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The fundamental and practical distinction between coniferous and deciduous forests is useful (and was used herein), but precise, non-arbitrary "lines" are impossible when presenting and discussing forest range types in the eastern half of the continent. This is especially the case when climax or potential natural vegetation is used as the basis for forest types (ie. when cover types, or the more specific management cover types, are discussed as being more or less synonymous with permanent forest types). As discussed in detail below, the epic work of Lucy Braun (1950) is still the definitive basis for the ecological discussion and classification of those North American forests which extend from the Atlantic Coast to slightly beyond the Missouri and Mississippi River drainages. Braun (1950) included all the coniferous forests (forest types, regions, etc.)-- the generic "southeastern pine region"--as part of her one Deciduous Forest Formation.
The forest range types included in the following section include coniferous, deciduous, and mixed coniferous-deciduous forests. This is confusing but unavoidable given the nature of the vegetation and the standard understanding (the Braun interpretation) of ecological relations and classification of this forest vegetation. Most of the southeastern pine types presented are management cover types maintained silviculturally as more economically valuable coniferous forests rather than as the climax mixed hardwood-pine forest types. In other words, efforts were made to fit the Society of American Foresters (1980) cover types with the climax types of Braun (1950) and the potential natural vegetation units of Kuchler (1966).
The major forest communities or forest zones of eastern North America are broad or wide in their spatial patterns unlike the narrow zonation characteristic of the forests of western North America. The "young" mountains of the western part of the continent are taller (in fact, still getting taller) and as a result have more elevation-based zonation of vegetation than do the geologically older and more eroded (lower) eastern mountains such as the Applachians or Ozarks. So too, are the soils of the Atlantic Coast more zonal (ie. major soil units are larger or broader in spational dimension like those of the vast continental interior whereas soils of the Rocky Mountains and the Pacific Slope ranges are more of the intrazonal spatial scale. See for illustration the national soil map of dominant soil orders and suborders (Soil Survey Staff, 1998).
Vankat (1979, p. 137) wrote that relief within the eastern deciduous forest "is quite variable" yet earlier Vankat (1979, p. 41) had also correctly noted that "low hills" were characteristic of much of this deciduous forest region. Again, contrast this with the extreme physiography of the Rockys or Sierra Nevada-Cascade Ranges.
The classic and still-definitive work on forests of eastern North America (approximately east of the 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:
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.
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".
1. 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).
2. 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).
3. 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.
4. Hemongous hickory- A huge white hickory (Carya tomentosa) lived long and healthy in an oak-hickory forest on a bluff above a small stream in the Ozark Plateau. Redbud (Cercis canadensis) and flowering dogwood (Cornus florida) had just begun to bloom but these shrubs of mature statue served to show size of this magnificant specimen.
Above Modoc Creek, Ottawa County, Oklahoma. April: could there be any doubt that this was the vernal aspect?
5. Pignut, bitternut, or (sometimes) white hickory (Carya cordiformis)- Identification of the hickory species besides the obvious ones like pecan or shagbark and black hickories is often a frustrating undertaking. This is especially the case in forests having several Carya species growing in close proximity such that leaves and nuts of the species are intermixed and where the trees are so large that identification relies heavily on bark. The immense specimen seen here was a good example of the bark of the pignut or bitternut hickory. This species grows on a variety of habitats including a diversity of soils. As such, pignut hickory grows to different sizes and with various trunk and crown shapes. It frequently attains it's greatest size on moist yet well-drained soils on upslope drainages such as the one seen here growing high on a bluff above a stream in the Springfield Plateau portion of the Ozarks.
Along Lost Creek, Ottawa County, Oklahoma. April.
6. Leaves and hickory nuts of pignut or bitternut hickory, one of the more common Carya species in the Ozark Plateau. Ottawa County, Oklahoma. July.
7. Reaching to the sky- Another large hickory presented to represent its species was this specimen of shagbark or shellbark hickory (Carya ovata). Most of the neighboring trees were white oak, but shortleaf pine was also well-represnted throughout this forest. A hearty specimen of poison oak (Rhus toxicodendron= Toxicodendron radicans= Rhus radicans) had claimed the trunk of this large shellbark for its own.
Lennox Woods, Red River County, Texas. May.
8. A good foundation- Trunk of the large shagbark hickory shown in the preceding photograph. Note the characteristic bark which in large trees often forms canoe-sized sheets or shelves projecting conspicouosly from the large trunk. A large specimen of poison oak was growing up the right side of this trunk.
Lennox Woods, Red River County, Texas. 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.
9. 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).
10. 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).
11. 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).
12. 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).
13. 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).
14. 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).
Mixed white-black-northern red oaks forests probably constitute the "umbrella" or general forest type of which forests dominated singularly by white, black and northern red oaks are affilitated with each of these three being a forest cover type--frequently subclimax or climax--itself on certain forest sites. According to cover type descriptions by the Society of American Foresters (Eyre, 1980, ps. 41-44) the white oak- black oak-northern red oak cover type (52) is clearly the most extensive or widely distributed forest type among these various cover types or forms of the oak-hickory association. Another rational perspective of SAF forest cover type 52 is that it is a transition zone among white oak (SAF 53), black oak (SAF 110), and northern red oak (SAF 55) types.
In the western portion of the Ozark Plateau upland forests of the white oak-black oak-northern red oak cover type were usually found by this author to have developed on predominately north and east slopes with better drained soils resulting in overall moderately mesic to somewhat dry habitats. This corresponded to the dry-mesic chert forest of Nelson (2005, ps. 125-130). The Society of American Foresters (Eyre, 1980, ps. 41-42) listed white oak as the first of three dominants for this forest cover type and indicated that white oak occurrs across the moisture gradient from dry to moist. This followed the conclusion of Braun (1950, ps. 167- 168) that white oak was more common on north slopes, but that it was a major species on "many slope forests" where black oak was often "its principal associate" except on more mesic habitats where northern red oak held this status. Sugar maple is also a component species on slope forests, especially on limestone soils, "where it may be associated with white oak, red oak, chinquapin oak, basswood, and hickory" (Braun, 1950, p. 168).
Where sugar maple is a dominant rather than associate species the forest vegetation is recognized as another form of Ozark Plateau forest (Braun , 1950, ps. 168-169) and a variant or form of the sugar maple cover type and/or the sugar maple-basswood cover type (Eyre, 1980, ps. 29-30, 31-32, respectively). These latter hardwood forests were interpreted as component communities--in effect, edapho-topographic climaxes--of the general oak-hickory association as was explained by Braun (1950, p. 33-34) and quoted in the introduction of this chapter above. The sugar maple types of the Ozark Plateau were treated later in this chapter of Range Types because they are climax forest types (associes vs. associations in the Clementsian model) of the oak-hickory association in Ozark Mountains forests.
It was necessary to mention and draw attention (briefly) to Ozark sugar maple forests at this juncture because these cover types blend into the white oak-black oak-northern red oak type (SAF 52) and the northern red oak cover type (SAF 55); that is, there is a continuum among these various cover types, each of which can be climax on specifie forest range sites.
15. The mixed oak type above a hollow- Local second-growth stand of the white oak-black oak-northern red oak forest type in the western Ozark Plateau. This forest range vegetation grew on a predominately east slope. It was a more mesic forest site with co-dominants being white and northern red oaks. Woody plants from left to right in fore- and midground: sugar maple saplings with northern red and white oaks behind these; then from right of center (still left to right line-up) white oak, northern red oak, white oak, two northern red oaks, and, the shrubat far right, flowering dogwood (Cornus florida). Saplings behind these trees and shrubs were mostly bitternut or pignut hickory that was generally the associate species. Summer grape (Vitis aestivalis) was present with some growing among the sugar maple saplings (to confuse viewers trying to distinguish plant species). No herbaceous layer in this local stand. Instead there was a leaf layer acting as mulch that formed the ground layer of this forest vegetation.
Ozarkers are fond of specifying that the hills are not as high as the "hollers" are deep. The composite mix of dominant trees just enumerated "ringed" the head of a hollow in the Springfield Plateau portion of the Ozark Plateau.
This forest had no large animals in it except free-ranging white-tailed deer (Odocileus virginianus). Currently, deer populations were below carrying capacity of this forest range. Approximately 50-60 years ago this had been legal open range resulting in severe overgrazing and overbrowsing by livestock, especially hogs.
Fisher Flats, Ottawa County, Oklahoma. June (vernal aspect). 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).
16. Mixing around the edge- A second-growth stand of white, black , and northern red oak with a diverse understorey formed a composite example of this forest cover type at the head of a hollow on a predominately east slope in the western Ozark Plateau (Springfield Plateau). The three Quercus species were "about evenly divided" as tri-dominants. The associate species was pignut, bitternut, or (sometimes) white hickory. The principal shrub was flowering dogwood. The largest tree (left margin, foreground) was white oak as was the tree at far right. Center (somewhat right of center) tree was a black oak.
There was some regeneration of all oak and hickory species in spite of considerable shade. It was concluded that for this was the climax vegetation for this forest range as Eyre (1980, p. 42) concluded for the Ozark Highlands.
This local second-growth stand of mixed oak was, like the stand presented in the last slide, also at the head of a hollow. Likewise grazing/browsing by large animals was limited to that by white-tail deer.
Fisher Flats, Ottawa County, Oklahoma. June (vernal aspect). 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).
17. Under the oaks- A photo-quadrant that featured local herbaceous constituents of the east slope, second-growth stand white oak-black oak-northern red oak forest described in the immediately preceding caption. As in the instance of that stand there was regeneration of all major tree species (the three oaks and bitternut hickory). Again, flowering dogwood was the dominant shrub. The dominant herbaceous species was big bluestem (Andropogon gerardii). In this photograph the most abundant forb was the introduced perennial legume serecia lespedeza (Lespedeza cuneata). Shrubs comprised a pronounced and varied layer including smooth shumac (Rhus glabra) poison oak or ivy (R. radicans= Toxicodendron radicans), summer grape, Virginian creeper (Parthenocissus quinquefolia), greenbriar (Smilax spp.), and flowering dogwood. The various species of woody vines formed a "unifying" layer from ground layer to canopy of tallest trees.
Fisher Flats, Ottawa County, Oklahoma. June (vernal aspect). 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).
18. Holler away- Two photographs of a second-growth mixed oak-sugar maple forests at head of another hollow in the Springfield Plateau. The forest range vegetation introduced here and described in the next series of slides was in the general vicinity of the head-of-hollow white oak-black oak-northern red oak-bitternut hickory forest presented immediately above. This was a north versus the predominately east slope of the preceding stands. Composition of the adult and regenerating tree layers (including sugar maple) was the same as east slope stands except that bitternut or pignut hickory was barely present (largely absent) from this north slope forest. The most pronounced differences between the north and east slope forests were in the herbaceous and shrub layers. The dominant herbaceous species on this north slope was the native perennial legume, tick clover (Desmodium nudiflorum), whereas the dominant herb on the slightly drier east slope was big bluestem. Flowering dogwood was common to both east and north slope forests, but on north slope stands the associate shrub was highbush or squaw huckleberry or deerberry (Vaccinium staminium) followed by eastern or common redbud (Cercis canadensis). Young individuals of sassafras (small sapling-size), although a tree species at maturity, comprised another constitutent of the shrub layer.
Two photographs showing the same local stand of mature trees of a second-growth mixed oak-sugar maple, Ozark Hihglands forest on a north slope (mesic) with very limited understorey (mostly shed leaf ground layer). The first of these two photographs (vertical) presented a textbook line-up of the species composition of this forest cover type. Tree species from left to right: white oak, northern red oak, white oak, northern red oak , and two white oaks. The second photograph presented most of the same trees in a horizontal plane (left to right): white oak, northern red oak, white oak, northern red oak, and white oak. In the shade of this dense stand there was little if any understorey except for a ground layer of shed leaves with an occasional plant of Desmodium nuiflorum, another unidentified (pre-bloom) Desmodium species, and sugar maple seedling.
This tract of forest had not been grazed/browsed by livestock for decades (probably about a half century). From the frontier era up to that time forests in this area had been subjected to free-ranging livestock (including swine) under legal open range status with abuse of forest resources being "the norm". Spring burning of these woods was also a common practice until about the same period. Subsequently the only ungulates-- large vertebrates at all for that matter--in this forest were free-ranging white-tailed deer which ranged throughout this and adjacent forests. At time of these photographs deer numbers had not reached the stage of overpopulation, not yet anyway.
Ottawa County, Oklahoma. July (estival aspect). 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).
