Southern and Central Forests II

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 typzes 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 98th 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:                 1. Mixed Mesophytic Forest Region,                 2. Western Mesophytic Forest Region,                 3. Oak-Hickory Forest Region,                 4. Oak-Chestnut Forest Region,                 5. Oak Pine Forest Region,                 6. Southeastern Evergreen Forest Region,                 7. Beech-Maple Forest Region,                 8. Maple-Basswood Forest Region, and                 9. Eastern Hemlock-Eastern White Pine-Northern Hardwoods Region.  Braun (1950, ps. 11-12) interpreted these same combinations of species as forest communities at the scale (both spatial, mostly, and, also, temporal) of climax association  from which, as quoted immediately above, Braun derived the names of forest regions. Braun (1950, ps. 11-12) distinguished between the association-abstract and the association-concrete, a distinction discussed in the review of the derivation of vegetation cover type from the concept of plant association. The Braun association is the association of F.E. Clements. Indeed the entire ecological paradigm on which Braun (1950, ps. 10-15) based her monographic treatment of the North American Deciduous Formation is Clementisan except allowance for and inclusion of edaphic and physiographic climaxes of Cowles, Tansley, etc.  Vankat (1979, ps. 137-150) and Delcourt and Delcourt in Barbour and Billings (2000, ps. 365-378) described eastern deciduous forest vegetation under the Braun (1950) associations of the Clementsian model. It is important to bear in mind that the Braun associations can occur in more than the one forest region bearing the name of the association (eg. the Oak-Pine Association commonly occurs and the Maple-Basswood Association infrequently occurs in parts of the Oak-Hickory Forest Region). Several of the species combinations that delineate deciduous forest regions and associations were also used as forest cover types by the Society of American Foresters (Eyre, 1980) as for example White Pine-Hemlock (SAF 22), White Pine-Northern Red Oak-Red Maple (SAF 20), Sugar Maple-Basswood (SAF 26), and Beech-Sugar Maple (SAF 60). The Society of American Foresters emphasized that it’s forest cover types were “based on existing tree cover” (… forest as they are today…”) and that some types may be climax while others are “transitory” (ie. seral stages leading to another climax). Braun (1950, p. xiii) specified: “Some of the communities for which composition is given are readily referable to ‘forest cover types’ as defined by the Society of American Foresters”. She then added, “However, an attempt to classsify all communities as to ‘cover types’ would be artificial” and often impossible. Undoubtedly this was due to the differences in classification by Braun’s climax basis (with seral communities clearly specified) versus the existing or present-day forest communities basis of the SAF.  The Society for Range Management (Shiflet, 1994, p. xi) also specified the criterion of “existing vegetation” and that some rangeland cover types are climax and others are seral. The author of this collection of photographs and descriptions repeatedly reminded readers of this situation, but specified that most of the rangeland and forest cover types included herein were climax vegetation. That criterion exist for forest range types of the Eastern Deciduous forest Formation with most photographs being of either old-growth or second-growth forest with climax species composition as described in the classic literature such as Braun (1950) or Shelford (1963, ps. 17-119). The nine forest regions of Braun (1950, ps. 35-37) were retained with little modification as series in the fairly comprehensive system of vegetation (primarily, climax; secondly, disclimax or subclimax) used in A Classification of North American Biotic Communities by Brown et al. (1998). Their organization of the Eastern Deciduous Forest Formation was: Oak-Hickory Series, Oak-Chestnut Series, Beech-Maple Series, Oak-Pine Series, Maple-Basswood Series, and Hemlock-White Pine-Mixed Hardwood Series within the Northeastern Deciduous Forest biotic community and Mixed Mesophytic Series and Pine Series within the Southeastern Deciduous and Evergreen Forest biotic community. The Brown et al. (1998) series were included below following SAF and/or SRM cover type designations. Additional designations as for forest wetlands were shown as required.

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”.

Bottomland Forest- Example along a stream in the Ozark Plateau

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. The first sequence of photographs of this bottomland forest presented the vernal aspect of the vegetation in which the herbaceous layers were at peak standing crop andjust prior to summer dormancy. Emphasis was laid on showing species composition and structure of this forest range when the greatest number and most important indicator species would be visible and flowering. This is the spring season, especially for the cool-season festucoid grasses that are dominants of the herbaceous layer of the understorey. Plant species were described and shown in their spatial relations to each other, but without reference to their successional status or the dynamics of this forest range vegetation. Development of the forest community and dynamics in response to disturbance were discussed in the section that followed this one, and at the estival aspect when many of the herbaceous species had gone into dormancy.

1. A lot of players along the old channel- During some extraordinary flood in decades (probably centuries) past Modoc Creek eroded an accessory or overflow channel. When this Ozark Plateau stream returned to regular or normal flow (receded back to its normal channel) the newly cut, extraordinary flood channel remained as a denuded area that served as a sere on which plant succession progressed. Eventually a late seral or climax forest developed on/in this overflow . channel that had long since been abandoned by Modoc Creek which flowed in its regular channel tha runs parallel to the now-forested former flood channel. The forest (stand might be a more precise term) that developed on bank and bed of the overflow stream channel was conterminous with forest vegetation along the regular channel of Modoc Creek and limestone bluffs above this stream as well as with the forest community (again, stand might be a more explicit term) on the floodplain between the two stream channels. There was a great diversity of tree species within this entire tract of floodplain forest including sycamore, eastern cottonwood (Populus deltoides var. deltoides), black walnut, red mulberry (Morus rubra), box elder (Acer negundo), chinquapin oak, northern red oak (Quercus rubra), Shumard oak (Quercus shumardii var. shumardii), white ash (Fraxinus americana), sugar maple, bitternut or pignut hickory, American elm (Ulmus americana), red or slippery elm (U. rubra), and western hackberry (Celtis occidentalis var. canina). The successional status of these sundry species (at least some of them) was discussed in another section below. Tree species on the high bank of the overflow stream channel (right side of photograph) were (left to right): Shumard oak, chinquapin oak, and pignut or bitternut hickory. Younger (smaller) trees on the low side of the channel (center and left of photograph) were hackberry and American elm. The woody vine was that of a species of grape (Vitis sp.). Dominant shrub was spicebush (Lindera bezoin); local associate shrub was buckbrush or coralberry (Symphoricarpos orbiculatus). Herbaceous dominant of this understorey was silky wildrye (Elymus villosus) with some Virginia wildyre (E. virgincus). Canada or hairy wood brome (Bromus purgans= B. pubescens) was also present but widely scattered and most frequent at bases of tree trunks. Ottawa County, Oklahoma. May (vernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). MesicBottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

2. Beneficiaries of many floods- Bottomland hardwood forest on the floodplain of Modoc Creek in western Ozark Plateau. Dominant of herbaceous layer(s) was silky wildrye with local patches of Virginia wildrye. Largest tree in both photographs was sycamore (right of center in midground of first photograph; right margin of midground of second photograph). Other trees included additional sycamore, hackberry, American elm, and chinquapin oak. Sapling in foreground of both photographs was American elm with Virginia creeper climbing in it. Spicebush and buckbrush were visible in both slides. Ottawa County, Oklahoma. May (vernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). MesicBottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

3. Spring verdure of floodplain forest- Early morning light showed herbaceous layer(s) of this botanically rich bottomland forest to good advantage. Silky wildrye (overall dominant herbaceous species) and Virginia wildrye at peak standing crop beautifully represented the feed value of this forest range and the climax cool-season grasses that provided it. Sycamore (big trunk in center midground) and northern red oak (big dark trunk at left margin) along with western hackberry and both American and red elm (smaller boles: poles and saplings) were the tree species present in this stand. Leaves in left foreground were hackberry. It was obvious in this photograph that elms and hackberries were replacing sycamore which had very little regeneration, certainly nothing approaching that of hackberry and the two elm species of which American elm was the more abundant. More discussion on this dynamics was given later in this section. Ottawa County, Oklahoma. May (vernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). MesicBottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

4. Composition and structure of an Ozarks bottomland forest- The beauty as well as the botanical makeup and internal architecture of a mixed hardwood floodplain Ozark forest in full array of "spring fashion". Peak standing crop for grasses, all of which were cool-season, festucoid species. Silky wildrye was the dominant with Virginia wildrye the associate. Canada or hairy wood brome was present as robust but widely scattered individuals. The second of these two photographs was a closer-in view of forest range vegetation presented in the first photograph. The smaller trees (pole-size) in foreground of both slides was hackberry which, along with American or white elm and red or slippery elm, made almost all of the young tree stock. (More on this development later in the show.) The largest tree (darker trunk at right midground of both slides) was northern red oak. The other two large trees (left midground) were sycamore. There was almost no reproduction of sycamore of northern red oak. Most abundant shrub in these views of the forest community was buckbrush (eg. large bush in foreground). Ottawa County, Oklahoma. May (vernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). MesicBottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

5. Shadows and spring green-This "photo-plot" featured a young western hackberry (of smaller or younger adult age class) in front of an old sycamore symbolizing replacement of the latter by the former. This same trend was introduced in immediately precdeding slides. The successional relations between sycamore and western hackberry (also American and red elms) were described in the subsequent section that described this bottomland forest. The shrub in right corner was a handsome specimen of spicebush. The more common and lower-growing shrub was buckbrush. The dominant (present everywhere) was silky wildrye. Local patches of Virginia wildrye accompanied it fellow Elymus species. Ottawa County, Oklahoma. May (vernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). MesicBottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

6. Verdant spring attire- Another view of the species composition and and interior structure of a mixed hardwood bottomland forest on the floodplain of an Ozark stream. Between the regular channel (for normal stream flow) and the ovrflow channel of Modoc Creek a species-rich forest range had developed. This photograph provided a different vantage point of some of the same vernal aspect vegetation introduced above. Point of peak standing crop and soft to hard dough stage of grain in the cool-season festucoid grasses that made up most of the heerbaceous understorey in the vernal society of this layer. Silky wildrye was the dominant with Virginia wildrye the associate herbaceous species. There were a few robust plants of Canada or hairy woody brome. Largest tree (dark bark at right midground) was a northern red oak. The two other large trees (one in right midground; the other in right background) were sycamore.The four pole-size trees (including two in foreground) were hackberry. Virginia creeper was creeping up the trunk of the foremost hackberry. Shrubs were mostly buckbrush in this shot of the understorey. Spicebush was generally the dominant shrub, but it and buckbrush were in two distinct woody layers: low-shrub and mid-to tall-shrub). Ottawa County, Oklahoma. May (vernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). MesicBottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

7. Sample of herbaceous sward- Vigerous specimen of silky wildrye The broad-leafed forb was pokeberry, pokeweed, or poke (Phytolaca americana). The trailing forb in foreground was cleavers or bedstraw (Gallium aparine), a minor (incidental) species. Ottawa County, Oklahoma. On the floodplain of Modoc Creek.

8. A pleasant and all-too-infrequent encounter- "Hale and hearty" individual of Canada or hairy wood brome (Bromus purgans= B. pubescens) growing near base of an equally hearty northern red oak. This decreaser grass is an indicator species. Its presence on this flood plain forest range indicated that any past overgrazing had been long enough ago so that this abuse-sensitive grass had recovered to the point of having some healthy individuals to show off to those appreciative of the finner things in life. Ottawa County, Oklahoma. On the floodplain of Modoc Creek.

9. A closer encounter- Panicle and spikelets of Canada or hairy wood brome of the plant shown in the preceding photograph. This is a climax grass of Ozark Plateau which has sadly become all too uncommon on forest ranges subjected to overgrazing by cattle (and in earlier years by hogs and horses). Pubescence on bracts of spikelets is a typical identification feature of this species. Ottawa County, Oklahoma. On the floodplain of Modoc Creek.

