant succession was dominated by western hackberry, American and red (slippery) elm, bitternut hickory, and black cherry. Old individuals of chinquapin and Shummard oaks were also present (persisted) to the climax stage.

Chinquapin oak was represented in this photograph by the tree with firescar on left bank of old channel in midground while two boxelders were in background immediately in front the old-growth cottonwood. These trees and this immediate area were presented in the immediately succeeding slide. Large lianas of summer and fox grape graced the old channel.

The dominant herbaceous species in the forest community presented in this photo-plot was silky wildrye. Also present in the herbaceous understorey as major forb species were Eurasian common chickweed (Stellaria media), the native chickweed (Cerastium brachhypodum), and false rue aneomone (Isopyrum biternatum).

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

61. The old channel in winter- Another photo-plot of the western hackberry-American elm-sycamore-boxelder bottomland forest described in preceding photographs. This forest community developed along the former channel of Modoc Creek after this stream cut a new channel. Featured here was the forest range vegetation shown in the midground of the preceding photograph. Largest tree in background with forked trunk was an old-growth eastern cottonwood that pioneered the former stream channel. The tree in left foreground with basal firescar was a chiquapin oak. The two trunks in front and slightly to left of cottonwood were boxelders. Large grape vines drapped down into the abandoned channel bed. Understorey was not prominent here, but was well-developed elsewhere in this forest. Details of understorey were presented below.

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

62. Browsing in the bottomland forest (nobody said that replacing the old order would be easy)- A pole-size (roughly less than seven inches basal diameter) young hackberry (western hackberry) growing in the bed of the former channel of Modoc Creek was felled by bark-feeding beaver. The hackberry had resprouted from the stump. On this bottomland forest range site hackberry was (is) the climax co-dominant (with American elm) or, sometimes, tri-dominant with bitternut hickory and/or box elder. In this mesic bottomland forest hackberry is the foremost ("first among equals") dominant and defining species of the climax (potential natural) forest vegetation.

Plant life in this "photo-plot" was a cornucopia of Ozark forest species including (besides sprouts of hackberry): poison ivy/oak, green-briar (catbriar or bullbriar), lopseed, pokeweed, silky wildrye, seedlings of chinquapin (chinkapin) oak, American bladdernut, and a caric sedge. Students can practice their botanizing skills by trying to identify these (probably other) species.

In the old stream channel of Modoc Creek, Ottawa County, Oklahoma. July (estival aspect).

63. Being dominant does not mean "being out of the woods"- In the bottomland forest described here a large pole- or small log-sized American elm had been girdled by beaver. Beaver had not completely girdled this elm and instead had left a sizable part of its bark intact (on other side of the trunk as presented here). Thus, there was still some hope for survival of this tree. Plus, it was still small (=young) enough that it probably had the capacity to sprout or sucker (produce shoots from portions of the shoot near ground or from the rootcrown, the latter of which seemed more likely given that beaver gnawed this tree to ground-level).

There were other (smaller) neighboring saplings of both American elm and other species (most notably, hackberry) ready to grown--and possibly escape beaver-feeding--should this pathetically debarked individual cash in its chips (made by beaver of course).

Authors 38 inch hickory walking showed height of bark-feeding by beaver.

IMPORTANT POINT: This and the preceding photograph showed that beaver fed readily and extensively on hackberry (western hackberry) and American elm, the two major climax (= typically dominant) tree species of this Ozark Plateau bottomland forest. Interestingly, beaver like graziers such as cattle, buffalo, elk, and horses preferred the dominant climax plants. On this forest range, under browsing by beaver, hackberry and American elm were decreasers. By contrast, on upland, mesic-limestone forests (including one adjacent--contiguous with except for the channel of Modoc Creek--to this tract of bottomland forest) beaver did not select sugar maple and white ash which were the climax dominant and associate tree species, respectively. Instead, beaver browsed selectively on northern red oak in preference to sugar maple and white ash which stood trunk-by-trunk to northern red oak.

Floodplain of Modoc Creek, Ottawa County, Oklahoma. February, 1990.

64. Now they've turned to woody vines- Browsing on fox grape (Vitis vulpina) by beaver in late autumn to early winter. The liana of a moderate-sized fox grape that was stripped of its bark and gnawed clear through into several sections by browsing beaver (first of these two slides) along the banks of Modoc Creek in the western hackberry-American elm-sycamore-boxelder-chinquapin oak-northern red oak floodplain forest introduced above. The second slide was a close-up view (from the opposite of liana) of the right-most section in the first photograph.

Greening shoots of silky wildrye, the dominant herbaceous species of this forest range, were visible in left midground of first photograph.

Ottawa County, Oklahoma. Early January.

65. Can't keep a good grape down- First-season stump sprouts (basal shoots) from the fox grape gnawed off by beaver in the two preceding photographs. Size and extent of resprouting of fox grape (regrowth from basal buds) about three months into the first warm-growing season following complete topkilling by browsing beaver. Also in this "photoplot" were many seedling of western hackberry, the climax dominant species of this bottomland forest range. Virginia creeper and Davis caric sedge (Carex davisii) were also well-represented in this sample of vegetation on the forest floor.

Ottawa County, Oklahoma. Mid-July.

66. Winter vegetation- Range vegetation of a bottomland (floodplain) hardwood dominated by western hackberry, American (some slippery) elm, and bitternut hickory (with persistent sycamore and eastern cottonwood) in winter. Boxelder was also present as both large (presumedly pioneering individuals) and medium-size trees. The large woody vines (lianas) were summer and fox grape. Larger tree at extreme right margin and midground of first photograph and again in midground of second slide was western hackberry. Almost all trees in midground were hackberry. The tree with bent trunk (to right of the larger hackberry in midground) was boxelder. Most of the trees in background were sycamore (readily distinguished by upper gray bark) growing along bank of Modoc Creek, a typical Ozark Plateau stream. There were not oaks or cottonwood in this stand.

Most of the shrubs were spicebush (Lindera bezoin), but some pawpaw (Asimina triloba) grew in shrub understorey closer to Modoc Creek. Both the shrub layers and herbaceous layers were described immediately below.

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

67.Lower layers in winter- Shrub and herbaceous layers in a bottomland forest dominated by western hackberry, American (and some slippery) elm, and bitternut hickory (with persistent sycamore and eastern cottonwood) in winter. There were two pronounced shrub layers: 1) tall shrub dominated and made up mostly of pawpaw and 2) lower shrub dominated by spicebush with buckbrush or coralberry (Symphoricarpos orbiculatus) as an associate of this layer. The conspicuous shrub in this slide was spicebush.

The herbaceous zone also consisted of a upper or taller layer and a lower of shorter layer. The herbaceous zone in winter (hibernal aspect) was presented and described in the next series of two photographs.

Floodplain of Modoc Creek, Ottawa County, Oklahoma. January; hibernal aspect. Vegetational units cited in immediately preceding caption.

68. The lowest layers- Details of the herbaceous zone of a bottomland, hardwood forest described above. At stages of maturity of these herbaceous species and peak herbaceous standing crop there will be at least two layers: 1) low herbaceous layer made up of two Eurasian annual forbs, common chickweed ( Stellaria media) and dead nettle (Lamium purpureum), the native chickweed (Cerastium brachhypodum), both bigleaf waterleaf (Hydrophyllum canadense), and Virginia waterleaf (H. virginianum), and false rue anemone (Isopyrum biternatium) and 2) tall herbaceous layer comprised of Virginia and silky wildrye and, later in summer, forbs the major of which were sweet, purple, purple-jointed, or green-stemed Joe Pye weed (Eupatorium purpureum) along with wood nettle (Laportea canadensis), slender nettle (Urtica gracilis= Urtica dioica ssp. gracilis ), and tall nettle (U. procera= U. dioica var. or ssp. procera).

Floodplain of Modoc Creek, Ottawa County, Oklahoma. January; hibernal aspect.

69. Use of the understorey dominant- A plant of silk wildrye, the dominant understorey herbaceous species in the bottomland forest in which hackberry and elm were replacing sycamore and cottonwood as dominant trees. This particular plant had been grazed (defoliation was evident in this photograph) by white-tailed deer, the only ungulate, on this winter range. Silky wildrye matures and goes dormant by mid to late June by which time the only remaining herbaceous species on this forest range were forbs including purple-jointed or green-stemed Joe Pye weed, wood nettle, and slender nettle. The herbaceous portion or component (layers) of the understorey of this bottomland forest clearly comprised winter and spring. Winter usage was more critical and valuable given general scarceness of native, cool-season herbs.

Virginia wildrye was almost as abundant locally as silky wildrye, but this latter was slightly more plentiful.

Ottawa County, Oklahoma. January.

70. Winter forb in the woods- The dominant perennial forb in understorey of bottomland hardwood forest was false rue anemone (Isopyrum biternatium) of the Ranunculaceae (buttercup family). The plants in the first two of these three slides were were growing and blooming in January and were photographed at the same time the author recorded tree damage from a severe ice storm (presented immediately below). The plant in the third of these slides was growing in this same location in March. I. biternatium is of necessity an opportunistic bloomer. This cool-season forb has a life cycle pattern (adaptative "strategy") that permits it to survive and reproduce in what would otherwise be a deeply, densely shaded understorey from spring through autumn leaf-fall.

Floodplain (terrace) of Modoc Creek, Ottawa County, Oklahoma. Full-bloom phenological stage.

71. Browsed by ice- Defoliation is the generic term for removal of plant parts by such factors or agents as grazing/browsing animals (including insects), diseases, hail, frost and freezing temperatures, drought, wind (in a myriad of forms), and sheet ice and snow. Obviously hail, ice (such as from freezing rain and sleet), and snow are all forms of frozen precipitation. Whereas hail defoliates as it falls, ice and build-up of snow defoliates only after accumulations reach weights that plant parts can no longer support thereby resulting in breakage and separation (to one degree ir the other) of those parts from the remaining body of the plant. Ice, itself, can fall directly as frozen precipitation in form of sleet (rarely, grupel), freezing rain which is rain that freezes on contact, or rain that subsequently turns to ice at temperatures of freezing and subfreezing over periods of time.

Damage (defoliation) from ice accumulations is both direct and indirect. The latter occurs when branches, limbs, or crowns of taller trees that were brought down directly by excessive weight in turn (secondarily) break off, crush, or uproot lower, smaller trees and shrubs when the heavier and larger tree material from above crashes down on "underlings". Likewise, there is both primary and secondary damage to trees and shrubs suffering direct and/or indirect defoliation. Primary damage occurs from loss of plant parts whereas secondary damage takes place in numerous ways including disease entry via wounds, general weakening from severe or excessive ice-prunning, and heat from any fires (such as those ignited by lightening from thunderstorms) which will burn much hotter due to greater fuel loads from downed timber.

Damage done by severe and extreme ice storms can rival that from smaller tornados and hurricanes. Ice storms sometimes vastly exceed in area that of the largest twisters so that the scope of and total damage wrought by ice storms can exceed that of violent tornados. Such was the case of this storm (9 December, 2007) which cut a swath about three hundere miles long (west to east) and one hundred and fifty miles wide (south to north).

These two photographs showed severe damage done by that storm in a bottomland (floodplain of Modoc Creek) hardwood forest that was pioneered by sycamore, eastern cottonwood, and boxelder and that was approaching climax state in which dominant trees were western hackberry, American (white) and slippery (red) elm, and bitternut (pignut) hickory. Most severely damaged was boxelder. Large limbs that broke under weight of ice typically split and down their center (pith and heartwood) so as to "peel" or "run" down the limb and even trunk for long distances thereby magnifying extent of tree injury. An example of that pattern of breakage was obvious in the large limb of boxelder seen in center midground of first slide and centr foreground of the second of these slides. Shumard and black oaks and eastern cottonwood suffered considerably less limb breakage. Tolerance to ice damage in eastern cottonwood seemed strange given its soft wood, but the author consistently observed this outcome in numerous forest and shade trees in this immediate area.

In the example of severe ice damage presented in these two photographs a large limb (making up almost half the tree's crown) of a boxelder crashed and broke over (rather than completely off) a smaller American or white elm in the lower tree layer. The severed bole of the was shown only in the first photograph. Break-over of the American elm was presented at closer distance in the second of these photographs. The tree in right midground that was completely topped and the toppled portion in foreground was an American elm that had previously been burnt clear through by a surface fire (fire scar distinctly visible). Damage from this fire had not extended upward to height at which this "born-loser" elm broke off. The point on this American elm at which regrowth occurs will be interesting. From intercalary meristem at point of breakage or stump sprouting?

The two saplings in foreground were western hackberry readily identified by the distinctive corkly ribbed bark.

On this bottomland hardwood forest western hackberry, American and slippery elm, and bitternut hickory were succeeding sycamore, eastern cottonwood, and boxelder as climax dominants. It seemed likely that damage from this ice storm would slow this successional development and perhaps create enough disturbance that pioneer and other seral tree species might have relatively more reproduction and reclaim more of their former crown cover. All-in-all another example of Frederick Clements "dynamic vegetation".

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

72. Iced down- Large limbs brought down by ice accumulations (3/4th inch to slightly less than two inches) resulting from a mixture of rain, freezing rain, sleet, and light snow (in that order) from a late autumn (9 December, 2007) storm produced when warm, moist air from the Gulf of Mexico overrode the heavy, cold, dry air of a Norther (an Arctic polar front). When the wet Gulf air cooled enough at higher altitudes it released the water which it could not hold at colder temperatures. Then as this precipitation fell through the heavy, cold Arctic air it went through varying frozen states that fell and accumulated on objects such as trees (power lines, roofs of houses, etc.) the added weight of which broke and brought down numerous of these things.

In the scene shown here damage to (defoliation of) sycamore, like the large limb at left, and boxelder, represented by big limb at right, was featured. Boxelder sustained disproportionately severe damage due to its soft, brittle wood. The standing trunk in right foreground was chinquapin oak, a species that withstood icing better than western hackberry, elms, and, of course, boxelder.

The understorey of this bottonland forest was presented and described in detail in several preceding photographs-captions.

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

73. By the sheer weght of it- Two western hackberry in a bottomland forest uprooted by weight of ice accumulation from a late autumn ice storm (9 December, 2007). A combination of heavy ice accompanied by high winds and saturated (or nearly saturated) soil, aided and abetted by falling limbs from neighboring trees resulted in partial uprooting and toppling of these two pole-size (DBH approximately 14 inches) hackberries.

In this creek floodplain forest in the western Ozark H:ighlands hackberry, American and slippery elms, and bitternut hickory were replacing sycamore, eastern cottonwood, and Shumard oak that had pioneered the old channel of Modoc Creek and adjacent land when this mid-size creek changed course sommetime over a hundered years ago. Other hackberry had lost major limbs and even their entire crowns (see next slide). Sycamore, Shumard and chinquapin oaks, and black cherry were also similarly impacted while bitternut hickory (and sugar maple in a neighboring creek bank stand) were least affected. Boxelder with its extremely brittle wood suffered the most ice damage (see photographs and discussion of boxelder breakage below).

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

74. Crowned- The upper trunk of a western hackberry was broken by ice accumulation from a late autumn (9 Decembr, 2007) storm of wind and mixed precipitation (that promptly froze and built-up on trees). In crashing, the crown of the hackberry broke and bent down a sapling of American elm (to left of hackberry) The most extensive damage was to boxelder, but sycamore, chinquapin oak, black cherry, and Shumard oak were all affected by the storm and suffered major loss of limbs and branches. Centermost tree in foreground was a hackberry that experienced only minor damage.

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

75. No, not a tornado; ice- Ice can be as effective as a tornado, hurricane, or straightline wind blowdown through several manisfestations. This includes uprooting, complete toppling by breakage near ground level, removal of entire crown by breaking below lowermost limb, or extreme breakage of individual branches and limbs. An example of the latter was provided by this photograph.

