Southern and Central Forests I

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”.

Live oak (Quercus virgiana) Hammock Forest

1. Live oak hammock (hammock is Indian for "shady place")- This is a unique land form and range vegetation type restricted to the southeastern portion of North America, especially Florida. Hammocks are characterized by being fairly level to gently rolling and higher than surrounding land. Yet they are poorly drained and fertile (soils are high in humus) with hardwood communities that are typically dominated by live oak canopy and an understory of cabbage palm (Sabal palmetto), skunk cabbage (Symplocarpus foetidus), carpetgrass (Axonopus affinis), and panicgrasses (Panicum spp.). FRES No. 16 (Oak-Gum-Cypress Ecosystem). There is no Kuchler equivalent for communities this small; the Kuchler system lumped this type in with some other larger unit of which K-81 (Live Oak-Sea Oats) seemed most fitting. SRM 817 (Oak Hammocks). Mixed Hardwood Series in Floridian Evergreen (Hammock) Forest biotic community of Brown et al. (1998). Southern Coastal Plain- Central Florida Ridges and Uplands Ecoregion, 75c (Griffith et al., undated).

2. Interior of a Florida live oak hammock with tell-tale raised yet wet ground- Mid-canopy shrubs include wax myrtle (Myrica cerifera) and American beautyberry (Callicarpa americana) with sparsely scattered saw palmetto (Serenoa repens).FRES No. 16 (Oak-Gum-Cypress Ecosystem). Subunit of K-81 (LiveOak-Sea Oats). SRM 817 (Oak Hammocks). Mixed Hardwood Series in Floridian Evergreen (Hammock) Forest biotic community of Brown et al. (1998). Southern Coastal Plain- Central Florida Ridges and Uplands Ecoregion, 75c (Griffith et al., undated).

Slash Pine (Pinus elliottii) Forest

Slash pine (Pinus elliottii= P. caribrea= P. heterophylla) grows--or at least under natural conditions (like pre-whiteman days)--in associateion with longleaf pine (P. palustris) with the latter growing on drier upland forest sites while slash pine commands control of more mesic habitats such as environments around ponded water (hence the other common, common name of pond pine); plus slash pine is between longleaf pine and loblolly pine (P. taeda) with regard to tolerance (Harlow et al., 1979, p. 97). For this reason these two native, southeastern, yellow pines have typically been included as the same forest ecosystem (Garrison et al., 1977, p. ps. 5-8) and same forest cover type (SAF 83; Eyre, 1980, p.s 52-53). In addition, however, slash pine occurs as a sole dominant of a subclimax cover type (SAF 84, SAF 111; Eyre, 1980. ps. 56- 58) and as the principal pine of the hardwood-pine type (SAF 85; Eyre, ps. 61-62). With human-directed fire exclusion (ie. replacement of natural fire regimes) slash pine gradually replaces longleaf pine on the more moist habitats as well as growing as pioneering tree species on cutover forest lands much like loblolly pine (Harlow et al., 1979, p. 97). Slash pine is a major pine of considerable commercial importance in southeastern North America, especially in Florida. Slash pine is valuable as both a commercial wood crop species in pine plantations and a dominant native tree of more natural forests. Under most such situations, there is slash pine-dominated/defined forest range that is usuable by both livestock (mostly cattle) and wildlife. As shown below, however, unless plantations of slash pine are thinned and/or aritfically pruned eventually a closed canopy forms resulting in exclusion of the previous, successional, herbaceous understorey. Under these conditions, slash pine range is transitory forest range--a fleeting forage resource--in the same manner as loblolly pine.

3. Florida flatwoods range site at climax- This is the most widespread range site in Florida. Slash pine (Pinus elliottii) upperstory with a nutritious and productive understory dominated by creeping bluestem (Andropogon stolonifer), chalky bluestem (A. capillipes), and shortspike bluestem (A. brachystachys). Osceola County, Florida. February. FRES NO. 12 (Longleaf-Slash Pine Ecosystem). K-106 (Subtropical Pine Forest [Southern Florida]). SRM 811 (South Florida Flatwoods). Variant of Pine Series in Southeastern Deciduous and Evergreen Forest biotic community of Brown et al. (1998). Southern Coastal Plain- Eastern Florida Flatwoods Ecoregion, 75d (Griffith et al., undated).

4. Florida flatwoods range with saw palmetto invasion on deteriorated range- Infrequent fire and possibly past abuse from overstocking has depleted the climax bluestem understory allowing domination by the unpalatable, less fire-tolerant, invading palmetto. Osceola County, Florida. February. FRES No. 12 (Longleaf-Slash Pine Ecosystem). K-106- (Subtroical Pine Forest [Southern Florida]), some departure from climax. SRM 811 (South Florida Flatwoods). Variant of Pine Series 123.12 of Southeastern Deciduous and Evergreen Forest biotic community Warm Temperate Forest and Woodland 123 of Brown et al. (1998, p.38). Southern Coastal Plain- Eastern Florida Flatwoods Ecoregion, 75d (Griffith et al., undated).

121254 and 121255.

Future Florida forest- Seedlings of slash pine in an herbaceous understorey of a bluestem range with major species including creeping (Andropogon stolonifer), Elliott's bluestem (A. elliottii), chalky bluestem (A. capillipes), shortspike bluestem (A. brachystachys), and toothachegrass (Ctenium aromaticum). There was some cover of pineland threeawn (Aristida stricta), but htis was a pine-bluestem range cover type not a pine-wiregrass range type. Understorey of this forest was climax vegetation. as was woody component except that trees were not at old-growth state. Avon Park Air Force Bombing Range, Highlands County, Florida. February. FRES No. 12 (Longleaf-Slash Pine Ecosystem). K-106- (Subtroical Pine Forest [Southern Florida]). SRM 811 (South Florida Flatwoods). Variant of Pine Series 123.12 of Southeastern Deciduous and Evergreen Forest biotic community Warm Temperate Forest and Woodland 123 of Brown et al. (1998, p.38). Southern Coastal Plain- Eastern Florida Flatwoods Ecoregion, 75d (Griffith et al., undated).

5. Slash pine plantation- Excellent management with thinning and prescribed burning to control saw palmetto. Understory is primarily wiregrass or threeawn, especially pineland threeawn (Aristida stricta). Avon Park Bombing and Gunnery Range, Florida. February. FRES No.12 (Longleaf-Slash Pine Ecosystem).Variant of K-106 (Subtropical Pine Forest [Southern Florida]), human-directed departure from climax and an example of agroforestry. FRES No. 12 (Longleaf-Slash Pine Ecosystem). K-106- (Subtroical Pine Forest [Southern Florida]). SAF 84 (Slash Pine). Variant of Pine Series 123.12 of Southeastern Deciduous and Evergreen Forest biotic community Warm Temperate Forest and Woodland 123 of Brown et al. (1998, p.38). Southern Coastal Plain- Eastern Florida Flatwoods Ecoregion, 75d (Griffith et al., undated).

6. Needles and cones of slash pine (Pinus elliottii= P. caribrea= P. heterophylla)- The attractive long leaves of this fast-growing pine (and of the longleaf pine) are likely an example of evolution and adaptation to specific habitats. The pines sporting long needles are native to the Lower South where heavy snow and ice storms are rare. The shortleaf pine with it's much shorter leaves is native to the Upper South where infrequent but severe ice storms are a winter fact of life. Ice storms in particular can destroy a crop of young long-needle pines just as surely as ice in the form of hailstones wipes out a wheat crop. Small (1933, p.5) remarked that slash pine produced the the softest wood of the eastern pines and had the most restricted biological range. Standard references for slash pine included Sargent (1933, ps. 15-16), Vines (1960, ps. 20-22), Harlow et al. (1979, ps. 97-98), and Burns and Honkala (1990).

7. Slash pine plantation-This stand was recently thinned to two-thirds of pre-thinning density; about 20 years post-planting. Slash will be disposed of primarily by prescribed burning which will do "double-duty" of reducing cover of saw palmetto and the likelihood of danger to trees from wild fire.Pine will be harvested at about 45 to 50 years post-planting. Slash pine stands such as this are clearly transitory forest range with a grazable understory persisting only until pines grow large enough that their crowns form a closed canopy which excludes light from the forest floor. Avon Park Bombing and Gunnery Range, Florida. February. FRES No. 12 (Longleaf-Slash Pine Ecosystem). Variant of K-106 as explained immediately above.SAF 84 (Slash Pine). Variant of Pine Series in Southeastern Deciduous and Evergreen Forest biotic community of Brown et al. (1998). Southern Coastal Plain- Eastern Florida Flatwoods Ecoregion, 75d (Griffith et al., undated).

Big Thicket Forest (including "islands" or isolated tracts of the Mixed Mesophytic Type)

8. Bayou range in the core of Texas Big Thicket- This frequently flooded bottomland site has high clay content soils that dry rapidly (due to high evapo-transpiration rates) creating an understory dominated by dwarf palmetto (Sabal minor). The tree layer defines this laurel oak (Quercus laurifolia)-overcup oak (Q. lyrata)-loblolly pine (Pinus taeda) forest range type. This is an edaphic climax as viewed from the perspective ofpolyclimax theory. Little Pine Island Bayou, Big Thicket National Preserve, Hardin County, Texas. May. FRES No. 16 (Oak-Gum-Cypress Ecosystem).One of many variants of K-103 (Southern Flood-Plain Forest). One of the numerous variants of SAF 82 (Loblolly Pine-Hardwood). Oak-Pine Series in Northeastern Deciduous Forest biotic community (even though in southeastern part of continent; an Oak-Pine Series for Southeastern Deciduous and Evergreen Forest biotic community would seen warranted) of Brown et al. (1998). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

9. Climatic (= regional) climax of Texas Big Thicket- American beech (Fagus grandifolia)-southern magnolia (Magnolia grandiflora)-white oak (Quercus alba)-loblolly pine forest. In Clementsian monoclimax theory this is the ultimate formation, the endpoint of vegetation development to which all vegetation of this region converges. It is the climax on mesic, level, upland areas. Trees left to right: southern red oak (Quercus falcata var. pagodifolia), magnolia, loblolly pine and beech. Lance Rosier Unit, Big Thicket National Preserve, Hardin County, Texas. May. This magnificant forest cover type was apparently too small for FRES or Kuchler to "pigeon-hole". As it is one of the southern forest types with a pine as one dominant species it would have to be included in FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem) with K-101 (Oak-Hickory-Pine Forest) being the Kuchler equivalent listed thereunder.However, the close "kinship"of this forest to the mixed mesophytic and mixed hardwoods forests (both in the Southeastern Evergreen Forest Region) and to the mixed mesophytic forest in the Southern Appalachians of the Oak-Chestnut Forest Region (Braun, 1950, ps. 199-205, 297-303) strongly suggested that this cover type most closely fit Kuchler-95 (Applalachian Oak Forest) which is an equivalent under FRES 15 (Oak-Hickory Forest Ecosystem). Frankly, it was surprising to this author that Kuchler did not give a more specific designation for this major, though restricted, type given that Braun (1950, p. 445) specified that it could be seen as a separate association the same as for the Oak-Hickory Association and the Beech-Maple Association both of which have FRES and Kuchler recognitions. (Braun [1950, ps. 443-445] placed the Beech-Magnolia climax in the Mixed Mesophytic Association as transitional between Deciduous and Broad-leaved Evergreen Formations.) The Society of American Foresters (1980) also missed this one. The closest SAF forest cover type is probably 82 (Loblolly Pine- Hardwood) but that is not close enough. Beyond any doubt beech and magnolia are the recognized dominants and the pine is the least of the dominants. Only with man-set prescribed fire could the loblolly pine be maintained at higher proportions of the climax community.This is primarily a hardwood type and certainly not a pine-oak type. Braun (1950, ps. 300-303, enclosed map) discussed and mapped the beech-magnolia forest as part of the Southeastern Evergreen Forest Region. Braun did not map at association levels. Perhaps western island of Mixed Mesophytic Series in Southeastern Deciduous and Evergreen Forest biotic community of Brown et al. (1998). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

10. Climax mixed mesophytic forest (the beech-magnolia-loblolly pine type)- This is the westernmost extension of a forest type dominant in the Great Smokey Mountains and Applachian region. Tree in foreground is American beech; trees on far left are magnolia. Lance Rosier Unit, Big Thicket National Preserve, Hardin County, Texas, May. Mesic site. FRES and Kuchler recognitions (lack thereof) discussed immediately above. Variant of SAF 82 (Loblolly Pine-Hardwood). Oak-Pine Series of Brown et al (1998) (but should be for Southeastern Deciduous and Evergreen Forest biotic community). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

11. Climax of the mixed mesophytic forest- In the Clementsian concept of climax (ie. a monoclimax or regional climax determined primarily by climate) the ultimate expression of potential natural vegetation of the deep pineywoods (general hardwood, namely oak,-pine forest) is a mixing of this regional climax with elements of the mixed mesophytic forest to the east. In this species-rich forest vegetation southern magnolia and American beech join with white oak (and several other oak species as shown below) and loblolly pine to form a unique forest range community that can be visualized as postclimax (ie. more mesic than the general climax of the area or region) as was explained by braun (1950, p.13). This photograph of a mixed mesophytic X white oak-loblolly pine forest (ie. an overlapping or ecotone of these two was on a locally wet site and featured a large, old southern magnolia (foremost tree in center foreground with bulging trunk) growing beside (to right of) an immense white hockory, mockernut hickory, hognut hickory, or white-heart hickory (Carya tomentosa) with readily identifiable soft gray bark and pointed compound leaves. These two "champines" of their respective spceis stood majestically in front of loblolly pine, white oak, laurel oak (Quercus laurifolia) and cherrybark oak (Q. prinus). The backside (side away from the photograph) of this specific southern magnolia was shown in detail in the next photograph. Lance Rosier Unit, Big Thicket National Preserve, Hardin County, Texas, May. Mesic site. FRES and Kuchler recognitions (lack thereof) were covered in the photo caption before the immediately preceding slide. Variant of SAF 82 (Loblolly Pine-Hardwood). Oak-Pine Series of Brown et al (1998) (but should be for Southeastern Deciduous and Evergreen Forest biotic community). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

12. Goosepen in the Big Thicket- The forester's term, goosepen, is "a place hollowed out by fire at the base of a standing tree" (Munns, 1950). This large fire scar (it extended to a height of nearly five feet) was on the backside of the large southern magnolia featured in the immediately preceding photograph. Such fire scars on old trees attest to incidence of surface forest fires in the pineywoods the same as obtains for all other regions of the eastern deciduous forest. Lance Rosier Unit, Big Thicket National Preserve, Hardin County, Texas, May.

13. Climatic climax vegetation type of Big Thicket form of Texas Piney Woods- The beech-magnolia-white oak-loblolly pine forest cover type. This is the ultimate expression, the mesophytic form, of the Big Thicket forest.Magnolia, far left; white oak; center right. Note lush understory of browse plants even under closed forest canopy at climax. Tree species visible include loblolly pine, laurel oak, and swamp chestnut oak (Quercus prinus).Lance Rosier Unit, Big Thicket National Preserve. May, vernal aspect; mesic site. Problems with unit recognition by FRES, Kuchler, and SAF discussed two slides above. South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

14. Southern magnolia (Magnolia grandiflora)- A beautiful flower of the stately southern magnolia against a backdrop of the broad sclerophyllus leaves of this climax pineywoods species. State Tree of Mississippi.

Water Oak (Quercus nigra) Forest

The first example of the water oak (Quercus nigra) forest cover type shown immediately below was in a commercial forest in the Big Thicket section of the Texas Pineywoods. This water oak forest vegetation was adjacent to loblolly pine stands and a forest dominated by loblolly pine, water oak, American holly both with a lower woody layer comprised primarily of yaupon or, often called, yaupon holly (Ilex vomitoria). Water oak has been regarded as Intolerant as to tolerance and as a subclimax species that is quite susceptible to fire damage (Fowells, 1965, p. 630; Burns and Honkala, 1990, Vol. 2, p. 703). Thus while light surface fires tend to maintain pines like the associated loblolly pine, major fire damage as with crown fires would select for regeneration of water oak. In absence of fire plant succession would progress to a climax of hardwoods, which in the Big Thicket would commonly be American beech, southern magnolia, American holly, and climax oaks such as white oak.

15. Water oaks in the Pineywoods- Exterior view of a local stand of water oak growing on a flatland forest site that frequently ponded water. Loblolly pine were growing around perimeter of the water oak stand. Yaupon grew as widely scattered individuals while most of the ground layer was oak leaves with scattered plants of longleaf woodoats (Uniola sessifolia), cottongrass bulrush (Scirpus cyperinus), and green flat sedge (Cyperus virens). These species (from this locale) were featured below under the loblolly pine-water oak-American holly form or subtype of loblolly pine-hardwoods forest. The largest--and also the most scarce-- herbaceouss pecies was bentawn plumegrass (Erianthus contortus) which was also featured below. Liberty County, Texas. February, late hibernal aspect. This is a component or subtype of the general hardwood-pine southern forest forest that has one of the southeastern yellow pines a dominant or, sometimes, an associate species with oaks, hickories, or even beech as the more common climatic dominant (in contrast to a fire-determined dominant). Overall this forest range vegetation would have to be included in FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem) with K-101 (Oak-Hickory-Pine Forest) being the Kuchler equivalent listed thereunder. SAF 88 (Willow Oak-Water Oak-Diamondleaf [Laurel] Oak). Oak-Pine Series of Brown et al (1998) (but should be for Southeastern Deciduous and Evergreen Forest biotic community). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

Inside the water oaks- Interior of the local stand of water oak presented immediately above. This was a local consociation of Quercus nigra with a "broken" (widely scattered) population of yaupon holly and local herbaceous cover composed variously of longleaf woodoats, cottongrass or woolgrass bulrush, green flat sedge, and panicgrasses (Panicum spp.). This isolated water oak stand was adjacent to a mixed forest of loblolly pine, wter oak, and American holly (covered below). Liberty County, Texas. February, late hibernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem) with K-101 (Oak-Hickory-Pine Forest) being the Kuchler equivalent listed thereunder. SAF 88 (Willow Oak-Water Oak-Diamondleaf [Laurel] Oak). Oak-Pine Series of Brown et al (1998) (but should be for Southeastern Deciduous and Evergreen Forest biotic community). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004)..

16. One of the more common forms or manifestations of oak forest in the Pineywoods of Texas and Louisiana is the Palmetto-Oak Flats (Ajilvsgi, 1979, ps. 12-13) or, when expressed as to topographic-edaphic rather than botanical features, Clayey Wet Upland Depressions (Diggs et al., 2006, ps. 97-98). Ajilvsgi (1979, p. 12) described overcup oak and laurel oak as dominants whereas Diggs et al., (2006, p. 98) emphasized willow oak and green ash (Fraxinus pennsylvanica) as major plants of the larger tree species. The Society of American Foresters (Eyre, 1980, p. 63) described the willow oak-water oak-diamondleaf or laurel oak type (SRM 88) as developing on a topographic-soil moisture gradient intermediate between the swamp chestnut oak-cherrybark oak type (SRM 91) and the overcup oak-water hickory type (SRM 96) with dominance of SRM 88 tending to change to non-oak hard spceies like green ash under heavy logging or high-grading. The photographs shown below were of a water oak-willow oak forest with a lower shrub layer made up almost exclusively of dwarf palmetto and a herbaceous layer(s) of sedges, rushes, bulrushes, and panicoid grasses. Views of the Oak-Palmetto Flats in these slides were presented so as to view this forest range vegetation going from exterior to deep interior as if the viewer were traveling to and then into it.

17. Coming onto the Oak-Palmetto Flats- Exterior view of an example of the willow oak-water oak-diamondleaf (laurel) oak type showing physiogonomy and overall species composition of this form of Pineywoods. Dominant species of this stand was water oak with willow (locally known as "pin" oak). Laurel oak was a distant third Quercus species. Blackgum or black tupelo (Nyssa sylvatica) was another associate tree species. The largest tree with the horizontal upper limb and fire-scarred basal trunk was an ancient water oak readily idetified by the sporadically scattered, prominent "warts" of bark. Loblolly pine was represented by one conspicuous tree in center midground. There were other infrequent loblolly pine throughout. Young trees grouped at right foreground were a mixture of water and willow oak and very black tupelo. Dwarf palmetto made up a lower shrub layer. Grassses and grasslike plants comprised one or two (rarely three) herbaceous layers in the forest understorey. Herbaceous plants were most common around perimeter of the forest vegetation. Individuals of broomsedge bluestem (Andropogon virginicus) were prominent in foreground understorey. Hardin County, Texas. February, hibernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem) with K-101 (Oak-Hickory-Pine Forest) being the Kuchler equivalent listed thereunder. SAF 88 (Willow Oak-Water Oak-Diamondleaf [Laurel] Oak). Oak-Pine Series of Brown et al (1998) (but should be for Southeastern Deciduous and Evergreen Forest biotic community). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

18. Edge of an Oak-Palmetto Flats forest range- Around perimeter of a stand of water and willow oak with dwarf palmetto were various local assemblages of herbaceous plants. The latter included cottongrass bulrush and miscellaneous sedges, both Carex and Cyperus species (eg. green flat sedge [C. virens]), along with panicgrasses, especially beaked panicgrass (Panicum anceps); paspalums like brownseed paspalum (Paspalum plicatulum), and both broomsedge and bushy bluestem. These latter two species are invaders. These same species also formed herbaceous strata beneath the oaks and pines though with less continuous cover and smaller plants, conditions likely resultant from fairly dense shade. Water and willow oaks are Intolerant species. Hardin County, Texas. February, hibernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem) with K-101 (Oak-Hickory-Pine Forest) being the Kuchler equivalent listed thereunder. SAF 88 (Willow Oak-Water Oak-Diamondleaf [Laurel] Oak). Oak-Pine Series of Brown et al (1998) (but should be for Southeastern Deciduous and Evergreen Forest biotic community). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

19. Into the Pineywoods flats we go- These three photographs were a pictorial "walk to the woods", a sequence of slides showing the range vegetation of a water oak-willow oak- loblolly pine-palmetto-herbaceous plants Pineywoods flats. Continually closer-in views allowed presentation of the herbaceous layer(s) of native vegetation that was better developed at outer edge of the forest stand. Some of the common herbaceous species of this vegetation were presented below in the section devoted to the loblolly pine cover type, specifically the loblolly pine-water oak-American holly form or subtype thereof. The smaller trees in foreground with unshed lower limbs (most of them still alive but senescing) were willow oak. Locals hereabouts apply the otherwise confusing and nonstandardized common name of "pin oak" to Quercus phellos. "Pin" in several oak species refers to any of the lower, usually dead, unshed limbs (ie. dying or dead limbs on species that do self-pruning, but instead become well-seasoned or preserved and, hence, persistent on the lower bole). There were a few scattered woody vines, the only one of which the author-photographer identified was rattan (= Alabama supplejack). Once inside the Pineywoods flats the interior of the water oak-willow oak-dwarf palmetto-herbaceous range community revealed a "closer-in" view of plant species composition and the lower woody layer of palmetto and the local vertical zone of herbaceous species. Largest trunk was that of a young to mid-age water oak with bark characteristic of an immature tree. At this stage of maturity bark of water oak and willow oak is so similar as to be indistinguishable, thereby making reliance on leaves and buds necessary for definitive indetification. "Warty" bark on older water oak bark was just forming on this straight-trunked specimen, but some smaller water oaks had larger "warts". Grass shoot (visible in both photographs) in front of this water oak was broomsedge bluestem, a common invader of Oak-Palmetto, which was common and conspicuous throughout this oak flats stand. Almost all herbaceous species were grasses or grass-like plants and, as this was dead of winter and this range had been grazed so that most species had to be identified by vegetative features, most herbs could not be identified by the author who was a "stranger to these parts". The tallest green herb was cottongrass bulrush (shown and described briefly below). There were no prominent forbs in this forest range vegetation. Dwarf palmetto comprised a single-species, lower, woody layer. Hardin County, Texas. February, hibernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem) with K-101 (Oak-Hickory-Pine Forest) being the Kuchler equivalent listed thereunder. SAF 88 (Willow Oak-Water Oak-Diamondleaf [Laurel] Oak). Oak-Pine Series of Brown et al (1998) (but should be for Southeastern Deciduous and Evergreen Forest biotic community). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

20. "Up-and-dicular" perspective of a Oak-Palmetto Flats- Structure and species composition of the water oak-willow oak-dominated Pineywoods flatwoods described under horizontal photographs above. Architecture of this stand was displayed to better advantage in these two photographs. Most hardwood trees were water oak and willow oak of sapling to small pole size. Those with persistent lower limbs were willow oak. There was an "occasional" black tupelo (also of sapling-pole size). Cover and density of palmetto was shown to good advantage in the first of these two slides while the frequent openings within the palmetto that were populated by grasses and grasslike plants were evident in the second slide. Tree in left foreground with live lower limb was willow oak. This stand was obviously a second-growth forest. A cohort of sapling to small pole size oaks had developed beneath larger, established (older) but very widely scattered, mature oaks of both species. Structure and, especially, botanical composition of this stand was typical of climax water oak-willow oak-laurel oak-palmetto vegetation. Both willow oak and water oak are classified as Intolerant and recruitment of these species had been possible under a mostly open sky (sparse canopy of oak and loblolly pine). Natural thinning of oaks had already commenced as evidenced by the dead toppled pole (visible in both photographs). This will undoubtedly continue resulting in more dead younger oaks and fewer, though larger, trees (fewer boles but more board foot/acre) and eventually greater oak crown cover (increased--though by no means closed--tree canopy). The apparent dominant herbaceous species was cottongrass bulrush. Numerous individuals of broomsedge bluestem were conspicuous with their tannish yellow shoots dispersed among bulrush and other grasslike plants. Hardin County, Texas. February, hibernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem) with K-101 (Oak-Hickory-Pine Forest) being the Kuchler equivalent listed thereunder. SAF 88 (Willow Oak-Water Oak-Diamondleaf [Laurel] Oak). Oak-Pine Series of Brown et al (1998) (but should be for Southeastern Deciduous and Evergreen Forest biotic community). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

21. Closing, composite shot of Pineywoods Oak-Palmetto Flats- All-in-one shot of species composition and structure (architeture) the water oak-willow oak-loblolly pine-palmetto-cottongrass bulrush-broomsedge bluestem community featured above. All of these species except loblolly pine, which dominated (generally and/or locally) their respecive layers of vegetation, were visible (if not obvious). In addition, rattan (= Alabamas supplejack) was featured prominently growing up trunks of oaks in left midground. Almost all oak trunks of any age are hosts to various crustose lichen, at least on north and east exposure. Hardin County, Texas. February, hibernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem) with K-101 (Oak-Hickory-Pine Forest) being the Kuchler equivalent listed thereunder. SAF 88 (Willow Oak-Water Oak-Diamondleaf [Laurel] Oak). Oak-Pine Series of Brown et al (1998) (but should be for Southeastern Deciduous and Evergreen Forest biotic community). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

crown hignight

Water oak (Quercus nigra)- Upper trunk and crown of water oak showing leaves and bark of intermediate maturity. Older or most mature bark of water oak often forms wart-like raised areas (basal trunk and stump area). Houston County, Texas. March.

22. Dwarf palmetto, blue palmetto, swamp palmetto, dwarf palm, blue palm, etc. (Sabal minor)- Large, mature swamp palmetto with previous season's floral stalk and spent inflorescence. This true palm is most commonly acaulescent (lacking a trunk or bole) though sometimes there are individuals that have a single, short woody stalk which would "pass for" a trunk. The shoot or stem does not branch and is characterized as woody or pithy in nature. The speciment portrayed here was growing in the water oak-willow oak-lobollly pine-palmetto-cottongrass bulrush-broomsedge stand featured above. Hardin County, Texas, February.

Ecotone between Water Oak-Willow Oak Forest and Gulf Coastal Marsh

Another form or subtype of the primarily water oak (with willow oak locally con-dominant) cover type in the Texas-Louisiana Pineywoods develops as a transition zone between the hardwood-pine forest and the Gulf Coast marsh with one or the other of the major range plant communities (forest or marsh) having apparent dominance or predominance as aspect dominance (ie. either widely scattered hardwoods and pine with profusely branching, open crowns growing in a marsh or, alternatively, grasses, sedges, rushes, bulrushes, and other herbaceous marsh species growing as a sparse understorey beneath twater oak, willow oak, laurel oak, overcup oak, blackgum or black tupelo, green ash, and loblolly pine). It seemed to this author that it was less confusing and more consistent to include full coverage of this ecotonal range vegetation at this location rather than with Gulf Coastal marsh which was covered under the chapter, Tallgrass Prairie (Coastal). This forest range vegetation typically develops as or into a savanna and should in general be interpreted as such. Occasionally, however, there will be local communities of this ecotonal vegetation that appear to be more of forest (at least woodland) physiogonomy. This latter range plant community develops on depressions or ponded local sites. Both expressions of this ecotone are wetlands and were included below.

23. Out of the woods and onto the marsh- Exterior view of ecotone of Pineywoods and Gulf Coast marsh. Extreme eastern edge of hardwood-pine forest and beginning of coastal sedge-bulrush-tallgrass freshwater marsh. Feature of ponded water with combination of wet mesophytic and hydrophytic trees and an understorey of grasses and grasslike plants. Trees clearly dominanted the forest- or woodland-phase of this savanna wetland at edge of the Pineywoods forest whereas the predominant marsh-phase or form of this ecotone was in the background (and featured in succeeding slides). Even in this slide that emphasized the tree-dominated form there were scattered individuals of cottongrass bulrush (larger green clumps), numerous species of Carex and Cyperus along with various grasses of genera Panicum, Paspalum, and Andropogon. Tree species visible in this photograph included water oak, willow oak, blackgum or black tupelo, green ash (Fraxinus pennsylvanica), and loblolly pine. Hardin County, Texas. February, hibernal aspect. This ecotone or transition zone perhaps should be given its own designation as a distinct potential natural vegetation, but such a classification unit of rangeland or forest vegetation as cover type either by Society for Range Manageament or Society of American Foresters), Forest and Range Ecosystem (USDA Forest Service) or by Brown et al., (1998). There was further--even more--confusion as to most precise desingation of the herbaceous wetland because it had features (species composition, structure, geographic location) of both SRM 711 (Bluestem-Sacahuista Prairie) and SRM 807 (Gulf Coast Fresh Marsh) as if it was a mixture of these two rangeland cover tyhpes. Closest overall vegetational designation with existing classification systems is a combination or "hybrid" of: FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem) with K-101 (Oak-Hickory-Pine Forest) being the Kuchler equivalent listed thereunder, SAF 88 (Willow Oak-Water Oak-Diamondleaf [Laurel] Oak), Mixed hardwood Series (223.13), Southeastern Swamp and Riparian Forest (223.1) of Brown et al (1998, p. 43) and FRES No.41 (Wet Grasslands Ecosystem) with K-70 (Southern Cordgrass Prairie) the Kuchler equivalent thereunder, SRM 711 X 807, Southeastern Interior Marshland (243.1), Warm Temperate Marshland of Brown et al. (1998, p. 45). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

24. Interior of the ecotone between a mixed hardwoods flat and freshwater marsh- Inside the woodland- or forest-form or phase of an overall or general savanna formed by the transition between eastern edge of hardwood-pine Pineywoods and Gulf Coastal marsh of sedges, bulrushes, and panicoid grasses. Although the local site shown here has ponded water much of the year it is not inundated for a long enought period to qualify as a swamp. Presence of blackgum or black tupelo, a swamp species, along with a few individuals of overcup oak (Quercus lyrata) attested to extended periods of saturated (= a hydric state) soil. Dominance of this plant community by water oak and willow oak showed this site to be intermediate between poorly drained sites of overcup oak-water hickory forest and better-drained sites dominated by swamp chestnut oak and cherrybark oak (Eyre, 1980, p. 63). Dwarf or swamp palmetto (eg. center midground of first slide) was the main shrub, but individuals of this species were too widely scattered to form a shrub or lower woody layer. Most common and largest herbaceous species in understorey beneath trees on this ponded habitat was cottongrass bulrush (visible as large green tufts and also a component in large dried clumps of herbage). Carex, Cyperus, Panicum, Paspalum, Andropogon (especially A. virginicus, broomsedge bluestem, and A. glomeratus, bushy bluestem), and bentawn plumegrass were present either under crown canopy or adjacent to crown drip line. Loblolly pine in this wetland vegetation was represented by the largest trunk in the second slide. Tree immediately to right of the loblolly pine was a green ash. Foremost tree (left foreground) was a water oak. Hardin County, Texas. February, late hibernal aspect. Tree-dominated phase of this savanna that was an ecotone of: FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem) with K-101 (Oak-Hickory-Pine Forest) being the Kuchler equivalent listed thereunder, SAF 88 (Willow Oak-Water Oak-Diamondleaf [Laurel] Oak), Mixed hardwood Series (223.13), Southeastern Swamp and Riparian Forest (223.1) of Brown et al (1998, p. 43) and FRES No.41 (Wet Grasslands Ecosystem) with K-70 (Southern Cordgrass Prairie) the Kuchler equivalent thereunder, SRM 711 X 807, Southeastern Interior Marshland (243.1), Warm Temperate Marshland of Brown et al. (1998, p. 45). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

25. Outer perspective of hardwood-pine flats X fresh Gulf Coast marsh ecotone- Two views of the transition zone (and quite a tension zone at that) of a water oak-willow oak-overcup oak-blackgum-green ash-loblolly pine-palmetto community and a cottongrass bulrush-sedge-bluestem-panicgrass-paspalum-cordgrass freshwater coastal marsh. In contrast to the tree-dominated form of this ecotone presented above this showed the more widespread--and more savannah-like-- form of a marsh with scattered trees of the just-listed tree species. The largest trees and with spreading crowns in the first of these two photographs were water oak. Trees in center of second photograph included black tupelo (three trunks together). Identification of grasslike plants and most grasses was impossible in the existing state of decay and/or shattered fruit. Large individuals of cottongrass bulrush and broomsedge bluestem were exceptions. The Illustrated Flora of East Texas (Diggs et al., 2006) indicated 42 species of sedge (Carex spp.) and 18 species of flatsedge (Cyperus spp.) for the small county of Hardin. It was unquestionable that many of these species, along with species of spikerush (Eleocharis spp.) and bulrush (Scirpus spp.) as well as panicoid grasses such as bushy and broomsedge bluestem and isolated large individuals of sacahuista or Gulf cordgrass (Spartina spartinae), were present on this botanically diverse savannah. Forbs were limited nearly to point of "nonexistence". Hardin County, Texas. February, late hibernal aspect. This ecotone was a combination of : FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem) with K-101 (Oak-Hickory-Pine Forest) being the Kuchler equivalent listed thereunder, SAF 88 (Willow Oak-Water Oak-Diamondleaf [Laurel] Oak), Mixed hardwood Series (223.13), Southeastern Swamp and Riparian Forest (223.1) of Brown et al (1998, p. 43) and FRES No.41 (Wet Grasslands Ecosystem) with K-70 (Southern Cordgrass Prairie) the Kuchler equivalent thereunder, SRM 711 X 807, Southeastern Interior Marshland (243.1), Warm Temperate Marshland of Brown et al. (1998, p. 45). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

26. Inside an east Texas transition zone- Interior views of a savannah formed by the ecotone of a wet flatwoods made up of water and willow oak (dominants), with some overcup oak and laurel oak, loblolly pine, green ash, and black tupelo and of freshwater marsh composed of cottongrass bulrush, sedges, flatsedges, spikerushes, panicgrasses, paspalums, bentawn plumegrass, bushy and broomsedge bluestem (beardgrasss), and common cattail. Most of the trees in the first photograph were water oak; trunks of trees at far right margin of second photograp were overcup oak (Quercus lyrata).Immediately to left of the overcup oak (near center foreground) were shoots of bushy bluestem (closesst to overcup oak) and broomsedge bluestem (to left of the bushy beardgrass). Conspicuous green clumps were mostly cottongrass bulrush. Hardin County, Texas. February, late hibernal aspect. This ecotone was a combination of : FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem) with K-101 (Oak-Hickory-Pine Forest) being the Kuchler equivalent listed thereunder, SAF 88 (Willow Oak-Water Oak-Diamondleaf [Laurel] Oak), Mixed hardwood Series (223.13), Southeastern Swamp and Riparian Forest (223.1) of Brown et al (1998, p. 43) and FRES No.41 (Wet Grasslands Ecosystem) with K-70 (Southern Cordgrass Prairie) the Kuchler equivalent thereunder, SRM 711 X 807, Southeastern Interior Marshland (243.1), Warm Temperate Marshland of Brown et al. (1998, p. 45). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

27. Pool in the Pineywoods X Gulf Coastal marsh ecotone- At spatial scale of local site (in this instance, a larger microhabitat or microsite) a pool in the transition zone between hardwoods-pine flats and freshwater coastal marsh supported an array of aquatic plants from bulrushes and sedges of the Cyperaceae to common cattail (Typha latifolia) of Typhaceae. In the first slide longleaf pondweed (Potamogeton nodosus) was floating on the pool surface. Trees surrounding pool were water and willow oaks. Hardin County, Texas. February, hibernal aspect. Range vegetation of this locale was described in captions of preceding slides.

Loblolly Pine (Pinus taeda) Forest

Loblolly pine is the mainstay of the forest products in the southeastern forest of North America. Loblolly pine (and hybrids thereof) is the single most important species across this general region though, of course, other species including pines are the major lumber/pulp species in portions of the southeastern forest region (eg. slash pine in peninsular Florida). The classic though dated reference for loblolly pine probably is still the monograph by Wahlenberg (1960) with the work of Schultz (1997) a fine successor.

Organization note: this section is a "sampler" of loblolly pine. Complete coverage of loblolly pine forest range was shown in its own chapter, Loblolly Pine Forests.

28. Trunk of loblolly pine- This is the typical pattern and color of bark on this the largest of the four major pine species in the Southern Pine Region. Appropriately the bark bears the burnished coloration of past cool surface fires. A Virginia creeper or woodbine (Parthenocissus quinquefolia) with newly emerged leaves was climbing the attractive trunk. Crocket National Forest, Houston County, March.

29. Cones of loblolly pine- Relative size, shape, and unique gross features of loblolly pine were presented in these two views of two cones of the dominant pine over much of the southeastern portion of the deciduous forest of North America in cluding the pineywoods of east Texas and western Louisiana. The reproductive (sexual) organs of conifers are borne on a woody or fleshy (berry-like) structure that is designated a strobulus (strobili is the plural). These stroboli are called cones by foresters and most regular folk other than "real" botanists. Conifers produce separate male (staminate) and female (ovulate) cones so that this group of gymnosperms are either diocecious or monecious. Moneciousness is the general arrangement for most genera in Pinaceae (Pinus, Abies, Picea, Tsuga, Pseudotsuga), Cupressaceae (Cupressus, Thuja, Juniperus), and Taxodiaceae (Taxodium, Sequoia, Sequoiadendron).. Montgomery County, Texas. February (most seed had shattered from the woody ovulatecone).

30. Seed in a cone- Two views of a seed in the ovulate cone of loblolly cone. A seed of loblolly pine was shown without a marker in the first slide whereas the base of a fasicle of loblolly pine marked one seed in the second slide slide.The brown, parchment-resembling wing of this single seed (one of a pair) was cut away to more clearly reveal the dry seed. Most other seeds, including the other one of this cone unit, had shattered (been shed) from this cone. The cone of conifers in the Pinaceae is a compound woody structure comprised of numerous units on which the seed, with its attendant parts, develops and is borne while developing before being shed. Each woody unit of the ovulate cone is is a primary appendage-- a woody bract-- that is called the ovuliferous scale. This woody scale is the ovule/seed-bearing part of the cone (strobolus). Typically a pair of ovules, each of which develops into a dry seed (complete with a winged part for wind dispersal), form on the adaxial (= upper) surface of each woody ovuliferous scale resulting in formation of two seed scars on this upper surface (in the axil where scale joins the central woody axis of the cone. Montgomery County, Texas. February.

31. Core and treasure of the cone- Another view of a seed near apex of cone of loblolly pine. Then two views of a pair of loblolly pine seeds on adaxial surface of ovuliferous scale (ie. in axillary area where woody scale attached to central shaft of the cone). In the first of these two photographs the wing on the left seed of the pair was twisted far to the right for better viewing of the wing, but the wing was in from of (covering up) the right seed. In the second photograph the wing of both seeds of this pair had been removed to more clearly reveal the two seeds. Montgomery County, Texas. February.

32. Future loblolly pines- Unshattered loblolly pine seeds taken from the cones shown above. Some seeds still had their wings attached whereas others were missing these wind-dispersal facilitting organs. Steel measure indicated the size of seed and wings. The two seeds with attached, intact wings were a pair attached on the same woody ovuliferous scale. Montgomery County, Texas. February.

33. Up-close look- Seeds of loblolly pine were shown at closer distance to present details of seed coat and texture of the fragile woody material making up wings. Pines are examples of seed dispersal by wind, anemochory (= aerochory), in which wings facilitate action of wind as an agent or facilitator of sexual reproduction. Wind also operates as facilitator during pollination. Montgomery County, Texas. February.

34. Loblolly pine forest- Second (or "third", "fourth", etc.)- growth, but natural revegetation with an open understory dominated by little bluestem. Associated understory herbs include slender-leaf wood oats (Uniola sessiliflora) plus species ofPaspalum, Panicum, and Sporobolus among grasses plus native legumes like tickclover (Desmodium spp.) and numerous composites. Davy Crockett National Forest, Houston County, Texas. March, vernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem). Variant of K-101 (Oak-Hickory-Pine Forest): the southern pine types have traditionally been interpreted as sub-climax fire-types and this seral stage is maintained by foresters in order to produce the more valuable pine wood products. SAF 81 (Loblolly Pine). Oak Pine Series of Brown et al. (1998). This type is clearly transitory forest range with total loss of understory as pines approach maturity as was shown below. South Central Plains- Southern Tertiary Uplands Ecoregion, 35e (Griffith et al., 2004).

35. Going into a loblolly pine pine- Stucture and botanical composition of a second-growth loblolly pine forest showing an herbaceous layer dominated by little bluestem followed by longleaf woodoats then beaked panicgrass with two annual panicgrass species sometimes locally functioning as associates, a tall shrub layer represented by flowering dogwood in full-flower, and a lower shrub layer (in this vegetation) of which yaupon holly was the major species. The two annual panicgrasses were warty panicgrass (Panicum verrucosum) and savanna panicgrass (P. gymnocarpon). Fire-scourched bark attested to use of prescribed or, at least, convenience burning in a Pineywoods oak-pine forest that was maintained primarily as a loblolly pine stand. Davy Crockett National Forest, Houston County, Texas. March, vernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem). Variant of K-101 (Oak-Hickory-Pine Forest): the southern pine types have traditionally been interpreted as sub-climax fire-types and this seral stage is maintained by foresters in order to produce the more valuable pine wood products. SAF 81 (Loblolly Pine). Oak Pine Series of Brown et al. (1998). This type is clearly transitory forest range with total loss of understory as pines approach maturity as was shown below. South Central Plains- Southern Tertiary Uplands Ecoregion, 35e (Griffith et al., 2004).

36.Transitory loblolly pine forest range- Loblolly pine forest range at either: 1) late seral stage with loblolly pine maintained in a hardwood (mostly an oak)-pine forest by frequent burning or 2) a mid-stage (more-or-less) of a wood crop in a thin (sparsely or weakly stocked) stand of loblolly pine. Either way there was low stocking of loblolly pine, the tree crop species, and a well-developed, high-yielding (by loblolly pine range standards) herbaceous understorey for grazing livestock and/or wildlife. This was the same stand of loblolly pine forest vegetation as introduced in the immediately preceding photograph. Dominant grass in this "photo-plot" of that loblolly pine forest range was little bluestem with longleaf woodoats, beaked panicgrass, savanna panicgrass, and warty panicgrass also present. Foremost tree (slightly to right of a conspicuous pine trunk) was sweetgum (Liquidambar styraciflua). A young sassafras (Sassafras albidum) with just-beginning-to-open leaves was growing at left margin of photograph. Yaupon of sundry sizes was widespread throughout the woody layers. Davy Crockett National Forest, Houston County, Texas. March, vernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem). Variant of K-101 (Oak-Hickory-Pine Forest): the southern pine types have traditionally been interpreted as sub-climax fire-types and this seral stage is maintained by foresters in order to produce the more valuable pine wood products. SAF 81 (Loblolly Pine). Oak Pine Series of Brown et al. (1998). This type is clearly transitory forest range with total loss of understory as pines approach maturity as was shown below. South Central Plains- Southern Tertiary Uplands Ecoregion, 35e (Griffith et al., 2004).

37. Growing doghair- In the same understorey as presented in the last two preceding photographs and captions there were some very localized (restricted) areas near mature loblolly pine trees on which there were extremely dense patches of pine seedings. Obviously not all of these seedlings could survive, but it was also obvious that these would develop into proverbial "doghair stands" unless something thinned them out. If natural agents of mortality such as fire and disease did not eliminate some of these woefully overstocked little blessings of Mother Nature then Man the Forester would have to intervene in order to achieve efficient management of forest resources. One such "doghair stand" of loblolly pine seedlings was presented in foreground of this photograph. Various Panicum species (major ones were listed in the preceding caption) were the major grasses on this "photo-plot". Davy Crockett National Forest, Houston County, Texas. March, vernal aspect. Leaves on hardwood species (such as one in left foreground) were just emerging from buds. FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem). Variant of K-101 (Oak-Hickory-Pine Forest): the southern pine types have traditionally been interpreted as sub-climax fire-types and this seral stage is maintained by foresters in order to produce the more valuable pine wood products. SAF 81 (Loblolly Pine). Oak Pine Series of Brown et al. (1998). This type is clearly transitory forest range with total loss of understory as pines approach maturity as was shown below. South Central Plains- Southern Tertiary Uplands Ecoregion, 35e (Griffith et al., 2004).

38. A crop of poles- Closed canopy stand of loblolly pine with no herbaceous (grazable) understorey and lower woody layers limited to Tolerant flowering dogwood and yaupon. As a wood crop this single-species stand of loblolly pine was approaching (within a few years of) maturity. This plantation stand was an example of industrial foresty. On this commercial forest the wood crop was a monoculture of fast-growing, hybrid loblolly pine. Such loblolly pine plantations are a form of even-aged silviculture (silvicultural system). This crop will be harvested in a few years by clearcutting, "a regeneration or harvest method that removes essentially all trees in a stand" (Helms, 1998). Harvest will result in release of many species of grasses, grasslike plants, forbs, shrubs, and Intolerant tree species like sweetgum. Several of the grass species such as those presented in preceding slides will persist for a number of years and through mid-sere of secondary plant succession. Tree species like sweetgum and numerous oaks and hickories will persist unless eliminated by application of selective herbicides or reduced by commercial livestock grazing and/or prescribed burning. Combinations of these silvicultural treatments may be used (as shown periodically throughout this chapter). This and preceding slides of loblolly pine forest vegetation illustrated transitory forest range that is typical of forest cropping systems on commercial (industrial) forests throughout much of southeastern North America. Davy Crockett National Forest, Houston County, Texas. March, early vernal aspect (time of the dogwood-redbud tours). This was an anthropogenic variant of the following vegetational units. FRES No. 14 (Oak-Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 82 (Loblolly Pine-Hardwood). Biotic community unit of Brown et al. (1998, ps. Oak-Pine Series, 1212.14 of Northeastern Deciduous Forest 122.1 except that there should have been an Oak-Pine Series, say 123.13, os Southeastern Deciduous and Evergree Forest 123.1.South Central Plains- Southern Tertiary Uplands Ecoregion, 35e (Griffith et al., 2004). Another interpretation of this highly human-modified (= non-natural) forest vegetation was FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem). The Kuchler designation would still be K-101 (Oak-Hickory-Pine Forest) while the Society of American Foresters (Eyre, 1980) would be a man-made forest cover type of Loblolly Pine (SAF81).

Clarification of terms: the following concepts and definitions were provided from The Dictionary of Forestry developed by the Society of American Foresters (Helms, 1998) to assist students in understanding management and production of grazing and/or browsing resources on transitory forest range. Silvicultural system- a planned series of treatments for tending, harvesting, and re-establishing a stand. Regeneration method- a cutting procedure by which a new age class is created. Even-aged regeneration methods regenerate and maintain a stand with a single age class. One even-aged method is clearcutting which is the cutting of essentially all trees, producing a fully exposed microclimate for the development of a new age class (by either natural or artificial re-eatablishment of the next generation, crop, of trees)

39. "Will the real Pineywoods please stand up?"- "You bet, and this is it." In contrast to the loblolly pine plantation introduced in the immediately preceding slide (and several used below to illustrate silvicultural methods) here were two views of the natural oak-hickory-loblolly pine forest vegetation. This was a second-growth forest recovering from the heady days of "cut-and-run" heady logging, but it had the structure (including several layers of vegetation) and species composition of the native mixed hardwood-loblolly pine. Flowering dogwood and lesser cover of redbud hearlded the early days of spring in this sandy land (note road) upland Pineywoods forest. Not exactly a lobolly pine plantation as shown immediately above and farther below. Water oak, accompanied adult and sapling to pole-size loblolly pine. This forest consisted essentially of the species compoisition indicative of the climax vegetation except that following initial frontier, destructive, non-scientific logging; overgrazing (including by free-ranging, mast and root-eating hogs); and underburning (more likely, total fire exclusion) there was only limited herbaceous understorey (mostly of bluestem, panicgrass, and paspalum species along with some sedges and flatsedges) Davy Crockett National Forest, Houston County, Texas. March, early vernal aspect (time of the dogwood-redbud tours). FRES No. 14 (Oak-Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 82 (Loblolly Pine-Hardwood). Biotic community unit of Brown et al. (1998, ps. Oak-Pine Series, 1212.14 of Northeastern Deciduous Forest 122.1 except that there should have been an Oak-Pine Series, say 123.13, os Southeastern Deciduous and Evergree Forest 123.1.South Central Plains- Southern Tertiary Uplands Ecoregion, 35e (Griffith et al., 2004).

40. Open understory(= permanent forest range) loblolly pine forest- This loblolly forest is growing immediately above the first terrace of the Sabine River. Recurrent fire (note small fire scar on base of first tree on the left) after establishment of loblolly pines maintained an open understory predominately of perennial wiregrass or threeawn species like woolyleaf threeawn (Aristida lanosa), longspike or slimspike threeawn (A. longespica), and purple or arrowfeather threeawn (A. purpurascens) with broomsedge bluestem (Andropogon virginicus) and splitbeard bluestem (A. ternarius) as associates. Pioneer annual composites like giant ragweed (Ambrosia trifida) and horseweed (Conyza canadensis= Erigeron canadensis) are also present, but as last year’s weathered-down stalks. Flowering dogwood (which is at peak bloom in this view) dominates the upper shrub layer with yaupon (green shrub beneath the blooming dogwood) as dominant of the lower shrub layer.   Under the current fire regime this is permaent loblolly pine forest range. It stands in contrast to the transitory loblolly pine forest range where there is denser tree stocking combined with discontinued use of prescribed burning to produce pulp wood instead of pine lumber as in the forest seen here. This range forest cover type persist as a result of the disturbance of repeated fire which kept out hardwood tree species like water and white oaks.The fire disturbance also made favorable habitat for old-field pioneer species such as the two weedy composites and annual threeawns like old-field threeawn (Aristida oligantha plus the similar A. desmantha) and churchmouse threeawn (A. dichotoma). The physiogonomy is that of climax Pineywoods but the absence of hardwood trees and the species composition of the herbaceous understory is clearly that of late seral (= subclimax) forest vegetation. It is a textbook example of “pine woods wiregrass range”. Sabine River, Harrison County, Texas. Vernal aspect, March. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem), man-modified variant of K-101 (Oak-Hickory-Pine Forest), SAF 81 (Loblolly Pine). Brown et al. (1998) Oak-Pine Series converted into Pine Series by human management. South Central Plains- Southern Tertiary Uplands Ecoregion, 35e (Griffith et al., 2004).

41. Climax Loblolly Pine-Hardwood-Pinehill Bluestem Pineywoods Forest- Although not old-growth forest, this is a classic composite Pineywoods community with the species composition of the virgin vegetation. Loblolly pine is the major dominant thereby establishing this as the loblolly pine form of the Pineywoods Complex, but water oak is co-dominant while sweetgum (Liquidambar styraciflua) is well-represented among the trees. The shrub layer is dominated by wax myrtle (Myrica cerifera). The herbaceous understory is remarkably diverse. Scattered small colonies of the rhizomatous decreaser pinehill bluestem (Andropogon divergens) serve as an indicator of what the climax dominant for this site should be. Other grasses include splitbeard bluestem, low or spreading panicgrass (Panicum rhizomatum),  longleaf uniola (Uniola sessiliflora), and Florida paspalum (Paspalum floridanum). Several Carex species are present with considerable cover. The conspicuous graminoid in the immediate foreground is a species of bulrush (Scirpus sp.).     This is a bottomland loblolly pine-hardwood forest on the flood plain of the Sabine River. It is the forest vegetation just below that seen in the previous slide. It is less apt to burn and has a more favorable soil moisture regime than the previous forest range type. Recent and recurrent fire had to have been part of the environment however to maintain this open understory and the fire-adapted grasses. Rather than wiregrass loblolly pine forest range this is the pinehill bluestem-Florida paspalum-low panicum understory. It is produces much higher quality and higher yielding forage. Both the bluestem and wiregrass Pineywoods range types are permanent forest range with a persistent grazable understory.This is climax loblolly pine-oak hardwood forest while the previous plant community was seral loblolly pine forest vegetation. Sabine River bottom, Harrison County, Texas. Vernal aspect, March. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem), K-101 (Oak-Hickory-Pine Forest), SAF 82 (Loblolly Pine-Hardwood). Anthropogenic Pine Series of Brown et al. (1998). South Central Plains- Southern Tertiary Uplands Ecoregion, 35e (Griffith et al., 2004).

42. Big Thicket loblolly pine woods- Exterior view of a loblolly pine-water oak subtype or variant of loblolly pine-hardwood cover type shjowing physiogonomy and botanical composition. At this edge (in a forest opening) young water oak, yaupon holly, and various herbaceous species and the layers they comprise provided an unusual composite "Big Picture" perspective of this forest range vegetation. The small oaks (pole-size) at base of pines were water oak. These young trees had not shed their dead leaves and even had a few persistent live leaves. Yaupon or yaupon holly composed almost all of the lower woody layer except for the regenerting oaks. Grasses were bushy and broomsedge bluestems and bentawn plumegrass. Liberty County, Texas. February, late hibernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem), K-101 (Oak-Hickory-Pine Forest), SAF 82 (Loblolly Pine-Hardwood). Given published biotic communities in the system of Brown et al. (1998) the closest designation would have to be interpreted as the Pine Series (123.12) of Southeastern Deciduous and Evergreen Forest although the Oak-Pine Series (122.14) of Northeastern Deciduous Forest (122.1) is closest in name. There should be a Brown et al. (1998) biotic community of Hardwood-Pine Series under Southeastern Deciduous and Evergreen Forest. South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

43. Edge of a loblolly pine-water oak forest- Detail of herbaceous and lower woody layers of the forest-clearing edge shown in the immediately preceding slide. Regenerated water oak saplings and yaupon made up a lower woody layer. Bentawn plumegrass and broomsedge bluestem and bushy beardgrass were the major herbaceous species. Pine seedlings bore witness to regeneration of loblolly pine. Liberty County, Texas. February, late hibernal aspect. Designations of this forest range vegetation were given in the preceding caption.

44. Inside with the pines and oaks- Interior of the loblolly pine-water oak forest introduced in the two preceding slides and their captions. American holly (Ilex opaca) was an associate tree species. A few individuals of regenerating American holly and dense yaupon holly made up most of the lower or secondary woody layer. Adequate light on the forest floor at this stage of vegetation development permitted regeneration of pine as well as persistence of an herbaceous understorey consisting of two or three layers. In the interior of this forest community that was more shaded or, same thing, less well-lite than the forest edges shown above, the dominant herbaceous plant was longleaf woodoats (Uniola sessiflora) which "bumped out" the bluestem or beardgrass species and bentawn plumegrass. Other common to locally dominant herbaceous species included cottongrass bulrush, green flatsedge, beaked panicgrass (Panicum anceps), redtop panicgrass (P. rigidulum), and brownseed paspalum (Paspalum plicatluum). These herbaceous species grew together on local habitats (microhabitats at about largest spatial scale) within this loblolly pine-water oak-American holly forest community. Most of these were not visible at scale of these two photographs, but they were featured below at scale of both herbaceous layers and individual plants. In the first of these two slides water oak and loblolly pine were visible as distinctive trees. The largest tree (left-of-center midground) was water oak. In the second of these photographs water oaks were relegated to midground and surrounded by loblolly pines. Yaupon and small saplings of American holly were widespread in the lower woody layer. Liberty County, Texas. February, late hibernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem), K-101 (Oak-Hickory-Pine Forest), SAF 82 (Loblolly Pine-Hardwood). Given published biotic communities in the system of Brown et al. (1998) the closest designation would have to be interpreted as the Pine Series (123.12) of Southeastern Deciduous and Evergreen Forest although the Oak-Pine Series (122.14) of Northeastern Deciduous Forest (122.1) is closest in name. There should be a Brown et al. (1998) biotic community of Hardwood-Pine Series under Southeastern Deciduous and Evergreen Forest. South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

45. Up close in the interior- Interior structure and species composition of a Big Thicket Pineywoods second-growth forest dominated by trees of loblolly pine, water oak, and American holly. Yaupon and small saplings of American holly made up bulk of lower woody layer(s). There was no reproduction of loblolly pine in the denser locale of the forest. Neither was there presence of herbaceous species. Foremost trunk (left foreground) and four pole-size trunks were loblolly pine. Largest tree with straight bole (left midground) was water oak as were the two small saplings still with green leaves growing between the foremost loblolly pine and the large water oak. Liberty County, Texas. February, late hibernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem), K-101 (Oak-Hickory-Pine Forest), SAF 82 (Loblolly Pine-Hardwood). Given published biotic communities in the system of Brown et al. (1998) the closest designation would have to be interpreted as the Pine Series (123.12) of Southeastern Deciduous and Evergreen Forest although the Oak-Pine Series (122.14) of Northeastern Deciduous Forest (122.1) is closest in name. There should be a Brown et al. (1998) biotic community of Hardwood-Pine Series under Southeastern Deciduous and Evergreen Forest. South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

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46. Loblolly pine-water oak-American holly subtype of Big Thicket Pineywoods- All-in-one shot of an example of this variant of a loblolly pine-hardwood forest cover type with yaupon and regenerating American holly comprising a lower woody layer and a local opening with longleaf woodoats and other herbaceous species. There was also much reproduction of water oak with numerous small saplings that had retained many of their leaves throughout the usually mild winter of the Big Thicket. Longleaf woodoats and local stands of other herbaceous species were also present though widely scattered. All of the larger trees in midground were water oak except for one loblolly pine. This was an example that the climatic climax of these forest type is oak and not pine. Both cover types of Loblolly Pine (SAF 81) and Loblolly Pine-Hardwood (SAF 82) (Eyre, 1980) are fire types (ie. only under conditions where fire overrides other components of climate) does this forest vegetation have loblolly pine as a dominant or co-dominant species into climax stage. Liberty County, Texas. February, late hibernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem), K-101 (Oak-Hickory-Pine Forest), SAF 82 (Loblolly Pine-Hardwood). Given published biotic communities in the system of Brown et al. (1998) the closest designation would have to be interpreted as the Pine Series (123.12) of Southeastern Deciduous and Evergreen Forest although the Oak-Pine Series (122.14) of Northeastern Deciduous Forest (122.1) is closest in name. There should be a Brown et al. (1998) biotic community of Hardwood-Pine Series under Southeastern Deciduous and Evergreen Forest. South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

47. Longleaf woodoats (Uniola sessiflora)- Five plants of longleaf woodoats adorned the floor of a loblolly pine-water oak-American holly variant of the Loblolly Pine-Hardwood forest cover type (SAF 82). Liberty County, Texas. February.

48. In a small clearing- A small clearing provided adequate light for local stands of herbaceous vegetation, woody layers made up of yaupon and regenerated water oak. At edge of clearing the woody shoot of rattan or Alabama supplejack was climbing a young loblolly pine (small pole in right midground). Species composition of the herbaceous layers was presented in the next slide. Liberty County, Texas. February, late hibernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem), K-101 (Oak-Hickory-Pine Forest), SAF 82 (Loblolly Pine-Hardwood). Given published biotic communities in the system of Brown et al. (1998) the closest designation would have to be interpreted as the Pine Series (123.12) of Southeastern Deciduous and Evergreen Forest although the Oak-Pine Series (122.14) of Northeastern Deciduous Forest (122.1) is closest in name. There should be a Brown et al. (1998) biotic community of Hardwood-Pine Series under Southeastern Deciduous and Evergreen Forest. South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

49. Something to graze- Two views of a local stand of herbaceous species in the understorey of the Pineywoods loblolly pine-water oak-American holly flatwoods forest shown and described immediately above. Largest and most common species was cottongrass bulrush followed by green flatsedge, redtop panicgrass, and longleaf woodoats. The latter species more commonly grows by itself as shown above, but it frequently grows in association with other grasses and grasslike plants. Individual plants of the cottongrass bulrush and green flatsedge introduced in these two photographs were shown in more detail in the succeeding four slides. Liberty County, Texas. February, late hibernal aspect.

50. Cottongrass bulrush, wooly-grass bulrush, or wool-grass (Scirpus cyperinus)- This was one of the most common and largest species of grasslike plants in much of the wetter environments of the loblolly pine-hardwood forest cover type in the Big Thicket area of east Texas. Cottongrass bulrush was especially common (and locally dominant) on mesic to wet forest and range sites such as those for loblolly pine and the various forms of "oak flats" or "flatwoods". The individual plant presented in this and the two slides above the next caption was one of several specimens growing on the loblolly pine-water oak-American holly forest community featured above Liberty County, Texas. February.