10. Morning light on spring green- Two-slide sequence showing a panaramic (panned) view and a more species-specific (zoomed) view of the range plant community of a bottomland forest that had developed on the foodplain of a stream (Modoc Creek) in the Springfield Plateau of the Ozark Plateaus physiographic province. The biggest (and rightward leaning) tree was an old-growth (and hollow) sycamore. The large tree with fine straight bole in right bacckground of first photograph and to right of leaning sycamore (but mostly concealed by leaves) in second photograph was a northern red oak (also an old-growth specimen). Other trees included more sycamore (too distant to be seen distinctly), black walnut (shown specifically in another photograph below), and chinquapin oak (right margin of the first of these two slides). Most of the smaller, younger trees (mostly sapling to pole age/size classes) were western hackberrn and American elm which were succeeding the pioneer and persistent sycamore, northern red oak, black walnut, and chinquapin oak. (This phenomenon was described in later parts of this section devoted to Ozark floodplain forests.) Good examples of an American elm were the three foremost saplings in first of these two slides and in left foreground in the second photograph. Most of the pole-size tree in both slides were hackberry. The foremost tree at left margin was a mid-size white ash which, along with hackberry, American and red elms, and pignut or bitternut hickory were ascending to dominancy as this bottomland forest approached the climax stage of plant community development. Grayback and fox grape grew up into crowns of all the old-growth trees. Virginia creeper was even more common (eg. American elm sapling in foreground ). The dominant shrub was spicebush with pawpaw surpassing buckbrush as the associate species in this closer to Modoc Creek. American bladdernut became a major shrub down closer to this stream. In the herbaceous zone shown in both of these "photo-quadrants" silky and Virginia wildryes, the dominant and associate species, respectively, in most of the general herbaceous layer, were replaced in dominance in local habitats by wood nettle (Laportea canadensis). Slender nettle (Urtica gracilis= Urtica dioica ssp. gracilis ) was also present, but it did form exclusive, single-species colonies in the matter of wood neetle. In regard to species diversity, plant families represented, forest structure, plant growth form, plant age distribution, microsite stands (eg. wood nettle), and even local disturbance due to blowdown and breakage from recent ice storms (covered below) this forest range was almost unparalled even by standards of a bottomland forest. In spring verdure it was of unsurprassed beauty, raw and sheer grandure as a sad reminder of what forests of the Ozark Plateau once were. Of course no photograph could capture the rapture envoked by such beauty tempered with crawing ticks, webs of orb spiders in the face, and britches sopping-wet from dew. Some things simply have to be experienced personally and, better yet, when alone but for the presence of God. Ottawa County, Oklahoma. May (vernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). MesicBottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

11. Forest garden in the morning- Enough morning light penetrated the canopy of a floodplain forest along an Ozark Plateau stream (Modoc Creek) to provide more detail of species composition and structure of vegetational layers of this diverse stand that was introduced in the preceding two-slide set. This present photograph was taken from a different vantange point so that the large northern oak is no longer visible except in distant background (right margin). Tree regeneration was represented by the sapling age/size class. These trees were all western hackberry and American elm. There was no reproduction of sycamore in this "photo-transect". Sycamore (most in background) were all arge,mature (over-ripe) trees. Just beyond (to right) of this "photo-transect" larger trees were chinquapin oak and box elder with most regeneration that of white ash. Prominent shrub species in foreground (eg. left margin, foreground) was spicebush. Pawpaw was also present as was buckbrush though this latter at much less cover than in the forest stand growing on the abandoned overflow channel of Modoc Creek shown in first photographs of this section. The dominant herbaceous species in this slide was (as in portions of the understorey shown in preceding two photographs) wood neetle. Silky wildrye and Virginia wildrye (dominant and associate herbaceous species, respectively, in other parts of this understorey) were much less abundant closer to the regular-flow channel of Modoc Creek. Slender nettle was also present, but not as local patches like stinging neetle. Ottawa County, Oklahoma. May (vernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). MesicBottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

12. A stinging selection- Grazing by white-tail deer on shoot apices of wood nettle (Laportea canadensis). This species grew in large local colonies in a floodplain forest of such species as sycamore, western hackberry, American elm, box elder, white ash, black walnut, northern red oak, chinquapin oak, and eastern cottonwood. This specific colony grew near base of a sycamore and northern red oak.

13. Dynamics of forest vegetation- The key features of plant community ecology tracing Frederic E. Clements, to the most prominent founder of the specialty, were: 1) the ever-changing, cyclical pattern of vegetation and 2) temporary stability (climax) before disturbance set the sere in chage again. This photograph was an example of Clementsian vegetational dynamics in the bottomland forest that developed on the floodplain of Modoic Creek, a stream in the western edge of the Springfield Plateau of the Ozarks Plateau physiographic province. Western hackberry and American elm were replacing pioneer tree species like sycamore, black walnut, and eastern cottonwood that had persisted to a late seral stage (subclimax). This photograph featured a mature black walnut with an amazingly tall bole in its prime being replaced by American elm (pole-size trunk to immediate right of the black walnut) and hackberry (represented by large shade leaves at top and right margin of slide), the ultimate climax tree species of this forest sere. The black walnut was actually overripe and already too far gone for valuable lumber. It was mostly hollow and serving as a bee tree (used as a hive by natualized honeybees [Apis mellifera]; entrance, hole from a dead limb, was on backside of tree as presented here). This walnut had just begun to shed limbs from its crown as shown by two dead ones leaning against or toward the left side of its trunk. There were no seedlings of saplings of black walnut. This walnut had established on the sere at an early stage and persisted into the subclimax as an over-mature individual while those species that would be climax dominants established all around it. Black walnut has a tolerance rating of Intolerant as compared to that of Tolerant for box elder and Intermediate for hackberry and American elm (Wenger, 1984.ps. 2-3). The dominant shrub was spicebush with buckbrush as the associate shrub. Pawpaw was also present. (Actually each of three species dominated a tall, middle, and low shrub layer (pawpaw, spicebush, and buckbrush, respectively). A small sapling of box elder was present (right margin of slide). There were no herbaceous species in this local habitat, the shade being too dense for species like the wildryes and wood nettle. Ottawa County, Oklahoma. May (vernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). MesicBottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

14. Bed on the bank: a forest forb and decreaser grass as indictor species- On part of a bank of an abandoned channel of Modoc Creek a "bed" (local stand) of southern or Virginia blue flag (Iris virginica) and silky wildrye had developed as part of the herbaceaous understorey of a bottomland forest composed of sycamore, eastern cottonwood, western hackberry, white ash, chinquapin oak, boxelder maple, northern red oak, black walnut, and American elm. This bank of the former overflow stream channel was made up of larger stones and gravel along with alluvial soil. The edaphic part of these species' habitat was fertile and well-drained. Duration of sunlight was limited to roughly half of the daily photoperiod only some of which received full-light intensity due to shade from nearby large trees. Iris virginica is extremely rare in this locale and its presence here attested to the virgin environment of this old-growth floodplain forest. Silky wildrye is a decreaser native grass. These two herbaceous species indicated the climax (or near climax) state of the forest range described here. On abandoned overflow channel of Modoc Creek. Ottawa County, Oklahoma. May.

15. A bottomland bouque (or everybody's blooming)- Close-in view of a local stand of Virginia or southern blue flag and silky wildrye, both species of which were in full-bloom, in a bottomland forest of sycamore, western hackberry, chinquapin oak, northern red oak, American elm, black walnut, white ash, and eastern cottonwood. This pretty "photo-quadrant" presented details of the local herbaceous plant community that was introduced in the preceding slide. On abandoned overflow channel of Modoc Creek. Ottawa County, Oklahoma. May. Phenological stages: peak-bloom for southern blue flag, soft-dough stage for silky wildrye.

16. Floral treasures in the forest- Details of southern or Virginia blue flag (Iris virginica) on part of the bank of an abandoned overflow channel of Modoc Creek, a stream in western edge of Springfield Plateau of Ozark Plateau Region. Silky wildrye accompanied the native iris. Was there ever a more beautiful bouque? Ottawa County, Oklahoma. May. Phenological stages: peak-bloom for southern blue flag, soft-dough stage for silky wildrye.

17. Maturing spikes- Spikes of silky wildrye (Elymus villosus) in bottomland forest on the floodplain of an Ozark Plateau stream (Modoc Creek). Ottawa County, Oklahoma. July. Hard-dough phenological stage of silky wildrye.

18. Study of an understorey grass- Three-slide sequence showing sexual shoots of Virginia wildrye (Elymus virginicus) in forest understorey in western Ozark Plateau. Peak standing crop with grain in mid-dough phenological stage. Ottawa County, Oklahoma. July.

19. Spikes in the shade- Examples of spikes of Virginia wildrye in understorey of western Ozark Plateau forest at mid-dopugh stage of phenology. Ottawa County, Oklahoma. July.

20. Plant succession "ain't" always straight forward- The concept of tolerance is a cornerstone of forest succession. A discussion of this fundamental concept and the phenomena involved in it are beyond purview of this publication (at least at this juncture). A concise definition for this factor (or group of interacting afctors) in context of plant succession in forest vegetation seemed sufficient. "The capacity of trees to grow satisfactorily in the shade of, and in competition with other trees; if intolerant of shade, they are temred light demanders; if tolerant, shade bearers" (Helms, 1998). Tree and shrub species are rated as to their tolerance and givern rankings ranging from Very Tolerant to Intolerant (Wenger, 1984.ps. 2-4). Generally speaking woody plants that are Very Intolerant tend to be early seral or even pioneering species. Intolerant species that were major trees growing on the floodplain forest being considered here were eastern cottonwood, black walnut, hickories, black cherry, and sycamore. At the other end of the tolerance spectrum Very Tolerant species (sugar maple on bluffs along this floodplain forest) and Tolerant species (box elder in this bottomland forest) are generally dominants of climax forests. Tree and shrub species rated as Intermediate (eg. northern red oak, hackberry, American elm, and white ash on this bottomland forest) are also frequently climax dominants (or associate species to those with greater tolerance). Forest Ecology is, of course, not a precise science in the fashion of Physics. Hence, forest succession and silvicultural application--including grazing/browsing management--are not as simple as spaceship design. Things are "more messy" in the biological sciences and their applied fields. With regard to the phenomenon of tolerance some species have tolerance ratings that are inconsistent or unclear or, satted more clearly perhaps, dependant on local site conditions. Sycamore, some hickories like bitternut or pignut hickory, hackberry, and American elm fall into this "shade of grey" category (Wenger, 1984, ps. 3-4). These species were dominant trees on the bottomland forest that developed on the floodplain of Modoc Creek used as an example of this forest range type. These two photographs (the second a zoomed-in view of the overall view in the first) showed a local floodplain forest stand in which plant succession seemed somewhat "confused". This forest range vegetation was a small grove of mature white ash with most revealing (and confusing) lower layers. Greater detail of the forest range of this ash grove was discussed in the photograph following this set. The first lesson of forest range vegetation presented here was in the first of these photographs. The sapling in foreground and the larger tree (pole-size) in right midground were slippery or red elm which had regenerated as part of the climax forest vegetation. There were also seedlings (not visible) of chinquapin oak in the herbaceous understorey which was dominated by silky wildrye almost exclusively in this floodplain forest except for patches of stinging nettle (shown above). Regeneration of chinquapin oak in this bottomland forest was consistent with existence of a sugar maple-chinquapin oak cover type and presence of chinquapin oak in bottomland forests where white ash is an associate species (Fralish and Franklin, 2002, p. 478-479). Burns and Honkala (1990) placed chinquapin oak in Intolerant rank, but they specified that young trees tolerated moderate shading (Intermediate tolerance rank) and became less shade-tolerant with advancing age. They concluded that chinquapin oak was a subclimax or even climax tree species on more mesic forest sites (Burns and Honkala, 1990). Red or slippery elm has generally been regarded as Tolerant so its general regeneration on this well-drained bottomland, including under cover of white ash, was consistent with silvics of this species. The second and major lesson of these two slides was presence of sycamore at small sapling size in the shade of a dominant tree of the climax forest. It was explained in the first paragraph of this caption that sycamore was ranked as an Intolerant species yet one of varying tolerance response and thus some uncertainity of tolerance rating. Sycamore is also a very long-lived tree (by longevity standards of species comprising flora of this region). Sycamore is one of the first tree species to invade habitats of severe disturbance such as stream scouring and cutting of new stream channels (ie. a pioneer species), but it subsequently persist to the stage of subclimax or even climax.forest. This silvic feature combined with rapid growth rate is why sycamore is the largest-diameter tree and one of the tallest-growing hardwood species in North America. Fralish and Franklin (2002, p. 483) published similar conclusions regarding sycamore. If this little sycamore sapling "played by rules" of plant succession it would not be here. But there it was: an Intolerant species had reproduced and was doing just fine beneath adults trees that were climax dominants of this forest cover type. Here was photographic evidence of the unclear tolerance ranking--and the successional status--of sycamore. The specifics of forest succession have yet to be determined conclusively. Perhaps they can never be because they do not aways "stick to the successional script". Plant succession may not always be direct and simple, but it is "cool". Sycamore remains a member--one of the biggest ones--of the climax bottomland forest. Floodplain of Modoc Creek. Ottawa County, Oklahoma. June

21. Climax pals- Silky wildrye, the overall dominant species of the herbaceous understorey, and white ash, one of the climax dominant trees, were the principal plants of a floodplain forest in the western Ozark Plateau (Springfield Plateau thereof). Sharing the "spotlight", that ever-shifting shaft of sunlight that was a (perhaps the) major abiotic factor in this forest, was Virginia creeper (climbing the left white ash trunk) and buckbrush (to immediate left of the left trunk). The most revealing "understudies" in this local lineup of species were seedlings of white ash and Americanaelm (foreground; in front of the left bole of white ash). Regeneration of these latter two tree species was evidence of their status as dominant trees of the climax bottomland forest. Continued presence of silky wiildrye inside the grove of white ash testified to role of this cool-season grass as the general dominant of the herbaceous understorey and main forage species in this forest. The principal forb was purple Joe Pye weed. All-in-all, a very revealing local "photo-quadrant". Floodplain of Modoc Creek. Ottawa County, Oklahoma. June.