19. A hollow cast of charaacters- At the head of a hollow on a north slope in the Springfield Plateau of the Ozark Highlands a second-growth, hardwood forest of white oak-black oak-northern red oak (tri-dominants) with sugar maple as the associate species had developed. This local stand provided a good example of this climax forest cover type and its range vegetation. Tree species in this photograph (left to right) were a mid-size white oak and two young pole-size northern red oaks. Between these three trees and in mid foreground were two small, crooked-trunk saplings of flowering dogwood. Eastern redbud was present though scarce. Scattered small saplings of sassafras were also present as in the woody layer that was dominated by shrubs (again, mostly flowering dogwood). Major herbaceous species were two species of tick clover (Desmodium nuiflorum and another Desmodium species in pre-bloom stage that could not be positively identified).
Ottawa County, Oklahoma. July (estival aspect). 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).
20. Principal players- Good species composition "quadrant" showing climax species of: 1) dominant trees, dominant shrub, and dominant forbs which also portrayed layers of vegetation and general forest structure. Plants in a left to right line in foreground of slide were: leaves of black oak (left margin), two black oaks, bent sapling of sugar maple (foreground), and white oak (extreme right margin).
Ottawa County, Oklahoma. July (estival aspect). 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).
21. Developing forest with the understorey coming first (or forest vegetation on the edge)- An edge of the north slope white oak-black oak-northern red oak forest featured here provided an opportunity to document a successional stage of forest development. All three oak species and sugar maple had ample regeneration at the gap-like edge of this mesic Ozark Hihglands second-growth forest. Summer grape was also well-represented by young and conspicuous plants as was highbush or squaw huckleberry (short woody plants with large linear shade leaves). Some sassafras was present as large seedlings.
The major herbaceous species was one of the rosette panicgrasses (one of the species in Dichanthelium subgenus), which in the post-spring bloom and grain-shatter stages could not be identified positively, but seemed most likely to be western panicgrass or western rosettegrass (Panicum lanuginosum= Dichanthelium acuminatum var. implicatum= Dichanthelium lanuginosum). Members of the Dichanthelium subgenus produce winter foliage in the form of a rosette (hence rosette Panicum spp.) from which infloresecences emerge to typically bloom in mid to late apring and then again in autumn. This rosette panicgrass species presented here had formed an extensive colony at edge of the north slope mixed oak-sugar maple forest. Poverty oatgrass (Danthonia spicata) was also abundant-- by standards of the shade cast by this forest even at its outer edge.
Ottawa County, Oklahoma. July (estival aspect). 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).
22. Highbush huckleberry, squaw huckleberry, or deerberry (Vaccinium staminium)- On oak-hickory forest range of the Ozark Hihglands where this shrub occurs it is a defining species because highbush huckleberry has nothing even approaching the commonness and widespread feature of other shrubs likc flowering dogwood, blackberries, smooth sumac, poison oak or ivy, and Virginia creeper. Highbush huckleberry is usually restricted to the more mesic, yet well-lite forest sites. This specimen was growing at the edge of the north slope, white oak-black oak-northern red oak-sugar maple forest shown in the four immediately preceding photographs.
Seedlings in lower corner of foreground were black oak and blackjack oak (Quercus marilandica).
Ottawa County, Oklahoma. July (estival aspect); immature fruit stage of phenology.
23. Under uneven-aged management- General view of an upland white oak-northern red oak second-growth forest in Boston Mountains. Shortleaf pine (Pinus echinata) is a very infrequent component of this forest range community, but not "anywhere near" associate species status. As such, this forest tract was presented as another example of the mixed oak type in which white oak ranks the edge as the "first-among-equals" dominant. Red maple was an important member of the understorey with its ultimate staus yet to be determined in this forest (see below).
Shrubs included flowering dogwood, redbud, buckbrush or coralberry, poison oak, poison ivy, lowbush huckleberry (Vaccinium vacilians), and various kinds of blackberry. Grasses and forbs were in short supply, but some species were noted in the next caption.. Most of the understorey consisted of various layers made up of the age classes of the co-dominant oak species and red maple.
Ozark National Forest, Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004)
24. White oak-northern red oak forest under uneven-aged management- Appearance "closer-up" of a second-growth upland mixed oak forest under selective harvest. Species composition and structure in the interior of this forest range was emphasized in this photograph. White oak was the most common (major) dominant, but northern red oak gave it a good run for the money. In this view there was tremendous regeneration of bittrnut or pignut hickory (Carya cordiformis as well as northern red oak (center foreground). Red maple at seedling and small sapling age classes was extremely abundant with its successional and domiance status unknown.
Most of the understorey was regeneration of oaks.bitternut or pignut hickory, and red maple with the various layers made up of the different age classes of these hardwood trees. Shrub species included flowering dogwood, redbud, buckbrush or coralberry, poison oak and poison ivy, Viraginia creeper, and various kinds of blackberry. Not much by way of grasses and forbs. Virginia and silky wildrye (Elymus virginicus and E. villosus, respective) were the main grasses, but they were sparse and widely scattered. Mayapple, one of the more characteristic herbaceeous dicots throughout the Ozark Plateau, was the most conspicuous forb. Clearly this was was a browse range with white-tail deer the game species of choice (the range animal managed for).
Obviously the main commodity from this forest tract was saw timber. It was instructive that wood from this forest included a major representative of the white oak group (Leucobalanus subgenus) and the red oak group (Erythrobalanus subgenus). The red oaks are especially prized for fuurniture manufacture whereas the white oaks have traditionally been used as rougher construction materials as well as cooperage, wooden containers (eg. barrels) made up of a round head (made of wood) on both ends and a body of one to many staves (wooden; long, narrow, and curved one-piece units) held together by hoops (usually metal). Historically one of the most important uses of oak was for timbers (by definition at minimum 5 inches by five inches), especially railroad ties (=crossties) which are eight inches by eight inches and eight feet long. Hickory is prized for tool handles because it is so resistant to shock, breakage, and deformity (tough wood fibers of hickory are very resilant or plastic), is much easier on workers' hands than metal or fiberglass, and aesthetically pleasing (at least to those with good taste and not obsessed with synthetics).
Ozark National Forest, Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004)
25. Interesting interloper in the understorey- Large seedling of red maple (Acer rubra) in understorey of an upland white oak-northern red oak second-growth forest with uneven-aged management. This species was very abundant in this developing forest, but the ultimate successional status and role of this species in an mixed oak-hickory was unknown.
Red maple is a fairly palatable browse for white-tail deer. All species of livestock will browse on hardwoods to some degree or the other. They are especially fond of mast, including acorns and hickolry nuts. Hogs generally browse less than other kinds of livestock, but their disturbance of soil (rooting and wallowing) and feeding on roots makes swine the most potentially destructive of all livestock in hardwood forests, perhaps especially the oak-hickory forest range types.
Ozark National Forest, Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004)
Note on location: An example of the white oak-black oak northern red oak forest in which black oak was a minor component (generally not present) was presented below at end of the section devoted to the variant forest cover type of American beech (Fagus grandiflora)-white oak-hickory.
In western parts of the Ozark Highlands, such as the Springfield Plateau, black oak-dominated forests are some of the most common remaining forest tracts as they are found on some of the driest, shallowest, rockiest soils and, hence, generally the harshiest of remaining forest sites. The more mesic forest sites with their deeper, less stoney soils (ie. the less marginal, more-or-less arable land) naturally developed old-growth forest that were dominated by northern red and white oaks (with black oak as an associate species). Most of these more productive mesophytic forests were converted to hill-side pastures, strawberry patches, fruit orchards, and even field or vegetable crops decades ago. Some of the cropland derived by clearing the original (post-Indian) white oak and northern red oak forests was retired from intensive cropping years later as a result of socio-economic developments (ie. loss of small, inefficient farms). Some of the old-fields (abandoned farmland) on these almost subsistence-level (and hobby) farms have gone back through secondary plant succession to the stage that they revegetated to seral or subclimax black oak forest. Also, some old-growth (climax) black oak forest have recovered and are routinely logged (usually high-graded or clearcut) at, say, quarter to half century intervals.
Some examples of these second-growth black oak forests and secondary plant succession on them following disturbance was presented in the following section.
26. 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 and white 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 with black oak the dominant and northern red oak the associate 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. For this forest site and this stocking rate these trees were probably of their maximum size. There were several snags. However most regeneration was hickory. This was a second-growth forest, yet species composition and structure of forest was climax. This forest approached old-growth status.
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).
27. 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).
28. 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.
29. 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).
30. 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).
31. 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).
32. 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).
33. 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).
34. 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.
An example of high-grading- The following short set of seven slides was an example of high-grade harvesting in a second-growth mixed hardwood forest of northern red oak, chniquapin oak (Quercus muhlenbergia), Texas or black hickory (Carya texana), black oak (Q. velutina), post oak (Q. stellata), hackberry (Celtis occidentalis),and American elm (Ulmus americana) that had developed on the western Ozark Plateau in northeastern Oklahoma. This was an ecotonal forest existing as transition vegetation between an upland northern red oak-chinquapin oak-black hickory forest and a bottomland hackberry-American elm forest.
This second-growth ecotonal forest had been subjected to high-grade logging the year before these photographs were taken. These images documented the ecological/economical evils of high-grading, the selective harvest of the most commerically valuable trees (high-grade trees) leaving a remaining forest made up of trees (individual- and species-wise) of lower value such that species composition of the residual forest is less desirable ecologically and/or economically (Helms, 1998).
35. High-graded; opened up- Man-made patch or gap in a second-growth Ozark mixed hardwood forest created by high-grade logging of two sub-adult (immature for harvest; too small to be felled for log production) northern red oak. Remaining trees were (three in fore- to midground, left to right) were chinquapin oak, northern red oak and Texas or black hickory. Leaves at upper right margin were of American elm. Forbs in center were pokeberry (Phytolacca americana) and the naturalized Asian weed known as beefsteak plant or, locally, rattlesnake weed (Perilla frutescens). Both of these forbs pioneer disturbance areas, but pokeweed is a native perennial whereas beefstake plant is a prolific seed-producing annual. The main shrub was the native buckbrush or coralberry (Symphoricarpos orbiculatus), a highly rhizomatous, low shrub of the honeysuckle family (Caprifoliaceae).
Details of this forest gap (including the two stumps of the logged northern red oak) and the pioneering forest vegetation in this second growing season post logging were shown in the next photograph.
Ottawa County, Oklahoma. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest)/K-.92 (Elm-Ash Forest). SAF 55 (Northern Red Oak)/SAF 93 ((Sugarberry-American Elm-Green Ash). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37)/and nothing fit for the bottomland hackberry-American elm component. Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
36. Climax trees replaced by pioneer forbs- Stumps of two sub-adult northern red oak, the potential climax dominant tree species for this site, that were improperly harvested (felled when too small for good lumber logs) two years prior to time of this slide. Surrounding first seral stage plants included pokeberry, beefsteak or rattlesnake weed, hairy crabgrass (Digitaria sanguinalis), prickly lettuce (Lactuca scriola= L. serriola), and coralberry or buckbrush as the now-dominant species.
Even though both of the northern red oak were young trees with a high proportion of sapwood and cambium, there was no coppicing (stump sprouting): both trees had been killed even when harvested prematurely. Premature in this usage referred to younger trees which are not only smaller but also more likely to resprout. In these two instances, however no asexual reproduction had taken place.
Remaining trees in this "photo-transect" of a second-growth Ozark forest were chinquapin oak and northern red oak (eg. the saplings left and right, respectively, in far midground; the same two trees as presented in the immediately preceding slide), black hickory, hackberry, and American elm.
These trees (along with others presented in this "ecological crime scene") had been felled by someone that knew nothing about Forestry except how to identify northern red oak, and he harvested almost every one of these bigger than a sapling. (Your author deduced who the logger was when he showed up at the writer's homeplace wanting to cut big northern red oaks there. He got sent packing in short order.)
Northern red oak lumber is quite valuable for lumber, especially for church pews, and makes the best oak firewood because it splits so easily (relatively speaking). Harvest of these two far-below-prime age/size oaks was a waste of the wood resource. Removal of these two northern red oaks ("its so easy to split" in the fellers words) and leaving behind inferior lumber species such as American elm, hackberry, chinquapin oak, black hickory, and post oak was a textbook case of highgrading as defined in the introduction of this short section.
Ottawa County, Oklahoma. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest)/K-.92 (Elm-Ash Forest). SAF 55 (Northern Red Oak)/SAF 93 ((Sugarberry-American Elm-Green Ash). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37)/and nothing fit for the bottomland hackberry-American elm component. Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
37. Those left behind- Inside a northern red oak-chinquapin oak-black hickory-black oak-post oak-hackberry-American elm second-growth forest two years after amateurish high-grade logging of northern red oak. Trunk of American elm (right margin) with Virginia creeper (Parthenocissus quinquefolia), blackberry (Rubus species), poison ivy (Rhus radicans=Toxicodendron radicans), beefsteak or rattlesnake plant, and buckbrush or coralberry coming up in a gap created by felling a young (immature for proper harvest) northern red oak two years ago.