This stand was a dry bottomland forest of western hackberry, American and slippery elm, and boxelder with black walnut, chinquapin oak, and honey locust as minor but consistent tree species. The largest bole in this stand was hackberry (right foreground). Trees in left foreground were (left to right): hackberry, American elm (tree with upper forked trunk), and boxelder. The large, split limb in front of largest hackberry was from a boxelder immediately to right (out of photograph) of the hackbery.

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

76. Details of defoliation by ice- Limbs and even half-crowns downed by heavy accumulations of ice (3/4th inch to slightly less than two inches) from the most severe ice storm (9 December, 2007) in over a century (determined from records and reported by Empire District Electric Company, Jopoin,Missouri). Most of the debris on the forest floor shown here was boxelder, but American and red (slippery) elm, western hackberry, and honey locust contributed to the tangle of downed timber.

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

77. Collapse of a patriarch created a gap- At edge of the bank of the former creek channel in an Ozark Highlands floodplain forest a large Shumard oak reached the end of its silvan rope. The base of this tree (immense by standards of the Ozarks and this species) had rotted away to "almost nothing" long before wind and gravity teamed up to bring it down. The tree species "standing to benefit" from this windfall of light included two sycamore (two largest trees), a sapling of American elm, hazelnut, and red mulberry.

The "hollow heart" of this Shummard oak was similar to that of the large (and largely hollow) northern red oak killed by bark-feeding beaver that was described above.

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

78. Left a hole- Example of a forest gap created by death of an old-growth Shumaard oak along the bank of the abandoned (former) channel of a stream (Modoc Creek) in the western portion of the Ozark Highlands. With its unbelieveably rotten trunk (for a still-standing tree) this long-abiding master of the woods crashed to create an opening in the forest canopy and set the stage for the phenomenon of patch dynamics.

The first of these wo slides of this forest gap provided an opportunity to present the exterion of the mesic bottomland forest in the Ozark Plateau just described. Other large trees whose crowns dominating the canopy were sycamore, eastern cottonwood, northern red oak, chinquapin oak and (though one less) Shumard oak.

The second slide showed details of part of the immense crown of the Shumard oak along with plant species at the outer edge of this bottomland forest. The oak fell outward from the forest so that its "final resting place" was a field that had once been a prairie that was exterior to the floodplain forest that developed along Modoc Creek. This was part of the Prairie Peninsula. The major shrub in the vegetation shown here was American hazlenut. The major (and rather large) herbaceous species were two composites: cup-plant or cup rosin-weed (Silphium perfoliatum) and yellow crown-beard (Verbesina alternifolia). These were the main ecotonal species and the ones to benefit immediately from this ecological "windfall".

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

79. Start of a forest (forest colonization)- A small old-field at edge of the botttomland hackberry-American elm-sycamore forest featured herein (and visible in background) midway through the second warm-growing season following abandonment from grain-farming. This was (had been) a little patch of a wheat field too small to pay for farming so tenant farmers just "let it go back". It was "go-back land" with a twist. Secondary plant succession followed the textbook sequence for such land in eastern North America (the vast region of the eastern deciduous forests) with hairy crabgrass (Digitaria sanguainalis), giant ragweed (Ambrosia trifida), and horseweed or mare's tail (Conyza canadensis)--the usual dominant, r-selected, pioneer, weedy plant species--being everywhere. However, seedlings of sycamore and box elder were the most common plants from the criterior of relative foliar cover. Also, there was a lot of cover of two perennial herbaceous species, one a native grass and the other and introduced and now widely naturalized legume. These were purpletop (Tridens flavus) and sericea lespedeza (Lespedeza sericea= L. cuneata), respectively:.

This small parcel of old-field showed that on this bottomland forest range sycamore and box elder were colonizing or pioneering species (along with the sterotypic annual weeds of abandoned farmland). This parcel of "go-back land" was contiguous with the former channel of Modoc Creek on which old-growth sycamore and eastern cottonwood still persisted from when they had pioneered that new bank at a time known only to God. This little, former, wheat field was the next stand of long-lived, persistent, and, ultimately, large trees which will eventually be replaced by western hackberry and American elm. The sycamores and box elders will--barring introduction of more diseases or a major disturbance like wind, fire, or ice--will persist as part (constituents) of the climax bottomland forest dominated by hackberry and American elm. It was noted above that for whatever reason American elm (along with some slippery elm) was not killed out by Dutch elm disease on this tract of forest.

It was not determined definitively why there were no eastern cottonwood seedlings accompanying those of sycmore and box elder. Most of the old-growth cottonwoods in the immediate vicinity of this "go-back land" were male plants which would have resulted in reduced availability of cottonwood seed. This fact did not seem to be the sole cause for lack of eastern cottonwood in the pioneer stage of this forest range vegetation.

Part of the greater floodplain of Modoc Creek, Ottawa County, Oklahoma. July, 1991.

In summary, a list of "sins" (and at least one crime, a criminal offense) committed in timber harvest of this forest included: 1) timber theft, 2) improper felling, 3) wood wastage, 4) needless destruction of future logs (via "hatchet-happy" sawing, wanton felling, and careless skidding), 5) removal of riparian trees). The old stand deserved better management, but maybe this sad case of failed stewardship can in some small way educate future foresters and rangemen (by inverse example) in the ways of wise use management. The "gospel" (certainly not the Gospel of Efficiency) lesson here is that forests are renewable natural resources. Even after such "mutilation" this forest can recover, even if this author will not live to see (the nature of Forestry). Ironically, felling of the young adult sycamore might have extended their lives through coppicing. Otherwise those genetic trees would have "stayed dead" after senescence at completion of their normal life cycle.Like the meanders of Modoc Creek such are the twists and turns of life.

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

Ottawa County, Oklahoma. March.

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

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

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

Ottawa County, Oklahoma. March.

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

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

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

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

Wahoo, not yahoo- Eastern wahoo (Euonymus atropurpureus) growing in front of a box elder (larger tree at left) and white ash (smaller diameter trunk at right) on a mixed hardwood floodplain forest in the western Ozark Plateau. This shrub is widely distributed throughout the Eastern Deciduous Forest Region and westward with a biological range extending from Quebec and Ontario south to Florida across to Texas and west to Montana. Eastern wahoo prefers moist, rich habitats such as bottomlands.

Terrace of Modoc Creek, Ottawa County, Oklahoma. Late June..

Large, green leaves and little, green fruits- Leaves and immature fruit of eastern wahoo on a plant growing on a mixed hardwood bottomland forest. These organs were on the same plant that was introduced in the preceding photograph and caption. Eastern wahoo is a member of Celestraceae (bittersweet family). Other photographs of eastern wahoo along with American bittersweet (Celastrus scandens) were included in the chapter, Oak-Hickory Forest I of Range Types of North America.

Terrace of Modoc Creek, Ottawa County, Oklahoma. Late June..

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

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

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

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

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

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

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

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

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

Ottawa County, Oklahoma. July.

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

Ottawa County, Oklahoma. July.

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

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

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

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

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

Species in tribe Eupatorieae have only tubular or disk flowers.

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

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

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

Ottawa County, Oklahoma. April.

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

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

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

Ottawa County, Oklahoma. May.

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

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

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

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

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

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

Ottawa County, Oklahoma. June.

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

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

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

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

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

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

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

Ottawa County, Oklahoma. September; ripening fruit.

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

Ottawa County, Oklahoma. July.

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

Ottawa County, Oklahoma. September.

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

Ottawa County, Oklahoma. November.

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

Ottawa County, Oklahoma. Novermber.

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

Ottawa County, Oklahoma. Christmas Day.

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

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

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

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

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

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

Ottawa County, Oklahoma. June.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Ottawa County, Oklahoma. December.

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

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

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

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

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

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

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

Ottawa County, Oklahoma. August.

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

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

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

Permanent spring, Ottawa County, Oklahoma. April.

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

115. Bottomland forest of western hackberry, sugarberry (Celtis laevigata), and American elm (Ulmus americana) with an understory almost exclusively of broad-leaf wood oats (Uniola latifolia= Chasmanthium latifolium) with scattered solidary plants of the shade-tolerant composite, elephant's foot (Elepohantopus carolinianus). Beautiful plant community and an example of permanent deciduous forest range in contrast to the typical transitory feature of these dense hardwood forest types.

Ottawa County, Oklahoma. September, late estival aspect. No obvious FRES or Kuchler designations as this is a smaller unit. Neither was this relatively unique community described by Braun (1950) though it corresponds to general creek and river bottom vegetation in the Ozark Mountains. SAF Cover Type 93 (Sugarberry-American Elm-Green Ash). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

117.Winter in the Bosque bottoms- Hardwood bottomland forest dominated by netleaf hackberry, pecan, and American elm with dominant varying from location to location. In this view the dominant was netleaf hackberry with the four foremost trees all of that species. Most trees in background were pecan (largest trees) and American elm.

The dead herbage in foreground was broadleaf woodoats, the dominant warm-season herbaceous species. The tallest dead shoots were of frostweed (Eupatorium ). The dominant cool-season herbaceous species was Canada or nodding wildrye which was featured in the next photograph. There was also abundant Texas wintergrass (Stipa leucotricha). There were scattered individuals of naturalized, annual, Eurasian grasses including rescuegrass (Bromus unioloides= B. catharticus), Japanese brome or Japanese chess (B.japonicus), cheatgrass (B. tectorum), and wild oats (Avena fatua). There were also winter, annual forbs such as shepherd's purse (Capsella bursa-pastoris) that was common at time of this photograph.

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

118. Still winter in the Bosque bottoms- This photograph was taken within a few minutes of the preceding slide, but here the dominant, cool-season herbaceous species, Canada or nodding wildrye, was featured. Almost all of green bunchgrass in foreground was Canada wildrye though some of the green clumps were Texas wintergrass. The biggest tree (leaning right) was pecan (Carya illinoinenisis). The three smaller trees far to left of this pecan were American elm. Most of the trees behind the elms were netleaf hackberry. The dominant warm-season herbaceous species was broadleaf woodoats which was featured in the immediately preceding slide.

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

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

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

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

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

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

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

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

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

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

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

The following section dealt with management (proper and improper) of oak-hickory forests. An example of natural revegetation (reforestation) of a clearcut climax black oak-dominated forest was included above under the section, Black Oak Forest. Discussion of the even-aged (clearcutting) silvicultural method and secondary plant succession of a clearcut black oak forest was placed there because that tract of climax oak-hickory forest had been used as an example of the black oak forest cover type. Secondary succession on the clearcut was part of the forest range vegetation of that cover (dominance) type.

Presentations and discussions in this section were more generic. Examples and principles taught from them were, regardless of the dominance type, applicable across forest range cover types in the oak-hickory forest (Oak-Hickory Region)

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

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

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

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

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

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

131. Greed and amateurs in the Ozarks- A second-growth, black oak-dominated forest in the western Ozark Plateau was high-graded for furniture lumber and pallat material (and the felled trees were immature for those uses) by a group of half-assed, clueless buffons armed with barely running chainsaws. Any black oak having the diameter of a soup bowl and maybe one 2X4 in it) was cut while blackjack and post oaks of twice the size were left standing (although many of these were "barked" and otherwise scarred by fun-loving boys driving make-shift skidders through the woods). The result was near-devastation of the forest, but Mother Nature is forgiving (especially in a 40-42 inch precipitation zone).

Appearance of this degraded black-oak Ozark Highlands forest in the first third of the third growing season post-high grading was presented in this and the remainder of the following set of slides. These photographs documented the remarkable--and the remarkably rapid--recovery of this black oak forest and chronicled the resilence of this forest cover type and ecosystem in the Ozarks. This series of slides provided a photographic chronicle of secondary plant succession in the Interior Highlands of the oak-hickory forest region (Braun, 1950,ps. 164-173).

This photograph presented the general outer appearance of the high-graded, black-oak-dominated forest early in the third growing season following amateruish logging. The largest tree along the left margin was a second-growth blackjack oak. The foremost tree on the right was a second-growth post oak. Both of these trees--immature though they were for logs--were larger and older than most of the high-graded black oaks. God's mercy and resilence of this forest vegetation was readily apparent with amble regeneration of black oak along with a typical pattern of secondary plant succession that was reset to approximately mid-sere. The dominant shrub in this "photo-plot" was any of several species of blackberry (Rubus spp.) followed by smooth sumac (Rhus glabra var. glabra), winged sumac (R. copallina var latifolia), poison ivy (Toxicodendron radicans), and wild grapes (Vitis spp.). The dominant herbaceous species was broomsedge bluestem, but there was considerable cover of silky wildrye (Elymus villosus), Virginia wildrye (E. virginica), purpletop (Tridens flavus), beaked panicgrass (Panicum anceps), and occasional large clumps of big bluestem. There were some sedges (Carex spp., Cyperus spp.) present at pre-flowering stages.

132. Recovery in spite of greed and amateurs- Two overall composite shots of vegetation recovering via secondary plant succession on a high-graded, black oak forest in the western Ozark Plateau early in the third warm growing season following destructive logging by a bunch of amateur woodcutters. There was almost unbelieveable regeneration of black oak along with that of tree species including Texas or Ozark hickory (Carya texana), blackjack and post oaks, black cherry (Prunus serotina), sassafras (Sassaras albidum), and American elm (Ulmus americana). Shrubs (for which high-grading was an manmade windfall) in the recovering forest vegetation included blackberry species, wild grape, smooth sumac, winged or shining sumac, and Virginia creeper (Parthenocissus quinqueolia). Dominant herbaceous species varied locally and included broomsedge bluestem, silky wildrye, Virginia wildrye, purpletop, beaked panicgrass, naturalized tall fescue (Festuca arundinacea), and big bluestem. Major forbs in the range vegetation presented in this photograph included woodland or hairy sunflower (Helianthus hirsutus), and slender mountain mint (Pycnanthemum tenuifolium).

In the forest range vegetation shown in these two photographs black oak had more overall reproduction than any other tree species, except at isolated spots where local dominants varied from black cherry to post oak to eastern red cedar (Juniperus virginiana). Overall the second most regenerate tree was Texas, black, or Ozark hickory.

133. Third growing season- Revegetation of a high-graded, second-growth black oak-dominated forest in western Ozark Plateau in first third or so of third growing season following high-grading. Black and a few choice post oak logs were taken leaving blackjack oak, Texas or black hickory, elm, black cherry, and eastern red cedar. Fortunately, from an economic standpoint as well as recovery of forest vegetation most tree regeneration had been of black oak, the climax dominant, although locally black cherry, sassafras, post oak, and American elm occasionally exceeded black oak reproduction.

The local forest range vegetation presented here was a forest gap where a few "half-grown" black oaks had been felled for small logs out of a local stand that also included Texas or Ozark hickory and post oak. The gap had become a blackberry patch that was surrounded by uncut small trees. The perimeter of the gap also included, smooth sumac, winged sumac, summer grape (Vitis aestivalis), poison oak, and Virginia creeper. Herbaceous species of the gap included grasses such as big bluestem, purpletop, beaked panicgrass, broomsedge bluestem, and a few spots of southern crabgrass (Digitaria sanguinalis), each of which was locally dominant. In this photograph purpletop was the local dominant and it was featured in center foreground. Important forbs included hairy or woodland sunflower, slender mountain mint, giant ragweed (Ambrosia trifida) and tick clover (Desmodium spp.).

134. Plenty of new trees and grass- Recovery via secondary plant succession of a high-graded black oak-dominated Ozark forest was shown in this photo-quadrant which offered several lessons for the student of range vegetation. In about the first third of the third warm growing season following improper logging native grasses and forbs had made tremendous recovery along with regeneration of tree species. At this local scale big bluestem was the dominant climax grass with a nice clump growing in front of a nice black oak sapling both of which were already growing at time of high-grade logging. Both of these plants were released by removal of overshadowing crowns of black oak.