This bottomland forest now being described had 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...

22. 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 with a supporting arboreal cast including hackberry or western hackberry, American elm, pignut or bitternut hickory, slippery or red elm, chinkapin (chinquapin) oak, black cherry, black walnut, box elder, red mulberry, northern red oak, and Shumard oak. 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: grayback or sweet winter grape (Vitis cinerea) 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.

23. 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).

24. 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 grayback or sweet winter 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).

25. 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 grayback 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), part of the Springfield Plateau, 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).

26. 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).

27. 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 cotton 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).

28. 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).

29. 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).

30. 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 grayback 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 mouse-ear chcicweed (Cerastium vulgatum), the native chickweed (C. 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).

31. The old channel in winter- Another photo-plot of the western hackberry-American elm-sycamore-boxelder bottomland forest described in preceding photographs. This forest community developed along the former channel of Modoc Creek after this stream cut a new channel. Featured here was the forest range vegetation shown in the midground of the preceding photograph. Largest tree in background with forked trunk was an old-growth eastern cottonwood that pioneered the former stream channel. The tree in left foreground with basal firescar was a chiquapin oak. The two trunks in front and slightly to left of cottonwood were boxelders. Large grape vines drapped down into the abandoned channel bed. Understorey was not prominent here, but was well-developed elsewhere in this forest. Details of understorey were presented below. Floodplain of Modoc Creek, Ottawa County, Oklahoma. January (hibernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). Mesic Bottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

32. 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).

33. 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 stick 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.

34. 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 grayback and fox grape. Larger tree at extreme right margin and midground of first photograph and again in midground of second slide was western hackberry. Almost all trees in midground were hackberry. The tree with bent trunk (to right of the larger hackberry in midground) was boxelder. Most of the trees in background were sycamore (readily distinguished by upper gray bark) growing along bank of Modoc Creek, a typical Ozark Plateau stream. There were not oaks or cottonwood in this stand. Most of the shrubs were spicebush (Lindera bezoin), but some pawpaw (Asimina triloba) grew in shrub understorey closer to Modoc Creek. Both the shrub layers and herbaceous layers were described immediately below. Floodplain of Modoc Creek, Ottawa County, Oklahoma. January (hibernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). Mesic Bottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

35.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.

36. 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 mouse-ear chickweed (Cerastium vulgatum) and dead nettle (Lamium purpureum), 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.

37. 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.

38. 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.

39- 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, a snag, of the American elm 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? (See later slides of plant and vegetation recovery below.) 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).

Follow-up photographs (five and seven months following ice devestation) of forest vegetation on this exact spot were shown below).

40. 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).

41. 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).

Follow-up photographs (five and seven months following ice devestation) of forest vegetation on this exact spot were shown below).

42. Crowned- The upper trunk of a western hackberry was broken by ice accumulation from a late autumn (9 Decembr, 2007) storm of wind and mixed precipitation (that promptly froze and built-up on trees). In crashing, the crown of the hackberry broke and bent down a sapling of American elm (to left of hackberry) The most extensive damage was to boxelder, but sycamore, chinquapin oak, black cherry, and Shumard oak were all affected by the storm and suffered major loss of limbs and branches. Centermost tree in foreground was a hackberry that experienced only minor damage. Floodplain of Modoc Creek, Ottawa County, Oklahoma. January (hibernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). Mesic Bottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

43. 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. Damage done by same ice storm as in preceding slides (9 Decembr, 2007). 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).

44. 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).

45. Browsed by ice (recovery efforts; phase 1) Early stage of recovery of vegetation in a mixed hardwood bottomland forest becoming dominated by western hackberry and American and red elm that had been severely damaged by a late autumn (9 December, 2007) ice storm. It was only six to eight weeks into the warm-growing season (that length since the last killing frost) so that plant growth and sesonal development was still at early stages of phenology and recovery. Both the larger completely topped snag and the "broke down and pert-n'ar broke off" sapling were American elm. It was wondered above (first slides of ice storm devestation) if the nearly benearly broken off American elm would resprout at point of breakage or at base of trunk (stump sprouting). The answer was provided here: at point of break-over and also along the incompletely severed bole distant (upward) from point of breakage. In other words, all parts of this trunk were still alive at this point in time (both season and plant age). Spicebush was th e shrub species that was most abundant and benefitted greatest (at least initially) from the forest gaps created by ice defoliation. Pawpaw and buckbrush were also favored by ice-created forest openings. Grayback and fox grape had sprouted profusely following some ice damage and creation of the forest gap. Major herbaceous species were silky wildrye, pokeweed, and purple Jope Pye weed. The large, split limb in foreground of these three photogrphs (best seen in right center foreground of third photograph) was box elder. It had been a major part of the crown of the largest tree in right margin of the first two photographs. Ottawa County, Oklahoma. May (vernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). Mesic Bottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

46. Browsed by ice (recovery efforts; phase 2)- Early versus mid-stage of recovery of vegetation in a forest gap created by severe ice storm (9 December, 2007) in a bottomland mixed hardwood forest that was developing toward a western hackberry-American and red elm-bitternut or pignut hickory climax. These first of these photographs was taken at same time as the three slides in the preceding set (six to eight weeks after last killing frost) whereas the second of these slides was taken sixteen to eighteen weeks into the warm-growing season (after last killing spring frost). Such paired temporal "photo-plots" showed extent and relative rate of revegetation in this forest. Hackberry (at several age/size classes) was the major tree species, hence the one to exploit to best early advantage increased quantities of light. American elm was the second most common tree, including the sheared off snag (far right) and the sapling with the broken-over-but-not-off trunk. The sapling had sprouted profusely right below the stress injury and also above and beyond this break (higher up on the shattered stem). The snag of the larger (pole-size) American elm had sprouted far below shear site close to base of the trunk. This larger American elm was "broke clean- off" by a falling limb of box elder. Part of the trunk of that box elder barely was shown in the first of these two photographs (upper right corner) but it was shown to good advantage the entire height of the second slide (right of American elm snag). Wood of box elder is extremely brittle and readily splits and "runs" for considerable length along cracks in its wood. Spicebush and fox grape were the major shrubs (those with most cover) in this forest gap. Silky wildrye (some Virginia wildrye), pokeberry, and purple Joe Pye weed were the major herbaceous species (in that approxomate order). Other forbs important elsewhere in this forest (eg. stinging nettle, slender nettle, and southern blue flag) were not present in this forest gap. Ottawa County, Oklahoma. may (vernal aspect) and July (estival aspect) for first and second slides, respectively. 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).

47. By the sheer weght of it (recovery efforts)- Early stage of recovery of vegetation in a mixed hardwood bottomland forest where two hackberry had been uprooted five months earlier (9 December, 2007) by a severe ice storm. Stage of plant development on this forest range as shown here was only six to eight weeks into the warm-growing season (following the last killing frost in late March or early April). Most of the remaining trees were hackberry (mature down to two-year-old seedlings) and American elm (two forememost saplings of two size classes). Major shrubs were spicebush and buckbrush with pawpaw forming an irregular mid-height shrub layer. The main herbaceous species was silky wildrye. The most common forb was pokeweed followed by purple Joe Pye weed. Ottawa County, Oklahoma. May (vernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). Mesic Bottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

48. Letting the light in- Old-growth white ash surrounding on limbs and general debris tron from box elder, American elm, and sycamore by an ice storm during the preceding late autumn (9 December, 2007). The ash was unscathed and being one of the tallest trees with a large crown it did not get as buch benefit of additional light and growing space as did other plants, especially those in lower shrub and herbacaeous layers. Some of the shrubs to benefit first from this new spot of denudation 9a forest gap) were grayback and fox grape. In addition, silky wildrye (dominant herbaceous plant), Virginia wildrye (general associate herbaceous species) along with perennial forbs including pokeberry or pokeweed, purple Joe Pye weed, tall nettle, and wood nettle plus pioneer plants like giant ragweed (Ambrosia trifida) also benefited from this forest gap or patch. It would be revealed in time shich tree and shrub species regenerated in this disturbance patch with most success. 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).

49. 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).

50. 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).

51. 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.

Moving upstream: An ephemeral stream that was a tributary of Modoc Creek entered immediately upstream from the bottomland (floodplain) forest that was just described. In summary that forest which was developing on the greater floodplain of Modoc Creek was progressing through processes of plant succession from a subclimax sycamore-eastern cottonwood- northern red oak-Shummard oak-dominated community to a climax western hackberry-American elm-white ash-bitternut hickory forest. Another tract ("branch" or "arm") of that bottomland forest had developed along the tributary of the "wet weather" drainage. This forest range community was contiguous with the one just presented. It bordered on the other side of the old fiield shown and described immediately above. The upstream bottomland forest range was slightly above the floodplain of Modoc Creek though, again, conterminous with it. Forest vegetation on the Modoc Creek tributary had developed on a less mesic forest site with shallower and, almost assuredly, less fertile soil. Many of the same plant species were common to both floodplain and tributory forest range; however, there were conspicuous and ecologically meaningful differences between the two tracts of bottomland forest including those in forest structure or architecture, age class structure of trees, density and cover of trees, and herbaceous layer (in addition to those in plant species composition). The most important and practical difference (at least from standpoint of this photographic presentation) was in light conditions, both intensity and duration. This was due to greater density of smaller and generally less mature trees in the tributary forest the result of which--from much of spring through to leaf fall during mid-autumn--in more complete and denser canopy cover. This vegetational structure excluded most light from the lower forest layers and permitted much shorter periods of light when photosynthetically active radiation could penetrate the upper forest cover. As such, photographing of this forest vegetation--especially the herbaceous layer-- and details of tree trunks could only or, at least, best be achieved during winter. Even then the angle of winter sunrise and sunset was so much farther to the south that there was never a period of time when direct light reached more northerly exposures. This resulted in dimlite appearances combined with shadows from tree trunks as the bane of this winter forest photographer. Viewers will have to suffer along with him at some shots.