Ottawa County, Oklahoma. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest)/K-.92 (Elm-Ash Forest). SAF 55 (Northern Red Oak)/SAF 93 ((Sugarberry-American Elm-Green Ash). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37)/and nothing fit for the bottomland hackberry-American elm component. Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
38. Textbook highgrading- Inside a second-growth northern red oak-chinquapin oak-black hickory-black oak-post oak-hackberry-American elm forest two years after it had been high-graded for northern red oak (logging off of mostly of this species and leaving behind trees of the less valuable lumber species). Almost all northern red oak had been too small--by at least 20 years on this forest site--to be of much value (some were barely above large sapling-size).
The stump of a northern red oak less than one foot in diameter at ground level--and which had not stump sprouted--was visible in foreground of the first slide. Plants surrounding this dead stump (in absence of coppicing two years post felling it was obvious that this young northern red oak had been killed) included individuals of buckbrush or coralberry, beefsteak plant, prickly lettuce, hairy crabgrass, and seedlings of hackberry. There were no seedling of northern red oak; in fact the felled tree was so young it was doubtful that it had reached sexual maturity and was capable of producing acorns.
Pioneer species in this man-made forest gap ranged from native perennial forbs like pokeberry to exotic annual forbs, including beefsteak plant and prickly lettuce, to hairy crabgrass, an annual grass, to shrubs species including buckbrush or coralberry, blackberry, Virginia creeper, and poison ivy up to seedlings of hackberry and American elm. Two forb species in the forest vegetation of these two slides and not presented in other slides included white vervain (Verbena urticifolia), a native perennial, and the Eurasian biennial weed, flannel mullein (Verbascum thapsus).
The tallest seedling/small sapling stage shoots in the second slide were of American elm. There were relatively few seedlings of northern red oak, but abundance of American elm in this ecotonal forest was heartening to the extent that the species had not been locally exterminatedy by the onslaught of Dutch Elm Disease that wiped out beautiful, huge Americn elms a third of a century prior to time of these slides.
The four trees in the first slide (left to right) were post oak, northern red oak, and two black or Texas hickories. The large tree in background of the second slide was a black oak.
It was obvious from this highgrading operation that northern red oak, the choice lumber species, was in decline relative to other tree species. This is the classic, defining result of highgrading. Highgrading is a form of overlogging just like overgrazing and overfishing in which the most valuable species (those most palatable to livestock and humans, respectively) decline relative to the pre-harvest composition, or they disappear all together. Ultimately, overharvest of any renewable natural resource results in decline or loss of the preferred species while leaving behind the trash or weedy (least preferred) species.
Ottawa County, Oklahoma. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest)/K-.92 (Elm-Ash Forest). SAF 55 (Northern Red Oak)/SAF 93 ((Sugarberry-American Elm-Green Ash). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37)/and nothing fit for the bottomland hackberry-American elm component. Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
39. Stump and gap- Stump of a northern red oak that had been felled in a second-growth forest of northern red oak, chinquapin oak, black hickory, black oak, post oak, hackberry, and American elm subjected to high-grade logging two years ago. This northern red oak was one of the very few of its species that was of a commercially useful size, and it had wood damage due to heart-rot (seen as the black disk--actually a hole--in center of the stump). Heart-rot is decay of portions of the heartwood caused by a number of native fungus species (Wagener and Davidson, 1954, Berry, 1973). Usually heart rot (known also a heart decay [Helms, 1998] is limited to the dead heartwood so that it does not adversely affect the host (infected) tree as it does not spread to living sapwood. Instead, damage is to value of logs from infected trees the lowered value of which depends on extent of damage (area of infection).
Heart rot-causing fungi are primarily indigenous species which enter the tree through points of injury such as those caused by fire or breakage due to wind and ice. Older, larger trees are, of course, more apt to have heart rot because they have been subjected to more injurious agents simply by having "been around longer".
Heart decay is common throughout the Ozark Pleateau being a natural feature of forests in this region. Experienced loggers can often tell if a tree is likely ot have heart rot by seeing injured areas (eg. broken limbs, fire scars). Fellers--many of whom are sportsmen--often leave such trees, which in time may become den trees for furbearers like coon (Procyon lotor) and a source of acorn mast for most animals as well as being a seed source for the next tree crop.
Plants growing in the forest gap formed by the harvest of this northern red oak included buckbrush, beefsteak plant, prickly lettuce, hairy crabgrass, and a few seedlings of northern red oak. As noted by the great author Carl Sandberg, “A baby is God's opinion that life should go on." Even with harvest of the parent tree there was regeneration of its species. Afterall, reproduction is God's vote that the world should go on.
Ottawa County, Oklahoma. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest)/K-.92 (Elm-Ash Forest). SAF 55 (Northern Red Oak)/SAF 93 ((Sugarberry-American Elm-Green Ash). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37)/and nothing fit for the bottomland hackberry-American elm component. Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
40. Harvested before its time with weeds left behind- Small, hence young (less than one foot basal stump diameter), northern red oak felled in a high-grade logging operation in a second growth forest of northern red oak, chinquapin oak, black hickory, black oak, post oak, hackberry, and American elm forest in the western Ozark Plateau. Even though thisnorthern red oak had been a young tree with a trunk consisting of least three-fourths sapwood, it had not (coppiced (stump sprouted) in this second year after felling.
Instead, a seedling of eastern red ceedar (Juniperus virginiana) was growing immediately adjacent to the stump (to left, immediate front of stump as seen here). Such shifts in tree species is the net result of highgrading. Other plants that pioneered this local disturbance included hairy crabgrass, beefsteak plant, prickly lettuce, and buckbrush or coralberry.
Ottawa County, Oklahoma. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest)/K-.92 (Elm-Ash Forest). SAF 55 (Northern Red Oak)/SAF 93 ((Sugarberry-American Elm-Green Ash). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37)/and nothing fit for the bottomland hackberry-American elm component. Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
Note: there were at least two positive outcomes of this otherwise "ecologically evil" highgrading. The logging-created forest gaps produced a boon of feed--both herbaceous and woody--for white-tailed deer . Likewise, the resultant gaps or openings within the canopy of this mixed hardwood forest resulted in creation of blackberry patches which was a boon to your berry picking author.
A sticky beggar-- Plant of Virginia stickseed or beggar's-lice (Hackelia virginiana) growing a cutover (= high-graded) uplnd transition forest of mixed oak (northern red oak dominant)-hackberry-American elm in the western edge of Ozark (Springfield ) Plateau. This native annual forb of the borage fami8ly (Boraginaceae) was growing close to the stump of a felled "sub-prime" (young adult tooo small for optimum harvest) northern red oak.
Annual forbs like Virginia beggar's-lice are generally weeds (even though they are native species), but in this casebeggar's-lice was a pioneer plant on the initial (pioneer) successional stage on the sere of this upland forest. The stage "had been set" for secondary forest succession. Under this condition, Virginia stickseed was a desirable plant. It was facilitating re-development of the forest vegetation via secondary plant succession. Stickseed was an indictor species. It indicated that the forest was slowly "healing itself", the forest community was in the first step toward re-establishment of the climax forest vegetation.
Ottawa County, Oklahoma. Mid-July; late-bloom phenological stage.
Lousy end of a sticky beggar- Inflorescence on lateral branch (first slide) and flowers and fruit (second slide) of stickseed or beggar's lice on the specimen introduced in the previous slide-caption set.
The fruit of Virginia stickseed, a member of the borage family, has been interpreted as being comprised of two united carpels each of which is divided into two locules (total of four) each enclosing a nutlet or achene for a total of four (Smith, 1977, p.195).
Ottawa County, Oklahoma. Mid-July; late-bloom phenological stage.
41. Black oak forest in an Ozark spring- Interior view of dry chert upland forest dominated by black oak with post oak, 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 (Symphoricarpos orbiculatus) 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 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"!
42. 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).
43. 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.
44. 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 ploe-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.
45. "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. June.
46. 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).
47. Savanna specimen- An old-growth black oak (Quercus velutina) on a one-time tallgrass prairie savanna. This old-age, (and now senescing) tree established and grew on a big bluestem-dominated prairie and matured into the "shade tree" (that is, open growth) form characterized by a fully developed crown. Black oak is primarily a forest tree species--and a major species in the forests and savannahs of the ancient Ozark Mountains--but black oak is also one of the major oak species, after the co-dominant post oak and blackjack oak, on the oak-hickory-tallgrass savannas and woodlands.
The old codger was the last remnant, about the last plant, of a former oak-hickory savanna that through overgrazing, overmowing, overseeding, haying, etc. was converted to a "trash pasture" with "a little of everything" besides the native grasses and palatable forbs. This specimen served as an example for both 1) its species at maturity and 2) a savanna form of its species.
Springfield Plateau portion of the Ozark Mountains, Ottawa County, Oklahoma. Late July.
48. Morphology of the lower end on a savanna specimen- Trunk and lower crown of the old-growth black oak introduced in the immediately preceding two-slide/caption set. This magnificant specimen grew on an Ozark (Springfield) Plateau savanna on which almost all other native plant species had been obliterated by overgrazing and related stock-farming activities. Relict stands of big bluestem along with such forbs as catclaw sensitive-briar (Schrankia uncinata) and Illinois bundleflower (Desmanthus illinoensis) remained growing in fencerows all around this tree which, in the spirit of James Fenimore Cooper, remained "The Last of the Mohigans" of the pre-white man savanna.
Sad, but this old-growth relict grew along the author's route to the one-room schoolhouse of his first eight school years and, up to this time, it still stood to greet him on adult walks to the yellow-brick schoolhouse which was in worse shape than this slowly senescing black oak.
Springfield Plateau portion of the Ozark Mountains, Ottawa County, Oklahoma. Late July.
A forester's or rangeman's Sherlock Homes observation: keen observers of the above four photographs will have noticed that there were no lower branches or leaves of the above old-growth black oak. This was the direct result of browsing by beef cattle (usually cows and calves, rarely stockers). This photographer observed cattle reaching up their necks and extending their tongues as far as possible to grasp leaves and small twigs which were readily eaten. A popular myth is that because cattle are, by general preference, grazers rather than browsers (scientific fact) cattle do not eat browse (leaves, buds, twigs, etc.) of trees (absolute balderdash). This conclusion is wrong and based on lack of observation-- both of 1) cattle feeding behavior and of 2) physical evidence of plant material removal by cattle.
Yes, white-tailed deer also ranged over this area and, most likely, deer did eat some foliage from this black oak, but the stocking rate of cattle in this small, fenced enclosure was much greater than that of free-ranging white-tailed deer. Furthermore, white-tailed deer were absent from this pasture (this entire general area in fact) for a period of at least a half century.
The absence of leaves and small twigs from the black oak down to heights at which cattle could reach constituted a broswe line, "a well-defined height to which browse has been removed by animals" (Kothmann, 1974). Development of a browse line is commonly known as highlining. Presence of a browse line indicates either 1) overbrowsing or, in cases where such tree material is very limited, 2) consumption of feed produced by ice cream species. Ice cream species are those plants that are so palatable they cannot be maintained under proper stocking rates. There is no evidence to indicate that black oak is an ice cream species to cattle.
It is axomatic that if cattle find black oak leaves and buds palatable enough that they will put forth extra grazing (browsing) effort to reach high to get this feed, cattle will certainly eat black oak seedlings that are at the grazing level of grass where less expenditure of energy and grazing motion is needed. Lack of recruitment of black oak on this former savanna was due, at least in large part, to consumption of black oak seedlings by cattle. Forget about "brush goats". Grass-preferring cattle are more than adequate to prevent regeneration of black oak on this climax savanna of which black oak was a major woody species (apparently the dominant tree species). Likewise, cattle were probably the major--though certainly not the only factor as lack of fire was also of paramount importance--in prevention of black oak reproduction and invasion of eastern red cedar on the second-growth black oak forest shown and described above.
Foresters know what they are talking about when they state that heavy grazing by livestock (including cattle) will retard regeneration of hardwood species. Oak reproduction (regeneration of any natural hardwood species) is undeniably the major management objective in maintenance, productivity, and use of oak-hickory forests just as in preservation and use of natural hardwood-tallgrass savannahs. Invasion by brush (any woody species present at excessive density and cover) can be another major--and often a commensurate--problem (eg. the above example of eastern red cedar invasion of a black oak forest). Brush invasion can be exasperated by overgrazing (eg. by beef cattle in the above black oak forest example), often accompanied by (and contributed to) cessation of fire. Wise (= proper) use management--and not the abuse and mismanagement shown in the two examples just presented--is essential if forest and range resources are to around for the next generation (or, maybe, even to end of the current generation).