Also present prior to (as well as after) logging was the eastern red cedar (behind and to right of the black oak sapling and big bluestem) and two black cherry saplings (crooked trunks at left midground; somewhat in line but slightly to farther back that the easter red cedar). Obviously, a prescribed burn (or a wild fire) was needed to improve this forest range and facilitate secondary plant succession because a fire would kill the invasive and nonsprouting eastern red cedar as well as "stimulate" the big bluestem to produce sexual shoots (ie. increase seed production of the potnetial natural climax grass for this forest cover type on this range/forest site).

Most tree regeneration in thios photo-plot was Texas or black hickory.

135. Another patch in another gap- In a forest gap produced by removal of all trees from a local stand of black oaks a small-scale but diverse forest plant community developed consisting of blackberry, smooth and winged sumacs, Virginia and silky wildryes , broomsedge bluestem, purpletop, beaked panicgrass, hairy or woodland sunflower with black cherry as the main regenerating tree species. There was not much reproduction of black oak in this local gap.

136. Next generation of black oak- Outstanding regeneration of black oak, the climax dominant tree species, on a high-graded oak-hickory forest in the western Ozark Plateau about one-third into the third growing season following the most recent forest harvest. There was also considerable reproduction of black cherry. Winged and smooth sumac, blackberry, and wild grape were the main shrubs in this local stand of forest vegetation. Herbaceous species included grasses (broomsedge bluestem, purpletop, beaked panicgrass, big bluestem, naturalized tall fescue), sedges (both Carex and Cyperus species), and tick trefoil (Desmodium rigidum), the forb (a legume) with the most cover.

137. Released- Recovering range vegetation in a microhabitat by stump of a black oak roughly one-third through the third warm growing season following high-grading of a second-growth black oak-dominated Ozark forest. There was no sprouting from this stump, but there was tremendous sexual reproduction of black oak, the climax dominant tree species in all directions of the mmediate microenvironment of the stump. There was also a limited selection of shrub species including blackberry and poison ivy. The local dominant herbaceous species was woodland sunflower.

The conspicuous heart rot in this rather small stump was typical of many harvested black oaks. Cause of this rot was unknown. There was not a fire scar on this stump. From a wood production perspective it was good practice to rid the forest of such trees as the logs from such trees are "pert nar" worthless, being limited to wood chip products or cheap pallats. This is particularily the case when there is so much regeneration of the climax dominant and economically most valuable tree species. Unfortunately, much of the log removal by "hatchet-happy" skidder drivers left scars on trunks of many remaining trees which provided modes of entry for future causes of heart rot and economic losses in the next wood crop.

138. Baby boom- Prolific sexual reproduction of black oak, the climax dominant, in a high-graded oak-hickory in the western Ozark Plateau roughly on-third through the third warm growing season following half-assed forest harvest. The second most successful reproducing (also sexual reproduction) was black, Texas, or Ozark hickory. There was also regeneration of black cherry from seed. Other regenerating woody species included summer grape and Virginia creeper. The clumps of green grass at right side of stump were poverty oatgrass (Danthonia spicata). The stump was, of course, black oak. Tan-colored grass culms were broomsedge bluestem. The main forb was woodland or hairy sunflower. The trunk of the sapling (behind and slightly left of the stump) was flowering dogwood.

The following photographs and capitons provided an example of the effects of a spring wildfire of moderate intensity in the western boundary of the Oak-Hickory Forest Region. The forest was a second-growth, climax, black oak-pignut or bitternut hickory upland forest that was uneven-aged and approaching old-growth age and structure. This was a dry-mesic chert forest (Nelson, 2005, ps. ) in the Springfield Plateau section of the Ozark Uplands.The wildfire occurred in late March or early April during which period all hardwood species were still dormant. Eastern red cedar was the only tree species with leaves during this early spring period. Photographs were taken in late May and early June (approximately eight to ten weeks following the spring wildfire).

The results of this wildfire appeared to be those resulting from natural or anthropogenic ignition during a typical moist early spring (based on the author's experience with this forest range vegetation over a span of a half century). The wildfire was not of the intensity, season, etc. that would induce major disruption or drastic alteration of the oak-hickory forest ecosystem. It was not a wildfire of such severity as to cause major retrogression of the forest plant community nor, therefore, to induce secondary plant succession at lower stages along the forest sere. Even though the surface fire was a cool burn with fire intensity being relatively low for this forest range type, fuel (consisting mostly of shed leaves) had accumulated to considerable depth and extent due to prolongued absence of fire and, in addition, was quite dry such that the wildfire burned long enough to remove essentially all of the leaf/litter layer and exposed the extremely stoney surface of the soil on this upland chert forest. The surface forest fire usually left only incidental visible burn marks (eg. blackened bark), but it generated sufficient heat and/or persisted long enough to topkill plants of tree species ranging in age/size classes from one- or two-year old seedlings up to saplings of over three to four inches DBH.

White-tailed deer, squirrels, cottontail rabbits (Sylvilagus floridanus), and perhaps species of birds that might eat buds, leaves, bark as well as mast were the only grazing/browsing animals in this forest. No livestock had ranged in this forest for at least the last three-fourths of a century-- if ever (other than strays). The rough terrain of this forest tract precluded fencing. One side of this private property included bluffs above a creek. Bluffs were steep and treacherous, but there were local strips of land free enough of limestone edges that served as natural game trails through which livestock could stray. It was not economically (and probably not even physically) feasible to fence this steep upland forest. As such livestock had been excluded from this forest range which was then accessable only to wildlife species.

This second-growth forest had the species composition, structure, etc. of a climax forest with many old-age, hollow trees that formed a closed canopy. Even with heavy canopy cover the forest vegetation included a well-developed understorey with two to four layers (varying by local stands) that provided herbage and browse. This was permanent forest range in contrast to the transitory forest range of the black oak-dominated upland forest that was clearcut and described above.

139. Freshened by fire- Interior of a dry-mesic chert foret (Nelson, 2005, ps. 125-130) in the western Ozark Highlands approximately eight to ten weeks following a cool, surface fire (accidental wildfire). This was a black oak-pignut or bitternut hickory climax upland forest with well-developed layers of forest vegetation including two herbaceous layers and one (sometimes two) woody layer(s) that comprised a grazable/browsable understorey.

Other tree species included northern red oak that dominated a few local stands (these were conterminous with and ecotonal to a sugar maple-pignut hickory climax forest), post oak, chinkapin oak (Quercus muehlenbergii), sugar maple (also ecotonal to the adjoining sugar maple-pignut hickory forest), American or white elm (Ulmus ameicana), a few red or slippery elm (U. rubra), black cherry, and white ash (Fraxinus americana). Probably all age/size classes of these tree species from seedling to large sapling were present in the understorey or lower tree layer if forest gaps were included. Most regeneration of tree species was mockernut hickory and/or black oak, which in conjuction with dominance of canopy cover and total accumulation phytomass, indicatdd that these were co-dominants of this forest that was interpreted as climax vegetation.

Shrub species included the dominant flowering dogwood plus eastern redbud, shadbush or eastern serviceberry (Amelanchier arborea), and buckthorn (Rhamnus lanceolata var. glabrata) in the interior of the forest. Common woody vines included summer grape (Vitis aestivalis), Virginia creeper, and greenbriar, known also as bullbriar or catbriar, (Smilax bona-nox). Poison ivy/poison oak was present, but it was much less common than in other oak-hickory forests in this general area. At forest edge (just out of camera range and adjoining a limestone glade of big bluestem-dominated tallgrass prairie) major shrubs included smooth and skunkbush sumac.

The main herbaceous species in interior of this black oak-pignut hickory forest was pointed-leaf, sticky, or cluster-leaf ticktrefoil or, also, tick trefoil, or tick clover (Desmodium glutinosum), but there were local habitats where pokeweed, carrion-flower (Smilax herbacea), woodland sunflower (Helianthus hirsutus), white vervain (Verbena urticifolia), Indian physic or American ipecac (Gillenia stipulata), and small skullcap (Scutellaria parvula) were locally dominant or, at least, common. There were also other Desmodium species, but these were not blooming and could not be identified positively at species level. Bigbract or large-bracted tick trefoil (D. cuspidatum) was tentatively identified based on vegetative characteristics. Grasses were absent from this understorey except at forest edges--ecotonal to an adjoining glade--were big bluestem, Canada or nodding wildrye, silky wildrye, purpletop, and three-flower (ed) melic (Melica nitens) were abundant.

Tree species in the first photograph included post oak (foremost tree at left with fire-charred/blackened basal trunk), a minor component species, northern red oak (big tree at right background behind five or six saplings) associate species, black oak (large tree behind and to right of the fire-charred post oak; rightmost foreground tree behind and to immediate left of dead foreground sapling), co-dominant species; chinkapin oak, and , pignut or mocker nut hickory (largest tree; right of a waist-high rotten snag; center), co-dominant species; sugar maple (tree closest on the right to the big pignut hickory), indicator species and transitional to adjoining sugar maple-mockernut hickory forest; and Anerican elm (many of the saplings including the one in left midground to right of fire-charred post oak). Also in the first of these two slides was summer grape.

The second of these two photographs was a closer-in view of the center of forest introduced in the first slide. Two large black oak(left and right margins) and pignut or mockernut hickory (the same big tree at right of a rotten snag now barley visible) represented the two co-dominants of this dry-mesic chert forest. The sugar maple (tree to right of the previously noted pignut hickory), was an indicator of the somewhat mesic nature of this forest range site. Also in this second slide were many plants of both clusterleaf ticktrefoil and what was tentatively identified as bigbract ticktrefoil. Species in the herbaceous and low woody layer were presented and described in detail in subsequent slides.

Most regeneration presented in these two slides was pignut hickory, but black oak was well-represented. Successional status of American elm and black cherry on this forest site was unknown, but these are generally regarded and treated as seral species in climax oak-hickory forests.

140. Layers in a recently burnt Ozarks oak-hickory forest- This second-growth black oak-pignut or bitternut hickory upland Ozark Plateau forest of climax composition and structure was burned by wildfire (a cool, surface fire) about eight to ten weeks prior to time of photograph (late March or early April when all hardwood species were dormant and only eastern red cedar had leaves). This slide presented the wildfire-modified structure of this forest range vegetation.

Although the surface fire was a cool burn that only occasionally left visible burn marks (eg. blackened bark) it generated sufficient heat or persisted long enough to topkill plants of tree species ranging in age/size classes from one- or two-year old seedlings up to saplings of over three to four inches DBH. Fire intensity was generally low by standards of this forest type, but fuel (primarily several years accumulation of shed leaves) was dry enough that the wildfire burned of a duration sufficient to consume almost all of the leaf layer thereby exposing the rocky soil surface.

A high proportion of oak and hickory seedlings or small saplings ranging in size up to about one or more inches in basal diameter and roughly eight to less than ten feet maximum height were topkilled, but almost all of these promptly resprouted. By contrast, saplings of American elm and black cherry up to the approximate four inch DBH-size and some up to 18 or more feet in height were topkilled. Almost all of these resprouted from the basal trunk (= "stump") or roots. The glaring difference in tolerance (survivability) to fire exhibited by climax oak and hickory species versus American elm and black cherry was in the larger size-saplings (over three to four times greater height and stem diameter) of the latter that were topkilled. Sprouting appeared to be the same among hardwood tree species because apparently about 100% of topkilled individuals of all hardwood trees resprouted. The difference was in how much more effective even a cool surface fire was in total shoot kill of hardwood trees of species other than oak and hickory. Simply put, in the same forest and when standing side-by-side much larger trees of the nonclimax, associate species were topkilled (ie. only smaller shoots like seedlings and young saplings of the dominant oaks and hickories were killed whereas shoots up to pole-size of black cherry and elm died).

Stark differences between dominant oaks and hickory versus black cherry and elms in response to fire of the same intensity, frequency, at same season (same time, same dormant stage of phenology) was treated in separate photographs later.

The response of co-dominant black oak at large sapling size was shown by the individual in left foreground. Lower limbs appeared to have been killed as they had blackened bark and did not have leaves. Higher limbs and the crown were unscathed and in full-leaf about two months following a fire that occurred dormancy in early spring. Presence of a few persistent leaves from last year on some of these lower limbs was proof that they had been alive going into winter dormancy (unless they died the previous summer which seemed unlikely). The most obvious response of this black oak to the cool wildfire was basal sprouting. The fire had been hot or lasted long enough to induce "stump-suckering" (production of long shoots from the basal trunk), but it had not--up to time of photograph--damaged upper limbs and branches in the crown.

The snag of an American elm of pole size was also representative of this species in the black oak-mockernut hickory climax forest. American elm demonstrated high sexual reproduction with many seedlings and saplings. Even when these were topkilled by cool, surface fire they promptly resprouted. Resprouts from crowns of American elm saplings were conspicuous as, for example, new leaves of one such elm sapling directly in front of the pole-size snag (at first glance it appeared that the sprouts were from the snag, but this was only an illusion produced by camera angle from which the photograph was taken). American (and a few slippery) elms grew throughout the understory of this oak-hickory forest, but there were no individuals of these species larger than small poles. Evidentally, elms had (could) not compete effectively with larger trees of oaks, hickories, and sugar maples all of which had regenerated--and were continuing to regenerate--under their own shade. Such tolerance was indicative of climax species on this forest range type. Elms were major constitutent species in a lower woody layer dominated by shrubs like flowering dogwood, eastern redbud, lanceleaf buckthorn, and shadbush, but elms could not compete with climax oak and hickory dominants nor live long enough to enter the crown canopy. American and red or slippery elm were lasting components of the shrub layer.

Other plant species in the lower layers included Virginia creper, summer grape (both seedlings and fire-induced, basal sprouts), carrion-flower, and ticktrefoils.

Mature trees in right foreground were black oaks having DBHs of 24 to 25 inches. The large tree in left background was a pignut or mockernut hickory of 23 inches DBH.

141. Details of the layers and responses to wildfire- A canopy-to-ground "photoplot" in a recently burnt (eight to ten weeks prior to time of photograph) black oak-bittternut hickory climax forest forest revealed varying responses of different plant species to a cool, surface fire modifications in forest structure. The relatively cool fire that occurred during dormancy of hardwood species (late March or early April) topkilled and stimulated basal trunk ("stump")-sprouting of smaller saplings of trees along with sprouting from live shoots of some shrubs in the lower layers of range vegetation. Responses among angiosperms differed greatly, but this photographer could not find a single individual plant of any shrub species that had been topkilled. This included the woody vines of summer grape like those shown reaching into the canopy of the large black oak in left foreground. This was also the responses of lanceleaf buckthorn and shadbush. An individual of shadbush was immediately behind the large black oak (its trunk bent to the left from behind this black oak. This individual shrub was about as large as any the photographer ever found in the this vicinity. It suffered no detectable injury (did not even have a smoked basal bole) yet it had sprouted profusely from its root crown portion. Flowering dogwood apparently was affected the least of any woody species of similar size. Some smaller plants of flowering dogwood did sprout whereas some larger individuals of this species did not sprout.

This was a dry-mesic chert forest (Nelson, 2005, ps. ) in the Springfield Plateau of the Ozark Highlands that had been burned by wildfire eight to ten weeks previously.

142. New sprouts in old shrubs- A cool, surface wildfire burned (late March or early April) through the dry-mesic, upland, chert Ozark forest dominated by black oak and pignut or bitternut hickory about eight to ten weeks prior to the stage of growth and recovery of forest understorey shown in these two slides. These two slides presented details of the interior of this post-burn forest following the first fire to have taken place here in 40 to 50 years. The first slife featured a large tree in right foreground that was a black oak which two vines of summer grape continued to use as a trellis following the fire. These two shoots of summer grape had responded to the wildfire by sending up new shoots from their bases even though these shoots were still alive and leafed out (clear to the top of the black oak canopy).