51. Inflorescence of cottongrass bulrush- Two panicles on the same individual plant of cottongrass bulrush introduced in the immediately preceding photograph (one of several such bulrush plants shown in the two photographs preceding that last slide). More recent taxonomic treatments of the Scirpus species such as that by Diggs et al. (2006) have rearranged many of these once-upon-a-time Scirpus members into such genera as Schoenoplectus, Bolboschoenus, Isolepis, and even Eleocharis! Interestingly, in this bulrush basket upset cottongrass bulrush remained as Scirpus cyperinus (ie. still a true bulrush as it were). Liberty County, Texas. February, late hibernal aspect.

52. Weathered flat- A highly weathered panicle of green flatsedge (Cyperus virens) in the local stand of herbaceous vegetation in the Pineywoods loblolly pine-water oak-American holly flatwoods forest described herein. In the Illustrated Flora of East Texas Diggs et al. (2006) showed and mapped 19 species of flatsedge as occurring in the small Texas county of Liberty. Green flatsedge is one of the more common Cyperus species growing within the Big Thicket of the Texas Pineywoods. Liberty County, Texas. February.

53. Plumes in the pines- Panicles of bentawn plumegrass (Erianthus contortus) growing on the loblolly pine-water oak-American holly flatwoods forest featured here. The Illustrated Flora of East Texas (Diggs et al., 2006) put Erianthus in with an expanded sugarcane genus (Saccharum spp.) and renamed bentawn plumegrass S. brevibarbe var. contortum. Diggs et al. (2006) listed and mapped four former Erianthus species as being in the Pineywoods, including Liberty County, Texas (county where these photographs were taken). Correll and Johnston (1979) in Manual of the Vascular Plant of Texas, still the statewide "bible" of plant taxa, listed three Erianthus species for this "neck of the woods". Erianthus species are some of the largest grasses native to the Pineywoods. E. contortus is probably the most common of these. Liberty County, Texas. February, hard-grain/shatter stage of phenology.

54. Beauty in the little things- Closeups of spikelets in the panicles of bentawn plumegrass shown immediately above. Liberty county, Texas. February, hard grains were shattering rapidly (lucky photographer preserved this lively scene for generations of grassmen).

55. Holly in the stand- Deeper inside the loblolly pine-water oak-American holly stand of flatwoods featured herein (and shown repeatedly above) American holly was becoming the local dominant tree species. Shrub-sized woody plants with persistent green leaves (foreground understorey) included young yaupon, water oak saplings, and American holly saplings and seedlings. Whereas tolerance rating for loblolly pine is Intolerant (Fowells, 1965, p. 366; Wenger, 1984; Burns and Honkala, 1990, Vol 1, p. 505) and water oak is Intolerant (Fowells, 1965, p. 630; Burns and Honkala, 1990, Vol. 2, p. 703) , American holly is Very Tolerant which is the same as American beech, sugar maple, and flowering dogwood (Wenger, 1984). In absence of fire, windthrow, logging, and other disturbances (natural or anthropogenic) American holly (or other Very Tolerant species in the Big Thicket like American beech and southern magnolia) eventually become dominant as development of forest vegetation reaches climax. That phenomenon was shown in this photograph where the only seedlings present were those of American holly. The largest and foremost trunk (right foreground; smooth bark) was a nice, shaply specimen of American holly. This specific plant was used below to describe its species, including another view of this same bole. The medium-sized liana growing up the American holly was some species of grape (Vitis sp.). Liberty County, Texas. February, late hibernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem), K-101 (Oak-Hickory-Pine Forest), SAF 82 (Loblolly Pine-Hardwood). Given published biotic communities in the system of Brown et al. (1998) the closest designation would have to be interpreted as the Pine Series (123.12) of Southeastern Deciduous and Evergreen Forest although the Oak-Pine Series (122.14) of Northeastern Deciduous Forest (122.1) is closest in name. There should be a Brown et al. (1998) biotic community of Hardwood-Pine Series under Southeastern Deciduous and Evergreen Forest. South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

56. "Hollyer" Pineywoods- On one local part of the loblolly pine-water oak-American holly dominated stand shown and described above, American holly was becoming the climax dominant tree species. Saplings of American holly were more common than those of water oak and the only seedlings were those of American holly. There were no saplings (only pole-size trees) of loblolly pine, the tree which clearly dominated the canopy (with water oak as the local associate species) at this subclimax stage. Yaupon, the overall community dominant of a lower woody layer, was common and shared this layer with saplings of water oak and both saplings and seedlings of American holly as was just remarked. The adult American holly was the same individual whose trunk (with grape vine) was featured in the slide immediately above. Liberty County, Texas. February, late hibernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem), K-101 (Oak-Hickory-Pine Forest), SAF 82 (Loblolly Pine-Hardwood). Given published biotic communities in the system of Brown et al. (1998) the closest designation would have to be interpreted as the Pine Series (123.12) of Southeastern Deciduous and Evergreen Forest although the Oak-Pine Series (122.14) of Northeastern Deciduous Forest (122.1) is closest in name. There should be a Brown et al. (1998) biotic community of Hardwood-Pine Series under Southeastern Deciduous and Evergreen Forest. South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

57. Happy holly happenings- Regeneration (from seed) of American holly on floor of a subclimax loblolly pine-water oak forest. A young sapling of American holly represented the future generation of the climax dominant of this Pineywoods forest range. Yaupon holly, an individual of which was behind the featured holly sapling, was the overall dominant of a lower woody layer. American holly, a Very Tolerant tree species, had successfully reproduced even with this competition. The only herbaceous species was longleaf woodoats. Another photograph of a small American holly sapling was presented below to show details of holly leaves. The was the same stand of loblolly pine-water oak-American holly-yaupon-longleaf woodoats-bulrush-flatsedge Big Thicket flatwoods forest as featured above. Liberty County, Texas. February, late hibernal aspect.

58. Dynasty in the making- This large, symetrical American holly and its progeny (ranging from seedlings to small saplings) were the climax tree species and future dominant of a loblolly pin-water oak-American holly-yaupon holly- longleaf woodoats-bulrush forest range community. American holly is regarded as Very Tolerant (Wenger, 1984) so that in absence of disturbances (eg. repeated forest fires, forest harvests) the Intolerant loblolly pine (Fowells, 1965, p. 366; Wenger, 1984; Burns and Honkala, 1990, Vol 1, p. 505) and Intolerant water oak (Fowells, 1965, p. 630; Burns and Honkala, 1990, Vol. 2, p. 703) will be succeeded by American holly. Successful regeneration of holly and absence of reproduction bf loblolly pine was shown in these two photographs. Numerous young plants of yaupon were also present. Besides showing regeneration of American holly, these two slides provided views of a mature American holly showing crown shape and branching pattern typical of this climax hardwood (angiosperm) species. Note that limbs and branches of the holly extended lower on the crown than did those of surrounding loblolly pine and water oak and that, overall, the crown of the holly was substantially larger and fuller than those of the two current dominants of the subclimax flatwoods forest. Liberty County, Texas. February.

59. Heir-apparent up close- Trunk of a mature American holly, the hardwood species that was successionally ascending to climax dominance with successful reproduction in shade of the loblolly pine-water oak-American holly Pineywoods forest. Another view of this same trunk was presented above (complete with the same grape vine). Extent of shading of forest floor was also typical and indicative of a lower-layer forest environment in which only Toleranmt and Very Tolerant species could regenerate. Liberty County, Texas. February.

60. Tale of trauma with a position to teach- For reason(s) unknow to the photographer this sapling of American holly was leaning to the stage of being nearly "flat".In spite of such posture the young tree was otherwise "alive and well". The first slide of this sapling provided a better composite view than if it had been vertical like it should have been. The second photograph was a view of branches and leaves on this flattened sapling. Spiney margins of holly leaves were presented to rangemen and foresters unfamilar with this species. Liberty County, Texas. February.

61. Shinny holly leaves against a background of pine straw- Another small sapling (or large seedling) of American holly (Ilex opaca) growing in shade of loblolly pine, water oak, and its parent which was the large, symmetrical adult holly introduced above. Another living bit of evidence as to the Very Tolerant rating of American holly as well as an example of leaves of this species. Leaves of American holly are vriable in shape, but the margins have spines and are usually scalloped. Leaf blades are shiny green "on top" (upper surface) and pale underneath. This small tree was in the same flatwoods forest stand as shown and described above. Liberty County, Texas. February.

62. Yaupon or yaupon holly (Ilex vomitoria)- Yaupon is one of the most aggressive shrubs in the understorey of the various pine and oak-pine types of the eastern deciduous forest in southeastern North America. It is usually not a dominant shrub in advanced stages of forest succession, but disturbances like logging (and subsequent regeneration methods) shift competitive advantage to this rapidly spreading, evergreen scrub holly allowing it to become a major brush species on regenerating forests and transitory forest ranges. The individual shown here had persisted late into the rotation of a loblolly pine forest. International Paper Company, Harrison County, Texas. May.

10247 and 10248.

Girls' time- Flowers at peak-bloom stage on a female yaupon. Yaupon is a dioecious species. Details of a female plant in full bloom were presented. International Paper Company, Harrison County, Texas.April.

63.Yaupon leader- Leaves and fruits (drupes) of yaupon. Yaupon is dioecious. The drupes are a favored food of numerous species of songbirds and even furbearers like coons. Browse value of yaupon is often rated as good for deer and fair for livestock though some dispute this. Heavy livestock grazing early in the forest rotation is often an effective means of yaupon control. International Paper Company, Harrison County, Texas. January.

64. Longleaf wood oats (Uniola sessiliflora)- This is one of the more common and important grass species in the shortleaf pine and pine-oak forests. It responds quickly with vigorous growth and reproduction to clearcutting and thinning operations in these forest cover types. Red River County, Texas. July.

65. Loblolly pine flatwoods- Example of the "pure" cover type of loblolly pine (SAF 82) made up this flatwoods forest community in the Big Thicket. Young, second-growth loblolly pines comprised the entire canopy (crown) layer while yaupon made up the lower woody layer and giant ragweed (Ambrosia trifida) comprised most of the herbaceous layer. Perennial grasses such as longleaf woodoats and grasslike plants like cottongrass bulrush and green flatsedge which were common on adjacent and close proximity forests of loblolly pine-hardwood cover type (SAF 88) were much less dense on this forest range stocked with younger (smaller) trees. On local areas most disturbed by forest harvest activities there were some individuals of broomsedge and bushy beardgrass with fewer plants of longleaf woodoats and even density of bentawn plumegrass. While there were widely scattered water oaks stocking of this species was so slight that this forest was a loblolly pine cover type. Selective (uneven-aged) harvest had taken place on this forest three or four years earlier. Hence, pioneer establishment of the giant ragweed and subsequent release of yaupon. Liberty County, Texas. February, late hibernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem), K-101 (Oak-Hickory-Pine Forest), SAF 88 (Loblolly Pine). Biotic community in the system of Brown et al. (1998) would have to be interpreted as the Pine Series (123.12) of Southeastern Deciduous and Evergreen Forest. South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

66. Not sharing much- Stand of young loblolly pine on a flatwoods form of Big Thicket Pineywoods resulted in a nearly exclusionary crop, a single-species stand, with almost complete canopy cover of this shade-intolerant subclimax species. Yaupon had developed into a lower woody layer that was sporadic ranging frrom nearly absent to local heavy cover/dense shoots (as in the second of these two slides). Major grass overall was longleaf woodoats though it was absent from local areas of more extreme disturbance. Broomsedge and bushy bluestem were present (some shoots were visible in these and the preceding photograph) on the more seriously disturbed areas, but their cover and density could be described as "few and far between". Plants of bentawn plumegrass were even more uncommon. Loblolly pines of about all one size (poles) and evidence of high degree of disturbance of soil surface indicated that the clearcutting method of regeneration had been used in silvicultural treatment. This was more obvious int the second of these two photographs. The largest tree in second photograph (right midground) was a water oak that had been spared in the recent clear-cutting operation because it was of no value for pulp or poles (ie. a trash tree). It presence and that of scattered water oak seedlings indicated that this was a subclimax loblolly pine-hardwood forest cover type (SAF 82) maintained by silviculture as a loblolly pine forest type (SAF 88). Liberty County, Texas. February, late hibernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem), K-101 (Oak-Hickory-Pine Forest), SAF 88 (Loblolly Pine). Biotic community in the system of Brown et al. (1998) would have to be interpreted as the Pine Series (123.12) of Southeastern Deciduous and Evergreen Forest. South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

67. Inside the loblolly pine stand- Interior of a single-species stand of young (pole-size) loblolly pine. Yaupon formed a sporadic woody understorey or lower woody (shrub) layer. Otherwise this vegetation was loblolly pine cover type (SAF 82) of an industrial or commercial forest where maximum financial return from the forest resulted from establishment and maintenance of single-species stands (= populations) of loblolly pine. In other words, this was a "rough" form of a loblolly pine planation established by natural regeneration. This form of silviculture (clearcutting method) produced denser stands of loblolly pine with more shade and therefore less herbaceous vegetation, especially less cover and lower density of grasses and grasslike plants, for grazing animals. Larger-size clearings and greater disturbance with more bare soil did, however, create better habitat for pioneer species like giant ragweed and this resulted in superior habitat for some kinds of wildlife like bobwhite quail (Colinus virginianus). Presence of saplings of water oak was proof that this was a loblolly pine-hardwood cover type (SAF 88) maintained as the more "pure" loblolly pine forest type (SAF 82) as an economic forest crop (ie. pine wood was more valuable than that from oak in the current market). This loblolly pine stand was in close proximity to the loblolly pine-water oak-American holly forest displayed and discussed above. Yaupon was common and formed a lower woody layer. Dwarf palmetto was also present though mostly as isolated plants. There was less longleaf woodoats, broomsedge and bushy bluestem, bentawn plumegrass, cottongrass bulrush, and sedges and flatsedges than on the nearby loblolly pine-hardwood (water oak and American holly) forest. The most common herbaceous plant on this recently harvestly forest was giant ragweed. Liberty County, Texas. February, later hibernal aspect. Classification units of this forest range vegetation were presented in the two immediately preceding photo captions.

68. Loblolly pine-dominated backwater forest- Another form or subtype(s) of loblolly pine forest range (though one with minimal grazing and browsing resources) in the Pineywoods Region is that (those) that develop on land having ponded water for prolonged periods though not for periods of time consistent with those of swamps. In fact, the common name of loblolly comes from the condition known as a loblolly, a term referring to a mudhole or deep mud puddle, which is an ideal edphic condition for this species (Harlow et al., 1979, p. 93). This is an exterior view of a loblolly pine-dominated forest that developed on a backwater of the San Jacinto River. Hardly visible on disturbed, bare soil in foreground are many pine seedlings indicative of extensive regeneration of the dominant tree species.Bare limbs and branches are those of water oak, overcup or swamp white oak, and black gum (= black tupelo), the associate species. This forest range vegetation would have to be described as a loblolly pine-mixed hardwood-dwarf palmetto forest. It was another form or variant of flatwoods forest. Montgomergy County, Texas. February, later hibernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem), K-101 (Oak-Hickory-Pine Forest), SAF 82 (Loblolly Pine-Hardwood). Given published biotic communities in the system of Brown et al. (1998) the closest designation would have to be interpreted as the Pine Series (123.12) of Southeastern Deciduous and Evergreen Forest although the Oak-Pine Series (122.14) of Northeastern Deciduous Forest (122.1) is closest in name. There should be a Brown et al. (1998) biotic community of Hardwood-Pine Series under Southeastern Deciduous and Evergreen Forest. South While this backwater forest was definitely not a swamp like cypress or tupelo it was at least a seasonal wetland and perhaps should be interpreted as part of Southeastern Swamp and Riparian Forest (223.1) of Brown et al. (1998) in what could be called a Pine-Hardwood Series (of say, number 223.15). Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

69. Next crop of the dominant native and a naturalized alien- At the edge (exterior) of the backwater Pineywoods forest introduced immediately above numerous large seedlings of loblolly (midground) illustrated regeneration by sexual reproduction of this conifer that was dominant in this forest range community. Although both loblolly pine and water oak are rated as Intolerant and subclimax (discussed above for a loblolly pine-water oak-American holly flatwoods forest), on this river backwater wetland these two tree species were climax dominants due to natural protection from severe fire and/or as an edaphic climax. Overcup oak and black tupelo were associate species. The green leaves in background were those of yaupon holly which comprised a lower woody or shrub layer. Dwarf palmetto (none present in this photograph) formed a second shrub lower in height than the yaupon. (These two lower woody layers of vegetation were presented in the immediately succeeding slide.) At local scale there were sapling- and pole size trees of water oak, overcup oak, and blackgum (indicative of regeneration of these hardwood species later than loblolly pine in this sere) that formed a second tree layer. This was not consistent throughout this vegetation. The large cespitose grass in foreground was a specimen of Vaseygrass (Paspalum urvillei), an introduced or agronomic forage species that is now naturalized throughout the Pineywoods of Texas and Louisiana. Over its much of its naturalized range Vaseygrass is a highly productive, welcome addition to the often sparse herbaceous understorey of the Pineywoods region. Vaseygrass was dealt with in the chapter, Introduced Forages, under Grasslands. Montgomergy County, Texas (backwater of San Jacinto River). February, later hibernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem), K-101 (Oak-Hickory-Pine Forest), SAF 82 (Loblolly Pine-Hardwood). Given published biotic communities in the system of Brown et al. (1998) the closest designation would have to be interpreted as the Pine Series (123.12) of Southeastern Deciduous and Evergreen Forest although the Oak-Pine Series (122.14) of Northeastern Deciduous Forest (122.1) is closest in name. There should be a Brown et al. (1998) biotic community of Hardwood-Pine Series under Southeastern Deciduous and Evergreen Forest. South While this backwater forest was definitely not a swamp like cypress or tupelo it was at least a seasonal wetland and perhaps should be interpreted as part of Southeastern Swamp and Riparian Forest (223.1) of Brown et al. (1998) in what could be called a Pine-Hardwood Series (of say, number 223.15). Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

70. Interior of backwater loblolly pine-mixed hardwoods-palmetto flatwoods forest- Inside look at the forest range vegetation introduced in the preceding two photographs. This "photo plot" provided a nearly comprehensive sample of the species composition of Pineywoods pine-hardwoods forest. There was no regeneration of the Intolerant loblolly pine beneath canopy of pine, water oak, overcup oak, and black tupelo in contrast to "doghair" stands of seedlings at edge or exterior of this stand as shown above. Saplings and pole-size trees of water, overcup oak and, to lesser extent, black tupelo were present indicating that these hardwood species had regenerated later than loblolly pine in seral development of this forest vegetation. Younger trees of these woody angiosperms did not form a continuous lower tree layer, but this vegetational strata was frequently present. Dwarf palmetto and yaupon made up two lower shrub layers in this vegetation. Two saplings in foreground were overcup oak (left) and water oak (right). Montgomergy County, Texas (backwater of the San Jacinto River). February, later hibernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem), K-101 (Oak-Hickory-Pine Forest), SAF 82 (Loblolly Pine-Hardwood). Given published biotic communities in the system of Brown et al. (1998) the closest designation would have to be interpreted as the Pine Series (123.12) of Southeastern Deciduous and Evergreen Forest although the Oak-Pine Series (122.14) of Northeastern Deciduous Forest (122.1) is closest in name. There should be a Brown et al. (1998) biotic community of Hardwood-Pine Series under Southeastern Deciduous and Evergreen Forest. South While this backwater forest was definitely not a swamp like cypress or tupelo it was at least a seasonal wetland and perhaps should be interpreted as part of Southeastern Swamp and Riparian Forest (223.1) of Brown et al. (1998) in what could be called a Pine-Hardwood Series (of say, number 223.15). Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

71. Structure and species composition of backwater loblolly pine-mixed hardwoods forest- Two views of lower layers of vegetation in a seasonal wetland forest of loblolly pine, water oak, overcup oak, black tupelo, yaupon, and dwarf palmetto. Large trunk was loblolly pine. On-going regeneration of palmetto was obvious from numerous seedlings of this shrub (eg. in front of pine trunk). Herbaceous species were absent from forest floor which was covered with leaves of tree species. Montgomergy County, Texas (backwater of the San Jacinto River). February, later hibernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem), K-101 (Oak-Hickory-Pine Forest), SAF 82 (Loblolly Pine-Hardwood). Given published biotic communities in the system of Brown et al. (1998) the closest designation would have to be interpreted as the Pine Series (123.12) of Southeastern Deciduous and Evergreen Forest although the Oak-Pine Series (122.14) of Northeastern Deciduous Forest (122.1) is closest in name. There should be a Brown et al. (1998) biotic community of Hardwood-Pine Series under Southeastern Deciduous and Evergreen Forest. South While this backwater forest was definitely not a swamp like cypress or tupelo it was at least a seasonal wetland and perhaps should be interpreted as part of Southeastern Swamp and Riparian Forest (223.1) of Brown et al. (1998) in what could be called a Pine-Hardwood Series (of say, number 223.15). Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

72. Twixt the pines- Among large loblolly pine dwarf or swamp palmetto made up a lower shrub layer (lower than yaupon which constituted another shrub layer) throughout a backwater flatwoods forest. Also below the older (larger) and more scattered pines, water and overcup oak (with occasional black tupelo) formed a discontinuous lower tree layer. Montgomergy County, Texas (backwater of the San Jacinto River). February, later hibernal aspect. Various units of forest range vegetation were listed in preceding photo captions.

73. Loblolly Pine-Mixed Hardwood Wet Forest- Loblolly pine is the most common and economically important pine in east Texas. It often grows on well-drained soils, but it is also the Pinus species best adapted to wet, even ponded, sites in the Pineywoods. On this regenerated second-growth forest loblolly pine is the dominant species but shares the forest with numerous associated angiosperm species including water oak, swamp chestnut oak, white oak, and sweet gum (Liquidambar styraciflua) in the overstory with an understory limited to a shrub layer of the small American holly (Ilex opaca) and a lower layer of pine seedlings with sedges and rushes. Liberty County, Texas. Vernal aspect, May. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem), K-101 (Oak-Hickory-Pine Forest). Forest cover type is best described as SAF 82 (Loblolly Pine-Hardwood), but it has elements of SAF 91 (Swamp Chestnut Oak-Cherrybark Oak). Oak-Pine Series of Brown et al. (1998), but in location of their Pine Series. South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

74. Loblolly Pine-Oak Hardwood Forest in Texas Pineywoods- On this wet, often ponded, site loblolly pine is co-dominant with numerous hardwood species including cherrybark oak (Quercus falcata var. pagodifolia), water oak, swamp chestnut oak, white oak, water hickory or bitter pecan (Carya aquatica), and sweet gum. The woody understory consist largely of regenerating species of the dominants just listed. Herbs consist of sedges, rushes, and scattered small individuals of the native bamboo (Arundinaria gigantea). Liberty County, Texas. Vernal aspect, May. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem), K-101 (Oak-Hickory-Pine Forest). Appears to be a transition or “hybrid” between SAF 82 (Loblolly Pine-Hardwood) and SAF 91 (Swamp Chestnut Oak-Cherrybark Oak). Oak-Pine Series of Brown et al. (1998), but in region of their Pine Series. South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

75. White umbrella sedge or white-top sedge (Dichromenta latififolia)- The striking bright corolla of this member of the Cyperaceae has prompted wild flower enthusiasts to regard this grass-like plant as a “wild flower”. It is restricted to wet open habitats as an understory to the more open loblolly pine-hardwood forests forms growing on poorly drained sites like that seen immediately above. Hardin County, Texas. May.

Although loblolly pine is well-adapted to wet soils (as suggested by the designation of "loblolly" in reference to deep mud hole or large mud puddle) is also occupies and even dominates less moist sites. On moist, but well-drained upland habitats throughout the extensive, eastern deciduous forest region loblolly pine frequently grows with various associated species. The Society of American Foresters (Eyre, 1980) explained that for the Loblolly Pine-Hardwood forest cover type (SAF 82) there is a "spectrum of moisture regimes and sites" with hardwood species varying according to these gradients. White oak is one of the more widespread component hardwood species on direr upland sites. Throughout much of the Pineywoods white oak is a major--often dominant--species on various forest cover types including shortleaf pine as well as loblolly pine. Forests of white oak and shortleaf pine (with associated species) were treated later on in this chapter. Natural vegetation of an upland (well-drained) forest subtype that was composed of white oak and loblolly pine was presented and described immediately below. This "vegetational sampler" was typical of the interrupted or variously scattered forest communities in the southcentral portion of the Oak-Pine Forest Region in which loblolly pine serves to charactrize this transition from Oak-Hickory Region to the Oak-Pine Region (Braun, 1950, p. 259, 278-279).

76. Upland loblolly pine-white oak forest- Composite view of an upland Pineywoods forest above a small stream in which loblolly pine and white oak were do-dominants of the the canopy layer with progressively lower vegetational layers formed by sweetbay (Magnolia virginiana) as a lower tree layer; yaupon holly as a shrub layer; Walter's greenbriar (Smilax walteri), some species of grape (Vitis sp.), and rattan or Alabama supplejack as multi-layer (ground to canopy) shrubs; and a sparse-barely present herb layer made up mostly of longleaf woodoats. Leavaes of tree and shrub species covered the ground level (soil surface) to such degree as to exclude most herbaceous species, including individuals of longleaf woodoats (generally the dominant herb). The small tree with green-tinged, smooth bark and arching over the stream (lower right corner) and the two two smaller trees on the opposite (left) bank were individuals of sweetbay. A woody shoot of grape was in center foreground. The green zone of vegetation was produced by green leaves of the evergreen yaupon. Liberty County, Texas. February, later hibernal aspect. Forest and Range Ecosystem (Garrison et al., 1977) was either FRES 14 (Oak-Pine Forest Ecosystem) or FRES 13 (Loblolly-Shortleaf Pine Forest Ecosystem). Either way Kuchler unit was K-101 (Oak-Hickory-Pine Forest). SAF 82 (Loblolly Pine-Hardwood). Not a good fit in the biotic community classification of Brown et al. (1998) has closest was Oak-Pine Series (122.14) of Northeastern Deciduous Forest (122.1) of the Cold Termperate Forest whereas this vegetation was clearly in the Warm Temperate Forest, Southeastern Deciduous and Evergreen Forest (123.1) for which there was not an Oak-Pine Series. Braun (1950, 1950, ps. 259-279) extended the Oak-Pine Region to the Coastal Plain which would include the Warm Temperate Forest and Woodland (123), Southeastern Deciduous and Evergreen Forest (123.1) of Brown et al. (1998, p. 38). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

77. Up and above a "crick"- Stand of loblolly pine and white oak on an upland site immediately above a small creek in Pineywoods of east Texas. Vertical view of the same forest introduced in the immediately preceding photograph (and from the same-- though closer-in-- vantagepoint) to better show structure and layering of this forest range vegetation. Woody vines of grape and smaller ones of rattan were in foreground. Limb in upper right corner was of sweetbay, the smaller tree species that constituted a lower tree layer of this forest community. Liberty County, Texas. February, later hibernal aspect. Forest and Range Ecosystem (Garrison et al., 1977) was either FRES 14 (Oak-Pine Forest Ecosystem) or FRES 13 (Loblolly-Shortleaf Pine Forest Ecosystem). Either way Kuchler unit was K-101 (Oak-Hickory-Pine Forest). SAF 82 (Loblolly Pine-Hardwood). Not a good fit in the biotic community classification of Brown et al. (1998) has closest was Oak-Pine Series (122.14) of Northeastern Deciduous Forest (122.1) of the Cold Termperate Forest whereas this vegetation was clearly in the Warm Temperate Forest, Southeastern Deciduous and Evergreen Forest (123.1) for which there was not an Oak-Pine Series. Braun (1950, 1950, ps. 259-279) extended the Oak-Pine Region to the Coastal Plain which would include the Warm Temperate Forest and Woodland (123), Southeastern Deciduous and Evergreen Forest (123.1) of Brown et al. (1998, p. 38). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

78. Lumberman's view of a loblolly pine-white oak upland forest- Structure and species composition of an upland white oak-loblolly pine form or subtype of loblolly pine-hardwood dominance type. Larger trunks in both photographs are loblolly pine. Ph;otographed immediately following heavy rain shower so bark over some long strip-line areas of pine trunks was darker. Sweetbay formed an interrupted second tree layer. These were present as smaller, shorter trees (large saplings or pole-size: two in front of and to left of foremost pine in first slide; whitish trunk in foreground of second slide). An upper shrub (lower woody) layer comprised of yaupon was not distinct in these slides, but was shown in the six slides of immediately succeeding three sets of slides. Ground surface was covered with mulch or duff layer formed from shed leaves of all species. There was a very sparse understorey made up mostly of Walter's greenbriar that was more shrub than herb. A very sparse, intermittent herbaceous layer was composed mostly of longleaf woodoats (an individual of this species was to immediate left and upslope of the sweetbay in foreground of second slide). There were even more scattered individuals of some unidentifiable Carex species. Liberty County, Texas. February, later hibernal aspect. Forest and Range Ecosystem (Garrison et al., 1977) was either FRES 14 (Oak-Pine Forest Ecosystem) or FRES 13 (Loblolly-Shortleaf Pine Forest Ecosystem). Either way Kuchler unit was K-101 (Oak-Hickory-Pine Forest). SAF 82 (Loblolly Pine-Hardwood). Not a good fit in the biotic community classification of Brown et al. (1998) has closest was Oak-Pine Series (122.14) of Northeastern Deciduous Forest (122.1) of the Cold Termperate Forest whereas this vegetation was clearly in the Warm Temperate Forest, Southeastern Deciduous and Evergreen Forest (123.1) for which there was not an Oak-Pine Series. Braun (1950, 1950, ps. 259-279) extended the Oak-Pine Region to the Coastal Plain which would include the Warm Temperate Forest and Woodland (123), Southeastern Deciduous and Evergreen Forest (123.1) of Brown et al. (1998, p. 38). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

79. Ground level view of a lobolly pine-white oak upland forest- Species make-up and layer arrangement of an upland loblolly pine-mixed hardwood forest was shown to good advantage. This forest community was in the unique Big Thicket portion of the Pineywoods and was featured in this segment of discussion devoted to loblolly pine. Loblolly pine is typically a subclimax stage of forest in the vast eastern deciduous forests of North America. Successional status of white oak varies considerably. Liberty County, Texas. February, later hibernal aspect. Forest and Range Ecosystem (Garrison et al., 1977) was either FRES 14 (Oak-Pine Forest Ecosystem) or FRES 13 (Loblolly-Shortleaf Pine Forest Ecosystem). Either way Kuchler unit was K-101 (Oak-Hickory-Pine Forest). SAF 82 (Loblolly Pine-Hardwood). Not a good fit in the biotic community classification of Brown et al. (1998) has closest was Oak-Pine Series (122.14) of Northeastern Deciduous Forest (122.1) of the Cold Termperate Forest whereas this vegetation was clearly in the Warm Temperate Forest, Southeastern Deciduous and Evergreen Forest (123.1) for which there was not an Oak-Pine Series. Braun (1950, 1950, ps. 259-279) extended the Oak-Pine Region to the Coastal Plain which would include the Warm Temperate Forest and Woodland (123), Southeastern Deciduous and Evergreen Forest (123.1) of Brown et al. (1998, p. 38). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

80. Shrubs beneath the pine and oak- Yaupon or yaupon holly made up a shrub layer in the upland loblolly pine-white oak dominated (canopy or cover dominance) forest that had developed along a small stream in the Big Thicket portion of Texas' Pineywoods. This specific forest stand was shown and described in detail above and below. Liberty County, Texas. February. Various classification units of this forest vegetation were presented in photo captions herein.

81. Walk through the upland woods- A series opf three photographs taken from about the same angle presented structure and species composition of vegetation in an loblolly pine-white oak-sweetbay-liana-yaupon- longleaf woodoats upland forest in the Big Thicket portion of the Texas Pineywoods. This "photo stroll" centered on a large white oak (largest tree trunk) near center of photographs. The bark on this old and still alive white oak had sloughed off in patches which were accentuated by a heavy rain moments before this series of photographs was taken. Smaller, shorter trees were sweetbay which made an interruped lower tree layer.Yaupon formed a shrub layer throughout. A "top-to-bottom" (ground-to-crown canopy) shrub layer consisted of grape, rattan or Alabama supplejack, and Walter's greenbriar. Leaves of all species produced a ground cover layer so thick that there were very few herbaceous species. The most common herb was the perennial grass, longleaf woodoats. Liberty County, Texas. February, later hibernal aspect. Forest and Range Ecosystem (Garrison et al., 1977) was either FRES 14 (Oak-Pine Forest Ecosystem) or FRES 13 (Loblolly-Shortleaf Pine Forest Ecosystem). Either way Kuchler unit was K-101 (Oak-Hickory-Pine Forest). SAF 82 (Loblolly Pine-Hardwood). Not a good fit in the biotic community classification of Brown et al. (1998) has closest was Oak-Pine Series (122.14) of Northeastern Deciduous Forest (122.1) of the Cold Termperate Forest whereas this vegetation was clearly in the Warm Temperate Forest, Southeastern Deciduous and Evergreen Forest (123.1) for which there was not an Oak-Pine Series. Braun (1950, 1950, ps. 259-279) extended the Oak-Pine Region to the Coastal Plain which would include the Warm Temperate Forest and Woodland (123), Southeastern Deciduous and Evergreen Forest (123.1) of Brown et al. (1998, p. 38). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

82. Fire-scarred forest veteran- An old white oak on a loblolly pine-white oak dominated upland forest bore testimony to the frequency of past fires. As a general rule, fire in the hardwood-pine cover types of the North American or eastern deciduous forests selects for greater proportions (crown cover, density, dominance, etc.) of pine rather than hardwood species such as the various oaks and hickories. This is most true for the extremely fire-tolerant longleaf pine, but even less fire-tolerant pines like loblolly generally benefit at competitive expense of the angiosperm trees. Furthermore, susceptibility to fire is greater for seedlings, saplings, and small poles than adult hardwood trees. Once hardwoods become established and grow larger they are less susceptible to fire-damage and death. The ole patriarch of this upland loblolly pine-hardwood forest had obviously been through several surface fires. Past surface fires had burned through the bark of this large white oak which, however, survived quite well as most of its cambium tissue had not been injured. White oak has a variable tolerance response, but is generally rated as Intermediate (in contrast to Intolerance of loblolly pine). White oak is somewhat more tolerant than loblolly pine to drought (Moderate-tolerant vs. Moderate, respectively) while loblolly pine is much more flood tolerant (Moderately vs. Intolerant of white oak) (Wenger, 1984, ps. 2-8). Frequent fire shifts the forest environment in favor of loblolly pine. Leafy plants at base of the white oak were small individuals of yaupon. Sapling behind and to right of white oak was sweetbay, the major species of the lower tree layer. Liberty County, Texas. February.

83. Walter's greenbriar, coral or red-bead greenbriar,, or sarsaparilla (Smilar walteri)- One of several Smilax species in the Pineywoods Region. This one generally prefers moist to wet habitats, including sandy soils along streams such as that in the loblolly pine-white oak-sweetbay-yaupon upland forest described here. Liberty County, Texas. February.

84. Nitty gritty of Walter's greenbriar- Details of leaves and stems of Walter's greenbriar which is only one of various Smilax species in the eastern deciduous forest of North America. This was growing in the understorey of a loblolly pine-white oak-dominated upland forest that developed along a small stream in the Big Thicket. Liberty County, Texas. February.

Agriculture in southeastern North America (the Southeast) has changed continuously-- often radically-- over the course of history. Without going back to pre-Columbian (prehistoric) times and Indian days and the colonial period it was sufficient to remark that much of what was known generally (and simplistically) as the Cotton Belt has been one of the continent's agricultural regions most changed by socioeconomic and biological forces. Where it was first the "precious stink of tobaccy" and later King Cotton "the southeastern sector" is now one of the most diverse agricultural regions in North America. Probably nothing as so changed in the agriculture of the Old South as the shift from field, mostly row, crops to industrial forestry and, usually integrated with forest cropping, beef cattle production. Development of permanent pastures based on improved, introduced tropical and subtropical grasses like bermudagrass, bahiagrass, dallisgrass, and vaseygrass along with tall fescue and Kentucky bluegrass went side-by-side with scientific forest management and a sustainable forest industry producing wood for pulp, post, poles, naval stores, and about anything else imaginable. Other agricultural changes that at first might appear to be unrelated to those of the old Cotton Kingdom were in reality direct "directing forces" in the diversified agriculture of the South that slowly "rose again". Development of a concentrated, intensively managed cattle-feeding industry in the Great Plains of Texas, Oklahoma, Colorado, New Mexico, and Kansas was one example. This post-World War II phenomenon provided a ready made market outlet for Okie feeder calves from Florida, Georgia, Alabama, Louisiana, etc. while the surrounding wheat fields were pastures without peer for the Okie "threes and fours" which were to be the stockers not yet ready for "prime time" in the feedlots. Development and ever-newer improvements in the Forestry and Agronomy of the South (including the Border States like Kentucky, Tennessee and southern Missouri) were joined with improvements in Range Management. The following section presented in photograph and discussion some of the major parts of this integrated agriculture that was a region-wide form of agropastoralsilviculture long before such terms became popular or were even recognized as such.

Recall from the introduction to this section that Braun (1950) recognized nine sections of the eastern deciduous forest formation. Two of these comprise-- more or less-- the forests of the Southeast that include sizable components of pine (Pinus) species: 1) Oak-Pine Forest Region and 2) Southeastern Evergreen Forest Region. This part of the Southeast could be viewed conveniently as the "pineywoods" part of the eastern deciduous forest formation or "super-region", as indeed it has long been known in Texas. This has also been known traditionally as the Southern Pine Region. It is mostly this part of the Southeast, especially in less economically developed areas, in which industrial or commercial forestry is a-- often, the-- major industry and certainly the more important part of agriculture in general. It was loblolly pine (Pinus taedna), including hybrids thereof, more than any other single species that literally revolutionized commercial forestry in the Southeast. Slash pine (Pinus elliottii) was the other major commercial conifer of this region, specifically the southeasternmost part including Florida and the Carolinas. In the more northern and western portions of the Southeast shortleaf pine (P. echinata) was the native species and it has remained the major commercial conifer in certain locations. The fourth native pine having some commercial value in industrial forestry is longleaf pine (P. palustris). Following logging of the virgin forest, the slower-growing (but often having the superior wood) longleaf pine was in most areas the least economical species for commercial wood production. Shortleaf pine is the least likely of the major pines of the Southern Pine Region to suffer damage or death from ice storms that are only all too frequent especially in the more northern zones of the region. Slash pine is typically the most extensively damaged by ice storms. Ice damage also limits commercial production of longleaf pine, as does it's slow growth rate relative to loblolly and slash pine and silvicultural problems associated with propagation/planting. Slower growth rate is also a factor limiting the industrial production of shortleaf pine. A good-- thorough but concise-- guide to the feasibility of the various pines for production in industrial forestry can be found in Fowells (1965) and Burns and Honkala (1990), the definitive reference on the silvics and silviculture of forest trees in North America. Wahlenberg (1960) and Schultz (1997) are probably the two comprehensive treatments of loblolly pine forestry. Another classic reference, whose name says just what it is (one hallmark of a classic), would have to be The Genetics and Breeding of Southern Pines (Dorman, 1976). The definitive authority on southern pine ranges and their management is Pearson et. al. (1987).

The following sequence showed the harvest of loblolly pine by the clearcutting silvicultural method and subsequent site preparation for the next pine crop. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 81 (Loblolly Pine).

85. Landing in loblolly pine forest- A landing is the point to which logs, poles, pulpwood, etc. is brought for delivery to the mill. In this scene loblolly pine logs were skidded and loaded on the truck by a log self-loader operator. Miller County, Arkansas. July.

86. Skidding logs to the truck- Skidding (also called yarding or forwarding) is the hauling of logs, poles or so on to the landing which is the collection or loading point for transport of logs to the mill. A rubber tired-skidder brought loblolly pine logs to the landing for loading onto the logging truck. Miller County, Arkansas. July.

87. Loading logs with a self-loader- A knuckleboom loader mounted on the logging truck was being used to load logs on the truck for transport to the mill. A knuckleboom (= hydraulic loader) is a loader that swings when used for loading and has hydraulic controls that activate boom members (the tongs or jaws). Miller County, Arkansas. July.

88. Slash at the landing- Slash is the logger's term for the limbs, branches, tops, parts of roots, waste logs, etc. remaining on the land surface after harvesting saw timber, pulpwood, etc. It is the forestry equivalent of stubble, straw, stalks, etc. remaining on the soil of a farm field. In both cases this residue must be managed in preparation for planting the next crop. It can be beneficial in reducing soil erosion, adding organic matter, and protecting the new seedlings, but such trash can interfere with planting or seeding operations. Miller County, Arkansas. July.

89. Oak-hickory-pine forest after harvest of loblolly pine- Where the potential natural forest vegetation is a hardwood-pine cover type (ie. "pineywoods") pines are usually the designated or intended crop and the hardwoods are weed trees. Weed trees are any species of tree (or large shrub), and the individual plants of these species, that have little value (or their value is less than-- cannot offset-- their disadvantages); used especially when these woody weeds compete with the intended forest crop. This latter is known as crop tree, any individual tree or tree species chosen to be part of the future forest harvest. Some weed trees grow even under the most intensive forest management practices. These weed trees must be controlled before the next tree crop can be planted. This is true from standpoint of both 1) competition to planted seedlings or seedings from pre-existing, established, and much larger trees (for light, water, soil minerals, etc.) and 2) operating planting equipment, movement of tree-planter, workers, and so on. One of the major points of contention among conservationists involved management of weed trees. Foresters who favor less intensive forest management (especially less radical or disruptive silvicultural methods), wildlifers, some watershed experts, and lay people who enjoy the varying and seasonal color changes of the deciduous species want a higher proportion of hardwood species growing with the pines. Foresters who favor the more intensive management of industrial forests (eg. shorter rotations, higher degree of control of pathogens and insect pests, and generally "cleaner" or "neater" forests opt for fewer weed trees). Forests managed from the latter perspective are often plantations of one (or very few) species, especially pines, hence pine plantation. These monocultures (where there is one crop tree species such as loblolly pine) are literally tree farms. (This should not to be confused with the registered Tree Farm trade mark of the American Tree Farm System although some of these may be registered or generally comply with those guidelines). They are the forestry equivalent of the agronomic single species of field crop (corn, cotton, wheat). As with every other form of commercial agriculture in industrialized democracies, market economics has consistently favored ever-increasing intensification of food and fiber production with the result that wood crops have been produced under more intensive foret management, especially tree breeding and silviculture. This has not been completely successful with any form of agriculture whether tree farming, fish farming, game bird farming, or broiler and pork production under total confinement systems, but this industrial agriculture has produced the most economical food and fiber for the consumer as she has continued to "shell out" less of her disposable income for food, clothing, lumber, and computer paper with each passing year. More recently certain factors, both biological, chemical-physical, and socioeconomic, slowed the trend of increasing intensification of production, at least somewhat. Endangered species management, more emphasis on aesthetics and recreation, and less concern about price of goods and services may well create a demand in the market place of forest products (not just wood commodities) for a "New Forestry" or a "Forestry for the 21st Century". If consumers are willing to put their money where their mouth (and their vote) is and "play fair" by the rules of a democratic free market so as to pay for the increased cost of products and services in order to enjoy more "natural" forests, foresters and forest firms will rise to the occasion as they did in wise-use conservation for economically efficient wood production. If, on the other hand, the consumer is an irresponsible customer and seeks to be a free-rider and stupidly expects the forest products industry to continue providing economical wood products plus other forest amenities at the same prices (ie. the addded costs are paid entirely by the industry) the system will fail. Natural resources are not free goods; all who use them must pay a fair price. That is a fundamental rule of the woods. That is the system, economic and ecosystem. Back to the weed trees: they must be removed for proper site preparation. Miller County, Arkansas. July. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 81 (Loblolly Pine).

90. Site preparation- A small loblolly pine clear-cut with weed trees (hardwood species, mostly oak) removed. This was a portion of the pine-harvested forest with hardwoods left standing shown in the preceding slide. This part of the forest was in a later phase of site preparation for planting the next crop of loblolly pine. In this part of the "pine field" silvicultural scarification has been completed. Scarifiction in this usage means the physical removal of slash (or related debris) and competing plants (eg. weed trees, shrubs) and/or physical disturbance or disruption of the land surface as one of the practices in site preparation. Site preparation is the manipulation, usually limited in meaning to physical (manual or mechanized) and chemical treatment, of the site (the land) on which reforestation is to occur. Site preparation is the equivalent of seedbed preparation in production of field crops. Site preparation treatments include all those associated with scarification and readying of the soil surface for planting or seeding (eg. brush or weed tree control, disking, chopping, brush raking, tile drainage, terracing). These modifications to soil, land shape, slash (litter), and remaining plants are performed to improve microsite conditions in which the new trees will began their growth. The forest management objective is enhanced regeneration of the crop trees (recall definition from preceding slide caption). The land seen here is not in the final stage of site preparation. Slash has been piled for burning, but the piles contained enough soil that combustion will be incomplete, and the soil will have to be redistributed back over the land. This procedure exposed soil to erosion but there was probably enough slash residue to prevent accelerated erosion. Such tillage of the soil can be beneficial (it was part of the scarification procedure) and is often essential for successful regeneratioin of species that evolved under disturbances like flooding (eg. coast redwoods) or fire (pines with serotinous cones like some lodgepole genotypes). This was explained in discussions of these species elsewhere in this publication. Miller County, Arkansas. July. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem. K-101 (Oak-Hickory-Pine Forest). SAF 81 (Loblolly Pine).

91. Proper site preparation- Appearance of a "field", the site, ready to be replanted to loblolly pine. This land is in the final state or last stage of site preparation. Replanting of loblolly pine seedlings will begin soon. Note the amount of "crop residue", slash, remaining on the soil surface. Erosion will be minimum with this residue covering the soil and acting as small "dams" or "windbreaks" to slow movement of water and wind that could carry soil particles. Also, this modifies the microenviornment where replanted conifers will grow and develop, thereby increaing the chances of regeneration of the crop trees (ie. improve the odds of making the next crop). Hardwood tree and shrub species have already resprouted or germinated. This means that conifer seedlings will be in intense competition as soon as they are planted and/or germinate. Efficient regeneration of loblolly pine under these conditions will usually require some brush (weed tree) control either by herbicides or cattle-grazing. This was discussed below (and for several conifer species and different forest cover types such as the Sierra Nevada mixed conifer type). In the case of grazing by cattle (as well as native whitetail deer of course) the stage of reforestation displayed in this slide was the start of transitory forest range, native plants on forestland that can (will) be used as forage and/or browse only as long as the forest canopy formed by crowns of trees (loblolly pine in this instance) does not close enough to prevent light from reaching lower layers of the vegetation. Once enough light is intercepted by the crowns of the ever-growning tree layer(s) --resulting in elimination of grazable or browsable layers-- the forest no longer produces native vegetation available for use by range animals. An understorey (understorey layers) capable of being used as forage and/or browse by wildlife or livestock no longer exist. The forest range is gone. It has been replaced by another seral stage or by the climax of that sere. In other words, such forest range is a stage or a series of stages in secondary plant succession that produces vegetation that can be consumed by grazers and/or browsers until it is evenutually replaced by climax or a higher seral stage that is incapable of producing feed for range animals. Stages in the development of forest vegetation that do produce such forage and/or browse are thus in transition to a closed-canopy forest. In other words, it is successional, sub-climax range that is transitory-- in a state of transition-- to more successionally advanced vegetation that lacks an understorey. Transitory forest range is not permanent range. Some forest range types (eg. longleaf pine and parklike ponderosa pine forests) are permanent. A number of these were discussed throughout the Forests and Woodlands portion of this publication. Many of the forest ranges of the Southern Pine Region are transitory ranges. In these instances, such as the one shown in the next slide, man the forester facilitated succession or grew a disturbance climax that no longer had an understorey layer. Earlier generations learned from Plant Ecology (Weaver and Clements, 1938, ps.238, 271-273) that this was an example of human coaction, an advanced process in plant succession leading to stabilization. Vernon Parish, Louisiana. July. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 81 (Loblolly Pine).

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92. Pine nursery- Nursery stock for artificial regeneration. In tree nursuries, fields such as the one shown here have seeds of the desired species planted in rows. Seedlings are allowed to reach heights of about 10-15 inches (maybe slightly more) then they are lifted and bound in bundles of a certain number (commonly 50 seedlings at this nursery). The seedlings are transported to the site (the "field") on which they are to be planted. Planting can be done by tractor-drawn implements (commonly called tree planters) or by hand using various tools such as tree-planting bars, hoedads, or spades. Bundles of seedlings (typically of several species) are packaged together and routinely shipped by surface mail, especially to small private landowners for reforesting relatively small acreages. Commercial lumber and pulp companies often have their own nurseries, of their own selected germ plasm. This nursery field had both loblolly and shortleaf pine seedlings (growing in the center rows) and black locust and bois-d'arc or Osage orange (these latter two species visible at far left as the taller seedlings). This was a state forestry nursery funded by tax dollars and minimal prices for seedlings. It exist primarily to provide trees for replanting on private, non-commercial (ie. smaller) forest tracts. Oklahoma Forestry Division, State Department of Agriculture. July

93. Young loblolly pine seedlings growing in a state forestry nursery- These trees will be sold at minimal (tax-subsidized) cost to private landowners for reforestation. This nursery stock method is the standard form of artificial regeneration for species of southern pines. (An erratic stand by standards of corn and cotton fields.) Standard three and a half inch stockman's knife for scale. Oklahoma Forestry Division, State Department of Agriculture. July.

94. Loblolly pine plantation- This is a forest planting neighboring on the area shown in the preceding photograph. It is a six to (at most) ten-year-old loblolly pine plantation. The pines were large enough that they were experiencing relatively little competition from weed trees, brushy shrubs, or grass. The pines were also large enough that there was not much forage and/or browse remaining in the plantation. Cover of loblolly pines, which had limbs to the ground, was so nearly complete that forest plants usable for forage and/or browse had been reduced nearly to point of exclusion by interception of light by the planted pines. The loblolly pines had grown enough to exclude light from the ground around them and little of the grazable/browsable understorey remained. Here was an example of where man the manipulator of ecosystems accelerated the rate of secondary plant succession while producing a crop that could be characterized as a disturbance climax or a subclimax (Weavaer and Clements, 1938, p. 81, 86-88). This resulted in a marked degree of stabilization, the most advanced process in plant succession (Weaver and Clements, 1938, p. 234). The Society of American Foresters (Helms, 1998) defined a plantation as "a stand composed primarily of trees established by planting or artificial seeding" and specified that "a plantation may have tree or understorey components that have resulted from natural regeneration" and that it "may be pure or mixed species, treated to have uniform or diverse structure and age classes". Most of the loblolly plantations like the one seen in this slide are monocultures with a single age class and one simultaneous harvest (ie. even--aged management with the classic pattern of clear-cutting). Vernon Parish, Louisiana. July. Western Coastal Plains ecological section. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 81 (Loblolly Pine).

95. Reproduction of loblolly pine by natural seeding- A dog-hair stand of loblolly pine developed on a road-cut from seeds in the soil seedbank and those newly shed by existing trees. Silvicultural systems for production of loblolly pine vary but, as is the case for industrial forestry in general throughout the Southern Pine Region, commercial regeneration (silvicultural establishment of the next tree crop) and the major regeneration or reproduction method became the pine plantation sysem of even-aged clearcutting and replanting. Replanting of nursery stock (young tree raised in tree nurseries from seed) became established quite early in the development of scientific forestry in the Southeast because it provided the opportunity to use improved, faster-growing genotypes (including hybrids) as well as increasing chances of tree establishment and a future wood crop. Smith (1986, p. 372) explained that production of loblolly and slash pine by clearcutting and planting was "the most important American manifestation of the general use of this silvicultural method". Loblolly pine does, however, reproduce readily from seed as shown in this photograph. This is even more the case when competing hardwood species are reduced by prescribed burning or mechanical site preparation (as was the inadvertant result of the road construction that eliminated all competing plants and exposed mineral soil). Loblolly pine has also been established by artificial aerial seeding, but this practice was discontinued and was replaced by planting nursery stock to get properly spaced trees of superior germ plasm. Regeneration of loblolly pine from seed (naturally or artificially) results in stands of densities greater than those that are optimum for rapid tree growth and shorter rotations. It not only creates such dog-hair stands that pines are crowded and stunted, it also results in develop of a closed tree canopy early in development of vegetation and exclusion of an understorey capable of being used as transitory forest range. Natchitoches Parish, Louisiana. July. Red River Alluvial Plain ecological section. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 81 (Loblolly Pine).

96. Thinned out- Stand of loblolly pine thinned for optimum growth of individual trees and wood yield. Yaupon holly comprised a lower, second woody layer. Herbaceous understorey consisted of a diversity of grasses including especially longleaf woodoats, panicgrasses (Panicum and Dichanthium spp.), and paspalums (Paspalum spp. including the naturalized Vaseygrass) along with sedges (Carex and Cyperus spp.) and spikerushes (Eleocharis spp.). Within a relatively short period (probably less than 10-15 years) this stand will become a closed canopy monoculture of plantation loblolly pine like that shown immediately below. Hardin County, May, estival aspect. Pyric or anthropogenic variant of K-101 (Oak-Hickory-Pine Forest) that would terminate in dominance by hardwood species. SAF 81 (Loblolly Pine). Pine Series of Brown et al. (1998). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

97.Loblolly forest at mature stage relative to harvest (mainly pulp wood)- Closed canopy forest that is totally devoid of understory. Stands of loblolly pine such as this essentially single-species stand are transitory range that is grazable/browswable only until the upperstory of trees closes thereby depriving understory layers (often even shrub stories) of light. Houston County, Texas.March, vernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem). Pyric or anthropogenic variant of K-101 (Oak-Hickory-Pine Forest) that would terminate in dominance by hardwood species. SAF 81 (Loblolly Pine). Pine Series of Brown et al. (1998). South Central Plains- Southern Tertiary Uplands Ecoregion, 35e (Griffith et al., 2004).

98. Small clear-cut of loblolly pine like that pictured in previous slide- What slash is not used locally as fire wood following pole and/or pulp wood harvest will be burned and the site prepared for replanting which is typically artificial propagation (planting of nursery grown seedlings rather than natural regeneration by seeding from adjacent or scattered remaining trees). Liberty County, Texas. March. FRES No.13 (Loblolly-Shortleaf Pine Ecosystem).K-101 (Oak-Hickory-Pine Forest).SAF 81 (Loblolly Pine). Pine Series of Brown et al. (1998). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

99. A loblolly pine plantation eight years following planting of seedlings in a clear-cut like the one immediately above— The plantation understory is being grazing by cattle as a means of biological control of the fiercely competitive weed tree, sweetgum (Liquidambar styraciflua), as well as utilization of the native grasses that vary from climax Andropogon and Panicum species to the the threeawns or wiregrasses and crabgrasses (Digitaria spp.). Little bluestem and broomsedge bluestem (Andropogon virginicus) are the dominant grasses shown here. Native hickory and oak species have regenerated by both coppice sprouting and seedling emergence yet are less a brush problem than seral sweetgum.This illustrates that oak and hickory species are natural dominants of the climax and that the human inputs of forest management are essential to economically raise the crops of pines which are minor climax dominants relative to the hardwoods.Note preferencial grazing first of grasses and secondly of browsing on hardwoods. Pines have not been browsed. Loblolly pine transitory range.Weyerheuser-contracted crop. LeFlore County, Oklahoma. May. FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem). K-101 (Oak-Hickory-Pine Forest) SAF 81 (Loblolly Pine) or SAF 82 (Loblolly Pine-Hardwood). Pine Series of Brown et al. (1998). Ouachita Mountains- Athens Plateau Ecoregion, 36a (Woods et al., 2005).

100. Transitory forest range in a 10 to12 year-old hybrid loblolly pine plantation showing ungrazed understory that is tallgrass prairie of little bluestem, big bluestem, broomsedge bluestem, Indiangrass, and side-oats grama. Hardwood species like oaks, hickories, and sweetgum are totally absent due largely to previous heavy browsing by cattle.Weyerheuser trees.McCurtain County, Oklahoma. July. FRES No.13 (Loblolly-Shortleaf Pine Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 81 (Loblolly Pine) or SAF 82 (Loblolly Pine-Hardwood). Pine Series of Brown et al. (1998). South Central Plains- Cretaceous Dissected Uplands Ecoregion, .35d (Woods et al., 2005).

101. Loblolly pine regeneration (and competition) in cut-over forest- Natural regeneration of loblolly pine from seed following clearcutting (even-aged regeneration method). The pines are in competition and face the threat of wild fire from the lush growth of native grasses including broomsedge bluestem (the most abundant grass), splitbeard bluestem, big bluestem, plus various species of Panicum, Paspalum, and Sporobolus.

102. Vulnerable to fire- Close-up view of a loblolly pine seedling in the cut-over forest shown immediately above. This silvic baby is about as vulnerable as a newborn lamb. The seedling was surrounded by broomsedge bluestem (the Andropogon species closest to it), splitbeard bluestem, little bluestem with dead herbage of Panicum and Paspalum species not far away. This new pine was produced by natural reproduction (seed production, germination, and emergence) and there are frequently too many such seedlings produced so that killing of the vast excess by fire is essential management for efficient, economical production of forest products. On this cut-over forest, however, regeneration was not excessive thus necessitating protection of the next cohort of loblolly pines from fire at this vulnerable stage of their life cycle. Grazing by cattle (the kind of range animal most likely to consume grass and not browse on pines) would reduce the fuel load produced by grasses (and some grasslike plants and forbs)thereby reducing chances of wild fire that would eliminate the barely adequate stocking of loblolly pine. Grazing of such rank, dormant, and, in instance of broomsedge and splitbeard bluestems, unpalatable grasses at this stage of full plant maturity and dormancy is not feasible. Cattle will not graze such herbage as show here (at least not at levels of voluntary forage intake that would be profitable to cattlemen). Rather, grazing should have been done back when these herbaceous species were immature and less unpalatable (ie. get on top and stay on top of the potential fuel). Harrison County, Texas. December.

103. Waiting to burn and die- A fine loblolly pine seedling in an ocean of grass herbage. Grass material was mostly from broomsedge and splitbeard bluestems, species of extremely low palatability. Getting loblolly pine to enough height to withstand a wild fire on this cut-over pine forest is "ify" and one time when even rangemen (if they done a forester's hardhat for a time) find common cause with that otherwise deplorable bruin, Smokey Bear. The forest range shown here is in the heart of the loblolly pine region and to the west of longleaf pine forest. Thus, wise use of prescribed fire is not as (perhaps not) feasible. Grazing of this regenerating loblolly pine forest by beef cattle would be one of the best--if not the best--practices to maximize the chance of establishing the next crop of wood. Of course, grazing of unpalatable species like broomsedge and splitbeard bluestems has to begin when herbage is young and more acceptable to cattle. Cows and calves would be preferable to steers on low-quality feed such as that seen here because stocker cattle must achieve higher levels of individual performance to be profitable under the almost-always negative price structure (heavier cattle fetch lower prices per cwt.). See there, the ole range professor slipped in one of the Cardinal Principles of Range Management: Proper Kind and Class of Range Animal (in this case, class as to sex of animal). Harrison County, Texas. December.

104. Longleaf pine (Pinus palustris)- Seedlings and young trees of longleaf pine are seen here in at least four age classes. Longleaf pine is one of the most fire-tolerant trees in North America. Natural and Indian-set fires contributed to persistence of Pinus species in the deciduous forests of eastern North America. This was particularly the case for longleaf pine where fire maintained parklike forest of almost pure longleaf with a grassy understory much like the case for the ponderosa pine forests of western North America. In addition to reduction of a woody understory (and thus likelihood of a crown fire) fire may have helped control southern blister or fusiform rusts such as Cronartium fusiforme= C. quercuumf. ssp. fusiforme. (In addition, longleaf pine is more resistant to fusiform rust than is loblolly or slash pine [Baxter, 1943].) Fire has definitely been proven to be useful in control of brown spot disease (Septoria acicola) when longleaf is in the grass stage (Wright and Bailey, 1982, ps. 369, 415). Fire cost the burnt grass stage longleaf a year’s growth because it consumes the needles (fire destroys the brown-spot spores in the fallen leaves), but in absence of this fire the young longleaf trees would die from brown-spot disease. Winter burns at three year intervals result in doubling the growth of longleaf. Longleaf pine is much more tolerant of fire than are loblolly and slash pine. Natural fires at two to three year intervals maintained longleaf whereas a reduced fire frequency results in loblolly and slash pine becoming the dominant Pinus species. Absence of fire results in succession to the climatic climax mixed pine-deciduous (= hardwood) forest. (Wright and Bailey, 1982, ps. 368-371). In other words, all the southern pine forest types are fire types and this is most true for the longleaf pine type. Stand of young longleaf pine in background. Hardin County Texas, May. FRES N0. 12 (Longleaf-Slash Pine Forest Ecosystem), K-102 (Southern Mixed Forest, Seral Stages), SAF 70 (Longleaf Pine) of the Southern Yellow Pines.

105. The grass stage of longleaf pine- A longleaf pine seedling in the grass stage (front foreground) and a loblolly pine seedling of comparable age (behind the longleaf) shows conclusively why frequent firing will maintain longleaf pine instead of the more competitive loblolly pine.  A surface fire will burn off the leaves of the longleaf seedling and set it back a year’s growth, but the fire will kill the loblolly seedling. In absence of fire brown-spot would likely kill or retard growth of many young longleafs. The actual mechanism by which grass-stage longleaf survives is simple: the needles grow in a dense pattern around the terminal bud (apical meristem), which is the actively growing tissue of the seedling and whose hormones regulate growth of the tree, and protect it (and thus the seedling’s life) from the heat or consumption by the flames. A most remarkable evolutionary adaptation for a desirable forest tree valuable for both its lumber and naval stores. Hardin County, Texas, May. FRES No. 12 (Longleaf-Slash Pine Forest Ecosystem), K-102 (Southern Mixed Forest, Seral Stages), SAF 70.

Organization note: more coverage of loblolly pine forest types are in a seperate chapter, Loblolly Pine.