52. Seral and climax- Second-growth mixed hardwood forest on tributary of Modoc Creek (western border of Ozark Plateau) immediately above creek's floodplain with western hackberry (the climax dominant) and American elm, slippery elm, pignut or bitternut hickory, mockernut hickory, and white ash (climax associates or, locally, co-dominants in various combinations with western hackberry) succeeding black walnut and honey locust. Chinquapin oak, boxelder, and red mulberry were also present at meaningful porportions of relative cover. The successional status of these latter three species was unclear. The major shrub was buckbrush or coralberry on land having less tree density and canopy cover and fox or frost grape where tree density and crown cover were greater.Common greenbriar, poison ivy, and trumpet creeper were present sporadically. Other than local habitats where buckbrush was dominant, the preeminent part of the understorey of this tributary forest was the herbaceous layer which consisted almost exclusively of Canada or nodding wildrye. This was strikingly different from the more mesic forest on the Modoc Creek floodplain which supported a more diverse understorey (see above) including an herbaceous layer dominated by silky and Virginia wildrye. The first of these two photographs was a second-growth bottomland forest dominated by pignut or bitternut hickory (two foremost trees in center to right foreground, the clump of three trunks in center background on bank of the ephemeral stream channel) and hackberry (largest tree that was also on stream bank [at left margin of midground], large sapling in front of the clump of pignut hickory in background). Both pignut hickory and western hackberry were climax species. There were also a few trees of mockernut hickory (Carya tomentosa= ) that were indistinguishable in background. There was regeneration of both Carya species represented by various age classes from large seedling through sapling and pole to mature tree. Dead shoots from last year and green shoots of the current cool-growing season of Canada or nodding wildrye (both visible in foreground) indicated the sole dominant of the herbaceous layer of the understorey. In the second photograph young hackberry (foremost pole-sized tree in left foreground; sapling in right midground; "sliver" of a trunk at left margin) were succeeding two log-sized, mature black walnut (diagonally opposite each other in midground) while white ash (three large trees in right background), pignut or bitternut hickory (foremost tree situated between the two walnuts) and boxelder (tree to left of the pignut hickory in far background) persisted (probably as climax) into this subclimax state forest. Canada wildrye was present (and generally conspicuous) as both last year's dead and current cool-growing seson shoots. Ottawa County, Oklahoma. Early January (hibernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). Mesic Bottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

53. Seral forest on a tributary of an Ozark creek- A second-growth mixed hardwood forest dominated by black walnut and honey locust had been successfully invaded by hackberry, slippery or red elm, and American or white elm which were succeeding the seral black walnut and honey locust. Fox grape (conspicuous speicmen climbing black walnut in right foreground of first photograph) and buckbrush or coralberry (inconspicuous in these photographs) were the major shrubs. Herbaceous layer was the predominant layer of this forest understorey. It was dominated almost exclusively by Canada or nodding wildrye. Winter annual grasses, especially Japanese chess (Bromus japonicus), was present, but stood no chance at widespread invasion given dominance by vigerous plants of the the robust native, Canada wildrye. Winter forbs were absent. Dead shoots of elephantfoot (Elephantopus carolinianus), the major warm-season forb, were present. Elephantfoot is a native perennial composite. In the first photograph the three large trees on the right bank of this ephemeral channel were black walnut. The sapling on the opposite (left) bank was an American or white elm. Large tree at right margin in midground, sapling appearing to left of this tree and right of the three walnuts, and the largest tree in this scene (orn right bank behind the black walnuts and to front of a rotting snag ) were western hackberry which is the number climax dominant and the species with greatest recruitment (= regeneration). A large boxelder was near center of channel in distant background. Dead shoots of a dense colony of Canada wildrye were visible on the left channel bank in midground. The second photograph presented a textbook example of forest succession with one seral stage replacing another and displayed dramatically by two seral species of aging trees being succeeded by two climax species with saplings. Black walnut, an Intolerant (Wenger, 1984, p. 2) seral species, represented by the two adult comparatively large trees in right foreground and one behind them, and honey locust, an Intolerant (Wenger, 1984, p. 3) seral species, represented by the two largest trees in center midground were being succeeded by western hackberry (foremost tree, a sapling, at far right; several small trees in badkground including the one behind the fox grape in center), American elm (sapling directly in front of second or center black walnut), and slippery or red elm (sapling behind foremost hackberry sapling and in front of foremost black walnut). Shoots, both dead and live, of Canada or nodding wildrye (the dominant herbaceous species) were conspicuous. Greater detail of this vegetation at local scale was presented in photograhs below. Ottawa County, Oklahoma. Early January (hibernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). Mesic Bottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

54. Being replaced (more-or-less) while othr stay- Vegetation dynamics in a bottomland mixed hardwood forest on an ephemeral drainage in the Ozark Plateau. Dramatic example of elm replacing in successional order black walnut while younger trees of black walnut along with older trees of white ash, chinquapin oak, and western hackberry remain. Through all this turnover among tree species Canada or nodding wildrye (a climax decreaser) remained the overwhelming dominant of the herbaceous layer which was the predominant layer of the forest understorey. The first photograph presented a broader view of a "photographic sample" of a local stand in which largest (and, by inference, oldest) trees were black walnut as represented here by the largest tree (left midground). There had been regeneration of the Intolerant (Wenger, 1984, p. 2) black walnut in the past as shown by trees of intermediate size such as foremost tree in immediate right foreground, but recent regeneration (recruitment) was of slippery or red elm as represented by the two saplings in front of big black walnut (left foreground), American or white elm, and western hackberry. Chinquapin oak (tree with light-colored bark to immediate right of big black walnut) and white ash such as the one opposite the chinquapin oak (right-center midground) persisted into what was developing into the climax forest vegetation. There were three large white ash in left background beyond the large black walnut. Big tree in center of distant background was an immense boxelder. Conspicuous shoots of Canada wildrye (several with clearly defined spikes such as those at base of big black walnut) proclaimed this species as the dominant of the herbaceous layer which was the most prominent part of the forest understorey. Second photograph was a "nested photoplot" within the larger "photosample" jsut described. The two saplings of red or slippery elm in front of the large, mature black walnut that was "escorted" by a small log-size chinquapin oak. Three nice mature (log-size) white ash as well as the white ash opposite the chinquapin oak represented this climax tree species. Small pole-size tree behind and to immediate of the white ash in right foreground was American elm. The very large boxelder in center of distant background was also still shown. Various plants of Canada wildrye, which was the herbaceous dominant, were obvious (due largely to the prominence of last year's dead shoots). These two photographs provided a textbook example of sucession and transition from one species to others of higher successional order. Black walnut, an Intolerant (Wenger, 1984, p, 2) and hence seral species (on this bottomland sere) was slowly being replaced through tree reproduction by red or slippery and American or white elm which, though somewhat variable, are generally Imtermediate in tolerance (Wenger, 1984, ps. 2-3). Likewise, white ash was also listed as listed by Wenger (1984, ps. 2-3) as Intermediate. For this forest cover type (SAF 93: ugarberry-American Elm- Green Ash) the Intermediate species comprise a variant form of the climax eolm-ash forest of Kuchler (1964, p. 101) and as recognized in Garrison et al. (1977, ps. 15-16). This forest range vegetation was in process of achieving species composition and structure of the terminal forest vegetation on this bottomland site . Ottawa County, Oklahoma. Early January (hibernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). Mesic Bottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

55. A lesson in plant succession- Close-up "photoplot" of the large black walnut and one of two slippery elm saplings introduced in the two immediately preceding photographs. This is "a picture worth a thousand words". Intolerant black walnut being replaced by Intermediate tolerance elm (Wenger, 1984, ps. 2-3) as elaborated in the preceding caption. No further comment deemed necessary here. Canada or nodding wildrye was presented in clariity and promience appropriate for its presence as the overall dominant of the understorey. This natiive, perennial, festucoid grass was sole dominant of the herbaceous layer which was the most consistent and prominent stratum of this forest's understorey. Ottawa County, Oklahoma. Early January (hibernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). Mesic Bottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

56. Another stand, a different seral dominant, the same successional story- Honey locust, as represented by two "really big" and senescing trees, was being replaced by younger trees of western hackberry and American and red elms on this sere of a bottomland forest site just above the floodplain of Modoc Creek near the western border of the Ozark Plateau. The first slide was at distance to give some idea as to physiogonomy as well showing internal structure and species composition of this form of a mesic bottomland forest in the ecotone between eastern deciduous forest and the vegetational mosaic of tallgrass prairie and tallgrass-hardwood savannah immediately to the west. Second slide presented sufficient detail for description of species composition, structure (arrangement) of forest vegetation, and progression of plant succession. The two largest trees (center foreground and standing diagonal to one another) were comparatively ancient (over mature) honey locust with dead and dying limbs (complete with woodpecker nesting cavities) in their crowns and just enough thorns to proclaim their species identy. These "long-past-their-prime" senescing trees still looked out on younger honey locust trees (just beyond camera range) proving that their species' moment in the "successional sun" (span of time on this sere) was not completely over, but it was clear that their floristic reign as local dominant was past as honey locust was being replaced by both American or white elm and slippery or red elm which were dsstined to share climax dominance with western hackberry, the overall forest dominant. American elm was represented by the largest tree and one small pole or large sapling to right-rear of the two honey locusts in both slides while slippery elm claimed the other large sapling or small pole by the big American elm in the first slide. Western hackberry was present as the young adult tree behind and immediate left of foremost honey locust (both slides), young adult tree to left of big American elm (second slide), and as small sapling (immediate left foreground of second slide). The aray of age classes of the Intermediate in tolerance elms and hackberry and Intolerant honey locust (Wenger, 1984, ps. 2-3) bespoke the inevitable replacement of honey locust by the hackberry-elm climax. This inexorable successional pathway, this progression along the sere, depends on absence of disturbance(s) severe enough that western hackberry and elm species are not halted in their advance so that honey locust (or species even more seral) reclaim their transitory throne. A plant of fox grape provided a continuous "thread" from lowest (ground) layer to highest layer (foret canopy) of this forest plant community. Here, as throughout the rest of this forest range vegetation, Canada or nodding wildrye dominated (essentially, exclusively so) the herbaceous layer which was the prominent one of the understorey. Buckbrush or coralberry was also present, but its leafless, short woody shoots (foreground of both slides as in front of foremost honey locust trunk) did not adequately proclaim it to be the most widespread shrub on this forest range. Ottawa County, Oklahoma. Early January (hibernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). Mesic Bottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

57. An opposite view of the stand just described- From the opposite end of the honey locust-dominated seral stand presented in the preceding two-slide set and caption the laina of fox or frost grape along with the short, woody shoots of buckbrush or coralberry (absent leaves and most fruit) presented the shrub layers of this bottomland forest range. Canada wildrye was present of course, but less conspicuously featured than in preceding slides. Successional replacement of Intolerant honey locust by Intermediate tolerance (and climax on this forest site) western hackberry American elm, and slippery elm was again the central theme of this photograph. Hackberry was represented by the large sapling or small pole to the forward and left of the big honey locust. Both Ulmus species were well-represented by saplings and small pole-sized trees to left of and behind the honey locust. There were also older, larger elm trees in distant background. All this forest vegetation was on the banks of an ephemeral stream that was a tributary to Modoc Creek, the floodplain forest vegetation of which was described in detail above. Ottawa County, Oklahoma. Early January (hibernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). Mesic Bottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

58. Calf pen trees- Numerous shoots of chinquapin oak that developed as stump (epicormic) sprouts from the previous tree on a tributary of Modoc Creek in the western Ozark Plateau. The first two photographs (side-by-side) are of the same set of young adult clones, the second (right slide) being a closer view than that shown in the first. The third photograph was of a second set of epicormic shoots from a second previous tree. Agent of defoliation that removed the preceding tree trunks was unknown. There were no remains of stumps or logs. Based on surmise and fragmentary history of land use it seemed most likely that the former trees had been cut for firewood, but this was not known with certainity. It remained irrelevant because the original (at least preceding trunks or shoots) of both chinquapin oaks had been topkilled and the two genetic individuals resprouted. Burns and Honkala (1990) cited work which concluded that chinquapin oak readily resprouts resulting in shoots that are usually younger than their root systems. Resprouts (resgrown shoots) from sawed stumps are common. Such epicormic or water shoots are clonal organisms. These coppices or clones are ramets of the original genet (the genotype or unique genetic individual). Such coppicing can result in shoots developing into trees that originated from trees of genetypes that appeared several "generations" (cycles of shoot cutting and regrowing) earlier. Burns and Honkala,(1990) also explained that chinquapin oak is shade-intolerant, but young trees are less so than older ones. This means that regeneration of chinquapin oak is disproportionately greater than than indicated by tolerance of established individuals. Hence, chinquapin oak can reproduce asexually from stumps and, by relying to some extxent on reserve food stored in roots, grow into mature trees under some degree--perhaps substantial--of shading. These same authors interpreted chinquapin oak as climax on soils derived from limestone parent material even when these are droughty natured soils. Even on mesic forest sites chinquapin oak is "subclimax to climax" and present as part of the climax forest vegetation (Burns and Honkala, 1990). Shoots originating from previous trunks (phenomenon of coppicing) can grow into mature trees as large or larger than the original or, at least., preceding shoots (trees). Guargantian clumps of resprouted coast redwood (Sequoia sempervirens) have clonal shoots which grow to such immense size that they are known reverently as "cathedral trees" (examples presented in the chapter, Coast Redwood Forests, under Woodlands and Forests). This Okie author has also been to the National Catheral in our Nation's Capital, but he feels more at home in the smaller Oakdale churchhouse. Having had experience with both beef and dairy cattle in the Ozark Region this photographer felt it appropriate to dub these clonal clumps of chinquapin oak "calf pen trees". Eat your evergreen, redwooded hearts out prune-pickers. Ottawa County, Oklahoma. Early January; dormancy phenological stage.