In center foreground of the first photograph a nice shadbush or eastern serviceberry had also responded to the fire by basal sprouting even though its crown had also leafed-out and was apparently unaffected by the fire. The full-grown flowering dogweed behind the shadbush appeared to have been completely unaffected by the fire as it had not even sent up new shoots from the trunk base (ie. no stump sprouts). This is about as large as (the maximum size reached by) individuals of flowering dogwood and shadbush growing in upland forests in this portion of the Springfield Plateau, Ozark Mountains. Did the fire induce stump-sprouting in some individuals that will prove beneficial to these old-age (as indicated by large size) shrubs? Or was as-yet unseen damage done to these shrubs such as these two examples of two species? Proliferation of new shoots from the basal trunk cannot be viewed as neutral--even if sprouting and fire are natural phenomena.

The second photograph presented the sprouting shadbush shown in center foreground of the first slide accompanied by the woody vines of summer grape. The shoots of neither of these shrubs revealed any signs of fire-damage all of which having leafed out and growing elongating leaders. The only signs of fire on shoots of these plants (besides smoked bark) was their basal sprouting response. All basal sprouts of the shadbush had been browsed by white-tailed deer so that the terminal bud of each had been removed. None of the newly sprouted shoots of summer grape had been browsed.

143. Top to bottom in a fired upland forest- Forest opening (gap) in front of a small, local stand of black oak in a dry-mesic chert upland forest in the Ozark Highlands eight to ten weeks after a cool, surface fire burned through this forest range vegetation while all woody angiosperm species were still dormant (late March or early April). The large black oak had a DBH of 24-25 inches and a severely damaged crown so that this individual was long past its "prime" as a tree. Several of the larger black and northern red oaks in this climax (though second-growth) forest had moderate to severe crown damage meaning that these individuals were long past the rapid growth stage in their life cycle. Some larger trees were still-living (barely alive) snags with their crowns completely gone. If this forest was being managed for a wood crop over-ripe trees like these were "long over-due" for harvest. They probably had such heart-rot that their wood was unfit for lumber and useful only for fuelwood. Alternatively, they were extremely valuable for cavity nesting animals.

Much of the green (the vascular plant) surface layer in the gap and up to the trunks of mature black oaks was made up of seedlings and both small and larger saplings of pignut hickory and black oak (with slightly more hickory). All of these had resprouted since the wildfire (otherwise there would have been six to eight foot saplings in front of the big black oak).Other lower-growing species included pointed-leaf or cluster-leaf ticktrefoil and, probably, bigbract or large-bracted ticktrefoil (though this latter species was in pre-bloom stage and could not be identified with 100% accuracy), carrion-flower, Virginia creeper, summer grape, and seedlings of American elm.

144. Regeneration of the climax- In a local stand of black oak in a black oak-bitternut hickory forest there was ample reproduction of both of these climax co-dominants following a wildfire (in late March or early April; a cool, surface fire). These two photographs showed survival of both 1) year-old seedlings and 2 of older, topkilled saplings which had respounded by resprouting. In the foreground of both photographs there was a single trunk of a mature black oak. In left midground of the first slide a mature northern red oak represented this more mesic of the two oak species. Both slides showed the abundant sexual reproduction of oak and mockernut hickory as well as that of American elm along with much cover of pointed-leaf or cluster-leaf ticktrefoil, white vervain, summer grape, and Virginia creeper. Some small trees of sugar maple were growing in the more densely shaded background (which was too dark to photograph and reveal anything other than shadows).

A high proportion of the saplings in foreground of both slides was American elm. All of these saplings that had been alive at time of the wildfire had been topkilled and resprouted by time of photographing; however, many of the elm saplings had already died (presumedly via competition with adult oaks and hickories as Dutch elm disease had not reached this tract of forest).

145. Fire in the woods: past and present- Eight to ten weeks after a wildfire (cool, surface fire even though it was an accidental burn) burned (late March or early April) through this climax black oak-pignut hickory forest the spring vegetation on this forest range showed evidence of the recent fire as well as previous fires. The large fire scar at base of the center (largest) tree was unequivacol evidence of past fire(s). This tree was a pignut or bitternut hickory with a DBH of 23 inches. The tree to the left and the smaller trunk to the right of the fire-scarred hickory were black oaks.The shoot of the smaller black oak (the one on the right) showed no affect of the wildfire other than prolific and rapid growth of new basal long shoots. The leaning (to the left) tree behind the hickory and black oaks was a northern red oak. The dead sapling behind and to right of the bitternut hickory was also a northern red oak which had been taken out by extreme competition from adult trees or disease or...

There was much regeneration of the oaks and bittternut hickory along with that of American elm and summer grape. The woody shoot of an adult summer grape was growing between the hickory and left-front black oak. Much of the lower layer of prominent green leaves consisted of pointed-leaf or cluster-leaf ticktrefoil, the most abundant herbaceous species in this tract of forest.

Follow-up timber tale: the year after this wild fire and the set of slides for this section lightening killed the large pignut hickory (basal fire scar and 23 inch DBH) shown here. It was not just top-killed, but total death including of basal stump and root system. An account of vegetation dynamics in the forest gap created by opening in the canopy with loss of this tree was provided below following treatment of this wild fire.

146. Burnt but still coming (or blessing of fire in an oak-hickory forest)- Synopsis shot of the understorey as well as adult trees in a black oak-bitternut hickory dominated upland forest in the western Ozark Highlands about eight to ten weeks following a cool, surface burn (wildfire) that moved through this forest vegetation while all hardwood species were still in dormancy (late March or early April). Eastern red cedar was the only woody species that had green leaves and was capable of photosynthesis at time of the wildfire.

Adult or, even, mature trees in midground were northern red oak (two dark trunks at left) and chinkapin oak (gray or pale-colored trunk at right). The partial trunk of a sapling at right corner foreground was also a northern red oak. Sapling in left corner foreground was an American elm which was not topkilled by the wildfire. The topdead sapling that had rapid growth of basal sprouts (long shoots) was a slippery or red elm. There were also large seedlings or small saplings of black cherry and sassafras that had been topkilled and quickly sprouted back. Shrubs included the dominant flowering dogwood plus eastern redbud and lanceleaf buckthorn.

Herbaceous species included the dominant pointed-leaf or cluster-leaf ticktrefoil and what was tentatively identified as bigbracted ticktrefoil. This local habitat was deeper in the forest and more densely shaded so that Desmodium species were the only herb of any consequence.

147 Adult fire sprouts- Shoots sprouted from a burnt out stump of black oak decades ago and grew into four mid-size tree trunks. These black oak shoots were approaching adult size in these two photographs. In the first slide there was a fifth tree trunk of black oak (to left and rear of the four stump sprouts) that is a separate tree, a genetically distinct (genotype) tree. The four trunks of black oak that sprouted from the stump of the previous black oak that was killed by the fire are the same tree (same genotype). Each of these four tree trunks is a clone, module, or ramet (synonyms) of the original genetic individual just as the fifth black oak trunk is that of a separate, genetically unique tree. In sum, these five tree trunks (boles) are of two trees one (the older/larger) of which had four boles and one that had a single bole like a "normal" tree. The second slide showed basal portions of the stump shoots, including the hole by basees of the four boles which was left when remains of the burned off stump rotted away, proving their common point of origin by asexual reproduction which in this case was suckering, sprouting, or coppicing.

The four boles of one tree are the progeny of fire-induced coppicing. Coppicing in black oak was introduced, described, and references cited in regard thereto above in the section, Black Oak Forests. Examples of two-year-old stump suckers (long shoots that arose from the root crown and/or cut sutmp) were presented in that portion of the Oak-Hickory Forests chapter.

These two photographs were taken about eight to ten weeks after a cool, surface fire (accidental cause; late March or early April) burned off most of the understorey of a second-growth black oak-mockernut hickory upland forest in the western Ozark Highlands (Springfield Plateau section of the Ozark Plateau or Mountains). Recently resprouted yearling and two-year old seedlings and small saplings included those of mockernut hickory (the co-dominant with most regeneration overall), summer grape, Virginian creeper, pointed-leaf or cluster-leaf ticktrefoil, white vervain, small skullcap, and even a young plant of some species of wild lettuce (Lactuca sp.).

Ottawa County, Oklahoma. June (vernal aspect).

148. Sweet root sprouts- Base of a sugar maple in a dry-mesic chert upland forest (Nelson, 2005, ps. 125-130) dominated by black oak and bitternut hickory eight to ten weeks following a cool, surface wildfire (back in late March or early April). Sugar maple was one of several other hardwood tree species including chinkapin, post, and northern red oak along with black cherry, American and slippery elm, and white ash. Major shrubs included flowering dogwood, eastern redbud, shadbush or eastern serviceberry, and lanceleaf buckthorn. The main herbaceous plants were ticktrefoil (Desmodium spp.), the major species of which was . All of these species have the capacity to sprout from stumps, roots, rhizomes, root crowns, etc. In fact the only species in this forest lacking the sprouting response was eastern red cedar. The wildfire topkilled many individual plants that ranged in size from only inches-high seedlings of oaks and hickories to 25 foot tall black cherry with four inch-diameter trunks. The great majority of these topkilled plants--except for those of eastern red cedar--resprouted almost "immediately".

The small, new shoots of sugar maple shown in this slide were root suckers or root sprouts in cotrast to stem or stump sprouts of most of the other resprouting species (and individuals thereof) in this forest. Root suckering is apparently relatively rare in sugar maple, but it obviously took place in this instance. These young, living (this year's) shoots of root origin from the parent plant were themselves fire-induced resprouts from previous season's shoots thar arose from roots in absence of fire. In other words, the root shoots from this sugar maple had arisen as clones or modules in previous growing seasons. These ramets or clones of root origin were topkilled by the surface forest fire and current growth (the only living root shoots) were resprouts of fire-killed root suckers (fire-caused resprouts of normal root sprouts). Both normal (the original) root shoots and regrown root shoots following death of original root suckers were products of asexual reproduction.

Original shoots coming off of roots of the genetically unique mother tree (clones or ramets of the genet) was an example of natural cloning the same as was coppicing in black oak described above. In the case of sugar maple root shoots this is apparently one adaptation (survival feature) of the Very Tolerant sugar maple. Rather than relying strictly on food storage in a maple seed and gambling on risky, high-mortality rate sexual regeneration (reproduction) sugar maple can use photosynthate directly to reproduce asexually with fewer risk and lower mortality rate among its progeny. In this way sugar maple remained a constitutent species in the black oak-mockernut hickory climax forest.

Ottawa County, Oklahoma. June (vernal aspect).

149 Rejuvenated the base (after it killed the top): an example of a tree species with low adult fire-tolerance- Understorey of a black oak-bitternut hickory upland forest in the western Ozark Plateau about eight to ten weeks following a cool, surface wildfire in late March or early April. This was a dry-mesic, chert forest (Nelson, 2005, ps. ) that developed adjacent to a sugar maple-mockernut hickory climax north slope forest. A black cherry of pole size (about 25 feet tall with a trunk diameter over four inches) was completely topkilled, but sprouted long shoots from the root crown of basal shoot (center midground). Oaks and hickories less than half this size were unscathed, but black cherry and elm species showed much less tolerance to fire than did climax hardwood species. Seedlings and small saplings of both American and slippery elms were abundant in this portion of the forest understorey.

Shrubs visible in this "photo-quadrant" included flowering dogwood (small. resprouted seedling in right foreground), Virginia creeper, and summer grape. About the only grass present in this forest (other than at forest edge adjacent to a prairie glade) was hairy woodland brome (Bromus purgans= B. pubescens) seen here at early growth stages.

For a comparison of the higher fire-tolerance of black oak, co-dominant of this climax-composition upland forest, in contrast to high fire-susceptability of black cherry, local associate species, the next slide in this series was offered. This black cherry and the black oaks grew within about 20 to 30 steps of each other.

150. Springing from and staying on newly fired ground: plus an example of a tree species with relatively high adult fire-tolerance- Interior of a western Ozark Highlands upland forest dominated by black oak and bitternut hickory eight to ten weeks following a surface wildfire in late March or early April. Many (in fact, most) individual plants of several species were topkilled even though this was a relatively cool fire.

This photograph featured several black oaks of large sapling to small pole size that were largely unaffected by the wildfire. Most such young trees lost lower limbs, but blackoak with trunk diameters greater than approximately three inches (perhaps slightly smaller) were neither topkilled nor induced to produce long shoots (= resprouts= suckers) from basal trunks or root crowns. This lack of sprouting response was evident in the two small pole-size black oaks in center of this slide. Neither of these (or other nearby) black oaks sent up new basal shoots even though the fire at this local environment burned long and/or hot enough to remove all leaf and litter material from the ground surface and burn much of the bark on trunk bases.

This relatively high fire-tolerance in black oak contrasted sharply with that of black cherry. By way of comparison/contrast readers were encouraged to "roll up" to the immediately preceding slide in which a black cherry much larger than (at least half again as large as) these black oaks was completely topkilled and promptly sprouted from the shoot base and/or root crown. All of these sapling or pole-size trees were growing within 20 to 30 steps of each other.

There were many resprouted plants were of numerous species the most common or abundant of which included seedlings and small saplings of both co-dominants, summer grape, Virginia creeper, ticktrefoil (Desmodium spp) most of which was pointed-leaf or cluster-leaf ticktrefoil, and carrior flower. Viewers should take note that the young, previous seasons' shoots of summer grape that had twined up into the foremost (foreground) black oak were killed by the surface wildfire and had resprouted from their root crown(s). As was the case with trees and other shrubs, the larger the shoot the greater the probability that the shoot would less or little affected by fire (other things being equal, of course).

151. What fire wrought in the woods- Two views of an extremely instructive "photoplot" of the understorey in the interior of dry-mesic, chert, upland forest (Nelson, 2005) eight to ten weeks after a cool, surface wildfire (in late March or early April) burnt off much of the forest understorey. This climax-composition forest range was dominated by black oak-bitternut hickory (locally accompanied by northern red and chinkapin oaks, American and slippery or red elm, sugar maple,and white ash) with two species-rich shrub layers and one or more herbaceous layers. The dominant shrub was flowering dogwood with local associate shrub species varying from lanceleaf buckthorn, shadbush, eastern dogwood, Virginia creeper to summer grape.

The most obvious shrub effects due to forest fire in this forest involved the dual impacts or interaction of defoliation, first by the surface fire followed by white-tailed deer browsing. Deer must have either "camped out"on this black oak-mockernut hickory forest range or else included it as part of their daily feeding route. There were few sprouts on any individual plant of most of the shrub species that had not been browsed by deer. The one exception was the dominant shrub overall: flowering dogwood. There were several (a fairly high percentage) of dogwood plants on which some or all fire-induced sprouts were unbrowsed by deer. By contrast there were almost no fire sprouts on any plant of lanceleaf buckthorn that had not been browsed, and browsed heavily. Deer-browsing on shadbush and summer grape was intermediate between that of buckthorn and dogwood.

This author/photographer could not find a single sprout of black, northern red, chinkapin, or post oak or of bitternut hickory that had been browsed. The same feeding situation obtained for sassafras. Sprouts of the dominant oaks and hickory might as well as been steel posts or creosoted power poles as regards deer feeding. Sassafras would have been the equivalent of asbestos. The most highly preferred tree species for browse was elm, most of which were American or white elm. All plants of Ulmus spp. that sprouted following fire were browsed; usually there were few, if any, shoots of any elm that were unbrowsed. Black cherry was also highly prefered based on proportion of shoots browsed and relative degree of use on these shoots. White ash was the least abundant tree species in this forest being mostly a transitional and only occasional species from an adjoining bluff forest community of sugar maple, mockernut hickory, and white ash.White ash received very little browse use by deer, but relative rarity of this species might have been a factor in this regard (availability being one component of browsing selectivity).