Swamps and Related Wetland Forests

Dispersed widely, though sometimes extensively, the eastern deciduous forest complex there are various forest cover types on wetlands. Most commonly these forested wetlands are swamps or riparian forests or woodlands. Swamp was defined by the Society of American Foresters (Helms, 1998) as "a tree- or tall shrub-dominated wetland, characterized by periodic flooding and nearly permanent subsurface water flow through mixtures of mineral sediments and organic materials, essentially without peat accumulation". Much of the wetland forest vegetation furnishes little or no herbage or woody material for forage and browse due to either absence of an understorey or nearly permanent water inundation. While such forests are of limited value (at best) as grazing land per se their vegetation is part of the overall range landscape and does provide water and shade for livestock; serves as sources of water, cover, and space as habitat factors for wildlife, contributes biodiversity to the general forest range ecosystem; and, probably most important of all, serves as essential watershed including the role of flood protection. A short sample of these wetland forest types was provided below.

106. Pine Island Bayou- Portion of bayou along which bald cypress dominated (almost exclusively) the riparian zone. Example of a slough-swell system. Manco soil series. Big Thicket National Preserve, Pine Island Bayou, Hardin County, Texas. May, late vernal aspect. FRES No.16. (Oak-Gum-Cypress Ecosystem). A co-dominant variant of K-103 (Southern Flood-Plain Forest). SAF 102 (Baldcypress-Tupelo). Bald Cypress Association (if and when recognized), Tupelo-Cypress Series in Southeastern Swamp and Riparian Forest biotic community of Brown et al (1998). South Central Plains- Floodplains and Low Terraces Ecoregion, 35b (Griffith et al., (2004).

107. Bald cypress (Taxodium distichum) backwater swamp-Consociation of bald cypress but local associates are water oak (Quercus nigra) and water elm (Planera aquatica). No understory at all; standing water most of the year. Big Thicket National Preserve, Maple Creek, Hardin County, Texas. May, late vernal aspect. FRES No.16 (Oak-Gum-Cypress Ecosystem). The bald cypress variant of K-103 (Southern Flood-Plain Forest). SAF 102 (Baldcypress-Tupelo). Tupelo-Cypress Series in Southeastern Swamp and Riparian Forest biotic community of Brown et al. (1998). Would be Taxodium distichum Association (if such is recognized). South Central Plains- Floodplains and Low Terraces Ecoregion, 35b (Griffith et al., (2004).

108. Bald cypress (Taxodium distichum)- Leaves and cones of bald cypress. Hardin County, Texas. September.

Presented immediately were a series of eight photographs of bald cypress, bald cypress-water tupelo (Nyssa aquatica), and bald cypress-red gum (Persea borbonia) swamps in Big Thicket National Preserve, Hardin County County, Texas. These various stands were used to represent forest cover types recognized by the Society of American Foresters (Eyre, 1980). The overall forest vegetation was bald cypress-water tupelo bottomland forest on soils that are more-or-less permanently inundated with water. Such wetlands that are dominated by trees have traditionally been defined and described as swamps.

109. Bald cypress- Consociation of bald cypress, including knees and regeneration of bald cypress. Water tupelo was an associate species in this stand of slightly deeper water. Water oak was also "among the numbered", but the number of species was extremely limited. Slough of Beech Creek, Beech Creek Unit, Big Thicket National Preserve, Hardin County, Texas. May, later vernal aspect. FRES No.16. (Oak-Gum-Cypress Ecosystem). A variant of K-103 (Southern Flood-Plain Forest). SAF 101 (Baldcypress). Taxodium distichum Association (if and when recognized), Tupelo-Cypress Series (223.11) in Southeastern Swamp and Riparian Forest biotic community (223.1) of Brown et al (1998). South Central Plains- Floodplains and Low Terraces Ecoregion, 35b (Griffith et al., (2004).

110. Big Thicket Cypress-Tupelo Swamp- General view of a bald cypress-water tupelo (Nyssa aquatica) swamp deep in the Big Thicket portion of Texas Pineywoods. Structure, architecture, and species composition typical of this forest type.Second-growth forest so that trees lack size of old-growth patriarchs, but species composition was that of the climax forest. There was abundant regeneration of these two climax tree species. Understorey shrub was swamp cyrilla (Cyrilla racemiflora) which was also regenerating. Did not take long to describe this simple wetland forest community. Obviously the only range feed available was browse provided by the swamp cyrilla. Along margins of this tract of swamp an adjoining forest on slightly higher land and less hydric soil another forest community had developed that consisted of swamp chestnut oak or, as also known, cow oak and basket oak (Quercus michauxii), shortleaf pine, loblolly pine, sugar maple, red maple (Acer rubrum), sweet gum, and the small tree or shrub of the tallest lower layer known variously as musclewood, blue beech, or American hornbeam (Carpinus caroliniana). Soil association was a Caneyhead-Kenefick. Big Thicket National Preserve, Maple Creek Unit, Hardin County, Texas. May, late vernal aspect. FRES No.16. (Oak-Gum-Cypress Ecosystem). A co-dominant variant of K-103 (Southern Flood-Plain Forest). SAF 102 (Baldcypress-Tupelo). Nyssa aquatica Association (if and when recognized), Tupelo-Cypress Series (223.11) in Southeastern Swamp and Riparian Forest biotic community (223.1) of Brown et al (1998). South Central Plains- Floodplains and Low Terraces Ecoregion, 35b (Griffith et al., (2004).

111. Deep in the swamp; thick in the Big Thicket- In the deep interior of the legendary Big Thicket bald cypress and water tupelo formed a forbidding, mysterious, erie, etc. (adjectives and explectives abound) wetland forest. These two vertical photographs showed representative samples of this forest vegetation. The shrub in center foreground of second slide was swamp cryilla. Big Thicket National Preserve, Maple Creek Unit, Hardin County, Texas. May, late vernal aspect. FRES No.16. (Oak-Gum-Cypress Ecosystem). A co-dominant variant of K-103 (Southern Flood-Plain Forest). SAF 102 (Baldcypress-Tupelo). Nyssa aquatica Association (if and when recognized), Tupelo-Cypress Series (223.11) in Southeastern Swamp and Riparian Forest biotic community (223.1) of Brown et al (1998). South Central Plains- Floodplains and Low Terraces Ecoregion, 35b (Griffith et al., (2004).

112. Water tupelo-bald cypress swamp- Healthy natural regeneration but perennially standing water undoubtedly prevents other than rare browsing by white-tail deer (Odocoileus virginianus). Big Thicket National Preserve, Maple Creek, Hardin County, Texas. May, late vernal aspect. FRES No.16. (Oak-Gum-Cypress Ecosystem). A co-dominant variant of K-103 (Southern Flood-Plain Forest). SAF 102 (Baldcypress-Tupelo). Nyssa aquatica Association (if and when recognized), Tupelo-Cypress Series (223.11) in Southeastern Swamp and Riparian Forest biotic community (223.1) of Brown et al (1998). South Central Plains- Floodplains and(223.11) Low Terraces Ecoregion, 35b (Griffith et al., (2004).

113. Local stand of bald cypress and redbay (Persea borbonia)- There was some water tupelo present, but redbay was clearly co-dominant with water tupelo a "dim and distant" third among tree species while red maple limped in at fourth place. The major species of shrubs were sqamp cyrilla and swamp or dwarf palmetto. Little Pine Island Bayou, Big Thicket National Preserve, Hardin County, Texas. May, late vernal aspect. FRES No.16. (Oak-Gum-Cypress Ecosystem). A variant of K-103 (Southern Flood-Plain Forest). Variant of SAF 104 (Sweetbay-Swamp Tupelo-Redbay). Tupelo-Cypress Series (223.11) in Southeastern Swamp and Riparian Forest biotic community (223.1) of Brown et al (1998). South Central Plains- Floodplains and Low Terraces Ecoregion, 35b (Griffith et al., (2004).

114. Black tupelo (Nyssa sylvatica) swamp- Blackgum dominated a small swamp formed by backwater of the San Jacinto River. Yaupon and swamp or dwarf palmetto formed one to two shrub layers (depending on height of yaupon at different locations) in this wetland forest. Adjacent to this swamp were larger areas of less wet soils on which loblolly pine-mixed hardwood-palmetto forest developed. That forest range vegetation was covered under loblolly pine forests earlier in this chapter. Montgomergy County, Texas (backwater of the San Jacinto River). February, later hibernal aspect. FRES No.16. (Oak-Gum-Cypress Ecosystem). A variant of K-103 (Southern Flood-Plain Forest). SAF 102 (Baldcypress-Tupelo). Nyssa sylvatica Association (if and when recognized), Tupelo-Cypress Series (223.11) in Southeastern Swamp and Riparian Forest biotic community (223.1) of Brown et al (1998). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

115. Blackgum in the backwater- More detailed view of a black tupelo swamp showing species composition and structure of a fairly restricted cover type in the Pineywoods. In addition to the trunks of blackgum "spotlighted" there were various twisted woody vines of rattan or Alabama supplejack, yaupon holly, and swamp or dwarf palmetto in this wetland forest vegetation. Montgomergy County, Texas (backwater of the San Jacinto River). February, later hibernal aspect. FRES No.16. (Oak-Gum-Cypress Ecosystem). A variant of K-103 (Southern Flood-Plain Forest). SAF 102 (Baldcypress-Tupelo). Nyssa sylvatica Association (if and when recognized), Tupelo-Cypress Series (223.11) in Southeastern Swamp and Riparian Forest biotic community (223.1) of Brown et al (1998). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

Spanish moss or black moss (Tillandsia usneoides = Dendropogon usneoides) is a common--and conspicuous--component species of the wetland forests (both swamps and riparian forests) as well as surrounding upland forests and savannahs. It was included at this juncture for purposes of interest, variety, and as it seemed "fitting".

116. "Moss"-festooned limb in the Pineywoods- Trees in more southern portions of the eastern deciduous forest complex "sport" numerous species of epiphyte, " a plant that uses another plant, typically a tree, for its physical support, but which does not draw nourishment from it" (Allaby, 1998). Perhaps the most common and widely distributed epiphytic species of forests in southern North America is Spanish moss or, sometimes, black moss or blackmoss, or old-man's beard (Tillandsia usneoides = Dendropogon usneoides). Contrary to the misleading designation "moss" this epiphyte is not only not a moss but it is, in fact, an advanced vascular plant, a monocotyledon in the Bromeliaceae (the pineapple family). Other forests--the Olympic Peninsula rain forest is the classic case--do have true mosses elegantly draped (="festooned" is the popular word) from limbs and branches of trees. Other forest communities have hanging wisply from their branches so-called "moss" that are species of lichen. The California oak woodland is the classic example. Still yet other forest range types (eg. Oregon white oak forest and the Olympic rain forest) have both actual moss and lichen species as distinct components or even layers of their vegetation. Where Spanish moss is a member of various eastern deciduous forest cover types it is a conspicuous, even prominent, botanical component of the vegetation, especially given the species' rather indistinctive, "bland" arrangement of thread-like, gray-colored leaves and stems. In popular imagination Spanish moss figures more picturesquely in erie, deep woods like cypress and tupelo swamps as in the mystic Big Thicket. Actually the densest populations of this bromeliad are in trees growing not in forests but in the open and that have large, spreading crowns and where there is plenty of light for this chlorophyllous epiphye. This set of photographs was taken from ancient post oak and willow oak growing in open fields (but still in the area of the historic Big Thicket of Texas' Pineywoods). Liberty County, Texas. February.

117. Making a habit ot it- Habit (general or outer physical form) of Spanish moss. Strands of thread-like (filiform), grayish stems and leaves of Spanish moss form festoons of considerable size (attaining lengths measured in feet or even yards or meters) as they hang from branches of trees and sway in the slightest breeze. This lichen-resembling species generally lacks roots and instead uses scaly hairs on leaves to absorb water and mineral nutrients from the air (hence, another common name of "airplant"). Spanish moss is regarded as an atmospheric or atmospheric-type eipphyte. It is also a xerophyte (plants living in arid or extremely dry [xeric] habitats) having such xerophytic features as the Crassulacean Acid Metabolism pathway of photosynthesis and multicellular hairs on leaves that reflect excess light and reduce water loss in addition to capturing air-borne nutrients (Diggs et al., 2006, ps. 478, 480). Ephphytes provide one of the textbook examples of commensalism, a symbiotic relationship in which one organism or species benefits (positive effect) from the association (the commensal; in this instance, Spanish moss) while the other "pardner" (the host), trees or other plants functioning as support and growing space, is unaffected (neutral or no effect) by the relationship. The Spanish moss specimen introduced in the first photograph was growing on a willow oak whereas the Spanish moss plant in the second photograph was hosted by an ancient post oak. Liberty County, Texas. February.

118. In the thick of it- A sample of portions from three plants of Spanish moss showing the tangled arrangement of gray, filiform (thread-like; linear, slender and circular in cross-section) stems and leaves within the festoon produced by plants of this epiphytic and xerophytic monocotyledon. Spanish moss is frequently used as nesting material by various species of birds, reptiles, and mammals (from smaller rodents to farrowing sows of free-ranging, feral swine). Indians in both North and South America made miscellaeous uses of this widely distributed bromeliad. Industrial Age man has used Spanish moss for everything from floral decoration to stuffing and packaging material. Anecdotal and empirical evidence has indicated that Spanish moss can be a component of deer diets. Liberty County, Texas. February.

119. Strands of thread-like leaves- Two close-up views of Spanish moss to show the filiform shoots and individual leaves of this xerophytic epiphyte. These adult plants were of large, mature size but at pre-bloom phenology. Liberty County, Texas. February.

120. Two distinct plant communities comprising locally restricted vegetation types: 1) a swamp of water oak with bald cypress as an associate and 2) a maidencane (Panicum hemitomon) marsh designated as a lowlands range site. These two types together constitute a flatwoods pond. FRES No. 16 (Oak-Gum-Cypress Ecosystem) and corresponding K-101 (Southern Flood-Plain Forest) and FRES No.41 (Wet Grasslands Ecosystem) with with no Kuchler units small enough to pick up the maidencane type. Maidencane would be included with Kuchler-83 (Everglades) in Florida. The maidencane marsh type is SRM 819 (Freshwater Marsh and Ponds). Mixed Hardwood Series in Southeastern Swamp and Riparian Forest biotic community and Maidencane Series (if and when such is recognized) in Southeastern Marshland biotic community, respectively, of Brown et al. (1998). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

121. Edge of two wetland range communities- Boundary between the water oak-bald cypress swamp and maidencane swamp introduced in the preceding slide. The swamp portion of this flatwoods pond was FRES No. 16 (Oak-Gum-Cypress Ecosystem) and corresponding K-101 (Southern Flood-Plain Forest) while maidencane marsh was FRES No.41 (Wet Grasslands Ecosystem) with with no Kuchler units small enough for this region so that instead maidencane would be included with Kuchler-83 (Everglades) in Florida. Maidencane rangeland cover type was SRM 819 (Freshwater Marsh and Ponds). Swamp with water oak dominant and bald cypress the associate species comprised a combination or "hybrid" of SAF ted

122. Maidencane in the spring- Hardin County, Texas. May, late vernal aspect. Maidencane Series (if and when such is recognized) in Southeastern Marshland biotic community, respectively, of Brown et al. (1998). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

123. Bay-Gall Bog or Titi in the Texas Big Thicket- This vegetation is the most impenetrable “jungle” or “tangle’ in the Big Thicket. The local mound-and-intermound relief creates a bog ecosystem. The soil series of the mound microrelief (on the mound) has the spodosol soil series Babco. This is currently the only spodosol mapped in Texas. The dominant plants are red bay (Persea borbonia) and sweet bay or swamp bay (Magnolia virginiana) among the hardwood trees and shortleaf and loblolly pine from the conifers. Gall, swamp cyrilla or, by the Indian name, titi (Cyrilla raecmiflora) is the dominant species of the shrub layer along with gallberry (Ilex coriacea; not to be  confused with the preceding gall), bull-briar (Smilax bona-nox), saw-brier (S. glauca), buttonbush (Cephalanthus occidentalis), and wax myrtle (Myrica cerifera) dominate the shrub layer. Completing this “tangle” is the herbaceous understory often dominated by rather rank-growing ferns. The largest trunk (in center) is a loblolly pine, the trunk immediately behind and to the right of it is a water oak, the two trees immediately behind and to the right of the water oak are sweet bay magnolias, and the left foreground tree is a red bay. Most of the shrubs in the foreground understory are swamp cyrilla or titi. Hardin County, Texas. May. There is no specific FRES or Kuchler for this local community that grows within the FRES No. 13 (Loblolly Pine-Shortleaf Pine forest Ecosystem). Mixed Hardwood Series in Southeastern Swamp and Riparian Forest biotic community of Brown et al. (1998). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

124. Interior of a Texas Big Thicket Bay-Gall Bog- Detail of the shrub layer described in the preceding slide caption. Note the seedling or young tree stage of loblolly pine in the foreground and the adult loblolly pines in background indicating that this is the dominant conifer for this unique local community. Hardin County, Texas. May.

125. The floor or herbaceous layer of a Bay-Gall Bog dominated by ferns. Over 20 species of ferns are native to the Big Thicket and there are another four or five species that may have naturalized here. The ferns are growing on a mound of Babco soil. Hardin County, Texas. May.

126. Profile of Babco soil (the only spodosol mapped in Texas)- Spodosols comprise the  soil order characterized by having  a light gray eluvial horizon over a reddish aluminum- and/or iorn-enriched horizon. They typically occur in humid areas.  The Babco pH ranges from 3.1 to 3.6. Hardin County, Texas. May.

127. Climax Loblolly Pine-Oak Hardwoods Forest- This bottomland Pineywoods is deep inside the Big Thicket and at or, at least, approaching the state of old-growth. It is on the first terrace above Beech Creek and is an edphic climax community of the region with a characteristic open, sometimes bare, understory of grasses in the Panicum, Paspalum, Uniola, and Andropogon species. The three mature trees are (front to rear) water oak, loblolly pine, and cherrybark oak (Quercus falcata var. pagodifolia) whose big limbs form a spreading crown. The adult tree in the background and appearing immediately to the right of the loblolly pine and the small tree adjacent to and, from this angle,  appearing to sprout from the water oak are swamp chestnut oaks (Q. michauxii). The small tree at far left opposite the branched cherrybark oak is a young willow oak (Q. phellos) whose branches are interwoven with those of another water oak just to the left of the field of view. The trunk immediately to the right of the loblolly pine whose upper portion is adjacent to the water oak is the rotting snag of some tree that lost the struggle for the most limiting resource, light. The background vegetation is a Bald Cypress Swamp in the floodplain of Beech Creek. About 200 yards from this site there is a sandjack or bluejack oak-sandhill bluestem scrub type that formed from aeolian sand carried up out of the Beech Creek bottoms over geologic time. The sandjack or bluejack cover type (a variant of SAF 72) was covered in the following section. Beech Creek Unit, Big Thicket National Preserve, Hardin County, Texas. Vernal aspect, May. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem), K101 (Oak-Hickory-Pine Forest), SAF  82 (Loblolly Pine-Hardwood). South Central Plains- Floodplains and Low Terraces Ecoregion, 35b (Griffith et al., 2004).

Sandjack= bluejack oak (Quercus incana) Scrub Forest or Woodland

Sandjack or bluejack oak is one of several scrub oaks that constitute a forest cover type (SAF 72). The example of this range cover type presented here had developed on an upland approximately 200 yards from the bottomland Pineywoods presented in the preceding section.

128. Sandjack= bluejack oak (Quercus incana)-sandhill bluestem scrub type- The bluestem is a taxonomic complex of little bluestem, including the taxa often shown as Andropogon divergens or Schizachyrium scoparium var. divergens, and slender bluestem (Andropogon tener= Schizachyrium tenerum). A few post oaks are associates of bluejack oak. Composites and various prickly pears (Opunia spp.) are scattered throughout the bunchgrass sward. An aeolian ("blowsand") ridge community. Beech Creek Unit, Big Thicket National Preserve, Hardin County, Texas. May. FRES No. 14 (Oak-Hickory Forest Ecosystem). A variant of K-72 (Oak Savanna). One of the many forms of Southern Scrub Oak, a variant of SAF 72 (Southern Scrub Oak). A Scrub Oak Series of Brown et al (1998), but one was not shown for this region. Sandy upland variant of South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

Shortleaf Pine (Pinus echinata) Forests

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

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

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

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

132. Shortleaf pine-oak-hickory forest- These two views are of a shortleaf pine-dominated forest with the physiogonomy and morphological form characteristic of old-growth (or approaching that stage). The author interpreted this state of forest vegetation development as, most likely, subclimax or perhaps even climax for this site. The crowns and general habit of the large pines had the appearance of very old trees. The understorey was of oaks (primarily the less mesic species like post, blackjack and black oak rather than the more mesic species like northern and southern red oaks). The hickory species could not be identified positively, but it was likely black hickory (Carya texana), a species adapted to drier sites and a local dominant in this area. Sweeetgum (Liquidambar styraciflua), one of the pioneer species on disturbed sites, was very common. The oak, hickory, and sweetgum species appeared to have been suppressed by periodic fire. Perhaps this forest developed following a major fire and the fire-adapted shortleaf pine out-grew the oaks and hickories. The suppressed state of sweetgum would not be explained by the latter alternative. There was regeneraation of shortleaf pine. It was an uneven-aged stand or population of P. echinata, but pine reproduction was less than that of the hardwood species. Understorey shrubs included smooth sumac (Rhus glabra), trumpet creeper (Campsis radicans), Virginia creeper (Parthenocissus quinquefolia), redbud (Cercis canadensis), and poison oak (Toxicodendron radicans). There were two pronounced woody plant layers (two shrub understories). The herbaceous species comprised a single layer (excluding ground-surface species like mosses and lichens). This was the most species-diverse layer of vegetation. It included eastern gamagrass (Tripsacum dactyloides), longleaf wood oats (Uniola longispicata), beaked panicgrass (Panicum anceps), little bluestem (Andropogon scoparius), big bluestem (A. gerardii), and the composites, compassplant (Silphium integrifolium) and gayfeather (Liatris elegans). Obviously this was an amazingly species-rich and structurally diverse plant community. It had for dominants climax species from both the tallgrass prairie and the oak-hickory-pine forest. Ouachita National Forest, Scott County, Arkansas. July. Ouachita Mountains physiographic province. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Natural Vegetation Classification System of Arkansas for GAP Analysis Project, Natural Terrestrial Cover Unit 1.A.9.b.2 Evergreen Forest With Rounded Crowns Pinus echinata-Quercus marilandica-Quercus stellata. Oak-Pine Series of Brown et al. (1998). Ouachita Mountains- Fourche Mountains Ecoregion, 36d (Woods et al., 2004).

133. Shortleaf pine-oak-hickory forest near old-growth stage- Another view of the shortleaf pine-dominated forest with hardwood and herbaceous understories. Species were listed and the structure of this forest vegetation was discussed in the preceding caption. It will be emphasized again that the pine population was un-even aged, but recruitment of shortleaf pine was less than that of the oaks and hickories. The vegetation was obviously developing into a oak-hickory-shortleaf pine forest, but one with well-developed shrub and herbaceous layers composed of climax species. The large old-growthlike shortleaf pine on the far right side of this photograph had numerous horizonally spreading limbs and bare spire-form of top which was typical of mature trees growing on savannas. This condition, taken in conjuction with the evidence of a tallgrass understorey (big bluestem, little bluestem, eastern gamagrass plus prairie composites), suggested that this vegetation might have been a shortleaf pine-tallgrass savanna. There were several other older individual pines in this and the preceding slides that had crowns made up of several major limbs which in turn branched diversely, a pattern or arrangement which indicated that these trees had developed in full light and not in a forest. This habit or morphological form indicated that these oldest trees lacked the apical dominance-development shape typical of trees growing up through an established forest. Some of these older trees had limbs below the terminal extension of the bole that were actually larger than this central apex. These trees had well-developed boles and would produce such fine logs as to self-start a Stihl, but the crown form and structure resembled that of wolf trees (trees whose big, spreading crowns occupy more canopy space than that of neighboring trees with larger, more valuable trunks). Vegetation shown in these slides appeared to be a model example of the oak-hickory-shortleaf pine climax type at the subclimax stage which is most productive of high-quality conifer timber and forest range for wildlife and livestock. Ouachita National Forest, Scott County, Arkansas. July. Ouachita Mountains physiographic province. FRES No. 13 (Loblolly-Shortleaf Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forests). SAF 76 (Shortleaf Pine Oak). Natural Vegetation Classification System of Arkansas for GAP Analysis Project, Natural Terrestrial Cover Unit 1A.9.b.2 Evergreen Forest With Rounded Crowns Pinus echinata-Quercus marilandica-Quercus stellata. Oak-Pine Series of Brown et al. (1998). Ouachita Mountains- Fourche Mountains Ecoregion, 36d (Woods et al., 2004).

134. Arkansawyer pineywoods- Second-growth shortleaf pine (in effect, a planttion) established as artificial regeneration after the native shortleaf pine--white oak-northern red oak-bitternut hickory forest had been logged. Artificial regeneration (also, artificial reproduction) refers to a stand of young trees established by human seeding or planting of seedlings or cuttings (Helms, 1998). In this forest tract in the Boston Mountains the U.S. Forest Service planted shortleaf pine and managed the forest for that single species with the result presented here: shortleaf pine plantation. The natural pine-oak forest was converted into a one-species forest (at least, more-or-less so; there was clearly a sole dominant tree species). The natural co-dominant oak and hickory component had been supressed by silvicultural practices so that only the natural co-dominant shortleaf pine pine held forest dominance. The hardwoods had reproduced and were growing well in this forest, but at this point (which was approaching harvest age-size for pine) oaks and bitternut hickory were not of size to share in dominance and, thus, in defining this forest community. This tract of forest managed so as to have an unnaturally or atypical high cover and community dominance by one species, the economically most valuable species under current marketconditions, was an example of a forest plantation. The Society of American Foresters specified that a plantation was "a stand composed primarily of trees established by planting or artificial seeding" and with the specific notations that 1) plantations could have tree or understorey components from natural reproduction and 2) they could consist of single- or mixed spceies composition with either uniform or diverse structure, age classes, etc. (Helms, 1998). The example of shortleaf pine plantation shown here was a textbook example of those criteria. In short, this was a man-modified forest. It was a forest with native species that had been manipulated to produce the forest crop species of choice while other species were managed for secondary considerations (eg. habitat of white-tail deer, a secondary crop). In this sense, this forest range community was transformed by management from a unit of potential natural vegetation (climax) into a politico-socio-economic unit of vegetation, an Arkansawyer forest. Even though this was a forester's choice of forest, a forest modified by minds and hands of forestry professionals, at least one--a a glaring one at that--of these modifications was due to neglect or, perhaps more precisely, to human action that had "backfired". Exclusion of fire and/or, same thing with emphasis of inaction rather than action, lack of prescribed burning had allowed invasion of eastern red cedar (Juniperus virginiana). The high relative canopy cover of sweet gum was typical of forests recovering from disturbances such as fire, blowdown, or, in this instance, timber harvest. Sweet gum is a seral tree species and one with a tolerance rating of Intolerant. It is, however, very effective in growth competitiveness on a wide range of soils and soil moisture conditions (Burns and , 1990). Sweet gum can readily come to dominate forests that were reestablished through artificial regeneration like planting as well as those that are redeveloping through secondary plant succession. This is simply part of the forest sere whether natural or aftificial reproduction. Sweet gum cover and density can be reduced substantially by grazing (even grazers like cattle are remarkably effective), herbicidal treatment, and prescribed burning. Such silvicultural practices may or may not be desirable and included as part of forest management. Shrub species in this tract were listed in the next caption. Ozark National Foraeest, Newton County, Arkansas. July, estival aspect. FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Pinus echinata Association (if recognized) in Oak-Pine Series of Brown et al (1998). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004).

135. Pine plantation in Boston Mountains- Interior of a shortleaf pine plantation showing composition and structure of a man-modified shortleaf pine-oak-hickory forest. Silvicultural practices had produced a forest in which one conifer species dominated the vegetation rather than the natural state in which three or four tree species (and both hardwoods and conifer) are dominants. This situation was explained in the immediately preceding caption. Most of the understorey consisted of woody layers that were primarily different age classes of such climax dominants as white oak, northern red oak, black oak, bitternut or pignut hickory, mockernut hickory (Carya tomentosa) along with sweet gum, generally a pioneer tree species, and sassafras (Sassafras albidum), a large shrub or small tree species. Shrubs included flowering dogwood, eastern redbud, Carolina buckthorn (Rhamnus caroliniana), lowbush huckleberry (Vaccinum vacilians), poison oak, pison ivy, Virginian creeper, and blackberry (this latter probably of several species). Herbaceous species were generally absent (at least from a practical standpoint) which was undoubtedly due to the dense canopy produced by woody species. This photograph included some of the larger trees of native hardwood dominants for comparison to the larger individuals of shortleaf pine. For example, the two northern red oak that appeared at this angle to be growing side-by-side in left foreground. The next succeeding slide showed large diameter of some of the pines. Ozark National Foraeest, Newton County, Arkansas. July, estival aspect. FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Pinus echinata Association (if recognized) in Oak-Pine Series of Brown et al (1998). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004).

136. Oaks, out; pines, in (or foresters' choice)- Interior of a shortleaf pine-oak forest, a plantation, produced by artificial regeneration. This forest had obviously been managed at (or near) time of planting pines resulting in their noticably greater size than almost all individuals of other tree species. This greater size (both height and diameter) was more than could be accounted for by growth. Sweet gum, the most widespread and abundant hardwood species, could easily have outgrown shortleaf pine on such sites as could stump sprouts from coppicing oaks and root-sprouting hickories. Two of the conspicuous shrubs in foreground of this photograph were sassafras and Virginia creeper. Other shrub and tree species were listed in the preceding caption. Viewers attention was directed to the large diameter of shortleaf pine (about time for logging) and in comparison to that of the much smaller hardwoods. A similar comparison was presented in the preceding slide. Ozark National Foraeest, Newton County, Arkansas. July, estival aspect. FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Pinus echinata Association (if recognized) in Oak-Pine Series of Brown et al (1998). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004).

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

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

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

139. Virgin shortleaf pine-oak-hickory forest— One of the few remnants of old growth forest left in Texas is this shortleaf pine-white oak-chinkapin oak (Q. muhlenbergii)-shellbark hickory (Carya ovata)-pignut hickory (C. cordiformis) community seen here. There are several layers of vegetation including a second tree layer of young climax tree species and species like winged elm (Ulmus alata) and boisd'arc or Osage orange (Maclura pomifera) and a shrub layer of flowering dogwood, Arkansas traveler or pepperwood (Ampelopsisarborea), blackberry, gooseberry, and various wild grape vines. The prominent herb layer(s) include little bluestem, rosette panic grasses (Panicum spp.), slender- or longleaf wood oats, and scattered clumps of the native bamboo, giant cane (Arundinaria gigantea). Lennox Woods (donated by Kirby Lumber Company to The Nature Conservancy), Red River County, Texas. May, vernal aspect. FRES No. 14 (Oak-Pine Ecosystem).FRES No.14 (Oak-Pine Forest Ecosystem). Classic example of K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Oak-Pine Series of Brown et al. (1998). South Central Plains- Tertiary Uplands Ecoregion, 35a (Griffith et al., 2004).

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

140. Physiogonomy and external architecture- Overall views of a second-growth, uplnd shortleaf pine-mixed hardwood stand from surface of forest floor to near top of forest canopy. Dominant woody vines in this particular local stand were Small's, lanceleaf, or coral greenbriar (Smilax smallii) and an unidentified grape (Vitis sp.?). Smooth sumac (Rhus glabra), American beautyberry (Callicarpa americana), and yaupon. Common grasses were longleaf woodoats, the dominant herbaceous species overall, purpletop (Tridens flavus), and beaked panicgrass (Panicum anceps). Forbs were extremely scarce so as to be of no notable relevance except to record presence of bracken fern (Pteridium aquilinum). Davy Crockett National Forest, Houston County, Texas. October. FRES No. 14 (Oak-Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Pinus echinata Association (if recognized) in a Oak-Pine Series of Brown et al. (1998, p. 37) or, alternatively, Warm Temperate Forest and Woodland (123), Southeastern Deciduous and Evergreen Forest (123.1), Pine Series (123.12) of Brown et al. (1998, p. 38). South Central Plains-Southern Tertiary Uplands Ecoregion 35e (Griffith et al., 2004).

141. Internal structure and species composition- Two " photoplots" at progressively closer distance to range vegetation of a second-growth, upland shortleaf pine-mixed hardwood forest. The same shortleaf pine of relatively large size was in foreground of both slides. Dominant shrub overall as based on cover, density, and frequency was American beautyberry. Yaupon was a distant second based on these criteria. There were two shrub layers (upper and lower) in the forest range vegetation presented. The upper or higher shrub layer was very sporadic consisting of "here-and-there" isolated plants of flowering dogwood (left side of first slide), red maple (Acer rubrum) which, though a tree species, was represented as a shrub in this stand (left side of first slide), eastern redbud (right-center foreground of first slide and same plant at far-right in second slide). Common greenbriar (Smilax bona-nox) and an unidentified species of wild grape (Vitis sp?) formed a fairly continuous "botanical connection" from ground level to tree canopy. Davy Crockett National Forest, Houston County, Texas. October. FRES No. 14 (Oak-Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Pinus echinata Association (if recognized) in a Oak-Pine Series of Brown et al. (1998, p. 37) or, alternatively, Warm Temperate Forest and Woodland (123), Southeastern Deciduous and Evergreen Forest (123.1), Pine Series (123.12) of Brown et al. (1998, p. 38). South Central Plains-Southern Tertiary Uplands Ecoregion 35e (Griffith et al., 2004).

142. Species composition of a second growth shortleaf pine-mixed hardwood stand with emphasis on shrub layers- In this forest range vegetation as shown in the first photograph hortleaf pines (in background) were joined by sweetgum (foremost tree at far right margin) and white oak (center midground) with an understorey dominated by the low shrub American beautyberry with yaupon as associate shrub species. An irregular or sporadic herbaceous layer consisted of longleaf woodoats (overwhelmingly the dominant herbaceous species) accompanied by splitbeard bluestem (Andropogon ternarius) as the associate herb. Second and closer-in "photoplot" (second slide) presented this forest plant community from roughly the same camera point. Tree trunk in immediate foreground (left-center) was the sweetgum in far right margin of first slide. There was a sassafras seedling to right of this sweetgum. American beautyberry and yaupon (first and second, respectively, major shrubs in lower woody layer) were presented to better advantage than in the first of these two slides. Other ("also-ran") shrub species included common greenbriar; Small's, lanceleaf, or coral greenbriar, smooth sumac, winged sumac (Rhus copallinum), and an unindentified Vitis species. Red maple was present as large seedling and small sapling age classes so as to be part of this lower woody layer. Red maple was present as a small tree along margin of this forest tract (see below). Davy Crockett National Forest, Houston County, Texas. October. FRES No. 14 (Oak-Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Pinus echinata Association (if recognized) in a Oak-Pine Series of Brown et al. (1998, p. 37) or, alternatively, Warm Temperate Forest and Woodland (123), Southeastern Deciduous and Evergreen Forest (123.1), Pine Series (123.12) of Brown et al. (1998, p. 38). South Central Plains-Southern Tertiary Uplands Ecoregion 35e (Griffith et al., 2004).

143. Synopsis views of vegetational layers- Interior of a second-growth shortleaf pine-mixed hardwoods forest showing the lower woody (shrub) layer. There were more shrub species and consequently greater diversity in understorey structure than in the local forest vegetation presented in the immediately preceding "photoplots". Hardwood species present in forest vegetation presented here included white oak, sweetgum, sassafras, and, especially, red maple. This latter species was present primarily as large seedlings to small saplings except on perimeter of forest where it grew to small tree size (shown bwelow).Here both smooth sumac and winged sumac were present at height intermediate between tall shrubs such as flowering dogwood and eastern redbud and lower shrubs like American beautyberry, the dominant shrub overall. Shrub layers of the forest range vegetation in this local area was presented in the immediately succeeding pair of photographs at shorter focal length to better "sample" species composition and arrangement of this forest stand. Herbaceous species consisted mostly of grass species of which longleaf woodoats was "head-and-shoulders" above all others to rank as the dominant herb. Other important grass species included beaked panicgrass, rosette panicgrasses (Panicum species of the Dichanthelium section), purpletop, and splitbeard bluestem. Forbs were too limited to warrent remarks other than to note presence of an occasional plant of bracken fern, perhaps the most widely distributed plant species on Earth (greatest species range). Davy Crockett National Forest, Houston County, Texas. October. FRES No. 14 (Oak-Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Pinus echinata Association (if recognized) in a Oak-Pine Series of Brown et al. (1998, p. 37) or, alternatively, Warm Temperate Forest and Woodland (123), Southeastern Deciduous and Evergreen Forest (123.1), Pine Series (123.12) of Brown et al. (1998, p. 38). South Central Plains-Southern Tertiary Uplands Ecoregion 35e (Griffith et al., 2004).

144. More emphasis on shrubs- Interior of a second-growth, upland shortleaf pine-mixed hardwoods forest presented so as to show greater detail of the shrub layer. This local area was the same "as introduced in " photoplots" represented in the immediately preceding pair of slides. More details of the lower woody and herbaceous layers (s) of this local stand were visible in these "photoquadrants". Dominance was a matter of extremely localized groups because some of the major species grew as clonal colonies. Smooth and winged sumac were most pronounced of these, but Small's or coral greenbriar (immediate center foreground in second of these slides) was also an obvious clonal plant spreading by "rootstocks" (woody rhizomes). Likewise, American beautyberry (the overall dominant shrub) grows in dense populations due perhaps to its typically abundant yields of fruits (immediate center foreground of second slide). Other shrubs represented in these two photographs included poison ivy, an unindentified species of wild grape, and yaupon.Seedlings and small saplings of sweetgum and red maple were plentiful in this stand (they were especially noticable in the first slide). Herbaceous species were primarily grasses and included the dominant, longleaf woodoats, splitbeard bluestem, purpletop, beaked panicgrass, and rosette panicgrasses. the most conspicuous forb was bracken fern, but it was present as incidental, individual plants and did not form colonies or brakes. Davy Crockett National Forest, Houston County, Texas. October. FRES No. 14 (Oak-Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Pinus echinata Association (if recognized) in a Oak-Pine Series of Brown et al. (1998, p. 37) or, alternatively, Warm Temperate Forest and Woodland (123), Southeastern Deciduous and Evergreen Forest (123.1), Pine Series (123.12) of Brown et al. (1998, p. 38). South Central Plains-Southern Tertiary Uplands Ecoregion 35e (Griffith et al., 2004).

145. On the outskirts- The outermost edge of an upland, second-growth shortleaf pine-mixed hardwoods forest featuring a large red maple (immediate center foreground). Most tree trunks were shortleaf pine except for an occasional white oak and sweetgum. Shrubs included American beautyberry, yaupon, and Small's or coral greenbriar. Herbaceous species were not dectible in this photograph taken at such distance as to show most of the nice specimen of red maple. (Details of lower layers of understorey were featured in the next two two-slide sets.) Davy Crockett National Forest, Houston County, Texas. October. FRES No. 14 (Oak-Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Pinus echinata Association (if recognized) in a Oak-Pine Series of Brown et al. (1998, p. 37) or, alternatively, Warm Temperate Forest and Woodland (123), Southeastern Deciduous and Evergreen Forest (123.1), Pine Series (123.12) of Brown et al. (1998, p. 38). South Central Plains-Southern Tertiary Uplands Ecoregion 35e (Griffith et al., 2004).

146. Lower layers- Two "photoplots" of herbaceous and lower shrub layers of an upland, second-growth shortleaf pine-mixed hardwoods forest in the Texas Pineywoods. The first photograph presented beaked panicgrass and purple as local dominants of the herbaceous stratum (longleaf woodoats was the overall dominant herbaceous species; next set of slides). A species of rosette pancigrass was represented at far left. Yaupon, the overall associate shrub species, accounted for almost all cover in background of this first photograph. An unidentified species of wild grape was also present. The second photograph included seedlings of red maple and sweetgum along with smooth sumac and American beautyberry at less-than-its-usul abundance for the lower woody layer. A plant of bracken fern, the only forb of much abundance, was also included for viewers' interest. Davy Crockett National Forest, Houston County, Texas. October. FRES No. 14 (Oak-Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Pinus echinata Association (if recognized) in a Oak-Pine Series of Brown et al. (1998, p. 37) or, alternatively, Warm Temperate Forest and Woodland (123), Southeastern Deciduous and Evergreen Forest (123.1), Pine Series (123.12) of Brown et al. (1998, p. 38). South Central Plains-Southern Tertiary Uplands Ecoregion 35e (Griffith et al., 2004).

147. More detail of lower layers- Smaller "photoquadrants" of the lower woody and herbaceous layers of an upland, second-growth shortleaf pine-mixed hardwoods forest in the Texas Pineywoods. Longleaf woodoats was the star of this lineup and merited center-stage by nature of it being the dominant herbaceous on this forest range (across much of the Pineywoods Region for that matter). American beautyberry and yaupon with their characteristic leaves also were obvious in both photographs. Also present in the first photograph were beaked panicgrass and a rosette panicgrass as well as smooth sumac, all locally common species in this stand of forest vegetation. Davy Crockett National Forest, Houston County, Texas. October. FRES No. 14 (Oak-Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Pinus echinata Association (if recognized) in a Oak-Pine Series of Brown et al. (1998, p. 37) or, alternatively, Warm Temperate Forest and Woodland (123), Southeastern Deciduous and Evergreen Forest (123.1), Pine Series (123.12) of Brown et al. (1998, p. 38). South Central Plains-Southern Tertiary Uplands Ecoregion 35e (Griffith et al., 2004).

148. Older pines, more hardwoods, and denser shade- Another stand of upland, second-growth shortleaf pine-mixed hardwoods forest that was adjacent to the stand featured in this section was made up of somewhat larger (resumedly older) shortleaf pines at conspicuously greater density and canopy closure. The downed pine was the victim of a hurricane six weeks prior to time of photograph. The tree closest to this laid-low pine (in front and at left of the trunk) was a sweetgum. Smaller tree with light gray bark in center midground was a Texas or black hickory (Carya texana) readily distinguishable to and handily identified by the photographer (not discernable in photographs) due to presence of 13 leaflets (the only hickory in this area to "sport" so many leaflets). The only shrubs present with any remarkable cover were American beautyberry, dominant shrub of this forest plus flowering dofgwood and eastern redbud, those stewarts of the upper shrub layer across much of the eastern deciduous forest. All three of these shrub species provide mast for eastern wild turkey, the critical animal species for which this forest tract was being managed. Shade was too dense to permit much grass cover. Even the longleaf woodoats, the dominant herbaceous species, had been pretty much excluded from this "sylvan party". Davy Crockett National Forest, Houston County, Texas. October. FRES No. 14 (Oak-Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Pinus echinata Association (if recognized) in a Oak-Pine Series of Brown et al. (1998, p. 37) or, alternatively, Warm Temperate Forest and Woodland (123), Southeastern Deciduous and Evergreen Forest (123.1), Pine Series (123.12) of Brown et al. (1998, p. 38). South Central Plains-Southern Tertiary Uplands Ecoregion 35e (Griffith et al., 2004).

149. Sunlite sample of structure- A small, local opening in a shortleaf pine-mixed hardwood second-growth forest afforded a view of forest structure and botanical composition of this foret range vegetation. In addition to the tall shortleaf pines there was a hardwood component of saweetgum, white oak, and black or Texas hickory in the tree layer. In addition there were two shrub layers: 1) taller layer of flowering dogwood and eastern redbud and 2) lower layer dominated by American beautyberry with yaupon as associate species of this stratum. There was an interrupted herbaceous layer dominated by longleaf woodoats. Davy Crockett National Forest, Houston County, Texas. October. FRES No. 14 (Oak-Pine Forest Ecosystem). K-101 (Oak-Hickory-Pine Forest). SAF 76 (Shortleaf Pine-Oak). Pinus echinata Association (if recognized) in a Oak-Pine Series of Brown et al. (1998, p. 37) or, alternatively, Warm Temperate Forest and Woodland (123), Southeastern Deciduous and Evergreen Forest (123.1), Pine Series (123.12) of Brown et al. (1998, p. 38). South Central Plains-Southern Tertiary Uplands Ecoregion 35e (Griffith et al., 2004).

150. Small's, lanceleaf, or coral greenbriar (Smilax smallii)- Common greenbriar (S. bona-nox) was comparatively more abundant than Small's greenbriar in the second-growth shortleaf pine-mixed hardwood forest featured in this section, but S. smallii was well-represented and afforded an opportunity to introduce viewers to another Smilax species. Davy Crockett National Forest, Houston County, Texas. October; autumnal aspect.

Miscellaneous Forest Types

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

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

The four slides presented immediately below were taken of an unusually mesic form of climax oak-hickory forest in the Boston Mountains section of the Ozark Plateau. The photographs were in the location specifically identified by Braun (Braun, 1950, ps. 170-172) as being an outlier or island of the Mixed Mesophytic Association (Braun, 1950, p.11) of the Mixed Mesophytic Forest Association but found in the Western Mixed Mesophytic Forest Association (Braun, 1950, p. 35). Braun (1950, p. 170-172) concluded that this specific forest vegetation was typical of that in the Cumberland and Allegheny Plateaus. Braun's conclusions were based on species composition, specifically of key species like beech (Table 33, p. 172) and local dominance into the climax by species like sweetgum. Beech was largely extirpated from this locale, but the combination of species mentioned by Braun including Kentucky coffeetree (Gymnocladus dioicus), American or white elm (Ulmus americana), and chinquapin oak (Quercus muhlenbergii) along with the typical sassafras, persimmon, and flowering dogwood as shrubs or understorey trees distinguished this as a unique community. Ecologically significant by their absence were post and blackjack oaks, and even black oak (Q. velutina), this latter the dominant species and key species over much of the Ozark Plateau. Commonness of black locust (Robinia pseudoacacia), classified as Very Intolerant, along with Intolerant species like sweetgum and Kentucky coffeetree were also indicators of a "choice blend" of the oak-hickory "brand". This was further verified by presence of northern red oak (Q. borealis= Q. rubra), southern red oak (Q. falcata), shagbark hickory (Carya ovata), and bitternut hickory (C. cordiformis), one of the more tolerant hickories Deemed by the author of substantial indicator value was commonness of wild hydrangea (Hydrangea arborescens), an understorey shrub limited to the most moist habitats such as seeps, springs, and north slopes. Relative abundance of this species and of hop hornbeam (Ostrya virginiana) along with the more typical poison oak and ivy, Virginia creeper, and pawpaw (yet nearly complete absence of herbaceous species) indicated an understorey that also varied from the typical Ozark Mountains oak-hickory forest. The conclusion reached by Braun (1950, p. 172) was :"These isolated mixed mesophytic communities are related to past forest migrations. Their preservation here, in a region whose physiographic history is similar to that of the Cumberland Plateau, is significant." This was an example of the point made by Braun (1950, p. 34) that each of the climax associations which characterize a specific forest region also occur in other forest regions characterized by, and thus named after, another climax forest association. This illustrated the dual nature of a Clementsian association: it was both an abstraction (abstract concept) and an actual climax plant community depending on both 1) the context in which association was applied and 2) the precise spatial and temporal location of the vegetation. The specific forest vegetation shown in this three-slide series illustrated a forest outlier, "an area of forest separated from the main occurrence of its type generally because of some local variation in ecological conditions or past migration of vegetation associated with major climatic changes" (Helms, 1998). Braun (1950, p. 172) specified that this forest outlier was largely a product of "past forest migrations". The following three photographs were taken on the upper terraces of the Mulberry River south of the community of Cass in Franklin County, Arkansas on a moderately steep northeast slope. July. The closest reference for native plant communities in Arkansas is that of neighboring Missouri (two counties north of the vegetation shown in this series) by Nelson (1987) who named and described forest natural communities as to either upland or bottomland forests. These two general groups were then divided on edaphic features such as depth, soil moisture and parent material. From this base the white oak-red oak-hickory forest introduced below would be either Mesic Forest (Nelson, 1987, p. 21) or Mesic Sandstone Forest based on the geologic aspect of the sandstone-capped Boston Mountains and absence of shortleaf pine (Pinus echinata) found on Dry-Mesic Forest (Nelson, 1987, ps. 37-38. According to the elaborate (and confusing, to this author) Natural Vegetation Classification System of Arkansas for GAP Analysis Project the Natural Terrestrial Cover of this forest was:1.B.3.a.6 Quercus alba-Carya tomentosa-C. ovata listed under Temperate Lowland and Submontane Broad-leaved Cold-Deciduous Forest. Ahh, right. The U.S. Forest Service Forest Type and Management Type Code designation was White Oak-Red Oak-Hickory and 53 for Type Name and Code, respectively. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). Society of American Foresters general designation was SAF 52 (White Oak-Black Oak-Northern Red Oak) (Eyre, 1980), BUT this was much less accurate than the SAF 1954 designation of White Oak-Red Oak-Hickory. The SAF (Eyre, 1980, p. 42) explained that "as hickories seldom make up more than 10 percent of the stocking, they have been dropped from the type name and black oak, a more common component, has been added". This was a true statement if applied at a landscape or regional scale (ie. across the Ozark Mountains where this type is climax according to the SAF and where black oak is a common dominant), but it most certainly is not a true statement if applied at the stand scale. The stand scale was used in the current publication of photographs and descriptions because stands-- and not landscapes or larger units-- are all that can be photographed with any detail to show vegetation. As shown below, hickories were often not only the obvious dominant but the most tolerant species and those accounting for most regeneration. As such, the SAF number was used below with the specification that hickory was co-dominant. Furthermore, as noted seven paragraphs above, black oak was not common on this Boston Mountains location but instead was generally absent from this more mesic area whose forest vegetation was an island of the Western Mesophytic Forest Association.

153. Mesic white oak-red oak-shagbark hickory forest- A stand of shagbark hickory within the specific mesic form or community indicated. Tolerance of shagbark hickory-- as for all Carya species-- is apparently open to debate. Harlow et al. (1979, p. 251) rated shagbark hickory as "moderately tolerant" while Burns and Honkala (1990, p. 222) regarded it as "intermediate". Both authorities agreed that shagbark hickory produces a deep, rapidly penetrating taproot and that younger trees of the species respout prolificly. Shagbark hickory is a minor component of six forest cover types recognized by the Society of American Foresters (Eyre, 1980) and probably of others including the more mesic Beech-Sugar Maple Type (Burns and Honkala (1990, p. 221). Local dominance by shagbark hickory throughout this specific oak-hickory forest community in the Boston Mountains was one indication that this forest vegetation was a geologic-determined remnant or relict of the more eastern and mesic Western Mixed Mesophytic Forest Association as discussed immediately above. The trunk with the bleached-color bark in left background was one of many of the red or black oak species (subgenus Erythrobalanus) killed by an outbreak of the red oak borer (Enaphalodes rufulus). The center and foremost tree was red mulberry (Morus rubra) that, while not a rare species in the Ozarks, in combination with the other species of this community was yet another indicator of the botanical diversity and uniqueness of this specific vegetation. Understorey species were strictly woody and included flowering dogwood, sassafras, persimmon, wild hydrangea, poison ivy, smooth sumac, and Virginia creeper. Black locust as a small tree was present just to the right of the photograph. Interestingly, and ecologically significant, was the fact that the most common tree species that was regenerating in the understorey was shagbark hickory. This indicated that this species was indeed relatively tolerant. (See also the slide below of a white oak stand in this forest-- Ozark National Forest, Franklin County, Arkansas-- where regeneration beneath large, mature Q. alba was shagbark hickory.) Boston Mountains of Ozark Plateau. Ozark National Forest, Franklin County, Arkansas (near Cass). July. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak), but with hickory and not black oak. Oak-Hickory Series of Brown et al. (1998). Boston Mountains- Upper Boston Mountains Ecoregion, 38a (Woods et al., 2004).

154. Mesic white oak-red oak-shagbark hickory forest- Large and, thus, quite old shagbark hickories (two center, obvious trees) and white oak (two trees at far left) grew alongside an also-old sweetgum (leaves visible on far left, background trees behind and to left of hickories) for an unusual combination of species in the Ozark Plateau (Boston Mountains section). Understorey species were all woody plants with Virginia creeper dominant on the ground and with poison ivy growing up every tree trunk of any size. Shrubs throughout the community of this and other photo-plots in this series included persimmon, pawpaw, sassafras, flowering dogwood, and hop hornbeam. Wild hydrangea was common indicating the mesic nature of the general habitat. There was considerable sexual reproduction by shagbark hickory. Locally the red oak borer had destroyed many trees in the Erythrobalanus subgenus. Oaks in this group included both northern red oak (Quercus rubra= Q. borealis= combinations of both epithets) and southern red oak (Q. falcata). Q. velutina was conspicuous by its absence in this forest community as were post and blackjack oaks, but chinquapin oak was present in small numbers and cover in localized spots. In general, white oak was-- as is typical-- relatively more abundant on less mesic sites like south slopes while the various red/black oak species were more common on the more mesic sites, but there were many examples where all were "fully integrated" and grew side-by-side. Boston Mountains of Ozark Plateau. Ozark National Forest, Franklin County, Arkansas (near Cass). July. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak), but with hickory and not black oak. Oak-Hickory Series of Brown et al. (1998). Boston Mountains- Upper Boston Mountains Ecoregion, 38a (Woods et al., 2004).

155. Mesic white oak-red oak-hickory forest in Ozark Plateau- Here is a "sample" of the Mixed Western Mesophytic Forest Association "lost" a "fur piece" from it's Cumberland Plateau region here in the Boston Mountains section of the Ozark Plateau. It is a remarkably species-rich community in a small "plot". The half of a trunk on far left is of shagbark hickory. The four trees to the right of it and in center background were white oak. The largest tree on the right was bitternut hickory (C. cordiformis), often regarded as intermediate in tolerance and more tolerant than it's associates (Harlow et al. (9179, p. 263; Burns and Honkala, 1990, p. 194). It will be seen that there were several lower small branches coming directly off the trunk of this large tree (leaves on these and interlacing furrows on bark made identification possible) suggesting relative tolerance in a dense forest. Understorey species included Virginia creeper all over the ground and poison ivy growing to tops of large trees. All the usual understorey shrub/small tree species of this area grew on or close to this photo-spot, including smooth sumac, persimmon, flowering dogwood, and wild hydrangea. Hop hornbeam was least common. None of the early spring forest forbs, like mayapple for example, were visible. Grasses and grasslike plants were absent. Boston Mountains of Ozark Plateau. Ozark National Forest, Franklin county, Arkansas (near Cass). July. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak), but with hickory and not black oak. Oak-Hickory Series of Brown et al. (1998). Boston Mountains- Upper Boston Mountains Ecoregion, 38a (Woods et al., 2004).

156. White oak stand representing the white oak phase of mesic white oak-red oak-hickory forest- Here all the large trees were white oak. The largest tree was approaching size of old-growth and was ripe for harvest. Regeneration was almost exclusively hickory, mostly shagbark and some bitternut. This hickory reproduction dominated the understorey and excluded most of the shrubs and small trees of the lower woody layers. For understorey species see captions for three slides of mesic white oak-red oak-hickory in this same forest (near community of Cass in Franklin County, Arkansas) shown above. Ecological implications of this were unknown, but in this local area the Carya species appeared to be tolerant enough to regenerate in what was obviously a dense forest and crowded understorey. Boston Mountains of Ozark Plateau. Ozark National Forest, Franklin County, Arkansas (near Cass). July. FRES No. 15 (Oak-Hickory Froest Ecosysstem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak), but with hickory and not black oak. Oak-Hickory Series of Brown et al. (1998). Boston Mountains- Upper Boston Mountains Ecoregion, 38a (Woods et al., 2004).

157. Shortleaf pine-oak forest with a canebrake understorey, a pine-oak-giant cane savanna- This was a most unique composite vegetation. It was an oak-pine forest with tree and shrub layers that were diverse in their own rights to which was "added" a canebrake of giant or southern cane, the native bamboo (Arundinaria gigantea subsp. gigantea). There was no layer of vegetation (at least of macroscopic vascular plants) beneath the canebrake which is the typical condition for canebrakes of this species. Transeau et al. (1940, p. 762) specified that canebrakes were "often invaded" by evergreen oaks. As described in the next paragraph evergreen species in this community were pines and the oak species were not "live oaks". This community was similar, however, to that described in a standard Botany text widely used decades ago (ie. old textbook knowledge). Prior to settlement by European man, giant cane formed vast "brakes" or colonies (as in endless miles of bamboo) along water courses ranging in size from major rivers to creeks throughout southeastern North America. These seemingly infinite canebrakes occurred as far north as the Ohio River and westward to central Texas. Giant cane also grew (and still grows) as isolated plants of one or a few shoots up to small colonies of several hundred square feet as part of the understorey vegetation in oak-hickory and pine-hardwood forests. An exclusive (a homogenous or "pure") stand of uninterrupted canebrake understorey beneath an adult tree canopy was-- in this vegetation hunter's experience-- quite rare. The species composition, structure, and physiogonomy of this forest-woody grass vegetation was likely that of the climax. This was relict vegetation. Even though it was not virgin forest this was interpreted as a climax community tht lacked the component of very old trees of the species present. It was one of the most unique assemblages of native plants encountered by your photographer-author. Shortleaf pine was the dominant tree, but there were a few individuals of loblolly pine (or perhaps of the widespread natural hybrid of these two species). Other trees included white oak, southern red oak, water oak, and sweet gum in about that order. There was a shrub layer composed of small individuals of persimmon and sassafras as well as smooth sumac. These shrubs and/or short trees grew in the canebrake, but were shorter than the taller shoots of the bamboo except at the edge of the canebrake where there was light of greater intensity and frequency. Pensacola bahiagrass (Paspalum notatum var. saurae) and hurrahgrass (P. pubescens= P. muhlenbergii= P. setaceum var. muhlenbergii) also grew on these extreme edges where they got adequate light for survival. The development of spreading, horizonal branches on all trees, including the dominant pines that are usually strongly apically dominant, indicated that these trees had developed on a savanna which was exactly what this community appeared to be: an oak-pine-giant cane savanna. The soil was moist enough to support water oak and the scarce loblolly pine (or a hybrid thereof), but soil water conditions were not periodically ponded let alone swampish. The grassland expression of Arundinaria gigantea canebrake (exclusive, homogenous stands or colonies of giant cane with widely scattered or infrequent trees) was covered in the Miscellaneous portion of the Grasslands section. Miller County, Arkansas. July. West Gulf Coastal Plain portion of the Coastal Plain physiographic province. Pineywoods vegetation, part of the Southern Pine Region. According to U.S. Forest Service Ecological Divisions (Forest Service, Final Environmental Impact Statement, 1999) this was in the Southeastern Mixed Forest Province (231), Mid Coastal Plains, Westen Section (231E), South Central Arkansas Subsection (231Ea). FRES No. 14 (Oak-Pine Forest Ecosystem). K-102 (Southern Mixed Forest). SAF 76 (Shortleaf Pine-Oak, variant of). No Kuchler or SRM units for canebrake. South Central Plains - Tertiary Uplands Ecoregion, 35a (Woods et al. 2004).

158. Shortleaf pine-mixed oak-giant cane savanna or a shortleaf pine-mixed oak forest with a giant cane understorey- A more interior view of the composite pineywoods vegetation displayed in the preceding slide. Interpretation of this vegetation was subject to many subjective factors including disciplinary background or professional allegiance. It could be viewed as a mixed oak-southern pine forest with an understorey comprised exclusively of a "brake" of giant cane, the one species of native North American bamboo. Alternatively this could be viewed as a combination of diverse pineywoods forest and giant cane or bamboo grassland. Beyond any debate these two "elements" (components) were present. Also beyond debate is the ecological fact that both of these vegetational elements do occur (they formerly occurred on massive scales in the virgin vegetation) in their own forms separate and distinct from each other as potential natural vegetation. It was a different form of canebrake; it was a unique developmental expression of oak-pine forest growing as it were "out of" or in the midst of a canebrake. The characteristic leaves of southern red oak appeared very distinctive along both the left and right borders of this photograph. Smooth sumac and shrublike or scrubby trees of persimmon and sassafras were overwhelmed in a "sea" of native bamboo. Canebrake of Arundinaria gigantea as a grassland form devoid of other than occasional trees was presented and discussed in some detail in the Miscellaneous portion of the Grasslands section herein. Miller County, Arkansas. July. West Gulf Coastal Plain unit of Coastal Plain physiographic province. Gorest Service ecological divisions given in preceding caption. FRES No. 14 (Oak Pine Forest Ecosystem). K-102 (Southern Mixed Forest). SAF 76 (Shortleaf Pine-Oak, variant of). No Kuchler or SRM unit for canebrake. South Central Plains - Tertiary Uplands Ecoregion, 35a (Woods et al. 2004).

159. Bottomland (floodplain) gallery oak-hickory forest- A "finger" of the eastern deciduous forest projects into the climatic or regional climax tallgrass paririe here in the Cherokee Prairie in the Osage Plains division of the Central Lowlands physiographic province. This gallery forest community is classified by the Missouri Natural Areas Committee (1987) as wet-mesic bottomland forest. It is dominated by pin oak (Quercus palustris) represented here by the largest tree with the light-colored trunk (center). Associated species also visible include: western hackberry (Celtis occidentalis), black cherry (Prunus serotina), persimmon (Diospyros virginiana), bois d'arc, red mulberry (Morus rubra), and silver maple (Acer saccharinum). Dominant shrub is Missouri gooseberry (Ribes missouriense). Herb layer is absent. Missouri State Prairie Park, Barton County, Missouri. June, early estival aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). One riparian form or part of K-73 (Mosaic of Bluestem Prairie [K-66] and Oak-Hickory Forest [K-91]). Variant of SAF 65 (Pin Oak-Sweetgum). Central Irregular Plains- Cherokee Plains Ecoregion, 40d (Chapman et al., 2002).