59. Stand of pignut or bitternut hickory (Carya cordiformis)- Local stand of pignut hickory on bank of an ephemeral stream that was a tributary into Modoc Creek in the western Ozark (Springfield) Plateau. All trees in photographs were pignut or bitternut hickory showing reproduction of this species as represented by various age classes. Downed branches resulted from an ice storm in late autumn of the previous year. Most plants in understorey were of buckbrush or coralberry and Canada wildrye. The interlacing (anastomising) network of bark yet with a rather smooth appearance, especially, on upper trunk and limbs was characteristic of Carya cordiformis. Positive identificaation of these trees (trunks) was easy and foolproof given plentiful specimens of lower, terminal buds which in C. cordiformis are a giveaway sulphur-color. This is the only species of hickory to have this coloration. Some specimens of mockernut hickory (Carya tomentosa) also grew on the banks of this ephemeral drainage. These trees were just beyond depth-of-field in these photographs, but they were presented later (below) in this section. These two Carya species are readily distinguished by features of winter twigs, especially buds, as was described immediately above as well as below. Ottawa County, Oklahoma. Late December; winter dormancy.

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

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

62. Bunch of bitter nuts- Nuts of bitternut or pignut hickory gathered beneath the trees (and their buds) introduced above growing on the bank of an ephemeral stream draining into Modoc Creek. Ottawa County, Oklahoma. Late December; ripe fruit stage, but they live up to their name (and the fact that humans would just as soon their hogs fattened on them).

63. Trunk of mockernut or white hickory (Carya tomentosa)- Specimen of mockernut growing on bank of an ephemeral stream that drained into Modoc Creek in western Ozzrk Plateau. This hickory species was much less abundant than pignut or bitternut hickory which was a minor dominant on a western hackberry-elm-white ash tract of forest that was part of a larger bottomland forest that included sycamore, eastern cottonwood, boxelder, chinquapin oak, and black walnut along with the more consistently present climax dominants. Bark of mockernut hickory is noticably smaller and more scalelike in general appearance than that of pignut or bitternut hickory with which it was closely associated in this bottomland forest. The only foolproof distinguishing difference, at least in late autumn through early spring prior to leafing-out are twigs. Both buds and lenticels differ drastically between mockernut and pignut hickories. Proceed to next two sets of slides and then compare back to pingnut twigs shown above. Ottawa County, Oklahoma. Late December; winter dormancy.

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

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

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

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

67. Bidding winter and successional farewell- Final view of species composition during early winter of a bottomland forest in the western Ozark Plateau in which western hackberry, the "number one" climax dominant tree species (two leftmost trunks: big trunk in foreground and smaller trunk to left and behind it) was successionally replacing black walnut (four diagonically aligned trunks from right foreground extending to left midground). Most of the smaller (indistinguishable) trees in background were American and slippery elm (return to previous slides in this series for details concerning these two Ulmus species). The most consistent understorey dominant was Canada wildrye. It was the only herbaceous species in the understorey of any consequence, Japanese brome or chess being present only as rare, isolated individuals. Buckbrush and fox or frost grape were the major shrubs, and of two sporadic layers. Canada wildrye, by constrast, was omnipresent comprising a nearly universal layer in this bottomland forest that developed along an ephemeral stream draining into Modoc Creek, an Ozark stream. Ottawa County, Oklahoma. Early January (hibernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). Mixed Hardwood Series, 223.1, in Southeastern Swamp and Riparian Forest biotic community Brown et al (1998, p. 43). Mesic Bottomland Forest (Nelson, 2005, ps. 147-150). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

68. Carrying the banner- Canada or nodding wildrye was--as shown and described throughout this section--the dominant and basically the sole species of the herbaceous layer (the predominant stratum of the understorey)--of a second-growth bottomland forest currently dominated by black walnut, honey locust, and white ash but progressing along its sere to a western hackberry-elm (American and slippery)-white ash climax. Canada wildrye was present as both dormant shoots from last year and new (and very young) shoots of the current cool-growing season. Livestock were never present on this forest range, but white-tail deer as well as cottontail rabbits were abundant throughout and at all seasons. Squirrels, coons, and 'possums also called this tributary forest home, but of course fed only on fruits and buds. One of the interesting differences between this bottomland forest range and the floodplain forest range (treated above), with which it was contiguous and immediately unstream from, was presence as local dominants and, sometimes, associates of silky and Virginia wildrye (Elymus villosa, E. virginica) downstream on deeper alluvial soil versus nodding wildrye (E. canadensis) on higher, shallower, less mesic soil. Also, the herbaceous layer (s) of the floodplain forest was (were) much more diverse in species and structure (see again above). Canada wildrye was presented at level (the photographic scale) of local population in these two slides and at levels of individual plant and individual tiller immediately below. Ottawa County, Oklahoma. Late December; all (at least, most) basal leaves stage (four to seven-inch length).

69. A real cool grass- The cool-season, perennial, native grass, Canada or nodding wildrye, was the dominant--indeed, sole species of any consequence--in the herbaceous layer of a bottomland forest on a tributary of Modoc Creek in the Ozark Plateau. Canada wildrye was present in almost every forest community photograph and description presented in this section (above). This species was presented "up close" at scale or level of individual, cespitose plant in these two slides. Canada wildrye was shown at the organizational level (scale) of local population in the immediately preceding set of slides and at level of individual tiller in the succeeding two-slide set. Both current, cool-season shoots and dead, last year's shoots were present on this forest range. Ottawa County, Oklahoma. Late December; all (at least, most) basal leaves stage (four to seven- inch length).

70. New growth in winter- Tillers of Canada or nodding wildrye in early winter in the herbaceous layer of a bottomland forest (ephemeral drainage into Modoc Creek, western Ozark Plateau) beneath black walnut and honey locust (Intolerant species) with western hackberry, American elm, and slippery elm in process of achieving domiance. These two slides presented Canada wildrye at photographic scale (plant organizational level) of individual shoot. Scales (organizational levels) of local population and individual plant were presented immediately above as two two-slide sets. Canada wildrye is a strictly cespitose (bunchgrass) species meaning that the only shoots produced by this grass are tillers. These vertical (intravaginated) shoots do, however, grow in a somewhat decumbant form just below the soil surface as they grow away from the base of last year's shoots. That morphology was presented in these two photographs. Both current, cool-season shoots and dead, last year's shoots were present on this forest range. Ottawa County, Oklahoma. Late December; all (at least, most) basal leaves stage (four to seven-inch length).

71. 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.

72. Comparison of two major elm species- Distinction between American or, sometimes, white elm and slippery or red elm-- especially in young trees--is difficult (and not always correct even by "experts"). This can be a perplexing and frustrating problem in forests where both species are found (often trunk-by-trunk) such as on the bottomland forest discussed here and the sugar maple-bitternut hickory foresst covered previously. There is one brief period, however, when identification and separation of these two species is "a chinch". When these two species are in fruit (for ever so brief a time) they can be distinguished instantly by the distinctively different features of their samaras. The first of these two photographs was a comparison of twigs heavily loaded with samars. In this first slide the twig of red or slippery elm was on top while that of American or white elm was on the bottom. These were typical spring twigs with the classic differences between these two species quite evident. The samaras of slippery elm are noticeably larger and the wings are "wavy" or "crinkled". By contrast samaras of American elm are smaller with more-less "flat" wings. The buds of red or slippery elm are covered with orangish-red (dull rust-colored) short hairs and at time of opening these reddish buds are much smaller (often less than half the size) of American elm which often have an elongated appearance (visible here). The second photograph was a comparison of samaras of these two species. The smaller samara on the left was American elm and the larger samara with the "wavy" wing on the right was that of slippery or red elm. Ottawa County, Oklahoma. March.

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

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

75. Woody climber in the bottoms- Fox, frost, or chicken grape (Vitis vulpina) in the western hackberry-elm (mostly American; some slippery)-white ash- boxelder (with persistent sycamore and eastern cottonwood) bottomland forest in the floodplain of Modoc Creek. This liana was brought to ground by the ice storm treated above which facilitated its visual reproduction here. This grape was growing in the forest gap described previously in various stages of growth and recovery Terminal portions of this woody shoot subsequently began the plant's ascent back into the forest canopy (but not before it left a contribution to educational of the next generation of rangemen, foresters, and wildlifers. Frost or fox (the literaal interpretation of vulpina) grape is one of the more common Vitis species in the Ozark Plateau as well as on this particular bottonlamd forest in the Springfield Plateau portion of the Ozark Region. Floodplain of Modoc Creek, Ottawa County, Oklahoma. May; peak bloom phenological stage.

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

77. 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..

78. 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..

79. 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.

80. 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.

81. 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.

82. 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.

83. 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.

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.

84. 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.

85. 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.

86. 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.

87. 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.

88. 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.

89. 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.

90. 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.

91. 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.

92.. 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.

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

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

93. White ash (Fraxinus americana)-sycamore (Platanus occidentalis) wet bottomland forest- A natural spring in this bottomland at the western edge of the Ozark Plateau supported a unique combination of mesophytic and hydrophytic species. The dominant tree was white ash based on both relative forest canopy cover, number of mature trees, and regeneration of young trees. Sycamore was conspicuous by the white, exfoliating bark of large tree trunks, but there were no trees of immature age classes (ie. no reproduction). From perspective of tree size and apparent age, sycamore was interpreted as semi- co-dominant to ash (an ecological-successional status between that of a dominant and an associate species). Black oak (Quercus velutina) and northern red oak (Q. rubra) were obvious associate (tree with retained dead leaves in right background); hackberry (Celtis occidentalis), boxelder (Acer negundo), and red or slippery elm (Ulmus rubra) were also present in several age classes with regeneration of hackberry and boxelder prouonced. Larger elms had succomed to Dutch elm disease (Ceratocystis ulmi). Black walnut (Juglans nigra) and chinkapin oak (Q. muehlenbergii) grew on better drained soils at outer edges of this community. The lower shrub layer was dominated by buckbrush or coralberry which occurred throughout the understorey. Pawpaw and gray dogwood (Cornus racemosa) grew at restricted, local scale. Woody climbers, which grew abundantly on trunks and high into the crowns of almost all the larger trees, consisted of trumpet creeper (Campsis radicans), Virginia creeper, and poison ivy. Herbaceous species were covered in succeeding photographs. The green patches of understorey visible in this slide were composed of two sedge species (Carex lurida and C. lupulina). The author observed use of this vegetation-- specifically that immediately adjacent to the spring-- in autumn by migrating woodcock or timber doodle (Philohela minor). There were many probe holes (earthworms were most likely the sought prey) in the mud of this habitat. All-in-all, a most unique combination of biotic range community. Ottawa County, Oklahoma. Hiebernal aspect, December. One of numerous forms or variants of wet bottomland forest based on classification of natural communities by Missouri Natural Areas Committee (1987). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). Combination of SAF 93 (Sugarberry-American elm-Green Ash ) and SAF 94 (Sycamore-Sweetgum-American Elm). Too small for Kuchler units. Mixed Hardwood Series in Southeastern Swamp and Riparian Forest biotic community of Brown et al. (1998). Wet Bottomland Forest (Nelson, 2005, ps.153-157). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

94. White ash-sycamore wet bottomland forest in summer verdure- Same vegetation as seen in winter aspect in the preceding slide. The dominant herb averaged over entire herbaceous layer was cyber or lurid sedge (Carex lurida). It was frequently "accompanied" by hop sedge (C. lupulina). The dominant forb in the vernal aspect was wake robbin (Trillium sessile); dominant forb in estival aspect was spotted jewelweed (Impatiens capensis) and halbertleaf mallow (Hibiscus militaris). Other forbs present were cardinal flower (Lobelia cardinalis) and swamp milkweed (Asclepias incarnata). Dominant grass in and along the spring was rice cutgrass (Leersia oryzoides) while woodreed grass (see slides above) occurred farther from the aquatic habitat and scattered among the trees. The floating green plant on the water surface was lesser duckweed (Lemna minor). Ottawa County, Oklahoma. Estival aspect, July. Designation of wet bottomland forest was determined from classification by the Missouri Natural Areas Committee (1987). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). Combination of SAF 93 (Sugarberry-American Elm-Green Ash) and SAF 94 (Sycamore-Sweetgum-American Elm). No Kuchler unit at this small scale. Mixed Hardwood Series in Southeastern Swamp and Riparian Forest biotic community of Brown et al. (1998). Wet Bottomland Forest (Nelson, 2005, ps.153-157). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