Cattle and deer (as well as fox squirrels [Sciurus niger] and other vertebrates, including birds) readily ate acorns produced by this black oak. Such consumption of acorns (the sexual propagules) was obviously a limiting factor in local regeneration of black oak. Conversely, free-ranging wildlife were also agents of dispersal as, for example, when squirrels or woodpecker species removed, transported, and stored acorns at considerable distance from the parent tree. Squirrels buried acorns and, hence, planted trees at relatively long distances from the parent black oak. This was most likely the explantion for establishment of a black oak seedling, offspring of this old-growth black oak, at three or four times the length of tree height away from the seed source (see below).
49. As they grew- Leaves and immature acorns on the black oak specimen shown and described in the two immediately preceding slide/caption units. This is the density, overlapping cover, and arrrangement of leaves and acorns as they appeared growing on the old-growth black oak.
Springfield Plateau portion of the Ozark Mountains, Ottawa County, Oklahoma. Late July.
50. Pattern and shape- Pattern of arangement of leaves and young acorns on the apical (terminal) end of twigs (first slide) and shape of a typical leaf (second slide) in (of) black oak. These organs were off of the old-growth black oak specimen introduced above.
Springfield Plateau portion of the Ozark Mountains, Ottawa County, Oklahoma. Late July.
51. Leaves and acorns- Characteristic leaves and immature acorns of black oak on the old-growth specimen shown and described above. The acorns were still immature, but would complete growth and ripen in about two months henceforth. This tree was in the far western edge of the Ozark (Springfield) Plateau.
Ottawa County, Oklahoma. Late July.
52. Out on the end-- Immature (but rapidly growing and ripening) acorns of black oak (Quercus velutina). These acorns were
Springfield Plateau portion of the Ozark Mountains, Ottawa County, Oklahoma. Late July.
53. The proof of any fruit; the only measure of survival- "A chip off of the old block". "That acorn didn't fall far from the tree". Both of these old (and, perhaps, hackneyed) expressions aptly expressed the fact that this large (about a yard tall) seedling of black oak was the sexual progeny of the immense old-growth specimen presented and explained above. This seedling was roughly three to four times the height of the adult old-growth black oak away from its parent. It was growing safely in a fencerow where cattle and, so far, a rotary shredder had not eaten or broken it off. This seedling most likely grew from an acorn planted by a fox squirrel.
There were no offspring trees (that means zero)--of any age or size--from the old-growth black oak anywhere (at any location) that cattle grazing/browsing or mechanical mowing/shredding had occurred at any time over roughly the last century.
It was shown and explained above that beef cattle (and dairy cattle in decades before that) had highlined (removed all tree material within their reach, that is to this height).
The "proof of the pudding' for any plant propagule is that it produce more plants (or plant parts). One acorn had "done its thing" in this case shown here. The final, the only, measure of any species is that it reproduce (regenerate, recreate) its kind. The only measure of the "genetic fitness" of the individual genotype is that it leave progeny to replace it. This old-growth, parent black oak as a representative of its species had done this (ie. it "measured up").
In the instance presented here, failure (of genotype or species) in regard to reproduction, to regeneration, would not have been a failure to natural phenomenon (not failure of natural selection), but rather failure in an unnatural set of conditions created by contemporary man (failure to have fitness to a manmade habitat, to an environment in which man was an artificial and not a natural part).
In final analysis, not only had the native herbaceous species been lost from this savanna, but the woody component was being lost-- only at a slower rate--such that in only a few years none of the savanna will remain. Specifically, the savanna will remain only in fencerows--and only that until hatchet-happy, make-work county road workers destroy it.
Springfield Plateau portion of the Ozark Mountains, Ottawa County, Oklahoma. Late July.
Oak-hickory forest dominated by northern red oak, northern red oak cover type 55 (Eyre, 1980, ps. 43-44), tend to develop on some of the most mesic forest sites and, therefore, to be some of the most productive forests throughout theoak-hickory association. Unfortunately for these forests (and the foresters and rangemen who would manage them) many were converted into farmland and, later, unceasing concentrations of highways, locations for houses (in all price ranges), and the final consequences of unabated, never though-through, run-amuck suburban sprawl.
From a more positive perspective some northern red oak forests remain and some of the degraded (by farming and improper logging, grazing, burning, etc.) tracts have recovered to considerable degree. Perhaps most impressive and inspiring have been efforts made by some states, local governments, and non-government organizations that resulted in preservation of remaining relicts of natural vegetation including northern red oak forests. Missouri has been one of the leaders in this marvelous development in preservation of its natural heritage. Nelson (1987, 2005) described many of these natural communities and listed various units of land and water set aside by Missouri and other cooperating organizations. These included various oak-hickory forests, woodlands, and savannahs in which northern red oak is a constitutent species.
54. Stand of young northern red oak- On a north slope of a hollow in the Springfield Plateau of the Ozark Highlands a second-growth of northern red oak was developing to the species composition and vegetational structure of a climax red oak-dominated forest. Sugar maple, also young trees, was the associate species. A good example of young sugar maple was the rightmost tree (small pole or large sapling). The major shrub was flowering dogwood which comprised most of the shrub layer although it was accompanied by its usual shrub associate, eastern redbud. Numerous shrub-sized or small tree-class sassafras were also part of this intermediate woody plant layer. A lower woody layer was made up of regenerating hardwood tree species includnig white oak as well as sugar maple and northern red oak. An herbaceous layer was also present. It was dominated by tick clover.
Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
55. Recovery of northern red oak-A second growth forest dominated by young trees of northern red oak with sugar maple as the associate species on a mesic north slope in the western part of the Ozark Highlands (Springfield Plateau). A few older and larger trees of white oak were present locally in this tract (two larger trees at far right, the rear one of which was dead with exfoliating bark) and there was regeneration of white oak (seedling age class). Most regeneration of all younger age classes was northern red oak with sugar maple (eg. large saplings to front and left of the two large white oaks) being second in this regard. In other words it seemed obvious that white oak was seral to northern red oak and sugar maple.
The major shrub was flowering dogwood which was growing all around the foremost northern red oak.in center foreground Tick clover dominated the sparsely populated but continuous herbaceous layer.
Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
56. Restoration of northern red oak and sugar maple- Interior of a second-growth stand of northern red oak with sugar maple as associate species on a moist north slope in the Springfield Plateau portion of the Ozark Plateau (= Ozark Highlands). The shrub layer was dominated by flowering dogwood with eastern redbud and young sassafras saplings functioning as associate shrub species.
Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
57. Tick clover in the understorey- Two species of tick clover or tick trefoil dominated the herbaceous layer of a second-growth northern red oak-dominated (sugar maple was associate) forest on a moist north slope in the Springfield Plateau of Ozark Hihglands. This photo-plot provided a close-up, detailed view of the herbaceous zone of the understorey of this forest that was recovering from logging several decades previously. One species of tick clover was was in full bloom and handily identifed as naked-flowered ticktrefoil (Desmodium nudiflorum) whereas the other species was obviously a late summer- or autumn-blooming species whose identity could only be guessed from its round leaves. The latter was perhaps smooth tick trefoil or beggar's lice (D. marilandicum) which is common on wooded slopes and uplands throughout the Ozark Highlands.
Also present at herbaceous level was s seedling of summer grape and a larger seedling of flowering dogwood.
Species composition of this forest range vegetation was that of (at least, closely approaching) the climax northern red oak and sugar maple associate forest. The Society of American Foresters description of the northern red oak cover type (Eyre, 1980, p. 44) explained that this forest range type was "probably subclimax", especially on north and east slopes. One species of Desmodium back) s
Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak).Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
58. Seeing red in the western Ozarks- Stand of second-growth northern red oak with various age/size classes ranging from large seedling to small sapling to large pole. There was a dead sapling of white oak, but otherwise no evidence of this often associated climax species. Trees in background included a high proportion of black oak along with northern red oak. A shrub-dominated lower woody layer was dominated by flowering dogwood with some scattered individuals of eastern redbud and large seedings and small saplings of sassafras. The major herbaceous species was the papilionaceous legume, naked-flowered tick trefoil. Individuals of another species of tick trefoil or beggar's lice was present which could not be positively identified and may have been smooth tick trefoil.
Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).Northern red oak is a component (ranging from dominant to minor species) in several forest cover types and lower hierarchial-level forest communities. Northern red oak has historically been regarded as Intermediate in tolerance, in particular with regard to shade (Burns and Honkala, 1990). This adaptation is similar to that of black oak (Q. velutina) although optimum habitat (growing site) for northern red oak is considerably more mesic than that for the frequently associated black oak. Other tree species that grow as associates of northern red oak such as sugar maple (Acer saccharum), basswood or American linden (Tilia americana), and the hickories (Carya species) are considerably higher than northern red oak in Tolerance rating.
For this reason, Tolerant and Very Tolerant tree species associated with northern red oak are typically dominants while northern red oak is an associate species in climax forest. The northern red oak cover type (SAF 55)--northern red oak is dominant--is most likely subclimax (Eyre, 1980, p. 44). This ecological condition is most pronounced on more mesic and fertile forest sites (eg. north and east slopes, in coves) where forest (canopy) cover is consequentially more dense or closed.
Examples of these more mesic (more favorable forest site) forest cover types that are dominated by Tolerant and Very Tolerant species (eg. sugar maple, basswood, hickory) and with northern red oak as a suboordinate or associate species were included in chapters, Oak-Hickory Forest-II and Southern and Central Forests-I (eg. Sugar Maple-Dominated Forests thereunder). An example of a mixed hardwood bottomland (floodplain) foresst in which northern red oak was an important associate species was treated under Southern and Central Forests-II.
The discriptive treatment that follows was of a second-growth forest with mature trees of northern red oak ranging from inidivduals in their "natural prime" (peak adulthood or optimum size for quality red oak lumber), older trees that were senescing ("past their prime"), snags (trunks of dead trees), down to a few saplings and fewer seedlings. In this second-growth stand the proportion, cover (crown and stump), plant vigor, and, most of all, reproduction (=regeneration) of northern red oak was declining. Northern red oak was persisting through minimal sexual reproduction, but as a species its time (years) of dominance were limited. The grand moment of northern red oak in this forest stand was coming to an end. As a species in this forest, northern red oak was in decline. By contrast, pignut or bittternut hickory was on the ascent and on its way to becoming the dominant of the developing climax forest. Almost all regeneration and, in fact, most increase in population of all trees was that of pignut hickory. Individuals of pignut hickory ranged from a few mature trees (still in prime adult stage of their life cycles) down through young adults, saplings, and seedlings. Most tree seedlings in this forest were those of pignut hickory. If--as seemed to be the case--the current successional trajectory held through to impending climax, pignut or bitternut hickory would be the climax dominant while northern red oak would slip to the successional status of associate species. Fewer of the older northern red oaks were being replaced by the youngest (seedling) generation of its aboreal kind. Regeneration (recruitment of new trees) in this subclimax oak-hickory forest was greatest for pignut or bitternut hickory. The successional game was still in play.
This tract of northern red oak-pignut (bitternut) hickory upland forest was second-growth, but most of the adult trees in this natural forest vegetation were essentially "fully mature". This meant that these individuals had "for all intents and purposes" achieved their maximum growth (greatest accumulation of wood and other plant tissue at any stage of their lives) and were at senescence, the final stage of their inidvidual life cycles. This was explained in the preceding paragraph. This condition was more the case for adult northern red oak that were older (apparently) trees and, certainly, of a subclimax species. As such, this second-growth forest had the now-dominant and forest type-defining tree species at the senescent stage of old-growth with regard to most individual trees. In effect this was an old-growth northern red oak forest. Likewise, the largest (and, presumedly, oldest) pignut hickory also were old-growth trees. In net sum, there was probably about as much annual loss of wood as of wood production. In essence, this tract was more an old-growth--though clearly not virgin--forest than it was not old-growth. It was a relict forest at subclimax to early climax stage of vegetation development.
From perspectives of lumbermen and professional foresters most of the northern red oak and the largest pignut or bitternut hickory should already have been harvested and the next wood crop being grown, and at a faster rate of wood accumulation. Alternatively, if the forest community on this tract of timber continued to grow and reach climax forest vegetation the upland forest site will be a hickory-oak forest (with pignut hickory the dominant and northern red oak the associate species) while the lowland site portion of the tract will progress to a American elm-hackberry climax.