160. Oak-hickory forest- Landscape scale view in central Ozark Mountains of a white oak-black hickory (Carya texana)-black oak forest that is the mesophytic or climatic climax of this western-most extension of the deciduous forest proper of eastern North America. Shortleaf pine is an associate that is locally dominant. Post oak and scarlet oak (Q. coccinea) are also common upperstory associates. Flowering dogwood, persimmon, sumac (Rhus spp.), summer grape (Vitis aestivalis), and lowbush huckleberry (Vaccinum vacillans) dominate the shrub layer. The herb layer is composed of prairie grasses and forbs of the tallgrass prairie to the west. In addition to typical prairie species, a major legume component is present including tick clovers (Desmodium rotundifolium, D. nudiflorum), wild indigo (Baptisia leucophaea), and native Lespedeza spp. Classified as dry-chert forest by the Missouri Natural Areas Committee (1987). Christian County, Missouri. July, estival aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). Classic example of K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- White River Hills Ecoregion, 39c (Chapman et al., 2002).

161. Dormancy afforded a good "look-see"- With leaves on the ground instead of in the canopy an all-in-one view was provided of structure, arrangement, species composition, and lumber crop of a climax black oak-northern red oak forest full of mature trees. Black or Ozark pignut hickory and white or mockernut hickory (Carya texana and C. tometosa, respectively) were associate tree species. There were some post oaks, but this was clearly a forest site for the black oak species. There was also mentionable cover of black cherry, which probably indicated infrequent surface fires in this stand. Grasses were few in understorey, but dominant was broomsedge bluestem. There had been a history of cattle grazing in this stand, but it was generally light (mostly breechy ones looking for better pasture and finding worse than they left). Canopy was too dense for much herbaceous understorey other than for that of early season species like Mayapple (Podophyllum peltatum). Many of the larger trees in this stand were over-ripe and dying or even dead. There were several snags. However most regeneration was hickory. Ottawa County, Oklahoma. FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak). Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

162. Now a summer view- With leaves back up in the canopy a vastly different perspective was afforded of the same climax black oak-northern red oak of mature trees as introduced immediately above. There was ample regeneration of hardwoods, but hickory predominated. Ottawa County, Oklahoma. FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak). Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

163. Interior of the climax black oak-northern red oak forest presented in the immediately preceding two sets of two slides each. Abundant reproduction of hickory so that this stand was becoming a hickory phase or variant of the black oak cover type. Ottawa County, Oklahoma. FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak). Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

Forest remains: harvest of the black and northern red oaks-black and mockernut hickories forest- The tract of upland black and northern red oaks-black and white or mockernut hickories forest shown immediately above was highgrade-logged and basically clearcut in winter (and on the sly) 15 years after the preceding photographs were taken. The cutover land was re-photographed in July of the second spring-summer growing season post-harvest. Redevelopment of forest vegetation on this improperly logged, degraded tract was a combination of typical old-field plant succession beginning with pioneer or colonizing species including giant ragweed (Ambrosia trifida), horseweed or mare's tail (Conyza canadensis), hairy crabgrass (Digitaria sanginualis), and the naturalized annual, beefsteak plant (Perilla frutescens) along with climax dominants that were regenerating both sexually and asexually. Most asexual reproduction was in black oak (and some northern red oak) by coppicing from mid-size stumps. The other three species besides climax dominants that had appreciable cover were black cherry (Prunus serotina) and sycamore (Platanus occidentalis). Trees with noticeably less cover were , Kentucky coffee tree (Gymnocladus dioica), American elm (Ulmus americana), black walnut (Juglans nigra), and eastern red cedar (Juniperus virginiana). Surprisingly, the typical pioneer tree species, sycamore had less cover than would be expected judging from recovering vegetation on other severely disturbed sites in the immediate vicinity. Other tree species commonly found as pioneer or early seral plants included sassafras (Sassafras albidum) and persimmon (Diospyros virginiana). These two species were also less abundant than in the typical situation observed for disturbance in this vicinity. This cutover land had not been denuded to the extreme old-field state. Most regeneration of dominant climax trees was of the hickory species present as saplings before logging (described and ahown above). Most reproduction of oak (especially black oak) at this point in forest recovery was by stump-sprouting. There were few oak or hickory seedlings at this stage of secondary succession.Most seedlings were sycamore, black cherry, American elm, and sassafras. Some smaller though sexually mature trees that were left standing in the clearcut as well as uncut adult trees on the perimeter of the forest (see below) served as seed trees. Otherwise, re-establishment of oaks and hickories depended on coppicing and/or the soil seed bank. Shrubs were very important in this recovering vegetation. Far and away the most important shrubs were blackberry (probably including several Rubus species) that formed immense thickets that were summarily invaded by this author adequately equipped with buckets at berry picking time. Another important shrub (though less productive of tasty fruit) was smooth sumac (Rhus glabra). Fox grape (Vitis vulpina) and summer grape (V. aestivalis) had spread quickly on the cutover land. Eastern redbud (Celtis canadensis) and flowering dogwood (Cornus florida) that had already been growing in this forest remained minus minimal mechanical damage. This remarkably diverse recovery vegetation also included numerous species of forbs, both native and naturalized. The latter included the biennial, flannel mullein (Verbascum thapsus), and introduced legumes, both perennials such as white clover (Trifolium repens) and red clover (T. pretense) and annuals like Korean or Japanese lespedeza (Lespedeza striata). Other forbs included numerous natives like biennial evening primrose (Oenothera biennis); pokeweed (Phytolacca americana); and various composites, especially giant ragweed (Ambrosia trifida), common ragweed (A. artemisiifolia), Carolina elephant foot (Elephantopus carolinianus), Baldwin ironweed (Vernonia baldwinii), black-eyed susan (Rudbeckia hirta), Canada wild lettuce (Lactuca canadensis), and wingstem crown-beard (Verbesina alternifolia). The pioneering composite, horseweed or mare's tail was widespread and locally dominant although generally not in dominating proportions that might have been expected (and, probably, present in the first season following harvest). There was also some immature smartweed or knotweed plants (Polygonum sp.) that could not be identified as to species along with toothed spurge (Euphorbia dentata). Pokeweed was the most widespread and overall most important forb, but giant ragweed was a close second forming dense stands from which almost all plants of other species were excluded. The only common grass was hairy crabgrass. The perennial grasses were represented almost exclusively by broomsedge bluestem (Andropogon virginicus). Poverty oatgrass (Danthonia spicata) already present in the understorey persisted by larger stumps where it was not "overrun by pioneer species. There was also an occasional plant of purpletop (Tridens flavesus) next to shelter (eg. uncut tree).There were also some plants of green and/or yellow foxtail (Setaria viridis, S. glauca, respectively) which could not be distinguished at vegetative stages present at time of photographs. Likewise there were incidental plants of Carex and Cyperus species. White-tailed deer where the only species of large vertebrate that had access to this was black oak-northern red oak forest which had not been grazed/browsed by cattle or hogs for several decades. In the climax oak-hickory forest that approached old-growth state there were very few plants available for grazing and/or browsing in the understorey. These were limited to leaves and buds on regenerating oak and hickory species, flowering dogwood, poison oak/ivy, and incidental plants (trace amounts in absolute and relative cover) of blackberry, poverty oakgrass, and sedge. This tract of black oak-dominated upland Ozark Plateau forest was clearly transitory forest range. Once (after) canopy cover reached almost 100% almost no light that could penetrate throught the dense foliage to reach the ground (soil) surface. After leaves were fully grown in the forest canopy each spring light could only reach to the level delineated by lower leaves on shrubs like flowering dogwood and saplings of oak, hickory, black cherry, etc. In this climax oak-hickory forest with mature ("over-ripe") trees spring forbs like mayapple (Podophyllum peltatum) were not present except in natural forest gaps and at forest edges. Organization Note: a black oak-pignut or mockernut climax forest that served as permanent forest range was presented and described in the Use and Abuse of Oak-Hickory Forests section of Oak-Hickory Forests-II. Harvest of this forest was an example that socioeconomic factors often override biological ones, and that many if not most human endeavors (not excluding silvicultural operations) involve human emotions including greed and jealosy. Logging of this oak-hickory forest was a case of timber theft. A thieving son snuck in and stole this standing timber from his aged father who for romatic and aesthetic reasons wanted the forest with its many mature trees left as it was. The conniving son left uncut the trees around the edges of this forest to hide what he was doing inside the tract. The timber-buyer left some of the smaller trees inside the perimeter that he felt were not worth felling (probably to the chagrin of the greedy son who worked up all the slash to sell as fuelwood). Not only was this dysfunctional family relations, but it was also improper forest practices: cut-and-run, sloppy, (and illegal) logging of the worst form, the kind that gives the forest products industry a bad reputation. Thief of "free grass" and "timber for the taking" was standard fare in early history of use and abuse of forest and rangeland resources. The important thing for rangemen and foresters to bear in mind is that these resources are, to large degree, renewable (even with improper harvest methods and other forms of abuse). The following series of slides was of the black and northern red oak-black and/or Texas and mockernut hickories upland forest in the second growing season following highgrade logging that was, in effect, land-clearing.

164. Missing trees, stolen timber- The second-growth black oak-dominated (northern red oak, associate) forest shown and described here was secretly clearcut with a few smaller trees left around the perimeter so that a son could hide the timber theft from his aged father. The crime scene and second-year successional vegetation was presented in two photographs that gave a general view of the cleared forest. The wasted sound logs in the first of these slides added resource waste and abuse to the weed patch effect. In both slides remaining trees (a young northern red oak in the first; black and northern red oaks, Texas or Ozark and mockernut hickories in the second) "looked on" cutover land supporting a pioneer stage of recovering forest vegetation dominated by such colonizing species as the annual composite, giant ragweed, along with common ragweed, hairy crabgrass and such disturbance-loving perennials as pokeweed, black-eyed susan, Baldwin ironweed, wingstem crown-beard , Carolina elephant's foot. Foreign weeds that benefitted from this disturbance included flannel mullein and beefstake plant. Common shrubs included several species of blackberry, summer and fox grape, and smooth sumac. This stage of secondary plant succession on a black oak-dominated Ozark Highlands upland forest was about half-way into the second warm-growning season following clearcutting of a second-growth forest that approached old-growth status. Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak). Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

165. Into the weeds (and the ticks and chiggers)- Two close-in views of seral vegetation on a forest site that two winters or one an a half warm-growing seasons before had been a climax black oak-northern red oak (dominant and associate species, respectively) upland forest that was harvested by clearcutting (in effect a land-clearing operation). The most abundant herbaceous species were giant and common ragweeds. Other composite forbs included black-eyed susan, common horseweed or mare's tail, an Carolina elephant's foot. Pokeweed and flannel mullein represented native perennial and naturalized Eurasian biennial forbs, respectively. Passionflower (Passiflora incarnata), a perennial herbaceous vine or twining forb, was conspicuous along right margin (about half-way up) in the first slide. The only grass with cover and density worthy of note in the seral range vegetation presented in these two photographs was hairy crabgrass. Shrubs included blackberry, smooth sumac, and summer and fox grape. Tree species present as pre-existing (present before logging), stump sprouts, or seedlings included black oak, northern red oak, post oak, Texas or Ozark hickory, mockernut hickory, black cherry, American elm, red elm, sassafras, and persimmon. Young trees (either too small to make a saw log and/or serving as a cover to conceal the crime sceene from outside the clearcut) included all of the oak and hickory species as well as a few black cherry. Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak). Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

166. Edge effect, a benefit of forest harvest- Two more general views of a clearcut in a black oak-dominated climax forest in the Springfield Plateau. These two photographs high-lighted the effect of edge, the union of standing trees at perimeter of the now clearcut forest and seral range vegetation developing on the clearcut through secondary plant successtion midway through the second warm-growing season post-harvest. Some of the more abundant and conspicuous herbaceous species on this clearing included native forbs common to disturbed habitats: smartweed, Baldwin ironweed, pokeweed, giant ragweed, common ragweed. Most of these were of little or, at least, very limited forage value. The forb of most forage value was red clover, a naturalized, Eurasian, perennial legume (barely visible in the first photograph as pink clusters). How such abundant cover of this valuable forage plant developed by the second growing following logging remained a mystery. Farmers in this local vicinity have overseeded permanent pastures to red clover. Clearcutting had converted a climax forest with limited understorey (most of that regenerating hardwoods, especially hickory species) into a cutover pasture or range that, though a far cry from standards of high-quality tame pasture, provided valuable forage plants for livestock and wildlife including an introduced, perennial legume; a palatable, annual grass, and forbs of diverse palatability). This was transitory forest range. The second of these slides featured the numerous woody species that invaded the clearing the first growing season following clearcutting. Most conspicuous were two-year-old seedlings of sycamore (center of photograph). There were stump sprouts and seedlings of black oak, Ozark and mockernut hickories and black cherry. Most of the latter were seedlings or pre-established saplings. Prominent forbs in successional range vegetation shown in the second slide included Baldwin ironweed, giant ragweed, and common ragweed. Young vines of summer and fox grape trailed and wound their way around newly established large patches of blackberry out of which grew black cherry, sassafras, and elm (both American and red). In this arrangement of seral vegetation there were edges within edges:. edges where blackberry patches met weed (eg. colonizing composites, pokeweed, flannel mullein) patches within the edge that was the perimeter of the climax black oak-red northern oak forest. This was shown in more detail in the next set of two slides. In Nature any disturbance--no matter how traumatic or disturbing of existing species, communities, ecosystems, landscapes, etc--is a boon to some other species, communities, ecosystems, and so on. Clearcutting the climax, near old-growth black oak-northern red oak forest that occupied this site was an extreme perturbation, a drastic alternation of that forest ecosystem, that had dire consequences on many living things ranging from old trees and their dependent fungi species, squirrels, pileated woodpeckers (Dryocopus pileatus), and humans due to dissolutionous family relations between a father and a theiving son. That action, that extreme ecological disturbance, also created greately improved habitat for white-tailed deer and northern bobwhite (Colinus virginianus). One of numerous reasons why populations of bobwhite quail have declined throughout much of the Ozark Plateau Region is that their prime habitat of old-fields, recovering cleared forests, and small farm fields had revegetated back (had secondary plant succession progress closer toward) the pre-white man forest. Advanced seral, subclimax, and climax stage oak-hickory forests provide marginal to poor habitat for bobwhite. "Setting back" forest vegetation to pioneer and other early seral stages was (is) of immense benefit to seral species like bobwhite quail and white-tailed deer. When this climax forest (with its high proportion of rotten trees that were "overripe" for quality hardwood lumber) was logged pileated woodpeckers that lived off of insects that thrived on decaying wood as well as cavity nesting mammals including squirrels, 'possums (Didelphis virginiana), and coons (Procyon lotor) came out "loosers", at least in the short run. Critical parts of their habitat were eliminated. Perhaps this was the situation for wild turkey (Meleagris gallopavo) and raptors like great horned owls (Bubo virginianus) and red-tailed hawks (Buteo jamaciensis), all species that commonly lived in the black oak-northern red oak forest with its old age, over-mature trees. However, turkey and coon soon benefitted from increased production of blackberries, pokeberries, etc. Reduction of canopy cover facilitated predation by raptors although owls are still going to have to find dense tree canopy to escape their arch enemy, the annoying, sleep-depriving crow (Corvus brachyrhynchos). The 'possum will miss her favorite hollow tree, but see what a crop of persimmons will be produced for her descendants in a few years. The edge where uncut, timber theft-blocking trees meet and merge with the new weed and brush patch provided a new corridor as well as ample cover for new feed sources for various species of animals and new growing environments for plants. Use of natural resources always producers "winners"and "loosers". Wise use to conservation prophets like Gifford Pinchot consisted of finding that "blend" which, in the spirit of Jeremy Bentham's utilitarianism, "provided the greatest good to the greatest number for the longest period of time". As was shown in later descriptions and discussions of clearcutting this climax black oak-forest, it was quite likely that in the "long run" Pinchot's standard for wise use was fulfilled on this land, in spite of the villainous crime committed by a treasonous son against the rest of his family. Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak). Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

167. A closer look at edges and species make-up of seral vegetation- Two photographs showing seral forest range vegetation about half-way through its second warm-growing season growing adjacent to adult black and northern red oak trees that were left uncut to hid from view a forest clearcut of a climax black oak-dominated upland forest in the western Ozark Plateau. The plant communites that developed along the uncut forest perimeter occured spatially as "rows" (narrow zones) of woody plants (both tree saplings and shrubs) while there were larger patches of "weedy" colonizing species interior to the "belts" of wody vegetation. In the first photograph a narrow belt of adult black and northern red oaks was in the background while the foreground was an area dominated by giant and common ragweed. The bulk of the recovering forest vegetation (midground of photograph) was a blackberry patch with numerous saplings of black cherry, American and red elms, and sassafras (less persimmon) along with fox and summer grapes. The second photograph also showed a "weed patch" that was almost exclusively giant ragweed with some common ragweed and mare's tail orhorseweed (foreground) contiguous with a zone or "natural row" of blackberry patch with tree saplings and shrubs the major ones of which were smooth sumac, the two grape speceis, black cherry, American elm with some climax hardwoods (more hickories than black or red oaks). Such strips of seral woody vegetation are almost impenetrable to humans as the blackberry picking photographer attested. This is bobwhite habitat par excellence. Even pointers experience some problems getting through such brushy barriers. Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak). Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

168. Sere colonizers, both pioneer newcomers and climax holdovers- Two close-up "photo-plots" of seral range vegetation on a climax black oak-dominated upland forest site in wesstern Ozark Highlands that had been clearcut two winters previously. Recovering forest vegetation in these photographs was about half-way through the second warm-growing season after logging. The plant community of this pioneer or initial seral stage was a combination of colonizing species (giant ragweed, common ragweed, mare's tail or horseweed, hairy crabgrass, and beefsteak plant) native perennial composite forbs (Baldwin ironweed and Carolina elephant's foot), biennial forbs (flannel mullein and biennial evening primrose), native shrubs (blackberry, smooth sumac, summer and fox grape, Virginia creeper), colonizing trees or generally less tolerant tree species (sycamore, black cherry, sassafras, American and red elm), and seedlings and stump srpouts of climax trees (black and red oaks, Ozark or Texas and mockernut hickories). An amazing array of plant species in small local habitats. Seral range vegetation shown in the first slide consisted of a weed patch dominated by giant ragweed with Canada wild lettuce, biennial evening primerose, and greeen and/or yellow foxtail mixed with woody species ranging from black oak and hickory seedlings and stump sprouts to summer grape that had developed in front of an edge of woody vegetation made up of taller shrubs and young tree saplings. Shrubs included blackberry, smooth sumac, fox and summer grapes while saplings ranged from black oaks and two hickory species to American and red elms, black cherry, and sassafras. This was another example of the edge where different plant communities meet producing the edge effect that is so important for certain species of wildlife. Northern bobwhite quail was a species of great local importance that benefits immensely from this kind of seral range vegetation.. The second slide was of seral forest range vegetation dominated by forbs and seedlings and stump sprouts mostly of black oak and the two hickory species. Conspicuous forbs included giant ragweed, horseweed or mare's tail (many of which had upper parts of shoots grazed off by white-tailed deer), Canada wild lettuce (also grazed by deer though less so than mare's tail), toothed spurge, and biennial evening primrose. Red clover was also present but less conspicuous in spite of its showy inflorescneces than the taller-growing "weedy" colonizers. Tha annual grasses, airy crabgrass and green or yellow foxtail, were the representatives of the Gramineae in this photograph There was considerable cover of young Virginia creeper. The tree trunk in center midground was a black oak. Sprouts surrounding this black oak were mostly Ozark and mockernut hickories. Also in this slide was a hollow butt portion (roughly one ana a half foot diameter) of a black oak log that was later used for fuelwood. Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak). Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

169. Forgiving oaks- Four examples of stump sprouting or coppicing in black oak on a black oak-northern red oak upland forest in the western Ozark Highlands (Springfield Plateau section) midway through the second growing season for hardwood species following clearcutting. Coppice was defined by the Society of American Foresters (Helms, 1998) as: "1. the production of new stems from the stump or roots 2. to cut the main stem (particularly of broadleaved species) at the base or to injure the roots to stimulate the production of new shoots for regneration 3. a plant derived by coppicing 4. any shoot arising from an adenvtitious or dormant bud near the base of a woody plant that has been cut back". All four meanings were appropriate for the morphological/physiological phenomenon presented in the four black oak stumps and their second-season suckers. Helms (1998) also provided: "stool- a living stump (capable of) producing sprouts". Coppice shoots or stools (and related forms of vegetative reproduction including water shoots, suckers, and sprouts) are, of course, clones, ramets, modules or modular units of the parent plant, genet, entire vegetative plant, respectively. A simplified explanation is that stump sprouts are natural grafts or naturally formed scions arising from the stock (the stump or root). Forest regeneration by coppicing is a major silvicultural system (or component treatment thereof) for certain hardwood species. Black oak has not generally or typically been viewed as a coppice species or black oak lumber as a coppice crop to the extent as for some other species (eg. black cherry), but on the clearcut described in the above captions a high proportion of the felled black oak did coppice (sprout back from living stumps). In fact, this is the usual regeneration response of black oak, at least of the younger trees. Stump sprouting in black oak was described in Silvics of Trees of the United States (Fowells, 1965, p. 560; Burns and Honkala, 1990, Vol. 2.p. 747). The latter reported that roughly 95% of black oak regeneration in clearcutting was by sprouts from either stumps or "advance reprodution" (new sprouts from dormant buds near juincture of shoot and root). An interesting inverse relationship exist between stump size and successful stump sprouting. Shoots (sprouts= stools) from black oaks that were larger at harves grow faster than those of smaller cut trees; however, stump sprouting was inversely related to stump size, tree age, and forest site with larger stumps and those of older trees (often "same difference") having reduced sprout perduction. In other words stumps of bigger trees (ie. larger diameter stumps) and, hence, usually older trees are less apt to sprout, but if they do sprout these suckers (stump shoots) grow faster (have more rapid grow rates) than suckers from smaller stumps. This is a common--if not nearly universal--physiological response in hardwood species. Black oak stump-sprouts less readily than norther red oak which, as on this oak-hickory forest, is commonly associated with black oak. Fowels (1965, p. 591) cited research that reported over two-thirds of black oak reproduction in the Missouri Ozarks was "of sprout origin". Nonetheless, coppicing is the best possible means to replace 1) "original" (genetically identical) trees and shrubs and 2) climax woody species. In this regard, sawing logs amounts to the same thing as mowing shoots of perennial forage species for hay. With properly harvest the species composition of the tree component of the clearcut forest is essentially the same as it was before logging. In the instance of the upland Ozark Highlands black oak-dominated forest that served as the example for this lesson there was a cruel irony to the clearcutting operation in which a theiving son stole timber from his trusting father. The criminal act of forest harvest actually benefitted the black oak-northern red oak forest. The felled trees were still of an age and/or size that a high proportion of them coppiced (stump-sprouted) resulting in regeneration of both 1) the climax tree species and 2) reproduction of the same genetic individuals (ie. restoration of the identical harvested trees). Obviously the shoots (trunks= boles) of the mature (actually, over-mature) trees will not be restored because they were hauled of to the sawmill for pallats and framing lumber, but genetically these identical trees will regrow (grow back) so as to be be replaced as they were before forest harvest. There will be the same original root systems and basal trunks, the stumps, but morphologically new trunks. Based on canopy cover the proportion of black and northern red oaks in the recovering (seral) forest was less than in the climax forest, but climax tree species (and the same individual trees) were in the seral forest vegetation from the initial (pioneer) stage of plant succession (forest restoration). With asexual reproduction (coppicing) climax trees were present in the initial plant community following clearcutting rather than entering the forest sere later in the sequence of secondary plant succession as would be necessary if climax tree establishment depended solely of sexual reproduction (production of seedlings). Net result will be (barring other or continued severe forest disturbances) a more rapid return to the terminal stage of plant succession (ie. a shorter time interval to replace the climax black oak-northern red oak forest). Ironically, if the near old-growth black and red oaks (many of them already "overripe" with trunks partly hollow with heart rot) had remained standing for more years most of them would have been too old to coppice. Thus regeneration of the climax oak species almost assuredly would have been much lower and slower. Sexual reproduction (seedlings from acorns) of black and red oak is much less likely (much lower probability of successful tree establishment) than is asexual reproduction by coppicing. Furthermore, recall (from photographs and descriptions of the unlogged, nearly old-growth forest) that most tree reproduction in this climax black oak-dominated forest was of the associates, Ozark hickory (a small tree at maturity) and, secondly, mockernut hickory. In this forest of mature oak trees and in absence of natural disturbances like windthrow (blowdown) and fire the climax oak species were, through a combination of natural death due to old age and lower rates of regeneration, being replaced by replaced by hickory species. Clearcutting this black oak-dominated forest resulted in 1) regeneration of the climax oaks along with that of the associate hickory species, 2) increased plant biodiversity due to a combination of regeneration of climax species and colonization by pioneer ("weedy") species, 3) increased forage and browse production for livestock and deer, 4) improved habitat for bobwhite quail, and 5) caused loss of habitat for plant and animal species dependant on climax forest vegetation.

170. The black oak cover type- Interior view of dry chert upland forest dominated by black oak with post oak, northern red oak, blackjack oak, and black hickory as associates. There are two obvious shrub layers: 1) an upper one dominated by flowering dogwood (State Tree of Missouri; conspicuous here), redbud (State Tree of Oklahoma), and shadbush= eastern serviceberry (Amelanchier arborea) and 2) a lower one dominated by buckbrush= coral berry (Symphoricarposorbiculatus) and blackberry. Wild grape and Virginia creeper (Parthenocissus quinquefolia) form an aboreal shrub layer while poison ivy (Rhus radicans = Toxicodendron radicans) occurs in both shrub layers ranging from lianas extending to tops of trees to non-climbing thickets.The herb layer is usually limited to early spring species that complete their annual cycle before greening of the forest canopy. Mayapple (Podophyllum peltatum) is the conspicuous dominant herb. Head of hollow on a chert upland. April, early vernal aspect. Ottawa County, Oklahoma. FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak). Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005). IMPORTANT: As a general rule, browsing animals find deciduous shrubs and trees considerably more palatable than coniferous ones with the general response being that deciduous forests are much more prone to suffer damage, especially retarded regeneration, by overbrowsing than are coniferous forests. Understandably, foresters are reluctant to recommend (typically adamantly oppose) stocking livestock on hardwood forest types.Swine with their incessant rooting and feeding on mast are the livestock species that cause the most damage to these remarkably fragile range types.Proper livestock stocking rates on hardwood range are those described generically as “conservative”. These are forests that are usually most valuable for watershed and whose main crop or commodity is wood.HARDWOOD FORESTS ARE NOT "STOMP LOTS"!

171. Dormant but healthy- An Ozark Plateau upland forest (south slope) co-dominated by black oak and post oak that had received no livestock grazing for decades supported various age classes of trees. Other major trees included northern red oak, black hickory, black walnut, Kentucky coffeetree (Gymnocladus dioicus) and black cherry (Prunus serotina). Typical understorey shrubs included redbud, flowering dogwood, and woody vines such as various species of greenbriar and grape along with Virginia creeper and poison ivy (oak). This forest stand was so dense and had a nearly closed canopy so as to exclude development of an herbaceous understorey other than for early growing season species like mayapple. In most precise terms, the potential natural vegetation of this tree-dominated plant community was more woodland than forest per se. Climax vegetation would most likely consist of a more open or incomplete canopy cover (ie. tree crowns would not be interlocking). Nelson (1987, 2005) made a rational, well-written distinction between forest and woodland vegetation of the Ozark Plateau. The stand of black and post oak-dominated vegetation described here and immediately below were Dry-Mesic Chert Woodland (Nelson, 2005, ps. 190-193). Otawa County, Oklahoma. January. An upland forest of mixed oak and hickory species, but given overall dominance of this and adjoining forest stands the Society of American Foresters (Eyre, 1980) cover type that most closely fit this forest vegetation was Black Oak (SRM 110). Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Chapman et al., 2002).

172. Also dormant but not healthy- An Ozark Plateau upland forest (south slope) co-dominated by black oak and post oak that had been grazed by beef cattle for decades. This stand (if that term could be used loosely for comparitive purposes) was about 150 yards down a county road from the stand shown in the preceding slide. In addition to mature black oak, post oak, and, fewer, northern red oak (some of each species were on the ridge crest in background) there was a pole-size black walnut. There were also numerous and very conspicuous seedlings to small saplings (say, two to eight years in age) of eastern red cedar, an eastern juniper (Juniperus virginiana). There was zero regeneration of hardwoods of any species including the strongly smelling, usually unpalatable black walnut. This stand was a degraded Dry-Mesic Chert Woodland (Nelson, 2005, ps. 190-193) with potential natural vegetation for this forest site being an open or sporadic (vs. closed or complete) canopy of an actual forest having interlocking crowns. Foresters and rangemen would still management this as a stand of hardwood trees capable of producing high-quality oak, hickory, black walnut lumber as as having a grazable understorey for light stocking of livestock and habitat for wildlife including white-tailed deer, bobwhite quail, and squirrels. Ottawa County, Oklahoma. January.

173. Older hardwoods and younger cedars (or Where are the young hardwoods?)- The same "stand" of Ozark Plateau upland forest (south slope) co-dominated by black oak and post oak as presented in the immdeiately preceding slide. Large, two-trunked tree in center foreground was northern red oak. Extreme overrgrazing/overbrowsing for unknown decades (probably half a century or longer) had prevented regeneration of hardwood species. This management of a hardwood stand as a "stomp lot" had taken place over such a long time that even pole-size trees were absent from these "woods". Overgrazing/overbrowsing by cattle had not only been responsible for failure of hardwood reproduction, but this poor forestry (and animal husbandry) practice had also prevented periodic surface fires in what would otherwise have been an oak-hickory-walnut forest. Removal of essentially all herbage and prevention of production of fine woody material pre-empted light forest burning (ie. there was no fuel). Meanwhile birds that had eaten the fleshy seeds of eastern red cedar on rocky north slopes and bluffs above a nearby creek perched in and defeacted cedar seeds from the mature oak and black walnut trees. This avian behavior resulted in establishment of young cedar seedlings and saplings in the understory. In absence of fuel for light surface fires eastern red cedar was becoming established as the new forest cover type (SAF 46, Eastern Redcedar). Cattle will not eat eastern red cedar even inside corrals (or "cowpens" as such enclosures are called by many Ozark hillfolk). Barring disease these eastern junipers are safe-- at least until they become so large and close together (adequate canopy cover) that an accidental fire can spread almost instaneously through their crowns (ie. a crown fire, which is what any self-respecting rangeman would be hoping for in this degraded forest range site). This is horrid mismanagement of resources resulted in anthropogenic vegetation that was textbook case of grazing disclimax. In fact, this stand fit perfectly the description of the Eastern Redcedar forest cover type (SAF 46) by the Society of American Foresters (Eyre, 1980, p.50-51). Ottawa County, Oklahoma. January.

174. "In the Good Ole Summertime" but still "sick"- Sleek cattle and green leaves do not change the fact that this should-be or one-time forest is a degraded plant community (again, notheing but a "stomp lot"). What should be an oak-hickory forest with miscellaneous hardwoods such as black walnut, black cherry, and Kentucky coffeetree become a degraded pasture of mostly Eurasian annual grasses and a few mature trees of climax species from the previous forest stand. Cattle through overgrazing and overbrowsing prevented regeneration of the hardwood trees and also precluded light surface fires that would have killed these non-sprouting junipers and benefitted former climax understorey grasses. Yes, it is true that a climax or late seral stage oak-hickory forest like the stand that was about 150 yards down the road from this "mess" (and that was used as the control plot to present this lesson) would have little grazable/browsable understorey. A comparison of that late successional stage of forest vegetation with the "cow pasture" shown here would suggest to the neophyte that there is more "cow feed" on this degraded former forest. That is not true, not the case at all. The near-climax forest of the control plot is ready for logging. Following harvest of oak, hickory, and walnut logs, native grasses (big bluestem, Indiangrass, and purpletop are the main ones) and numerous shrubs (including blackberry, sumac, buckbrush, and wild plum) as well as regenerated hardwoods (mostly seedlings with some stump sprouts) will soon become re-established and provide range forage and browse for livestock and wildlife (notably white-tailed deer and bobwhite quail). Most importanlty from a forest perspective is the fact that the wood crop (hardwood logs are the agricultural commodity) is a source of revenue along with cattle (feeder calves and cull cows) and wildlife (either as recreational products or a sources of income from egress fees). Over the longterm, a properly managed oak-hickory forest will generate more revenue and produce more resources and commodities than this degraded barnyard with shade trees. Even the latter will eventually die to be replaced by juniper which will be a fire hazard by that time. Of course, agricultural producers are the world's greatest and most sustained optimists. In that spirit one can always hope that the eastern red cedar will escape crown fires and live long enough to grow into lumber that will fetch a fair price and that can be made into fragrant cedar chests to sell to tourists in flea markets. If this owner wanted nothing but pasture for these cows and calves the proper farming practice would be to plant this land to introduced (agronomic) pasture grasses like bermudagrass (Cynodon dactylodon) or tall fescue (Festuca arundinacea), both of which are well-adapted to these shallow rocky Ozark hills. The owner could then properly manage this tame pasture for economical production of beef cattle. Instead and as it was this landowner had nothing but "bragging rights" to running some cows and a lot less income than if he had wisely managed his forest, range, and livestock resources. This joker had not done justice to the revered title of "hillbilly" (just plain "hick" about covered it). Ottawa County, Oklahoma.

175. Some respectible-size hardwoods on a rock pile- A climax oak-hickory forest with black oak (two foremost trees on the left; two left foreground trees) the most common tree species along with black hickory and bitternut hickory (Carya cordiformis) and even one blackjack oak (rightmost larger tree). The large tree with the high scar (tree behind foremost tree on left) was a bitternut hickory with a DBH of 24 inches. Not much herbaceous understorey but big bluestem and broomsedge were main species. Flowering dogwood (left margin; just coming into bloom) was major species of the upper shrub layer. A second or lower shrub layer consisted of buckbrush, blackberry, and Virginia creeper, this latter of which covered much of the ground surface and also reached up into tree crowns so as to be in both shrub layers. Grape (right foreground) also extended in both shrub layers. Very marginal land (Land Capability Unit #8). April, early vernal aspect. Ottawa County, Oklahoma. FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak). Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

White Oak (Quercus alba) Forests

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

176. The white oak cover type- As stated by the Society of American Foresters (1980), the white oak forest cover type is "pure". In classic Clementsian terms this primarily a consociation (certainly in the photo-plot presented here). Seen here is a stand of vigerous young white oaks on a moist north slope in the Missouri Ozarks. The dominant herb is the widespread composite, black-eyed susan (Rudbeckia hirta). Also visible is the unique natural spiderwort hybrid (Tradescantiaozarkana X T. ernestiana). The main shrub growing amidst the oaks is flowering dogwood. Roaring River State Park, Barry County, Missouri. May, vernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). White Oak form of K-91 (Oak-Hickory Forest). SAF 53 (White Oak). Quercus alba Association (if recognized as such) in Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Dry-Mesic Chert Forest (Nelson 2005, ps. 125-130). Ozark Highlands- White River Hills Ecoregion, 39c (Chapman et al., 2002).

177. Members of an upland white oak-dominated forest- A species-rich upland forest community had developed on this upland Ozark Highlands location. At some local sites white oak formed a consociation. At other local sites white oak was joined by shagbark hickory (Carya ovata) and sugar maple (Acer saccharum) as associates. Other tree species included black oak, northern red oak, post oak (of course as nearly always present), western hackberry, sycamore, and honey locust (in that approximate order). Flowering dogwood comprised most of a lower woody layer other regenerating young trees of above listed species. An herbaceous layer at this particular location included shooting star or American cowslip (Dodecatheon meadia) and Christmas fern (Polystichum acrostichoides) along with the hybrid spiderwort specified in the preceding caption. Roaring River State Park, Barry County, Missouri. May, vernal aspect.FRES No. 15 (Oak-Hickory Forest Ecosystem). White Oak form of K-91 (Oak-Hickory Forest). SAF 53 (White Oak). Quercus alba Association (if recognized as such) in Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Dry-Mesic Chert Forest (Nelson 2005, ps. 125-130). Ozark Highlands- White River Hills Ecoregion, 39c (Chapman et al., 2002).

178. Deep in "them thar hills"- White oak-dominated upland forest in Ozark Mountains.Associates of white oak were (locally or at local site scale) shagbark hickory , sugar maple, black oak, and the ever-present post oak. Flowering dogwood was present throughout as the principal shrub species though it was not in bloom during this mid-spring season. Redbud was also present, but it was much less common than flowering dogwood. In these two "photo-plots" herbaceous were sparse and limited mostly to the hybrid spiderwort noted above. Roaring River State Park, Barry County, Missouri. May, vernal aspect.FRES No. 15 (Oak-Hickory Forest Ecosystem). White Oak form of K-91 (Oak-Hickory Forest). SAF 53 (White Oak). Quercus alba Association (if recognized as such) in Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Dry-Mesic Chert Forest (Nelson 2005, ps. 125-130). Ozark Highlands- White River Hills Ecoregion, 39c (Chapman et al., 2002).

179. Scond-growth white oak et al. forest with amazing species diverstiy- At head of a hollow deep in the Ozark Plateau white oak dominated a north slope of a Dry-Mesic Chert Forest (Nelson, 2005, ps. 125-130). Associate tree species were of the red or black oak group (Erythrobalanus subgenus): black oak and northern red oak. Hickory was common as was (though less so) black walnut. Sugar maple was also present in more mesic microsites. Dominant large woody understory species (large or tall shrub layer) was flowering dogwood. Understorey shrubs of a second or lower layer included wild hydrangia, blackberry, poison oak, grape, Virginia creeper, and lowbush huckleberry (Vaccinium vacillans) . The most abundant grass was hairy, downy, or silky wildrye (Elymus villosus) while the dominant forb was pokeweed (Phytolaca americana). Several species of tickclover (Desmodium spp.) were also widely distributed. Flag Springs State Park, McDonald County, Missouri. June, late vernal aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Most plausible Society of American Foreters (Eyre, 1980) cover type was SAF 52 (White Oak-Black Oak-Northern Red Oak) and this was predominant cover type of this locality: white oak was not as much as obvious dominant and defining species as was case for SAF 53 (White Oak) which this closely resembled. Oak-Hickory Series of Brown et al. (1998). Dry-Mesic Chert Forest (Nelson 2005, ps. 125-130). Ozark Highlands- Elk River Hills Ecoregion, 39b (Chapman et al., 2002).

180. White oak-black oak-noethern red oak forest- White oak clearly dominated this north slope of a western Ozark Plateau forest, but it was not cover or density domiance to the degree that white oak dominated the north slope shown in the preceding set of slides (Roaring River State Park, White River Hills Ecoregion). Development and species diversity of two prominent woody layers below canopy layer was greater and different from that of a more overwhelming dominance by white oak as described for the immediately preceding Ozark Plateau forest.. This north slope and forest hollow was substantially more mesic and had less light exposure than the preceding forest. Flowering dogwood was the dominant of the taller shrub layer. A lower and more diverse shrub layer included wild hydrangia, blackberry, poison oak, grape, Virginia creeper, and lowbush huckleberry. The herbaceous layer(s) was not as diverse as the white oak-dominated north slope forest described above, but the grass component (mostly downy, silky, or hairy wildrye) was much more productive and of far greater foliar cover on this mixed oak Ozark forest. Most common forb was pokeweed (which is more commonly a forest forb on locally disturbed areas). Flag Springs State Park, McDonald County, Missouri. June, late vernal aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Most plausible Society of American Foreters (Eyre, 1980) cover type was SAF 52 (White Oak-Black Oak-Northern Red Oak) and this was predominant cover type of this locality: white oak was not as much as obvious dominant and defining species as was case for SAF 53 (White Oak) which this closely resembled. Oak-Hickory Series of Brown et al. (1998). Dry-Mesic Chert Forest (Nelson 2005, ps. 125-130). Ozark Highlands- Elk River Hills Ecoregion, 39b (Chapman et al., 2002).

181. White oak-shortleaf pine forest- White oak is frequently the dominant oak in the shortleaf pine-oak type (SAF 76), especially on more mesic forest sites. By definition and description dominated more by pine than by oak (even with co-cominance). The forest shown here, and included under the White Oak section of this chaper, was clearly dominated by white oak with shortleaf pine ranging from being lesser of two co-cominants to the main associate species. In local stands of this forest in which white oak and shortleaf pine were co-dominant post oak was associate tree species. A list of shrub species in the forest community presented here was a long one. Flowering dogwood, sassafras, persimmon, grape, smooth sumac (Rhus glabra), shining or winged sumac (R. copallina), blackberry, redbud, and poison oak were good "for starters". Indiangrass, little bluestem, and Canada wildrye were dominant grasses in that order. The Eurasian orchardgrass (Dactylis glomerata) was also present. Hobbs Wildlife Management Area, Benton County, Arkansas.October, autumnal aspect. FRES No. 14 (Oak-Pine Forest and Woodland Ecosystem). K-101 (Oak-Hickory-Pine Forest). White oak-dominated variant of SAF 76 (Shortleaf Pine-Oak). Oak-Pine Series 122.14, Northeastern Deciduous Forest 122.1 of Brown et al., 1998, p. 37). Ozark Highlands- Dissected Springfield Plateau-Elk River Hills 39b (Woods et al., 2004).

182. Views of an Ozark "pinnery"- Hillfolk in the Ozarks traditionally refered to hardwood (most commonly oak-hickory) forest with pines (usually scattered individuals rather in groups) as a "pinnery". "Setting the woods afire" (often for the expoused purpose of "killing them *#&%** ticks" by white hillbillies was a lesson well-learned from the Indians and such flaming rituals of spring undoubted gave some competitive advantge to the more fire-adapted conifers. The sundown autumn scenes shown here from the western Ozark Highlands accurately represented a typical "pinnery". There were enough adult shortleaf pines and they were reproducing adequately to add a "pine flavoring" to the white oak-dominated form or phase of the Ozark oak-hickory forest. As the sun sets there's just time to do the chores, eat a leisurely supper, and then load up the hounds to spend an evening listening to the mountain music as Black and Tans, Blueticks, and Redbones inform us of their progress in pursuit of coon or fox. Bring plenty of crackers and sardines, boys. It'll be a fine fall night in the pinnery. Hobbs Wildlife Management Area, Benton County, Arkansas.October, autumnal aspect. FRES No. 14 (Oak-Pine Forest and Woodland Ecosystem). K-101 (Oak-Hickory-Pine Forest). White oak-dominated variant of SAF 76 (Shortleaf Pine-Oak). Oak-Pine Series 122.14, Northeastern Deciduous Forest 122.1 of Brown et al., 1998, p. 37). Ozark Highlands- Dissected Springfield Plateau-Elk River Hills 39b (Woods et al., 2004).

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

Bur Oak (Quercus macrocarpa) Forest

Bur oak as a forest cover type (SAF 42; Eyre, 1980, ps. 39-40, 236) is the most northward and westward of the eastern and central continental tree oaks. Bur oak comprises the only oak forest cover type in more northern portions of the Central Lowlands and Great Plains physiographic provinces. Bur oak is extremely tolerant of drought and fire and forms forests, groves and savannahs within (and within) the central grasslands of North America from the eastern edge of the Prairie Peninsula (Transeau, 1935) through to the Black Hills (Eyre, 1980, p. 39). Bur oak is to this vast region what post oak and blackjack oak are in the south (eg. Cross Timbers). These two forest range types meet near northern reaaches of the Flint Hills Region. Presented in this section was an example of the bur oak forest cover type (SAF 42) found on a north slope and outward to a ridge top in the Nebraska Sandhills. This was about as dry (least mesic) an environment as bur oak is adapted to, at least to the extent of forming forest communities. This example was a forest and not a woodland or savanna although at its outer edges it did form small groves with tree density and understorey more suggestive, or even typical, of woodland. This contiguous bur oak-dominated community (both forest and grove phases, or two communities if so interpreted) was the climax (ptential natural) range vegetation on this sere except where frequent, close, mechanical mowing had converted the native herbaceous understorey to a manmade one made up of Eurasian perennial grass species adapted to such intensity and frequency of defoliation. Examples of both native and naturalized herbaceous layers were presented following a brief section that introduced the foliage and fruit features of bur oak. This relict stand of bur oak forest had developed on what was at this time the Valentine, Nebraska city park (and almost any city park says it all). This forest range vegetation occupied both upland, including and especially a mesic north-slope, and bottomland site. Vegetation on all but the north-slope was highly modified by yard-mowing, manicure-it urban man (even in this over-whelmingly rural region). The natural herbaceous understorey of the upland phase was Canada wildrye (Elymus canadensis) except on north-slope upland forest range where the dominant herb was—as on the bottomland phase—long-beak(ed) or Sprengel’s sedge (Carex sprengelii). It was likely that there was also some bristleleaf or ebony sedge (C. eburnea) was growing in association with long-beak sedge, but C. eburnea could not be positively identified given absence of inflorescences. Under frequent, close mowing Kentucky bluegrass (Poa pratensis) dominated the lowland understorey. With less frequent mowing smooth brome (Bromus  inermis) dominated both bottomland and upland phases. Both of these introduced Eurasian grasses have naturalized widely and under a farmer (vs. forester or rangeman) frame of mind, and management consistent therewith, the native grasses and sedges were crowded out by aggressive, highly competitive agronomic forages. The timeless story since Cain and Abel. The farmer (cornhusker in the Cornhusker State) again won out over the pastoralist.

184. Big tree on the prairie- Beautiful and massive specimen of bur oak that made its home in the floodplain of the Bosque River in the Western Cross Timbers of northcentral Texas. Bur oak is obviously a minor member--especially when compared to the dominant post and blackjack oaks--of the scattered forest and woodland communities in southern portions of the tallgrass prairie region. In more northernly parts of this vegetational realm bur oaks replace blackjack and even post oak as the dominant oak. In these northern parts of the Prairie Peninsula and tallgrass-true prairie region bur oak would be second--if that--only to eastern cottonwood as a dominant tree both on grassland and isolated forest communities such as gallary forests. Bur oak is the most widely distributed of all the oak species that have the role of hardwood dominant across the North American central prairies (McGregor et al., 1977, ps. 39-41). Post oak and blackjack oak barely extend northward to Iowa whereas bur oak extends to Ontario. Bur oak forms forests, woodlands, and groves (distinguished by tree density, extent of canopy cover, and size of stand resulting in varying degrees of understorey development) in more northern areas of North America as in the Nebraska Sandhills, an example of which was used for photographs and corresponding descriptions and explanations in this section. Hamilton County, Texas. July.

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

186. Leaves and fruit of buroak- Burr oak (either one or two "rs" are used) is regarded as having the largest acorn of any Quercus species in North America. Like other white oak species buroak bears fruit every year (versus a two-year cycle) in the red or black oaks. The combination of large acorns and production each year makes this species one of the most valuable producers of mast in the eastern deciduous forest (ie. the eastern part of North America). Burroak is also known as mossy-cup oak, both common names in reference to the conspicuous tapered tips of the scales of the cup which present a fringe-like appearance. The geographic range of. Q. macrocarpa extends from eastern Canada (New Brunswick and Quebec) to Texas. Hamilton County, Texas. September.

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

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

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

Technical Note: The following sequence of photographs of the Bur Oak forest cover type (SAF 42) was taken in early summer (late-June). Two aspects were presented and described: 1) north-slope and 2) hilltop. These represented two distinct aspect-based bur oak forest communities even though both were: 1) dominated by bur oak, 2) the same cover (dominance) type, and 3) upland forest. Both aspect-determined bur oak communities were photographed at the same time (about 0730 to 100 hours Central Standard Time). This time of day was most advantageous for pphotographing the north-slope forest which, having a general northeast orientation (hillside alignment), received most light in the understorey (maximum land surface area exposed to full-sun lighting; greatest understorey coverage of greatest light intensity) in early to mid-morning. Obviously time of daylight hours (excepting very early morning or late evening) was largely irrelevant for photographing vegetation of hilltop forest. Relatively high density of adult bur oaks on the north-slope forest coupled with dense individual crowns of this large-leafed species resulted in one of the darkest understorey habitats ever seen by this photographer. Most of the understorey area (square footage, yardage, meterage, etc.) of the north-slope forest range received somewhere between half to three-fourths of an hour of direct light daily. During the remainder of daylight time plants--especially herbaceous species--were in relatively dense shade and obtained only indirect light. Herbaceous species in bur oak forest have to be some of the best adapted and most extreme examples of sciophytes (skiophyes) or sciaphlic (skiaphilic) plants, "shade-loving" or plants that have evolved to shade (shady habitats), in the eastern deciduous forest. Viewers should realize that the shady (generally dark; poorly lite) images presented below, while of relatively poor viewing quality (compared to range types in other biomes), "caught" exactly normal light conditions as they existed, and at their brightest (time of day in which most light penetrated forest crown canopy to reach ground level). No artificial lighting was used. In other words, intensity and quality of light seen in photographs were those that the viewer himself would have experienced, the images that would have traveled through his cones, rods, and optic nerve, had he moved through this forest environment.One can but marvel at the adaptation of the herbaceous plants in bur oak forest range.

190. Sylvan scene in Nebraska Sandhills (Are you kiddin"?)- Send this as a postcard from the Nebraska Sandhills and those who deal in sterotypes will asert, "You're kidding me". Nope, the real thing.. Bur oak Sandhill forest on a north slope in early summer. Most of the trees in these two "dendrographic" photographs were bur oak (including the largest tree and the pole-size sampling in left front of the "big 'un"), but white or American elm (Ulmus americana) and green ash (Fraxinus pennsylvanica) were associates that were present (as in background of these two slides). (Incidentially the commerative specific epitthet pennsylvanica means literally "Penn's woods" [ in honor of the Quaker founder, William Penn] ; " sylva from the Latin and Greek roots is in reference to "wood" or "forest"). This was an old-growth stand of bur oak with the climax herbaceous species still dominating the understorey. This was clearly virgin range vegetation. The graceful herbaceous understorey in these two "photo-plots" was almost exclusively made up of long-beak or Sprengel's sedge (Carex sprengelii). Canada wildrye (Elymus canadensis) was the associate herbaceous species overall though very little of this grass was present in forest range vegetation presented here. Eastern red cedar (Juniperus virginiana) had in absence of fire begun to encroach as an invader into this sylvan Eden. Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Juchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001).

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

192. Forest and sunflecks- Nebraska Sandhills bur oak forest and shade-adapted herbaceous species. Light conditions of early morning, the major part of day during which light could reach ground level of this north-slope forest, provided a textbook illustration of the sunfleck phenomenon. In dense-canopy plant communities like forest (and even grassland swards) sunlight penetrates through foliage at varying lengths of time. The shortest of these light periods or bouts (time lengths or durations of light at whatever light intensity) which can last only a few seconds up to several minutes are known as sunflecks (Smith et al., 1989; Chazdon and Pearcy, 1991). The converse of sunflecks are shadeflecks which are shade fluctuations due to cloud cover between shaded spots and the sun. In vegetation, such as that of a bur oak forest featured here, wind movement of leaves, travel of Earth relative to sun, and any cloud cover are responsible for the ever-changing state of light that penetrates to lower strata of the range plant community. The general condition or phenomenon of changing or dynamic light as to duration and related intensity or "fluctuation in irradiance" as it was described by Chazadon and Pearcy (1991, p. 760) is an all-important abiotic factor in growth and survival of plants in lower layers of range vegetation-- the understorey of this bur oak forest. Light dynamics interacts with other factors (abiotic and biotic) such as defoliation, precipitation, and, as in this north-slope forest, aspect. Dense shade produced by the large leaves and close interlocking crowns of bur oak on the aspect most shielded from sunlight resulted in severe conditions of light deficiency related stress for understorey plants in this bur oak forest. Range plants in lower layers of this north-slope bur oak-dominated forest community were sciophytes (sciophytes), "shade-loving" species of the highest adaptation. There were a few, though extremely patchy or irregular, layers of woody plants below the forest canopy. These vegetational strata were made up primarily of regenerating bur oak (seedlings, saplings, and poles), American elm, green ash, and, least of all, eastern red cedar. This photographer did not see any shrub species. The herbaceous layer (which was also the ground layer) of this forest range was dominated mostly by long-beak or Sprengel's sedge with Virginia wildrye the associate species. The major--and about the only--forb was anise root (Osmorhiza longistylis). This virgin range vegetation was a fine representation of old-growth bur oak forest (even if there were too many shadows to show it to best advantage). Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Juchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001).

193. A study in shadows- Closer-in photograph of the two big bur oaks and associated herbaceous layer introduced in the last slide (second slide of the preceding pair of photographs). This will be about all--and clearly the most (greatest intensity and longest duration of)--light that the understorey of this north slope bur oak forest will receive, and this was close to longest day of the year. Early morning and late evening were the major (about the only) times that sunlight could fall on the ground at this extreme north aspect. The phenomenon of sunflecks as explained in the preceding caption was obvious. Understorey was herbaceous, except for basal sprouts of bur oak (see below), and consisted primarily of long-beak sedge with some Canada wildrye. The only forb of note was anise root (Osmorhiza longistylis). This was virgin vegetation in structure and composition and in all layers; a good representation of climax bur oak forest range. Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Juchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001).

194. About as bright as it ever gets- The slant of earlier morning light on this north slope is just about as much photosynthetically active radiation as the understorey of this north-slope, Nebraska Sandhills bur oak forest ever gets. These paired photographs presented an example of light dynamics in the understorey layer of this densely shaded forest range. The second of these two photographs was at a closer distance to trees, but the major difference in pattern of light and shade on the forest floor was due to changing radiation that occurred with passage of time. These photo-dynamics resulted from 1) variation due to the diurnal (day-and-night) cycle of light and 2) annual cycle due to travel of Earth on its yearly orbit around the sun. Sunflecks lasting only a few minutes or, perhaps more commonly, mere seconds (or fractions of seconds) are largely due to wind movement of leaves in the forest canopy. This rapid photo-dynamics--very short-term spots of alternating sun and shade--could not be captured in photographs or sequences photographs, but the prevalence of light dynamics, hence the concept of sunflecks, was represented photographically. Trees and saplings in these slides were all bur oak. The herbaceous layer was mostly long-beak sedge with some Canada wildrye and, even less, anise root. Old-growth bur oak forest serving as a reminder of what a virgin forested range was like in a most unlikely grassland region. Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Juchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001).

195. Shade-adapted range plants- This two-slide set of a north-slope, Nebraska Sandhills bur oak forest featured the 1) herbaceous understorey dominated by long-beak sedge with Canada wildrye as associate species and anise root as principal (essentially sole) forb and 2) base of a mature bur oak with stump sprouts and a sapling. The sapling was at such distance (about a foot and a half) away from the trunk of the adult tree that it was apparently a separate tree derived from an acorn (most likely the adult bur oak it was closest to). Thus this bur oak sapling appeared to be a sexual offspring of the adult oak, a separate and unique genetic individual produced by an adult bur oak through fruit. It was possible that this sapling grew from an acorn produced by a neighboring bur oak and transported next to the big oak shown here by a combination of gravity, wind, and steep slope or by a food-storing squirrel. There was some probability that this sapling was actually a basal sprout, secondary shoot, that arose from the adult tree, but this seemed not to be the case. By contrast, the small stump sprouts to the right of the sapling were asexual offspring from the large bur oak. In this case the new oak shoots (stump sprouts or stump suckers which are secondary shoots) were clones (clonal progeny) of the parent tree. These were genetically identical to the big bur oak from which they arose. Stump sprouts were secondary shoots of the existing bur oak tree the same as new limbs, branches, and buds. Such clonal shoots are often called offshoots. This asexual reproduction is vegetative reproduction (synonyms), human horticultural forms of include grafting and budding. More (and more clear) examples of suckering from healthy, uninjured, mature bur oak was shown below near end of this section. Almost all tree species are capable of both sexual and asexual reproduction, but sexual regeneration is less successful in forest of dense shade unless the species are relative in tolerance (Tolerant or Very Tolerant). Vegetative regeneration is not limited (at least not nearly as much so) because the parent tree can translocate photosynthate and other nutrients to its own shoots (clonal organs). By contrast, sexually produced progeny are their "own plants" and must be able to survive under the shade of their parents at age classes ranging from seedling through sapling to pole-size. Most of these individual smaller trees--at least in species having lower tolerance--die. An example of this reality was the stick (at left of trunk and sapling) which was the remains of a small sapling tht could not survive in the shade of (compete with) the aduld bur oak. Burns and Honkala (1990) reported that bur oak has usually be interpreted as being Intermediate in tolerance rating although certain observers viewed bur oak as more tolerant than other climax oaks such as northern red and white oak. In the Prairie Peninsula Region however bur oak stands have been invaded by other oaks including white oak, and bitternut hickory (Burns and Honkala, 1990). This was far to the east and on more mesic habitats than in the Nebraska Sandhills where the more mesophytic oaks and hickories are absent. These same authors specified that in forests to the east and north bur oak died from suppression (due largely to shading?) after reaching sapling size. That same response was frequent in the north-slope Sandhills bur oak forest described here. Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Juchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001).

196. Dominants of a deep, dark lower layer- Local sunlit patch of the herbaceous layer in a north-slope bur oak forest in Nebraska Sandhills. Plant species seen here were the climax herbaceous species that dominates the understorey of a sandhills bur oak forest..Long-beak or Sprengel's caric sedge (Carex sprengelii) was the dominant--almost exclusive--species with Canada wildrye the associate species. Anise root was the only forb of consequence in the herbaceous layer. These were reported as characteristic (indicator) herbaceous species in bur-oak dominated range vegetation (Barker and Whitman, 1989, ps.17-18). The first of these two photographs was of a single-species stand of long-beak sedge. Second photograph included a shoot of Canada wildrye accompanying the long-beak sedge. Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Juchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001).

197. Contaminated understorey- Sward of herbaceous layer of a north-slope bur oak forest in Nebraska Sandhills consisting of long-beak or Sprengel's sedge and of naturalized Kentucky bluegrass (Poa pratensis). Kentucky bluegrass and smooth bromegrass (Bromus inermis) are two widespread Eurasian, perennial festucoid grasses introduced by the white man that have natrualized widely in North America. These two introduced grasses are commonly used throughout the greater Northern Great Plains Region (as well as areas of adjoining regions) for grazing and, in case of smooth brome, for both pasture and hay. They continue to have ample opportunities for naturalization and establishment across a vast acreage, especially on disturbed sites in humid through semiarid zones, so they are properly regarded as naturalized range plants.. Continued heavy mowing up to edge of the north-slope, old-growth bur oak forest described heein enabled both Kentucky bluegrass (as shown here) and smooth brome (covered below) to invade outer parts of the native herbaceous understorey of this bur oak stand. This invasion allowed displacment, to varying extents, of native graminoid species like long-beak sedge and Canada wildrye. These two agronomic, Eurasian grasses were able to outcompete natives only where there was continued disturbance by repeated, heavy mowing over specific locations or of adjacent areas from which the exotics could spread for short distances into unmowed native sward. The sward shown here was an example of the latter situation. Cherry County, Nebraska. Late June, peak flowering stage of both long-beak sedge and Kentucky bluegrass.

198. Long-beak or Sprengel's caric sedge (Carex sprengelii)- Three photographs showing progressively greater detail of sexual shoots on the dominant herbaceous species in the understorey of a north-slope Nebraska Sandhills bur oak forest. Cherry County, Nebraska. Late June, peak flowering stage just after anthesis.

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

200. Invasion at the edge- Lower and outer edge of a north-slope bur oak forest in Nebraska Sandhills with an individual eastern red cedar prominently and happily growing in a mid-layer of vegetation. This immediate local was just above Minnechaduza Creek. This accounted for the lush stand of herbaceous riparian vegetation in the foreground. which "crept in" into this "photo-plot" useed to show size of the cedar relative to typical size of adult bur oak (to immediate right of eastern red cedar). Human suppression of fire had permitted eastern red cedar to establish sporadically both along edge and in interior of the bur oak-dominated forest. Most of the herbaceous riparian vegetation was the introduced Eurasian grass, smooth brome which invaded the bur oak understorey under frequent, heavy mowing. Also present with much cover and density, though lacking height of the bromegrass,was another Eurasian grass, Kentucky bluegrass. Other riparian species were native grasslike plants (Cyperaceae) that were still only in vegetative stages and unidentifiable by this worker. These had, however, been overwhelmed by the introduced grasses. The only native plants that could persist under cover of the agronomic grasses (which were competitive with native herbaceous species only under heavy mowing) were bur oak, green ash, American elm, and eastern red cedar. And mowing wiped out all plants of these tree species except saplings and adult trees. Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Juchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001).

201. Bur oak stand at the edge and changed by man- Part of the perimeter of the north-slope Sandhills bur oak forest adjacent to a field of smooth bromegrass. Extension of close (short-height) mowing, over a number of years, into the edge of the bur oak forest had either directly killed out long-beak sedge, Canada wildrye, and anise root and/or permitted invasion of smooth brome and Kentucky bluegrass that "choked out" (outcompeted, overshadded, etc.) these natives. Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Juchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001).

202. Advance guard of bur oak-dominated vegetation being invaded by enemy legions- Two views of a bur oak forest with herbaceous layer(s) that were highly man-modified. This local small stand of bur oak stood in stark contrast to the virgin climax bur oak forest featured above. These two photographs showed the outer- and lowermost perimeter of the bur oak community covered in this section. The adult bur oaks and a few patches of the natural understorey beneath mature trees had persisted (where oak trunks were too close together to permit entry and operation of rotary mowers [=shreaders]) along this perimenter. Generally, however, continued close mowing of the range vegetation had done two things: 1) prevented regeneration of bur oak by cutting off all bur oak seedlings and 2) largely converted the natural herbaceous understorey to an anthropogenic or manmade (=artificial) layer of smooth brome with local stands of Kentucky bluegrass. Change in species composition of the herbaceous layer(s), replacement of native species with naturalized Eurasian species, was due to either direct killing of long-beak sedge, Canada wildrye, and anise root by defoliation or indirectly by reducing competitiveness of these range species and/or favoring the taller-growing smooth bromegrass along with denser-growing, more cutting-toleraant Kentucky bluegrass. It was explained above that both of these introduced, agronomic forage grasses have naturalized throughout much of the North America range country. Where mowing disturbance (= abnormal defoliation regimen) occurred throughout much of the growing season the understorey was converted from an herbaceous layer of native grasslike, grass, and forb species to one of naturalized agronomic grasses (ie. conversion from native to naturalized range plants). Also, the lower woody layer(s) was totally eliminated because mowing prevented regeneration of all tree species. Beyond doubt or debate, continued close mowing as had been practiced on this forest range for years will eventually eliminate the bur oak forest (as soon as the already old trees die off) and convert the plant community into a manmade grassland of smooth brome and Kentucky bluegrass, a farm field of domestic forages. Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Juchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001).