95. White ash-sycamore wet bottomland forest- The rice cutgrass and sedge understorey (in and immediately adjacent to the spring feeding this forest range community) was shown here at peak standing crop. The saplings were all green ash. Regeneration of green ash established this as the dominant species of this vegetation. Green ash was rated as Intermediate in tolerance and Moderate in flood tolerance; sycamore had these same ratings (Wenger, 1984, ps. 3, 7). Pawpaw is visible in far left foreground. Trunk of mature tree is green ash with 32 inch DBH (shown immediately below). Ottawa County, Oklahoma. Estival aspect, July. FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). Combination SAF 93 (Sugarberry-American Elm-Green Ash) X SAF 94 (Sycamore-Sweetgum-American Elm). Mixed Hardwood Series in Southeastern Swamp and Riparian Forest biotic community of Brown et al. (1998).Wet Bottomland Forest (Nelson, 2005, ps.153-157). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

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

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

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

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

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

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

101. ce ain't nice- Severe damage from an ice storm to trees in a green ash-sycamore wet bottomland forest in the Ozark Plateau. A late autumn precipitation event resulted when warm, moist air from the Gulf of Mexico rose over cold, dry Arctic air (a Norther) resulting in the "winter mix" of, in order, rain, freezing rain, sleet, and snow that accumulated as ice to depths of one to four inches. The heavy accumulations of ice added so much additional weight to trees, power lines, roofs, etc. that numerous of these objects crashed to the ground. Such precipitation events, known as ice storms, can cause some of the most incredible damage in Nature to trees. Locally, extreme damage to trees and shrubs may rival that from tornadoes and hurricanes. Area of impact is frequently larger than that from tornadoes. The ice storm responsible for the damage shown here and in subsequent slides cut a swath about three hundred miles long and a one hundred and fifty miles wide. The wet bottomlad forest shown here was the green ash-sycamore community introduced above that developed around a natural spring. Most of the downed crowns and individual limbs at this site were from sycamore, the locally dominant tree, but Shummard oak, black oak, and green ash also suffered major damage. Ottawa County, Oklahoma. Hiebernal aspect, December. One of numerous forms or variants of wet bottomland forest based on classification of natural communities by Missouri Natural Areas Committee (1987). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). Combination of SAF 93 (Sugarberry-American elm-Green Ash ) and SAF 94 (Sycamore-Sweetgum-American Elm). Too small for Kuchler units. Mixed Hardwood Series in Southeastern Swamp and Riparian Forest biotic community of Brown et al. (1998). Wet Bottomland Forest (Nelson, 2005, ps.153-157). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

102. The double-edged sword of water- A hollow and nearly-dead-but-still-standing sycamore was toppled by heavy ice accumulations resulting from late autumn ice storm caused by the atmospheric phenomenon of overriding (warm, moist air rising above cold, dry, heavy air resulting in mixed precipitation of rain, freezing rain, sleet, and snow). This old sycamore and its cohorts, including its side-by-side "sister" tree, became established and survived as facultative phreatophytes along a natural spring in the western edge of the Ozark (= Springfield) Plateau. The ice-downed sycamore was the left sycamore (leftmost, conspicuous, white-barked trunk in left midground) shown in the first slide above that introduced this local green ash-sycamore wet bottomland forest. Ponded water from the spring was visible in the left midground of the first of these two slides. Ironically it was liquid water that permitted establishment and continued survival of sycamore and it was solid (frozen water in form of ice) that finally ended the life of the old, toppled sycamore. Details of the hollow trunk and its downfall were covered in the immediately following set of photographs and caption. From perspective of plant succession and development of this wet bottomland forest the most obvious change in this forest community was the on-going replacement of sycamore by western hackberry. The characteristic netted or deeply furrowed bark of the latter species was distinctive on the trunk of the pole-size tree in left foreground of the first photograph and in the three centermost saplings (included the leftmost one on which the sycamore had fallen and bent). The sapling on the right of the first slide and in left midground of second slide was a sapling of Shummard oak (the same sapling in both photographs incidentially). These species dynamics were already in play several years before the old sycamore fell. They will certainly be accelerated thereafter. This pattern of species replacement (the succession of plant species) was taking place at the edge of the wet bottomland forest. In closer proximity to the pool of spring water sycamore and green ash were still the species having greatest recruitment and, thus, were an edaphic climax (in polyclimax theory), postclimax (in monoclimax theory), or a climax that developed along a gradient in which soil moisture was determinative (in climax pattern theory). The lack of sycamore reproduction and, instead, reproduction of western hackberry was consistent with changes in the hackberry-American elm-bitternut hickory floodplain forest along Modoc Creek described in detail above. These two forest range communities were about 350 yards apart. Ottawa County, Oklahoma. Hiebernal aspect, December. One of numerous forms or variants of wet bottomland forest based on classification of natural communities by Missouri Natural Areas Committee (1987). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). Combination of SAF 93 (Sugarberry-American elm-Green Ash ) and SAF 94 (Sycamore-Sweetgum-American Elm). Too small for Kuchler units. Mixed Hardwood Series in Southeastern Swamp and Riparian Forest biotic community of Brown et al. (1998). Wet Bottomland Forest (Nelson, 2005, ps.153-157). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

103. Hollow-grounded- Details of the hollow and almost-dead-though-still-standing sycamore shown above that was felled by heavy accumulation of ice in late autumn ice storm. Hollow trees (trees with hollow trunks), contrary to the erroneous popular view, are usually about as strong as trees with solid trunks. Not quite, but close. This is the same basic physics as the fact that under certain kinds of strain a steel or iron pipe is quite strong compared to a solid piece of steel or iron of the same metal and diameter. This is why shafts of bird feathers, the hollow culms of many grasses (think bamboo), and hollow airplane wings (used as fuel tanks with different degrees of fullness) are very strong, especially when compared to their weight. In the same manner hollow trees can stand much strain and stress. In the case of this sycamore the added weight of accumulated ice (about two inches) and accompanied with gusty winds was more than the trunk that was 9/10ths or more hollow could bear up under. In this same ice storm there were large, completely intact trees (including sycamore, American elm, and hackberry) that also crashed. It was no wonder that this over-ripe oldster ended its life cycle thusly. There was a small, but very deep fire scar on the base of the fallen sycamore that appeared to the point of entry of decay agents (fungal mycelium were present) that permitted decomposition of the dead wood (heartwood) inside the trunk. The lower end of this fire scar was visible in the second of these photographs (at extreme left margin of the photograph about 3/8ths of the vertical distance from bottom of slide). Referral to any freshman Botany text will remind viewers that the layers tissue of the tree trunk moving from interior to exterior were pith (very little of that); xylem of heartwood; followed consecutively by living xylem, cambium, and live phloem; and, finally, bark. It was the heartwood, which in a tree of this age comprised vast bulk of the trunk, that was susceptible to decay by microorganisms and fungii. In fact, in this tree there was remarkably little sapwood and bark to keep this old fellow upright. It was a testament to the strength of hollow trees that this more-dead-than-alive, rotten sycamore stood as long as it did and that it took such extreme stress to bring it down. Even down this sycamore remained a critical component in the forest range ecosystem of which it was part for decades. As this trunk continues to rot it will release nutrients stored in wood to enter the biogeochemical cycles of carbon, nitrogen, etc. It hollow will undoubtedly serve as a den for coon, possum, feral house cat, etc. Ottawa County, Oklahoma. December.

104. Sycamore (Platanus occidentalis)- Distinctive bark, branching pattern, and habit of sycamore. This massive old-growth specimen of sycamore was growing along the flood plain of Modoc Creek (Ottawa County, Oklahoma) and was over 7 feet DBH (where the branch-like sprout originated). Authorities regard the sycamore as one of the hardwood (angiosperm) species that attains largest mature size (especially in diameter) in North America. Sycamore is interpreted as a pioneer species that persist into the climax forest, often as ancient behemenths like this splendon specimen. Such trees often live for decades as a member of climax riparian vegetation where they function as facultative phreatophytes. Even at some distance from streams sycamores like the one presented here get a continuous water supply from the water course by lateral flow. April.

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

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

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

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

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

110. Part of sycamore's secret- In contrast to detrminate growth of many other--probably most-- hardwoods (eg. oaks, hickories, walnut) and conifers with preformed buds at shoot (leader) tips, sycamore and some other angiosperms like sweetgum and willows have indeterminate growth. Whereas extension of the branch in determinate growth is in one seasonlong bout of growth (at least as growing conditions are favorable and at rates consistent with growth favorability) indeterminate growth occurs in phases or "flushes". These flushed are obvious when new leaves appear periodically throughout course of the growing season Examples of new leaves at tips of sycamore can often be seen throughout summer months. The three photographs provided examples of such recurrent or flushes of growth at ends of lower branches in sycamore growing on banks of Modoc Creek, Ottawa County, Oklahoma. June.

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

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

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

114. Colony of sedge in understorey of gren ash-sycamore wet bottomland forest- Hop sedge is the species most abundant here but there were some "strays" of cyber sedge present. Ottawa County, Oklahoma. April.

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

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

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

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

119. Halbertleaf rose mallow and cardinal flower as local dominant forbs in green ash-sycamore wet bottomland forest- This spring-fed unique form of the eastern deciduous forest (combination SAF 93 X SAF 94 forest cover types) was botanically diverse in both species and structure. The herbaceous layer was a mosaic of microsites composed here-and-there variously of tall, robust forbs; hydrophytic sedges; or grasses of varying water requirements. In this view two of the most mesic (and strinkingly photogenic) forbs native to the deciduous forest of eastern North America grew as pals in the rich mud of a bottomland forest. This forest community was at the extreme western edge of the Ozark Plateau where it contacts the Cherokee Prairie of the Central Lowlands physiographic province. Forbs like cardinal flower are more typical of "wild flowers" found in the Applachians or Catskills. Wetland vegetation was an interesting assemblage of species usually not growing together. They also occur more commonly on habitats different from the environment seen here. Ottawa County, Oklahoma. Late estival-early autumnal aspect, September.

120. Cardinal flower (Lobelia cardinalis)- This large specimen (over a yard in height) was growing at the bank edge of the spring that was the basis for the wet bottomland eastern hardwood forest presented in this series of photographs. Cyber and hop sedge were visible in the background as was the trunk of the large green ash shown previously. Ottawa County, Oklahoma. Late estival-early autumnal aspect, September.

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

122. Halbertleaf rose mallow (Hibiscus militaris)- The flowering shoot of this native forb portrayed the staminal column that immediately marked it a member of the Malvaceae (mallow family). This remarkable family includes not only the greenhouse or indoor Hibiscus species but such yard beauties as Rose of Sharon (H. syriacus) and hollyhock (Althaea rosea) as well as okra or gumbo (Hibiscus esulentus), the beloved southern garden vegetable, and the world's most important fiber crop, the Gossypium species. Those familar with King Cotton will immediately note the close resemblance of the unopened flower buds on the rose mallow shown here with the cotton "square". The descriptive name of halbertleaf is in reference to the halbert, a combination spear or pike and battle-axe used by soliders in the Fifteenth and Sixteenth Centuries. Hence, the Latin militaris. This high seed-yielding prairie and forest forb should be a favorite for those landscaping with native plants in humid regions. Ottawa County, Oklahoma. August.

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

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

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

Bottomland Forest- Example on an Ephemeral Stream in Ozark Plateau

The following example showed a forest co-dominated with both hackberry or western hackberry (Celtis occidentalis) and sugarberry (C. laevigata) along with American elm (Ulmus americana) with an understory almost exclusively of broad-leaf wood oats (Uniola latifolia= Chasmanthium latifolium). Details of the two Celtis species and broad-leaf wood oats was provided. Part of this forest had suffered severe damage due to windthrow by straight-line winds. Uprooting of trees by combined action of wind and saturated soil from recent heavy rains furnished an example of secondary plant succession in the "tree cradles" formed by tree blowdown (uprooting).

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

127. Side-by-side seedlings- Hackberry or western hackberry and sugarberry frequently grow in close proximity to each other. This complicates identification in the field, especially when there is only one species present and even more so with large trees whose leaves are considerable distance from the observer. When Celtis occidentalis and C. laevigata grow next to each other leaves can be compared and differences between the two species are more obvious. Such was the instance presented here and in subsequent slides. Larger (two- to three-year-old) seedlings of hackberry and sugarberry germinated and continued to grow side-by-side on a tree cradle created when several adult trees were uprooted from saturated soil by straight line winds. In an example on one of those cradles in the Ozark Plateau a study in species was presented with a seedlings of hackberry on the left and one of sugarberry on the right. Ottawa County, Oklahoma. July.