Note on common names: the common name for Carya cordiformis has been shown variously as either pignut or bitternut hickory depending on which source one refers to (Vines, 1960, p. 132; Steyermark, 1963, p. 516; Harlow et al., 1979, p. 261; Eyre, 1980, p. 137; McGregor, 1986, p. 132) with bitternut hickory being more frequently used. In the section below (and throughout this chapter) the present author used both common names interchangably or often both were presented seperated by "or" or, alternatively, with one common name enclosed in parentheses (behind or following the other).
59. Outer edge- Margin of a second-growth, subclimax northern red oak-pignut (bitternut) hickory mesic upland forest in the western Ozark Plateau. This exterior edge of the forest was shown as a two-slide "nested photoplot" whereby the second of these two photographs was a closer-in view of part of the forest vegetation presented in the frame of the first (larger view) photograph. The large trees are northern red oak (two trees on left including the cneter one with forked, two large limbs, trunk) and pignut hickory (large tree on right). There were large seedlings and small saplings of both of these species, but there were far more young trees of pignut (bitternut) hickory which was the tree species that was winning the race to climax. Here along the exterior of the forest there were some small saplings of sassafras (Sassafras albidum), an erlier seral tree species, and seedlings of eastern red cedar (Juniperus virginiana). The latter is a non-sprouting conifer kept to a very low portion of oak-hickory forests in the Ozark Plateau by fire. This forest, near the boyhood home of the author (ie. he knew when it burnt), had not burned in over a half century.
Other woody plant species included two Rubus species, including common or black raspberry (R. occidentalis) and the other tentatively identified as Pennsylvania blackberry (R. pennsylvanicus). Virginia creeper (Parthenocissus quinquefolia) was climbing the trunk of the large pignut hickory. The major grass species were nimblewill (Muhlenbergia schreberi) and silky wildrye (Elymus villosus). The most abundant grasslike species were quill caric sedge (Carex tenera) and bur-reed caric sedge (C. sparganoides). Forbs were, in net effect, absent from this forest range.
Ottawa County, Oklahoma. Mid-June; late vernal aspect in an Extreme Drought (Palmer Index). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37). Dry-Mesic Chert Forest (Nelson, 2010, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
60. Inside the forest- Interior of a subclimax northern red oak-pignut (bitternut) hickory dry mesic upland forest in the Springfield Plateau, the western part of the Ozark Plateau. These two views--one under a full-sun sky (first slide) and the other under an overcast sky (second slide) feaatured the same trees and parcel of forest from two different camera angles. The foremost adult tree (center and right margin, first and second slide, respectively) with a healing wound from lost limb on upper trunk was an old-growth (= senescing) pignut (bitternut) hickory. Large trees behind (first slide) and behind and to left (second slide) were adult--and mostly senescing (ie. old-growth or overmature) individuals of northern red oak. Young trees from seedling to poles were almost exclusively pignut (bitternut) hickory, meaning that most regeneration (= reproduction) was of pignut (bitternut) hickory and nor northern red oak. It was obvious which tree species was winning the forest succession race and which "tree team" ) was going to be at the climax finish line as the dominant species.
There was a sapling of pignut (bitternut) hickory in immediate center foregtround of the first photographic view. By the way, from standpoint of forest development this sapling--and not the senescing old-growth adult--was the "star of the successional show". The second "photoview" featured numerous saplings and poles of various sizes of pignut or bitternut hickory while almost all visible northern red oak were old-growth ("long-past-prime") adults.
Plant species in the understorey were buckbrush or coralberry and Virginia creeper, co-dominant shrubs, and silky wildrye.
Ottawa County, Oklahoma. Mid-June; late vernal aspect in an Extreme Drought (Palmer Index). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37). Dry-Mesic Chert Forest (Nelson, 2010, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
61. Other inside views- Two "photoplots" of a subclimax northern red oak-pignut (bitternut) hickory dry mesic upland forest in the Springfield Plateau portion of the Ozark Plateau. The first slide was taken under an overcast sky while the second had a full-sun sky. This pair of "photoplots" featured forest regeneration--as did the immediately preceding pair of "photoquadrants". In the forest plant community shown in both sets of paired slides almost all forest reproduction (tree recruitment; on-going tree invasion) was by pignut or bitternut hickoy. There were some young trees (seedlings through poles and young adults) of northern red oak, but not enough to maintain northern red oak as the dominant tree (plant) species. By contrast, with pignut (bitternut) hickory there were more young trees (seedlings through young adults) than old-growth (mature to senescing) trees. This successional trend or pattern of species replacement indicated beyond any doubt that pignut (bitternut) hickolry was going to be the dominant plant of the climax forest with northern red oak likely the associate species on this upland slope forest site. (Progression of forest succession to a American elm-hackberry climax on the lowland portion of this forest tract was shown below.)
The first "photoplot" included three pignut hickory saplings in left to center foreground (the center one of which could be interpreted as a large seedling) along with a senescing adult pignut (bitternut) hickory. In comparison, the senescing northern red oak with a basal sprout (asexual reproduction) in right midground had no visible progeny. An interesting sidenote in this first slide was found with the small pole in right foreground. This was a black cherry (Prunus serotina), a species with a tolerance rating of Intolerant (Wenger, 1984), which had died. (The wonder is that the black cherry lived long enough to attain this size.)
The second "photoplot" justaposed the senescing adult pignut or bitternut hickory (in the "spotlight" and the same one as in the first slide) and a pignut hickory sapling with numerous senescing northern red oak and very few northern red oak saplings. Same conclusion as stated before: pignut hickory was going to be the climax dominant with northern red oak relegated to associate status (if that).
Buckbrush or coralberry and Virginia wildrye were the dominant woody plants of the understorey while silky wildrye was the major herbaceous species which was not plentiful enough to constitute an herbaceous layer in this portion of the forest.
Ottawa County, Oklahoma. Mid-June; late vernal aspect in an Extreme Drought (Palmer Index). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37). Dry-Mesic Chert Forest (Nelson, 2010, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
62. Leadership change- Two "zoomed-in" (closer-to-main-subject range) "photoplots" of a subclimax northern red oak-pignut (bitternut) hickory dry mesic upland forest in the western Ozark Plateau. These views of an upland slope portion of this forest tract provided a "textbook example" of species replacement along a forest sere with on-going forest development (plant succession). The large trees (some over two and a half foot DBH) were all old, senescing northern red oak. Most of these oaks most likely had some degree of trunk rot, perhaps some were largely hollow. There were numerous shed branches and, even, smaller limbs that were visibl in both "photoplots". All the young trees (ranging from small seedlings to saplings) were pignut (bitternut) hickory. In fact, there was almost complete foliar cover of the land surface by seedlings, many of them less than a foot in height.
Some of these shoots could have been root sprouts given that pignut or bitternut hickory is a prolific root and stump sprouter (Burns and Honkala, 1990). "Same difference" as the expression goes. The physiological fact of life was that pignut or bitternut hickory was reproducing--sexually and/or asexually--much more than northrn red oak.
Bitternut or pignut hickory was replacing northern red oak as the dominant of this upland oak-hickory forest. It was obviously becoming a hickory-oak forest. There were some saplings and small poles of northern red oak such that this species would apparently maintain a presence in the climax forest. At best, cover and density of northern red oak would be that of associate species. At current successional "trajectory" this forest was becoming a consociation of pignut hickory.
Carya species vary in their degree of tolerance. Burns and Honkala (1990) rated C. cordiformis as Intermediate, the same as northern red oak. Tolerance is, of course, site-specific to some extent. There was no doubt that on this upland slope form of a forest site pignut or bitternut hickory was much more tolerant than norethern red oak.
Ottawa County, Oklahoma. Mid-June; late vernal aspect in an Extreme Drought (Palmer Index). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37). Dry-Mesic Chert Forest (Nelson, 2010, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
63. Dominant grass by downed dominant tree- Rather sparse, local stand of nimblewill growing with some plants of common or black raspberry, possibly Pennsylvania blackberry, and buckbrush or coralberry on edge of tree cradle created when a large northern red oak was uprooted from water-soaked soil by strong winds. A tree cradle is a small crater or hole in the ground formed when a large tree is uprooted (or, at least, partially uprooted) as it is "brought low" by gravity or wind. The hole (which is often two to four feet deep) is a more favorable microhabitat for growth of certain plant species (often including the same species as that of the uprooted tree) due to a combination of temporarily ponded water and, thus, greater soil moisture content for longer periods, shelter from dessicatting winds, and protection from some grazing/browsing animal species (obviously not feral hogs who enjoy the nature-made wallow).
Nimblewill was a locally dominant native grass in the understorey of a subclimax northern red oak-bitternut hickory upland forest in the Ozark Plateau. Silky wildrye was the other major grass species on this forest range.
There was another tree cradle on this same forest range about about 40 yards away. It was shown below in another slide-caption set.
Ottawa County, Oklahoma. Mid-June; late vernal aspect in an Extreme Drought (Palmer Index). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37). Dry-Mesic Chert Forest (Nelson, 2010, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
64. Grass plot in the woods-"Photoquadrant" of the herbaceous understorey in a more open area of a subclimax northern red oak-bittternut (pignut) hickory upland forest in the Ozark Plateau. Nimblewill and silky wildrye swappd places as the local dominant herbaceous species on this forest range. Two important grasslike plants in the herbaceous layer were quill caric sedge and and bur-reed caric sedge. On this forest range forbs were "conspicuous by their absence".
Ottawa County, Oklahoma. Mid-June; late vernal aspect in an Extreme Drought (Palmer Index). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37). Dry-Mesic Chert Forest (Nelson, 2010, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
65. Forest on the move- Edge of a degraded tallgrass prairie and a subclimax northern red oak-bitternut (pignut) hickory upland forest in the western Ozark Highlands (Springfield Plateau). The subclimax forest community was expanding its perimeter outward and encroaching into the prairie (ie. the forest invading the grassland). Woody plant species in this local ecotone (transition zone between two range plant communities at local scale) included two species of Rubus, including R. occidentalis or black raspberry (and possibly R. pennsylvanicus), buckbrush, poison ivy/poison oak (Toxicodendron radicans= Rhus radicans=R. toxicodendron), sasafras seedlings and saplings, and lots of seedlings and small saplings of bittternut (pignut) hickory. For whatever reason(s) there were no seedlings or saplings of northern red oak.
Ottawa County, Oklahoma. Mid-June; late vernal aspect in an Extreme Drought (Palmer Index). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37). Dry-Mesic Chert Forest (Nelson, 2010, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
66. On the changing perimeter- Edge of a second-growth subclimax northern red oak-bitternut (pignut) hickory upland forest in the western Ozark Plateau. These two slides comprised a "nested photoplot" with the second photograph being a closer-in sample ("subplot") of the forest vegetation presented in the first photograph. These "nested" views were at a different location in the same transition zone between the oak-hickory forest and degraded tallgrass prairie introduced in the immediately preceding slide-caption set. The largest tree in right midground of the first slide was shown at a closer-up (shorter) distance in the second slide as the featured forest denizen-- along with its woody sidekick, a sapling of bitternut (pignut) hickory to immediate front of the northern red oak trunk, and a supporting cast of shrubs, including black raspberry ( possibly, also, Pennsylvania blackberry) and a seedling of flowering dogwood (Cornus florida) and herbaceous species, of which silky wildrye and quill caric sedge were local dominant grass and grasslike plant, respectively. In this second "photosample" there were also some plants of nimblewill and bur-reed caric sedge, associate grass and grasslike plant species, respectively (the order of herbaceous species as found in this view of developing forest vegetation).
The senescing adult of northern red oak and the sapling of bitternut hickory "personified" (encapsulated) the essence of plant succession on this forest sere. It was explained above that northern red oak was subclimax in this dry mesic chert forest whereas bitternut hickory was the tree species "ascending the seral staircase" to be climax dominant.All of the tree seedlings and saplings in these two slides were of bitternut hickory with none of northern red oak. This seemed to be especially successionally significant here on the forest perimeter (actually the local ecotone between this forest and degraded tallgrass prairie) where the subclimax species--and the tree species with most of the older adult trees--was not regenerating.
Several dead limbs and large branches of this northern red oak specimen were on the forest floor along with some that were hanging loosely from or still attached at base of the tree crown.
Ottawa County, Oklahoma. Mid-June; late vernal aspect in an Extreme Drought (Palmer Index). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37). Dry-Mesic Chert Forest (Nelson, 2010, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
67. Wood changes in the woods- Paired views of vegetation on the perimeter of a subclimax northern red oak-bitternut (pignut) hickory upland forest adjoining a degraded tallgrass prairie in the western Ozark Highlands. These two "photoplots" were centered on the same senescing northern red oak and bitternut hickory sapling that was introduced in the two slide-caption set immediately above. The sapling was to the immediate right of the oak trunk. There were three other ssaplings of bitternut hickory in right midground of the second slide (in front of several spike-bearing shoots of silky wildrye). There were no seedlings or saplings of northern red oak--no regeneration (reproduction) of the subclimax tree species--whereas there was abundant reproduction of the climax bitternut hickory. Successional replacement of the subclimax by the climax species was glaringly obvious.