203. Artificial understorey- Continued close mowing by roatary shredders killed out--directly or indirectly--the climax herbaceous species (long-beak sedge, Canada wildrye, anise root) of a bur oak forest understorey and converted the sward to a single-species layer of smooth brome.A small remnant of smooth bromegrass remained uncut where this beautiful trunk of an old-growth specimen of bur oak prevented mowing (an inexperienced or lazy shredder operator did not get close enough to the oak). The other glaring outcome of mowing was absence of regeneration of bur oak, dominant tree species, and green ash, the associate species. If present mowing practices continue (and they obviously had been in effect for a number of years) there will eventually be no bur oak forest left. Instead, the plant community will be a monoculture of smooth brome (with local spots of Kentucky bluegrass). All native vegetation will be wiped out (ie. the forest range will be no more and a hay field will have replaced it). Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Juchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001).

204. Genuine-article understorey- Two trunks of large bur oak prevented close mowing of the native herbaceous understorey comprised of long-beak or Sprengel's caric sedge, the dominant, and Canada wildrye, the associate species. Survival of these climax species permitted some of the composition and structure of a bur oak forest range to persist. This part of the old-growth bur oak forest that had developed on a north slope in the Nebraska Sandhills was mowed with less frequency than some of the other forest understorey with the result that the climax species composition of the sward (herbaceaous layer) persisted. Tree regeneration (bur oak, green ash, American elm) was still completely prevented as any seedling was mowed off where density of adult trees and saplings did not prevent entry of rotary mowers. Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Juchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001).

205. Nebraska Sandhills bur oak grove- A bur oak grove, the outermost edge of which was shown here, had developed on a sunlit ridge conterminous with the north-slope bur oak forest described immediately above. Both phases--open, fullsun grove and dense, deeply shaded forest--were climax range vegetation and represented the extremes physiographic and structural manisfestations of the bur oak forest cover type. The greatest differences were in the understorey vegetation, as to vegetational structure and species composition of herbaceous and woody (shrub and smalltree) layers. The bur oak grove phase (or community, if the two extremes were to be viewed as two different range plant communities) had no species of woody plants except bur oak (in contrast to American elm, green ash, and eastern red cedar in the north-slope forest). Also absent from the bur oak grove on the drier and more sunlit ridge was long-beak cric sedge which made up most of the herbaceous biomass on the north-slope forest. Instead, Canada wildrye, associate on the north-slope forest, was the overall dominant of the herbaceous layer in the bur oak grove. Giant ragweed (Ambrosia trifida) was a locally dominant herbaceous species and the most abundant forb in the oak grove. Anise root, the only forb, on the north-slope forest was absent in the grove vegetation. This bur oak grove was on the border of a field of the Eurasian perennial grass, smooth brome, which, as shown above, had invaded the understorey of a north-slope bur oak forest under repeaded close, mechanical mowing when the native long-beak sedge, Canada wildrye, and anise root were killed out by such extreme (intense and frequent) defoliation. Where trunks of bur oak were too close together to permit intrusion of this bloody equipment and overmowing the native, cool-season Canaada wildrye dominated the understorey other than in microsites where giant ragweed and bur oak stump sprouts held this honor. Physiogonomy and structure of this Sandhills bur oak grove was presented in this two-slide episode. Details of the herbaceous layer of this grove was presented in the next two-slide episode. Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Juchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001).

206. Edge of a bur oak grove- Lower zone of tree trunks and herbaceous understorey of a Nebraska Sandhills bur oak grove. This stand of climax vegetation was conterminous with the north-slope bur oak forest featured earlier. In contrast to the deeply shaded understorey of the bur oak forest in which long-beak caric sedge was dominant and Canada wildrye was the associate species, the herbaceous layer of the grove phase of the bur oak cover type was dominated by Canada wildrye except in small areas (microhabitats) where giant ragweed held supremancy or where basal sprouts of bur oak overtopped the dominant cool-season, festucoide grass. Canada wildrye was featured in both of these photographs with the local sward in left foreground of first slide shown at close range in the second slide. Some of the dead (light brown or buckskin-colored) shoots of Canada wildrye included some of the previous year's growth (slightly more faded) as well as some of those of the current growing season. Almost all of the current year's shoots of Canada wildrye were still in the boot. It was not known why a few of the current growing season's shoots had already matured, died and gone into dormancy. Did make for a nice contrasting picture with Canada wildrye very conspicuous. Giant ragweed, a warm-season annual composite of mature size that lives up to its common name, was still in early growth and thus not shown in proportion to its yield of biomass at peak standing crop. This herbaceous understorey was the vernal society even though it was summer and bur oak were representative of the estival aspect. Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Juchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001).

207. Parents, progeny, and playmates- Sprouting (suckering), the production of secondary shoots, from base of trunks in non-injured adult bur oak trees. Bur oak in both of these "photo-plots" were on a ridge at edge of a bur oak grove where light could penetrate from all directions throughout most of the day. This was a different condition of radiation than that of the north-slope bur oak forest described at beginning of this section. Differences in understories of the bur oak grove considered here and the bur oak forest described previously were explained in the immediately preceding caption. It was strikingly evident that full-grown (adult) bur oak produced basal sprouts in both deep shade and full sunlight (forest and grove stands, respectively). This was an understandable characteristic when it was born in mind that these secondary shoots were being supported by the parent shoot (ie. sprouts were a photosynthate sink from the parent tree that was the a nutrient (food) source for the basal suckers. (Sprouting frrom stumps and basal trunks of bur oak was considered in more detail in the next succeeding set of photographs.) What was substantially different in understories of bur oak forest versus grove was in herbaceous species and tree species other than bur oak (see again preceding caption). Mature sexual shoots of Canada wildrye, the overall dominant herbaceous species in the bur oak grove, with conspicuous spikes were present in the first of these two p;hotographs. Also prominent in this first slide was the pioneering, annual composite, giant ragweed. The second slide presented a fullsun shot of a basal sprout of bur oak at edge of the grove where frequent mowing had killed out the native Canada wildrye and its replacement with the introduced Eurasian Kentucky bluegrass. Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Juchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001).

208. Living stumps- Two stumps of bur oak with prolific and vigorous sprouting or suckering. These trees were at edge of a bur oak-dominated forest in the Nebraska Sandhills. Stump sprouting is an adaptation to fire, a key abiotic factor of grasslands. Bur oak is one of the most fire-adapted Quercus species in North America with sprouting from stumps or injured tree trunks one of its most important means of regeneration. In fact, bur oak sprouts prolifically even from old trees and those not subjected to injury. This was shown in several preceding photographs for bur oak growing in deep shade and fullsun environments. Bur oak has long been recognized for its "thick fire-resistant bark" (Burns and Honkala, 1990), but prolific sprouting (suckering) is an accompanying adaptation to recurrent fire that was evolved in the generally fire-prone habitat of this species biological range. Burns and Honkala (1990) explained that prolific sprouting was common following burning or cutting of bur oak only up to pole-size with such secondary shoots being of poor quality except those of seedlings. This latter conclusion was confusing to say the least given that shoots of seedlings are not sprouts (secondary shoots) at all but rather primary shoots derived from acorns. These same authors noted that larger trees (mature or full-grown age class?) also produced basal sprouts but that vigor and quality of these sprouts had not been evaluated relative to age, size, etc. of parent trees. The two examples shown here were pictorial evidence that seemed to contradict findings reported in Burns and Honkala (1990). The profuse (and plenty vigorous) stump sprouts shown off here were from mid-size trees (a foot or more in basal diameter). This was considerably smaller than the full-grown bur oaks presented in the immediately preceding set of two photographs, and these appeared vigorous enough too. Cherry County, Nebraska. Late June.

209. Fire ran through the bur oaks- At the outer edge of a grove form of a bur oak forest along a small stream through a hot fire with high flames had burnt just two to three prior to time of photograph. This forest range was in the Dissected Till Plains of the Central Lowlands physiographic province of northeast Kansas which is about the southern limit of forest dominated by bur oak. Thee two slides presented an outside-the-forest view to show physiogonomy and structure of the plant community. The lush growth of the herbaceous layer was dominated by seedlings of giant ragweed (Ambrosia trifida) with scattered plants of Canada wildrye, Canada wood nettle (Laportea canadensis), and pokeberry or pokeweed (Phytolacca americana). Giant ragweed is a pioneering annual; in fact, it is the most abundant pioneer or colonizing forb in this area. The hot fire had been a disturbance of such severity as to create an ideal habitat for invasion by giant ragweed which "eclipsed" the other herbaceous species which were perennials. These perennials had been present years before the burn, but they, too, appeared to have benefitted from the fire jucging by growth of other plants of these species in a few isolated spots that did not burn. This phenomenon was explained in captions below. The saplings in foreground of these (and subsequent) photographs were all hackberry and American elm. Almost every one of these saplings (and seedlings that were too small to show up in the slides) were top-killed by the heat of the obviously hot fire which burned with intensity adequate to partially kill lower branches of bur oak. This latter feature of the fire was shown and described in greater detail below. In this region most climax bottomland forests are dominated by various combinations of hackberry, American elm, slippery elm, and green ash (the Tolerant climax species) with old trees of such Intolerant, colonizing species as eastern cottonwood, sycamore, black walnut, and honey locust persisting into climax as associates. This is Society of American Foresters (Eyre, 1980) cover types 93 and 94. These mixed hardwood forests are the climatic climax. By contrast, bur oak forests, Society of American Foresters (Eyre, 1980) cover type 42, are a pyric climax. Differences in alluvial soils of these floodplain forest are not so different as to account for differences in forest range types. In the absence of periodic fire the bur oak forest would eventually proceed through plant succession to a hackberry-American elm climatic climax. That is, it would be the potential natural vegetation along streams in this climatic regime as interpreted from perspective of polyclimax and climax pattern theories. Or, what amounts to the same thing, bottomland forests dominated by hackberry and American elm would be the edaphic climax, the natural termination of such stream bottom seres. Either way, it is only periodic hot fires that maintain bur oak, which is Intermediate in tolerance (Burns and Honkala, 1990), as climax along stream bottoms (ie. bur oak is a fire type, a pyric climax). In monoclimax theory all such forests are postclimax to the climatic (= zonal or regional) climax of tallgrass prairie. Individuals of bur oak would undoubted persist into a hackberry-American elm climax as bur oaks are relatively long-lived. Burns and Honkala, (1990) reported that some bur oaks bear fruit up to ages of 400 years, the longest of any Quercus species in North America. Progression through plant succession from a seral bur oak-dominated forest to a hackberry or hackberry-elm climax forest was the pattern of vegetation development predicted for gallery or floodplain forests in northeast Kansas by Bellah and Hulbert (1974) and Abrams (1986). Abrams (1985) concluded that fire frequency (mean fire interval) bur oak-chinquapin oak gallery forests in the Flint Hills ranged from about eleven to twenty years with an average "somewhere between that range" but with a historic interval of two to three years. Clearly the historic frequency of two to three years would have maintained the oak forests and prevented progression to hackberry or hackberry-elm. It is likely that even an average fire return interval of roughly fifteen years in contemporary time would maintain the bur oak-chinquapin oak forest. Again, bur oak forest (SAF 42) is a fire type or pyric climax. Spring Creek, Marshall County, Kansas. Mid- June, vernal aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Kuchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Western Corn Belt Plains, Loess and Glacial Drift Plains Ecoregion 47i (Chapman et al., 2001).

210. Late spring after fire-Apperaance of a bur oak grove two to three months following a hot fire that effectively top-killed seedlings and saplings of hackberry and American elm that had established beneath the bur oak canopy. Fire intensity was adequate to create a disturbance that favored rapid establishment of a dense population of giant ragweed, the most common and prolific pioneering plant in this region of the Dissected Till Plains of northeast Kansas. Rapidly growing seedlings of giant ragweed, an annual composite, accounted for the greatest herbaceous cover, but there was also an abundance of Canada wildrye and pokeweed (though widely scattered with most cover being local) along with local exclusive colonies of Canada wood nettle (see photograph and caption below). At outer margin (just under forest canopy) there were large individuals of what appeared to be--without much familarity of early growth shoots--giant sumpweed (Iva xanthifolia). The first of these two photographs was a closer-in view of the bur oak at center-left in the second photograph of the preceding two-slide set. The trunk of this tree had split almost to ground level yet was still standing and healthy. Heartwood of a tree is dead tissue anyway so as long as the tree was not weakened to point of crashing and did not have heart-rot it remained as healthy is if the trunk was intact. The second of the photographs of this caption presented a summary view showing several top-killed saplings of hackberry and American elm. The rightmost bur oak (center-right) sported an old fire scar and ws in every way healthy from all outward appearances. This fire-scarred tree was featured below. Spring Creek, Marshall County, Kansas. Mid- June, vernal aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Kuchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Western Corn Belt Plains, Loess and Glacial Drift Plains Ecoregion 47i (Chapman et al., 2001).

211. Old and the new after a fire- Close-growing and very old bur oaks (first slide) along with seedlings of giant ragweed and this season's shoots of Canada wood neetle, pokeweed, and Canada wildrye as an herbaceous layer plus some saplings of hackberry and American elm (see especially second slide) comprised the range vegetation of a bur oak forest two to three months following a hot surface fire. In the background there were several pole-sized trees and saplings of red mulberry (Morus rubra) that formed an erratic lower tree layer. One of these saplings was shown in center midground of the second slide. The sapling in center foreground of this second photograph was one of the few young hackberries that was not top-killed by the fire. Other saplings of hackberry and American elm that were not topkilled outright had already started to "peel bark" and appearedlikely to die. The hackberry sapling featured here defied the odds and appeared ready--at this juncture anyway--to take its place ultimately as an adult in the grove of bur oak. It was explained in the immediately preceding caption that bur oak is a fire type forest. In absence of periodic fire bur oak--other than as a few persistent old-age trees--would be replaced by hackberry and American elm which would be the climax dominants on this bottomland sere. The latter two are Tolerant species whereas bur oak is Intermediate in tolerance rank (Burns and Honkala, 1990). Of course, fire is as much as a part of climate (or a consequence of climate if one prefers) as drought, wind storms, and floods. More precisely, lightening is as much part of the atmosphee as precipitation and temperature and fires from lightening with fuel made possible by climate are major factors responsible for maintenance of bur oak forests within the surrounding zonal vegetation of tallgrass parairie. Various studies such as those by Kucera (1960) and Bragg and Hulbert (1976) showed that woody plant communities, including forests, expanded in the tallgrass prairie region since European settlement (due in large part to fire suppression). Abrams (1985) estimated that the mean fire interval for bur oak-chinquapin dominated gallery forest in the Flint Hills of Kansas was two to three years and roughly eleven to twenty years with intervention by white settlers and contemporary man. Abrams (1986) documented this phenomenon and predicted that hackberry, elm, and redbud (Cercis canadensis) would gradually replace bur oak and chinquapin oak--at least in absence or reduced frequency of fire--as the climax forests for bottomland forest sites in the Flint Hills and Dissected Till Plains in northeast Kansas. Abrams (1992) issued a later report based on his earlier findings and concluded that for some areas of Kansas the presettlement vegetation of tallgrass prairie had become either a bur oak-hackberry overstorey with a hackberry-elm understorey or a chinquapin oak overstorey with a elm-redbud-chinquapin oak understorey. This analysis struck the current author as an interim report because it seemed likely to him that the ultimate state would be a climax hackberry-elm forest. Spring Creek, Marshall County, Kansas. Mid- June, vernal aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Kuchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Western Corn Belt Plains, Loess and Glacial Drift Plains Ecoregion 47i (Chapman et al., 2001).

212. Just another encounter with fire- Outer edge of a bur oak grove two to three months following a surface fire that burnt through this bottomland forest. Judging from fire scars and general understanding of bur oak forest and the natural envioroment of this area it was obvious that this was the most recent of many such fires in these old bur oaks. The fire had been hot enough to create a disturbance so severe that giant ragweed, the most abundant pioneer species in this area, had already established populations that pretty much excluded other herbaceous species except the climax perennials, Canada wildrye and Canada woodnettle (wood nettle). Even these shade-tolerant and already present perennial herbs appeared to have benefitted greatly form the fire. Without this fire a dense layer of bur oak leaves would functioned as mulch and largely smothered most herbaceous plants. Such a layer of large bur oak leaves (= mulch) was present in several small spots that did not burn. Giant ragweed was absent from these and Canada wildrye plants were smaller than those growing on land that had burned. Note that the lowermost branches of bur oak were largely leafless, but with enough leaves to prove that they were still alive (barely) and that heat from the last flames injured (appeared to have more-or-less killed) these smaller, lower limbs. This was an example of fire-pruning, but such defoliation did not necessarily kill these organs. It was likely that new shoots would arise from intercalary meristem on these lower branches. The sapling to right of foremost bur oak was hackberry. It leafed-out following the blaze, but was nonetheless severely injured by the heat from had to be an unusually hot fire. This one of the few hackberry saplings that was not totally top-killed and it had entire strips the total height of the shoot that already had peeling bark. All American elm sapling in the understorey were top-killed. Abrams (1985) studied fire frequency within gallery forests of northeast Kansas (the Fllint Hills) and found a natural fire frequency of two to three years and an extended mean fire interval of roughly fifteen years during time of occupation by European man. Either the natural or anthropogenic fire frequency would probably maintain bur oak groves and prevent development of hackberry-American elm forest on the forest range site featured here. The fire scar on the foremost bur oak was an old wound that was already present and in process of healing before the recent blaze. This "honor scar" was presented and described further in the next slide. Spring Creek, Marshall County, Kansas. Mid- June, vernal aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Kuchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Western Corn Belt Plains, Loess and Glacial Drift Plains Ecoregion 47i (Chapman et al., 2001).

213. New beauty in an scar- An old fire scar on an ancient bur oak that was renewed by a surface fire only two to three months prior to time of photograph. The tree had started started healing the wound caused from previous fire(s) and the most recent burn did not harm the new bark tht was slosly growing over the scar. This was a direct view of the scar that was shown from the side in the foremost bur oak featured in the immediately preceding two slides. Forbs growing in front of the oak were young seedlings of giant ragweed, the herbaceous species that most benefitted from the recent fire. This is the "price" ("rent" so to speak) to be paid by the fire-adapted bur oak for living in a pyric habitat in which less fire-adapted and more tolerant hardwoods such as hackberry and elms cannot survive. In absence of periodic fire hackberry, green ash, American elm, and perhaps boxelder would eventually replace bur oak through plant succession. These more tolerant species are the climatic climax whereas bur oak is a pyric climax (ie. bur oak forest is, in effect, a fire type in the overall environment of the Dissected Till Plains or, sometimes, Glacial Drift Plains of northeastern Kansas). Spring Creek, Marshall County, Kansas. Mid- June.

214. After the latest in a long history- Inside a bur oak grove through which a hot surface fire burned two or three months before. This bottomland forest range had developed along a small stream at edge of a tallgrass prairie in the Dissected Drift Plains of northeastern Kansas. There were several "featured attractions" in these two photogrphs. The most prominent of these was the large fire scar on one bur oak trunk that extended from the ground to over 15 feet in height. this was an old scar from previous fire(s) that was "freshened up" by the most recent burn. Another, though much smaller, fire scar on another bur oak tunk was presented and discussed in the immediately preceding photograph and caption. The current author interpreted bur oak forests in the Dissected Till Plains as a fire-climax type with bottomland forests on this forest site in absence of recurrent fire developing through plant succession to a climax hackberry-American elm forest. Effects from fire wounds and scars on bur oak are probably not know with any certainity. Entry of disease pathogens (eg. various forms of fungal rot) and insects through tree wounds, including those caused by fire, is always a possibility. One common suspicion--though perhaps without scientific proof--is that fire (heat or charring of wood) has something of a cauterizing effect so that disease entry is less likely with fire wounds/scars than with similar wounds (eg. those ranging from ax blazes to debarking by falling trees). The charred (charcoal) surface of a fire wound, even a fresh one) is a different (drier, perhaps chemically) environment than the moist, peeled-surface wound left by mechanical injury. What is certain from even brief scanning of the literature is that bur oak fire scars as records of fire regimes, especially fire frequency, have been analyzed in considerably more detail than the impacts of fire damage on tree health. It is also certain that fire in bur oak-dominated and influenced communities (forests, open groves, savannahs, even prairies) is a natural part of the habitat of this range vegetation. Furthermore, it follows that bur oak is adapted to fire and, was the species not so adapted to and tolerant of fire to the degree it is, this species would not be a member of these range plant communities. It is axiomatic that the overall impact of fire on bur oak is minimally adversive, neutral at least, and almost assuredly positive for survival in range vegetation of which it is an important member. Peterson and Reich (2001) concluded that on bur oak was a fire-resister, a designation given by Rowe (1983) to shade-intolerant tree species that suffer little or no damage from low-severity fres. Peterson and Reich (2001) explained that "[f]ire rarely killed mature bur oaks, even those in the smaller size classes ...". They (Peterson and Reich, 2001) noted further that even saplings of bur oak grew corky bark of such thickness as to protect the cambium from most fires. Burns and Honkala (1990) cited a volume of literature attesting to the thick fire-resistant bark of bur oak, which along with its general drought-tolerance, accounted for presnece of bur oak on xeric sites as well as mesic ones where bur oak was an associate on sugar maple-American beech forests. Another "featured attraction" in this pair of photographs, especially the second one, was the death (or near death) of shoots of hackberry and American elm saplings standing in stark contrast to the survival of adult trees of bur oak even with fire scars where sizable portions of their trunks were removed. This is a photographic lesson showing that it is primarily periodic fire (with browsing obviously less important) that maintains bur oak groves and forests which would otherwise develop into hackberry-elm forest with only persistent and senescing adult bur oaks (at status of associate species at most). Bur oak forests and woodland as pyric climax (a climax fire type) was discussed in above photo captions (complete with citation of relevant literature) so that further discussion was not deemed necessary or desirable in this current caption. This was an opportune point to acknowledge that in more xeric environments (eg. those to the west the Dissected Till Plains such as the Smoky Hills or Sandhills in subhumid to semiarid precipitation zones) it is probable that limited soil moisture, especially in drought, rather than fire is the primary variable responsible for maintenance of bur oak and restriction of Tolerant tree species like hackberry, elm, box elder, etc. Also featured here was presence of red mulberry (lower branch with large leaves extending downward from upper-left corner of first photograph). Red mulbery was usually observed to be a smaller tree of the second woody (lower tree) layer, but as one of the--if not the single most--consistent tree species in bottomland forests in humid and subhumid zones extending from the Cross Timbers and Central Prairies in Texas, Ozark Mountains in Missouri and Oklahoma, and through to the Great Plains in Nebraska. In southern mixed hardwood forests, such as those in northcentral Texas, red mulberry frequently grew to relatively large size (eg. 16-18 inch DBH) with straight boles. Red mulberry was found as a consistent member of bottomland forests were dominance varied from that by eastern cottonwood, sycamore, pecan, bur oak, elm, green ash, and hackberry or sugarberry and where soil texture ranged from primarily sandy to predominantly clay. In progressing northward, dominance by members of Ulmaceae goes from exclusively sugarberry (Celtis laevigata) as in north Texas to side-by-side co-dominance of sugarberry and hackberry (C. occidentalis) in northern Oklahoma to exclusively hackberry in northern Kansas and Nebraska. Red mulberry grew with an array of dominant tree species. Again this was usually, though not always, as an understorey or lower-height tree species. Species in the herbaceous layer shown in these two photographs included giant ragweed, Canad wood nettle, Canada wildrye, pokeweed, and giant sumpweed in that order based on estimated relative cover. Giant ragweed, an annual composite, was the number one pioneer species of denuded land in this area and far-and-away the dominant of the herbaceous layer except where there were local colonies of Canada woodnettle farther in the interior of this bur oak forest. Canada wildrye, a festucoid grass of the barley or wheat tribe (Hordeae or Tritaceae) that responds to disturbance as a decreasere, was able to "hold its own" against the rapid-growing and rank seedlings of giant ragweed. Giant sumpweed was present as sparse though very conspicuous individuals. Spring Creek, Marshall County, Kansas. Mid- June, vernal aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Kuchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Western Corn Belt Plains, Loess and Glacial Drift Plains Ecoregion 47i (Chapman et al., 2001).

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

Sugar Maple-Dominated Forests

Sugar maple-dominated forest in the Ozark Plateau constitute a more mesic phase(s) of the Oak-Hickory Forest Region (Braun, 168-170). Specifically, sugar maple-dominated forests in the Oak-Hickory Forest Region were maple climaxes (climax sugar maple-dominated forests) the same as (at least, equivalent to) the maple climaxes that define the Maple-Basswood Forest Region of Braun (1950, ps. 327-336). The quote from Braun (1950, p.34) cited in the introduction to this chapter bore reciting: "Each of the several climaxes, although characterizing a specific region, nevertheless occurs in other regions.”. Braun (1950,p. 164) also stated: "The forest of the most mesophytic slopes usually contain sugar maple and an admixture of other mesophytic species". In other words, sugar maple is a defining, dominant species of some upland, slope forest in the oak-hickory forests. Thus there were the climax sugar maple-dominated forests in the Ozark Plateau that were presented in this portion of the Oak-Hickory Forest chapter. Three forest tracts dominated by sugar maple were described in this portion of Oak-Hickory Forests-II. These separate tracats of forest range were similar and comparable to published descriptions of sugar maple forests in the Ozark Highlands (Ozark Plateau). Two forms, phases, variants, (or whatever they would be designated) of climax sugar maple were recognized for the Ozark Mountains by forest ecologists. These were the Acer saccharum-Quercus alba associes of limestone slopes and the Acer saccaharum-Carya cordiformis associes" that Braun (1950, ps. 168-169) recognized from the preceding work of Steyermark (1940). Both Steyermark (1940) and Braun (1950) used the Clementsian monoclimax vegetation system (Clements, 1916) in which associes "is the developmental equivalent of the association" so "used where the community is not permanent" but seral (Weaver and Clements, 1939, p. 99). In the geologic time scale of monoclimax theory the limestone bluffs and hills of the Ozarks would be worn down to a peneplane so as to eventually become the regional climax (monoclimax) of oak-hickory forest. Forests of sugar maple (with co-dominant tree species) on more favorable, mesic sites (eg. north and east slopes) are in monoclimax theory, postclimax in oak-hickory regional (climax) forests. In the polyclimax theory of Tansley or climax pattern theory of Whittaker the Clementsian associes of Steyermark (1940) would be associations. The Acer saccharum-Quercus alba associes and Acer saccaharum-Carya cordiformis associes would be interpreted as variants of the Society of American Foresters (Eyre, 1980, p. 30) forest cover type, Sugar Maple (SAF 27). The forest cover type description of SAF 27 in Eyre (1980, p. 30) included four subtypes, two of which corresponded to the two forms of the moist slope Ozark Highlands forest: the sugar maple-bitternut hickory "restricted to deep soils in the southernmost part of Quebec" and the sugar maple-basswood-white ash subtype "found in the lowlands of the St. Lawrence Valley." Obviously authors of the sugar maple cover type description were unaware that the same subtypes (the associes of Steyermark [1940]) also existed in the Ozark Mountains of Missouri. This sugar maple-dominated forest range type three variants or forms) was described as mesic-limestone forest by the system of classification and designation in Nelson (1987, p. 28; 2005, ps. 122-125). Two of these variants of the sugar maple cover type had developed on limestone bluffs along (above) Modoc Creek and one was on a steep north slope (limestone parent material though not near a stream) at the western boundary of the Springfield Plateau of the Ozark Highlands. These were presented and described below as sugar maple-bitternut hickory (north slope), sugar maple-white ash- northern red oak (east slope), and sugar maple-northern red oak (north slope).

216. Postclimax of the Oak-Hickory Forest Region- A mesic, north slope of limestone bluffs above Modoc Creek supported a sugar maple-bitternut hickory (Acer saccaharum-Carya cordiformis) forest that was--when viewed from the Clementsian perspective--postclimax for oak-hickory (hardwood, in general) forests of the Ozark Highlands. On this limestone bluff forest sugar maple was the tree species with most regeneration, but this was followed closely by bitternut hickory. Associated tree species were (in this general order) white ash (Fraxinus americana), basswood or American linden (Tilia americana), hackberry or, sometimes, western hackberry (Celtis occidentalis), northern red oak (Quercus rubra= Q. borealis var. rubra), chinquapin (=chinkapin) oak (Q. muehlenbergii), black walnut (Juglans nigra), slippery or red elm (Ulmus rubra), honey locust (Gleditsia triacanthos), Kentucky coffeetree (Gymnocladus dioicus), and sycamore (Platanus occidentalis) . There were two shrub zones: 1) upper (higher on bluffs; farther from creek) and 2) lower (farther down on bluffs; next to creek). The higher or upper zone had one major shrub layer that was dominated by eastern dogwood (Cornus florida) with eastern redbud (Cercis canadensis) as the associate shrub. The lower zone on bluffs had two major or prominent shrub layers or strata: 1) taller shrub stratum made up almost exclusively of pawpaw (Asimina triloba) and 2) lower (although still relatively high) shrub stratum composed solely of American bladdernut (Staphylea trifolia). There was some buckbrush or coralberry (Symphoricarpos orbiculatus), but this otherwise widespread Ozark Plateau shrub was present only in sub-trace quantities in this mesic, relatively cool, north slope forest. Locally common to dominant forbs included Jack-in-the-pulpit (Arisaema atrorubens), green dargon (A. dracontium), wake robin (Trillium sessile), trout lily (Erythronium americanum), columbine (Aquiegia canadensis), dutchman's breeches (Dicentra cucullaria), bloodroot (Sanguinaria canadensis), and false Solomon's seal or false spikenard (Smilaceina racemosa). Rue anemone (Aneomella thalictroides) and false rue anemone (Isopyrum biternatum) were the earliest vernal forbs in this forest. Mayapple (Podophyllum peltatum) was present at extremely low cover (it was "barely there") which was "sign-significant" for this climax forest vegetation given the nearly ubiguitous presence of this forb in forest, even savannahs, of the Ozark Plateau. Other forbs were mosses and ferns, including walking fern (Camptosorus rhizophyllus= Asplenium rhizophyllum) and Christmas fern (Polystichum acrostichoides). Ferns were not as abundant (general density and cover parameters) as on an east slope sugar maple forest farther downstream (see below). Dominant grass overall was silky wldrye (Elymus villous) though locally broadleaf woodoats (Uniola latifolia) was common to dominant. Also local stands of Virginia wildrye. Naturalized Kentucky bluegrass (Poa pratensis) was also common at local scale. Higher up on bluffs the dominant climax grasses were Canada brome or hairy, wood brome (Bromus purgans= B. pubescens ) and woodland bluegrass (Poa sylvestris). Two photographs provided a longer glance (first slide) followed by a shorter glance (second slide) of the range vegetation of this mesic-limestone forest (Neldon, 1987, p. 28; 2005, ps. 122-125). Snag (right side near margin in first slide; right of center in second slide) was sugar maple. Large tree to its left with trunk injury or damage blaze (both slides) was white ash. Sapling to left of blazed big white ash with missing crown and profuse generation of long shoots was white ash. Sprouts in front and slightly to left of blazed big white ash were American elm with a few individuals of slippery elm. Shrubs were pawpaw (taller; more sparse) and American bladdernut (forming colonies; shorter that pawpaw). Smaller trees (not shrubs) at edge of dense forest stand with conspicuous leaves (partial crown in first slide; only one or two limbs in second slide) was sugar maple. Along Modoc Creek, Ottawa County, Oklahoma. Late May (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

217. Another view the next year- Another vantage point provided this farther-off exterior view of the same mesic-limestone forest introduced in the preceding two-slide sequence. This sugar maple-bittrnut (pignut) hickory-basswood-white ash-hackberry-northern red oak-chinquapin oakhad developed on a north-slope of limestone bluffs just above Modoc Creek, a smaller though typical Ozarks Plateau stream. This photograph was taken one year after the two in the immediately preceding set.. The large, tall tree in left foreground was the same white ash with blazed scar on upper trunk. One year later the crown of the sugar maple snag (tree just to right of white ash) had lost another major limb. Most of the maller trees (not shrubs) at perimeter of dense forest to left of big white ash were sugar maple. Shrub layers as seen in foreground (as under fallen sugar maple limb) included American bladdernut, pawpaw, and spicebush (in that order of cover and general importance or dominance). Saplings to right and slightly behind (higher on the bluffs) were basswood (larger sapling) and American and slippery elm (farther to right). Along Modoc Creek, Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

218. From stream bank to tree tops- Ground to crown "photo-transect" of the mesic-limestone mixed hardwoods (sugar maple, bitternut hickory, white ash, hackberry, basswood, black walnut, northern red oak, chinquapin oak, american and slippery elm) Ozark Plateau forest. American bladdernut, pawpaw, and spicebush made up two (or three) distinct shrub layers while sugar maple (biggest, leftmost tree with holes used as dens by coon; many saplings in background), American elm (tree immediately right of maple), and chinquapin oak (background), which were most of the larger trees presented here, made up most of the uppermost canopy. Most regeneration was sugar maple with that of bitternut hickory, a distant runner-up. Along Modoc Creek, Ottawa County, Oklahoma. May (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

219. Tall hardwoods that like it cool and moist- A mesic-limestone forest on bluffs above a creek in the western Ozark Plateau. A fairly comprehensive list of major species (trees, shrubs, forbs, and grasses) in this tract of forest was given in the immediately preceding caption. Coverage in this caption was limited to specific featured species. Tree at left margin was black walnut. Sugar maples were to the close-by right and also behind the black walnut. Tall tree in foreground left of center was basswood. A young Kentucky coffeetree was to the immediate right of tall basswood and with crowns of the young tree passing in front of the bsswood. The three trees in right foreground were (left to right): American elm, hackberry, and white ash. Shrub layer (mostly in foregroune) were mixture of pawpaw, American bladdernut, and some flowering dogwood. The only herbaceous species visible at this distance were a large plant of silky wildrye and some leaves of the otherwise scarce Mayapple. Along Modoc Creek, Ottawa County, Oklahoma.Late May (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

220. Tall hardwoods later (and drier) in the summer- Two views of the same tract of north-slope, mesic-limestone mixed hardwood forest that developed along Modoc Creek, a stream typical of those in the Ozark Plateau. Overall dominant, including of regeneration, was sugar maple with bitternut hickory, basswood, white ash, northern red oak, chinquapin oak, American elm, and hackberry local associates. Sycamore and black walnut, pioneers of freshly denuded stream channels and banks, had persisted into this climax forest along its exterior. Forest range vegetation was presented extending from the creek bank to top of limestone bluffs in both of these slides. In the first of these two slides sugar maple (biggest tree; right-center), northern red oak (big tree left-center), basswood (big tree in right margin; background), and American elm (pole to left of big maple) were represented. In the second photograph northrn red oak (biggest tree; right background), chinquapin oak (foremost tree; left-center foreground), white or American and red or slippery elm (one sapling of each left and in front of northern red oak), and bitternut hickory (seedlings in right foreground), and sugar maple (most regeneration in foregoround) were represented (one way or the other). These photogrphs did not feature shrub layers, but American bladdernut was present and visible in second slide. Along Modoc Creek, Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

221.. At the base of a bluff- By the banks of Modoc Creek in the Springfield Plateau section of the Ozark Plateau (Mountains) some large trees of a mesic-limestone forest created a peaceful sylvan scene that belied the extreme competition for light (and undoubtedly, other resources) in this postclimax forest range vegetation. Tree in left foreground was an American or white elm (no umbrella-shaped crown on this forest dweller). The tree behind (and largely hidden by) the tall American elm with its upper bole leaning right was basswood. To the right (and behind at some distance) of the right-leaning basswood was a smaller sugar maple. Largest tree in this photograph (center foreground) was white ash. Smaller trees to left and right of large white ash were hackberry, including the large tree in right background. (Incidentially, real woodsmen would have noticed the black spot of a varmit in the right background hackberry, and we didn't even need our Black and Tans or Blueticks to tree it.) Smaller trees at right were sugar maple. Most of the young shoots in the understorey were also sugar maple. Regeneration in dense shade is a trademark of this Very Tolerant hardwood. Most shrubs were American bladdernut. Some pawpaqw were present. The spike of silky wildrye (lower right corner) was conspicuous (and a rangeman's signature on this slide). If any greenhorns were viewing this who could not locate the coon you might have better luck finding the large rotting log, the species of which could not be determined, that was close by the large white ash (in right center midground). Downed timber, especially big trunks and limbs that require long time periods to rot away, are part of the forest. A tree does not cease to be part of the forest range ecosystem just because it dies. There is life after death in the forest. Along Modoc Creek, Ottawa County, Oklahoma. Late May (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

222. A logger's view of trees at the base of a bluff- Vertical "photo-transect" of the larger trees introduced in the preceding slide and caption. The greenhorns' log served as focal point. Largest tree to left of log was the large white ash previously noted. Small tree to immediate left of large white ash was hackberry. Foremost tree (near lower left corner) was American elm. Tree at far left margin (only upper bole in photograph) was basswood. Smaller tree with dark trunk between (and behind) the American elm and the upper bole of basswood was sugar maple-- as were most of the seedlings and saplings in the understorey. Also visible in understorey were large plants of silky wildrye. Most shrubs (foreground) were American bladdernut. Along Modoc Creek, Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

223. As much light as you get in this woods if you're limited to ground-level- This poorly (barely) lite photograph showed the most light that this immediate location (local habitat) receives once trees have fully leafed-out in this north slope, sugar maple-dominated forest. Biggest tree (21 inches DBH) that was slightly left of center was a dandy sugar maple. Tree to its left (lower trunk at lower left margin) was northern red oak. Small sapling to left of northern red oak was white ash. Dominant shrub was flowering dogwood with eastern redbud the associate shrub species. Main herbaceous (not visible) was the lebuminus forb pointed-leaf or cluster-leaf ticktrefoil or tick clover (Desmodium glutinosum). Along Modoc Creek, Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

224. Regeneration at the bottom- Two side-by-side views of lower layers of the climax range vegetation of a mesic-limestone mixed hardwood (sugar maple, bitternut hickory-white ash-basswood-northern red oak-chinquapin oak) forest that developed along limestone bluffs above a typical Ozark Plateau stream (Modoc Creek). Understorey of the same tract of climax forest treated above and below. Tree trunk in these slides (upper left corner in first photograph; center of second photograph) as well as smaller trunk in second slide and fallen limb in first slide were sugar maple. Almost all sapling and tree seedlings in both "photo-plots" were either sugar maple or bitternut hickory with maple the more abundant of these generally co-dominant tree species. A sapling of American elm was present in upper left corner of second photograph.Trumpet creeper (Campsis radicans), a native liana, and Virginia creeper had notable regeneration. The broadleafed forb conspicuous in foreground of both slides was slender nettle (Urtica dioecia= U. gracilis). Its "excort" in lower left corner of first photograph was brown-eyed Susan (Rudbeckia triloba). It is only the lowerrmost leaves of this species that are three-lobed , and these were not visible in this photograph. The other forb in this sample of forest understorey was columbine (Aquiegia canadensis). Columbine was shown to better advantage in photographs below. Many of the other native forest forbs (see list in introduction to this forest range type) were present earlier in the spring growing season, but these had either entered dormancy or were not readily visible at time of photographs. There were two climax decreaser grasses that formed an almost exclusive herbaceous layer: 1) Canada brome or hairy, wood brome (Bromus purgans= B. pubescens ) and 2) woodland bluegrass (Poa sylvestris). These two species were clearly key indicator species of this entire climax forest community.

225. Deep inside a bluffs forest- Wide-angle view of the lower layers of a mesic-limestone forest dominated by sugar maple with bitternut hickory, white ash, northern red oak, basswood, and chinquapin oak local associates. Dominant shrub was American bladdernut with pawpaw the associate shrub. These two shrubs formed two distinct though discontinuous lower woody layers. Virginia creeper was quite common and occupied almost all layers of this forest including lower tree canopy.Flowering dogwood was present and, while conspicuous during "dogwood and redbud time", confined to upper reaches of bluffs and not common at this level on the bluffs.. There was some regeneration of all of tree species, but sugar maple outdistanced all others. Bitternut or pignut hickory was second in regeneration. Regeneration was both sexual and asexual from basal sprouting. Columbine (right foreground) was the most prominent herbaceous species at this time and from this camera station. Other common forbs in this photograph (though not visible) were Jack-in-the-pulpit, green dragon, dutchman's britches, rue anemone, false rue anemone, and bloodroot. The more abundant (and less showy) herbaceous species were Canada or hairy, wood brome and woodland bluegrass. Virginia wildrye and broadleaf woodoats formed locally dense stands. On limestone bluffs above Modoc Creek, Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

226. Climbing the bluffs- Climax vegetation of this mesic-limestone mixed hardwood forest (sugar maple dominant with bitternut hickory, white ash, northern red oak, basswood, and chinquapin oak "swapping places" as local associate species) changes over short vertical distances in progression up limestone bluffs. At this near-to-the-top zone sugar maple dominated the canopy and tree regeneration (various age/size classes as, for example, the sapling featured here). American bladdernut was still the dominant shrub in in this sample of vegetation (foreground of both photographs, especially prominent in lowr left corner of the second). Canada or hairy wood brome and woodland bluegrass were the dominant and assocaite species, respectively, of the grass(the taller) layer of the herbaceous understorey. Columbine was conspicuous in the second of these slides. Moss- and fern-covered ledges of the limestone bluffs to "top-off" this calendar cover-like view of pristine Ozark Plateau sugar maple-dominated forest range. On limestone bluffs above Modoc Creek, Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

227. These already climbed the bluffs- At ledges atop the limestone bluffs of this mesic-limestone, mixed hardwood forest Canada brome and a northern red oak seedling (lower left corner) had staked their claim to space, moist and shaded soil, and what little sunlight filtered through or fleetingly blazed at full intensity on their small spots. They were joined by columbine and wind flower or thimbleweed (Anemone virginiana). At top of limestone bluffs above Modoc Creek, Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

228. In its own shade, and more some- Two "photo-quadrants" of the understorey of a mesic-limestone, north slope forest dominated by sugar maple (with bitternut hickory, white ash, basswood, northern red oak, hackberry, and American elm other locally important tree species). There was no doubt as to which was the dominant on this forest range (and most range hungry animals had better like it or go elsewhere). The adult tree (center), saplings (small and large), seedlings, and even root or trunk sprouts were sugar maple. There was some American bladder, distinguised by its compound (largely trifoliate) leaves, buckbrush or coralberry, and silky wildrye (prebloom). Otherwise, this was an "all-ages gathering" of sugar maple. The second slide featured shoot or stump sprouting in a sugar maple sapling that lost its crown in a severe icestorm in February (three to three and a half months before time of photograph). This never-say-die small sapling simply started over at the bottom as it were. The ability to reproduce (sexually and/or asexually) in "its own shade" is the defining feature of a species with high tolerance. Sugar maple has a tolerance rating as high as it goes: Very Tolerant. This was according to the Society of American Foresters (Wenger (1984, p. 3) which gave ratings of Tolerant for basswood while white ash, bitternut hickory, hackberry, northern red oak, and American elm were Intermediate. Along Modoc Creek, Ottawa County, Oklahoma. Late May (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

229. Not dead yet (at least, not quite)- Stump suckering (= coppicing) from an otherwise dead sugar maple on a sugar maple-dominated forest that developed on a bluff above Modoc Creek in the western portion of the Ozark Uplands. These were long or heterophyllus shoots (in contrast to fruit-bearing short shoots). Production of stump suckers is a form of asexual reproduction that is well developed in many hardwood trees and shrubs that are interpreted as having higher levels of Tolerance. Sugar maple has the highest rank of Very Tolerant (Wenger, 1984, p.3). In this tract of north slope, mesic-limestone forest sugar maple had various tree species that were local associates including bitternut hickory, white ash, basswood, northern red oak, and American elm. Note the Jack-in-the-pulpit and various ferns in front and to side of the trunk and long shoots. Along Modoc Creek, Ottawa County, Oklahoma. April (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

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

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

232. And Jill the basswood came tumbling after- When the crown of the white ash (discussed with the immediately preceding two slides and caption) crashed in a February icestorm it took an neighoring (and even larger) basswood down with it. In fact the basswood was brought even lower as the additional weight combined with wet soil resulted in complete toppling of the basswood. The first of these two photographs presented immediate local habitat (microsite) by the toppled basswood (including the stump of trunk from which the landowner had cut firewood). The owner had not worked up the small side shoot of this basswood that was on the ground surface (center foreground to left lower corner). The basswood had sprouted profusely all along this secondary bole. (A morphological feature of basswood is on-going or continued production of secondary shoots off of the main (original) tree trunk.) Also visible in this first slide were resprouting and seedling plants of white ash, sugar maple, and American elm along with American bladdernut and several pre-bloom-stage plants of silky wildrye, the dominant grass in this mesic-limestone, north slope, Ozark Mountains forest. The second of these slides showed stump sprouting (=suckering= coppicing) of the basswood in more detail. Along Modoc Creek, Ottawa County, Oklahoma. Late May (estival aspect; coppicing stage in basswood).

233. Leafy layers- Follow-up "photo-plot" of stump sprouts of basswood in middle of second summer of growth following downing in an ice storm (two winters earlier). This photograph also presented associated herbaceous species including silky wildrye, broadleaf woodoats, and lopseed (Phryma leptostachya). The latter, a forb, is in its own family, Phrymaceae. On limestone bluffs just above (on upper banks of) Modoc Creek, Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

334. Past-prime autumn coloration (in sugar maples), but still instructive- Late autumn in a north slope, sugar maple-dominated Ozark forest was still an appropriate time to describe the postclimax climax vegetation of this forested range. This was the same tract of forest (along and above Modoc Creek) that was presented and discussed in the preceding slides of the Sugar Maple Forest section. (See again first photo-caption for comprehensive list of major plant species.). Plants featured in the autumn scene were sugar maple (big trees at extreme left and right margins, Big tree in center midground (midway up the bluff) still bearing green leaves was a northern red oak. Tree with green leaves at far left (left margin) was a chinquapin (chinkapin) oak. Trees with retained dead leaves were sugar maple. Along Modoc Creek, Ottawa County, Oklahoma. Late November (autumnal aspect; leaf fall for most trees and shrubs except the oaks). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

235. Another instructive view of gone-past-autumn color- Vertical look at a sugar maple-dominated forest on a limestone bluff (above Modoc Creek) in the Springfield Plateau of the Ozark Highlands. Largest tree (center foreground; midway up the bluff) and the larger tree on top of the bluff (to left of largest tree) were sugar maples readily distinguished by their large, crooked limbs. Tree in front of and to left of largest tree (and still bearing green leaves) was a northern red oak. The several trees at far right mid- to background included sugar maple, basswood, and chinquapin oak. Most seedlings and saplings of understorey were sugar maple, the Very Tolerant, climax dominant tree species of this mesic-limestone forest (Nelson, 2005, ps. 122-125). Attention was drawn to the large, flat rock in left-of-center foreground which was limestone. Along Modoc Creek, Ottawa County, Oklahoma. Late November (autumnal aspect; leaf fall for most trees and shrubs except the oaks). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

236. Spring slope- "Photo-transect" on the north slope of the sugar maple-bitternut hickory in early spring. The spring flora of the mesic-limestone forest (Nelson, 2005, ps. 122-125), especially the lower herbaceous layers, that was described in this portion was one of the most unique or conspicuous features of this climax forest range vegetation. This early vernal aspect showed the temporal and spatial variation of this mesic Ozark Highlands forest. Species details were presented in the two immediately succeeding photographs. The farmer has to "make hay while the sun shines", and so do most of the florest floor-dwelling plants in this sugar maple-dominated plant community. Once the leaves are fully developed on the maples and associated tree species, along with the shrubs, there is inadequate light for most shorter plant species (this includes almost of the herbaceous species). To survive, forest forbs (which dominate the herbaceous layers) must complete their annual cycle of life (= growth cycle) as early as spring temperatures permit their growth and reproduction. Ottawa County, Oklahoma. April.

237. Details of spring slope- Two "photo-plots" of the vernal herbaceous understorey on a north slope, climax sugar maple-bitternut hickory forest that was introduced in the preceding slide. Species on the north slope of this forest floor included Jack-in-the-pulpit, wake robin, Solomon's seal, false Solomon's seal, wild geranium (Geranium maculatum), green dragon, trout lily, mayapple, Dutchman's breeches, and the fungus known simply as morel (Morchella rotunda.. The first and last of these forest range plant species were conspicuous. Students should try to find some more of the just-listed species in these two photographs. (Like a good matching question not all named species were present, and there could have been a species or two present in the photographs that was not listed. Good Luck.) Ottawa County, Oklahoma. April.

Beginning and end of plant succession in the mesic-limestone, mixed hardwood forest of the Ozark Plateau. The following short section showed the beginning (pioneer stage) and ending (climax forest range vegetation) of a sere of mesic-limestone forest on a stream floodplain in the western Ozark (Springfield Plateau) Mountains. The sample of this range cover type was the same location and tract of forest treated immediately above. This was a continuation of the north-slope, limestone bluffs drainage of Modoc Creek, Ottawa County, Oklahoma.

238. Started on a rock bar- Two "photo-transects" of the exterior of a mesic-limestone forest that developed along a typical Ozark Plateau stream (Modoc Creek). These photographs were centered on the currently dry shallow creek (half on the stream bed and half on the stream bank) and extended upward taking in lower portions of a limestone bluff. The first furnished a distant view while the second of these two slides was a nearer view that showed details of streambank vegetation. Most of this stream bank had been been scoured down to bare stream stones and gravel (chert and limestone) following removal of these creek bed materials in the foreground by county road crews (ie. land in the immediate foreground had been used as a borrow pit for roadbed material). This lowered land surface (bar pit) was refilled with more stones, gravel, sand, mud, and other alluvium the next time Modoc Creek flooded. These photographs were in taken in mid-spring two years after flood waters restored the creek bed and low stream bank (start of third warm-growing season following last flooding and stream channel restoration). Vegetation that pioneered the fresh rubble of the stream channel included both annual and perennial, herbaceous and woody, and native and naturalized (or adventive) species. Sycamore seedlings (more obvious in foreground of the second of these photographs) from the dying adult sycamore (center midground in both photographs) had pioneered the disturbance site. This was the only tree species that had regenerated on this denuded area. There were seedlings of sugar maple in rocks along the creek bank, but these grew mostly under shade (to rear of the parent sycamore) and not on the disturbed part of the channel.. There were only a few individuals of annual pioneer forbs like giant ragweed (Ambrosia trifida). The most abundant (density and cover) annual pioneer was hairy crabgrass (Digitaria sanguinalis). The native festucoid grass, Virginia wildrye, was one of the climax grasses in this forest vegetation and it was the most grass even exceeding hairy crabgrass. Weedy perennial forbs were also better represented than annual species. The more common forbs included the Eurasian curly dock (Rumex crispus) and the native, colony forming Pennsylvania smartweed or pinkweed (Polygonum pensylvanicum) plus big blue strar (Amnosia tabernaemontana).. Woody vine on the dying sycamore was trumpet creeper (Campsis radicans). There were many seedlings of this linana.that had established on both bed and bank of the stream channel (near base of sycamore). In exterior (physiogonomic) view of the mesic-limestone forest in the first of these slides the tallest tree (big crown in upper left corner) was northern red oak. The crown to left of the tall red oak was basswood. The next crown to the left was of chinquapin oak. All of the shorter trees and saplings (in front of the tall northern red oak and to right of the dying sycamore) with densely leaved crowns at outer edge of this stand were sugar maple. Details of this forest understorey, including grasses and forbs on the limestone bluffs was treated above. Stream channel of Modoc Creek, Ottawa County, Oklahoma. May (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple-bitternut hickory forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

239. Later in the summer on the same rock bar- Same rock-gravel stream bed and bank of Modoc Creek described immediately above. These two photographs were taken two months later as follow-up "photo-plots" to the preceding two transect views of this pioneer stage of a climax sugar maple-bitternut hickory-white ash-northern red oak-chinquapin oak forest, a north-slope, mesic-limestone mixed hardwood forest in the Springfield Plateau portion of the Ozark Plateau Region. This seral vegetation was developing on newly laid-down, channel material (limestone and chert) in early summer of the third growing season following deposition. The species composition was obviously the same as that of two months earlier (see above caption) except that some cool-season species (eg. curly dock, Virginia wildrye) had entered summer dormancy while warm-season species (giant ragweed, hairy crabgrass) were still growing toward full maturity and sexual reproduction. A single plant of brown-eyed Susan was blooming (at base of dying sycamore and not readily visible). The most conspicuous feature of this range vegetation was growth of sycamore seedlings which were now the tallest plants on the gravel bank of Modoc Creek. The trumpet creeper on trunk of dying sycamore was in bloom, and its seedlings around its trellis had also grown considerably in the two months since first "photo-plots" were recorded. Tree crown outline of the forest in background (second of these photographs) were from tallest crown at right margin moving leftward: northern red oak, basswood, chinquapin oak, and dying sycamore. Shorter trees in front (many were saplings) with crowns of especially dense leaves were all sugar maple. Stream channel of Modoc Creek, Ottawa County, Oklahoma. May (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple-bitternut hickory forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

240. Followup visit to the rock bar- Development of mesic-limestone forest vegetation on the same rock-gravel stream bed and bank of Modoc Creek introduced above except in the subsequent summer. after the above. This was still the pioneer (or, perhaps, second seral) stage on a sere of climax sugar maple-bitternut hickory-white ash-northern red oak-chinquapin oak forest. The sycamore had advanced to young sapling class, but this pioneer tree species was now joined visibly with seedlings and small saplings of American elm, white ash, and bitternut or pignut hickory. Elm had the apparent greatest cover of these three three species all of which are components of the climax forest. The alluvium (stones, gravel, coarser sand, alluvial soil particles) that had been laid down three years earlier by flood waters still supported populations of "new land" pioneer annuals including giant ragweed and hairy crabgrass that had been so conspicuous one year earlier. In addition, common cocklebur (Xanthium strumarium) had joined the list of pioneer plants. Also noticed for the first time was bracted plantain (Plantago major), a naturalized Eurasian annual forb. It was on the more recently laid-down alluvial material in the stream bed itself. Curly dock was still present on the now three-year-old alluvium, but it was being overshaded (and, consequently, losing vigor, cover, and density. The population of Pennsylvania smartweed or pinkweed was (pretty much) stable while the perennial forb, big blue strar, had increased in cover and density. Apparently most of the many seedlings of trumpet creeper that had established last year were larger and thicker "than ever". Poison oak/ivy had noticably joined previous occupiants on the channel bed and bank. Most telling succession-wise was migration of broadleaf woodoats down from higher up (older alluvium) on the shaded channel bank down onto the three-year-old alluvium where it joined Virginia wildrye, the other native, climax grass of this forest range site, that had established a year earlier. The introduced pasture grass, tall fescue (Festuca arundinacea) had also joined the cast of grasses, but it was moving in from the outer or sunlite edge whereas the two native grasses were dispersing and migrating in from the shady or forest side. Stream channel of Modoc Creek, Ottawa County, Oklahoma. Early July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple-bitternut hickory forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

241. Climax of the mesic-limestone, mixed hardwood forest- Deep inside a bottomland (floodplain of Modoc Creek) in western Ozark [Springfield] Plateau) forest of white ash, bitternut hickory, sugar maple, sycamore, northern red oak, black walnut, and honey locust. The second=growth forest vegetation in this photograph developed on a meander of an Ozark stream. Large inward (leftward) leaning tree at far right was sycamore as was end of limb in upper right corner. The other left-leaning trunk that ran from lower right corner to top center was a white ash, the trunk of which was covered with Virginia creeper. Sapling in center foreground and seedlings in immediate foreground were all bitternut or pignut hickory (Carya cordiformis). The sapling above this lower layer of regenerated bitternut hickory with sunlit-bright leaves leaning from the extreme left margin rightward to center of photograph was also a bitternut hickory. The cluster of young well-lit young leaders and leaves were resprounts when the crown of this young hickory had been sheared "clean-off" by a large limb of the whte ash that was brought crashing down in an ice storm. The smaller (but now taller) sapling to left of the ice-topped hickory was northern red oak, the species that was second to bitternut hickory in reproduction. Twigs/leaves of black walnut in upper left corner. White ash, bitternut hickory, and northern red oak are climax dominants of this forest site. Black walnut sycamore are pioneer (usually Intolerant species, especially walnut) species that live long enough to persist into the climax (at least subclimax) forest. There were no sugar maple on this creek overflow parcel of bottomland in the meander of Modoc Creek. Instead sugar maple was limited to the north slope of limestone bluffs above the creek where it was the dominant tree species. Regeneration of bitternut hickory and northern red oak was so dense that there was almost no herbaceous growth other than scattered silky wildrye and broadleaf woodoats. Shrub cover were also very sparse and limited mostly to American bladdernut. On a meander of Modoc Creek, Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax sugar maple-bitternut hickory forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

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

242. Leaves and fruit of sugar maple (Acer saccharum)- Details of leaves and nearly mature schizocarp of sugar maple. The fruit of maple has been interpreted as either a samaroid schizocarp having two winged mericarps or as two samaras joined together (Smith, 1977, p. 165). Schizocarp is a dry fruit with carpels separated from each other into single-seeded indehiscent segments called mericarps, which in Acer species are winged; samara is also a dry fruit and a winged one that is indehiscent with a single-seed (Smith, 1977, ps. 66, 307). Ottawa County, Oklahoma. August. Stage of phenology: premature but late fruit stage (immediate or near-term maturity of fruit).

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

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

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

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

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

248. Flowers of American bladdernut- Flowers of bladdernut grown in clusters that hang down in a racmeme-like arrangement. These inflorescences often occur on tips of smaller branches off of the main limb of the shoot. Bluffs above Modoc Creek, Ottawa County, Oklahoma. April.

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

250. Rolling in brome- Local stand of hairy wood brome (Bromus purgans= B. pubescens) on an upland, black oak-dominated forest that had a surface wild fire in spring of the preceding spring (ie. this stand was in the second growing season following a a spring wild fire). This upland forest was just above the limestone bluffs along Modoc Creek which was the forest site of the a mesic-limestone mixed hardwood (sugar maple-bitternut hickory-white ash-basswood-northern red oak-chinquapin oak) forest that was featured in this section. Hairy wood brome, which was shown as Canada brome in Barkworth et al., 2007, p. 220), is distributed sporadically throughout eastern North America ranging from central Manitoba across to and then south to Florida and westward to central Texas. B. pubescens has an interupted species (biological) range with local occurrence in in Arizona and in Colorado and Wyoming. Incidentially, the common name of Canada brome seemed to this author to be unwarrented and clearly inferior to hairy or hairy wood brome given the specifici epithet, "pubescens", rather than a commenerative one featuring Canada. The common habitat of hairy wood brome is forests and woodlands. Ottawa County, Oklahoma. Late May; peaking standing crop of herbage, anthesis stages of phenology.

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

252. Hairy spikelets- Spikelets of hairy wood brome in anthesis. These units of the panicle were produced on plants growing in the brome stand on a burned-off (spring wild fire) upland forest dominated by black oak shown above. Above bluffs of an Ozark Plateau stream (Modoc Creek)Ottawa County, Oklahoma. Late May; peaking standing crop of herbage, nthesis stages of phenology.

253. Shoot! Look at the shoots!- Culm details of shoots of hairy wood brome. Although organs of some shoots of this species are glabrous (Barkworth et al.(2007, ps. 220-221) they typically have a characteristic pubescence (hence the inclusion of "hairy" in common name and "pubescens" as specific epithet) as shown in these examples from the western Ozark Plateau. Another key characteristic of culms in this species is the prominently swollen nodes. Bluffs above Modoc Creek, Ottawa County, Oklahoma. Late May (peak standing crop, anthesis phenological stage).

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

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

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

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

The next series of photographs and their captions was of another sugar maple-dominated forest also along Modoc Creek in the Springfield Plateau section of the Ozark Highlands. This forest tract was an east slope forest located less than one mile downstream from the sugar maple-dominated forest just described. The east slope sugar maple forest had fewer species--except for ferns--overall. Most conspicuous was white ash (versus bitternut hickory) as the most frequent associate tree species followed by northern red oak and chinquapin oak. Basswood, bitternut hickory, and American elm were infrequent and did not approach associate species status even in local stands. This east slope, limestone bluff, sugar maple forest--another variant of mesic-limestone forest (Nelson, 2005, ps. 122-125)-- was postclimax in the general Ozark Plateau oak-hickory forest. The sugar maple-dominated forest described in the following portion was contiguous with a mesic bottomland forest (Nelson, 2005, ps.147-150), except for the separation by stream corridor of Modoc Creek, that was described later in this section on Miscellaneous Forest Types.

