Oak-Hickory Forests-IIA

Forest Range Types of Eastern North America

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.

The classic and still-definitive work on forests of eastern North America (approximately east of the 98^th meridian) is the life's work of Dr. Lucy Barun (1950). Braun interpreted this entire vegetation as one great forest formation existing as a mosaic of forest regions which in turn were made up of community units that she labeled variously as belts, areas, districts, sections, divisions, etc.

"The Deciduous Forest Formation of eastern North America is a complex vegetation unit most conspicuously characterized by the prevalence of the deciduous habit of most of its woody constituents. This gives to it a certain uniformity of phsiognomy, with alternating summer green and winter leafless aspects. Evergreen species, both broad-leaved and needle-leaved, occur in the arboreal and shrub layers, patticularly in seral stages and in marginal and transitional areas. They are not, however, entirely lacking even in some centrally loocated climax communities" (Braun, 1950, p. 31). "The Deciduous Forest Formation is made up of a number of climax associations differing from one another in floristic compositon, in physiogonomy, and in genesis or historical origin. While the delimitation of associations may be made on a basis of dominant species, and it is from these that the climax is named, dominants alone fo not suffice for the recognition of these units. …Although the delimitation in space of an association is difficult, if not impossible, it is entirely possible to recognize and to map forest regions which are characterized by the prevalence of specific climax types, or by mosaics of types. These regions are natural entities, generally with readily observable natural boundaries based on vegetational features. …Forest regions must not be confused with climax associations. Even though a region is named for the climax association normally developing within it, it should not be assumed that the region is coextensive with the area where that climax can develop. Each of the several climaxes, although characterizing a specific region, nevertheless occurs in other regions." (Braun, 1950, p. 33-34).

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

Mixed Mesophytic Forest Region,
Western Mesophytic Forest Region,
Oak-Hickory Forest Region,
Oak-Chestnut Forest Region,
Oak Pine Forest Region,
Southeastern Evergreen Forest Region,
Beech-Maple Forest Region,
Maple-Basswood Forest Region, and
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.

Historical Footnote and Editorial

The consistent and persistent use of the eastern deciduous forest associations of Braun (1950) by the foremost contemporary ecologists provides the beginning student of Ecology with a textbook example of the necessity of learning the fundamental concepts— and the language(s) thereof —that are the foundation of his selected field of Biology. No ecological monograph, including those of John E. Weaver or Victor E. Shelford, ever used Clementsian concepts and terminology any more consistently or with any more practical application than did Braun (1950). All three of these (and there were others besides these) patriarchal ecologists of North American vegetation left future generations with not only the seminal but also the definitive treatises of the communities to which they devoted their professional lives.

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".

Miscellaneous Forest Types Within the General Oak-Hickory Region

Forest regions of Braun (1950, ps. 33-37) were each named for the "climax association normally developing within it", but forest region and forest association are not always coextensive. Also, while "a specific climax association charactrizes a region" there are numerous forest communities within a region that have species compostion, forest structure, and physiogonomy that are more like (and floristically have more in common with) forest associations of other forest regions (Braun, 1950, p. 34). These forest assocations, forest dominance types, and perhaps other hierarchial and classification units of vegetation may be physiographic or edaphic climaxes rather than climatic climaxes whereas other forest vegetation might be postclimax, preclimax, or subclimax (Braun 1950, p. 13). Some of these "outlier" or "island" forest cover types within the Oak-Hickory Forest (specifically the Oak-Hickory Forest Region) and characteristic plant species of them were presented below.

Sugar Maple Forest

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.

Along Modoc Creek, Ottawa County, Oklahoma. 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).

Note on common names: the common name for Carya cordiformis has been shown variously as either pignut or bitternut hickory depending on which source one refers to (Vines, 1960, p. 132; Steyermark, 1963, p. 516; Harlow et al., 1979, p. 261; Eyre, 1980, p. 137; McGregor, 1986, p. 132) with bitternut hickory being more frequently used. In the section below (and throughout this chapter) the present author used both common names interchangably or often both were presented seperated by "or" or, alternatively, with one common name enclosed in parentheses (behind or following the other).

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.

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).

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.

Along Modoc Creek, Ottawa County, Oklahoma. Early June (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).

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 leguminus 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.

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).

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.

Along Modoc Creek, Ottawa County, Oklahoma. April (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).

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.

Along Modoc Creek, Ottawa County, Oklahoma. Late November (autumnal aspect; leaf fall for most trees and shrubs except the oaks). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax 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).

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.

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).

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

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).

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

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.

12585 and 12586.

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.

12587 and 12588.

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.

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

Ottawa County, Oklahoma. Late May.

Burdened down- Weight of heavy spikelets at anthesis resulted in the drooping of this panicle on hairy woodland brome. Bluffs above Modoc Creek in western Ozark (Springfield) Plateau.

Ottawa County, Oklahoma. Early June (this was another year than that in which preceeding photographs were taken; note difference in progression or stage of phenological development from one year to next).

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, anthesis stages of phenology.