Browsing selectivity on this site provided a textbook case. At left midground in the first of these two slides and featured exclusively in the second of these slides (front and center) there was a small pole-size sassafras that was topkilled and had resprouted. Less than halfway through the first spring-summer growing season this sassafras had new sprouts over five feet tall. These were untouched by deer (apparently the flavor of sassafras shoots to deer is not on par with that of the roots to human taste). There were numerous basal shoots (fire-caused resprouts) of lanceleaf buckthorn (eg. behind and to right of the sassafras) and American elm (eg. right foreground). In contrast to the untouched sassafras, buckthorn and American elm received moderate to heavy browsing by white-teiled deer. Note the almost extreme degree of use on the lanceleaf buckthorn. This feeding pattern undoubtedly reflected deer preferences and relative palatability of browse species to deer, given the same availability of sassafras, lanceleaf buckthorn, and Americn elm (ie. they stood side-by-side and/or back-to-back).

Note: another textbook example of browsing selectivity was presented later (below) in this series of slides on the climax black oak-mockernut hickory forest between hickory and lanceleaf buckthorn.

Most of the understorey in this "photo-plot" was pointed-leaf or cluster-leaf ticktrefoil. Contrary to common sense and human experience this nodulated (papiolionaceous), herbaceous legume received relatively scant utilization by white-tailed deer. There were a few plants of eastern redbud on this local site and these also received fairly limited utilization (given the heavy browsing of deer on accompanying species).

Organization Note: Examples and details of pointed-leaf or cluster-leaf ticktrefoil (known also as sticky tick-clover, sticky tick-trefoil) were presented above in the section entitled Sugar Maple Forests. Forest plants covered under Sugar Maple Forests included many of the same species found in the range vegetation of this black oak-pignut hickory forest. These two examples of those two forest types were contiguous with each other and even included a transition zone between them.

152. More released understorey and resprouting hardwoods- Another "photo-plot" of the second-growth (but climax-composition) upland, chert, Ozark Plateau forest range dominated by black oak and bitternut hickory eight to ten weeks after a cool, surface wildfire (late March or early April) spread through the understorey layers of this vegetation. Pointed-leaf or cluster-leaf ticktrefoil was the dominant herbaceous species at this location. Also locally (in the forest vegetation shown in this slide) low-growing or trailing shrubs, including Virginian creeper and summer grape, made up a lower woody layer while a taller or mid-height woody layer was comprised of lanceleaf buckthorn (three small, single-bole shrubs in center fore-to midground), eastern redbud, and flowering dogwood. The foremost (and smaller buckthorn) was topkilled and resprouted vigerously, but in an exceptional example these new shoot had received no browsing by white-tailed deer.

Large trees in background were mockernut or bitternut hickory (right margin) and black oak (left side).

153. Fire affects: some sprouted; some did not- Understorey in interior of a black oak-bitternut hickory upland chert forest in the Ozark Highlands about eight to ten weeks following a cool, surface burn (wildfire in late March or early April). This was a second-growth forest range with the species composition and structure of this climax forest type (for this forest site). Other important tree species included northern red, chinkapin, and post oaks along with sugar maple, black cherry, American and slippery or red elm, and white ash.

Major shrub species were present in this view featuring the understorey. Flowering dogwood was the dominant shrub species overall although locally shadbush or eastern serviceberry, eastern redbud, and lanceleaf buckthorn were dominant. Summer grape and Virginia creeper were dominant and almost universally present woody vines which following the wildfire resprouted to form a ground surface or lower woody layer.On this local site, resprouted seedlings and small saplings of American elm were associated with sprouts of these species of woody vines (lianas). (Fire-induced sprouts of elm seedlings and saplings were abundant throughout the field of view in this photograph.)

In the forest range vegetation presented here a seedling of white ash, a relatively rare species in this forest tract, had resprouted (lower left foreground). An example of a resprouted, young flowering dogwood (probably a two- or three-year-old seedling) was in right foreground.

The large sapling or small pole in center midground was a black oak that was unfazed by the wildfire and, hence, did not growh stump suckers (fire-induced long shoots).

154. Pyrogenic variety- "Photoquadrant" of the lower layer of the understorey of the second-growth, upland black oak-bitternut hickory forest with species composition and structure of climax vegetation at about eight to ten weeks following a cool, surface burn (wildfire in late March or early April). Main purpose of this photograph was to present the various herbaceous and woody species comprising the lower zone (vertically) that was typical of the outer zone (vs. interior) of this forest range vegetation. Herbceous species included carrion-flower, ticktrefoil (mostly pointed-leaf or cluster-leaf ticktrefoil with some large-bracted or big-bract ticktrefoil), and a few plants of hairy woodland brome. Other herbaceous range plants that were in this lower layer as well as a higher herbaceous layer included white vervain. small skullcap, pokeweed, and woodland sunflower.

Resprouts of shrubs and woody vines were also part of this lower layer (as well as the vertical layer or zone immediately above). Some of those species were greenbriar (known also as catbriar or bullbriar), Virginia creeper, summer grape, skunkbush sumac as well as seedlings and resprouts of saplings of the oaks, especially the co-dominant black oak, and mockernut hickory, the other co-dominant tree species.

155. Two two-time loosers (but still in the woods)- Two saplings of American elm that were topkilled by a fairly cool, surface wildfire (late March or early April), promptly resprouted, and then were browsed by white-tailed deer. These small trees were photographed approximately eight to ten weeks post-wildfire. In that interim period these two examples of fire-induced shoot/stump sprouts (long shoots) of American elm had received moderate to moderately heavy degrees of use by deer. Note that in both of these plants (examples) there few, if any, shoots that had some browsing .

These plants were in the upper shrub layer of range vegetation in a second-growth, upland, chert forest that had the species composition and structure of climax forest of this cover type and forest site. Black oak and pignut or bitternut hickory were co-dominants with other major tree species being northern red and chinkapin oaks, sugar maple, black cherry, American or white elm, red or slippery elm, and white ash. Major shrubs included flowering dogwood, overall major species, eastern redbud, shadbush, lanceleaf buckthorn, Virginia creeper, and summer grape. Sassafras was locally important,but sporadic in its distribution in this forest tract. At forest edges smooth and skunbush sumacs were important. The most important herbaceous species was pointed-leaf or cluster-leaf ticktrefoil. Grasses were non-existent except for very infrequent individuals of hairy woodland brome.

American elm and red elm were two of the more palatable and important browse plants on this oak-hickory forest range.

Ottawa County, Oklahoma. Early June (vernal aspect); less than midway through the first growing season following topkilling by wildfire.

156. Was loosing really winning or, alternatively, is there more waiting to be lost? - A third small tree of American elm served as another example of defoliation (and comparative degree of use) of resprouts of American elm by white-tailed deer on a climax, black oak-bitternut hickory upland forest in the Spring Plateau section of the Ozark Plateau about eight to ten weeks following a cool, surface burn (wildfire in late March or early April). The wildfire topkilled this young American elm (stem diameter was less than one inch) which quickly responded by prolific production of long shoots from root crown or base of bole and, which in turn, were just about as quickly browsed (moderate to heavy degree of use) by deer.

This example served to teach another lesson in management of ranges and ask an interesting question. The observant viewer will have noted that there was one (and only one) large, long shoot that received no defoliation by deer. The developing terminal bud with its growth-regulating apical meristem on this one shoot was still intact. If this one "Lone Ranger" long shoot continued to escape defoliation it would (presumedly) quickly replace the fire-killed young bole. The replacement shoot could then continue to grow and develop toward maturity; die in competition with climax oaks, hickories, and sugar maple; succumb to Dutch elm disease; or live to a ripe old age and return to the earth from which it grew. Any of these (or other) possible outcomes would depend on continued escape of this intact shoot from subsequent deer defoliation. If deer eventually browse this shoot (remove the terminal bud and apical meristem) then the fire-killed and subsequently browsed elm will likely remain a hedged bush rather than develop into a tree.

Whether a continually browsed (hedged) shrub or, alternatively, a tree capable of producing wood or habitat for birds, squirrels, and furbearers is the management goal will determine whether either final browsing outcome is desirable or undesirable. If a continual game browse range is desired then the hedged shrub would be preferred. If a fully developed, sexually mature elm tree is desired in order to propagate other elms and with the full-grown tree ultimately serving as a source of wood then escape of the remaining shoot would be preferred outcome.

If (for sake of this argument) the "sole, surviving heir" capable of developing into a replacement tree does survive it is possible (on the surface it is logical or rational to assume) that browsing by deer was a net benefit to this individual (unique genetotype of) American elm. This is plausible because browsing could have equated to "pruning by Mother Nature" in a way that facilitated more rapid growth, development, and sexual maturity of the remaing shoot. The remaining single shoot would have less stress from competition by the population--most likely an overpopulation-- of heterophyllous or long shoots (=suckers= sprouts). This in turn would increase the likelihood of survival by the tree (the original genetic individual) to maturity and successful sexual reproduction. It was conceivable that deer browsing amounted to beneficial or, perhaps, even essential pruning.

On the other hand, it was also quite likely that deer would "make another call" of this recovering American elm and remove the last intact shoot thereby reducing the former single-stem sapling to a continually cropped, circular hedge (ie. a persistent shrub with axesual reproduction the only alternative to regeneration). Time will tell. Another excuse to visit the woods.

Ottawa County, Oklahoma. Early June (vernal aspect); less than midway through the first growing season following topkilling by wildfire.

157. Leaves still at the top, but now at the bottom too- Two plants of lanceleaf buckthorn that featured selectivity by two defoliators in tandem. In a climax black oak-bitternut hickory upland forest in the Springfield Plateau section of the Ozark Plateau (Highlands) a cool surface burn (wildfire in late March or early April) induced basal stem or trunk ("stump")-sprouting of numerous species many of which were subsequently browsed (sometimes heavily) by free-ranging white-tailed deer.

In the first example (slide) a nice, single-trunk lanceleaf buckthorn exhibited no signs of the wildfire other than profuse sprouting from the trunk base or, perhaps, the root crown. In the second exampe (slide) a much smaller, two-trunked lanceleaf buckthorn had prominently charred and heat-cracked bark on its boles beside numerous long shoots (=suckers=sprouts) that were induced by the fire. This smaller buckthorn was, however, also not topkilled. It, too, showed no signs of fire-damage other than burnt bark and basal sprouting.

The second of these photographs furnished another lesson in browsing selectivity. A resprouted seedling of mockernut or bitternut hickory with zero defoliation (no deer-browsing) was growing beside the browsed lanceleaf buckthorn (hickory was in front and to the left of the buckthorn). It was mentioned previously that the author could not find any seedlings, fire-induced resprouts, or saplings of oak and hickory species, climax tree species, whereas buckthorn, apparently a climax shrub species, had been consistently consumed by deer (frequently at fairly high degrees of use). In the concept of selective browsing the hardwood tree species had low or poor palatability for deer and, for their part, deer demonstrated their preference for buckthorn and their lack of preference for mockernut hickory. Students should pay attention to the fact that, by definition, palatability is a characteristic or feature of plants while preference is a feature of animals. Availability of a plant species, a dual function of presence of the plant species at time and place that animals could utilize it, was obviously not a factor in the total combination of the phenomenon called selection. Selection or selectivity= palatability + preference + availability.

Note: another textbook example of browsing selectivity by deer was presented earlier (above) in this series of slides on climax black oak-mockernut hickory forest between sassafras and both lanceleaf buckthorn and American elm.

Ottawa County, Oklahoma. Early June (vernal aspect); less than midway through the first growing season following topkilling by wildfire.

158. When selectivity by one defoliator is enough- Two general views of the climax, upland black oak-bitternut hickory forest in the Springfield Plateau section of the Ozark Mountains described under several photographs presented above. These two slides were taken approximately eight to ten weeks following a cool, surface fire (anthropogenic wildfire; late March or early April). These photographs presented the general structure, including layering, and overall species composition of the forest range. Spermatophytes (seed-bearing plants) in this second-growth, climax-composition and structure forest were limited to angiosperms (fruit-bearing or flowering plants) except for one species of gymnosperm (spermatophytes bearing naked seeds or seeds not encased in fruits): eastern red cedar (juniper). Eastern red cedar was the only woody species with leaves present at the time of the wildfire. This conifer was also unique on this forest range site in being the only species consistently incapable of producing resprouts (long or heterophyllous shoots) from trunk or roots (root crown). Thus topkilling of eastern red cedar was synonymous with killing eastern red cedar.

Eastern red cedar grew beneath the nearly closed canopy of oak species (black, northern red, chinkapin, and post oaks), mockernut hickory, sugar maple, black cherry, and an occasional white ash and elm (American and red) and in (as a constitutent of ) a higher shrub layer made up of flowering dogwood, shadbush or eastrn serviceberry, lanceleaf buckthorn, and eastern redbub (and with summer grape and Virginia creeper) growing from ground to canopy layer. In this oak-hickory forest that had not burned in several decades (probably not in the last three or four) eastern red cedar had begun to invade.

The wildfire that rejuvenated numerous plants of various species and merely set back younger plants of other species was death to eastern red cedar. The wildfire was not hot enough to even consume juniper needles, but it was hot enough to give "complete" or "total" kill (vs. only topkill) of this nonsprouting juniper even if heat was just enough to scorch leaves. (And the dead coniferous invaders in this climax hardwood forest were "purtier than a Christmas tree" to foresters and rangemen.)

More discission on invasion and control of eastern red cedar in oak-hickory forests was provided in the next series of slides.

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

159. Four years after- The lower woody layer and herbaceous layer(s) of a climax dry-mesic upland forest in the western Ozark Plateau four years following a lightening-set surface fire that burned theough this vegetation (in mid-spring). Plants present included seedlings and saplings (of various ages) of climax tree species including black oak, pignut hickory, white ash, American elm, and post oak as well as shoots of the shrubs redbud, flowering dogwood, shadbush, and summer grape. Many of the shoots of these shrub species were rootcrown or stump sprouts that resulted from either topkill or damage-induced growth due to the wild fire.

Herbaceous species, all of which were unaffected by the fire, included hairy wood brome (Bromus purgans= B. pubescens), pointed-leaf, sticky, or cluster-leaf tick-trefoil (Desmodium glutitinosum), carrion-flower (Smilax herbacea), black-eyed susan (Rudbeckia hirta), and heartleaf skullcap (Scutellia ovata).

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

160. Long look for some long plants- Plants of bur-reed caric sedge (Carex sparganoides) in herbaceous layer of understorey in upland climax black oak-bitternut hickory) forest in Ozark (Springfield) Plateau. Plants of bur-reed caric sedge grew together in local populations of sparse cover. This cover was comprised of comparatively large, weak-stemmed individuals having a resultant prostrate growth-form. Such plants are easily overlooked by the casual observer. Rangemen with disciplined concentration and keen eyes should have no trouble in detecting these plants. The first of these two slides presented a typical rangeman's view from standing position. The second photograph presented two (2) typical plants of bur-reed sdege from a bent-over or mid-height distance.

Bur-reed caric sedge was the major grasslike plant in the rock-strewn soil of this comparatively xeric forest range community. The dominant graminoid of the herbaceous layer was hairy wood brome. The dominant herbaceous species was pointed-leaf, sticky, or cluster-leaf tick trefoil. Other associated forbs included black-eyed susan (Rudbeckia hirta) and heartleaf skullcap (Scutellia ovata) which were featured below.

In an unusual growing season (bitter-cold, wet winter followed by wet, warm spring and then unusually hot, dry summer with exceptional drought) bur-reed caric sedge grew larger and produced more fruit than was typical for this forest range site.Such circumstances provided fine photographic fodder and an educational opportunity for students of forest range vegetation.