257. East slope sugar maple forest- Another example or phase of sugar maple-dominated forest in the Ozark Plateau was also found along Modoc Creek. This forest tract had developed on an east slope and differed from the sugar maple-bitternut hickory forest type or subtype (variant) that developed on a north slope of Modoc Creek located only 3/4ths to one mile upstream from the forest vegetation described here. This sugar maple-dominated community was much less species-rich except that it had more ferns including maidenhair (Adianthum pedatum) and Christmas ferns (Polystichum acrostichoides). In this forest communitysugar maple was sole dominant with northern red oak and chinquapin oak (Quercus muehlembergia), and white ash being local associate species. Mature trees of these two oak species were generally larger than all but a few of the slower-growing sugar maples. There were also larger, less abundant trees of sycamore, bitternut hickory, and black walnut. Most regeneration was of sugar maple with somewhat less reproduction in box elder and northern red oak with these two species swapping places depending on microhabitat. Co-dominant shrubs overall (and of two different layers)were American bladdernut (taller) and spicebush (shorter-growing). The tallest shrubs (those of the taller shrub and/or lower tree layer) were flowering dogwood (more abundant) and eastern redbud. Pawpaw was present, but was primarily on the adjacent (and other side of the creek hackberry-American elm-sycamore-eastern cottonwood bottomland forest). Lanceleaf buckthorn (Rhamnus lanceolata) grew in association with American bladdernut, and though usually overtopping the latter, R.lanceolata was uncommon. Almost no herbaceous species other than ferns. Occasional plants of Virginia or silky wild-ryes or beakgrain (Diarrhena americana) were present, but these-like woodreed grass (Cinna arundinacea)-were usually confined to rock or gravel bars away from the sugar maple forest. Grasses were much less common than on an adjacent hackberry-elm-box elder forest on the stream floodplain (described below). The two wide-view photographs shown here presented species composition and structure of a postclimax sugar maple forest that developed on an east-facing limestone bluff along a typical stream in the Ozark Plateau. In the first of these photographs a large hackberry (largest trunk; dead center of photograph) and black walnut (tree to right of big hackberry) added some diversity to an otherwise solid stand of uneven-aged sugar maple. In the second of these slides an immense sugar maple (25 inch DBH) overlooks a nearly "pure" stand of its species (undoubtedly many of which were its own offspring). Even smaller adult trees at right were mostly sugar maple. Lanceleaf buckthorn (Rhamnus lanceolata), an exmple of which was tallest shrub in front (and slight to right) of large sugar maple, American bladdernut, flowering dogwood (conspicuous in right foreground), and American hazlenut (Corylus americana) were the primarily shrubs vbisible in the second photograph. Along Modoc Creek, Ottawa County, Oklahoma. July (aestival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

258. Broad view of an Ozark Plateau bluff and its forest- A wide-angle "photo-transect" on an east slope of a limestone bluff along a typical stream in the western part of the Ozark Highlands on which a postclimax sugar maple-dominated forest (with white ash and, to lesser extent, northrn red oak were associate tree species) had developed. This was a mesic-limestone forest (Nelson, 2005, ps. 122-125) that in monoclimax theory (Clements, 1916) was postclimax to the regional (climatic) climax oak-hickory forest. This photograph introduced forest range vegetation that was viewed at increasingly closer-in scale in subsequent slides.The huge sugar maple at the right served as a "landmark" and featured topic in the subsequent slides. The smaller tree in left midground with the right angle-like trunk leaning sharply to the left was another sugar maple. The tree in upper left margin (upper right corner) was a northrn red oak. Shrub species included American bladdernut (generally the dominant shrub), lanceleaf buckthorn, flowering dogwood, shadbush or, as it is also known, eastern serviceberry, and eastern redbud. There were only occasional herbaceous species that were not forbs, most of which were ferns, as most grasses and sedges grew closer to the creek. Along banks of the stream (Modoc Creek) there were occasional individuals of silky or Virginia wildryes, beakgrain, or giant woodreed. Most common ferns were Christmas fern and maidenhair fern. Regeneration of climax dominant trees (mostly sugar maple, white ash, and northern red oak) comprised much of the lower layers of the understorey as the young of these trees ranged from seedlings to large saplings. Along Modoc Creek, Ottawa County, Oklahoma. July (aestival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

259. Postclimax and "pure"- An east slope of a limstone bluff above an Ozark Plateau stream provided the setting for a "nearly pure" stand of sugar maple. In the Clementsian monoclimax theory this was a consociation of Acer saccharum that was postclimax vegetation where the climatic =regional or zone) climax was oak-hickory forest, the Oak-Hickory Forest Region (an association) of Braun (1950, ps. 35, 162-191). The mighty sugar maple in center foreground was the sugar maple described as "huge" in the immediately preceding photograph. This large maple had a large (by standards of its species) lanceleaf buckthorn to the right and in front of it. Shrub in lower right corner was flowering dogwood. Shrub species in left foreground included American bladdernut, pawpaw, and flowering dogwood. Many of the shrubs higher up on the bluff were shadbush or, as is also known, eastern serviceberry. Almost all tree regeneration was sugar maple with traces of northern red oak and white ash. Details of understorey of this immediate (local) site was given in the next slide and caption. Along Modoc Creek, Ottawa County, Oklahoma. July (aestival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

260. On the bank- On the bank of a typical stream in the Ozark Highlands there was a multi-layer understorey in a sugar maple-dominated forest on an east slope of a limestone bluff. Tree trunk in center midground was that of the mammoth sugar maple that served as the "landmark" in the immediately preceding slide. Shrub to right (and with its upper shoot growing to the right)was lanceleaf buckthorn. Most of the other shrub cover was that of American bladdernut. Ferns in edge of stream bank were Christmas fern, a major forb in this mesic-limestone, postclimax forest. Also present was a ground layer composed of luxuriant cover of moss, the species of which this bryophyte-ignorant author knew not. Along Modoc Creek, Ottawa County, Oklahoma. July (aestival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

261. A bluff full of beauties- An east-facing limestone bluff aligned along Modoc Creek in the western part (Springfield Plateau) of the Ozark Highlands supported a mesic forest dominated by sugar maple (with white ash and, secondly, northern red oak as associate tree speceis). This forest range was postclimax in the general oak-hickory forest of the Ozark Mountains. This photograph featured species composition (especially of dominants and associates) and structure of this southern and western outlier of the sugar maple forest cover type. In this stand white ash was the clear associate to sugar maple, but with northern red oak a close "runner-up". Large tree at right midground (and midway up the bluff) was sugar maple. The large tree to the right of this sugar maple (along right margin of photograph) was northrn red oak. The adult tree closest to the big sugar maple (left of it with its upper bole in the crown by large left limb of the maple) was white ash as was the largest tree in this photograph which was largely obscured by two smaller hackberry trees. The largest tree, the white ash, had lost much of its crown which retained a large left-leaning limb. Major shrub was American bladdernut, but flowering dogwood, American hazelnut, shadbush, and eastern redbud were present to locally common. Along Modoc Creek, Ottawa County, Oklahoma. July (aestival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

262. Sugar maple-white ash stand- Composite view of species composition and structure of a mesic-limestone (east slope; bluff habitat) with sugar maple the dominant and white ash the associate species. Biggest tree (right side) was sugar maple. Second largest tree ((left of center midground) was white ash. Most regeneration was sugar maple and white ash, including the larger saplings in baqckground. Hackberry at far left. There were two shrub layers: 1) taller layer dominated by flowering dogwood, lanceleaf buckthorn, and American hazelnut (the large shrub in front of the big sugar maple) and 2) lower layer almost exclusively American bladdernut but with considerable poison oak/ivy. Grape vines grew to top of canopy and formed a "unifying wooden thread" among the various layers of forest range vegetation. Herbaceous layer(s) consisted primarily of Christmas and maidenhair ferns. Mosses made up a lush ground layer (the lowest layer of vascular plants). The black circular "spot" in upper left was entrance to a small cave in the limestone bluffs. This is a common feature of the ancient Ozark Mountains. Along Modoc Creek, Ottawa County, Oklahoma. July (aestival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

263. Hole in the canopy- Example of gap or patch dynamics in a sugar maple-dominated east slope bluff forest in Ozark Plateau. A postclimax forest range in the general oak-hickory forest had extensive feeding by beaver (Castor canadensis) 17 years prior to this follow-up photographic session. Beaver killed a high percentage of the largest northern red oak (along with lesser browsing on American elm, box elder, sycamore), but fed on very little sugar maple. This gap was created 17 years earlier when beaver girdled three massive northern red oak (all were between two and a half and three feet DBH) which died in the spring following winter browsing a few months earlier (the three oaks never really leafed-out). The light-colored trunk in right background was the snag of the only one of the three northern red oaks that had not fallen. Death of the three giant oaks left such a gap in the canopy that the stage was quickly set for the process of recovery of the forest vegetation through secondary plant succession. Such vegetation dynamics--to use the term made famous by F.E. Clements who adopted it from H.C. Cowles--is studied as patch or gap dynamics which is unique from the perspective of small spatial scalet as in, say, the larger spatial of a large forest clearcut, blowdown, fire, or old-fields as in cut-over lands. Most tree regeneration was of sugar maple (including the sugar maple sapling in lower right foreground) and white ash. The large limb, which was shown at close range in the next (succceding) slide, was off of a large northern red oak (outside of camer range at left) brought down by a February icestorm. The falling red oak limb brought down canopies of two medium-sized black walnut trees (left midground). Beneficiaries of this tree damage were understorey plants, espceially herbaceous species the most common of which were maidenhair fern and the pioneer composite, giant ragweed (Ambrosia trifolia). American bladdernut was the most common shrub in the gap. Along Modoc Creek, Ottawa County, Oklahoma. July (aestival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

264. Ground level and ground zero- Lower and mid levels of forest range vegetation in a seventeen-year old gap in a sugar maple-dominated forest that was made when beaver girdled three massive northern red oak. In the 17 years following beaver-felling of old-growth trees the "dynamics of vegetation"--to borrow a title for selected writings of F.E. Clements (Allred and Clements, 1949)--had resulted in re-establishment of sugar maple and white ash, the two major (dominant) climax tree species for this forest site. Of course trees of these two climax domiants were still small, but the species composition had quickly progressed to that of the climax forest vegetation. The dominant shrub was American bladdernut which, while not as abundant as on the banks of the stream (Modoc Creek), still had appreciable cover and density. The largest and one of the most common herbaceous species was giant ragweed. It was not known whether this pioneering annual composite was more plentiful due to (or even present only after) disturbance due to tree damage from the February icestorm .There was very occasional cover of Virginia wildrye, but in more shaded local habitats the most common forb was maidenhair fern. The sugar maple sapling (right-of-center foreground), that became conspicuous within a couple or three years following death of three massive northern red oaks by beaver girdling, had in the interim become well-established and received limited beaver feeding (note blaze on lower trunk). Obviously beaver abandoned this feeding station quickly and before the sugar maple sustained enough feeding damage to be life-threatening, at least in the short-run. (Reduced performance or even death might be possible via disease entry through the debarking wound where beaver fed, however limited that was.) Along Modoc Creek, Ottawa County, Oklahoma. July (aestival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

265. Life among the fallern on the forest floor- A severe February icestorm brought down a large part of the crown of a northern red oak (which in a chain reaction brought down crowns of neighboring black walnut as shown in a preceding photograph). About six months later, as shown in this photograph, maidenhair fern and regenerated (seedling and/or root-sprouting) sugar maple were apparently benefiting from the ecological windfall caused by ice. Along Modoc Creek, Ottawa County, Oklahoma. July (aestival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

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

266. A lingering death and the tragic end of their reign- The first act of the unfolding forest drama (tragedy or comedy was left up to viewers' discretion) that was revealed above began with the empty stomach and ever-growing incisors of beaver(s). Bark-feeding by beaver during winter months left three very large northern red oaks (each over two and a half foot DBH) girdled and soon-to-be-dead in an east slope, mesic-limestone, sugar maple-white ash climax forest. The tree in left background as well as the trunk in midground, and partially concealed by the foremost tree, were all northern red oaks and completely girdled. Death would "officially" wait until spring when, with rising sap, these once-magnificant oaks would fail to leaf-out. (As it turned out, the preformed, terminal buds of these oaks did not fully open before they fell off indicating that trees were dead.) None of the three trees sent up any stump sprouts. Lack of suckering (production of basal shoots) is the typical or normal physiological response of old hardwood trees. Death of the entire tree (not just top-kill) was "sure and swift" for all three mature "timber-ripe" oaks. These three northern red oak were some of the largest trees in this east slope, limestone bluffs forest. Only a few sugar maple and white ash reached similar size. Trees of these latter two species were not browsed. This was a textbook example of feeding selectivity (= selective browsing). Locally these three oaks were (had been) "far and away" the largest trees that controlled the largest portion of the canopy (ie. had the greatest impact on how much and for how long light reached lower levels of the forest). Loss of the "masters" of the forest canopy set the stage for secondary plant succession on the bluffs above Modoc Creek. Oange inner bark is characteristic of several of the red oak species (Erythrobalanus subgenus) , especially black oak, and it showed prominently on the three girdled northern red oaks featured here. "Reading sign" revealed that beaver gnawed higher (almost three feet above ground level) on the upslope side of the trunks showing these rodents (largest rodent species in North America) fed on their hind legs on the high side of trunks while "standing on all fours" on the downhill side. Tthe author's 38-inch, hickory walking stick was placed lengthwise at base of the foremost (and largest) oak for scale. Historically there have been high population densities of beaver along Modoc Creek and other streams in this local area. As a boy the photographer's younger brother trapped beaver for several years along these streams. He caught a few adult male beaver that exceeded 50 pounds in weight. These were (are) bank-beaver. The beaver do occasionally build high dams on streams causing localized flooding of bottomland forests, but as far back as local observers can remember beaver never built stick-and-mud lodges. Rather, long tunnels dug far back into creek banks served as beaver dens. As was shown in this photograph adult beaver often fed at considerable distances from streams. This was never much more than half-way to top of bluffs (usually feeding was limited to approximately the lower one-third of bluff height). In other words, trees that grew high up on bluffs were safe from beaver browsing or, from a forester's perspective, beaver depredation. Above Modoc Creek, Ottawa County, Oklahoma. February, 1990.

267. Standing tall, proud, and dead- On east-facing, limestone bluffs above Modoc Creek the tallest and largest-diameter (over 30 inches DBH) of three northern red oaks was quickly dying (after only partially leafing out) following bark-feeding by beaver during the immediately preceding winter months. Death of the three northern red oaks was complete in a month or month and a half following bud-opening. Most of the neighboring (surrounding) trees--all of which were considerably smaller and, presumedly, younger--were sugar maple and white ash plus a few black walnut. Death of the largest--and locally dominant--northern red oaks created a forest gap or patch almost as rapidly as windthrow (blowdown) or lightening strike. Beaver were not only hungry and "eager" they were lethal. The stage was preparing for the second act of patch or gap dynamics. Above Modoc Creek, Ottawa County, Oklahoma. August, 1991.

268. The new living among the old dead- Two local "photo-quadrants" showed details of and adjacent to the largest of three northern red oaks that died after being girdled by bark-feeding beaver over a period of roughly two to four months prior to an aborted effort by oaks to leaf-out in spring. The first fact to "leaf-out" from the "crime scene" at this photographer was that the large oak (three foot DBH) was largely hollow. The trunk of this otherwise healty northern red oak (prior to beaver strike) consisted almost solely of sapwood. This condition of "hollow-heart" (by which it is known among Ozark sawmill operators) probably had little, if any, influence on this tree because 1) heartwood is dead and provides no life-supporting function and 2) a hollow tree, like many hollow cylinderical things in nature (grass culm, bird quill) is almost as strong as trees with solid (non-hollow) trunks. Therefore, beaver did not bring down a tree that was dying (they killed a sound, healthy tree). On the other hand, beaver did not destroy a valuable lumber tree and cause economic loss to the firm or landowner. The huge tree was useless for lumber and was fit only for fuelwood (the trunk would not have made a saw log and loggers would have wasted their time felling the tree). In fact, beaver conducted their own version of "site preparation" by clearing the way (making resources available and conditions more favorable) for younger trees that could grow sound boles for future (and more valuable) wood products. Let's here it for the beaver! Plus, the browse (bark) of the oaks contributed to the production of more beaver plew (pelts), another product of the forest. Why, it might even stir that younger brother to re-run his trap line (at least renew his subscription to Fur, Fish and Game). The hollow-trunk northern red oak was an example of "over-ripe" timber known technically as overmaturity or overmature which when applied to individual trees refers to the condition of having reached the "...stage of development when it is declining in vigro and health and reaching the end of its natural life span" or "...one that has begun to lessen in commercial value because of size, age, decay, or other factors" (Helms, 1998). It was possible (probably likely) that once it was dead, and with weakened and soon-rotting sapwood, the hollow oak crashed sooner than if it had been solid. In point of fact, one of the three girdled northern red oaks was still standing 17 years following its death (the snag standing conspicuously on the bluff in two of the photographs presented above). The solidness of this trunk was not determined by the author because cutting into the tree to determine its state of soundness could have caused it to fall when it might otherwise have continued to stand for decades. (Besides only a hollow-headed idiot would risk life and limb--catch the pun--to tote a double-bit up such a steep, slippery slope to cut a tree that was useless for wood. Now slinging a 35mm SLR Nikon for educational purposes--that is, to chop out ignorance--is a different grade of lumber.) The other--and more relevant--fact from forest vegetation and succession standpoints was the tree species that were replacing the former "mighty oak". Saplings and seedlings of sugar maple began almost instantly to fill the gap created by the beaver. Obviously, maple saplings had been growing beneath this northern red oak prior to beaver "attack". Furthermore, sugar maple is rated as Very Tolerant (Wenger, 1984, p. 3) so that continued survival and growth of sugar maple was likely even with continued canopy cover of the potential still-living oak. Nonetheless, more resources and modified conditions (including increased light, space, and soil moisture) undoubtedly was of some benefit to sugar maple growth and regeneration. There were also several age-size classes of white ash (and of northern red oak) in the gap formed by death of the large northern red oak. The tree behind the fallen oak was one of the other two northern red oaks killed--but still standing--by beaver. Above Modoc Creek, Ottawa County, Oklahoma. September, 1991.

269. Down and out (and after only one "beaver strike")- The "long view" of the largest of three northern red oak killed by beaver girdling in the preceding winter. This hollow-trunk old patriarch fell across the channel of Modoc Creek creating a forest gap in which already active-growing sugar maple and white ash (even a few young northern red oak) were ready to utilize more available resources and different conditions (not least of which was more light, space, and soil water). It would be humanly impossible to know all of the ways in which death and crash of this old-growth speciment affected life in this east slope climax forest, or of the creek below it or the bottomland forest (another forest cover type and separate tract of forest) separated from it by the stream. Even the trunk and crown of the oak across Modoc Creek could conceivably have some impact on some organisms. For instance, it could permit some animals to cross to other side of the stream. The potential barrier to movement, which though not exactly a grand canyon, had been breached. This was of no momemt to a coon, but it could be determinative to a biped (such as the hillbilly who took this photograph). Above Modoc Creek, Ottawa County, Oklahoma. September, 1991.

270. Bridge to a new forest and return of the final order- Two views from opposite ends of the rotting log of a large northern red oak that was killed by bark-feeding beaver and fell in the first summer following its death in early spring. The tall tree in left foreground of first slide was a sugar maple. Much, probably most, of the tree regeneration was also of sugar maple though this was accompanied by that of white ash, northern red oak, bitternut hickory, hackberry, and black walnut. Sycamore, the dominant pioneering or colonizing tree species along stream channels and freshly scoured bottomlands, was not released or encouraged by loss of northern red oak. Shrub species along this stream bank and the sides (slopes) of a limestone bluff above included American bladdernut, flowering dogwood, lanceleaf buckthorn, pawpaw, eastern redbud, and shadbush. These species were aligned along the slope from stream bank to top of bluff with bladdernut, lanceleaf buckthorn, and pawpaw typically restricted closer to water while shadbush grew highest up on the bluffs farthest from the stream. Flowering dogwood and eastern redbud generally grew all across this elevational gradient. Understorey herbs ranged from colonizing species like annual giant ragweed and mare's tail (and where did that seed come from?) and the hugh tap-rooted, perennial pokeweed to maidenhair and Christmas ferns.There were a few individuals of the two grass species, woodreed and beakgrain. Mosses grew on rocks, logs, tree trunks, etc. Above Modoc Creek, Ottawa County, Oklahoma. July,1997.

271. The young beneficiary (an immediate heir after assination)- A small pole of white ash was the largest tree above sapling size closest to the beaver-killed (and soon crashed) northern red oak described above on the east slope of a bluff above Modoc Creek that supported a climax sugar maple-white ash forest. Sugar maple (sole climax dominant), northern red oak, bitternut hickory, hackberry, and black walnut were generally the most abundant tree species and, not surprising, these species had the most regeneration in the gap created by death of three large northern red oaks. Of these, sugar maple was followed by white ash in general abundance (cover, density, etc.) as the major species to benefit from death of large, mature (actually, overmature) northern red oak. Not only were did sugar maple followed by white ash furnish the largest individual trees and greatest canopy cover, they were also the species which had greatest apparent (quite obvious) reproduction. Ergo, a sugar maple-dominated forest in which white ash was the associate species. Above Modoc Creek, Ottawa County, Oklahoma. June, 1997.

Concluding observtion with regard to browsing by beaver- On this stream-bordering, bluff forest (an east slope form of mesic-limestone sugar maple forest) feeding by beaver had a major impact on existing forest composition and structure and on future forest development (revegetation). Beaver were a major biotic factor that at local scale were a major cause of denudation and subsequent changes in the forest plant community via secondary plant succession. Beaver preferentially browsed northern red oak while largely leaving sugar maple and white ash untouched. Defoliation of some of the largest and oldest northern red oak by complete girdling resulted in quick death of some of those individual trees that had greatest control of canopy (forest) cover. Elimination of forest trees with some of the largest canopies produced forest gaps and patch dynamics. The main tree species to benefit from death of northern red oak by beaver and openings in the forest canopy created by such death were sugar maple and white ash, the climax dominant and associate of this forest which was a subtype of the sugar maple cover type, SAF 27 (Eyre, 1980, p.30). Some northern red oak--typically smaller and younger trees--was unbrowsed and remained in this forest as an important (though not an associate) species. Northern red oak was most likely a subclimax tree species having some large, long-lived (and lucky!) trees that persisted into the climax forest vegetation dominated by sugar maple with white ash as its associate (at least, major associate) species. Selective browsing by beaver hastened dominance by and general increased cover of sugar maple and white ash. Thus it appeared that sugar maple-dominated forest in the greater oak-hickory forest of the Ozark Highlands were partly a result of zootic influence and not just an edapho-topographic climax. Conversely, there were other tracts of sugar maple forests along this same stream (including a sugar-maple-bitternut hickory subtype of SAF 27) which supported no beaver populations and in which northern red oak was a major constituent but not an associate species. While beaver ignored or even avoided sugar maple in these forests they readily consumed bark and harvested saplings and smaller branches of box elder, the other Acer species on this tract. Box elder was uncommon on the sugar maple-white ash forest on the east-facing bluff above Modoc Creek, but box elder was locally a major species on the adjacent bottomland forest (just across the channel of Modoc Creek) where it was readily and regularily fed on. In contrast to browsing selectivity by beaver on a subclimax species (and avoidance of climax tree species) in this forest, a representative tract of SAF 27, beaver had a preference for and browsed selectively on climax American elm and hackberry on a bottomland forest, a representative of cover type SAF 93. These two tracts (and representatives of two forest range types) were contiguous except for separation by the narrow stream channel of Modoc Creek. On the bottomland forest the climax tree species were decreasers under beaver bvrowsing whereas on the adjoining bluff forest (and a different forest cover type) the climax tree species were avoided while beaver exhibited browsing selectivity for an important subclimax tree species which was a decreaser species.. Apparently there has been little evaluation or even recorded observations on the affect of beaver browsing on the development and compostion of forest range communities. Two interesting and extremely detailed studies on impacts of beaver on North American forests were included as chapters 18 (Donkor) and 19 (Smith) in the reference, Plant Disturbance Ecology, edited by Johnson and Miyanishi (2007, ps. 579-671).

Organization note: a climax, bottomland, mixed hardwood that developed on the floodplain of Modoc Creek and that was conterminous with the sugar maple-northern red oak-white ash-pignut hickory-white ash mesic forest on an east slope along bluffs above this creek was treated separately under Southern and Central forests-II.

A thrid series of slides and captions presented a third tract of sugar maple-dominated forest. This was also a third variant of form of the sugar maple forest cover type (SAF 27) in the Ozark Highlands which is the more westerly extent of the oak-hickory forest in general, the Oak-Hickory Forest Region of Barun (1950, ps. 35, 162-191). In monoclimax theory (Clements, 1916) sugar maple-dominated forests are postclimax vegetation in the climax oak-hickory forest region. Viewed from the polyclimax or climax pattern theories proposed by Tansley (1926) and Whittaker (1953), respectively, sugar maple forests are edaphic, topographic, etc. climaxes within the general or more widespread climatic climax (oak-hickory forest). By any of these interpretations sugar maple forests are the potential natural vegetation (Kuchler, 1964). Just as sugar maple-dominated forests in the general oak-hickory forest are variants of this regional forest (sugar maple forests usually include an oak and/or hickory species as the associate) so too are there variants (variant forms) of the generic sugar maple forest. The first series of photographs and their captions in this sugar maple section of Oak-Hickory Forests-II was devoted to the sugar maple-bitternut hickory variant and the second series of photographs-captions dealt with the sugar maple-white ash variant which were subtype 4 and subtype 3, respectively, of the sugar maple forest cover type, SAF 27, (Eyre, 1980, p. 31). The current (third) series of slides dealt with a sugar maple-northern red oak variant. This subtype was not noted by the Society of American Foresters description (Eyre, 1980, p. 31). Nor was it described by Steyermark (1940) or Braun (1950). Likewise, the SAF description did not include the sugar maple-white oak subtype of Steyermark (1940) and Braun (1950). It was somewhat surprising that Steyermark (1940) did not identify and describe the sugar maple-northern red oak variant because northern red oak is more mesic in its general habitat requirement than bitternut hickory and, even more so, than white oak. In moisture regimes of these tree species habitats, sugar maple is closer (more mesic) to northern red oak than to white oak or bittrnut hickory. In fact, among the three tracts of sugar maple forests described herein (western edge of Springfield Plateau of Ozark Highlands) northern red oak was the most consistently represented associate species. Nelson (1987, p. 28) listed the first three dominant plants of mesic-limestone forest as (in order): northern red oak, sugar maple, and white oak. In the first two tracts of sugar maple-dominated forest described above northern red oak was of lesser importance than bitternut hickory and white ash, but northern red oak was present at greater cover, density, frequency, etc. in all three tracts than the two associate species (other two species, if northern red oak was also an associate species). The sugar maple-northern red oak was the least mesic and most exposed of the three forest tracts described herein. This third variant form os sugar maple-dominated forest was a north slope (and limestone) forest, but it was not near a stream and more upland than bottomland forest. It was about two miles (or less) from the second tract (east slope) of a sugar maple-dominated forest, and with white ash as overall associate species. This third example of a sugar maple-dominated forest, or in this case co-dominated by northern red oak (Quercus rubra= Q. borealis), was on a steep north slope less than 2 miles from the two sugar maple forest stands along bluffs of Modoc Creek. The forest community featured next was a dry-mesic limestone forest (Nelson, 2005). This one was also in Ottawa County, Oklahoma; July, 2007). In strictest sense sugar maple was sole and defining dominant because most regeneration-especially in deeper shade-was of Acer saccharum and not Quercus rubra even though the latter was reproducing on some better-lite microsites (sometimes by established maples). Other species represented by occasional mature (and large) trees included bitternut hickory (Carya cordiformis) and white ash (in roughly equal proportions) along with lesser cover of basswood, all of which are climax species on this cover type and site. At upper (and drier) habitats in this forest chinquapin oak had both mature trees and regeneration. This species was regarded as a local climax member of this forest. Black walnut survived in more open spaces where it probably established in some gap and, having captured its share of canopy, survived to adulthood. At other spots in this forest black walnut succumbed to competition (was shaded out) by sugar maple and northern red oak. Major understorey (sub-canopy) shrubs included both eastern redbud and flowering dogwood throughoutalong with American bladdernut and pawpaw these latter two of which were limited to lower, more mesic habitats. The lower ground shrub layer was comprised mostly of Virginia creeper (Parthenocissus quinquefolia), but like the common, well-distributed summer grape (Vitis aestivalis), this woody climber ascended into the canopy. Poison oak/poison ivy (Rhus toxicondendron) was present, but was less common than in drier forests. Dominant forbs were tick clovers (Desmodium nudiflorum and D. glutinosum). Forbs that were locally abundant (in a relative sense) included Jack-in-the pulpit, Bloodroot (Sanguinaria canadensis), and wild ginger (Asarum canadense). An uncommon (and uncommonly striking) forb was blue cardinal flower of giant lobelia (Lobelia siphilitica). The only graminoids were unidentified species of Carex (only non-fruiting specimens present). These photographs were taken at abut 1330 hours Central Standard Time to get maximum direct light, but even then there was too much shade (too many shadows) to be able to catch key features such as bark. Ergo: not many good shots. Will try "next year" (wait until next summer) using with light cloud cover to get diffuse light.

272. Looking in on more sugar maple in the Ozarks- Exterior view of a north slope, mesic sugar maple-northern red oak forest at western extreme of the Ozark Plateau, and of the Oak-Hickory Forest Region (Barun, 1950, ps. 35, 162-191), in which this island of the sugar maple cover type (SAF 27) is postclimax (Clementsian monoclimax theory) or an edaphic or topographic climax (Tansley polyclimax and Whittaker climax pattern theories). The small pole-size trees were sugar maple. The tree species higher upslope (behind maples) included three oaks: chinquapin (chinkapin), northern red, and black oak with the latter more abundant on the drier, higher part of slope. Shrub species included American bladdernut (the most abundant shrub in understorey shown in this photograph), pawpaw (second most common shrub here), flowering dogwood, and eastern redbud. The latter two shrub species were more abundant in the tract overall just not in the sample of the forest plant community presented here. Virginia creeper formed the bulk of a lower shrub laye,r but along with the less common summer grape, this woody climber ascended into the canopy layer. Relatively little cover of herbaceous species, but the major forbs two species of ticktrefoil (Desmodium nudiflorum an D. glutinosum). Other forbs that were locally (and relatively) abundant in this forest vegetation were Jack-in-the-pulpit, mbloodroot (Sanguinaria canadensis), and wild ginger (Asarum canadense). An uncommmon (and uncommonly striking) forb wqas blue cardinal flower of giant lobeila (Lobelia siphilitica). The only graminoid observed by the author was an unidentifiable (pre-bloom) caric sedge (Carex sp.?). Ottawa County, Oklahoma. July (aestival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

273. On a steep north slope- The mesic north slope (with the fertility of a limetone-derived soil) furnished an ideal environment for this sugar maple-northern red oak forest in the Ozark Highlands.Northern red oak was the associate species overall, but local stands were dominated by white ash or bitternut hickory. In this first of these two photographs the four trees in foreground were sugar maple. Trees to left and downslope were white ash. In the second (vertical) photograph four sugar maples were shown at closer distance, but the more critical feature of this view was the understorey which included a diverse array od species including Virginia creeper, tick trefoils, a caric sedge, and seedlings to small saplings of sugar maple, bitternut hickory, and northern red oak. Regeneration of these tree species comprised most of the low and middle shrub layers in these two "photo-plots". Overall sugar maple, the tree with the high tolerance rating of Very Tolerant (Wenger, 1984, p.3), was the primarily reproducing tree species followed closely by bitternut hickory (eg. large seedling in front of and to left of the foremnost sugar maple). In this forest tract as a whole most reproduction was sugar maple and not northern red oak, white ash, or bitternut hickory. On some local sites (microsites) most of the seedlings and sapling stocking was of associate tree species. Examples of such stocking of younger age-size classes was shown in these and some of the following slides. There was very little herbaceous growth in this tract (including the "photo-samples" presented here) based on density and relative cover. Ottawa County, Oklahoma. July (aestival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

274. In the shade of their parents- A large sugar maple--by standards of this forest site in the Ozark Plateau--"looked down" on its progeny (and that of other sugar maples) as this climax dominant tree successfully reproduced in the mature soil and final stage of development of this forest sere. In this climax forest (potential natural vegetation) tree regenertation was primarily of the same species as the dominant trees (ie. same species of trees in the understorey layers as tree species in the canopy). Plant succession had stopped. The sere had reached its final "destination", climax stage. The final plant community for this natural vegetation--termination of plant succession on this forest site (this naturally occurring habitat of north slope, steep, limestone hillside in the Ozark Highlands)-- was complete until some disturbance (eg. fire, icestorm, disease or insect outbreak, logging, blowdown) reset vegetation on this sere back to a earlier state of vegetational development (plant succession). This was not, however, a "perfect match" of canopy and regeneration age-size classes in lower layers of this range vegetation. This was clearly a stand of sugar maple with all larger (adult) trees, including the one in foreground and those to its left in midground, as well as many larger seedlings and small saplings in foreground being maples. Also in foreground (and growing side-by-side with sugar maples seedlings) were seedlings of bitternut hickory, northern red oak, and chinquapin oak (these three hardwood species were discernable in lower right corner). It was uncertain as to which, if any, seedlings of these latter three species (all of approximate Intermediate tolerance) would grown to adulthood in shade of the mature sugar maples. It was a "good bet" that seedlings of the Very Tolerant sugar maple (Wenger, 1984, p. 3) would survive in the shade of their parent species. Two species of ticktrefoil were the dominant herbaceous plants (eg. a plant at prebloom stage to left of the center of sugar maple trunk in immediate foreground). Other herbaceous species were not identifiable in this photograph. Ottawa County, Oklahoma. July (aestival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

275. Layers and niches in climax sugar maple forest- Beneath a large, forked-trunk sugar maples saplings of bitternut hickory (left of the sugar maple) along with pawpaw (eg. shrub with light-colored, crooked trunk to the right and slightly behind the maple), eastern redbud (not discernable in this slide), flowering dogwood (not able to be detected in this photograph), and American bladdernut (shrubs of intermediate-height in lower right foreground) comprised much of the understorey of this climax sugar maple-northern red oak forest. There were actually up to three layers of woody plants in the understorey of this north (and very sttep) slope in the western perimeter of the Ozark Highlands. Pawpaw, flowering dogwood, and eastern redbud made up a high (the taller of two below-canopy) shrub layers while American bladdernut with seedlings and small saplings and ground-level Virginia creeper comprised the lower of these two shrub layers (those that did not extend to the canopy). Summer grape and some (lesser cover) poison oak/ivy and a few woody vines of Virginia creeper climbed into the canopy so as to extend the shrub layer into tops of tree crowns and form a third shrub layer in this forest plant community. Herbaceous species were sparse in this denser part of the understorey and limited mostly to two species of ticktrefoil. Ottawa County, Oklahoma. July (aestival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

276. "They have their exits and their entrances" (Shakespeare, As You Like It, II.7)- In a north slope, climax sugar maple-dominated e forest it was curtains for a black walnut, but the snag of the old tree was still serving a function in the forest ecosystem. A large summer grape used this snag of a long-dead black walnut as a Nature-provided trellis. The large bole behind and to left of the snag and that of the big bole behind and to right of snag were northern red oak each with a straight, standing log of clear lumber on the stump. Northern red oak produces some of the finest, clearest, straight-grained wood available for durable, attractive lumber that is commonly used in church furniture. This is a most appropriate use of this renewable natural resource especially in the Ozark Highlands section of the Bible Belt where hillfolk praise God and good timber. Of course there are heathens everywhere, but fortunately the owner of this timber land appreciated the aesthetic value afforded by the beauty of natural forest vegetation that blessed his property. This tract of climax, sugar maple-northern red oak forest was safe (at least for the foreseeable future) to perpetuate the species composition and structure of this potential natural vegetation, and re-develop that forest range vegetation should natural disturbances restart plant succession on this north slope, limestone sere. Most tree regeneration on this climax Ozark forest was sugar maple (including saplings in the background), but there was also some reproduction of bitternut hickory, white ash, and, of course, northern red oak. Herbaceous species in this vertical "photo-transect" were mostly ticktrefoils. Ottawa County, Oklahoma. July (aestival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

277. The players at play's end- Species composition of a climax sugar maple-dominated forest was furnished by this photograph of a north slope, mesic-limestone forest in the western Ozark Highlands. All visible trees were sugar maple except the tree with single (left center midground) and largest trunk (slkightly right of center background)both of which were northern red oak. In this tract of forest a high proportion of sugar maple had multiple trunks. Boles of the same tree were different sizes (hence of apparaently different ages). These were secondary shoots (basal trunk sprouts) which demonstrated the importance of vegetative (asexual) reproduction of this species (especially in this specific forest). The part of this forest's understorey was unusually sparse compared to most of the rest of its vegetation. Ticktrefoils and regeneration of sugar maple, northern red oak, and bitternut hickory comprised most of the lower strata. Ottawa County, Oklahoma. July (aestival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

278. Entrance of some new members of the cast- The next generation of sugar maple and northern red oak assured perpetuation of this climax, north slope, Ozark Hghlands forest range vegetation. Individuals of sugar maple, the dominant species of this forest (and the forest type it was representative of), included the larger tree at right foreground and the two-trunked, mature tree in center background as well as the three or four saplings with light-colored bark (left foreground back to background). Individuals of northern red oak, associate species, were the large pole-sized tree at left margin and larger tree in right-of-center background. Many of the larger seedlings and smaller saplings were bitternut hickory which were interspersed with those of sugar maple and northern red oak. A considerable number of flowering dogwood and eastern redbud were present, but these were in the background of this photographic view. A high percentage of the ground level layer of vegetation was Virginia creeper, which in this tract of forest range, tended to be more of the ground cover form than the vertical, tree climbing form. Ottawa County, Oklahoma. July (aestival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

279. The last set of players in the play of a sugar maple forest- Two photographs in the interior of a north slope, mesic-limestone forest in the western portion of the Ozark Plateau provided strong ecological evidence that northern red oak, the general associate tree species of this tract of forest, was subclimax to the climax sugar maple. Both of these slides featured a local stand of uneven-aged northern red oak, including some seedlings of this associate species. Most tree regeneration, however, was that of the Very Tolerant (Wenger, 1984, p. 3) sugar maple. While all the large, mature and pole-sized trees were northern red oak almost all of the seedlings and saplings (up to near pole-size) were sugar maple. Northern red oak, bitternut hickory, and white ash were represented in seedling and small sapling classes, but in such small (probably closer to trace) proportions that sugar maple was undeniably going to dominate almost all of the canopy of this forest in the not-too-distant future. These three species are generally relatively long-lived (on mesic, north slopes) and grow into large trees (comparatively speaking for trees in the Ozark Highlands) so that mature trees of these hardwoods persist into the climax sugar maple-dominated forest type. Any of these three tree species plus, on a more restricted basis, basswood are local associates of sugar maple. The latter has much slower growth rates on forest sites (even the most mesic and favorable of them) in the Ozark Plateau, but it exceeds all other trees in forest tolerance so as to regenerate in the forests it dominates, and at rates of reproduction that so "out-distance" its associate species that sugar maple is the last of the cast on the forest stage before curtains close. Sugar maple "steals the show" in the final act as it becomes the sole dominant of the climax forest. Shrubs in the understorey of this northern red oak stand included eastern redbud, flowering dogwood, and summer grape, the latter of which extended up through all layers of this climax forest. The most common, abundant herbaceous plants were Desmodium species. Ottawa County, Oklahoma. July (aestival aspect). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

One year (almost to the week) after photographing this third tract of sugar maple-northern red oak-dominated forest (one of old-growth status on a steep north slope with mesic, limestone-derived soil) the forest was clearcut by independent (gypo) loggers. Nothing was done on the land or with the forest vegetation except to remove logs (for pallats, including the sugar maple though a log of black walnut might have found other uses). Slash was not flat cut to enhance rotting but merely left after bucking up what logs were wanted. Some limbs of crowns protruded several feet in the air. Several larger logs were hollow and where left as they had fallen. There did appear to have been any fuel wood recovery although some portions of loaded logs might have eventually found their way into fire wood. Logging operations were completed in August and there was some plant growth prior to first killing frost in late October. Some cool-season species germinated (mostly annuals and biennials) or sprouted (mostly trees and shrubs; some perennial grasses) that autumn and/or early winter. In late May and early June (late spring), the beginning of first full growing season for warm-season species (including all tree and shrub species), a series of photographs was taken of the clearcut forest. Several of these slides were shared below. They presented the essence vegetation development, the process forest community recovery through secondary plant succession. This was an example of the action of "dynamic vegetation" as described by Henry Chandler Colwes and Frederic Edwards Clements. To reiterate specifics basic to forest range type and potential natural (climax) vegetation it was again noted that this forest was at western edge of the Springfield section of the Ozark Plateau, which was part of the Oak-Hickory Forest Region (Barun, 1950, ps. 35, 162-191).

280. Slicked off and starting over (or What hath man wrought?)- Clearcut old-growth sugar maple-northern red oak-dominated forest (with pignut or bitternut hickory, white ash, and basswood as associates). The north slope, limestone soil, mesic, mixed hardwood forest at old-growth state explored above was clearcut (just plain ole highgrading for pallet wood) in July and August of the year after the above photographs were taken. (God's guidance that your hillbilly professor got there a mere year before the white trash with their chain saws.) In these first two and a series of slides below the recovering (redeveloping) forest range vegetation was described and analyzed at season of late spring (late May to early June) of the first full-growing season (first spring) after logging (about nine months post-logging). There had been approximately six to eight weeks immediately after logging before the first killing frost and then one autumn and winter before this series of photographs was taken. These first two wide-angle "phototransects" gave overall views of the logged forest from a ridgeline (top of the north slope) vantage point. Finer details were presented in subsequent "photoquadrants" at closer camera range. Atop the ridgeline (hilltop) the species most visible on this new clearcut included mostly pioneering annual composites such as the horseweed or mare's tail, giant ragweed, and daisy or whitetop fleabane (Erigeron strigosus) as well as naturalized (from Eurasia) common sow thistle (Sonchus oleraceus) and prickly lettuce (Lactuca serriola= L. scariola). Japanese brome or soft chess (Bromus japonicus), a naturalized, cool-season, Eurasian annual grass, was also abundant. Japanesse brome was visible as several straw-colored, dead, small bunches (ie. it had completed its life cycle, and produced copius grain for next year's crop sunlight permitting). Another annual forb with considerable cover was common bedstraw (Gallium aparine). Numerous plants of poke or pokeweed (Phytolacca americana), a native perennial forb, were also present. Pokeweed was the first forb species to successfully invade the clearcut forest. Plants of pokeweed appeared within a few weeks following clearcutting and made normal-size growth in the remaining time of the first late summer-autumn following logging. There were a few saplings of redbud that survived the "woody massacre" (first of these two slides) and a fortunate-indeed small sapling of pignut or bitternut hickory (second photograph). Blackberry (Rubus sp.) and a seedling of smooth sumac (Rhus glabra) were other shrubs present on the ridgeline (second photograph). The shallower soil of and the more exposed (hotter, drier) land surface of the hilltop was more-or-less an old field environment, the classic cutover forest or bare ground point of denudation on this sere. This was in contrast to the greater area of land that was on the north slope that provided a "successional medium" which produced stump-sprouting and seedling establishment of the climax forest tree species along with release of native decreaser grasses in addition to the annual colonizing species that dominated the comparatively area of the hilltop or ridgeline. Only one of four oak stumps on the hilltop had coppiced (second slide). Ottawa County, Oklahoma. Late May (late spring). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

281. New life amid manmade devestation- Two views of the north slope sugar maple-northern red oak (dominants)-bitternut hickory-white ash-basswood (associates) mesic forest that had been clearcut about nine months prior to time of these two photographs. These shots were from the base of the north slope extending up to the hilltop (ridgeline) so as to provide a summary or overall view of the clearcut forest and recovering forest vegetation about six to eight weeks after the last killing frost. Species in the first of these two slides included sugar maple, both suckers (stump sprouts; coppice shoots) and seedlings of which were present only two months into the first full-growing season after logging. Stump-sprouting basswood or American linden (lkeft foreground). Coppicing might have begun in the last six to eight weeks right after logging and before the first killing autumn frost. Shrub species present included American hazlenut (Corylus americana) that was beside the basswood, greenbriar (Smilax tamnoides), and fox grape (Vitis vulpinia), sassafras (Sassafras albidum), and redbud, this lattter was the overall most abundant shrub on the clearcut. Herbaceous species ranged from native and introduced annuals like giant ragweed and common sow thistle to the tall-growing (left side) figwort or carpenter's square (Scrophularia marilandica). In second slide there many resprout shoots (from stump) of sugar maple plus a few of northern red oak. Redbud, most common shrub, was again conspicuous. In this second "phototransect" the herbaceous component was dominated by the native perennial woodland brome (eg. lower right and center foreground). Hairy wood brome is the dominant decreaser grass on north slope, mesic, limestone-soil forests, especially those in high successional status. The plants of woodland brome had been present prior to clearcutting as confirmed in a pre-logging photographic session by the author one year prior to clearcutting. Also present--as in left midground--were a few plants of tall fescue (Festuca arundinacea), an introduced, cool-season, perennial that is managed as an agronomic pasture grass and that has naturalized throughout this region. Tall fescue had undoubtedly established over the entire time period from end of logging until the present (a complete autumn through winter to late spring). There were a number of plants of pokeweed, a native perennial that was the first forb to successfully invade the clearcut (within weeks after logging). The most abundant forbs, however, were annual, colonizing, composite species: common sow thistle (naturalized Eeurasian weed) and mare's tail or horseweed (native) the standard pioneer of old-fields and cutover forests throughout this region.. Saplings in the background (both slides) were of northern red oak, bitternut hickory, and American elm along with redbud. Ottawa County, Oklahoma. First slide, early June (late spring); second slide, late May (late spring). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

282. All manner of new life- Two views midway up the steep north slope of a clearcut mesic, limestone-soil sugar maple-northern red oak forest in the western Ozark Plateau only two months into the first full-growing season after careless (as in "don't give a damn") logging. In the first of these two "photoplots" there was a copoicing American linden or basswood (right foreground) and two stumps of northern red oak neither of which produced sprouts (adventituous shoots or suckers). There was, however, a new seedling (not more than two-months-old) midway between these stumps. Most new phytomass (plant biomass or "weight") was of horseweed or mare's tail and common sow thistle, native and naturalized annual composites, respectively. There was also much common dandelion (Taraxacum officinale), another naturalized, annual, Eurasian composite. There was also fair representation of hairy wood brome, the dominant decreaser grass for this forest site. The small adult tree or large pole in center background was a northern red oak that somehow miraculously survived the onslaught. (This clearcutting was strictly harvest for saw logs, and for pallets only, such that many pre-log size trees were not cut. This was a logging not a forest-clearing operation. Nonetheless, most trees of any size with only a few exceptions like the small adult of northern red oak, were destroyed by careless logging. Most of this by felling saw timber-trees on top of the next crop and breaking them off or over.Loggers were careless, apathic, and hatchet-happy.) There were a number of seedlings of pignut or bittternut hickory surrounding the remaining young northern red oak. There was a single seedling of eastern red cedar (Juniperus virginiana) that was clearly visible in front of the young adult northern red oak. This young eastern red cedar had to have been present prior to clearcutting. The plant in lower left corner was leather-flower (Clematis pitcheri). The second "photoplot" featured a stump sprouted American elm amid much pokeberry, a perennial and the first forb to successfully invade the clearcut having grown to full size within two months after logging. There was also much hairy wood brome as in the previous "photoplot". Other common forbs were whitetop or daisy fleabane and mare's tail or horssweed, both native annuals. Another forb was golden Alexanders (Zizia aurea). Most of the remaining saplings in background of this second photograph were sugar maple. There were fewer northern red oak. Saplings of both species had been battered (including broken crowns) by careless logging; time will tell how they progress. Ottawa County, Oklahoma. Late May (late spring). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

283. Diversity of life on a clearcut- Two more "photoquadrants" roughly midway up a north slope of a limestone-derived soil from which a mesic sugar maple-northern red oak (two dominants)-basswood-bitternut hickory-white ash forest of old-growth state been clearcut about nine months previously. Present forest vegetation had developed from about six to eight weeks before first killing frost after logging through winter until late spring (three weeks before summer solstice). The first photograph was in a local spot where there were numerous stump sprouts and seedlings of American elm and much hairy wood brome, the dominant climax grass for this forest range site. Woodland bromegrass was a peak standing crop with maturing grain; in fact, some plants were senescing and entering dormancy. Redbud which was the dominant shrub across the clearcut was conspicuous in the foreground. Ther was also quite a bit of pokeweed or pokeberry, the first forb to successfully invade the clearcut, having established in the two months remaining of summer after logging. The most abundant forb in the first full-growing season after timber harvest was mare's tail or horseweed. There was also much giant ragweed. Both of these annual native composites are pioneering species. The yellow composite in background was the perennial composite, wingstem (Verbesina helianthoides). The second photograph centered on stump of a young northern red oak that had not commiced (stump sprouted). It was surrounded and being overwhelmed by fox grape and some tick-clover or tick-trefoil (Desmodium sp.). There was much of the dominant decreaser grass, hairy wood brome (almost everywhere). In addition there were several nice (and quite large) plants of woodland bluegrass (Poa slyvestris) which was readily distinguished by its cespitose habit, tall height, and dormant state as seen by flaxen, amber-colored straw. A specimen of woodland bluegrass was to upper right of the oak stump. Woodland bluegrass was the climax associate grass species for this range site. The tallest forb (eg. to left of stump) was a goldenrod (Solidago sp.) The white-flowered composite was daisy or whitetop fleabane. There were a lot of American elm seedlings to rear of this second "photoquadrant". Redbud, the major shrub, was also common. Ottawa County, Oklahoma. Early June (late spring). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

284. Up from the stumps- Coppicing of sugar maple two months into the first full growing season following high-grade clearcutting of a sugar maple-northern red oak dominated forest on a north slope, limestone-derived soil. Coppicing is the foresters' term for stump sprouting Two different sugar maple stumps in these two photographs. In the first slide there was also stump sprouting of American elm along with elm seedlings (ie. both asexual ans sexual reproduction of American elm). Also sprouting (from rhizomes) was the shrub, smooth sumac (Rhus glabra) which was very obvious at left of this first slide. Two annual native forbs were present in this first "photoplot": bedstraw and daisy or whitetop fleabane. This sugar maple stump had a profussion of stump sprouts. The second slide presented a second coppicing sugar maple stump with less prolific sprouting, but with an asexual shoot having mrvelously red-pigmented leaves (autumn-like leaves in late spring). In this second "photoplot" there were seedlings of pignut or bitternut hickory. Seedlings of the native pioneer species, mare's tail or horseweed (Conyza canadensis), were all around the sugar maple stump (eg. a big one at upper right of stump). A second annual composite--though a naturalized Eurasian one--was common sow thistle. Wild ginger (Asarum canadensis) which is a native, climax, perennial forb of mesic north and east slopes in the Ozarks was thriving with its new-found habitat of full light. A second, native, perennial forb--though a much more common one and one that thrives best in "full sun"-- was the mint, wild bergamot (Monarda fistulosa). In Silvics of North America Burns and Honkala (1990) reported the following findings: 1) sugar maple does have asexual reproduction via stump sprouting (ie. coppicing), 2) percentage of stumps that coppice decrease with increase in tree size, stand density, and years since cutting 3) average number of sprouts per stump decreased with years following cutting, 4) coppicing is more prolific in northern parts of the species range, and 5) coppicing is less in sugar maple than associated hardwoods in the southern part of its biological range. It appeared that, all factors considered, the degree (proportion or percentage) of coppicing bysugar maple and other dominant tree species on this clearcut was fortunately very high. It was underscored that there was abundant regeneration--axexual and/or sexual--of the climax dominants. Students of forest succession should note well this phenomenon. There certainly was an abundance of colonizing or pioneer plants (a number of plants of several weedy annuals, including the naturaalized aliens), but of native climax trees there was also abundant regeneration axexually (= regrowth of the same genetic individual) and sexually (seedlings of new genotypes). In regards the latter, on to the next slide, please. Ottawa County, Oklahoma. Late May (late spring). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

285. Up from seeds- Seedlings of sugar maple that were becoming established on a mesic, north slope forest dominated by sugar maple and northrn red oak that had progressed to old-growth state and was then clearcut. Logging amounted to nothing but high-grading by a bunch of local hicks (hillbilly was much too fine a term for this scalawags) and yet by the second month of the first full growing season after clearcutting seedlings of the climax dominant tree species had become established. it was possible that these sedlings had sprouted immediately following clearcutting in late summer of the previous year. However, there was only a period of six to eight weeks after clearcutting before the first killing frost that killed any remaining leaves of warm-season species (this included all woody species on this forest). Burns and Honkala (1990) reported that sugar maple seedlings are quite sensitive to mositure-stress near the soil surface because they have "a shallow, fibrous root system that lies between the litter-mineral soil interfaces". On this clearcut there had been much wetter than average moisture conditions in both O and A horizons of the soil due to heavy continuous rains in late summer-autumn immediately after logging through winter and spring of the following year. Such an abnormally wet edaphic environment favored establishment of seedling (and probably also stump sprouts) of sugar maple. Pignut hickory and American elm may not be as sensitive to dry soil habitats as the more mesic sugar maple, but abundant precipitation on rocky, well-drained soils could not have hurt these climax dominants either. With increaed light root systems of sugar maple grow down through the organic matter layer and into mineral soil which permits shoot growth and favors plant establishment (Burns and Honkala, 1990). Thess little sugar maples were "off and running". It was possible that the limb of a felled tree that lay to the west (upper portion of this photograph) of these seedling provided a key factor of shade from late afternoon sun that otherwise might have tried out litter and soil spelling doom to these delicate seedlings. This was an example of microclimate or a microsite habitat. Conspicuous other species in this "photoplot" included climbing wild buckwheat (Polygonum scandens), wake robbin (Trillium sessile), wild ginger, an unknown perennial composite, and a lichen on the downed limb. This was a combination of climax forbs (wake robbin and wild ginger) and weedy ones (climbing wild buckwheat). Reestablishment of existing trees (same genetic individuals) had also taken place by coppicing as shown in the two immediately preceding slides. Ottawa County, Oklahoma. Early June (late spring). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

286. Basswood from the base- Basswood or American linden coppicing from a stump following clearcut logging (exploitative highgrading) of an old-growth north slope forest in the Springfield section of the Ozark Plateau. Sugr maple and northern red oak were co-dominants of this mesic, limestone forest, but climax associate tree species included pignut or bitternut hickory, white ash, basswood, black walnut, and chinquapin (chinkapin) oak. Examples of coppicing (stum sprouting) in basswood or American linden were presented above at which juncture this form of asexual reproduction was explained in context of this species which is one of the more readily coppicing trees in Ozark forests. Other plant species shown in this "photoquadrant" included several forbs. Among these broadleaf herbs were the native annual composites, giant ragweed and horseweed or mare's tail, and the naturalized Eurasian biennial common or bull thistle (Cirsium vulgare) all three of which aree pioneer or colonizing species. Shrub species visible in this photograph included redbud, the most common ro abundanrt shrub, one of the wild grapes (Vitis sp.), and Virginia creeper. Grasses and grasslike plants were uncommon on the ground of this photograph or otherwise covered up by broadleaf species, forb and/or woody species. Oh, but you (like me) wanted grass? OK, next slide please. Ottawa County, Oklahoma. Early June (late spring). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

287 Released and growing - Hairy woodland or woodland brome (Bromus purgans) growing robustly on the north slope of a clearcut sugar maple-northern red oak-pignut or bitternut hickory-white ash-basswood mesic climax forest. Woodland brome was at its peak standing crop (soft-dough grain) stage in the first spring following highgrade logging. B. purgans is the only native perennial bromegrass in this region. This festucoid grass is restricted to the most favorable (especially most mesic; north slopes) habitats of hardwood forest ranges. Woodland brome is the dominant decreaser grass for this north slope, limestone soil forest site. The major (most important) climax grass benefitted from clercutting, at least initially. (Woodland brome was treated above in this chapter under the first of three sugar maple forests discussed.) Accompanying woodland brome in this "phytograph" were seedlings and/or stump sprouts of American elm and pignut or bitternut hickory, redbud (sprouts or already established seedlings released by loss of forest canopy) and Virginia creeper. Redbud was the most common shrub (had most foliar cover) on the clearcut. Some species of wild grape and poison oak/ivy were also common on this clearcut though not visible in this photograph. Students should note again in this "photoquadrant" (as in several of those above) that clearcutting this climax sugar maple-northern red oak forest created an initial forest range community that was a "blend" of plant species ranging from typical or standare colonizing species (including naturalized aliens) to sprouted clones of pre-existing climax trees as well as their seedlings (new genotypes) along with decreaser grasses. This forest was not going to have to regeneration from the bare ground stage of the sere. Ottawa County, Oklahoma. Early June (late spring). FRES No. 15 (Oak-Hickory Forest Ecopsystem), but a postclimax maple forest therein (Braun, 1950, ps. 168-169) so closest Kuchler unit was K-90 (Maple-Basswood Forest). SAF 27 (Sugar Maple). Either sugar maple postclimax in Oak-Hickory Series, 122.11, or Maple-Basswood Series, 122.15, of Northeastern Deciduous Forest in Brown et al.(1998, p. 37). Mesic-limestone forest (Nelson, 2005, ps. 122-125). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

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

288. Little patch- A small stand of prickly lettuce (Lactuca serriola= L. scariola) growing near a north-slope mesic forest that had been clearcut (highgraded) in western Ozark Plateau. This was a sugar maple-northern red oak-pignut or bitternut hickory-white ash-basswood climax community. Prickly lettuce is widely abundant in this general locality being found in overgrazed pastures, road ditches, cutover forest, parking lots, school yards, and you get the idea. Prickly lettruce is in the Liguliflorae, the subfamily of Compositae (tribe Cichorieae) whose members produce a milky (latex) sap. This sap does not deter consumption of the plant-- either by beast or man. This photographer enjoys a good bait of tender "spring greens" of which prickly lettuce is a "prime target species". Sow thistle (Sonchos oleraceus), another member of the Cichorieae, also eats well. Throw in lambsquarters (Chenopodum album), which is probably best pot herb overall, and poke salad (Phytolaca amricana), which should always be included, and any self-respecting hillbilly (eg. your author) is ready to devour a pot of "good eats". For pot herbs these forb species must be used earlier in the spring when shoots are relatively small, say less than a foot and a half tall (and then just take the first (top or upper) half foot or a little more).Not even deer would eat prickly lettuce at adult stages of maturity such as the plants shown in this slide. Cooking note: always pour out the first pot of water that such "weed greens" have been boiled in and then add fresh water and finish boiling to tenderness. Some of the Lactuca species are cool-season biennials with overwintering rosettes, but L. serriola is a cool-season annual (Diggs et al., 1999). L. serriola does have a basal leaf rosette, but this develops and persist through only one winter versus two winters in biennial Lactuca species. Most members of the Liguliflorae growing in forests of the Ozark Plateau are Eurasian species that have naturalized. This includes Lactuca and Sonchos species such as those featured in this section. oak-hickory. Species of these genera are certainly weeds--in both agronomic (cultural) and ecological (successional) meanings--that can be locally abundant, but they require fresh disturbances for establishment and, other than locally, do not form such extensive populations as to be serious pests. On clearcuts, such as the one presented here, Lactuca and Sonchus (treated shortly below) species are beneficial from standpoint of soil protection against erosion, recycling soil nutrients, providing green feed for grazing animals during the "off-season" (they are cool-season species), and even serving as less-than-choice bee plants. Ottawa County, Oklahoma. Early July.

289. Prickled shoots- Shoots of two plants of prickly lettuce growing near a clearcut north-slope mesic forest in western Ozark Plateau. Prickly lettuce is a rank-growing biennial. These shoots had the diameter of posthole digger handles and stood over ten foot tall. These plants were in peak-bloom stage with lower leaves already senescing and dying as this sporophytic generation was completing its phase of this species' life cycle. At this stage shoot are usually as hollow as a length of water pipe. Also at this stage of plant maturity, shoots of prickly lettuce have lost their palatability. For example, in this vicinity white-tailed deer feed on prickly lettuce shoots with the more apical portions being cropped. At the stage of "rank-maturity" seen here prickly lettuce is no longer taken by deers or other herbivores (ie. it has grown past its stages of vulnerability, at least to vertebrate grazers. Ottawa County, Oklahoma. Early July.

290. Prickly particulars- Details of stem and, especially, leaves of prickly lettuce. Organs seen here were on plants in the stand shown above that introduced this naturalized composite. These are full-grown and fully mature leaves. Leaves of Lactuca species such as prickly lettuce are extremely vriable in shape depending of taxonomic form (Steyermark, 1963, p. 1638). Ottawa County, Oklahoma. Early July.

11949.

291. How it grew back- New shoots growing from leaf axils of prickly lettuce in response to removal of apical meristem by grazing white-tailed deer. These axillary branches will go on to become sexual shoots (ie. produce a multi-headed flower cluster). This plant was in the same location as that of other slides in this section (near a clearcut mesic mixed forest in the Ozark Plateau). Deer frequently consume upper parts of prickly lettuce shoots. The author has yet to see one such grazed prickly lettuce that did not regrow and produce a plentiful crop of achenes, the dry fruit of composites. Speaking of which kindly advance the carousel. Ottawa County, Oklahoma. Early July.

11951.

292. Sexy lettuce- Sexual shoot of annual prickly lettuce with unopened heads, blooming heads, and heads of achenes all on the same cluster of sexual units. Upper portion of a floral assembly (= all heads of total flowering shoot) on one plant of prickly lettuce. Ottawa County, Oklahoma. Early July.

11954.

293. Impressive in numbers- Three tiny heads of prickly lettuce on one limb of the floral assembly shown immediately above. Any one of these heads or capitula may not strike the typical, citfied field-walker as being all that impressive, but such neophytes should observe just how many of these little heads are on one plant. To complete this mental calculation one should bear in mind just how many seeds (inside of achenes) are produced by each smallish capitula. For that part of the exercise please press the remote Forward button... Ottawa County, Oklahoma. Early July.

294. Loaded and ready to fire- Mature achenes (the fruit of composites) on the former heads or capitulae of prickly lettuce growing near a clearcut forest in the Ozark Plateau. There is one seed per achene (think sunflower "seeds"), and at least ten achenes per head. (Well, if you don't beleive me count them yourself!) This author has counted from 80 to over 200 heads per prickly lettuce plant. (If you think this is a Ozark talltale push the remote Reverse button and count the number of achene clusters, ripened heads, on just the upper portion of one flowering shoot.Damn fureners, anyway!) Ottawa County, Oklahoma. Early July.