The senescent stage in the life cycle of this aging northern red oak was vividly evident from tits numerous dead limbs lying on the ground as well as four "pins" (stumps or remnants of dead limbs) of the lower crown still attached to the trunk plus a dead branch balanced on the "pins". The stark contrast of the slowly dying (completing its life cycle) northern red oak and the relatively robust sapling of bitternut hickory exemplified the progression of plant succession as one tree species of lower seral order (subclimax northern red oak) was being replaced by a tree species of higher succeesional order (the climax bitternut hickory).
This forest vegetation included a lower woody layer consisting of black raspberry (and maybe some Pennsylvania blackberry) and a seedling or small sapling of flowering dogwood, this latter plant was growing to the immediate left of the large oak trunk. There was also a well-developed herbaceous layer here on the outer edge (perimeter) of this forest range that included silky wildrye (the locally dominant herbaceous species), nimblewill, quill caric sedge, and bur-reed caric sedge. There were no forbs in this herbaceous layer, which was shown in greater detail in the next slide...
Ottawa County, Oklahoma. Mid-June; late vernal aspect in an Extreme Drought (Palmer Index). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37). Dry-Mesic Chert Forest (Nelson, 2010, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
or with a slightly different sky (no overcast) 
68. Changes in the wood of the woods- A northern red oak in late senescence (and near death) being succeeded by sapling of American elm on a the lower and more mesic forest site in a dry mesic upland forest in the western Ozark Plateau. A hackberry-American elm-bitternuit (pignut) hickory forest was the apparent climax vegetation for this part of a tract of timber. On the lower (low-lying) portion of this tract the climax plant community was a hackberry-American elm-bitternut hickory forest.
The understorey of this climax mixed hardwood forest consisted of an upper shrub layer, lower shrub layer, and an herbaceous layer. The herbaceous layer was the best developed horizontal zone in this "photo-dendrogram". The dominant herbacous species was nimblewill. Silky wildrye and quill caric sedge were associate herbaceous species.
Northern red oak was a subclimax species on the less-sloping, lower-lying portion of this second-growth forest just as it was on the steeper, upper slope part of the same tract of relict forest.
Ottawa County, Oklahoma. Mid-June; late vernal aspect in an Extreme Drought (Palmer Index). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). No appropriate unit in Northeastern Deciduous Forest biotic community 122.1, Cold Temperate Forest and Woodland 122 of Brown et al. (1998, p. 37). Dry-Mesic Chert Forest (Nelson, 2010, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
69. Nimble if you will- Local colony of nimblewill (Muhlenbergia schreberi) on the floor of a low-lying, more mesic portion of a relict tract of dry mesic upland chert forest on which American elm and hackberry were succeeding northern red oak, the dominant of the subclimax forest. This unusually vigerous local population of nimblewill was growing in a tree cradle, a hole in the land surface formed when a snag of northern red oak fell (perhaps it was blown over by wind) and upon being uprooted left a yard-deep crater. Some seedlings of sassafras and several mature canes of black raspberry were also present in this extra-mesic microsite.
Another tree cradle in this same forest tract about 40 yards from the one shown here was presented above.
Ottawa County, Oklahoma. Mid-June; late vernal aspect in an Extreme Drought (Palmer Index). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash). SAF 93 (Sugarberry-American Elm- Green Ash). No appropriate unit in Northeastern Deciduous Forest biotic community 122.1, Cold Temperate Forest and Woodland 122 of Brown et al. (1998, p. 37). Dry-Mesic Chert Forest (Nelson, 2010, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
In the off-season- The tract of second-growth subclimax northern red oak-pignut hickory dry mesic upland forest shown immediately above in its vernal vestiture was next shown in "the dead of winter". In North America there is probably no other major unit of natural vegetation that changes as much in appearance with the seasons (ie. seasonal aspect), especially in color, as do forests of the Eastern Deciduous Rorest Region, the Deciduous Forest Climax:Querus-Fagus Formation (Weaver and Clements, 1938, p. 481).
Note on materials and methods: The Epson Perfection 700V scanner used to scan the following slides of Fujichrome Provia 100F did not, could not, would not, whatever scan them correctly, properly, or even "half-way right". Somehow this scanner repeatedly overexposed and produced a crystalline (or "frosted") appearence to the image. Creation of this crystal-like feature was a constant flaw in the Epson 700V scanner. The author did not have this problem with Hewlett-Packard scanners, but the HP equipment consistently turned green shades into blue ones. Apparently there has not been the economic incentive to "perfect" scanners to the degree achieved by digital cameras. Apologies to viewers for less-than-"perfect" images, but such is the state of technology (and of the market place) for presenting slides.
Slides still produce photographs that are superior to anything digital, although obviously not when viewed on the internet. Scanners are the equipment that limit quality of on-line photographs. Photographs on celluloid film when still be around--still intact and viewable--long after imaging technology has advanced to the next state of the art and the internet craze has passed. Conventional photographs, including slides, are an archivable (or archival, whichever adjective is correct), readily retrievable medium. that will remain available when computers capable opening a portable hard drive have passed into oblivion and the dust bin of obsolete inventions. Human eyes will still be able to see images on "hard-copy" photographs when the last compact disks become flakes of lacquer. This photographer was not willing to sacrifice photographic quality, permanent archives, and the convenience of storing information on slide mounts in order to "go digital" and present better images for his www. congregation. Be satisfied with half a load.
70. "Mature" in winter- A second-growth forest dominated by senescing (overmature) northern red oak with young adult bitternut hickory as the associate species in the western Ozark Highlands (Springfield Plateau) in early winter shortly after complete leaf fall. This view presented composition and structure of this second-growth, subclimax oak-hickory forest including representative plants of tree age/size classes including seedling, sapling, and pole-sized trees (as well as adults) of both of these tree species. Most of the larger trees were northern red oak except for the largest tree with lighter-colored bark in right midground. Most regenerationwas of bitternut hickory while reproduction of norther red oak was inadequate to permit continuing domiance by this species. Regeneration of northern red oak was ample to maintain some individuals into the climax stage, but as the then-will-be associate rather than the dominant species. Instead dominance was gradually shifting to bitternut hickory.
This successional development was consistent with the interpretation that the northern red oak cover type is generally subclimax (Eyre, 1980, p. 44). In this forest northern red oak was on "successional trajectory" to remain a major component of the climax forest though not as the climax dominant which, if this part of the forest is not disturbed by logging, tornado, or ice storm, will be bitternut hickory ( a slightly more mesic and certainly a more tolerant species).
Shrub and herbaceous species of the generally sparse and patchy understorey of this forest stand was described above and in the immediately following caption in which the accumulated mulch of leaf cover was also discussed.
This second-growth forest (based on recollections of long-dead local hill folk this forest had been logged over eighty and probably closer to ninety or a hundred years prior to time of photographs included here) was approaching the climax stage with the subclimax northern red oak being overmature such that this was almost an old-growth forest. In this locality most trees of northern red oak start to decline in their seventh and, certainly, in their eight decade of life. Fourscore years is old for man and red oak trees in these ancient hills ("wore-down-to their- roots" mountains).
Ottawa County, Oklahoma.Late December; early hibernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37). Dry-Mesic Chert Forest (Nelson, 2010, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
71. Inside in early winter- Interior of a subclimax northern red oak-dominated second-growth forest in western Ozark Plateau. On this slightly upslope oak-hickory forest almost all adult northrn red oak had advanced to old age (early to mid-senescence stages) with branches of their upper corwns dying, dead, and falling to ground (some of the latter visible at base of largest oak in center of slide). There were still some yuoung adult northern red oak as well as a few individuals in seedling, saplings, and pole age/size classes. In spite of representative northern red oak in these cohort categories, there was not enough regeneration of this species to permit it to remain the dominant forest species.
By contrast, bitternut hickory, the current associate species of this forest community, was the present major regenerating tree species with relatively greater numbers, cover, and density of trees in the various age/size classes. A few adult bitternut hickory were almost as large as senescing northern red oak, but these bitternut hickory were not yet at senescent stage. Bitternut (sometimes designated pignut) hickory was the obvious potential climax dominant of this subclimax northern red oak forest cover type. A nearly middle-aged bitternut hickory was in left foreground of this 'photoplot".
Examples of overmature northern red oak were presented at closer distance and greater density below where the terms overmature and old-growth were defined and explained.
In parts of this northern red oak-bitternut hickory forest there was a well-developed shrub layer dominated variously by buckbrush or coralberry, poison oak/poison ivy,and black raspberry in that order of overall shrub dominance. Herbaceous species were generally lacking in the sparse, sporadic understorey except along edges were either silky wildrye and quill caric sedge were dominant and associate species, respectively, or nimblewill was sole dominant graminoid species. Forbs were lacking.
It was explained below that it was not known if the accumulated dense ground cover by oak and hickory leaves (ie. leaf mulch) or nearly complete, light-excluding canopy cover was most responsible for elimination of herbaceous species. It had been a little more than half century since this forest tract had experienced a fire. In the author's boyhood years the forest had human-ignited, light surface fires every spring. He remembers that about forty years ago some plants of big bluestem grew to heights above his college-age head. (Of course, these senescing northern red oak and adult bitternut hickory trees were smaller and there was less canopy cover in those years.)
Ottawa County, Oklahoma.Late December; early hibernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37). Dry-Mesic Chert Forest (Nelson, 2010, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
72. Wintertime window- With shed leaves on the ground bare crowns of old-growth (and senescing) northern red oak and "prime adulthood" of bittrnut (pignut) hickory accompanied by saplings and pole-sized young trees of bitternut hickory were visible in early winter. This winter-decuduous view showed physiogonomy and structure of a second-growth stand of the subclimax northern red oak and climax bitternut hickory form of the northern red oak forest cover type, SAF 55 (Northern Red Oak). Species composition of this upslope Ozark Plateau oak-hickory forest was given above under photographs of this forest range in spring as well as in the immediately preceding caption.
Ottawa County, Oklahoma.Late December; early hibernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37). Dry-Mesic Chert Forest (Nelson, 2010, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
73. Oldsters and youngsters in early winter- Trees of all age/size classes of northern red oak and bitter or pignut hickory were present in this example of a second-growth Ozark Plateau oak-hickory forest presented in early winter to show (at least try to show given inferior scanning equipment) structure and composition of this example of the subclimax northern red oak forest cover type (Eyre, 1980, ps. 43-44). Most regeneration was of bitternut hickory which was obviously the climax dominant tree species for this upland forest site. Northern red oak regeneration appeared to be adequate to maintain some individuals of this species in the climax forest.
The large tree with ascending limbs (center of first slide) and left margin of secon slide) was a nice northern red oak just "past prime" with early progression to senescence including dyring upper limbs and apparent local decay in its lower trunk. Two close-together large tree in right midground of second slide were also northern red oaks though at smaller and what seemed to be younger ages (no signs of senescence). The state of overmaturity in forest trees was explained two captions below.
To the best of author's knowledge this forest had not had a fire for roughly a half century. The photographer as a woods roaming boy remembers this forest burning annually (each spring it was burnt off by the keper of a small city dump). Fire scars were still at bases of trunks of most of the larger northern red oak. As memory served there was far more herbaceous undergrowth--including of big bluestem--up to about 30 or 35 years prior to these photographs. It was not known if dense leaf cover or a closed crown cover was primarily responsible for loss of an herbaceous layer. This forest had not been grazed by any livestock during at least the last two-thirds of a century. White-tailed deer were common.
Ottawa County, Oklahoma.Late December; early hibernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37). Dry-Mesic Chert Forest (Nelson, 2010, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
74. Entering another winter- Species composition and structure of an upslope subclimax northern red oak forest in which bitternut hickory as associate species was in process of becoming dominant of the climax forest. The large tree in center was an old-growth specimen (an overmature tree) of northern red oak. This tree had grown a secondary shoot (left side of tree) from its trunk base, but this "sprout" had died. Small branches at base of this tree had fallen from the senescing crown of this showing-its-age northern red oak. The tree at left in midground or near background (left and behind northern red oak) was a bitternut hickory, the tree species that was producing most of the tree regeneration in this forest stand.
The terms old-growth and overmature as applicable to this forest were explained in the immediately succeeding slide-caption set. Composition of the sporadic understorey was also presented there as well as in slide-caption sets above.