This was a dry-mesic, chert forest by Missouri standards of forest community classification (Nelson, 2005, ps. 125-130).

Ottawa County, Oklahoma. Mid-July; fruit-ripe phenological stage.

161. Long caric shoots in a dry forest- Closer looks at bur-reed caric sedge in understorey of a black oak-pignut hickory upland forest. In contrast to an upright and a bent-over or leaning view (two immediately preceding slides) the current two perspectives (camera distances) presented rangemen with squatting-distance views of bur-reed sedge on a chert outcrop, dry-mesic Ozark Highlands forest.

The broad leaves of neighboring plants were American elm, which was locally an associate species to the co-dominant black oak and pignut or bitternut hickory whose leaves made almost all of the litter layer beneath the sedge.

Ottawa County, Oklahoma. Mid-July; fruit-ripe phenological stage.

162. Shoots and spikes under oaks and hickories- Basal shoots and a spike of ripening achenes (first slide) and shoots with leaves along with a spike of maturing achenes (second slide) of bur-reed cric sedge in the herbaceous layer of a black oak-pignut hickory upland chert forest in western Ozark (Springfield) Plateau. Some senescing sedge leaves were prominent in both of these photographs.

Ottawa County, Oklahoma. Mid-July; fruit-ripe phenological stage.

163. Burred tops- Four different views of spikes of bur-reed caric sedge that developed in the herbaceous layer of a black oak-pignut hickory climax forest. Many authorities describe/define the inflorescence (whole flower/fruit cluster) type of Carex species as a spike with individual groups being spikelets and each unit of the spikelet being a floret. C. sparaganioides spikelets are androgynous meaning that staminate florets are above lower pistillate florets, that is female florets subtend the male floral units (Steyermark, 1963, ps. 311, 335; Kaul et al., 2006, p. 379-380). Distinct spacing (well-delinated spaces) among spikelets is a readily seen feature in this Carex species (Steyermark, 1963, ps. 333, 335). The rich brown to golden color of florets (spikelets) is another charactristic of this species (Fernald, 1950, p. 308).

Interestingly, the same figure in both Fernald (1950, p. 308) and Steyermark (1963, p. 333) did not show the prominent often--but not always present--bract at base or beneath lower spiklets that are present as seen here, in photographs "on the web", and illustrated in Flora of North America (Flora of North America Editorial Coommittee, 2003, p. 296).

These specimens were in a dry-mesic chert upland forest (Nelson, 2005, ps. 125-130), and bur-reed sedge was the major grasslike plant species in this Ozrk Highland forest range.

Ottawa County, Oklahoma. Mid-July; fruit-ripe phenological stage.

164. Two forbs on the floor- Black-eyed susan (Rudbeckia hirta), left, and heartleaf skullcap (Scutellia ovata), right, growing of the floor of a climax black oak-pignut hickory four years following a lightening-ignited surface fire four years ago. Heartleaf skullcap was a dominant forb and black-eyed was an associate forb in this forest both before and after the fire (fire aspects had no observable impact on either forb species.

Ottawa County, Oklahoma. Late July (estival aspect).

165. Heart of the forest- Heart-leaf skullcap (Scutellaria ovata), a dominant forb, in the herbaceous understorey of a climax black oak-pignut hickory five years after a lightening-ignited surface fire burnt across this forest range. Although there is much variation in morphological features of this species (Steyermark, 1963, ps. 1271-1272), heart-leaf skullcap is one of the largest-growing Scutelleria species in the oak-hickory forests of the Ozark Highlands. This dominant forb grew in association with such herbaceous species as bur-reed caric sedge, hairy wood brome (Bromus purgans= B. pubescens), sticky (= pointed-leaf, or cluster-leaf) tick-trefoil or tick clover (Desmodium glutinosum), and carrion-flower (Smilax herbacea).

Plants and plant parts shown in this series of photographs were growing and flowering on extremely rocky, shallow soil in the second consecutive summer of Extreme to Extraordinary Drought (Palmer Scale). Heart-leaf skullcap is reasonably drought-tolerant. (most native species are).

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

166. Leave your heart in the Ozarks- Two views of upper shoots (of two plants) featuring the inflorescences and leaf of heart-leaf skullcap on a western Ozark (Springfield) Plateau climax upland forest, a dry-mesic, chert forest (Nelson, 2005, ps. 125-130) dominated by black oak and pignut hickory.

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

167. Flowers after your heart- Inflorescences of heart-leaf skullcap--presented so as to feature individual flowers--on plants growing in the haerbaceous understorey of a climax black oak-pignut hickory upland chert forest five years following a lightening-ingited surface fire.

Readers could handily see from these photogrphs that heart-leaf skullcap makes a nearly ideal species for folks desiring native plants in their yards and gardens. As increasing water shortages increase acceptability--not to mention desirability--of natural landscaping there will be on-going, trial-and-error experiments to find native plants best adapted for such uses. Heart-leaf skullcap is a a "real looker" in this regard. It is a comparatively large forest forb with pretty, eye-catching leaves to accompany its beautiful flowers. What is more, based on observations made on the climax upland oak-hickory forest being described here, white-tail deer do not graze this species. This woods-watcher did not find any evidence of defoliation of heart-leaf skullcap even though a small herd of white-tail deer had been attracted to and grazed some of the other herbaceous species on this previously burnt-over forest range. Finally, it was noted above that this species is quite drought-tolerant.

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

168. Nimble on the floor- Single plant of nimblewill (Muhlenbergia schreberi) on the floor of a dry-mesic, chert outcrop upland black oak-pignut hickory dominated forest. in the Ozark (Springfield) Plateau. This forest had burnt off (spring-season surface fire) four years prior to this growth of nimblewill. There were a few scattered plants of nimblewill in the herbaceous zone of this forest, but the dominant--though sparsely scattered--grass was hairy woodland brome.

Nimblewill is a native perennial that is probably better descriped as a bunchgrass; however nimbleweill grows adventituous roots and forms modules (clonal units) along nodes of the decumbent shoots such that it has a semi-sodforming habit (Tyrl et al., 2008, p. 136-137). In this manner nimblewill frequently grows as "dense colonies" (Steyermark, 1963, p. 160). Nimblewill has a unique habit or growth form in that early season shoots are characteristically lower-growing with wider leaf blades whereas by late late-season shoots take on a more up-right form with ascending to arching culms from which contracted panicles emerge (Hitchcock and Chase, 1950, p. 397).

Nimblewill is a common forest grass in forests of the Ozark Highland Region, and undoubtedly is a member of the climax plant community, but it is generally an increaser or even invader (when present at higher cover and density) particularily on local denuded (disturbed) areas. In facat, nimblewill is so adapted to and readily populates disturbed patches that it is a weed in lawns where domesticated turfgrasses like the bluegrasses (Poa species)or bentgrasses (Agrostis species) are preferred (Fernald, 1950, p. 169). In the author's experience and observations in the Ozark Plateau nimblewill is readily eaten by all species of livestock.

Ottawa County, Oklahoma. Late July; advanced stage of plant growth, pre-bloom.

The next set of slides and captions were of a second-growth, chert, upland oak-hickory forest dominated by (in approximate general order): black oak, post oak, blackjack oak, northern red oak (rarely), and Texas or Ozark hickory. Oldest/largest trees in this mixed oak-hickory forest were of mid-age (in contrast to the mostly mature trees of the black oak-bitternut or pignut hickory described immediately above). Both of these tracts of oak-hickory forest had burnt off at about the same time (late March or early April) in separate wildfires that were of about the same overall fire behavior and impact on forest vegetation. Both wildfires were generally cool surface fires that did not leave fire scars on trees that had mature bark, but that did topkill sprouting species and kill "outright" nonsprouting eastern red cedar.

The black oak-post oak-blackjack oak-Texas or Ozark hickory forest had not been grazed/browsed by livestock for about three decades. Prior to that time beef cattle had accesses to this second-growth forest range which, in earlier stages of forest recovery following logging had substantial quantities of herbage, browse, and mast. White-tailed deer ranged freely through this forest.

This tract of second-growth forest was best described when using existing cover types of the Society of American Foresters (Eyre, 1980) as a combination ("hybrid") of Post Oak-Blackjack Oak (SAF 40) and Black Oak (SAF 110). Black oak was generally the most abundant tree that commanded the most cover of the forest canopy, but post and blackjack oaks were almost as common and locally dominant (as was black oak on other local habitats). Both SAF 40 and SAF 110 have been regarded as climax or at least subclimax in the Ozark Highlands (Eyre, 1980, ps. 39, 43) where, incidentially, these types (and combinations thereof) might be edaphic, topographic, or pyric climaxes (forest types).

169. Fire at the edge- Appearance of the edge between a second-growth mixed oak (black, post, and blackjack oaks)-Ozark or Texas hickory upland forest and a remnant of degraded tallgrass prairie about eight to ten weeks following a cool, surface burn (wildfire; late March or early April). Black cherry is a limited component species of this mixed oak-hickory forest (SAF 40 X SAF 110) which has traditionally been interpreted as climax vegetation throughout much of the Ozark Plateau Region. Some plants of eastern red cedar (juniper) had invaded this forest (a major topic below).

Flowering dogwood and eastern redbud were co-dominants of the upper shrub layer. Various species of woody vines (lianas) extended from the ground layer to crown canopy: Virginia creeper, poison oak.ivy, greenbriar (mostly Smilax bona-nox), and both summer and fox grapes. Herbaceous species in the interior of this oak-hickory forest were very limited (mostly to ticktrefoils and caric sedges) so that an herbaceous layer was not present except for local colonies of Mayapple. Herbaceous species at edges of this forest and an adjoining grassland of former tallgrass parairie that was presently invaded by introduced Eurasian species, pioneer or colonizing annuals, and afew perennial broadleaf weeds. Forest-grassland edge plant communities were presented and discussed in the immediately following two photographs and caption.

There were two obvious effects of the recent wildfire on range vegetation of this oak hickory forest. The most general effect was reduction in cover of the multi-layered understorey. The visual effect of this reduction in understorey cover was due to topkilling of many (most) smaller, individual plants of shrubs and seedlings/saplings of the tree species. Essentially as many of these plants as were topkilled quickly sprouted back from basal meristem of shoots ("stumps") or root crows (ie. even those plants which had all of their aboveground, perennial biomass killed initiated new growth from perenniating buds at or below ground level).

The second and, to the eye of this photographer-rangeman, most striking or immediately visible outcome of this fire was the completely scorched (and thus "completely dead") eastern red cedar (juniper) that had invaded this ecotonal forest range site to which juniper is not native. Eastern red cedar is native to this general area (Springfield Plateau section of the Ozark Plateau or Ozark Highlands), but this conifer is not native to, not a natural component species of, oak-hickory forests, tallgrass prairie, of savannahs (ecotones) where the first two of these general range types merge or overlap.

In the Ozark Highlands and parts of the Central Lowlands (eg. Cherokee Prairie) eastern red cedar naturally exists (is a natural component of the native climax vegetation) only in habitats (eg. range and forest sites) naturally secure from fire. Such habitats include environments with shallow, stoney (low water-holding capacity) soils where plant cover is sparse and, therefore, fuel is inadequate for fires (fire regimes) that would kill the extremely fire-sensitive juniper. Such harsh, water-limited environments have fires of such limited intensity, frequency etc. that some (generally only a few) eastern red cedar survive to ages that enable them to grow large enouth to escape future infrequent, low-intensity fires.

Invasion of eastern red cedar in to climax or subclimax oak-hickory forests and tallgrass prairies is a sure sign of mismanagement of forests and grasslands (sloppy husbandry of the range, poor stewardship of the land). So much so in fact, that absence (or presence) of eastern red cedar is a "litmus test" for proper (or improper) management of forests and ranges. Death of eastern red cedar on this forest range was celebrated--Hip! Hip!, Hip! Hurray!--in more detail below.

These two photographs displayed the physiogonomy, general structure, and conspicuous species (eg. dominants) at edge of this Ozarks oak-hickory forest and tallgrass prairie in part of the Prairie Peninsula first described by Transeau (1935). The large tree in this two-slide sequence was blackjack oak. The sapling to left of this blackjack was black oak. The topkilled, smaller sapling to left of the black oak (larger) sapling and most of of the other saplings of various ages/sizes under and surrounding the black oak sapling and the adult blackjackwere saplings of Carya texana known variously as black, Ozarks, or Texas hickory. This is one of the smaller Carya species in this region. It is also one of the best adapted tree species for shallow, infertile soils, frequent drought conditions, tolerance to defoliation, and rapid regeneration on disturbed habitats. The fire-scorched eastern red cedar was obvious.

Low-stature (at time of photographs) plant species in foreground of the edge included tree seedlings, shrubs, grasses, grasslike plants, and forbs. This layer of the vegetational edge and these plants were shown and described in detail in the immediately following two-slide set. The most conspicuous species in these slides were the two forbs that were the major herbaceous species: 1) giant ragweed, the overall most abundant (apparent relative cover, biomass, density) species and 2) woodland sunflower, second most common (second greatest apparent relative cover, biomass, and density). The most common grass in edge vegetation was broomsedge bluestem, but big bluestem was well-represented (see slide set immediately below). Naturalized Eurasian grasses in this ecotonal range vegetation included tall fescue (Festuca arundinacea), orchardgrass (Dactylis glomerata), perennial ryegrass (Lolium perenne) and common bermudagrass (Cynodon dactylon). Two naturalized Eurasian legumes included white clover (Trifolium repens) and red clover (T. pratense). These naturalized introduced grasses and legumes grew at outer-most edge of the transition zone. They did not grow in either the extreme "dripline edge" or the interior of the mixed oak-Ozark hickory forest.

170. Details of the fired edge- Two "photo-plots" featuring species composition and general biodiversity of the outer zone of the edge of range vegetation at ecotone (transition) between mixed oak (black, post, blackjack oaks)-Ozark or black hickory, upland, chert forest and degraded tallgrass prairie about eight to ten weeks after a cool, surface fire burned through dormant range vegetation (man-caused wildfire; late March or early April; dry fuel). This was that part of the edge vegetation adjoining the grassland so that herbaceous species made up a plurality of this zone of the ecotonal range plant community.

Forbs dominated this local range vegetation. Major species (in approximate overall order of species makeup based on estimated cover and/or biomass): giant ragweed, woodland sunflower, black-eyed susan (Rudbeckia hirta), lopseed (Phryma leptostachya), Carolina elephant's foot, annual ragweed, and pointed-leaf or cluster-leaf ticktrefoil. Lopseed and elephant's foot were the most abundant forbs in the range vegetation that was recorded in the first of these two slides. The second of these slides displayed species biodiversity of this edge range vegetation. Species visible in this second photograph included saplings of tree (eg. the resprouted black cherry in upper right-hand corner), shrubs (summer grape, Virginia creeper) and forbs (giant ragweed, annual ragweed, woodland sunflower, lopseed, elephant's foot, ) in addition to the conspicuous clumb of big bluestem (left center , and

171. Cleaned up underneath (for now)- Interior of a second-growth, chert upland, black oak-post oak-blackjack oak-Ozark hickory forest approximately eight to ten weeks after a cool, surface wildfire (late March or early April; hardwood species dormant). There were two conspicuous effects of this wildfire. Most glaring and of most immediate importance was the easy and effective removal of eastern red cedar (an invader on this range site) from this Ozark mixed oak-hickory forest. Secondly--and of most moment from the management standpoint of browse production on this forest range--was proliferation (asexual reproduction) of long or heterophyllous shoots (=suckers= sprouts) from roots or trunk bases ("stumps") of topkilled seedlings and saplings of oaks, black or Ozark hickory, black cherry, and both American and red elms. The latter three species are not climax dominants of this forest type nor are they as tolerant of fire as the climax oak and hickory species. The fire selectively reduced these seral tree species more (to a greater relative degree) than climax plant species.