Now for a regrouping or composite shot of the "business end" of prickly lettuce:

295. Inflorescence of a naturalized import- Blooming heads and unopened floral buds (first slide) and clusters of achenes (second slide) of prickly lettuce (Lactuca serriola= L. scariola). These were on plants on an oak-hickory-tallgrass savanna about three/fourths of mile (as the crow flies) from the clearcut sugar maple-northern red oak-pignut hickory-white ash-basswood forest shown in the preceding photographs. One or two branches of one entire inflorescence (flower cluster) on a prickly lettuce were displayed. This is one of three Lactuca species known generically as "wild lettuce" that became well-established in the Ozark Highlands. All these species of "wild lettuce" do best in newly and frequently disturbed ground where they commonly thrive. In so doing they provide both forage from their foliage and small concentrates from their tiny achenes. The "new ground" of the clearcut forest, especially on the ridgetop, was ideal habitat for this pioneering annual of the Liguliflorae, the subfamily of Compositae whose members secrete a "milky" latex in their sap in contrast to those of subfamily Tubuliflorae. Ottawa County, Oklahoma. First slide, mid-July and flowring stage; second slide, early July fruit-ripe stage. Note that a week's difference was irrelevant as some branches of one inflorescence was more advanced phenologically than adjacent or closely neighboring branches, all on the same plant.

296. Flowers to fruits in detail- Flowering heads and just emerging achenes (first slide) and the intricate pattern of ripe achenes just as they were being shed (second slide) in prickly lettuce. Lactuca as well as Sonchos (see immediately below) species are in Liguliflorae, the lataex-secreting subfamily of Compositae. The best known species in this subfamily is common dandelion (Taraxacum officanale). It was featured in the Alpine chapter of Range Types. Another similar species is goat's beard (Tragopogon dubius) that was treated in the Palouse Prairie chapter. Compare these species to note the extremely similar disks and plummed achenes of these composites, all of which are alien to North America and yet all very much naturalized in their new "Home on the Range". Ottawa County, Oklahoma. Early July.

297. Another naturalized commoner- Common sow thistle (Sonchos oleraceus) on an oak-hickory-tallgrass savanna about three-fourths of a mile from this clearcut. These plants (two plants in first slide; upper shoot with inflorescence of right plant in sescond slide) were on the edge of the same black oak-dominated forest in photographs of prickly lettuce immediately above. It is common for several species of Lactuca and Sonchos to grow side-by-side (or nearly so) in the Ozark Highlands, especially on harsher habitats such as those of shallow, rocky soils. The ridgetop of this clearcut forest was just such an environment (in contrast to the deeper and moister soil of the shaded north slope below this high point of the forest. Like prickly lettuce (and other "wild lettuces"), sow thistles (there are at least two species in the Ozarks Region) are commonly eaten by white-tailed deer as well as cattle, sheep, goats, and eave horses on occasion. Ottawa County, Oklahoma. Mid-July.

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

299. Alien pioneer- Flannel mullein (Verbascum thapsus) is a Eurasian, biennial range forb of the Scrophulariaceae, the snapdragon family, that has completely adapted (=naturalized) to disturbed habitats in North America. Flannel mullein is one of the more widely distributed weeds on its adopted continent. That is, weed by definition (ie. an alien not grown under domestication, at least not purposely) but not weed by function or in an ecological sense. By functioning as a pioneer or colonizing species of drstically disturbed land this cool-season biennial--albeit it an invasive--is often a highly beneficial range and forest plant. Its broad-leafed, overwintering, basal rosettes covering more-or-less bare land serve as protection against accelerated soil erosion. Upon decay, rotting organic matter from flannel mullein serves in the role Clements describead as reaction (Wever and Clements, 1938, ps.234-247 passim) or what others have since labeled facilitation.(Connell and Slatyer, 1977) Plants of flannel mullein presented here were growing on the upper slope of a two-year old barrow ditch along a county road. These members of an exotic species were preparing the way for range plant species of higher succcessional order. In immediate time scale they were retaining some soil from being washed away from its natural resting place and covering a freshly paved road surface. Such plants were even more beneficial on the forest clearcut described here. As a secondary benefit this naturalized range forb and weed of tilled soil is a wildflower that always struck this author is quite attractive. Flannel mullein is afterall a tall "snapdragon of sorts", and one that plants itself. Furthermore, when drought comes early this biennial weed will still flower and add some beauty to a "burnt-up" country. Ottawa County, Oklahoma. Mid-July, full-bloom stage.

300. Flower stalk and flower- Part of the often much-branched flowering shoot of flannel mullein (first slide) and some individual flowers (second slide) of this anturalized Eurasian member of the snapdragon or figwort family. If a nurseryman sold a clueless city slicker the seed of this plant the urbanite would, quite rightly, view it as a garden flower. Yes, a large, rank-growing one, but a desirable (at least, acceptable) gardenflower nonetheless. If a farmer somehow missed a plant of flannel mullein until it reached this stage he would almost assuredly hoe it out cussing it as he did so. Now who has the more impartial perspective. There is often a common bond of sorts between the rangeman and/or grazier and the "little ole lady in tennis shoes" who enjoys "pretty wildflowers". Certainly there is also a bond between farmer and stockman. They may be one and the same. There has also been an antagonism between the herdsman and sod-busting plowman, one going back Cain and Abel. Ottawa County, Oklahoma. Mid-July, full-bloom stage.

Savanna of Oak-Hickory Forest and Tallgrass Prairie

The tallgrass prairie-hardwood forest savanna or transition was given a chapter of its own in this publication entitled, Tallgrass Savvanna, plus a section on Prairie Peninsulac in the chapter, Tallgrass Prairie (Interior). One photograph and caption was included at this point to highlight locations for these important deciduous forest and eastern prairie grasslands.

301. Oak-hickory forest and savanna- This is the widespread transition zone or intermediate community type between actual oak-hickory tallgrass prairie understory savanna and the western edge of the oak-hickory forest proper.Post oak and black hickory dominate the tree layer while black and red oak are associated species. Understory is primarily nodding or Canada wildrye and purpletop (Tridens flavus). Virginia creeper covers much of the understory and climbs into tree tops. Blackberry, buckbrush, Mayapple, and black-eyed susan dominate various microsites depending on cover of tree canopy. Chert savanna-chert forest composite type with fire determining which form prevails. Ottawa County, Oklahoma.June, late vernal aspect. FRES No. 15 (Oak-Hickory Ecosystem). K-73 (Mosaic of Bluestem Prairie [K-66] and Oak-Hickory Forest [K-91]). SAF 52 (White Oak-Black Oak-Northern Red Oak); SRM 801 (Dry Savanna). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

Dominant, Associate, Widespread, and Just Plain Interesting Plants

of the Central and Southern Forests

302. Composite view of Oak-Hickory Forest- Sample of species richness and biological diverstiy of eastern deciduous forest in central and southern regions. Tree on left is pignut or bitternut hickory, the forked trunks with burl on left trunk is black cherry, a young pignut hickory to its immediate right, and the two larger white-barked trunks in background visible between the hickory and cherry are post oaks. The barely detectable short shrub dominating the understory is buckbrush which in this winter season has shed its fruit. Hibernal aspect . Ottawa County, Oklahoma. Hibernal aspect, Late December. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 40 (Post Oak-Blackjack Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

303. Cherries in the woods- Leaves and ripening fruit of black cherry (Prunus serotina). The fruit type of Prunus species is pome. Black cherry is the source of commercial cherry wood, one of the most beautiful of all North Americn hardwoods. Black cherry is not a domiinant or even associate tree species in climax oak-hickory forests, but in absence of fire it increases quickly and can overwhelm the climax dominant and associate oak and hickory species. Ottawa County, Oklahoma. Late June.

304. Fruit of buckbrush or coralberry (Symphoricarpos orbiculatus)- This fleshy fruit is widely held to be valuable for bobwhite quail, squirrels, and rabbits. It is perhaps the most dominant species of the lower shrub layer (taller shrubs like dogwood, persimmon, sassafras, and redbud being local dominants of the taller shrub layer). Ottawa County, Oklahoma. Early December (and fruits may be shed within the month or persist on twigs in dried form until spring depending on the year).

305. Flowers of flowering dogwood (Cornus florida)- This is probably the most widespread understorey shrub or small tree in the oak-hickory forest forms and, therefore, is typically the dominant of the lower woody plant canopy. In the spring flowering dogwood (often accompanied by redbud) turns the dark, drab-colored, bare woods into a colorful botanical banner announcing start of another growing season. This species with it's delightful spring and, as shown immediately below, fall display is the State Tree of Missouri. Ozark Plateau. Newton County, Missouri. April.

306. Leaves and mature fruit of flowering dogwood in fall coloration- It is an age-old argument as to whether the spring or autumn is the more colorful in the deciduous forests of eastern North America. The argument cannot be resolved by admiring the flowering dogwood which contributes to the beauty of the timberlands in both seasons. The fruit is a major food source for wild turkey (Meleagris gallopavo) and bobwhite quail (Colinus virginanus), especially in the south as in the Ozark Mountains, while the twigs are important browse for white-tailed deer (Odocoileus virginianus). Indians had several uses for this shrub (eg. dyes, herbal remedies).The hard, tough, tight-grained wood has many non-construction uses. Newton County, Missouri. October.

307. Shadbush or eastern or downy serviceberry (Amelanchier arborea)- Another shrub of the oak-hickory-- and general eastern deciduous-- forest is downy serviceberry. The common name of shadbush comers from the hill folk who noted that blooming of this species often coincided with the spring runs of shad (Alosa spp.). Downy serviceberry is often confused with flowering dogwood because the flowering periods of these two species often overlap and that of one species may precede or lag behind the other depending on conditions in any one spring. (Dogwood gets all the credit by the way.) Serviceberry is one of the many members of the rose family which is the single most important family of range browse plants in North America. Amelanchier species are in the Pomoideae, the pome fruits subfamily, of Rosaceae. Shadbush undoubtedly provides some browse and the fruits are eaten by birds and furbearers, but it is not common enough to be a major feed plant. The bluffs of Modoc Creek, Ottawa County, Oklahoma. April.

308. Downy serviceberry in full bloom- In the opinion of your author when this species is flowering it is often the most striking and showy shrub in the oak-hickory forest. Much of the enjoyment of the spring blooming in the hills is due to serviceberry and not flowering dogwood, but a human population of predominantly city dudes does not know the difference and just calls everything other than redbud a "dogwood". Ottawa County, Oklahoma. April.

309. Flowering shoot tip of downy serviceberry- The inflorescence and two newly emergent leaves of shadbush or eastern serviceberry. Ottawa County, Oklahoma. April.

310. Goatsbeard or Goat's Beard (Aruncus dioicus var. pubescens= A. pubescens)- Goatsbeard is another striking member of the Rosaceae. It is far from common in the oak-hickory forest, but when it is encountered it causes the most hill-hardened hillbilly to pause and "take a gander". Aruncus literally means "goat's beard" and according to various manuals this name can be traced to Pliny and later applications to Eurasian species of this genus. This North American shrub has a range from the extreme northeastern part of the continent south to Alabama and west to Oklahoma. Springfield Plateau section of Ozark Plateau; base of limestone hill in Newton County, Missouri. June.

311. Inflorescence of Goat's Beard- Goats's Beard is a dioecious, but the flower clusters are similar for both sexes. Newton County, Missouri. June.

312. Catkins (male flowers) and mature fall fruit of common hazelnut (Corylus americana)- Another (but usually sparsely populated) shrub of the oak-hickory forest is the hazelnut. This nut was highly prized by the early American backwoodsmen, the proper historic name for the first wave of frontiersmen who moved westward beyond the Fall Line into the dark, forbidding, and mysterious deciduous forest of eastern North America. For millinia prior to the first footprint of man (God only knows what shade of skin or species of Homo it was) hazelnut was feed for wildlife ranging from wild turkey to furbearer to native ruminant. Male flowers of this monecious species are borne in catkins that are some of the first flowers in the late winter or early spring in the oak-hickory forest. The specimen shown here is from an individual of the variety C. americana var. indehiscens (formerly interpreted as C. cornuta) delineated as a separate taxon by the united fruit bracts that make the fruit to appear unopened. Fruits appear singularly up to as many as four in a cluster like the one pictured here. The family to which this little nutbearer belongs is another source of controversy. Some treat it as a member of the birch family (Betulaceae) with separate tribes of Betuleae and Coryleae while other workers place it in it's own hazel family (Corylaceae). On second terrace of Modoc Creek in Ozark Plateau. Ottawa County, Oklahoma. Catkins, February; fruit, October.

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

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

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

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

317. Wild hydrangaea (Hydrangea arborescens)- This (and the fine specimens shown in the next set of two slides) grew on a moist north slope by a wet-weather srping. Wild hydrangea is not only one the most showy of the understorey woody species in the eastern deiduous forests, but also one of the most mesic shrubs of this formation with occurrence of this member of the Saxifragaceae limited to moist or even wet habitats. Newton County, Missouri. June

318. More beauty- Details of wild hydrangea inflorescence. The larger, conspicuous, three to four-petaled flowers on the periphrey of the flower cluster are sterile (without sex organs). These appear to serve as attractants to insect pollinators. Newton County, Missouri. June

319. Lowbush huckleberry (Vaccinium vacillans)- Ericaceae Vaccinioideae Flag Springs State Park, McDonald County, Missouri. June.

320. Highbush huckleberry, squaw huckleberry, or deer berry (Vaccinium stamineum). Ottawa County, Oklahoma. June.

321. Poverty oatgrass (Danthonia spicata)- This cool-season bunchgrass often forms rather extensive carpet-like stands made up of these green clumps in the understorey of oak-hickory forests. This occurs mostly on the drier habitats like the steeper of south slopes and the more shallow, stoney, and acidic soils where the larger grasses (eg. the bluestems, panicgrasses, beakgrain, and woodreed grass), forbs, and shrubs like blackberry (Rubus spp.) are absent or stunted. Poverty oatgrass is also more common on "edges" between deciduous forest and tallgrass prairie where the more drought- tolerant trees predominate. For example, in the habitat shown here oak leaves in the background belonged to blackjack (Quercus marilandica). Ottawa County, Oklahoma. June.

322. Poverty oatgrass at more advanced maturity- Several cespitose plants of poverty oatgrass on the same precise microsite as shown in the immediately preceding slide (first of these two slides) and of a single plant of poverty oatgrass (second slide) just a short distance removed from those in the first photograph. These shoots were photographed in another year and at more advanced maturity (phenological age) from those presented immediately above. The rocky (chert limestone) nature of the microhabitat of these examples was representative of the environment on which plants of this species excell. It was shown above, however, that big bluestem will sometimes grow beside plants of poverty oatgrass such that these two species are range "pals" on oak-hickory/tallgrass savannas. Ottawa County, Oklahoma. Early July; grain at maturity just prior to its shatter stage.

323. Rock-poor or rock-rich?- Spikelets of poverty oatgrass at grain-shatter phenological stage. Number of florets per spikelet ranges from three to seven, rarely up to nine in poverty oatgrass (Steyermark, 1963, p. 142; Diggs et al., 1999, p. 1260). The rock on which these spikelets were laid was common chert, the erosion-resistant remains of Mississippi limestone that formed much of the parent material of the Ozark (Springfield) Plateau. Ottawa County, Oklahoma. Late May. (Maturity of caryopses varies considerably in poverty oatgrass even within a local population. These spikelets were at grain-shatter stageat this time whereas othr spikelets in the same local population had not reached grain-ripe/grain-shatter stage six weeks later.)

324. Not impoverished- Local dense stand (population) of poverty oatgrass in the western Ozark (Springfield) Plateau. This example of a single-species stand (or colony) at peak standing crop was a particularily--though not at all unusually-productive sward growing on a extremely rocky yet deep soil on the perimeter of a mixed oak-black or Texas hickory second-growth forest. Agrostologists cannot make up their minds (and that is not unusual either) as to proper taxonomic placement of Danthonia species. This genus has been placed in its own tribe, Danthonieae, within either subfamily Arundinoideae (Gould, 1975, ps. 56-58; Gould and Shaw, 1983, ps. 329-331) or Eragrostoideae (=Chorideae) (Diggs et al, 1999, 1259-1260). Ottawa County, Oklahoma. Late May; grain at mid-maturity stage.

325. Panicle of stout woodreed or woodreed grass- This festucoid grass (tribe Aveneae) is a major provider of nutritious forage in the understories of carefully managed (ie.conservatively stocked) oak-hickory forests. It is a major species only on a local basis because years of overgrazing have greatly reduced it’s abundance and even occurrence. Woodreed is adapted to diverse sites within the oak-hickory forest range type thriving on both bottomland and stream bank habitats as well as shallow upland soils. Bank of Modoc Creek, Ottawa County, Oklahoma, September.

326. Stout woodreed is a perennial grass that has an obvious prolonged growing season and pronounced allocation of resources. In the Ozark Plateau flowering occurs in late summer or early fall (August to October), but new basal shoots appear soon after completion of the annual life cycle (death) of current-season shoots. Thus there are usually green shoots in some phenological stage yearlong, but the growth and development of these shoots is extremely slow. In this photograph young shoots (tillers: intravaginated shoots) of two to four inches in length are growing among last year’s tillers which reached their maximum mature height of three to four feet. Shallow savanna range site (a shallow upland soil overlying a solid layer of chert). Ottawa County, Oklahoma, December.

327. Close-up of the woodreed tillers of the previous slide- Allocation of resources so as to maintain some live tillers yearlong is likely an adaptation to the dark shade existing from spring through autum due to  dense canopy of the forest during this frost-free period.(mid-April to mid-October in the Ozarks). Woodreed requires a full year under these conditions of limited light to fix enough carbon to complete the annual life cycle and store root reserves to initiate next year’s growth. Any plant living in habitats with such a low light intensity and a protracted regime of photosynthesis and growth will obviously be quite vulnerable to defoliation and require careful grazing management.     Not only is conservative stocking important for regeneration of shrubs and lumber trees but it is also essential to prevent overgrazing of grass species growing in the dimlite, stressful understory. Given the shallowness of soils on some sites in deciduous forest range in conjunction with the commonness of periodic summer drought it is likely that grasses like stout woodreed and beakgrain suffer from the stress of water shortage. This would be “true in spades” for these summer-flowering festucoid grasses which must compete with the more heat- and drought- tolerant dominant panicoid grasses like big bluestem and beaked panicgrass (Panicum anceps). Combinations of dense shade, shallow soils, drought, competition, etc. make for a harsh environment even in humid precipitation zones. Excerbation of this condition with overgrazing or improper season of use is one reason why deciduous forest ranges have become deteriorated. Furthermore (and contrary to popular misconception) these deciduous woods ranges were the last open range (the true meaning of open to the public, a grazing commons, and not just absence of fences) in the United States. The actual Public Domain ranges of the Intermountain West were closed and came under some scientific regulation by the Grazing Service (later, Bureau of Land Management) with attempted enforcement of the Taylor Grazing Act in the 1930s. By contrast, state laws officially closing the range (passage of mandatory “fence-in” laws) in Missouri, Arkansas, Oklahoma, and Texas were not passed until the 1950s. Free-ranging, acorn-eating, razorback, rooter hogs could be found sleeping and rooting on county courthouse squares through the first half of the 1950s throughout portions of the Ozark, Boston, Quachita Mountains in Oklahoma, Missouri, and Arkansas and in the Big Thicket of Texas. The account for the latter can be found in Sitton (1995, ps. 194-273 passim). Rowley (1985, ps.239-241) noted that control of livestock grazing on National Forest “… in the South [Region 8] remained an unfinished business in the early 1960s…”.and that in “… the 1950s range continued to be the southern region’s ‘ bastard child’. Closing the southern ranges had always been a battle in intself (King, 1982) because (and this is another scientific, economic fact that is contrary to popular, and incorrect, opinion) livestock raising was far more important economically and culturally than it was credited with. This was true even for the antebellum South when the planter class was at its zenith (McDonald and McWhiney, 1975). It is no wonder why professionally trained foresters still harbor strong prejudices against even proper (sustainable) livestock grazinig in the eastern deciduous forests.

Location Note: more detailed coverage of woodreed grass was offered in Range Types of North America in the chapter, Central and Southern Forests-II.

328. Beakgrain ( Diarrhena americana)- The understory of oak-hickory forest and its ecotone or leading edge with tallgrass prairie supports species of grasses which are not commonly discussed in standard range plant references (even though they often produce appreciable amounts of palatable, nutritious forage). Beakgrain is one of these species which was included here as an example of a situation commonly confronting the range practitioner (especially one just starting his career). A relatively obscure species which is given but short-shrift in handbooks or range guides is nonetheless of local or occasional importance. Rangemen must learn these important “locals” much like they must learn the locally powerful politicians.

329. Spikelets in a panicle of beakgrain- This member of the Festucoideae is in its own tribe (Diarrheneae). Floodplain of Lost Creek, Ottawa County, Oklahoma, July.

330. Nimblewill (Muhlenbergia schreberi)- This eragrostoid grass (Eragrosteae tribe) is another widely distributed grass in understories of oak-hickory forest ranges. Nimblewill grows on a diversity of habitats ranging from fairly dense, mesic woods to open prairies on shallow soils and as a component of stremside vegetation to a common species on abandoned fields (= go-back land) or even old city lots and schoolyards. It is generally accepted that nimblewill is probably the best-adapted of any muhly in the southern oak-hickory forests and savannahs, including adaptation to disturbances like overgrazing and trampling. Nonetheless, nimblewill was interpreted by Tyrl et al., 2002, p. 109) as "... characteristic of the mid to late stages of plant succession". Although nimblewill is neither rhizomatous nor stoloniferous it frequently grows as localized colonies especially in forest clearings, lanes, old barnyards, etc. This habit is due to adventituous rooting at nodes of the low, decumbent tillers from which short-branched, largely unspread panicles arise. Nimblewill has a unique feature (unique among North American Muhlenbergia species) in that morphology or growth form differs greatly from the early season, short plants with broad leaf blades to the late-season form with ascending to even sweeping shoots. Nimblewill specimens presented here were growing in shade of post and blackjack oaks at extreme western edge of Ozark Plateau where relatively large-sized tallgrass prairies and oak-hickory forests developed in a complex vegetational mosaic. This patchwork of range vegetation was part of the southern extension of the prairie peninsula. Ottawa County, Oklahoma. December: hibernal aspect, dormancy.

331. Shoot tips of nimblewill- Photograph of shoot apices of nimblewill with remnants of mature inflorescences (seed shatter stage) many of which were either reduced to their bare floral axisis or still partially to largely enclosed in the boot,. These are common or even characteristic late-season features of the spike-like panicles of this species. Ottawa County, Oklahoma. December: hibernal aspect, dormancy.

332. Local stand of nimblewill- This small colony or population of nimblewill shoots (along with a few shoots of Carex molesta) was growing along the edge of an oak-hickory Ozark forest. This provided an example of typical habit and morphology of this eragrostoid grass. Nimblewill is not a heavy or high herbage-producer, but it is one of the best adapted native grasses to stress caused to a combination of defoliation and shade. Ottawa County, Oklahoma. July; immediate pre-bloom stage of phenology.

333. Shooting in the shade- - Shoot of nimblewill growing in its common and, apparently, "prime habitat" in the Ozark Plateau. In shade of post and blackjack oaks, black cherry, and hackberry and amid blackberry canes this nimbleweill plant had produced a robust (for this species) shoot. Shoots of nimblewill are, as revealed here, distinctive among grasses. Ottawa County, Oklahoma. July; immediate pre-bloom stage of phenology.

334. Shooting a panicle- Sexual shoot of nimblewill with largely exerted panicle. Typically the panicle of nimblewill remains partly enveloped in the boot (encased in the leaf that subtends the inflorescence) so that branches off of the central floral axis appear appressed and ascending so as to resemble a semi-condensed panicle. Traditionally the fruit of Muhlenbergia species has been interpreted as an achene rather than a caryopsis. Spikelets in this panicle were pre-anthesis. Ottawa County, Oklahoma. July; early summer.

Most of the herbaceous understorey species in the various eastern deciduous forest cover types are forbs. Forb is a term used by foresters, rangemen, and wildlifers in reference to any herbs (=. herbaceous plants) that are not grasses or grasslike plants. In other words, forb refers to all soft-stemed dicots and to any monocots that have conspicuous petals. Forb is not a precise botanical term , but rather one used by professionals in the natural resource management fields. Range and forest plants are either 1) woody or 2) herbaceous. The woody plants are either trees or shrubs, the distinction between which is not always obvious. The herbaceous species (herbs) are either grasses, grass-like plants, or forbs. (Together these are generally the five "kinds"-- as in categories or groups not species-- of vascular range and forest plants.) Some of the more common and conspicuous forbs of the eastern deciduous forest communities were included immediately below. All of these were growing in the Springfield Plateau section of the general Ozark Plateau or Ozark Mountains.

335. Jack-in-the-pulpit or Indian turnip (Arisaema atrorubens)- This unique and picturesque forest forb is in the arum family. It has one of the more expansive species ranges of any understorey species in the eastern deciduous forest formation. Jack-in-the-pulpit is not always plentiful where it grows, but it does indicate a general lack of abnormal disturbance and botanically diverse forest community when present. This is a very interesting species. Indians made much use of it. It does cause some animal poisoning. Of most interest perhaps are the basic botanical aspects of this monocotyledonous wild flower. The inflorescence is a spadix in which "Preacher Jack" is the fleshy spike of imbeded flowers "who" is inside of an enveloping bract known as a spath. Individual plants are either monoecious or dioecious and, reportedly, individual plants can change the sex of flower from one year to the next. Jack-in-the-pulpit also reproduces vegetatively from creeping rootstocks. Plants perform the most "kinky sex". This is but one of countless interesting things about Range Management and Forestry. In bottomland forest of Modoc Creek, Ottawa County, Oklahoma. April.

336 Green dragon (A. dracontium)- This Arisaema species is another woodland forb which is also enjoyed as a wild flower like it's "cousin" jack-in-the-pulpit. These two species sometimes occur together in rich, moist forest soils. Green dragon spreads by creeping rootstocks, a form of asexual reproduction that is apparently quite efficient in the dim-lite forest floor of the deciduous forest. Arisaema species begin growth and flower early in the spring before the deciduous trees leaf-out and form their light-excluding canopy. First terrace of Modoc Creek, Ottawa County, Oklahoma. May.

337. Detail view of green dragon- Leaves and the spadix and spath of the "flower" of green dragon were presented here. Modoc Creek, Ottawa County, Oklahoma. May.

338. Deer's tongue, adder's tongue, or dog-tooth violet (Erythronium albidum)- This is a forest lily that emerges early in the spring through the deep layer of shed oak and hickory leaves on the forest floor. It then promptly flowers before the overstorey trees grow their annual canopy that affectively blocks most light from reaching the ground. Hill-tromping hillbillies enjoy this delightful forb as a harbinger of spring. Ottawa County, Oklahoma. March.

339. Trout lily (Erythronium americanum)- This even showier species is more restricted in its environments in the Ozark Plateau oak-hickory forest than is E. albidum. The trout lily featured here was growing on a bench on the north slope of limestone bluffs that supported a sugar maple-bitternut hickory (Acer saccaharum-Carya cordiformis) mesic-limestone forest (Oak-Hickory Forest- II). On bluffs above Modoc Creek, Ottawa County, Oklahoma.April.

340. Wake robin (Trillium sessile)- This is another member of the Liliaceae that emerges early in the spring when it can get enough light to complete it's annual cycle of life. Wake robbin grows best and to it's largest size on rich, moist soil. Ottawa County, Oklahoma. April.

341. Walking up on wake robbin- Two succcessively closer views of T. sessile in the western Ozark Highlands. These fine specimens were growing on the boggy soil of a green ash (Fraxinus pennsylvanica)-sycamore (Platanus occidentalis) wet bottomland forest (see Oak-Hickory Forest-II).

342. May apple or mandrake (Podophyllum peltatum)- May apple is probably the most widespread and, as it grows in sizeable colonies from extensive roostocks and sports large, spreading, uniquely shaped leaves, the most conspicuous forb of oak-hickory forests. The fruit is edible, but the foliage does not appear to be grazed by vertebrates. This mandrake should not be confused with the other species of that name that is native to the Mediterranean Region (the mandrake of the Holy Bible). This forb is often the locally dominant species of the upper herbaceous layer of the multi-layered deciduous forest. Ottawa County, Oklahoma. May.

343. Berry on the bottom- A ripe fruit of may apple still attached to the crotch or point of branching in this species was presented in the first of these three photographs. The second and third slides showed the fruit cut open with seeds remaining attached and then with seed cluster removed and set aside, respectively. May apple fruit is a berry. Though edible, this berry is pretty much tasteless (unless one is extremely hungry-- as in starving). A bona fide hillbilly would eat his specimen. The one who gave you these shots fulfilled this hill culture requirement. Ottawa County, Oklahoma. Early July.

344. Dutchman's Breeches (Dicentra cucullaria)- The range of this forest forb extends from Quebec westward to Oregon and Washington and south to Missouri and Oklahoma. As seen from this photograph it "does it's thing" early in the spring before the trees leaf out and exclude light from the forest floor. Even then, as also clear from this slide, light is limited by trunks and larger limbs. This species was discussed in Notes on Western Range Forbs (Dayton, 1960, p. 220-222) where it was noted that the bulbs are poisonous to cattle when they are pulled from the soil and eaten along with the shoot. Burrows and Tyrl (2001, ps.706-709) provided a recent synthesis. They noted that Dicentra species contain several isoquinoline alkaloids that affect the nervous system and cause trembling and stggering, but they added that the plants were so uncommon as pose no serious problem, and besides animals usually completely recovered. Your author noted that this is another application of the Cardinal Principles of Range Management, Proper Season of Use in this case. When forbs like the ones shown in this section are growing there is very little valuable forage in the oak-hickory forest because what grasses and valuable forbs (eg. legumes) do grow have not produced feed. Ergo: stay the heck off the oak-hickory forest ranges at this time. Perhaps of even more importance is the fact that twigs and buds are very susceptible to browsing injury at this season and growth such that regeneration of hardwood species is easily adversely affected. The wood products from these oak-hickory forests are a greater source of revenue than is the 10-30 acres per AUM valuation. It does not require a post doc to figure this out folks. Base of limestone bluff along Modoc Creek in Springfield Plateau section of Ozark Plateau; Ottawa County, Oklahoma. March.

345. Detail of Dutchman's Breeches- The source of the colorful common name for this range forb can be seen in this close-up shot of Dicentra cucullaria. Limestone bluff along Modoc Creek, Ottawa County, Oklahoma. March.

346. Bloodroot (Sanguinaria canadensis)- Here is yet another picturesque range forb from the early spring floor of the oak-hickory forest. And it comes with another colorful common name. This one derived from the red- or orange-colored exudate from the rhizomes that was used by backwoodsmen as a dye to add a little color to their drab garb of buckskins and hunting shirts and gingham or flour sack dresses. Bloodroot is a flowering neighbor of it's pantalooned "cousin" just presented. Both bloodroot and Dutchman's breeches bloom at about the same time and in close proximity to one another. Bloodroot seems to grow slightly higher on hillsides, especially those adjoining streams, than does Dutchman's breeches. Yes, these two species are "kissin' cousins" of a sort. Some taxonomists placed Dicentra species in the Fumariaceae (fumatory family) while others interpreted Dicentra as belonging to the subfamily Fumarioideae in the Papaveraceae (poppy family). Bloodroot has commonly been interpreted as a member of the Papaveroideae subfamily of Papaveraceae. On a limestone bluff alongside Lost Creek in the Springfield Plateau section of the Ozark Plateau physiographic provinece. Ottawa County, Oklahoma. March.

347. One shoot (from a rhizome) of bloodroot at peak bloom and accompanied by a detailed view of it's inflorescence- Take note of the fruit immediately above the leaf. Fruit of bloodroot is a one-locular (locule= cavity of, in this case, the ovary) capsule. This fruit is very similar, obviously, to the capsule of poppy. Bloodroot is one of the first native forbs to flower in the spring on the leaf-covered floor of oak-hickory forests. To be able to live through another hard winter and share the joy of the bloodroot in bloom is one of Nature's blessings to the hill-billy. Bluff above Lost Creek, Ottawa County, Oklahoma. March.

348. Some other seasoning to go with oak and hickory- Wild ginger (Asarum canadense) grows farom New Brunswick westeward to Minnesota and south through Kansas and Oklahoma to Arkansas and Alabama thereby encompassing most of the eastern deciduous forest formation in its range. Its best habitat is rich wooded slopes, especially calcareous soils associated with bluffs like the ones above Lost Creek in the western part of the Ozark Plateau on which this lovely plant was growing. The common name of wild ginger indicated the use that Indians and white frontiersman made of the rhizomes (either fresh or dried) as a seasoning and substitute for the real thing (Steyermark, 1963, p. 572). On east-facing bluffs above Lost Creek, Ottawa County, Oklahoma. April.

349. In rocks and earth tones- Wild ginger growing amid rocks on the east side of limestone bluffs above Lost Creek in the western part of the Ozark Plateau (Springfield Plateau section). The purplish-brown color of flowers of this species blend in with fallen leaves and moist soil almost as if camoflage was the objective. Wild ginger propagates asexually from long rhizomes that give off a pungent aroma (Fernald, 1950, p. 562) which as explained in the preceding caption were used as a seasoning. On east-facing bluffs above Lost Creek, Ottawa County, Oklahoma. April, full-bloom stage..

350. Solomon's seal (Polygonatum commutatum) - This and the next species are members of the asparagus tribe of the lilly family (Liliaceae) are found in the vast deciduous forest region of eastern North America. The pictured specimens of these two species were growing in the oak-hickory forest association in the Ozark Plateau. They were conspicuous plants and though of no commercial value often elicit favorable responses from people visiting the "woods", in particular native plant and wild flower buffs. Young spring shoots of Polygonatum and Smilacina species were eaten by Indians and backwoodsmen. The range of P. communatum is from New England westward to Oklahoma. Ottawa County, Oklahoma. May.

351. Flowers and leaves of Solomon's seal- The bell-shaped flowers and rounded to broadly acute leaves were obvious between views in these two photographs. Polygonatum species are strongly rhizomatous. The origin of the common name was from the seal-resembling scars on the rhizomes. Ottawa County, Oklahoma. May.

352. Synopsis views- More summary images of shoots of Solomon's seal. Thee plants were growing on a transition zone between tallgrass pariaire and oak-hickory forest in the western part of the Ozark (Springfield) Plateau. Shoots of these plants, including a super-abundant crop of flowes, were the result of a record wet spring (fololowing a record cold winter). Solomon's seal has been treated consistently as a member of the asparagus subfamily (Asparagoideae) of the Liliaceae (Dayton, 1960, 23-24; Smith, 1977, p.256). Parallel veins of the Monocotyledoneae were obvious in these photographs. The author never witnessed directly any grazing of Solomon's seal, but the best examples of this species (both specimens in these two and the next three slides as well as in the immediately preceding two slides) were in fence rows just outside of (on the other side of fences) extremely heavily grazed pastures. The inescapable, self-evident conclusion was that beef cattle had grazed out shoots of Solomon's seal that otherwise would have spread via robust rhizomes into the two adjoining pastures. It was more likely that plants (or shoots) of Solomon's seal had existed in these pasture until they had been killed out by heavy cattle-grazing with only remnants persisting in outer fence rows (from which cattle had been excluded). It is amazing what plant species will be eaten by grass-preferring Bos taurus and B. indicus. Ottawa County, Oklahoma. Late May; peak stage of blooming..

353. Musta' been a record- An extremely abundant crop of inflorescences on plants of Solomon's seal at edge of an oak-hickory forest-tallgrass prairie transition zone. After the coldest temperatures ever recorded for this area in February (in excess of -20 degrees Fahrenheit) and one of the wettest springs in history these monocotyledonous forbs produced this remarkable yield of flowers. With a recognition of the rarity and raw beauty of this pehnomenon the Nikon FM was brought into action for some heavy "photo-artillery shelling". This action in combination with the ground-clutching posture of the infantry trained photographer and the green color-capturing quality of Fujichrome 100F produced textbook images for those who appreciate God's own wildflowers and picturesque range forbs. Solomon's seal is first thought of as a forest plant, but in the Springfield portion of the Ozark Plateau this forb is equally at home in dense grassland, both tallgrass prairies and even thick patches of Johnsongrass (Sorghum halapense) which will not have not reached great heights by the time Solomon's seal bears flowers. Ottawa County, Oklahoma. Late May.

354. Cluster of woodland beauties- Example of the inflorescence (consisting of numerous flowers) arising from the leaf axil in a plant of Solomon's seal in an open oak-hickory forest in the western edge of the Ozark Highlands. The drooping (pendulant) feature of the clustered flowers is characteristic of this forb. Kaul et al. (2006, ps. 562-563) provided an especially good description of one of the more distinctive herbaceous forest plants in the Ozark Plateau. Ottawa County, Oklahoma. Late May.

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

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

357. Plant on a Pineywoods pond- Grassleaf arrowhead (Sagittaria graminea) on a local ponded habitat in natural forest clearing in the Big Thicket of Texas Pineywoods. There are several Saggitaria species in North America. These are in the monocotyledon family, Alismaceae (arrowhead family). Big Thicket National Monument, Hardin County, Texas, April.

358. Flowers and young fruits on a Pineywoods pond- Inflorescence on the specimen of grassleaf arrowhead introduced in the preceding slide. This provided a very good example of an indeterminate inflorescence, one that matures "top-down and outside-in" (flowering begins at apices of inflorescence, or on the upper- and outer-most floral tips, al-most and progresses downward and inward). Big Thicket National Monument, Hardin County, Texas, April.

359. Common pitcherplant or yellow trumpets (Sarracenia alata)- Habit and general appearance of common pitcherplant (also shown as pitcher plant) one of the more unique and distinctive of moist forests in the southeastern portion of the North American deciduous forest formation. This specimen featured was at the edge of a colony of its species on highly acid soil in the Big Thicket part of the Texas-Louisiana Pineywoods. Big Thicket National Monument, Hardin County, Texas, April.

360. Pitchers in the Pineywoods- Pitcherplants are these largest of Earth's carnivorous plants and are most readily distinguished by "modified leaves that form hollow, water-containing vessels that are adapted to trapping and digesting animal prey"(McPherson, 2007, p.3). These modified, cylinderical leaves that usually form complete enclosures open only at the top are referred to as "pitchers". Hence, the most commonly used common generic name. The pitcherplant featured here is the species most common in western portions of the southeastern region of the North American deciduous forest formation. It is also typically a generally large and showy species (even when it is out of season for its light lemon-colored corolla). Sarracenia alata is a highly variable species, especially given it rather limited species range. Definitive description of this species (and other species of the pitcher plant family native to the Americas) was McPherson (2007, ps. 195-203). As of this writing, the definitive work on carnivorous plants on ranges and forests of North America is that of Schnell (2002). Big Thicket National Monument, Hardin County, Texas, April.

361. Fruit pitcher- Shoot apices of the yellow trumpets pitcherplant bearing the leaf-enclosed fruit. The fruit has been interpreted as a loculicidal capsule, which is that dehisces due to or through openings in the locules between partitioning tissue (Smith, 1977, ps. 122, 300). Pitcher plants are in the family Sarraceniaceae. Big Thicket National Monument, Hardin County, Texas, April.

362. Plant eats bugs- Portion of lower pitcher of yellow trumpets pitcherplant revealing a partly digested insect being used as a source of nitrogen for a plant highly pted to impoverished soils. Big Thicket National Monument, Hardin County, Texas, April.

Pterophyte is the term for spore-bearing vascular plants. These are "intermediate" (in evolutionary development, taxonomy, etc.) between the traditional units of Byrophyta and the Spermatophyta (seed-bearing plants, usually a taxon at the division level). Traditionally the pterophyte taxonomic level (division) has been called Pteridophyta. All three of these traditional units or taxa were included in the Embryophyta (subkingdom of plants in which the zygote develops into a multicellular embryo while enclosed within the female sex organ or within the embryo sac). In the traditional taxonomic hierarchy the vascular plants (those possessing xylem and phloem including both Pteridophyta and Spermatophyta) were distinguished from the Bryophyta by being placed in the Tracheophyta (the taxon, variously a division or superdivision, of vascular plants). In other words, Tracheophyta minus Pteridophyta leaves Spermatophyta (Gymnospermae and Angiospermae). The Pteridophyta included the club mosses (Lycopocineae), horse-tails or scouring rushes (Equisetineae), and the ferns (Filicineae). Some taxonomic schemes had a taxon designated Pteropsida that grouped (largely on basis of complex, relatively large leaves) the ferns, gymnosperms, and angiosperms. Spore-bearing plants from the thallophytes through and including the pteriophytes have been called cryptogams (pteridophyts are vascular crypotgams). Plants have thus traditionally been grouped taxonomically in various arrangements depending on what features were of concern for different purposes. In effect, this is a form of taxonomic bilingualism. It is desirable, even essential, in instances where several features are of interest simultaneously. This is often the case when studying vegetation and, even more so, when applying this study to production agriculture. Those individuals are uninformed who view terms and taxa like the ones just discussed as meaningless or useless because they are archaic or, even, obsolete (two different things). If these individuals arrogantly persist in viewing such words as meaningless these folks are stupid (incapable of learning). There are several taxa of pteridophytes in range and forest vegetation. Pteridophytes are especially common in the various deciduous forest cover types. A few were selected for inclusion in this section devoted to the oak-hickory forest.

363. Colony of smooth scouring rush (Equisetum laevigatum)- Equisetum is, as obvious from the prefix, the genus of pteridophytes given the common name of horsetail. Many of the Equisetum species do not have the namesake horsetail arrangement of primitive leaves encircyling the shoot. Those Equisteum species whose leaves are reduced to rudiments were given the common name of scouring rush, a common name which appeared to aptly describe even those having a "horsetail". In fact, given that most Equisetum species in many locales are "bald" tails, scouring rush is often the more apt common name (genus designation notwithstanding). Lack of well-developed leaves that have a typical leaf form was a criterion used in classifying ferns as more advanced and closer to spermatophytes than are the club mosses and scouring rushes (ie. ferns are the least primitive of the vascular cryptogams). Within the oak-hickory forest many of the vascular cryptogams grow best on the more moist habitats.This colony of smooth rush was growing along a high bank of Lost Creek flowing through the extreme western edge of the Ozark Plateau section of the oak-hickory forest. Ottawa County, Oklahoma. July.

364. Shoots of smooth scouring rush- The texture of the shoot and a spore-bearing strobilus of smooth scouring rush were shown in this photograph. Strobilus (plural, strobolii) in this usage refers to a group or unit of sporophylls (including their sporangia) more or less densely encirclying the central sexual axis (the shoot apex in this genus). Sporophylls are modified leaves or leaf-like organs that bear spore-containing sproangia (singluar, sporangium; the case in which spores are formed and stored before release). Specimen from the stand shown immediately above. Bank of Lost Creek, Ottawa County, Oklahoma. July.

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

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

367. Colony of rattlesnake fern (Botrychium virginianum)- A high proportion of forest pteridophytes require relative moist (= mesic) soils (ie. they are at the "higher end" of mesphytes). One exception is rattlesnake fern which grows on soils having average quantities of soil water. The general common name for Botychinum species is grape fern, but B. virginanum is usually called rattlesnake fern. This colony of rattlesnake fern was thriving on a microsite that had a deep layer of rotting oak and hickory leaves in a second-growth oak-hickory forest along the extreme western edge of the Ozark Plateau. Newton County, Missouri. May.

368. Shoot of rattlesnake fern- An example of the large and intricately patterned leaves and the sori-bearing sporophyll of rattlesnake fern were "captured" in this slide. The sporophyll is the spore-bearing leaf of the pteridophytes. Sori (plural of sorus) are the "fruit structures" (typically borne on undersides of leaves) which bear numbers (usually large numbers of) spores. Newton County, Missouri. May.

369. Christmas fern (Polystichum acrostichoides)- This fern allegedly got it's common name from the fact that it is characteristically green (and often producing new fronds) in the middle of winter (ie. evergreen fronds). Christmas fern prefers moist (but typically well-drained) habitats especially along banks of shaded, flowing streams. This specimen was one of numerous plants of this species growing along the bank of Modoc Creek in a bottomland forest of sycamore, sugar maple, and box elder (Acer negundo), but here beneath huge black oaks. Beaver (Castor canadensis) had cut off many of the fronds and carried them into their lodges which were burrows in the creek bank (ie. bank beaver). What use beaver made of them was not be determined. Ottawa County, Oklahoma. January.

370. Put-together morphology- Details of frond of Christmas fern shown in a three-slide sequence from general morphology of frond blade (first slide)to details of ets or secondary leaves (second and third slides) of the frond. Leaflets are alsso known as pinnae (pinna, singluar). Students were referred to Diggs et al. 91999, p. 178) and standard Botany texts such as Weoer et al. (1982, ps. 586-589) for explanations of morphological structure of fern fronds, rhizomes, and roots. Christmas fern is aptly named (in the northern hemisphere) as it thrives in winter when it gets the greatest amount of sunlight with leaves shed from trees and shrubs. Bank of Modoc Creek, Ottawa county, Oklahoma. Late December.

371. Maidenhair fern (Adiantum pedatum)- This is perhaps one of the most graceful of all forest forbs. This was one of several individuals of this species that grew in close proximity to the Christmas fern shown in the preceding slide. Maidenhair ferns grew several feet higher up (on a bluff) from the Christmas ferns on the bank of a slow-moving stream in the far-western Ozark Plateau. Unlike the evergreen Christmas fern, maidenhair ferns were dormant and died back to the ground surface in winter. The maidenhair ferns shown in this slide were were growing in late summer in dense shade and on a moist east slope. They received less than three hours of direct sunlight during the longest days and had to survive on that and/or what diffuse light "filtered" through the leafy overstorey of black oak and sugar maple. On a limestone bluff above Modoc Creek, Ottawa County, Oklahoma. August.

Bryophytes are those range and forest nonvascular plants of the general groups of mosses, hornworts, and liverworts. (Recall that "plants" was used herein as in the older or more traditional usage of the two kingdoms of organisms that the author viewed as more practical for discussion of vegetation.). More specifically bryophytes are members of the Bryophyta (a division of plants in the Plantae); in short, they are nonvascular spore-producing plants. Bryophytes have alternation of generations (gametophyte and sporophyte), but in contrast to the vascular plants the dominant and conspicuous generation of bryophytes (ie. the so-called plant) is the gametophyte (gametophytic generation). While bryophytes lack differentiation into true roots they have root-like structures called rhizoids which anchor the gametophytes to their substrate (bryophyte rhizoids do not function in absorption). Bryophytes, like thallophytes (thallus-- the type of plant body that is undifferentiated into root and shoot-- plants such as algae, bacteria, and fungi), are primarily important in Range Management and Forestry as reducers or decomposers and therefore in processes like soil formation and plant succession (ie. the Clementsian process of reaction now more commonly known as facilitation). This was discussed above immediately before presentation of fungus species. Raven et al. (1992, ps. 298-316) provided an excellent introductory discussion of the Bryophyta. Shaw and Goffinet (2000) wrote a comprehensive and the recent classic text on the Bryophyta for the "really serious" student.

372. Gametophye of a "true moss" (order of Bryales)- "Moss" is one of the most confusing and most misleading names in all of Botany. "Moss" is applied to every sort of "plant" from reindeer moss (a lichen eaten by caribou and reindeer) to Spanish moss (a monocotyledonous epiphyte often eaten by native and domestic ruminants). Obviously such "mosses" are range plants by definition because they are native plants that serve as feed for grazing/browsing animals. Absent this axiomatic and self-evident definition, these and other "plants" growing on range are still range plants even if they function in less conspicuous roles such as decomposition, soil formation, nutrient cycling, and plant succession. True mossses provided such examples of these roles or functions in forest and range ecosystems. "True mosses" has been the designation traditionally used for the one (of three) class in the division of Bryophyta know as Musci. In some of the more recent works (eg. Raven et al., 1992, p. 308) Bryophyta division still consist of three classes one of which is that of the true mosses but known as Bryidae (instead of Musci). The Musci or Bryidae bryophytes are partially saprophytic being dependent on decaying organic mattter. The specimen photographed here (the species of which was not identified ) was attached to a piece of chert that had a deep facet which had acumulated rotting oak leaves. Oak-hickory forest in Ozark Plateau. Ottawa County, Oklahoma. January.

373. Gametophytic and sporophytic generations of a species of true moss (Funaria hygrometrica), Musci- In these two macrolense shots the leafy gameotphyte is visible at the base of the moss while the mature sporophytes are very prominent as the apex of the moss body. These shown here were female sporophytes known as archegonia (singlular is archegonium) or archegonial heads which consist of a capsule (= sporangium; plural is sporangia) that are borne on a seta (stalk). The first of these two slides presented a view from the top to show the overall appearance of moss in the sporophytic stage. The second slide was a side-view of the moss which showed more clearly both the leafy gametophytic generation and the stalked capsule of the sporophytic generation. The covering of the capsule is the calyptra the sharp, pointed tip of which is the operculum. This is the lid of the capsule which when shed opens up the capsule, the inside of which contains spores which are released upon this opening. While these are not vascular plants and are not differentiated into roots, stems, and leaves there is some differentiation of tissues. This occurs in the stalk as well as in the leafy gametophyte. At the base of this sectional sample of moss are the rhizoids which are the root-like structures (more like root-hairs actually) of the gametophytic generation that function only to anchor or hold the plants (absorption of water and mineral nutrients occurs directly through the gametophyte). Oak-hickory forest in Ozark Plateau. Ottawa County, Oklahoma. March.

374. Colony of juniper hair cap moss (Polytrichum juniperinum)- This well-developed stand was on the floor of a post oak flats site of the oak-hickory forest in the Ozark Plateau. Post oak flatwoods is a very mesic form of the oak-dominated deciduous forest. It is not a swamp or any form of wetland, but it is poorly drained flatland of primarily clay soils. In the Ozark Mountains and adjoining savanna of the Prairie Peninsula post oak is usually the only species that can survive the poor drainage of this site (hence the name of post oak flats). The high moisture condition of the soil and shade from post oaks provided a microhabitat (microsite) satisfactory for juniper hair moss. Newton County, Missouri. April.

375. Close-up of juniper hiar cap moss- This is the gametophyte (gametophytic generation) of this species. Newton County, Missouri. April.

The relatively high primary productivity of the oak-hickory forest results in heavy accumulations of detritis ("yields" of necromass) that are rich substrata for many kinds of reducers (= decomposers) among forest and range plants. (The author acknowledged the value and legitimacy of the five kingdom taxonomic system, but herein used "plants" in the traditional botanical usage for simplicity, utility, and practical application.) Botanical organisms functioning as reducers extend from bacteria, algae, fungi, lichens, and bryophytes to vascular plants. A sample of these was included below beginning with the fungi. Not all fungi are saprophytes. Some fungi are saprohytic, of course, but others form mycorrhiza or the mycorrhizal symbiosis with roots of vascular plants while others are parasitic (on vascular plants, insects, or even othr fungi). Some fungus species are a combination of saprophyte and parasite (examples were presented below). Likewise, not all saprophytes are fungi or bryophytes. An example of a saprophytic vascular plant (a dicotyledon) was presented below following presentation of several fungal species. Fungi are one of the three major groups of organisms traditionally regarded as plants (or plantlike taxa) that lack vascular tissue (ie. simplistically defined as nonvascular, undifferentiated lower plants). These "plants" (plantlike organisms) were historically regarded as thallus plants or thallophytes, members of the Thallophyta (usually listed as a subkingdom). Thallophytes are those organisms having plant bodies not differentiated into roots and shoots, lacking vascular tissue, and having gametes enclosed only by a cell wall such that their zygotes do not develop into embryos while inside the female sex organ (Wilson et al., 1971, ps. 447-451). With acceptance of the five kingdom classification system of organisms proposed by such stellar scientists as Robert H. Whittaker the older or traditional two kingdom scheme fell into disuse. While this development was generally an improvement (especially for the teaching of Biology to beginners) some of the taxonomic groupings (eg. thallophyte) still make sense especially for applied use in Agriculture. The author of this web publication periodically resorted to groups like the Thallophyta for practical use (ie. ease of teaching cencepts in conservation, agricultural production, etc.). Thallophytes and, as discussed below, bryophytes are essential to development of vegetation and soils. These lower plants typically form one or more layers of vegetation. Also, they are usually pioneer species. These primitive plants are among the first organisms to grow on the raw parent material of a sere such that they improve the habitat for higher plants of later seral communities. This was what Clements termed reaction, one of the processes in development of vegetation (Clements, 1916a, 79-96 passim; Weaver and Clements, 1938, ps. 234-241). Years later this process was termed facilitation (Connell and Slatyer, 1977) and in time became known as the facilitation model of plant succession (Begon et al., 1990, ps. 632-633, 635, 641-642). Clements' specific view (almost as much philosophical as theoretical) was that "[e]ach climax formation had its individual or ontogenetic development"... such that "it shows a phylogenetic development from a preceding climax or community". "If the phylogeny of the community comprises the same general process as that of the species, it should be recapitulated by the ontogeny as seen in the sere" (Clements, 1916a, ps. 344-345). An assumed phylogeny beginning with lower plants such as algae, liverworts and mosses, lichens, etc. (as determined from the fossil record) implied-- at least to Clements-- that development of vegetation on current seres began with these thallophytes and bryophytes. This recapitulation component of Clements' grand theory of plant succession was probably the least understood part of his complicated model of vegetation development, but undoubtedly all students of plant succession have observed that some of the earliest species to pioneer an area (especially a prisere, a fresh or newly created bare area of parent material or the area having to undergo primary succession) are the primitive plant forms. Whatever processes and paths of development are involved, the thallophytes and bryophytes do facilitate development of range and forest plant communities by the processes of plant succession. It was for this reason that a few of these species were included in this publication on range and forest cover types. There are numerous ouytstanding field guides to the fungi (ie. mushrooms) of North America including Krieger (1967), Orr and Orr (1979), Lincoff (1981), Pacioni (1981), McKnight and McKnight (1987), Metzler and Metzler (1992), and the massive Arora (1986). Mycology is the study of fungi. There are numerous outstanding texts and references for the fungi. Pritchard and Bradt (1984) was recommended for readability yet thoroughness. Carlile and Watkinson (1994) was a comprehensive text that covered basic biology, including ecology, evolution, genetics, etc., but for overall, concise reference (especially for beginners) the basic Botany textbook of Raven et al. (1992, ps. 208-243) seemed easiest to use.

376. Morel or yellow morel (Morchella esculenta= M. rotunda)- This fruiting (=fruit) body (reproductive structure; in higher fungi the fruit body is sometimes designated the carpophore) of this fungus is delicious and a delicacy to hillbillies who seek it out in the leafy forest floors of such ancient mountains as the Appalachians and Ozarks. Morchella species are ascomycetes (Ascomycotina is the largest subdivision of true or fleshy fungi) within the Discomycetes class. There are several species of morels across North America but the species presented here is the most common one in the oak-hickory forest of the Ozark Mountains. Banks of Modoc Creek, Ottawa County, Oklahoma. April.

377. Ink cap (Coprinus radians)- On this rotting log covered with decomposing oak leaves is the complete body of a fungus in the group, basidiomycetes (Basidiomycotina is a subdivision of the higher, fleshy, or true fungi), more specifically the Hymenomycetes class therein. The species shown here is one of the best or standard textbook examples of the true fungi, those species that produce fleshy fruit-bodies know generically as mushrooms and/or toadstools. The two general parts of a mushroom are 1) the filaments of hyphae which form a network known collectively as the mycelium or spawn and 2) the fruiting body, often called the carpophore, which bears the reproductive structures that produce spores. The fruit-body is the obvious, often conspicuous, part of the true or higher fungi (ie. the "mushroom") which forms from the mycelium. The latter part of the true fungi are not differentiated into roots and shoots and are typically not seen by human eyes. The only part of fungi usually visible (again, to humans) is the fruit-body, the mushroom. If some of the mycelium (hyphae filaments) remain attached to the carpophore when someone picks or knocks over the mushroom he would naturally think of the mycelium as the "roots" which is obviously incorrect. C. radians is unusual in having bright orange hyphae threads and, when these are accompanied by the carpophore (as shown here), students have the rare opportunity to see the whole "plant" (entire fungus body). In this slide there is one live or "fresh" and one dead (withered) carpophore. Some fungal species have a carpophore or mushroom that includes a prominent "cap" as its hymenium, the spore-bearing surface of the Ascoymetes and Basidiomycetes. In C. radians the spore-bearing underside of the cap is dark-colored hence the common name of ink cap (next slide). Obviously this fungus is a saprophyte. It was shown to offer students a classic example of a saprophyte or saprophytic "plant" which is one of the major categories within the reducer or decomposer group of organisms in forest and range ecosystems. First terrace of Modoc Creek, Ottawa County, Oklahoma. April.

378. Detail of the carpophore of ink cap and surrounded by the mycelium- The underside of the cap of Coprinus radians showing the gills that bear the spores of this species being supported or held by the stipe, the stalk or "stem" of the carpophore that bears the hymenium (in this species the latter developes as a cap). The substrate was a rotting oak log. Bank of Modoc Creek, Ottawa County, Oklahoma. April.

379. Close-up of the hyphae filaments comprising the mycelium of ink cap- This is the vegetative (non-fruit-body portion) of a fleshy or true fungus growing on a decaying oak log. An individual filament is a hypha (hyphae is the plural). The many hyphae form the mycelium which in this species is a brilliant orange. It was photographed in Kodochrome (hence without color-enhancement). Modoc Creek, Ottawa County, Oklahoma. April.

380. Wood ear or tree ear (Auriculularia auricula) on dead elm (Ulmus americana) limb- In the first slide, almost indistinguishable on this dead limb, are dry fruit-bodies (accompanied by equally dried-out lichens) of a species of the Hymenomycetes (class) of the Basidomycetes (subdivision) known by the colorful (and most appropriate) common name of wood ear. In the second slide, which had been taken 24 hours earlier when the fruit-bodies (the "mushrooms") were conspicuous, wood ear carpophores (and accompanying lichens) had swollen from a recent rain. This saprophytic species is but one of many fungi that quickly decompose the vast detritis or necromass (dead organic matter that was previously biomass) produced in the oak-hickory forest. This species of Hymenomycetes was growing in the "dead of winter", a common season for appearance of many carpophores due to abundance of water in detritis as a result of low evaporation rates. Ottawa County, Oklahoma. December.

381. Wood or tree ear (Auriculularia auricula) capophores fully hydrated- Close-up shot of wood ear immediately after a warm winter rain. Ottawa County, Oklahoma. December.

382. Wood ear fruit-body drying out- Appearance of the member of Hymenomycetes known as wood ear about six or seven hours after cessation of winter rain. The capophore was quickly returning to the more common dehydrated state as shown above. Ottawa County, Oklahoma. December.

383. Tooth fungus, old man's baerd fungus, or lion's mane fungus (Hydnum erinaceus= Hericium erinaceus) growing on dead wood of a fire scar- An old fire scar of black oak (Quercus velutina) in an oak-hickory forest of the Ozark Plateau was the substrate for this primarily saprophytic fungus. This fungal species (and the next one in this line-up) can be viewed as partly parasitic because it frequently grows on wounds of living deciduous trees as well as on totally dead wood. Like the preceding species this fungus is in the Hymenomycetes class of the Basidiomycetes subdivision. (As in the case for all groups of organisms there are different taxonomic schemes or arrangements and names for the hierarchial levels such that the class level may be a subclass in the hierarchy of another author). The fruit-body of H. erinaceus is edible when young, but it takes a long to cook the tough tissue. Newton County, Missouri. December.

384. Fruit-body of a tooth or comb fungus (H. erinaceus)- Detail of the carpophore on the fire scar of the preceding slide. The fruiting body of members of the Basidiomycetes has been designated the basidiocarp The basidocarp bears the basidia (singlular is basidium) which are the individual microscopic, spore-producing cells. H. erinaceus is a saprophyte, but it is also somewhat parasitic hastening the rotting of live wood adjacent to dead wood that is the main substrate. Oak-hickory in Ozark Plateau. Newton County, Missouri. December.

385. Basidiocarps of oak conk or cracked cap polypore (Fomes robustus= Phellinus robustus) on an ancient black oak- The trunk of this old black oak that had survived for decades on a rockpile of a site "sported" several carpophores (fruiting bodies which are basidocarps in this class of fungi) of this member of the Hymenomycetes (in the Basidiomycetes class). Oak conk is a shelf or cork fungus within the group known as polypores and/or bracket fungi. Polypore refers to the many pores (more precisely, spore-producing tubes) on the underside of the cap (carpophore, specifically the basidiocarp).The polypores or shelf mushrooms (conks are but one category of these bracket fungi) are in the general group viewed as true or fleshy fungi. Oak-hickory forest in Ozark Plateau. Ottawa County, Oklahoma. April.

386. Oak conk- The basidiocarp of a woody shelf fungus showing the underside with the layer of tissue from which the spores are released from the numerous spore-producing tubes. The typical cracked cap of the polypores (family: Polyporaceae) was also clear in this photograph. This was one of numerous carpophores on the trunk of the old black oak shown in the preceding slide.The shelf- or bracket-like basidiocarps of this and related species are just as woody as the substrata they grow on. Polypores have generally been viewed as more saprophytic than parasitic, but most species function in both capaacities. Arora (1986, p. 549) pointed out that the mushrooms of this group (Polyporaceae and related genera) "are absolutely indispensable to the forests of this world" in their function as decomposers. These are the major "wood-rotting fungi" causing 90% of the rotten-wood damage to standing timber in North America. This fact could be viewed as indicating a major forest pathology crisis, and at one time foresters who interpreted Forestry as mere tree farming (and thus sought to eliminate all "enemies of the forest") may have held such views. That was before application of the ecosystem concept brought back a more balanced perspective or maybe just some old-fashioned woods wisdom in new terms for original principles. Today's foresters and rangemen appreciate the essential role of decomposers in reduction of detritis (plus the ecological role of detritis itself), cycling and availability of nutrients, and provision of food for consumers (both vertebrates and invertebrates). Arora (1986, p. 549) stated the case for the polypore fungi and put the role of decomposition in proper perspective. "Without them there would be no logging industy in the first place: every cut stump, felled log, and lopped-off limb would be indefinitely on the forest floor, the woods would quickly become impenetrable, and the new trees would have neither room nor nutrients to grow". Oak-hickory forest in Ozark Plateau. Ottawa County, Oklahoma. April.