Ottawa County, Oklahoma.Late December; early hibernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37). Dry-Mesic Chert Forest (Nelson, 2010, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
75. Lineup of the old-timers- Three northern red oak at ages and stage of life known as old-growth. Old-growth is a term more typically applied to a forest at advanced stages of succession, especially climax or subclimax state, and characterized by individual trees "that are large for their species and site and somtimes decadent (overmature) with broken tops tops..." (Helms, 1998, p. 127). Overmature was defined in Helms (1998, p. 130) as "a tree or even-aged stand that has reached that stage of development when it is declining in vigor and health and reaching the end of its natural life span"; also" an individual or stand "that has begun to lessen in commercial value because of size, age, decay, or other factors...". In common usage, old-growth and overmature are frequently used synonymously when applied to individual trees as, for instance, an old-growth specimen.
All of the trees in the forest view seen here were overmature in all senses of the term. The trees were senescing and several years "past their prime" from standpoint of tree vigor, fruit production (sexual reproduction), and lumber value. Lumber of northern red oak--a main species comprising wood known generically by the lumberman's term of "red oak"--is prized for everything from rough construction (as in barns and sheds) to church furniture. Had a wood crop been the management objective (which it was not) of this forest, the private property of an absentee landowner,these trees should have been harvested and their quality lumber put to commercial use. As it was they remained in the forest where they continued to produce a few acorns, furnish cavities for birds and mammals, rotting wood for insects (in turn food for such species as pileated woodpeckers [Dryocopus pileatus]), and an educational example of overmature trees in a second-growth oak-hickory that was progressing to the climax stage. This forest was clearly an old-growth forest state even though it was subclimax northern red oak and a second-growth forest.
The tree in the left background (in the "gun sight" between the second and third, from left to right, northern red oak) was a bitternut or pignut hickory that was still "in its prime" though approaching overmature state. The tree in background at far right was another senescing (overmature) northern red oak. While there were a few seedlings and sapling of northern red oak in this subclimax forest the predominant regeneration was of bitternut hickory which was obviously the climax dominant tree species for this forest site.
There were two shoots of fox grape (Vitis vulpina) climbing the trunks of these old northern red oak. Buckbrush or coralberry and Virginia creeper were the overall co-dominant shrubs, and silky wildrye and quill caric sedge were the co-dominant herbaceous species in the poorly and sporadically developed understorey of this forest. Other locally dominant understorey species were black raspberry and poison ivy/poison oak.
Ottawa County, Oklahoma.Late December; early hibernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37). Dry-Mesic Chert Forest (Nelson, 2010, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
76. Amid shed leaves- A single sun-deprived, stunted plant of quill caric sedge (Carex tenera), the dominant grasslike plant species in the almost-nonexistent understorey of a subclimax northern red oak forest in the western Ozark Highlands. The dominant herbaceous species in this oak-hickory (bitternut or pignut hickory was the associate and soon-to-be dominant tree species) forest was silky wildrye (Elymus villosus). In the absence of the natural or, at least, pre-Columbian, fire regimen in this forest seedlings and saplings of bitternut hickory, the potential climax dominant for this forest site, were so dense that, in combination with the crown canopy of adult northern red oak and bitternut hickory, there was not an herbaceous layer (in fact, almost zero herbaceous plant life). This was in contrast to a well developed herbaceous layer of silky wildrye, nimblewill, and quill caric sedge at edges of this forest stand (see again above).
Examples of quill caric sedge from this forest edge were presented shortly below.
Ottawa County, Oklahoma.Late December; early hibernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak). Oak-Hickory Series 122.11 of Northeastern Decicuous Forest biotic community of Cold Temperate Forest and Woodland 122 of Brown et al. (1998; p. 37). Dry-Mesic Chert Forest (Nelson, 2010, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
77. Fertile so far- Two spikes on shoots of silky or downy wildrye at peak anthesis. One shoot was presented with its flag leaf (leaf immediately below inflorescence). These shoots were on one plant growing in the understorey of a subclimax northern red oak forest. They had grown after a preceding wet winter and early spring , but were now in a Severe Drought. (Palmer Index).
Ottawa County, Oklahoma. Mid-June
78. Herbaceous quills- Shoots (basal through upper to point of inflorescence) of quill caric sedge (Carex tenera) in the understorey of a mesic (though stoney soil) northern red oak-pignut or bitternut hickory forest. Quill caric sedge was co-dominant with silky wildrye (Elymus villosus). Nimblewill (Muhlenbergia schreberi) was the associate herbaceous species.
Ottawa County, Oklahoma. Early June; fruit-ripening phenological stage.
79. Quilled shoots- Sexual shoots and overall morphology of inflorescences of quill caric sedge growing in understorey of a northern red oak-pignut or bitternut hickory forest. This is an uncommon Carex species in the western Ozark Highland (Springfield Plateau) where this forest range vegetation had developed.
Ottawa County, Oklahoma. Early June; fruit-ripening phenological stage.
Note on equipment: For this section there were 28 slides scanned by the Epson Perfection 700V scanner. Four of the 24 slides (14.3%) were scanned correctly (based on original color of slide). In other words 86% (almost nine out of ten) slides required Adobe Photoshop to restore color of scanned images back to original color. Of course, quality such as inclusion of all of images (ie. none of images being fadded out) was much worse. This was a representative sample of scanning quality by this Epson 700 scanner. Warning to fellow authors/photographers: never buy Epson equipment!
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).
80. 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).
81. 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 cowslop (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).
82. 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).
83. 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).
84. White oak-black oak-northern 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).
85. 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).
86. 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).
87. 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.Throughout large areas of the Ozark Plateau there are upland forests with various proportions of red, white, and black oak all of which are dominants (ie. forests of co- or multi-dominants). This is in contrast to other forest cover types in which any one of these three species is the sole dominant. Forest cover types of these three single dominants (SAF 53, SAF 55, SAF 110) were treated immediated above. In the following section the forest cover type with various combinations of these three dominants (White Oak-Black Oak-Northern Red Oak, SAF 52) was covered. In this author's observation, over much of the Ozark Highlands the more mesic white oak and northern red oak tend to be frequently associated with each other whereas the less mesic black oak tends to dominate the more marginal forest sites (stonier, more shallow soil; south slopes), and as the sole dominant tree--at least, the oak--species.
All four of these cover types (SAF 52, SAF 53, SAF 55, SAF 110) frequently develop in close proximity with each other. There is commonly conterminous occurrence among the four forest types as recognized by the Society of American Foresters (Eyre, 1980). Making distinctions among these can be somewhat arbitrary. An example of this situation was given above in the White Oak Type (SAF 52) that tended to be an Ozark forest co-dominated by white oak and northern red oak.
The current section presented a climax forest (in species composition though not at old-growth state) in which northern red oak and white oak were obviously co-dominant and black oak was not present, period. This example was on a north slope above a deep hollow.
88. Mixed oak Ozark forest-Local stands of white oak-northern red oak as a second-growth forest populated by young adult trees just "shy" of being prime logs for lumber or timber. There was regeneration of both of these tree species adequate to maintain them as the co-dominants of this dry-mesic, chert, upland forest (Nelson, 2010, p. 125-130) in the Springfield Plateau of the Ozark Highlands physiographic province.
There was a limited understorey beneath the complete canopy closure of this climax (though not at old-growth state) forest. In addition to sexual regeneration of the two dominants there were seedlings and small saplings of sassafras, a seral (often, pioneer) tree species and pignut hickory (Carya cordiformis), a climax tree of more mesic habitats. Examples of sassafras seedlings and a large sapling of pignut hickory were visible in foreground of the second photograph.
The major low shrub in these two stands was buckbrush or coralberry (Symporicarpus orbiculatus) as seen in immediate foreground of the first slide.For all practical purposes there was not an herbaceous layer in this vegetation, the heavy accumulation of leaf litter serving as a mulch that prevented emergence of grasses, grasslike plants, and forbs. (Herbaceous plants were present in some portions of this forest as can be seen in accompanying photographs.)
This forest vegetation was growing on the upper perimeter of a hollow, a local feature of the landform form of some ancient mountains like the Ozarks. Hollow as applied to highly eroded mountains was defined in the caption following the next one (two captions from now).
Ottawa County, Oklahoma. Early June (vernal aspect). 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).
89. Ozark mixed-oak forest nearing its prime- Textbook example of a white oak-northern red oak forest forest of the dry-mesic, chert upland form (Nelson, 2010, p. 125-130). The second of these two photographs was a closer-in view of vegetation in right midground of the first photograph. This was a second-growth forest of adult trees that had grown to about their maximum size before senescence and interior decay (trunk hollowing) set in. This mixed-oak forest had the species composition (and was approaching the structure and physiogonomy) of the climax stage for this forest site. Most of the adult trees in these two views had DBHs in excess of 18 to 20 inches with some of the larger northern red oak exceeding two foot. Even the largest trees had intact crowns with few broken limbs that could allow entry of decay causing organisms.
There was sexual reproduction amble for maintenance of white and northern red oak as co-dominants of the climax forest. There were trees of these two species in various age classes. A few of the large saplings of both oak species had died, presumedly due to less tolerance to conditions of this climax forest (ie. competition for light, soil moisture, root space, or whatever variables). This phenomenon was represented by a small snag of northern red oak bearing a conspicuous broken bole in far-right of first slide and, more prominently, in the second slide (growing among several larger trees). There was, however, a root sprout or sucker of this particular tree at its base bearing the immense shade leaves typical of young oaks growing in the forest understorey. Shade leaves are leaves produced in microhabitats of shade (of varying degrees of darkness or shadows). Shade leaves are much larger yet thinner (reduced leaf thickness) so that they are able to utilize larger proportions of the light available to them.
Also growing among the adult oak trees--right in the thick of them nonetheless--was a small pole of pingut hickory, one of the most tolerant and mesic habitat-adapted Carya species in this forest cover type. This young hickory was prominent in the second slide.
There were numerous larger seedlings and saplings of red maple (Acer rubra) scattered throughout this forest. A red maple seedling was featured in the immediate center foreground of the first slide. The most conspicuous shrubs in these two "photoplots" was Virginia creeper (Parthoenocissus quinquefolia) and buckbrush or coralberry, bogth of which were visible in immediate right foreground of both slides. There were no herbaceous species, hence no herbaceous layer, in samples of forest vegetation seen in these two "photoplots".
Ottawa County, Oklahoma. Early June (vernal aspect). 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).
90. On one edge of a hollow- A large white oak (left) and northern red oak (right) growing at the outermost edge of a hollow in the Springfield section of the Ozark Plateau in a dry-mesic, chert upland forest (Nelson, 2010, p. 125-130). The large seedling to small sapling growing "smack dab" between these two adult trees was red maple, a tree species that was just making its appearance in this second-growth climax (or approaching climax) forest. The leader with narrower serrate leaves at left margin was the shrub or small tree species, hop hornbeam (Ostrya virginiana) which is one of the most tolerant of all North American hardwoods. Hop hornbeam produces wood surpassed in hardness only by that of flowering dogwood (Kurz, 2004, p. 160).
The lower shrub layer was dominated by seedlings of climax co-dominants, white and northern red oak, along with buckbrush or coralberry and Virginia creeper. The thick leaf mulch and dense forest canopy prevented development of an herbaceous layer in most parts of this forest.(A few exceptional local areas of herbaceous growth were presented below.)
Uppermost outer edge of the hollow was at the right third or so in these two photographs.
The term hollow is one which many readers may be either unfamilar with or have a misunderstanding of. In its Glossary of Geology (Gary et al., 1972) the American Geological Institute defined hollow in part as "a low tract of land surrounded by hills or mountains; a small, sheltered valley or basin, especially in a rugged area". OK, but more specifically the term hollow (often pronounced "holler") is used by hill folk in southeastern and southcentral North America to connote a deep, almost gourge-like (albeit smaller) depression in the land thereby giving the impression of a small or local valley nestled within mountains. In ancient mountains like the Ozark Plateau, the mountains of which have been eroded down to their roots, the hollows are actually deep indentions in the general land surface rather than being of the "typical elevation" with mountain peaks having high elevations. This is more of a perception than a definition, but it most accurately captures the ambiance of the landscape (and hillbillies like this author).
Ottawa County, Oklahoma. Early June (vernal aspect). 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).
91. Approaching another edge- Straight-on view on upper "lip"or "brim", (the edge) of the hollow that was introduced in the immediately preceding two-slide set. In the western Ozark Mountains (Springfield scetion of Ozark Plateau; Springfiield Plateau) a climax mixed oak-hickory forest (dominated by white oak and northern red oak) of the dry-mesic, chert upland form (Nelson, 2010, p. 125-130) had developed around and in the interior of a deep and relatively large hollow.