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

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

130. Leafy samples #1- Examples of leaves of hackberry (Celtis occidentalis). From the crade tree specimen introduced above. Ottawa County, Oklahoma. July.

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

132. Leafy samples #2- Examples of leaves of hackberry (Celtis laevigata). From the cradle tree specimen introduced above. Ottawa County, Oklahoma. July.

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

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

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

Windthrow or blowdown in a lowland forest of hackberry or western hackberry, sugarberry or southern hackberry, honey locust, chinquapin oak, and American or white elm provided an opportunity to study secondary plant succession at local scale through the phenomenon of forest gap or patch dynamics. Progression of revegetation, vegetation development by secondary succession, on a sere of climax lowland deciduous forest, SAF Cover Type 93 (Sugarberry-American Elm-Green Ash), in the western edge of the Ozark (Springfield) Plateau was preented in the following section of this chapter. Straight line winds of over 70 (probably closer to 80) miles per hour on Easter Sunday (15 April 2001) in conjuction with wet--though not saturated--soil resulted in uprooting of numerous mature trees of the species just listed. This blowdown occurred along the perimeter of a climax second-growth forest on slopes just above an intermittant stream. Deeper inside the forest there was minimal windthrow. This was a typical pattern resulting from the wind blockage affect of outermost trees on edges of the forest community. A series of sequential photographs over a period of years following blowdown of trees on the perimeter of this lowland hackberry-sugarberry-elm forest documented early recovery of forest range vegetation in the cavities and immediately surrounding areas of denudation caused by high winds and wet soil conditions. Such forest gaps with large holes or craters left by uprooting of mature trees provided initial bare ground for the process of plant community development (forest revegetation by secondary plant succcession) at local scale, known as patch dynamics. Foresters affectionately (by standards of stoic woodsmen) refer to such soil cavities as "tree cradles" due to the frequent germination and subsequent establishment of trees in the hole and/or surrounding area of tree fall. Cavities thus created provide bare soil and more mesic land (edaphic-microtopographic) habitat for tree seedlings or root suckers. As the following photographs showed development of vegetation (secondary plant succession) pm disturbance microsites caused by uprooting and related creation of bare soil (wind as "Nature's bulldozer") is not simply a matter of tree seedling pioneering the denuded microhabitat. Yes, this is sometimes the case as for example when seedlings of sycamore, eastern cottonwood, willow, sweetgum, and other hardwoods sprout up almost immediately on newly cut stream bnks, sand bars, roadcuts, or clearcuts. Douglas-fir (Pseudotsuga menziesii) and coast redwood (Sequoia sempervirens) are examples of western conifers that typically have the same pioneering or early seral stage establishment following creation of bare soil environments by disturbance. Coast redwood also sends up sprouts (vegetative reproduction). Classic studies of secondary succession on old fields or go-back land have shown unequivocally that such rapid establishment of late seral tree species (some persisting into climax) and/or climax tree species is not the typical pattern of vegetation development on the seres of many range and forest sites. The photographic study presented below showed the same phenomenon, the same pattern, of old field succession commencing with annual, pioneer ("weedy") species which were then followed in succession by short-lived, perennial, herbaceous species over course of several growing seasons before seedlings of tree species vegetated the "tree cradle". There was a "successional twist", however, as some root and stump sprouts from unearthed trees quickly emerged and grew alongside the herbaceous pioneers that sprouted from the soil seed bank and quickly populated "tree cradles" . Local microsites of denudation caused by uprooted trees were designated as "tree clods" (for the mass of soil, rock, tree roots) and their "holes" or "craters" (cavities left in the land surface by removal of "tree clods") were followed over spans of time beginning with initial plant communities in the first growing season starting after tree uprooting. A single "tree clod" typically consisted of two or more mature trees that had been uprooted together as a unit. A few "tree clods" were made up of a single large tree whereas other "tree clods" and their "holes" were made up of as many as five trees that were uprooted en mass. In this photographic treatment two examples of uprooted tree units ("tree clods") caused by the Easter Sunday blowdown were presented to show recovery of plants on these local spots or patches of disturbance. The first of these examples was of one large "tree clod" consisting of five individual mature tress. The second example included two somewhat smaller "tree clods" each of which was comprised of a single mature tree and adjacent immature trees (saplings and/or poles) These two examples of "tree clods" and their "root holes" was followed over a sequence of years to provide photographic documentation of secondary plant succession on (in and around) each denuded microsite (forest gap).

136. Broad view of windthrow- Uprooted trees (straight line winds of 70-80mph combined with wet soil) viewed as "tree clods" at outermost edge of a lowland forest of hackberry, sugarberry, hony locust, chinquapin oak, and American elm. A total of four "tree clods" and "craters" created by uplifted earth were visible in this wide-angle photograph. The first of these "tree clods" was caused by uprooting of a large chinquapin oak which broke off crowns of hackberry and two honey locust. The second (and largest) "clod" was created by uprooting of a large hackberry and very large black walnut which coame up together as a unit and which brought down a sugarberry, American elm and honey locust. The third "tree clod" consisted of a large honey locust, two young American elm, and two young (pole-size) hackberry or sugarberry (the two could not be distinguished without leaves). The fourth "tree clod" was comprised of a massive honey locust that brought down (uprooted with it) two smaller hackberry or sugarberry. "Tree clods" consisted of both: 1) direct victims or causalities (larger, mature trees initially uprooted through storm strain, combination of straight line winds and wet soil) and 2) indirect or secondary victims or causalities (smaller trees that survived storm stress of wind and wet soil but were brought down, and their roots partially up, by crashing larger trees uprooted by storm action). The large trees that were toppled by direct results of storm strain (high winds and wet soil) were almost entirely uprooted or else the major roots had been broken in the crash. As such, the main shoot of all large trees (chinquapin oak, honey locust, black walnut, sugarberry, and hackberry) had died by late spring. Some of these large trees sent up root suckers so that the actual genetic tree did not die completely and instead persisted as clonal offspring (asexual or vegetative reproduction), but these were relatively weak and lacking in vigor. In subsequent growing seasons some of these sprouts died. Some of the smaller trees that were brought down by uprooted windthrown trees were not completely uprooted with the result that they leaved albeit in the horizontal plane. The second and massive "clod" (directly uprooted large hackberry and even bigger black walnut with secondary causalities of Ameican elm, sugarberry or hackberry [whichever], and honey locust) served as one example described below. The third "clod" (directly uprooted honey locust and secondary causalities of elm and hackberry or sugarberry) and fourth "tree clod" (directly uprooted huge honey locust and two young or pole-sized hackberry as secondary causalities) were in close porximity and together served as the second example to be described. Photographs were taken periodically over a span of years in either late June or early July to present late spring or early summer aspects of successional range vegetation. Given that denuded microhabitats were created in mid April, these photographs represented the number of growing seasons (mid-way through) since denudation. Ottawa County, Oklahoma. June, 2001.

137. First growing season of first blowdown example- "Tree clod" and "crater", the second of four such forest gaps of "clods and craters" presented in the preceding photograph, caused when a very large black walnut and large hackberry were uprooted by straight line winds and wet soil. Plant species growing on this denuded microenvironment included such pioneers as giant ragweed (Ambrosia trifida) and hairy crabgrass (Digitaria sanguinalis) as well as surviving, long-established perennials such as silky wildrye (Elymus villosus). There was some mare'stail or horseweed (Conyza canadensis= Erigerion cnaadensis), but much less of this species than of giant ragweed (which was somewht surprising given that these two annual composites are frequently co-dominant in early years of old-field succession). There were also abundant root sprouts from the uprooted trees, but these were generally lacking in vigor so that most were dead by the second growing season. Most plants of poison ivy (Rhus toxicodendron= Toxicodendron radicans) and fox or frost grape (Vitis vulpina) survived blowdown and even benefitted from increased light. Ottawa County, Oklahoma. June, 2001.

138. Fifth growing season of first blowdown example- Successional vegetation in fifth growing season on a "tree clod and crater" caused by uprooting of a very large black walnut and large hackberry. Species of pioneer annuals--including giant ragweed, marestail or horseweed, and hairy crabgrass--were still common and dominant in bottom and sides of craters as in first growing season following windthrow. There was abundant cover of the Eurasian annual grass, Japanese chess or brome (Bromus japonicus) which had many straw-colored, dead or dying shoots at time of photograph (July). This grass had not been a prominent component in the first warm-growing season following blowdown (15 April, 2001) because it is a cool season species. The most abundant grass was silky wildrye (several spikes visible in foreground of second slide), the native, cool-seaaon perennial decreaser which had been present in understorey of the climax forest and survived on "tree clods" even in the first growing season following uprooting of trees. Another relative common and abundant annual that had been present since the first cool-growing season following denudation was the naturalized Eurasian forb, hedge parsley (Torilis japonica). Seedlings of American elm, hackberry, and sugarberry were now well-established. Sprouts of honey locust origining from roots of huge, uprooted trees were obviously larger. An example of one of these vigerous honey locust root sprouts was in lower right corner of first photograph. Note: it was very difficult to determine whether the young trees of hackberry, sugarberry, and elm were from seed or root sprouts, but based on plants in the first couple of growing seasons following blowdown it was determined that they were, in fact, seedlings and not resprouts off of roots of downed adult trees (ie. sexual not asexual reproduction had been responsible for regeneration of climax tree species. It was the opposite situation for honey locust in which regenerated trees were root sprouts of huge uprooted trees. These climax tree species did not dominate the denuded site on basis of density, cover, biomass, etc., but they were becoming more abundant. The biennial (and native) tall thistle (Cirsium altissimum) had become locally common (eg. lower left foreground of second photograph). Also, perennial forbs had begun to occupy the denuded site. This included an unidentified (not yet flowering) goldenrod (Solidago sp.) like the plant in the right corner of the second photograph. A common (and perennial) forb was climbing false buckwheat (Polygonum scandens). This species was conspicuous in the foregroung of both photographs, especially the second. Ottawa County, Oklahoma. July, 2005.

139. Eighth growing season of first blowdown example- Recovering vegetation in eighth growing season on a "tree clod and crater" caused by uprooting of a very large black walnut and large hackberry. Almost all foliar cover was that from seedlings of the climax tree species, the adults of which had been uprooted eight growing seasons ago. The two dominant species of seedlings were hackberry and sugarberry (with leaves clearly distinguishable and distinct). Seedlings of American elm were smaller and less common making this the associate species. In spite of vigerous root sprouts (resprouts) early on in the denuded areas, honey locust had become a distant "runner-up" among the climax tree species. Almost all forbs were gone from the sere at this stage of secondary plant succession. There were some scattered individuals of wingstem (Verbesina helianthoides). There were also individuals of silky wildrye which had survived windthrow. These were being overshadowed by seedlings and sprouts of climax tree species, but wildrye plants survived because they are a cool-season perennial species and make much of their growth during winter and early spring when hardwood tree species are bare (devoid of leaves). Ottawa County, Oklahoma. July, 2008.

140. Detail in eighth growing season of first blowdown example- Successional stage of developing range vegetation on a denuded microsite, a "tree clod and crater", caused by uprooting of a very large black walnut and large hackberry. This was appearance and composition of the successional plant community about midway through the eighth warm-growing season following blowdown. Almost all plant cover was that of seedlings and sprouts of the climax tree species of which hackberry and sugarberry were the dominants (followed by American elm and, as in distant fourth-place, honey locust). Note that the species composition--not structure or physiogonomy--was now that of the original climax hackberry-sugarberry-elm-honey locust lowland forest. In only eight years following denudation by windthrow, and old-field succession that began with weedy annuals like giant ragweed, marestail or horseweed, and hairy crabgrass, the original species composition of the forest had been restored. The rest of time taken for recovery of this natural forest range vegstoration on the sere will be maturing of established individual trees of the climax dominant species. In descriptive vegetational analysis (at least to this point on the revegetating sere), succession of patch dynamics created by tree blowdown was amazingly rapid with restoration of dominance by climax tree species completed by the large seeding stage (age class) of tree growth, and in eight years following initial denudation. Ottawa County, Oklahoma. July, 2008.