387. Sulphur shelf, chicken mushroom, or chicken of the woods (Polyporus sulphureus= Grifola sulphurea= Laetiporus sulphureus)- This colorful member of the Polyporaceae is edible (when young) and so distinctive that it is not likely to be confused or misidentified). Sulphur shelf is also both saprophytic and parasitic. Krieger (1967, p. 157) labeled the chicken mushroom as one of "the wood-destroying fungi" and "a most persistent enemy of coniferous as well as deciduous trees". He listed oaks, locusts, maples, alders, walnuts, pines, hemlocks, and spruces among it's host species. The fine specimen shown here was growing on an old fire scar at the base of a black oak in the Ozark Plateau. Ottawa County, Oklahoma. October.

388. Pinesap (Monotropa hypopithys)- This saprophyte is a vascular plant included by various treatments in such families as wintergreen (Pyrolaceae), Indian pipe (Monotropaceae), or as a separate subfamily (Monotropoideae) of the heath (Ericaceae) family. Color alone (absence of chlorophyll) indicated that this dicotylendous species cannot carry out photosynthesis and must be either parasitic or saprophytic. It is the latter by means of forming mutualism with mycorrhiza on its roots. This specimen was growing on rotting litter on the floor of a black-oak dominated oak-hickory forest in the Ozark Plateau. The dead stems from the previous growing season were photographic evidence of the perennial length of this species' life cycle. Ottawa County, Oklahoma. June.

389. Pinesap- Close-up photograph of the vascular saprophyte presented in the preceding slide. This specimen had less pubescence than is typical for this species. It was growing on the organic matter of rotting oak and hickory leaves in the western part of the Ozark Plateau. Ottawa County, Oklahoma. June.

390. Interior of an upland mixed oak (white, red, post, black)-hickory forest that has extremely lush and diverse shrub and herb layers. Understory plants include tick clover, blackberry, grape vine, Virginia creeper, poison oak (Rhus toxicodendron= Toxicodendron toxicarium), bluestem, and panic grasses. Second growth forest with excellent regeneration of climax tree species, especially hickories. Note that this excellent botanical diversity and forest regeneration is occurring under light or conservative grazing/browsing by cattle (Herefords are visible in center). Adair County, Oklahoma. May, late vernal aspect. FRES No. 15 (Oak-Hickory Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Dry-mesic chert forest. Oak-Hickory Series of Brown et al. (1998).

391. Uneven-aged management in upland oak-hickory forest- Selective cutting (= selective felling) in an Ozark Plateau chert forest dominated by black oak which was harvested for veneer to be used in making church furniture. Trees were felled in winter and this is the scene three months after logging. Most harvested trees were 70-90 years old. Note total absence of any soil disturbance or damage to young trees. Logs were carried— not drug —out by a rubber-tired skidder. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Selective cutting like clearcutting has both proper and improper applications. The benefits of proper selection-cutting (= selection felling) to maintain forest with all, or most all, age classes with minimal adverse impacts on the forest resouces is forest practice par excellence. Unfortunately, all too often selective cutting is not selection cutting (= the selection method) but simply amounts to high-grading, “the removal of the most commercially valuable trees (high-grade trees), often leaving a residual stand composed of trees of poor condition or species composition” Selective cutting is “a cutting that removes only a portion of trees in a stand… it is a general term that should not be confused with cutting done in accordance with the selection method” (Helms, 1998). Selection method (= selection felling) was defined under slides of the Sierran Mixed Conifer Type. Even-aged and uneven-aged management or regeneration methods are only one part of silviculture which in turn is but one component of forest management.      “Forest Management- the practical application of biological, physical, quantitative,      managerial, economic, social, and policy principles to the regeneration,      management, utilization and conservation of forests to meet specified goals and      objectives while maintaining the productivity of the forest” (Helms, 1998). Although the selective harvest seen here was not conducted according to a forest harvest plan (including harvest scheduling) or the guidelines of scientific forest management, the intensity and frequency of cutting was so low as to have minimal, if any, detrimental effects on the forest. Acorn production is so great as to overwhelm feed demands of wildlife and assure natural regeneration by sexual reproduction. Plus there is asexual or vegetative reproduction by :

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

Ottawa County, Oklahoma. August. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

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

Ottawa County, Oklahoma. Estival aspect, September. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

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

Organization Note: More examples of forestry practices (including harvest and regeneration) in the oak-hickory types were presented in the chapter, Oak-Hickory Forests-II (under the section, Use and Abuse of Oak-Hickory Forests).

Bottomland Forests

395. Bottomland hardwood forest on north slope in Ozark Plateau— Extremely species-diverse community shown here in early spring. Species include sycamore, extreme right foreground; western hackberry, foremost trunk; black walnut (Juglans nigra), two trees immediately behind preceding foreground trees; chinkapin oak, grey trunk just behind walnuts; sugar maple (Acer saccharum), two black trunks forming V-shape in background and one behind and partly concealed by the sycamore; basswood or American linden (Tilia americana), on upslope to left of maples; and white ash (Fraxinus americana), left-center midslope. Redbud and flowering dogwood are in the shrub layer but the dominant shrub is spicebush (Lindera benzoin) which comprises a lower shrub layer.The herbaceous layer, which is clearly visible in this early spring stage, includes bloodroot (Sanguinaria canadensis), trout lily or yellow dog-tooth violet (Erythronium americanum), broadleaf waterleaf (Hydrophyllum canadense), rue anemone (Anemonella thalictroides), false rue anemone (Isopyrum biternatum), and scattered colonies of the ever-present Mayapple. Following completion of their annual growing season these species are replaced in their location by great Indian plantain (Cacalia muhlenbergii) which then dominates a tall herb layer through mid-summer. This is a unique transect view going from the first terrace of a mid-size creek to the top of limestone bluffs. The upper-most forest is dominated by black and red oaks as shown in preceding slides. Downslope from the upland oak forest is a mid-slope sugar maple-basswood-chinkapin oak zone. The bottomland or floodplain forest is the sycamore-hackberry-black walnut-white ash forest. Thus this transect includes SAF cover types 61 (River Birch-Sycamore, variant form) or 94 (Sycamore-Sweetgum-American Elm, variant of), 26 (Sugar Maple-Basswood), and 52 (White Oak-Black Oak-Northern Red Oak), the latter is indistinguishable in background. Ottawa County, Oklahoma (bank and bluffs of Lost Creek). March, early vernal aspect. Mesic bottomland forest according to Missouri Natural Areas Committee (1987). FRES No. 15 (Oak-Hickory Forest). No obvious Kuchler unit. Mixed Hardwood Series in Southeastern Swamp and Riparian Forest biotic community of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

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

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

Organization note: a climax, bottomland, mixed hardwood that developed on the floodplain of Modoc Creek and that was conterminous with the sugar maple-northern red oak-white ash-pignut hickory-white ash mesic forest on an east slope along bluffs above this creek was treated separately under Southern and Central Forests-II.. That chapter also has other bottomland forest types or forms as well.

Pecan Bottoms- Example of Southern Bottomland Forests

The following section treated bottomland forests dominated by pecan (Carya illinonensis= Hickoria pecan) to furnish an example of a specific form or subtype of either Sugarberry-American Elm-Green Ash (SAF 93) or Sycamore-Sweetgum-American Elm (SAF 94). The latter forest cover type (SAF 94) had previously been designated as Sycamore-Pecan-American Elm (Erye, 1980, ps. 65-66). More detailed coverage of these forest range cover types was included with chapters, Miscellaneous Forests and Southern and Central Forests-II

398. Bottomland forest (actually more of a woodland physiogonomy as crowns overlap but slightly) of pecan (Carya illinoinenisis) with an understory devoid of shrub layers and with the herb layer dominated by Canada wildrye with frostweed (Verbesina virginica) an important associate. Indiangrass and eastern gamagrass add a tallgrass element. In the upper two post oaks (left center) entered this otherwise single-species stand or pecan consociation. In the second photograph two large pecans (only their lower trunks showing) shaded a carpet of the two cool-season perennial native grasses, Canada wildrye Texas wintergrass, while warm-season perennials like little bluestem, Indiangrass, and perennial dropseeds "waited in the wings" for their time on the shaded stage. This tree-dominated vegetation was an irregular gallery forest along the South Llano River. The pecan is the stately State Tree of Texas and the community shown here is known as "pecan bottoms" by locals who frequent it come nut season. Another beautiful permanent deciduous forest range type. Kimble County, Texas. June, estival aspect. An "island" or isolated part of FRES No. 15 (Oak-Hickory Forest Ecosystem) and K-91 (Oak-Hickory Forest). No SAF designation readily fit this range vegetation. This forest range appeared similar to the Cottonwood and Live oak Bottomland Types (SAF 63 and SAF 89), but most likely a Variant SAF 94 (Sycamore-Sweetgum-American Elm) that had the former SAF designation of Sycamore-Pecan-American Elm. Edwards Plateau- Edwards Plateau Woodland Ecoregion, 30a (Griffith eat al., 2004).

399. Texas-size- Though it was hard to show with all the shade, several widely scattered giant pecans formed a pecan bottom much like those that existed in Texas river bottoms before European man presumptuously assumed that he could improve things. For big trees like our "centerfold hero" here stocking rate has be relatively low. Prof. Benton Storey (Texas A&M Universty) felt that production of big pecan trees and high nut yields were best obtained with only one tree per acre. Pecan bottoms would thus be savannahs or, at most, woodlands because their crowns would not contact each other at such stocking. Studded T steel post serving as stakes for pecan seedlings and saplings and the top of one post by the pecan-picker's "centerfold" provided scale for size comparison. This part of the understorey of this river bottom woodland had been mowed (perhaps in anticipation of nut season). Kimble County, Texas. June, estival aspect. An "island" or isolated part of FRES No. 15 (Oak-Hickory Forest Ecosystem) and K-91 (Oak-Hickory Forest). No SAF designation readily fit this range vegetation. This forest range appeared similar to the Cottonwood and Live oak Bottomland Types (SAF 63 and SAF 89), but most likely a Variant SAF 94 (Sycamore-Sweetgum-American Elm) that had the former SAF designation of Sycamore-Pecan-American Elm. Edwards Plateau- Edwards Plateau Woodland Ecoregion, 30a (Griffith eat al., 2004).

400. East Texas pecan bottoms- Bottomland mixed forest in the Texas Pineywoods dominated by pecan with hackberry and water oak as associates. Foremost tree at left margin was a large water oak with a single bole of high-quality timber. The large tree at right-of-center with numerous major limbs and scaley, brown bark was pecan as were most of the trees (pole-size) in background. The smallest of three trunks in foreground (in front of and aligned along left edge of pecan was sugarberry (Celtis laevigata). Flowering dogwood (center green shrub) formed an interupted upper shrub layer. The invasive alien Japanese honeysuckle (Lonicera japonica), seen here as the green plant in left to center foreground, comprised a spotty though dense lower shrub layer. The exotic and now widely naturalized Japanese honeysuckle (a horticultural escape "gone wild") is a major noxious plant problem in forest understories, especially in pecan bottoms. There were only a few grasses (panicgrasses were most obvious), grasslike plants (limited to Carex spp.), or forbs in the understorey of this bottomland forest. The alien L. janpoica appeared to have crowded out herbaceous plants.Species of green briar and grape along with rattan vine or Alabama supplejacck formed a "jungle" of vines among the smaller pecans in bacdground. Local mowing at edge of this forest near a campground had apparently effectively controlled woody vines in this sample of lowland Pineywoods forest. Davy Crockett National Forest, Houston County, Texas. March, vernal aspect (post dogwood blooming stage ). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). Variant SAF 94 (Sycamore-Sweetgum-American Elm) that had former SAF designation of Sycamore-Pecan-American Elm. Closest biotic community designation of Brown et al. (1998, ps. 37, 38) was Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1, but there should have been a Oak-Hickory Series for Southeastern Deciduous and Evergreen Forest 123.. South Central Plains- Southern Tertiary Uplands Ecoregion, 35e (Griffith et al., 2004).

401. Another example of a pecan bottom- This example of bottomland forest dominated by pecan was along the small slow-moving Alarm Creek on extremely fertile alluvium. The four big trunks in right foreground and center midground were pecan. Other trees included both post and blackjack oaks. Hackberry was also well-distributed. Greenbriar was the dominant shrub. There was a well-developd herbaceous layer of Canada wildrye, little bluestem, and sand lovergrass (Eragrostis trichoides). Small isolated patches of giant cane (Arundinaria gigantea) grew along banks of the creek. Erath County, Texas. April. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-75 (Cross Timbers). Variant SAF 94 (Sycamore-Sweetgum-American Elm) that had former SAF designation of Sycamore-Pecan-American Elm. (Griffith et al., 2004). Closest biotic community designation of Brown et al. (1998, ps. 37, 38) was Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1, or more descriptively perhaps, a transition of this Oak-Hickory Series and Bluestem "Tall-grass" Series, 142.11, Plains Grassland 142.1 Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

402. Young pecans and tallgrasses in summer- Part of the floodplain of the Bosque River in the Western Cross Timbers supported this stand of "half-growed" young pecans with bur oaks of the approximate same age cohort as associate tree species. Understorey was locally dominated by Canada wildrye, especially prominent in the second of these photographs where that cool-season member of the Hordeae or Tritaceae tribe was taller than the top wire of the fence enclosing this nice sample of "pecan bottoms". The associate herbaceous species varied locally from such species as the native and-should-have-been-one-of-the- climax -dominants Indiangrass to naturalized Johnsongrass. Other grasses included big, little, and silver bluestems; Texas wintergrass, several perennial dropseeds, sideoats grama (Bouteloua curtipendula) tumble windmillgrass (Chloris verticillata), and tumblegrass (Schedonnardus paniculatus). Forbs were varied but, as to be expected, composites "ruled" with prominent and common species including western ragweed (Ambrosia psilostachya), Baldwin ironweed (Vernonia baldwinii), iand frostweed or white crownbeard (Verbesina virginica). The major shrub was common greenbriar or catbriar (Smilax rotundifolia). Other shrubs were trumpet creeper (Campsis radicans) and poison oak. Hamilton County, Texas. June. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-75 (Cross Timbers). Variant SAF 94 (Sycamore-Sweetgum-American Elm) that had former SAF designation of Sycamore-Pecan-American Elm. (Griffith et al., 2004). Closest biotic community designation of Brown et al. (1998, ps. 37, 38) was Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1, or more descriptively perhaps, a transition of this Oak-Hickory Series and Bluestem "Tall-grass" Series, 142.11, Plains Grassland 142.1 Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

403. Pecan timber- The forest form of pecan bottoms was represented by this stand of midddle-aged trees on the first terrace of a small creek floodplain in the patchwork of range plant communities where there is an intermixing of Western Cross Timbers and Grand Prairie in northcentral Texas. All trees were pecan whose tall straight trunks supported such woody vines as mustang grape, fiddleleaf or fringed greenbriar, and Alabama supplejack. Dominant understorey shrub was Carolina buckthorn. Most woody ground cover was the invasive, alien Japanese honeysuckle (Lonicera japonica), a treadful, horrid, etc. (adjectives fall short of vile description) woody weed. Major grasses included Canada wildrye, purpletop, Johnsongrass, and Texas wintergrass in that order. Major forbs were Baldwin ironweed (Vernonia baldwinii), and Indian plantain (Cacalia plantaginea). Erath County, Texas. April, early to mid-vernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-75 (Cross Timbers). Variant SAF 94 (Sycamore-Sweetgum-American Elm) that had former SAF designation of Sycamore-Pecan-American Elm. (Griffith et al., 2004). Closest biotic community designation of Brown et al. (1998, ps. 37, 38) was Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1, or more descriptively perhaps, a transition of this Oak-Hickory Series and Bluestem "Tall-grass" Series, 142.11, Plains Grassland 142.1 Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

404. Bottomland pecan forest- Higher stocking of pecan trees on bottomland produces a forest in contrast to the savanna oar open woodland form of this range type as shown above. All trees in these two photographs were pecan. Dense stocking resulted in tall, relatively straight boles in marked contradistinction from the widely spreading crowns of pecans with more spacing among trees. Pecans with stocking rates represented by forest vegetation presented in these two photographs produce wood and not fruit. This is the forest community of a lumberman not an orchardman, and the form of vegetation that develops in absence of fire. There was a well-developed woody vine layer that extended from ground level to tops of forest canopy which was comprised of mustang grape, rattanvine or Alabama supplejack, and fiddleleaf greenbriar. Carolina buckthorn grew as a smaller tree or larger shrub in the lower woody layer. The herbaceous layers included tallgrass species like Canada wildrye, purpletop, and the naturalized Johnsongrass along with composite forbs the two most common of which were Baldwin ironweed and Indian plantain. Much of the lower layer was composed of the invsive (and apparantely naturalized) Japanese honeysuckle. Erath County, Texas. April, early to mid-vernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-75 (Cross Timbers). Variant SAF 94 (Sycamore-Sweetgum-American Elm) that had former SAF designation of Sycamore-Pecan-American Elm. (Griffith et al., 2004). Closest biotic community designation of Brown et al. (1998, ps. 37, 38) was Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1, or more descriptively perhaps, a transition of this Oak-Hickory Series and Bluestem "Tall-grass" Series, 142.11, Plains Grassland 142.1 Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

405. Woody understorey of pecan bottom- From ground level and extending upward just shy of tree branching the range vegetation of a bottomland pecan forest was comprised of three woody vine species (mustang grape, fiddleleaf greenbriar, and Alabama supplejack) and the shrub or small tree, Carolina buckthorn. The wsidespread exotic invader, Japanese honeysuckle, was absent from this photoplot. The herbaceous portion of this forest vegetation visible in the background was presented in the next photograph. Erath County, Texas. April, early to mid-vernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-75 (Cross Timbers). Variant SAF 94 (Sycamore-Sweetgum-American Elm) that had former SAF designation of Sycamore-Pecan-American Elm. (Griffith et al., 2004). Closest biotic community designation of Brown et al. (1998, ps. 37, 38) was Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1, or more descriptively perhaps, a transition of this Oak-Hickory Series and Bluestem "Tall-grass" Series, 142.11, Plains Grassland 142.1 Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

406. Herbaceous layer (s) of a bottomland pecan forest- In addition to the woody vines of mustang grape, fiddleleaf greenbrier, and Alabama subblejack or rattanvine in right foreground and background of this photoquadrant important forbs of the herbaceous component included Indian plantain (left corner of foreground) and scattered smaller, apparently shade-stunted Baldwin ironweed. Many of the small green shoots with forb-like leaves were new sprouts of greenbrier and supplejack. Erath County, Texas. April, early to mid-vernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-75 (Cross Timbers). Variant SAF 94 (Sycamore-Sweetgum-American Elm) that had former SAF designation of Sycamore-Pecan-American Elm. (Griffith et al., 2004). Closest biotic community designation of Brown et al. (1998, ps. 37, 38) was Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1, or more descriptively perhaps, a transition of this Oak-Hickory Series and Bluestem "Tall-grass" Series, 142.11, Plains Grassland 142.1 Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

407. Pecan in winter- Examples of winter twigs, with emphasis on terminal bud, of pecan. The last photograph was a more detailed (closer-up) view of that shown in the third photograph. Upper bank above Bosque River, Erath County, Texas. January; winter dormancy.

408. Young leaves and catkins of pecan- A leader or shoot (first slide) and details of leaves and catkins (second slide) of pecan in West Cross Timbers of Texas. Erath County, Texas. March, pre-anthesis stage.

409. Fruit in the forest- Terminal end of a short shoot of native (hard-shell) pecan in the Grand Prairie of northcentral Texas. End of a short shoot on a pecan in the small bottomland forest range site along Richardson's Creek. Hunewell Ranch, Tarleton State University. Erath County, Texas. October; nearly fruit-ripe stage.

Eastern Cottonwood (Populis deltoides)-dominated bottomland forests

410. Floodplain forest of eastern cottonwood in the old bed of the Republican River— Seen here is a wetland forest site and a wetland range site (background and foreground, respectively). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). Cottonwood form ofK-89 (Northern Flood-Plain Forest).The forest site is an eastern cottonwood consociation (SAF Forest Cover Type 63) with an herbaceous understory of Indiangrass. The range site also has Indiangrass as the dominant herb, but Canada wild rye and prairie sandreed are important associated species. Dundy County, Nebraska. Populus deltoides Association (if and when such is recognized), Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). October, autumnal aspect. Dundy County, Nebraska.

411. Interior of a floodplain or wetland forest on the former bed of the Republican River— View inside the eastern cottonwood forest of the preceding slide, but showing local importance of scattered green ash (Fraxinus pennsylvanica). This vegetation is about 60 years old having begun development when the river flooded and changed course. The climax tree species were already established at time of that disturbance, but this perturbation removed most of the woody plants except the larger cottonwoods and ash allowing migration and establishment of prairie grasses as well as sedges and rushes more typical of the numerous marshes in the area. Dundy County, Nebraska. October, autumnal aspect. FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem).Cottonwood form of K-89 (Northern Flood-Plain Forest). SAF 63 (Cottonwood). Populus deltoides Association (if and when such is recognized), Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). October, autumnal aspect.

412. Semiarid gallery forest- Exterior view of a riparian zone forest of Rio Grande cottonwood (Populus deltoides ssp. wislizenii), netleaf hackberry (Celtis reticulata), Mexican walnut (Juglans major), and lanceleaf sumac (Rhus lanceolata) At outer (least mesic) edge of this gallery forest some of the more moisture-loving grasses formed a thick sward. The large cespitose grass was Wright's sacaton (Sporobolus wrightii) the dominant understorey species; Canada wildrye was the major associate species. The most common forb was horehound (Marrubium vulgare), a naturalized Eurasian weed. This range vegetation was in the Trans-Pecos Basin and Range vegetational area, the west Texas portion of the Chihuhuan Desert Region. It was in Chihuhuan Deserts 24 (ecoregion level III), Chihuhuan Montane Woodlands 24d (ecoregion level IV) based on Griffith et al. (2004). It was an example of mesic forest in a semiarid climate. FRES No 17 (Elm-Ash-Cottonwood Forest Ecosystem). No aqppropriate Kuchler unit for southern and.or southwestern cottonwood gallery forest. SAF 63 (Cottonwood). Populus deltoides Association (if and when such is recognized), Cottonwoow-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Trans-Pecos Basin & Range Vegetational Area: Chihuhuan Deserts- Chihuhuan Montane Woodlands Ecoregion 24d (Griffith et al., 2004). June, summer aspect. Calamity Creek, Jeff Davis County, Texas.

413. Great Plains floodplain forest- Eastern cottonwood and various species of willow formed a species-rich forest complete with prolific regeneration and a prominent herbaceous understorey. In these two slides the dominant herbaceous plant at local level ws American licorice (Glycyrrhiza lepidota), but grasses, sedges (Carex spp.), spike rush (Elocharis spp.) rushes (Juncus spp), and cattails (Typha domingensis) were the major understorey plants throughout most of the vegetation. Canada wildrye and bottomland switchgrass were most common with Johnsongrass (Sorghum halepense) the associate. Even the annual Eurasian grass, cheatgrass (Bromus tectorum) was common locally. FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). There is no Kuchler unit describing southern or southwestern cottonwood-dominated riparain forests. SAF 63 (Cottonwood). Populus deltoides Association (if and when such is recognized), Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Rolling Red Plains Vegetational Area: Central Great Plains- Red Prairie Ecoregion 27h (Griffith et al., 2004). June, summer aspect. Flood plain, North Fork of Red River. Wheeler County, Texas.

414. Foliage and fruit of prairie silvan queen- First of these two photographs was young shoots (two small seedlings or small saplings) of eastern cottonwood. They had been flattened by running water from recent spring (June) rains. (They straightened up.) Second photograph was the typical deltoid leaf with ripe fruit ( a capsule) of eastern cottonwood. Cottonwood capsules are described as comose, an adjective referring to having a coma (a tuft of hairs). Cottonwood derived its common name from the phenomenon of conspicuous, usually copious (and inconvenient) shedding of the comose ("cottony") fruit. All members of the Salicaceae are dioecious so that a tree is male or female and thus produces only staminate or only pistillate flowers. Of course only female plants produce fruit so that occurrence of the "messy fluff" comes only from the ladies. (Some range men find girls troublesome while others enjoy the mess.) Female catkins from this same plant were shown in the next photograph. Erath County, Texas. May.

415. Wild or American licorice (Glycyrrhiza lepidota)- Wild licorice is one of the more common legumes along watercourses throughout the Great Plains. It occurs in both the riparian zone and at outer (drier) margins of rivers, creeks, and floodplains. July, full bloom. Custer State Park, Custer County, South Dakota.

416. Licoriced sticks- Shoots with ripening legumes of wild or American licorice in the floodplain of the Platte River. The two closer-in (more detailed) shots when compared to similar photographs of other range species of Leguminosae served as examples of the great diversity in shapes, sizes, pubescence, etc. of legumes. Adams County, Colorado. Early July; advanced immature stage of fruit development.

417. Healthy regeneration of cottonwood and willow- One of the major problems in structure and sustainability of riparian and floodplain forests is reproduction under concentrated livestock grazing. This is especially so in arid and semiarid regions where livestock, especially cattle, tend to "hang in on water". The forest shown here was grazed periodically at light utilization (low degrees of use). There was extremely vigerous reproduction, both sexual and axexual, of both cottonwood and willow species. Generally members of the Salicaceae sprout vigerously particularily under moist soil conditions as on this subirrigated floodplain in the rolling red prairies. Successful reproduction of the salicaceous shrubs is a good indicator of proper management of the grazing resource and range ecosystem. Correct identifiction of willows (Salix spp.) to the correct species is a challenging task. It is often impossible to obtain positive identification without the catkins. There appeared to two and, probably, three species in this bottomland forest. This most likely "mixture" being some combination of black willow (S. nigra), peachleaf williw (S. amygdaloides), coyote willow (S. exigua), or sandbar willow (S. interior). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). No Kuchler unit for southern or southwestern cottonwood and/or willow riparian forest. SAF 63 (Cottonwood). Populus deltoides association (if and when such is recognized), Cottonwood- Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Rolling Red Plains Vegetational Area: Central Great Plains- Red Prairie Ecoregion 27h (Griffith et al., 2004). June, summer aspect. Flood plain, North Fork of Red River. Wheeler County, Texas.

418. Excellent regeneration of eastern cottonwood and willow species on a Great Plains bottomland forest- This is a close-up view of outstanding production of the "next generation" of eastern cottonwood (foreground) and willow (background) on a subirrigated river bottom in the Great Plains. Mixed prairie vegetation surrounded this wetland ecosystem.Trees were functioning as phreatophytes. FRES No.17 (Elm-Ash-Cottonwood Forest Ecosystem) .No Kuchler unit for southern or southwestern cottonwood and/or willow riparian forest. SAF 63 (Cottonwood) Populus deltoids association (if and when such is recognized), Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Rolling Red Plains Vegetational Area: Central Great Plains- Red Prairie Ecoregion 27h (Griffith et al., 2005). June, summer aspect. Flood plain, North Fork of Red River. Wheeler County, Texas.

419. Understorey of a cottonwood-willow bottomland forest- A lower woody layer of regenerating cottonwood and willow, an herbaceous layer of grasses and grass-like plants, and a multi-storied liana layer of greenbriar (Smilax sp.) and grape (Vitis sp.) added to the mature tree layer such that this range plant community formed a true forest structure on a wide expanse of river floodplain. Understorey graminoids included bottomland switchgrass, Canada wildrye, Johnsongrass, sedges, spike rushes, rushes, and cattail. This was another of the widespread and naturalized Johnsongrass. The most common forb was wild or American licorice. Proper range management with emphasis on light and periodic grazing/browsing (vs. the widespread practice of continuous heavy grazing) permitted perpetuation of this forest. This was an outstanding example of good stewardship and sustainability of range resources. FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). No Kuchler unit for southern and southwestern cottonwood and/or willow gallery forest. SAF 63 (Cottonwood). Populus deltoides association (if and when such is recognized), Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Rolling Red Plains Vegetational Area: Central Great Plains- Red Prairie Ecoregion 27h (Griffith et al., 2004). June, summer aspect. Flood plain, North Fork of Red River. Wheeler County, Texas.

420. Plains cottonwood bottomland forest in the far-western plains- Introduced in this photograph and treated further in subsequent slides was a forest range that developed along an *intermittant stream or, perhaps, interrupted stream (East Bijou Creek) in the Colordo Piedmont on which the dominant, the only, tree species was plains cottonwood (Populus deltoides ssp. monilifera). Plains cottonwood is another subspecies of eastern cottonwood native to the semiarid zone This forest range had been grazed by cattle for decades and was still in Good range condition class so as to approach the state of potential natural (climax) vegetation. There was limited reproduction of cottonwood, but this appeared adequate to maintain the uneven-aged population that dominated and defined this range plant community. Besides the upper woody (crown or canopy) layer there was a lower woody layer--though it was sporadic or "spotty" in its pattern of distribution-- made up of saplings and young poles of plains cottonwood. (This structural feature was shown in sebsequent slides.) A few seedlings of cottonwood were present in the herbaceous understorey. There were three herbaceous layers all of whch were dominated by grass species. These three were tallgrass, midgrass, and shortgrass layers so the understorey was mixed prairie. In fact this range vegetation could be visualized or even described as mixed prairie savanna with plains cottonwood as the woody element. Cottonwoods were dense enough, at least in numerous microhabitats where there were local populations (or, perhaps, individual stands would be a more apt term) that crowns typically interlocked. Thus, forest was the proper designation. Still, the overall or general vegetation could be conceived of as a savanna or, at least, as having a savanna-like physiogonomy, and a wetland savanna (or savanna wetland) at that. Portions of this range plant community actually comprised a gallery forest along the channel of this seasonally and spatially intermittant stream. Such a dispersion pattern and structure was shown in this photograph and the next photgraph where East Bijou Creek actually held water. Along other short reaches of this stream the creek channel was dry with only moister soil to distinguish the more-or-less interrupted riparian zone from the environment of the farther (outer) channel banks and adjacent floodplain. The herbaceous vegetation that was actually in the stream channel and extending outward on the nearest (lowr) banks was made up of prairie cordgrass (Spartina pectinata), bottomland ecotype(s) of switchgrass, and Canada wildrye (in that order of cover and biomass or standing crop). It should be noted that in addition to the light degree of use of tallgrass species there were a lot of cottonwood leaves and lower branches within browsing reach of cattle. This forest range was not being abused by the rancher who used. A well-earned tip of the hat and public praise to the faithful steward of this land. If the author had known he name of this commendable range manager he would have printed it. A note on stream terminology- The author was "only pasing through" this country and, as he did reside here, could not determine the nature of East Bijou Creek and use the most accurate designation. Specifically, the photographer could not distinguish whether it was an intermittent or intermediate stream. *Intermittent stream- "A stream or reach of a stream tht flows only at certain times of the year, as when it receives water from springs or from some surface source....A stream that does not flow continuously, as when water losses from evaporation or seepage exceed the available streamflow...Synonyms: temporaray stream, seasonal stream" and *Interrupeted stream- "A stream that contains perennial reaches with intervening intermittent or emphemeral reaches, or a stream that contains intermittent reaches with intervening ephemeral reaches" (Wilson and Moore, 1998). East Bijou Creek, Elbert County, Colorado. Mid-July (estival aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). No Kuchler unit for southern and southwestern cottonwood forest. No SRM. SAF 63 (Cottonwood). Populus deltoides association (if and when such is recognized), Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Southwestern Tablelands-Piedmont Plains and Tablelands Ecosystem, 26e (Chapman et al., 2006).

.

421. A little farther downstream- At another short reach with surface water of the channel of an intermediate or interupted stream (East Bijou Creek) along which a forest of plains cottonwood with a n herbaceous understorey had developed. In the reach of the creek viewed here the stream chaannel was smaller (narrower and shallower) so that water persisted for shorter periods than in the reach shown immediately above. Range vegetation along this segment of East Bijou Creek was more of a mixture of aquatic and also of less mesic grass species than in the reach with a bigger channel and greater riparian zone surface seen above. In this stream segment major range plants were Torrey's rush (Juncus torreyi) and western wheatgrass as well as Canada wildrye and prairie cordgrass which were relatively more abundant around the larger area of water shown in the preceding slide. Cheatgrass or downy bromegrass (Bromus tectorum) was present in minor down to trace amounts. Both smaller (presumedly younger) and full-sized adult trees of plains cottonwood were present in this portion of the floodplain forest. Only range vegetation of the general riparizn zone was described in this and the preceding photograph. Composition and structure of the floodplain away from the immediate riparian zone was treated in the slides and their captions that followed immediately below. East Bijou Creek, Elbert County, Colorado. Mid-July (estival aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). No Kuchler unit for southern and southwestern cottonwood forest. No SRM. SAF 63 (Cottonwood). Populus deltoides association (if and when such is recognized), Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Southwestern Tablelands-Piedmont Plains and Tablelands Ecosystem, 26e (Chapman et al., 2006).

422. Along a dry reach of the interupted stream- Range vegetation of a bottomland forest dominated by plains cottonwood along East Bijou Creek, an interrupted or intermittent (whichever it was) stream. This view across across (at a rough right angle to) the stream channel rather than a headon or inline view as in the two preceding photographs. Different age/size classes of trees were present showing this to be an uneven-aged population of plains cottonwood. There were relatively few cottonwood seedlings, but their number appeared adequate (though barely) to maintain the only tree species in this plant community. Density and spatial arrangement (dispersion pattern) of cottonwood varied from closed canopy forest along the riparian zone to such wide (sparse) spacing of trees as to constitute a savanna on the outskirts of this bottomland range plant community. Cottonwoods grew (occurred) as individual mature adult trees or small groups of two or three trunks up small groves of eight to ten trees. Groves characteristically included pole-sized as well as fully grown trees. These larger groups were closer to the stream (generally in the riparian zone) and tree density declined with distance from the creek until the savanna spacing at outer border of the bottomland community. Dead wood at base of the adult cottonwood across the creek was mostly debris washed in and lodged against the trunk although a minor amount of this wood pile was branches from the crown of the tree that served as a barrier to water-borne material on this floodplain. Although it was not obvious in this wide view there were three layers comprising the herbaceous understorey: tallgrass, midgrass, and shortgrass that were dominated by prairie sandreed, western wheatgrass, and buffalograss, respectively. Downy brome or cheatgrass was present (as it alway is) but it was minor on this range that was under superb management (see shortly below). Forbs present included wild licorice, the most common and important; prairie coneflower or Mexican hat (Ratibida columnifera); showy milkweed (Asclepias speciosa); annual sunflower (Helianthus annus); four o'clock (Mirabilis linearis); prairie sagewort (Artemisia frigida); western ragweed (Ambrosia psilostachya) and kochia (Kochia scoparia). There were sparsely scattered plants of plains pricklypear (Opuntia polycantha). Range condition class was Good. Cattle had most likely grazed this range for decades, certainly over the last several years to the present time as proven by dung piles of varying ages. In all these photographs there were cottonwood leaves and leaders within reach of cattle indicating that there had not been heavy browsing of of this species which has relatively high palatability to cattle. (High preference by cattle for Populus deltoides and the relative ease with which this palatable species is browsed out was described in detail below.) Presence of this palatable browse and fairly high abundance (author's subjective judgement based on cover, density, vigor, etc.) of prairie sandreed and prairie cordgrass along the creek was undeniable evidence that this range had not been abused (overgrazing/overbrowsing had not taken place) in the recent history of use by livestock. Accolades to the range manager. East Bijou Creek, Elbert County, Colorado. Mid-July (estival.aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). No Kuchler unit for southern and southwestern cottonwood forest. No SRM. SAF 63 (Cottonwood). Populus deltoides association (if and when such is recognized), Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Southwestern Tablelands-Piedmont Plains and Tablelands Ecosystem, 26e (Chapman et al., 2006).

423. On the higher and outer bank of the interrupted stream- View looking up the channel and along the bank of East Bijou Creek, an interrupted or intermittent (whichever it was) stream, in the semiarid Colorado Piedmont. From the physiogonomic and structural perspectives, an upper woody layer of plains cottonwood dominated this bottomland range vegetation. Tree density varied from that of a gallery forest in the riparian zones which remained wetter later into the summer to a savanna at outer margins of the floodplain plant community. This range community was, in essence, a mixed prairie with a tree element that varied greatly in tree density and dispersion pattern. From perspective of annual biomass production (standing crop productivity) the dominant plant species of this range ecosystem was almost assuredly prairie sandreed. Western wheatgrass was the overall community associate. Switchgrass held this honor within the riparain zone in which the dominant herbaceous species was prairie cordgrass. Thus it was that a tallgrass species was the herbaceous dominant of both the wetland (riparian zone) and the greater floodplain. Dominance of the greater floodplain (beginning at top of stream bank) by prairie sandreed and the close association of western wheatgrass, the major midgrass species, was depicted in this photograph. Buffalograss and blue grama were not conspicuously featured, but they were certainly well-represented. Mesic tallgrass species (notably prairie cordgrass and switchgrass) that dominated the lower stream channel were absent. Forb species included wild licorice, generally the most important; prairie coneflower of Mexican hat, showy milkweed, annual sunflower, kochia, prairie sagewort, and narrowleaf four o'clock. There were sparsely scattered plants of plains pricklypear, but nothing approaching density and cover to form even an interrupted shrub layer. East Bijou Creek, Elbert County, Colorado. Mid-July (estival aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). No Kuchler unit for southern and southwestern cottonwood forest. No SRM. SAF 63 (Cottonwood). Populus deltoides association (if and when such is recognized), Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Southwestern Tablelands-Piedmont Plains and Tablelands Ecosystem, 26e (Chapman et al., 2006).

424. A savanna form on the outer boundary- At outside margins of plains cottonwood-dominated bottomland range vegetation the plant community was actually a savanna or, at best from a crown canopy criterion, a woodland rather than a forest. This lower stand density or wider tree spacing was in contrast to the gallery forest that developed in the riparian zone of this interrupted stream as was shown above. Downed limbs immediately beneath cottonwood boles were a combination of those that fell from directly overhead off of senescing adult trees as well as those that had been carried by flood water to rest against the trunks. Such debris offered protection for the all-too-scant number of cottonwood seedlings available to replace their aging elders. The understorey consisted of three distinct herbaceous layers of tallgrass, midgrass, and (to a lesser degree) shortgrass species, the dominants of whch were, respectively, prairie sandreed, western wheatgrass, and buffalograss (blue grama was less abundant having lower relative cover than buffalograss). Forbs included wild licorice, the most common and important; prairie coneflower or Mexican hat; showy milkweed, narrowleaf four o'clock, prairie sagewort, annual sunflower, and kochia. East Bijou Creek, Elbert County, Colorado. Mid-July (estival aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). No Kuchler unit for southern and southwestern cottonwood forest. No SRM. SAF 63 (Cottonwood). Populus deltoides association (if and when such is recognized), Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Southwestern Tablelands-Piedmont Plains and Tablelands Ecosystem, 26e (Chapman et al., 2006).

425. Village meeting- Local assemblage of range plants on plains cottonwood-dominated bottomland range on floodplain of an interrupted or, possibly, an intermittent stream in the semiarid zone. Dominant plant species was prairie sandreed. Other range plant species included prairie sagewort, western ragweed, Mexican hat or prairie coneflower, and wild licorice. The second (vertical) slide was a closer-in, more detailed view of the left foreground of the first slide. This second photograph presented a better view of shoots of praiirie sandreed. East Bijou Creek, Elbert County, Colorado. Mid-July (estival aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). No Kuchler unit for southern and southwestern cottonwood forest. No SRM. SAF 63 (Cottonwood). Populus deltoides association (if and when such is recognized), Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Southwestern Tablelands-Piedmont Plains and Tablelands Ecosystem, 26e (Chapman et al., 2006).

426. At base- Basal portion of shoot of prairie sandreed with parts of sheath and blade of the four lowermost leaves. More details were presented in the next subsequent slide. This shoot was growing on East Bijou Creek, an intermittent or interrupted (whichever one), in the Colorado Plateau, the physiographic unit just west of the High Plains or Llano Estacho (Staked Plains) of the Great Plains province. Elbert County, Colorado. Mid-July.

427. Unit of a reed- Phytomer--fundamental, repeating unit of a grass shoot including upper half of a node, internode, and lower half of next highest node complete with leaf--of prairie sandreed (first slide) and details of culm, sheath, and colar of this phytomer (second slide). Floodplain of East Bijou Creek, an interrupted or, maybe, an intermittent stream in the semiarid tableland of the Colorado Piedmont. Elbert County, Colorado. Mid-July.

428. Atop the reed of a reed- The panicle--the entire structure--of prairie sandreed (first slide), upper or distal portion of this panicle (second slide), and one branch of this panicle at onset of anthesis (third slide). East Bijou Creek, Elbert County, Colorado. Mid-July; initiation of anthesis.

429. Unfulfilled florets- Spikelets of prairie sandreed showing florets at pre-anthesis stage. There is only one floret per spikelet in this species and the upper glume is considerably longer than the lower (Shaw, 2008, p.90). Both of these features were presented in these two photographs. East Bijou Creek, Elbert County, Colorado. Mid-July; initiation of anthesis.

430. Cluster of racemes- Inflorescences of prairie cordgrass growing on a plains cottonwood-mixed prairie savanna that developed along an intermittent stream in the Colorado Piedmont. The inflorescences of Spartina species are confusing. Traditionally the branches with units of spikelets aligned along them were interpreted as racemes or, maybe, spicate racemes, but as these spikelets are sessile this view was at least somewhat inconsistent with standard definitions. Hitchcock and Chase (1950, p. 508) regarded the Spartina flower cluster as being made up of "... appressed or sometimes spreading spikes racemose on the main axis". Later, more detailed (sophisticated) descriptions of grass inflorescences recognized specialized panicle inflorescences (Highnight et al., 1988). Using this scheme, Stubbendieck et al. (1992, p. 107) described the inflorescence of prairie cordgrass as a "panicle of 6-40 spicate pirmary unilateral branches". Shaw (2008, p. 183) described the inflorescence of prairie cordgrass as a panicle with racemosely arranged, appressed, alternate branches. East Bijou Creek, Elbert County, Colorado. Mid-July; peak-bloom stage (full anthesis).

431. Anthers, anybody?- Two close-up views of racemes in full anthesis on prairie cordgrass. The first photograph presented a standard side-view perspective of the raceme while still on the plant whereas the second photograph gave a ventral view of a raceme that was possible only by removal from the shoot. (The latter was laid on cattle dung in situ on a floodplain cottonwood-mixed prairie range that developed along an intermittent stream.) There are often so many spikelets (one-flowered and sessile) on the racemose, unilateral branches that these floral units overlap (ie. imbricate spikelets). The Spartina species often produce some of the greatest number of anthers of any grass genera native to North America which in combination with imbricate spikelets produces an extremely showy array of exerted anthers. That characteristic was seen in these photographs. East Bijou Creek, Elbert County, Colorado. Mid-July; peak-bloom stage (full anthesis).

432. Hanging lightly- Drooping leaders (woody shoots) with pendulous clusters of fruit on eastern cottonwood (Populus deltoides subsp. monilifera). The delta-shaped leaves, obvious basis of the specific epithet,.is a key identifying feture of this species. Vines (1960, p. 91) described leaf shape of P. deltoides as "broadly deltoid, margin crenate-serrate, apex abruptly acute or acuminate, base truncate to heart-shaped or abruptly cunete". Larimer County, Colorado. Mid-June (late spring), seed-shatter stage.

433. Lightly but effectively- Fruit (first slide) and individual seeds (second slide) of eastern cottonwood. Eastern cottonwood "wrote the book" on anemochory or aerochory, dispersion of plant disseminules or propagules by wind. Many prairie plants evolved (natural selection, survival of the fittest) anemochory as a means of effectively and efficiently dispersing pollen, but cottonwood went a step farther and has one of the best adaptations to dispersal of its fruit and seed via aerochory of any woody plant in North America. The liguliferous composites like thistles and dandelions have nothing on Populus species. Eastern cottonwood is a dioecious species. Its fruit is a capsule, numerous of which are borne in racemes (first slide) and each of which consist of 10 to 30 seeds (second slide) with each of these being equipped with pubescence in the form of numerous, long, cottony fibers (Vines, 1960, p. 91). It is the cottony pubescence that permits cottonwood seeds to become air-borne so as to float and finally settle by gravity into every available "nook and cranny". Larimer County, Colorado. Mid-June (late spring), seed-shatter stage.

434. Palatable resprouts- Root sprouts (suckers) from a felled, senescing eastern cottonwood on a prairie in the western Ozark Plateau. The three sets of root shoots presented here (first and second photographs) were two-year-old suckers or heterophyllous shoots produced from lateral roots (third photograph) of a relatively old, "half-rotten" eastern cottonwood after it was felled for fuel wood. The shoots, some of which in these photographs were over eight feet in height, had been completely defoliated (100% of leaves removed) by commercial beef cattle (cows and calves of mixed breeding) in both the first and second years of shoot development. The small leaves present on shoots at time of photographs were current-season's regrowth (early part of current--second--warm growing season). Several (a high proprotion) of cottonwood root suckers were broken off by browsing cattle so that this extreme or severe degree of use included both eating of leaves and upper twigs of current growing season as well as complete breaking off of many root suckers that were the production of two growing seasons. Several of the broken shoots and branches were visible in some of the photographs presented here and immediately below. Cattle defoliation in the first growing season was heavy, but not total as in the second season of growth. In the second growing season many of the first and current year's root shoots were broken off at their point of origin on the lateral roots. This was done by cattle who "rode down" the root suckers in order to reach the uppermost leaves and terminal buds (ie. cattle ate all leaves and much of the upper parts of shoots, terminal twigs, plus broke off entire shoots by the browsing technique of "riding down"). "Riding down" is the feeding behavior whereby browsing animals reach up as high as they can on plants and then commence walking forward so that forage or browse plants bend down beneath the animals' weight and pass progressively beneath the feeding animals' brisket and belly until the animals have fed as far as the plant will bend to within reach of the animals' mouth. In the process, some of the brittle shoots and branches are broken off while the more limber shoots spring back-- minus leaves, fruit, buds, twigs, etc. that were consumed by the animals. In this feeding pattern animals can remove plant material at much greater heights than they could otherwise reach by merely streatching their necks and extending their tongues. Herbaceous plant species growing adjacent to and beneath eastern cottonwood shoots was mostly the introduced pasture grass, tall fescue (Festuca arundinaceae), which has naturalized across most of the greater Ozark Region and had self-seeded onto this former tallgrass prairie. The second-most common herbaceous species on this local habitat was Japanese chess or Japanese brome (Bromus japonicus), a naturalized Eurasian annual grass. It appeared that none of the herbage of these two non-native grasses had been eaten by cattle although considerable quantities of biomass had been trampled. Cattle seemingly had quite a browsing bout on this spot as quantities of dung were left. There was no evidence of browsing by deer, rodents or rabbits. Total herbivory (=herbivorous action) that did and would continue to influence regeneration of eastern cottonwood on this pasture included at least four components: 1) complete removal of leaves from cottonwood shoots, 2) total elimination of some cottonwood shoots by breakage at ground level (ie. total loss of some two-year old trees or two growing seasons of plant regeneration), 3) feeding selectivity that severely injured or setback cottonwood while at the same time having much less defoliation impact on annual and perennial grasses (both non-natives), and 4) deposition of dung with importation of more potential plant nutrients than was recycled or otherwise generated on this local environment. Question to the "village idiot" (so that he can be "above average"): "will grasses or cottonwood derive most benefit for plant growth and/or survival from the combined influences of cattle feeding?" Eastern cottonwood (both eastern and plains varieties), like most other members of the Salicaceae, readily coppices or resprouts following major injury or defoliation. Fowells (1965, ps. 516, 521) and Burns and Honkala (1990, vol. 2, ps. 533, 539) described vegetative reproduction in eastern cottonwood. Eastern cottonwood sprouts vigerously from both stumps and roots. In the pattern of most sprouting species, vegetative (asexual) reproduction declines with increaseing age of trees while vegetative propagation, both natural and atrtificial, is vigerous in young plants. Fowells (1965, p. 516) and Burns and Honkala (1990, vol. 2, ps. 533) cited research which found that eastern cottonwood (P. deltoides var. deltoides) stump-sprouted up to ages of 25 years. Heterophyllous shoots (root sprouts or suckers) shown here were from a much older tree. Over half of the trunk (beyond the center growth rings) of this eastern cottonwood had rotted away so that it was impossible to accurately determine tree age. (Rotting appeared to have begun around an old fire scar.) Recollections by the author and his conversations with elderly observers firmly established age of this felled tree in excess of 80 years. Root suckering began in early spring following sawing of the old tree in the preceding winter. On various occasions this author has observed resprouting by various kinds of cottonwood (Populus spp.) in larger and older tindividual trees following sawing, bulldozing, f lood breakage, etc. Obviously resprouting in cottonwood is not limited to trees that are younger than a quarter century. It was also obvious that regeneration of eastern cottonwood depends upon protection from overbrowsing. It was possible that some of the herbivory described above benefitted (or had potential to benefit) eastern cottonwood. Dunging provided extra plant nutrients, especially nitrogen, available to surviving cottonwood shoots. Dung might also benefit grass proportionately more than cottonwood so that increased competition from grass resulting from soil fertilization was detrimental to cottonwood. It was conceivable that breakage of some cottonwood root sprouts (suckers) had the effect of beneficial prunning which would enable surviving shoots to grow larger faster and become too large for cattle to "ride down" and thereby escape most further browsing impacts (other than continued dunging if and when cattle rested in shade of growing cottonwoods). Overall, however, and up to the point of time recorded here, even the village idiot made 100% of this pop quiz by explaining (in simple talk) that the browsing shown here was excessive and could result in death of resprouts and ultimate loss of eastern cottonwood from this pasture. Ottawa County, Oklahoma. June.

435. Overutilization of palatable eastern cottonwood by cattle- Excessive (extreme or severe) defoliation of two-ear old root suckers of eastern cottonwood on a prairie in the Ozark Plateau. Beef cattle (crossbred cows and calves) removed all leaves and many shoot tips (including developing buds) from these heterophyllous shoots by early to mid-spring of the second growing season. This necessitated production and development of a new set of leaves (the leaves present in these and preceding photographs). Numerous branches were broken from shoots that survived "rding down" browsing by cattle, and lots of the two-year old shoots (root suckers) did not survive the aggressive feeding by cattle. This was an example of overuse which, if continued, might well result in killing and loss of eastern cottonwood from this pasture. These slides demonstrated the importance of protection of young eastern cottonwoods that are extremely palatable (even to animal species that are grazers more than browsers) from excessive browsing. Regeneration of eastern or plains cottonwood such as that shown above in a plains cottonwood-willow bottomland forest would not have been possible with continued overuse at the severity shown here and in the three immediately preceding slides. If degree(s) of use showh here continued for just a few more years (number unknown; would vary depending on many factors) the eastern cottonwood will die out on this pasture. The old felled cottonwood tree described in the preceding caption will be dead afterall. The main lesson from these photographs was the concept of proper degree of use (the first and most important of the Four Cardinal Principles of Range Management), often expressed as Proper Use Factors. Secondary lessons from these slides involved range animal behavior, including both the phenomenon of "riding down" and feeding selectivity. A degree of defoliation approaching "anililation" of eastern cottonwood coupled with non-eating of tall fescue and Japanese brome (though with some trampling and covering with dung of these grasses) provided a "textbook example" of feeding selectivity, the threeway interaction of animal preferences for certain plant species, relative palatability of different plant species, and availability of herbage and browse of certain plant species. Ottawa County, Oklahoma. June.

436. It'll do its part and its best, but eastern cottonwood cannot rise from the dead indefinitely- Close-up view of one of the severely browsed shoots that suckered roots of a felled eastern cottonwood described above. All leaves are regrowth that replaced earlier full-grown leaves that were eaten by browsing beef cattle. These were some shoots that survived extreme degrees of browsing. Many shoots did not survive. (This exterme defoliation might be expected from overpopulations of browsers like deer, but this was from just "plain ole cows".) If the rangeman takes care of the feed resource it will take care of him. If this cattleman wanted natural replacement of a nice shade tree a drastic change of management was in order. A quicky, makeshift exclosure of wovn wire would protect these young shoots until they could grown beyond reach of the cattle. If this cattleman had the goal of elimination of eastern cottonwood from this pasture he had a good start toward that objective. In other words, if eastern cottonwood was regarded as a brush species (a woody noxious plant) and a single-species stand of tall fescue was the desired result then cattle browsing appeared to be a very effective method of biological control to reduce (perhaps totally eliminate) this woody species from this grazing (or haying) unit of land. As it happened such was not the case. This cattle-owner was simply running cattle for tax purposes and social status (ie."dodging taxes" and "bragging rights"). This photographic case study was used to illustrate how excessive degree of use and/or general imporper animal browsing can prevent or greatly reduce regeneration of eastern cottonwood. Degree of use on eastern cottonwood would almost certainly have been much less if cattle-grazing had been restricted to late autumn through winter-early spring when cottonwood was dormant and nutritive value and palatability of the naturalized cool-season grasses was higher. Ottawa County, Oklahoma. June.

437. Standing in river water- Mature Rio Grande cottonwood trees were flooded by water from the Rio Grande River that flowed a short distance from this location. River water rose above the ground level via subsurface flow. Ponded water was visible because manicuring with power tools for a picnic area (in a national park) eliminated the understorey of what was (would have been) a bottomland or gallery forest in the Chihuhuan Desert. The herbaceous layer in this radically modified vegetation consisted almost exclusively of introduced perennial grasses including bermudagrass (Cynodon dactylon), King Ranch bluestem (Andropoogon ischaemum= Bothriochloa ischaemum), and Dallisgrass (Paspalum dilatatum). Natural vegetation of diverse structure, including a shrub layer below the tree layer and the invasive Eurasian salt cedar (Tamarix chinensis), was present in the background. FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). No Kuchler unit for southern or southwestern cottonwood and/or gallery forest.. SAF 63 (Cottonwood). Populus deltoides association (if and when such is recognized), Cottonwood-Willow Series in Southwestern Riparian Deciduous Forest botic community of Brown et al. (1998). Trans-Pecos Basin & Range Vegetational Area: Chihuhuan Deserts- Low Mountains & Bajadas Ecoregion 24c (Griffith et al. 2004). June, summer aspect. Floodplain of Rio Grande (water mostly from Rio Concho), Big Bend National Park, Brewster County, Texas.

Some "Odd Lots" Forests

438. Red oak-linden (basswood) mesic forest— This community was described by Weaver (1965, p. 14-17) as the most mesophytic forest in Nebraska. It extends from draws to north slopes and is one of the best-developed forest types at the edge of the vast grasslands of the continental interior. Ironwood or hop hornbeam (Ostrya virginiana) is the dominant species of the shrub layer, but Virginia creeper and wild grape vines climb trees to give an aboreal shrub component. The Missouri Natural Areas Committee (1987) described a similar community as mesic upland forest. The Society of American Foresters (1980) did not list or describe this type, but Braun (1950, p. 234) interpreted it as comparable to the red oak-sugar maple-basswood community in Pennsylvania and in the Mississippi Valley section of the oak-hickory region in Nebraska.Westveld (1939, pgs. 46, 51, 67-72, 77, 86, 114-117, 267, 268) recognized the red oak-basswood-white ash forest type for the New England white pine and central hardwood regions as well as for the mid-Atlanic oak region. Thus it seemed likely that the red oak-basswood mesophytic forest which occurs at the western-most edge of the immense eastern deciduous forest formations and the eastern origins of the vast interior grasslands is a variant of SAF 26 (Sugar Maple-Basswood) which was described as grading into other cover types throughout the deciduous forest zone.No obvious FRES or Kuchler designation, but basswood is a dominant of K-90 (Maple-Basswood Forest) to which it bears some taxonomic kin based on the Braun interpretation. Western Corn Belt Plains- Nebraska-Kansas Loess Hills Ecoregion, 47h (Chapman et al., 2001). Schramm State Park, Sarpy Cass County, Nebraska.August, estival aspect.

439. Floodplain (gallery) forest of Wichita River- Exterior view of a forest community composed of discrete zones of vegetation along the terrace of a meandering river in the . This was a gallery forest that lined the bank and terrace of a river bordering on the mixed-tallgrass prairie transition. Immediately adjacent to the river was a vegetation zone consisting of green ash as a tree layer with an understorey of Canada or nodding wildrye. Also in the tree layer were occasional individuals of eastern cottonwood, hackberry or sugarberry (Celtis laevigata var.reticulata), pecan, and very rarely, black willow (Salix nigra). The herbaceous understorey was "pure" or "solid" wildrye with essentially no other species present so as to constitute a consociation. Green ash approached consociation status. The vegetation zone outer (farther from the river bank) from the green ash-nodding wildrye zone was eastern cottonwood with cottonwood leaves forming a mulch or litter layer devoid of plant species except the lianas, poison oak and trumpet creeper (Campsis radicans). Both of these woody climbers reached tops of cottonwoods. Poison oak was present as both climber and "regular form" of shrub. Outward from the cottonwood zone was one dominated by black willow. The fourth or outermost (from the river bank) vegetation zone was composed of the arboreal (and almost exclusive) dominant, western soapberry (Sapindus saponaria var. drummondii= S. drummondii) with an herbaceous understorey dominated by western wheatgrass (Agropyron smithii) with purpletop (Tridens flavus) as the associate. Buffalograss (Buchloe dactyloides) and Texas bluegrass (Poa arachnifera) were present also, the former being an occasional "spot" (microsite) dominant. Lucy Park, Wichita County, Texas. October. Estival aspect. FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). Combination of SAF 63 (Eastern Cottonwood) and green ash-dominated variant of SAF 93 (Sugarberry-American Elm-Green Ash). Loamy Bottomland range site. Variant of Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Central Great Plains- Broken Red Plains Ecoregion, 27i (Griffith et al., 2004).

440. Deciduous riparian forest on terrace of Wichita River- Two interior views of the floodplain or gallery forest shown from the exterior in the preceeding photograph. The arboreal layer of this bottomland community was dominated by green ash with scattered trees of eastern cottonwood, sugarberry, and pecan making up trace amounts of the canopy cover. The herbaceous understorey layer was a consociation of Canada wildrye. There were no arborescent shrubs or woody climbers in this vegetation zone which bordered the river. Reason(s) for lack of adult trees and presence of even-age structure of immature trees could not be determined. There were no stumps, trunks, or other remains of previously live trees. Numerous sprouts from young stumps browsed by beaver (Castor canadensis) were present (see below).- Lucy Park, Wichita County, Texas. October. Estival aspect.FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). Green ash variant of SAF 93 (Sugarberry-American Elm-Green Ash). Loamy Bottomland range site. Variant of Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Central Great Plains- Broken Red Plains Ecoregion, 27i (Griffith et al., 2004).

441. Leader of green ash- Leaves and fruit along an internode of green ash. Length of internode was approximately 20 inches which indicated potential for annual increment of green ash on a bottomland site in a subhumid preacipitation zone. Lucy Park, Wichita County, Texas. October. Estival aspect.

442. Leaves and fruit of green ash- Fruit type is a samara (winged achene). Lucy Park, Wichita County, Texas.October.

443. Upper reaches of a deciduous bottomland (gallery) forest along Wichita River- Western soapberry dominated this forest community growing on the upper bench or reach of the river terrace. This was the uppermost zone of the riverine forest that was comprised of green ash, eastern cottonwood, hackberry, and pecan adjacent to the meandering river. At this greater elevation and less mesic soil western soapberry was the only tree species. Shrubs were extremely scarse with infrequent individuals of greenbriar (Smilax bona-nox) and skunkbush sumac (Rhus trilobata= R. aromatica). Note prolific regeneration of western soapberry. Lucy Park, Wichita County, Texas. October. Estival aspect.Perimeter of FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosytem). Given the western soapberry consociation and absence of other tree species in this local vegetational zone Kuchler and SAF units were deemed inappropriate; this vegetation was the edge of such units designated above for the bottomland forest. Loamy Bottomland range site. Central Great Plains- Broken Red Plains Ecoregion, 27i (Griffith et al., 2004).

444. Western soapberry zone of bottomland deciduous forest- Plant community of a gallery forest along the meandering Wichita River in an area transitional between mixed and tallgrass prairie in the Rolling Red Plains. Except for an incidental individual shrub like greenbriar or skunkbush sumac soapberry was the only woody species present. The understorey was primarily western wheatgrass, purpletop, little bluestem, and the naturalized Johnsongrass (Sorghum halepense). Buffalograss and hairy tridens (Tridens pilosus= Erioneuron pilosum) were locally abundant and added a shortgrass component so as to form a two-layer herbaceous understorey of a patchwork appearance. Lucy Park, Wichita County, Texas. October. Estival aspect. Perimeter of FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). Kuchler and SAF units were not descriptive but were given above for the more mesic and typical plant communities of this range cover type. Central Great Plains- Broken Red Plains Ecoregion, 27i (Griffith et al., 2004).

445. Western soapberry (Sapindus saponaria var. drummondii= S. drummondii)- Young plants of western soapberry in early fall (October, but still estival aspect) in fruit. Lucy Park, Wichita County, Texas.

446. Leader of western soapberry- Compound leaves and large inflorescence of western soapberry. This specimen was thriving in the western Ozark Plateau. July. Newton County, Missouri.

447. Infloresscence of western soapberry- An immense flower cluster of western soapberry on a tree that grew on the easternmost extension of this species' range. July. Newton County, Missouri.

448. Leaves and fruit of western soapberry- Lucy Park (just above Wichita River), Wichita County, Texas.

449.. White Christmas in the Ozarks- Snowfall (especially of major accumulation like a half foot or more) is rare in the oak-hickory forests of the Ozark Mountains. In fact, it is not unusual for there to be winters without snow and with winter precipitation being limited to rain or the treacherous "winter mix" of rain, freezing rain, sleet, and snow (ice storms). The "winter wonderland" of wet snow and an accumulation of 10 inches shown in these photographs demonstrated the ever-changing weather of the Ozark Region and the necessity of some winter adaptation for it's species. Besides it was "kinda purty". Post oak and black hickory were dominant tree species with northern red oak and black cherry the major associates of the canopy layer. Understorey was dominated by buckbrush and poison oak; Virginia creeper formed a uniform intermediate layer extending from ground level to tops of trees while wild grapes (Vitis spp.) were local dominants of the intermediate layer. In early spring the herbaceous layer was usually composed exclusively of Mayapple (Podophyllum peltatum) colonies. All plant life had settled in for a short winter nap in this scene which had all but entirely melted within 24 hours of these photographs. Ottawa County, Oklahoma. Christmas Day, 2002.