Tried for a closer-in view- These were beautiful (nearly perfect) slides of a portion of panicle with spikelets at anthesis in hairy woodland brome before Epson "Perfection" 700 scanner botched them up (total FUBAR). There are no really good scanners out there, Epson "Perfection" is a perfect Wall Street scam and confidence game. Do not buy Epson products. Too late for this author, but others stay away.

Anyway, maybe viewers can get the idea.

Ottawa County, Oklahoma. Early June.

Sagging under a heavy yield- Panicle of hairy woodland brome

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

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

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

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

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

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

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).

Dead and alive in winter- Last year's dead stalks (brown straw) and current year's young living shoots (short, green leaves) of hairy woodland brome growing on an upland, black oak-dominated forest, a mesic, limestone to chert forest site. Most of the shed leaves are those of black oak. This second-growth forest was at or closely approaching the old-growth stage

Above bluffs of an Ozark Plateau stream (Modoc Creek). Ottawa County, Oklahoma. Early January; early growth stage and onset of decomposition stage of dead shoots.

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.

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

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

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

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

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.

More peanut-flavored legumes- Flower cluster that progressed to fruit-stage (first slide) and both individual flower and legumes (second and thrd slides) of sticky tick-trefoil or pointed-leaf tick-clover in the understorey of an upland chert, black oak-pignut hickory forest in the Springfield Plateau.

This Ozark forest was grazed/browsed by white-tailed deer but "off-limits" to livestock. As such the understorey--both herbaceous and lower woody layers--was well-developed with a rich species diversity. The understorey had been burnt by a light surface fire five years ago (prior to time of these slides).

Ottawa County, Oklahoma. Mid-July; late bloom/ripening fruit stage of development.

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.

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

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

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

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

Just outward and upward from where sugar maple was dominant:

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

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

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

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

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.

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.

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.

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.

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.

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.

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.)

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.

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

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.

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

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.

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

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.

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

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.?).

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).

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.

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.

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.

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.

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.

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.

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.

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.

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.

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

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

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

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

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

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

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.

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

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.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Bottomland Forests

Within the Oak-Hickory Forest Region there are some bottomland forests in which oak and hickory species are subdominant or associate (vs. dominant) species (Braun, 1950, chapter 6). Somewhat surprisingly, however, Braun (1950, ps. 168-169, 174, 189) provided almost no coverage of bottomland forests in this region. On many of these bottomland forest ranges the dominant (climax or subclimax) trees include sycamore, eastern cottonwood, hackberry, sugarberry, bitternut hickory, black walnut, box elder, American elm, chinquapin oak, sugar maple, honey locust, and Kentucky coffeetree.

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).

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

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

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

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

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).

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

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

Start of a new black walnut-

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

How it starts in the soil-

Ottawa County Oklahoma. Mid-June; early seedling stage.

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.

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.

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.

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.

How not to do it- In August or September (probably August) 2010 amateurs with chainsaws high-graded part of the Modoc Creek forest plot that was at subclimax stage of the hackberry-elm-pignut hickory-sycamore-white ash cover type. This (former) relict forest of only three to four acres was an "amalgam" of three separate properties. Only one authorized timber sale. Wood-cutters (no way would they qualify for fellers) purposely stole two logs (a sycamore and a black walnut) off of the neighbors. This could not have been a mistake (even a careless one) because mostly unfenced properties were clearly marked by concrete fence posts on the land from which sale was authorized and by a railroad tie that was the corner post (and all that remained) of the property line fence where all three properties met. There was apparently only one (at most, two) chainsaw-users as verified by "signature" evidence of complete lack of undercuts on all except two stumps (ie. trees, boles, were flat-sawed). All except two stumps lacked stump stops (height of the hinge; difference between undercut and backcut). The only effective hinges were slight differences in chainsaw paths through tree boles. As a result all stumps (other than the two) had long strands (up to eight feet in height) of wood fibers pulled from the butt log (stump section of the trunk) thereby degrading value of the largest log (= most lumber). The buffoons who improperly felled the trees were clearly rank amateurs (ie. they were not fellers just duffuses who were handed chainsaws). In addition, they were thieves. Furthermore, they left some valuable lumber in the woods to rot, including a seven or eight foot section of clear, straight, mature black walnut that they had stolen from the neighbors. In addition (and perhaps most telling and important of all) was the "hatchet-happy" habit of needlessly felling young pole-size trees (ie. the next wood crop) including trees that were clearly not in any regards in the way of tree felling or skidding of logs to trucks). Finally, as if all this was not enough, chainsaw-using, "wood idiots" wasted effort and material to fell several patently obvious hollow trees (worthless for lumber). These were then left to rot or, worse yet, once the amateurs realized trees were hollow they withdrew saw blades leaving the trees standing as forest hazards. In fact, all of the sycamore logs were, to some degree or the other, hollow-except for the one stolen from the neighbors. Most of the sycamores were growing on and stabilizing the bank of Modoc Creek. Such logging of riparian trees is now banned on federal land other than to remove hazard trees in camping and picnic areas.

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.

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

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

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.

12062.

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..

12063 and 12064.

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..

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

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.

111459 and 111460.

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.

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

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.

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

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.

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