141. First growing season of second blowdown example- Appearance of two "tree clods" and "craters", the third and fourth of four such forest gaps of "clods and craters" that were presented in the introductory wide-angle photograph given above, about midway through the first warm-growing season after being caused by tree blowdown. The third "clod and crater" consisted of a large honey locust that was uprooted and took with it as it crashed two young American elm and two young (pole-size) hackberry or sugarberry (these two species could not be distinguished without leaves). The fourth 'clod and crater" was formed when an uprooted huge honey locust crashed taking with it to earth two young or pole-sized hackberry. The most conspicuous species to show benefit from denudation was giant ragweed and, later in the summer, hairy crabgrass. There was some horseweed or marestail, but much less than giant ragweed. As was the case for the first "tree clod and crater" example, liana species like fox grape and poison ivy survived and beneffited (at least initially) from increased sunlight. There were root sprouts from the uprooted, large, adult trees. These were generally weak and lacked vigor typical of sprouts (especially, it seems, those that are brush) except for those of honey locust. Ottawa County, Oklahoma. June 2001.

142. First growing season (close-in view) of second blowdown example- Detailed view of the "tree clod" on the :right as shown in the two immediately preceding photographs. These two "clods" and their "craters" served as the second example of old-field (secondary plant) succession on a local denuded microsite caused by windthrow (straight line winds up to 80mph and wet soil). This "tree clod" was caused by uprooting of a huge honey locust, the crashing of which caused uprooting (secondary impact) of two pole-sized hackberry. Many plants of giiant ragweed were the immediate successional "beneficiaries" of this devestation and denudation that left bare soil and a hole to temporarily pond and provide supplemental water to "lucky" plants. Giant ragweed grew so thick in this colony of seedlings that even hairy crabgrass was largely excluded. Existing shoots of uprooted trees, including the huge honey locust and two hackberry, were killed. These trees temporarily leafed out at the appropriate time in spring following toppling (which was on 15 April, 2001), but leaves soon died and fell off before achieving any appreciable size. The big honey locust did send up vigerous and persistent root sprouts. Honey locust was the only tree species to successfully reproduce asexually (vegetatively). Ottawa County, Oklahoma. June 2001.

143. Fifth growing season of second blowdown example- Appearance of recovering (successional) range vegetation about mid-way through the fifth growing season on two "tree clods" and their "craters" caused by uprooting of a one large to huge honey locust which on crashing ripped up or broke off smaller (sapling to pole-size) American elm and hackberry. These were the third and fourth "tree clods" of four such uprooted clumps of trees. They were the same "clods" shown above at the first growing season after uprooting. (More thorough descriptions of tree species that made up "tree clods" were provided above when "clods" described for the first growing season following blowdown.) At this stage of secondary succession on denuded areas seedlings and root sprouts of the climax trees were well-established. Most of these new shoots of American elm, hackberry, and sugarberry were seedlings, the initial resprouts off of roots of mature trees having died by end of second warm-growing season. Sprouts of the immense trees of honey locust were vigerous and rapidly growing. Fox or frost grape was the major woody vine growing in association with seedlings and sprouts. Fox grape was presented in greater detail in photographs below. Also present as robust and vigerous plants were pioneer annuals both cool- and warm-season species. In this fifth spring-summer period following denudation the dominant (and native), annual pioneer species remained giant ragweed (the same as in the first growing season following blowdown) with hairy crabgrass also common. Perennial herbaceous species included pokeweed (Phytolacca americana) which was the largest, most rank-growing, and dominant of this category of plant. Pokesalad was obvious in interior of denuded spaces in both of these photographs. Poke was featured in close-in, detailed slides shown and described below. Another herbaceous perennial (represented by scattered individuals) was goldenrod, the plants of which were not blooming and thus could not be identified as to species. Climbing false buckwheat was another perennial forb and olne that was abundant, especially around perimeter and near "craters". Another abaundant, perennial forb was giant yellow hyssop (Agastache nepetoides) which was presented as scattered individuals in the second of these two photographs. Also still present at this stage was Japanese chess or Japanese brome (a Eurasian, cool-season, annual grass which was visible as brown- or amber-colored straw in foreground of second slide) along with silky wildrye which was the major native grass (a cool-season perennial and climax decreaser species).Another Eurasian annual that was common on the more-or-less still bare patches of soil was hedge parsley. This species had been common since the first cool-growing season following uprooting of trees. Observation: much ado is often about biodiversity, especially species diversity, in regards to plant succession and stages of vegetation development. Some ecologists have argued that plant diversity is greater in this or that stage, often arguing in favor of climax or late (advanced) seral stages. On tree-lowdown denuded spaces (forest gaps and, thus, patch dynamics) in climax hackberry-sugarberry-elm-honey locust lowland forest in the western Ozark Plateau, biodiversity as viewed by species number and plant forms was greatest--far and away so--in mid-sere, and clearly not at stage(s) in which plant species composition is typical of climax. More will be shown of this successional fact in photographs and captions taken of denuded spaces in the eighth warm-growing season following denudation (see below). Ottawa County, Oklahoma. July, 2005

144. Fifth growing season (close-in view) of second blowdown example- More detailed perspective of successional, forest range vegetation on a "tree clod" and adjoining "tree crater" created when huge honey locust trees were uprooted by high straight line winds (70-80mph) eight warm-growing seasons ago (prior to time of these photographs). The denuded microhabitat (forest gap) was in a climax hackberry-sugarberry-American elm-honey locust lowland forest in the Springfield Plateau (western partion of Ozark Plateau Region). This was the "tree clod" on the right (see above photographs) in second blowdown example being used in this section (see again first warm-growing season after tree blowdown). This was the fourth of four uprooted tree clumps described herein. It was created with the uprooting of a huge honey locust, the unearthing of which tore out (ripped out) as a secondary impact two pole-sized hackberry. The first of these two photographs showed climbing false buckwheat growing on face (bottomside) of "tree clod". Pokeweed, the dominant herbaceous species growing on the "clod-crater" complex, was also featured prominently in this slide. Giant ragweed, the dominant annual pioneer species that was still abundant five years after denudation, was conspicuous in upper right margin. A spindly plant of marestail or horseweed (slightly right-of-center in foreground) represented another native, annual pioneer of the Compositae. A tall (almost 10 feet in height) root sprout of honey locust that had been present since the first growing season following blowdown (spring, summer, autumn after 15 April, 2001) of the large honey locust was in upper left portion of this slide. Several large seedlings or small saplings (all apparently from seed) of hackberry, sugarberry, and American elm were also present, especially in background, as were shoots of fox grape. The second of these photographs was of the "crater", the hole created when soil and other earth materials were lifted out by the uprooted trees. The large, robust shoots of pokeweed that dominated (overwhelmed was more like it) much of the denuded area were featured in this slide. Also visible in this slide was a vigerous plant of silky wildrye (the consist dominant grass of blowdown-denuded habitats since their creation), Japanese chess or Japanese brome, marestail or horseweed, giant ragweed, poison ivy, and gray-headed coneflower (Ratibida pinnata), perennial composite. Pokeweed, a large perennial with an immense and fleshy taproot, is capable of living for years. Pokeberry is readily "shaded-out" by growing trees. The prime habitat for pokeweed is disturbed ground a few years after initial creation of the denuded spot. Ash from burning large accumulations of wood, bulldozed piles of brush, cleared road sides, and fencerows are ideal environments for pokeweed, especially if there are perches for birds that recently feasted on pokeberry fruit. Ottawa County, Oklahoma. July, 2005

145. Eighth growing season of second blowdown example- Late seral (advanced successional stage) forest range vegetation roughly mid-way into the eighth growing season on a "tree clod and crater" caused by uprooting and cavity creation by straight line winds (70-80mph) and wet soil in early spring. Local sites of denudation (both of these "clod-crater" microsites) were caused when a single massive honey locust was uprooted and, in secondary consequences of the crashing honey locust, smaller hackberry and/or sugarberry and American elm were ripped up. Development of forest vegetation had achieved species composition of the climax hackberry-sugarberry-American elm-honey locust community. The plant community on "tree clods and craters" was dominated overwhelmingly at this stage by small saplings of hackberry, sugarberry, American elm, and as feaured in these two slides, black walnut. Almost all of the young trees of these species were from seed and not root suckers (ie. seedlings and not sprouts of uprooted trees) Young trees of honey locust were, in contrast, primarily root sprouts off of severed roots that remained in the soil when adult shoots (combination of trunks nad crowns) of large honey locust were uprooted. These honey locust saplings of root origin had been abundant and conspicuous by the fourth and fifth spring-summer growing seasons (see slides and captions above, especially the first example of blowdown). At this stage, eighth tree-growing season following windthrow, Celtis and Ulmus species had grown so thick that their larger leaves "blocked the view" of honey locust with their smaller, finner leaves. On these two denuded microsites black walnut seedlings and saplings had benefitted especially from the "craters" that temporarily ponded extra water. At this stage of secondary plant sucession the terminal (climax) forest community had been reached from standpoint of dominant species composition, but not from criteria of physiogonomy, architecture (structure), physical arrangement, or, perhaps, total number of climax species. No, ecosystem structure and function had not been restored. That point of ecosystem restoration would require presence of mature (adult-size or fully grown) trees of the dominant species along with species associated with them as, for example, such lianas as grape, poison ivy, and trumpet creeper. Gone from this plant community with its species composition comprised almost entirely of small trees of the climax dominant tree species were almost all annuals including giant ragweed, the dominant pioneer and later seral stage herbaceous species, as well as perennials of advanced stages like pokeweed. At edge or outer perimeter of this "tree clod and crater" there was one locally common, annual weed which wass shown in details of close-in views presented in the nest two slides immediately below. Ottawa County, Oklahoma. July, 2008.

146. Eighth growing season (close-in view) of second blowdown example- Detailed views of the second (rightmost) "tree clod and crater" of the two "clods" that served as the second example of secondary plant succession on local sites (microsites) of denudation caused by uprooted trees resulting from high straight line winds (70-80mph) and wet soil eight growing seasons ago. This "clod and crater" resulted from uprooting of a huge honey locust and two pole-sized hackberry. At this stage of forest recovery (vegetation development by old-field succesion on a lowland forest sere) the denuded site was almost completely populated by hackberry and sugarberry with some American elm and honey locust as associate species. Annuals, biennials, and herbaceous perennials had been "overtopped and overrun" by the climax tree species. For example, the large plants of pokeweed present from about the third to sixth year of secondary succession (recall from above) had either died out or survived as depauperate specimens (presumedly due to competition for light, growing space, soil water, or whatever from young saplings of climax dominants). In other words, species composition was restored to that of the climax hackberry-sugarberry-American elm-honey locust forest. Physiogonomy, structure, and function of the climax forest was far from restored. These features of the climax stage of forest range vegetation would not be restored, the forest would not be fully recovered (ie. the fully functional climax forest ecosystem will not be achieved or reacquired) until these large seedlings and small saplings reach adulthood and mature size. On basis of species makeup as determined by relative cover or biomass the botanical composition of climax had been reached. Decades would be required for complete recovery of the forest ecosystem and full development of forest vegetation. The phenomenon shown in this section devoted to forest gaps and patch dynamics was that the time span (temporal aspect or element) of secondary plant (old-field) succession from initial denudation back to the climax stage based on botanical composition of the plant community, the vegertation, was remarkably (indeed, almost miraculously) short or brief. Number of growing seasons was very few (only seven or eight) in the progression of vegetation development from bare soil to dominance of the disturbed site by climax tree dominants. Development of vegetation, progression along the forest sere, from denudation to termination of successional stages (advance of seral stages culminating in the climax community based on plant species) is rapid on the small spaces laid bare by uprooted individual large trees and/or small numbers of affilitated smaller trees. Secondary succession is rapid on the small microhabitats that are characteristic of forest gaps. Patch dynamics--secondary plant succession on small gaps (micro-spatial scale) created by local disturbances (tree blowdown in these examples)--is considerably faster than on larger areas such as old (abanodoned) farm fields or cutover foresst land. There is an interaction of temporal and spatial affects: temporal aspects operate (proceed) faster at smaller spatial scale (ie. vegetation development by secondary succession takes place more rapidly when areas of disturbance are smaller). Recovery of late seral and climax vegertation depends on size of disturbance (how much land, the area, was denuded) as well as time and plant-growing conditions related to time. Ottawa County, Oklahoma. July, 2008.