Even a cool, early spring fire when the dominant trees and shrubs (all were angiosperm species) were dormant resulted in a burn of sufficient heat and duration that seedlings and small saplings of hardwood species were topkilled. Killed "completely" (root as well as shoot) were most trees (up to large sapling size) of eastern red cedar, a non-sprouting conifer, that had invaded this forest range. Eastern red cedar was the only woody species in this forest that was "leafed out" at time of the early spring wildfire. It was also the only woody species that does not resprout (produce long shoots, suckers, sprouts). Killed crown= killed tree.

This wildfire--a convenience fire that burned out of control--"cleaned up the woods" (the initial purpose in parlance of the hillbilly who set it) and in the process rejuvenated browse species, reduced cover of seral species or, same thing, facilitated plant succession, and eliminated eastern red cedar, an invader (invasive species) that was a weed tree.

Smaller saplings in center midground and larger trees in right background were black oak; larger trees in left foreground were post oak. The largest and most common forb in the understorey in this scene was pokeweed. Most common plant species in lower layer (just above soil surface) was resprouting Virginia creeper. There was also extensive cover of summer grape and buckbrush or coralberry. Black cherry was the tree species with most seedlings in this lower layer of forest vegetation shown from this view (photo-frame).

172. Timber Stand Improvement- View inside a second-growth mixed oak (black, post, and blackjack oaks)-Ozark hickory on a chert upland in the Springfield Plateau section of the Ozark Highlands approximately eight to ten weeks following a cool, surface wildfire (late March or early April; eastern red cedar was the only woody species that was not dormant). Heat and/or length of burning time were adequate to topkill hardwood seedlings and small saplings and kill both root and shoot of most plants of the non-sprouting coniferous cedar (juniper). Since eastern red cedar does not sprout from roots, bases or sides of trunks, or branches (does not develop new shoots from existing woody tissue) death of existing live wood equates to death of the plant. If fire (or anything else) kills the shoot it kills the plant. Simply put: scorch the crown; kill the cedar. Over and done with.

The early spring, relatively low-intensity wildfire effectively ridded this oak-hickory forest of juniper. This included the largest plants in the forest. Saplings of eastern red cedar larger than ten feet in height and three inches in bole diameter were killed. It was all over except for the celebrating.

Fire--in this case, wildfire-- resulted in improvement in this forest stand. Fire eliminated a native tree species that was not a component species of the potential natural (climax) vegetation of this forest type. Fire, a natural factor in this forest range ecosystem (and an abiotic influence under which it evolved) removed an invasive plant species. Fire was God's and the Indians' form of brush control. In fact, fire of sufficient frequency serves as brush prevention, the preferred level of noxious plant management for forest or range. Fire is a tool in range and forest improvement.

For generations, Timber Stand Improvement was such a common and widely used term in the foresters lexicon that it was customarily shown as TSI. The Society of American Foresters (1Ford-Robertson, 1971) defined TSI as a "a loose term comprising all intermediate cuttings made to improve the composition, constitution, condition and increment of a timber stand". At time of this writing the Society of American Foresters (Helms, 1998) had decided that TSI was obsolete and used "stand improvement- an intermediate treatment made to improve the composition, strucrture, condition, health, and growth of even- or uneven-aged stands" with intermediate reatment defined as "any treatment or tending designed to enhance growth, quality, vigor, and compostion of the stand after establishment or regeneration and prior to final harvest".

Aside from the opinion of the current author-photographer that forest health is a substandard term, the concept of TSI or stand improvement (or whatever term will be used when the next SAF forestry glossary comes out) was appropriate for the effects of wildfire in this forest. Increased production of browse and herbage and elimination of eastern red cedar from this forest range were improvements by whatever term or definition was used.

The two large trees in right foreground and the one to their left in center midground were black oak, overall dominant tree species of this tract of mixed oak-Ozark or Texas hickory forest. Much of the lowest layer of vascular plants was seedlings of tree species, including black cherry, and American and red elms, though most sexual reproduction was of hickory and oaks. There was somewhat more regeneration of hickory than of the oak species. Sprouts (fire-induced shoots) of Virginia creeper, summer grape, and buckbrush or coralberry were common throughout the understorey. The grass highlighted against a dark background in the lower right corner was thin or thinseed paspalum (Paspalum ciliatifolium var. muhlenbergii=P. muehlenbergii= P. setaceum var. muehlenbergii, as now generally accepted to be a variety of P. setaceum the latter was most current at time this description was written). This grass grew only sporadically at forest edge. It could not survive in the forest interior.

173. More Timber Stand Improvement (a dead cedar is a good cedar)- Species composition and structure of a chert, upland mixed oak-Ozark or black hickory forest in the western Ozark Highlands as apparent eight to ten weeks after burning by an early spring wildfire (cool, surface fire; late March or early April). This second-growth forest was dominated (other than at local scale= localized stands) by black oak, but post and blackjack oaks and black or Texas hickory were general associates while black cherry and, less frequently, American and slippery or red elm were local associates. Eastern red cedar (juniper) was an invader and a weed tree.

Weed tree- "1. any tree of a species having little or no economic value on the site in quetion 2. a tree of little or no economic value, more particularly when competing with one or more desirable timber trees at any stage of development and therefore due for elimination from the crop" (Helms, 1998).

It should be emphasized that for many weed species there is an ecological basis in addition to the primary economic criterion for correct designation of weed or pest. This is clearly the case for eastern red cedar as exemplified on the oak-hickory forest described here. Some weeds (a la eastern red cedar) are alien species (or, at least, species that are not indigenous to the site they have invaded), that can migrate and become established due to disturbance (human-induced or otherwise). Disturbance certainly includes lack of fire in native plant communities in which fire was a natural abiotic factor under which natural (climax) vegetation developed and ecosystems evolved.

On most oak-hickory forests eastern red cedar is a species that is not a natural constitutent of this climax vegetation. Rather this conifer is an invader, an invasive species, that can only persist on such forest range in continued absence of fire. Eastern red cedar is a weed tree--both ecologically and economically--that becomes established under exclusion of fire. Conseversely, fire prevents invasion of this woody weed and/or controls it once it becomes established (if the trees are not too large to crown-scorch).

Even though the spring burn whose impacts were described here was a cool, surface fire (and all hardwood species were still in dormancy) there was sufficient heat generated or long enough duration of burning to kill almost all of the invading eastern red cedar. The surface fire did not consume the winter-dessicated needles of this juniper species nor did the fire blacken much of the bark on trunk bases, but a mere scorching of the crowns was sufficient to kill even the largest individual junipers (biggest trees were saplings about ten feet tall with stems of about three-inch diameter).

Numerous seedlings and saplings of oaks and hickory were topkilled as were those of the less fire-tolerant black cherry and elms which lost shoots of saplings larger than those of the climax dominants. For all practical purposes, all topkilled saplings and seedlings of hardwood trees promptly sprouted back (produced long shoots from roots or trunk bases) so that while the fire killed from one to several seasons of aboveground perennial growth it did not kill these plants. No flowering dogwood and eastern redbud were topkilled except for a few seedlings and very small saplings. Likewise, none of the larger plants of these two shrub species produced basal shoots (=sprouts= suckers).

Although this wildfire burned within a week of (and less than a mile from) the fire in the black oak-mockernut hickory upland forest described in the preceding slide series, the fire in the mixed oak-black hickory forest described here was a substantially cooler fire. It appeared from investigations subsequent to this fire that it burned downhill (ie. a backing fire) and on a day during which temperatures was considerably lower. Consequently the fire in the mixed oak-Ozark hickory forest had less impact on hardwoods (trees and shrugs). There was little if any difference in impacts on eastern red cedar between the two forests (forest types). It does not take a hot or persistent fire to kill an eastern red cedar. Fortunately, this conifer is a species that like the house cat (Felis silvestris catus=Felis domesticus=Felis catus) is easily killed. Apparently animal taxonomists are in as much disagreement over proper scientific names as agrostologists are of grasses, but the relevant thing is that house cats and eastern red cedar die easily if properly attacked. A good and faithful steward will keep both of these species out of the oak-hickory forest, and kill any he finds that are already there. Death to pests!

In sum, the wildfire selectively eliminated non-sprouting eastern red cedar, reduced the undercanopy of the shrub and lower tree layers, increased browse production, and facilitated plant succession in the forest.

The largest tree trunk (left foreground) and the one to left and mostly obscured by the largest trunk were black oaks. The tree behind these black oaks and left of the left cedar was northern red oak. The two trees in center foreground and midground to the left of the right scorched cedar and the pole-sized tree behind and to right of left scorched cedar were post oaks. The lowest layer of vascular plants (just above ground level) was comprised of seedling and resprouts of tree species, Virginia creeper, buckbrush or coralberry, and summer grape.

174. Fire is a selective feeder- "Photo-transect" showing selective defoliation by an early spring wildfire (cool surface fire; late March or early April) in the lower (herbaceous) layer of mixed oak-Texas or black hickory upland forest in the western Ozark Highlands. Shown here was the burn scene at local habitat scale about eight to ten weeks post-fire. In center foreground a caric sedge (Carex sp?) that was either dormant had but few leaves present at time of fire was thriving. All around the sedge were new shoots and leaves of Virginia creeper, and behind the sedge was a dead seedling of eastern red cedar. This young tree had been killed by the cool, surface fire which did not generate enough heat nor send flames high enough to incinerate this cedar (juniper). Instead the crown of this baby tree was merely scorched, but that was enough to kill a small plant of this non-sprouting coniferous species.

In this same forest and from the same fire much larger cedars (saplings with up to three-inch diameter boles) were also killed with no more visible damage than scorched crowns. Examples of this effect were presented in immediately preceding slides and captions. It was conceivable that some smaller, short-stature cedars might have escaped the fire due to passage of flames and/or heat above them. Such was not the case. The flaming angel of death did not pass over smaller junipers. Instead, eastern red cedar is so sensitive to fire affects that a broad range of age/size classes of cedar in this forest were susceptible to fire-kill.

Ottawa County, Oklahoma. Early June (vernal aspect).

175. More selective feeding by fire- Local scale (microsite, microhabitat, microenvironment) assemblage of plants in a mixed oak-Ozark hickory, upland, chert forest in western Ozark Plateau. Viewers should note that the cool, surface fire was not hot enough to harm moss growing on trunk of post oak, and at time of fire the moss was quite dry (in fact it was the type of moss carried by frontiersmen in their tenderboxes for tender when starting fires with their flints). Such a cool fire was sufficient to readily kill non-sprouting eastern red cedar. The point was made again: eastern red cedar is readily killed by even low-intensity fire.

The mossy tree trunk was post oak. Most of the leaves at or just above ground level were Virginia creeper (five-leaflet leaves) and buckbrush or coralberry or, frequently, coral berry (opposite leaves aligned along stems).

Management implication: eastern red cedar, an invasive weed tree on some range types, is so easily killed by fire that the safest (least-likely to escape) surface fire imaginable is adequate to eliminate most seedlings and small to mid-size saplings of this non-sprouting conifer from oak-hickory forest (and from tallgrass prairie and oak-hickory-tallgrass savanna). Therefore, fire these ranges and fire them frequently so as to improve chances of killing eastern red cedar (cedar trees will be younger and smaller under more frequent burning).

Ottawa County, Oklahoma. Early June (vernal aspect).

The next short series of slides-captions was of a natural (God-sent) disturbance in the second-growth, upland, black oak-pignut hickory forest in the western Ozark Highlands that was treated in considerable detail above in regard to impacts of a spring wild fire . This was a dry-mesic chert foret (Nelson, 2005, ps. 125-130) with the species composition and structure of a forest nearing the old-growth state of development (ie. for all practical purposes it was a climax forest upland oak-hickory forest). Our previous chapter in this "true tale from the timber" ending with rapid on-going recovery from a surface fire of moderate intensity.

The following spring was one with several severe thunderstorms and uprootings of many large trees due to a combination of high winds and saturated soils. Numerous strikes of sky-to-ground lightening accompanied all these storms. One such lightening bolt struck a large (23 inch DBH) pignut or bitternut hickory that bore a large basal fire scar and wounds from lost limbs attesting to previous stresses to which it was subjectd and survived. This stalwart veteran of many a disturbance did not survive the lightening strike. The impact of lightening (whatever specific phenomena of the electric discharge causes precise injuries or damage resulting in death) terminated the long and colorful life of this patriarch pignut hickory. This was no mere top-kill, spectacular though that would have been. Instead, the previously resilient hickory experienced the ultimate outcome. Death was dramatic, traumatic, and total: dead rootcrown or basal stump and root system as well as death of the crown. That tree was no more.

Removal of this living tree and the opening created in the forest canopy along with loss of competition, shelter, propagule source, etc. (ie. all combined influences and changes that such impact causes) was a readymade textbook example of a forest gap and its accompanying patch dynamics. As soon as the sound of the lightening strike and instaneous thunder faded away one could hear the echoing words of Henry Chandler Cowles and Frederic Edward Clements: "Vegetation is dynamic".

Some of these dynamic changes in the forest vegetation were presented below for the education and entertainment of viewers not so privileged to stand at the snag and see unfold the subsequent recovery--dynamic development--of vegetation on this Ozark Plateau forest range. Photographs were taken the second year after death of tree and creation of forest gap (ie. these slides were of recovering forest vegetation at beginning of year two of patch dynamics).

176. Blasted- Gap created in an upland black oak-pignut hickory forest by death of large (23 inch DBH) adult pignut or bitternut hickory due to lightening strike. Lightening knocked off one of two major limbs at base of tree crown and also resulted in total or complete death of the hickory. This was not just top-kill, but also death of rootcrown and roots. Saturated soil was perhaps a contributing factor to such thorough killing of tree through better grounding. Another factor was undoubtedly age given the physiological phenomenon whereby older trees sprout much less readily and vigerously than do younger trees. Amazingly, no other plants were affected by the lightening strike. One large single trunk of a flowereing dogwood was broken off by crash of the falling limb from the pignut hickory; however, this dogwood had alredy been top-killed by the moderate-intensity surface fire and resprouted from its rootcrown two springs earlier so that there was literally zero direct impact from the lightening strike on any plant except the hickory.

This was appearance of the forest gap in the early part of the second summer after death of tree and creation of the gap with initiation of vegetation dynamics. The initial or nearly immediate effect was release of all plants that had previously been shaded (for varying periods of time) by the canopy of the hickory, the holder of the high ground as it were. Tree species were the major beneficiaries of release in the gap. Tree release was as seedlings and saplings that were almost all of climax species including pignut hickory, the major released species, black oak, American elm, white ash, and northern red oak. The one seral tree species released was sassafras (Sassafras albidum). There were no grasses or grasslike plants in the forest gap because the soil surface was almost completely covered by seedlings and saplings of the various tree, shrub, or even forb species. The major forb species was pointed-leaf, sticky, or cluster-leaf ticktrefoil (also written as tick trefoil and known as tick clover). Other--though much less common--forbs were carrion-flower (Smilax herbacea) and pokeberry. Important shrubs in the gap itself included fox grape, flowering dogwood, Virginia creeper, and blackberry.

Pioneer (weedy) plant species were not present in thsi forest gap. The author could not find a single plant of an annual or biennial species. The closest species to early colonizers were pokeberry and sassafras. For all practical purposes secondary plant succession was not taking place. Patch dynamics consisted of establishment, growth, and maturation of young plants of climax species.

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

177. Kaboom!- A large (23 inch DBH) pignut hickory struck down in its prime by lightening that completely killed it (not just top-kill but total death). This hickory had a remarkably straight bole with relatively taper which forked into large limbs from which were the entire crown originated. One of these limbs was knocked off by the lightening strike. Saturated soil resulting recent heavy rains served as thorough grounding for the tree. No other plant was directly affected by the lightening strike. The crashing limb of the hickory broke off a large, dead trunk of flowering dogwood that had been topkilled three years earlier by a wild surface fire of moderate-intensity. The coppice or stump sprouts of this dogwood were unphased by the fallen limb.

The opening in the forest canopy that was created by death of the pignut hickory was a "windfall" for all other plants. These consisted primarily of seedlings and saplings of the climax tree species, especially pignut hickory of which there were more sexual offspring than any other tree. Other climax trees with both saplings and seedlings included white ash and black oak. There were seedlings only of northern red oak and American or white elm of the climax group and also seedlings of the seral (for this forest range site) black cherry (Prunus serotina). There was one already existing sapling of American elm that had been severely scourched by the wild fire and that had partly leafed out, but it was not going to make it (or amount to anything if it did).

Abundant shrub species in this still-young forest gap included fox grape, Virginia creeper, flowering dogwood, and blackberry. The most abundant forb was sticky, pointed-leaf, or cluster-leaf tick clover or tick trefoil, the herbaceous dominant for this forest range.site Other forbs included carrion-flower (an herbaceous green-briar), wingstem crownbeard (Verbesina helianthoides), and pokeberry or pokeweed. Grasses were not present in the thick "jungle" of this forest gap even though hairy mountain brome, Canada wildrye, and woodland bluegrass were scattered variously throughout this upland forest.

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

178. Shattered life and new life- Lightening-shredded trunk (23 inch DBH) of a pignut hickory was at center of a gap in a black oak-pignut hickory climax upland chert forest in the western Springfield Plateau. Complete (crown to root system) kill of this once-mighty hickory set into motion patch dynamics in this forest gap. Change in vegetation was nothing more than release of already present climax species of trees, shrubs, and forbs. The lightening strike affected directly only the one, mature, hickory tree. Neighboring adult black oaks, flowering dogwood, seedlings and saplings of other trees, etc. were unaffected by the lightening bolt, but clearly benefitted from the inpouring of light after the crown of the pignut hickory was reduced to a skeleton of bare branches. In fact, one of two limbs that formed the crown was knocked off by lightening strike.

There were saplings and seedlings of pignut hickory, the species that produced the majority of sexual progeny, along with those of black oak, and white ash. Reproduction of American elm and northern northern red oak, the other climax tree species in the gap, was limited to seedling classes. There was one partly leafed-out sapling of American elm that had been mostly top-killed by a surface wild fire of moderate intensity three springs ago, but it was not going to do anything. There were also some seedlings of wild cherry which is not a climax species for this forest type or site. The most abundant shrubs were fox or winter grape and Virginia creeper. Flowering dogwood was another important shrub many of which consisted of stumm sprouts following the wild fire. The most common and productive (greatest cover and density) forb was sticky, pointed-leaf, or cluster-leaf ticktrefoil (tick clover) which was a climax understorey dominant for this forest and forest site. Other forbs included carrion-flower (an herbaceous green-briar) and wingstem crownbeard, this latter visible in right midground. Grasses, such as hairy mountain brome and Canada wildrye which were present in various local microhabitats throughout this forest, were not in the gap.

The large sapling snag with falling bark and lichen growth to immediate right of the pignut hickory was northern red oak that had been dead for several years prior to lightening strike apparently having lost the war in competition for light (and/or whatever else) with the larger, older hickory. Just no justice in these woods: the nice northern red oak died too soon to grow to greatness. Its younger siblings will have to be the ones that become part of the climax. forest.

Plant succession was not occurring on this forest gap. Development of vegetation consisted of growth, maturation, and reproduction of individual plants of the climax species. Pioneeer and advanced seral plant species (other than a few sassafras plants) were not present.

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

179. "One generation passeth away,and another generation cometh..." (Ecclesiastes 1:4)- Several seedlings and one sapling of pignut or bitternut hickory at base of the trunk of an old-growth pignut hickory killed the previous year by lightening. Pignut hckory was the plant species with the greatest reproduction (both asexual and sexual) in the forest gap created by death of this single, adult hickory tree. Other climax tree species that had seedlings in the gap included black oak, northern red oak, american elm, and white ash. Also present (as in this photograph at far left margin) were seedlings of black cherry. The dead sapling at upper right corner was an American or white elm that was killed three years previously by a wild surface fire of moderate intensity. Leaves and woody vine of fox grape were conspicuous at lower right. Leaves of Virginia creeper were visible at left.

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

180. Same patch, next year-View of the lightening-created gap in a second-growth, upland, black oak-pignut hickory forest that was introduced above as it appeared in the next (and a drought) year. This was the third year, growing season, and part of summer after lightening blasted a comparatively large (23 inch DBH) pignut hickory and created a microsite of new forest land on which the forest community had to redevelopment. Development of new forest vegetation was primarily one of released existing plants and relatively little or quite limited appearance of typical pioneer or colonizing species. The most abundant of these early seral species was sassafras. By contrast there were several existing plant species that made rapid growth due to reduced competition, greater light, etc. (ie. that were released) with death of the large hickory and creation of an opening in the foret canopy. Most of these released species were climax or subclimax species including: seedlings and saplings of the tree species pignut hickory, black oak, American elm, and white ash; shrub species such as flowering dogwood, eastern redbud, and summer grape; and forbs like sticky tickclover or sticky tick-trefoi and carrion flower.

The shrubs and forbs were present before lightening killed the pignut hickory as were some saplings of the four tree species. In fact, most plants of the tree species had established prior to lightening strike, but there were seedlings that had emerged and established after lightening created the forest gap. None of the shoots of the four tree species were stump sprouts or suckers (ie. release and regeneration of trees resulted from sexual reproduction). By contrast, most (probably not all) of the increased cover and biomass of shrubs and herbaceous species wasasexual reproduction (ie. proliferation of shoots from rootcrowns, rhizomes and so on).

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

181. The early years have the big changes- The greatest changes on a redeveloping sere (ie. recovey of disturbed or perturbed vegetation following denudation of a parcel of land via secondary plant succession) take place in the earlier or younger stages. This is the case under textbook old-field or cutover forest succession. That general pattern or form of recovery--in other words, classic secondary succession as described by countless workers even before Clements (1916) reviewed this work and gave Ecology what became known as the Clementsian paradigm--is the typical or usual scheme of plant community redevleopment on parcels of land of such size created by farming (fields), intensive logging (clearcut forests), ranching (especially overgrazed ranges), large wildfires, road construction, etc.

Small gaps such as those in forests formed by death of damage to a few trees as caused by localized blowdown, lightening, uprooting, or beaver feeding and similar small areas of denudation or disturbance on shrublands or, even, grassland (eg. a small burnt patch after the flames burned out in short order) differ in at least one aspect of defoliation from these situations. That aspect or facet of defoliation or denudation is area. The impacts of the other defoliation (denudation) facets of intensity, frequency, seasonality, and selectivity are moderated by decreasing size of the affected (denuded) area. This is because existing plant propagules are close at hand as well as the phenomenon by which remaining vegetation lessens impacts of such variables as ambient temperature, wind, precipitation, even humidity by presence of shade, windbreaking, transpired water, etc. This might be partially counteracted by attraction of animal attraction or, alternatively, this might also moderate or even facilitate recovery of vegetation via seed dispersal, addition of soil nutrients via animal excretion, and so forth.

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

182. Growing on my mamma's grave- Pignut hickory seedlings that are approaching sapling size growing next to the shattered trunk of the parent hickory (23 inch DBH) that was killed by lightening gthree years ago. There were also young trees of this same size (and presumedly roughly same age) of black oak whose parents missed the lightening bolt and were standing on the edge of this forest gap. There were also seedling of similar size/age of white ash the parents of which grew farther away and lower down on an adjoining north slope as well as American elm the fruit-bearing trees of which grew in closer proximity to the three-year-old gap. There were also seedlings/saplings of slightly smaller size (based on general height) of sassafras. There were no adult sassafras in this forest so origin of propagules was unknown: could have soil seedbank, carried in by wind or animals, whatever.

There were young plants (four to six foot shoots) of flowering dogwood and redbud along with stems of Virginia creeper and summer grape. Herbaceous species were sticky tick-clover and carrion flower were presented in accompanying slides.

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

183. Where forest succcession is just more of the same- Climax dominant tree species immediated reclaimed forest gap created when lightening killed a large (23 inch DBH) pignut hickory. Member species of the climax forest represented by seedlings (some of which approached small sapling size) included pignut hickory (probably the progeny of the dead tree), black oak, American elm, and white ash. There were also seedlings of sassafras, a seral tree species the propagule source of which was undetermined. Climax understorey shrubs were also poresent including redbud, flowring redbud, Virginia creeper, and summer grape.

Herbaceous species were not descernible in this slide, but they were presented elsewhere including the next two-slide set.

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

184. Rotting corpse and burial attendants- One of two main limbs of a large pignut hickory that fell following death by lightening strike three years ago.Surrounding the limb were two major understorey forbs, sticky tick-clover or sticky tick-trefoil. Also conspicuous were yearling seedlings of pignut hickory and American elm showing that climax tree species were replacing the dead climax co-dominant tree species. Likewise, these two forbs had been present in the climax forest only in a suppressed state. No suppression in this lightening -created gap in the forest canopy. In essence there was very limited secondary plant succession in this small patch. Instead there was enhanced growth of already present climax forbs and seedlings of climax tree species. There was also sexual regeneration by seedlings thqat emerged following gap formation.

The decomposer (reducer) component of this small, desturbance forest ecosystem was represented by numerous fruiting bodies of the shelf-fungus, white cheese polypore (Tyromyces chioncus), growing along the fallen limb of the lightening-killed hickory. a pignut hickory killed three years earlier by lightening.

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

185. Being digested by a "cheesy" character- Fruiting bodies of the shelf fungus, white cheese polypore (Tyromyces chioncus), growing on and rotting (digesting) the limb of a pignut hickory that crashed three years ago after the large adult tree was struck by lightening. The gap created in the forest canopy by the lightening strike was treated in detail above (and summarized immediately below). These and and the two immediately preceding slides emphasized to students that decompoesrs are part of range ecosystems. Such fungal reducers can be especially "photogenic" in forest range ecosystems.

Tyromyces chioncus is a member of Polypodaceae (polypore family) within order Polyporales, class Hymenomycetes and in subdivision, Basidiomycotina (Basidiomycetes).

Ottawa County, Oklahoma. Late July.

186. "Cheesy" details- Two carpophores (fruiting bodies) of white cheese polypore (Tyromyces chioneus) on downed and slowly rotting limb of pignut hickory. A closer-in shot of tow of the many such spore-producing structures of this particular organism in a Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130) in the western margin of the Ozark Highlands (Springfield Plateau).

The involuntary, hardwood host (source of detritis) of this heterotrophic organism, a fungal saprophyte, was killed three years ago by lightening strike.This served as a reminder to students of Ecology that any disturbance serves as habitat for some living thing. The blast from above that ended the life of this magnificant pignut hickory created a forest gap that released the offspring of the killed hickory (described above) and that created a new forest environment for other species, including white cheese polypore.

Ottawa County, Oklahoma. Late July.

Big lesson from a big boom: The concept, the pehnomenon, that emerged from this example of patch dynamics in a forest gap was that secondary plant succession did not--and, by extension, to similar forest gaps does not--apply at such small spatial scale. Range scientists such as Harold Heady (1975, ps. 10 [Figure 1-3], 12-70 passim) quite properly defined and described the four defoliation aspects or components of: intensity, frequency, season, and selectivity. These aspects of defoliation have been expaned to those of herbivory in general. These four correspond roughly to the Four Cardinal Principles of Range Management: proper degree (intensity) of use, proper season (season and frequency, interaaction thereof) of use, proper distribution of use (interaction of frequency and intensity), and proper kind and class of range animal (selectivity). (See again Heady, 1975, p. 7, Figure1-2).

To these four aspects or components of defoliation fire ecologists more recently added a fifth: size of area or extent (of defoliation) versus intensity (as in degree of) defoliation. By way of example consider a large-scale forest fire covering hundreds of square mile as compared to a quickly extinguished spot fire that grew to only a fewhundred square feet. Opportunity for regeneration by dispersed seed will be greater for smaller areas. Size and severity of modified microclimate and surface for accelerated erosion will be less for smaller areas of disturbance.In final, smaller disturbed patches will recover more quickly--other things being equal--than will larger areas of defoliation disturbance.

Furthermore, other things will not be equal, they will be more severe for larger areas of disturbance (bigger defoliation patches). If the black oak-pignut hickory forest on which the lightening-created gap occurred had been clearcut there would have been local areas of pioneer annuals such as crabgrass and weedy composites (ie. r-selected species). In the lightening-caused patch seem above there were only climax or, at least, high seral stage species and no annuals or biennials (ie. primarily K-selected species).

Response also varies with plant species that comprise the range--in this instance, forest range--community. One of the most important variables in this context is sprouting characteristics of species, especially of those harvested. It was shown above, for example, how readily black oak coppiced (stump-sprouted) following clearcut logging. This was also shown earlier in this chapter for a sugar maple-northern red oak-pignut hickory-white ash-forest which was less than two miles distant from the lightening-induced forest gap. In that climax, mesic (north-slope), sugar maple-dominated forest most of the felled mature trees of sugar maple, American elm, basswood, pignut hickory, and northern red oak coppiced such that the climax tree species were still present in the recovering forest vegetation during the first year following clearcutting. In fact, most of the same genetic individual trees started regrowth and remained in the forest community though, of course, at the "infant" stage. On the clearcut forest there were all of the standard pioneering (weedy) grasses (hairy crabgrass, Japanese brome) , forbs (sow-thistle, mare's tail or horseweed, giant ragweed), and, even, trees (sassafras), but there were also climax, native, herbaceous species such as wildryes, hairy bromegrass, and woodland bluegrass.

Still, size or area of disturbance was a factor in species composition of the disturbed range vegetation. Larger areas of defoliation (eg. clearcut forest) was more apt to have seral plant species and undergo secondary plant succession (at least components thereof) than small localized areas of disturbance characterized by death or defoliation of a single or few individual plants, even when these individuals were comparatively large.

To summarize, changes in vegetation within small disturbed areas (patch dynamics) such as forest gaps are less dramatic and dynamic than changes in plant communities within larger disturbed areas like clearcuts. In extremely small forest gaps, for example, plant succession might not occur because injured or damaged individuals either regrow (eg. coppice shoots from stumps) much like grasse and forb regrow following mowing and/or sexual progeny spring up from the soil seed bank. Instead of the lengthy and complex pattern of secondary--not to mention primary--succession, recovery of range vegetation from denudation might be a simple matter of progression of the same climax species or, even, the same genetic plants through their life cycles. In the case of the forest gap created by death of an adult, lightening-struck pignut hickory, forest recovery was a simplier, shorter-term matter of seedlings of pignut hickory and, to somewhat lesser degree, of other climax tree species growing to adulthood. Instead of forest gaps being "new land" of a denuded area (an old field, "go-back ground", or cutover forest) and beginning plant succession at the initial seral (pioneer) stage, revegetation of the climax community ( black oak-pignut hickory forest in this instance) was a matter of progressing through the life cycle of a new cohort of the sporophytic generation of climax (especially, dominant) plant species.

In essence, if the disturbance gap is small enough plant succession does not occur. Instead, individuals (sexual and/or asexual in origin) of the climax species just grow to mature, adult-sized plants. In such cases, there is little if any change in plant species composition of the disturbed vegetation. Climax species composition remains over the course of recovery. Changes in vegetation on small denudation gaps (patch dynamics) are limited to those of plant community structure (seedling or stump sprouts through old-growth size and form). In such circumstances development is growth, tissue differentiation, maturation, and reproduction of plants (ie.species-specific plant life cycle) not development of vegetation (ie. plant succession).