The large tree in left foreground and the tree behind and closest to right of it were white oak. The tree in center midground and the larger tree at right midground with a lower leaning limb off right side of its trunk were northern red oak. The small pole-sized (or sapling-size) tree to the left and slightly in front of the lower, right-leaning limb was northern red oak.There were widely scattered shrubs and shrub-like oak saplings at mid-height throughout this forest range. One of these (ascending trunk in right front of the leftmost and largest white oak) was a hop hornbeam. There were several red maple of shrub-size to the right of the large leftmost white oak.
The lowest layer of this forest plant community was also made up of hardwood trees--as seedlings of both white and northern red oaks--but probably even more so of red maple seedlings as, for example, in lower right corner. Buckbrush or coralberry and Virginia creeper were common (though with sparse density and cover) lower-growing shrubs. The grape vine on the northern red oak could not identified to species.
Ottawa County, Oklahoma. Early June (vernal aspect). 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).
92. On the cusp of an Ozark hollow- Details of some of the forest vegetation introduced in the preceding slide were presented in these two slides that were taken a few steps off the rim and down into the hollow. The largest and foremost tree with a lower, ascending limb off right side of its trunk served as a "landmark" in reference to this immediately preceding photograph. This (and all the large trees, young to middle-aged adults) were northern red oak.
The shrub at far left margin (and with the trunk of a northern red oak serving as a backdrop in the second slide) was hop hornbeam, one of the most tolerant of all forest shrubs in North America. The low shrublike, sapling in center midground (to right of large, lower leaning-limb northern red oak) and several others behind it were all red maple. Sapling in right foreground in the first of these two slides was white oak. There were some adult white oak in background. All age classes of both climax Quercus species were represented although some saplings of these oaks had died (for whatever reasons; perhaps from competition).
Ground-level, the lowest layer of vegetation, was comprised of seedlings of both of these oak species plus some Virginia creeper, and buckbrush or coralberry as well as a few plants of Christmas fern (Polystichum acrostichoides) and some planrts of a Carex species (unidentifiable in its strictly vegetative state). Thus, in restricted microhabitats there was something of an herbaceous layer although it was sporadic.
Ottawa County, Oklahoma. Early June (vernal aspect). 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).
93. Growing on a cusp- Composition, structure, and partial physiogonomic perspective of an upland, climax white oak-northern red oak forest that developed at the head of an Ozark hollow. This second-growth forest was approaching the ultimte structure and physiogonomy of an old-growth forest (the state of a virgin climax forest). The largest trees were roughly middle-aged adults that in a few decades would be approaching their maximum size. These trees were thus nearing their "prime" from standpoints of quality lumber or timber products and still-rapid rates of growth. Species or botanical composition was clearly that of the climax forest vegetation. Old-growth structure had not been reached in this stand.
The three foremost (center foreground) trees in the first slide were northern red oak, the center and smallest (foremost tree) of which had died recently as evidenced by peeling bark at base of trunk and still attached dead leaves on lower branches.The adult tree in midground of both slides was a white oak. In the second photograph this white oak was positioned to the left of the big northern red oak in right midground. Three small saplings and one considerably larger sapling to the close-left of this white oak were red maple as was the large sapling situated between the two northern red oaks.
The conspicuous (in both of these slides) liana that had "clum to the top" of the left adult northern red oak was some grape that could not be identified from the lowest leaves which were at least 20 foot off the ground. This plant was likely summer grape (Vitis aestivalis). Most of the lower or ground layer of forest vegetation was seedlings of white oak and northern red oak.
Ottawa County, Oklahoma. Early June (vernal aspect). 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).
94. On around the "holler"- Another view of the large northern red oak (left margin) and white oak (as viewed here it positioned to immediate right of the northern red oak) and "attendant" plants seen in right midground of the immediately preceding slide. One more northern red oak and white oak were in the background (farther down in the hollow). The sapling in right foreground and another (and smaller) sapling at close-right to the foremost white oak were red maples.
Most of the plant growth in the lowest--the ground-level--layer of vegetation was made up of seedlings of the two oak species. There were even some seedlings of grape (unidentified Vitis species). Most of the ground layer was Virginia creeper. Buckbrush or coralberry was the next most common shrub. One plant of Christmas fern was to to close-right of the foremost northern red oak.
Ottawa County, Oklahoma. Early June (vernal aspect). 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).
95. Looking down deep- Head of a hollow in the Springfield Plateau section of Ozark Mountains on which a climax white oak-northern red oak forest had developed. These two more detailed ("closer-up") photographs presented the lower layers (understorey) of this forest range so as to show what few herbaceous species grew in the surface (ground-level) layer of vegetation. These two "photoplots" were of the same hollow shown previously except that in this local area there were a few plants of some caric sedge (Carex sp.), which in the strictly vegetative state could not be identified, and also of Christmas fern. A conspicuous plant of Christmas fern was featured prominently in center foreground of second slide. There were also several Christmas fern plants along with some of the caric sedge close to the trunk of the trunk of the northern red oak in the second slide. The photographer could not find any grass in this dry-mesic, chert, upland forest (Nelson, 2010, p. 125-130). The high tree density resulted in what, for all intents and purposes, was complete canopy closure (ie. 100% cover by tree crowns).
The two largest trunks (the only one in second slide) were northern red oak. The tree in center midground of both slides (and to left-rear of the big northern red oak in second slide) was white oak.
Ottawa County, Oklahoma. Early June (vernal aspect). 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).
General note: students should take note of (and make a mental note as to) general density or "spacing" of trees growing on the upper edge of the Ozark Plateau hollow featured above. Density is the number of individual organisms present in a specific area of space (land). An application of density in Forestry is the concept of stocking. In silvicultural usage stocking refers to occupancy of forest growing space relative to some preestablished standard. A closely related term used in Forestry is stand density which is either 1) a quantiative measure of stocking shown as number of trees, basal area, or volume of trees per unit of land area or 2) a measure of the degree of tree crowding in forests and shown as some unit relative to growing space (Helms, 1998).
The area of land per tree or relative proportion of sky (or canopy) occupied per tree was comparatively large in the white oak-northern red oak forest shown and described immediately above. Trees were not crowded. Instead they were of such density or stocking ("spacing") that crowns of individual trees were large yet without producing lower limbs in the fashion of shade trees. These trees grew as unbranched single trunks that were producing good sawlogs yet with abundant canopy so that they had relatively rapid growth.
With complete canopy closure the lower forest layers were limited to plants that have relatively high levels of tolerance. This included species rated as Tolerant to Very Tolerant (hop hornbeam, red maple, climax oak species, Virginia creeper, and among herbaceous species, Christmas fern).
In most soils of the Ozark Plateau rapid tree growth and production of large trees having high-quality wood requires low stocking or, in laymen's terms, wide spacing of individual trees.
A hybrid example-The following section presented and described a tract of second-growth forest that was part of the general Oak-Hickory Association (Clements, 1920; Weaver and Clements, 1938) or Oak-Hickory Region (Braun, 1950) in the westernmost Ozark (Springfield) Plateau. Although this second-growth forest was not a virgin forest, it had the composition and structure of the climax vegetation (for the oak-hickory Clementsian association or Braun region) including a high proportion of trees of the dominant species in or approaching the senescent stage of their life cycles. This senescence was likely hastened by three years of Moderate to Severe Drought (Palmer Scale). In other words, this forest was more-or-less at the old-growth stage of the forest community for this sere, for this forest site. It was the climax forest.
The plant community of this relict tract of semi-old-growth forest was a combination of two Society of American Foresters (Eyre, 1980) cover types: 1) SAF 52 (White Oak) and SAF 53 (White Oak-Black Oak-Northern Red Oak). More specifically, there were local stands or forest communities of white oak (SAF 52) within the overall mixed oak type (SAF 53). In parlance of Landscape Ecology, there were patches of white oak (SAF 52) within a matrix of white oak-black oak-northern red oak (SAF 53). There were, however, almost no black oak (Quercus velutina) trees in this relict forest.
Almost of slides of this relict forest community were general or overall views showing physiogonomy, structure, and composition of the approching old-growth oak-hickory. These photographs could be described or visualized as photographic profile diagrams or stratification diagrams (also sometimes referred to as a forest bisects) of the forest vegetation.
This forest had developed on gently sloping topography, but there was no slope aspect factor (no east versus west or north versus south slope variability). The forest was on a lower mid-slope above a valley (ie. a mesic, chert upland habitat that was nearly ideal for the more mesic oak species like white oak and northern red oak.
96. Oaks, white and red, in the southwestern Ozark Plateau- Four wide-angle views (20mm lens) of a white oak-northern red oak (co-dominant species) second-growth, mesic, upland forest in the westernmost Springfield Plateau. Most of the stocking in these four photographic forest profile diagrams was white oak with northern red oak was the lesser of two con-dominants overall. There was regeneration of these two co-dominants, but there were more seedlings and small saplings of bitternut or pignut hickory (Carya cordiformis) than of these two oak species. In some local areas there was seedling and small sapling cover of American elm.
There was a second woody layer or intermediate woody understorey of smaller trees (saplings and small poles) of the co-dominant oak species plus red mulberry (Morus rubra) and redbud (Cercis canadensis). There was a lower (basically ground) layer of woody species that included buckbrush (Symphoricarpos orbiculatus), Virginia creeper (Parthenocissus quinquefolia), poison ivy (Toxicodendron radicans= Rhus radicans= R. toxicodendron), and blackberry (Rubus sp.). Of course, the liana (woody vine) species like Virginia creeper and poison ivy sometimes grew to crowns of tallest trees such that these woody trailers occupied all three woody layers.
There was a very sporadic herbaceous zone (not really a layer per se) made up mostly of silky wildrye (Elymus villosus) and hairy or fuzzy wuzzy caric-sedge (Carex hirsutella). Forbs were lacking in this forest other than a local colony of Mayapple (Podophyllum peltatum) now and then.
For discussion purposes this second-growth forest was a climax forest in structure and composition. While the adult trees had not reached the classic habit (morphology or morphological form) of old-growth trees, many of the largest white oak and northern red oak trees in this forest were senescing at what appeared to be their natural end of life. Other trees of these two species were either dead or dying at what seemed to be younger stages of adulthood. It appeared as if this death and/or dying might have beeen caused--directly or indirectly--or at least exacerbated by Severe Drought (Palmer Index) following two years of Moderate Drought.
McDonald County, Missouri. Early June; late vernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF53 (White Oak). Cold Temperate Forest and Woodland 122, Northeastern Deciduous Forest 122.1, Oak-Hickory Series 122.11 of Brown et al. (1998, p.37). Ozark Highlands- Springfield Plateau ecoregion 39a (Chapman et al., 2002).
97. Sylvan scenes from the Ozarks-Interior of a white oak-northern red oak forest in the far-western Springfield Plateau of the Ozark Highlands (Plateau). The first (vertical) slide was a small, local stand of white oak the adult trees of which were either naturally senescing or dead/dying at old age which appeared to have been hastened by Severe Drought following two years of Moderate Drought (Palmer Scale). Interesting, however, was the fact that most regeneration at seedling to small sapling age/size classes was of bitternut or pignut hickory (and of all things!) American elm.
The second (horizontal) slide was a composite view with both northern red oak (the lesser of two co-dominant species) and white oak trees and with most--though not all--reproduction at seedling to small sapling age/size classes being of bitternut hickory and American elm. These two more mesic species (as a general rule) had produced these seedlings or shoots (perhaps offshoots from roots in case of bitternut hickory) during drought that appeared to be detrimental to adult trees of northern red oak and white oak. Time would tell as to survival of the 'baby" tres of bitternut hickory and American elm.
There was a prominent woody understorey in both of these "photoplots" that included red mulberry and redbud as larger shrub species as well as blackberry and buckbrush or coralberry as the main lower shrub and woody climbers, Both of the woody creepers, Virginia creeper and poison oak, grew from ground level to tree crowns so as to be present in all layers of this forest vegetation.
There was not an herbage layer per se, but there were scattered plants of silky wildrye and hairy or fuzzy wuzzy caric sedge in the understorey. The only forb of any consequence was Mayapple found in a few colonies.
These two slides were taken under a light overcast sky which eliminated shadows in contrast to slides taken under a full-sun sky as seen in the immediately preceding and succeeding slide-caption sets.
McDonald County, Missouri. Early June; late vernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF53 (White Oak). Cold Temperate Forest and Woodland 122, Northeastern Deciduous Forest 122.1, Oak-Hickory Series 122.11 of Brown et al. (1998, p.37). Ozark Highlands- Springfield Plateau ecoregion 39a (Chapman et al., 2002).
98. White oaks in the bright light of late spring- Local stand of white oak in a second-growth upland forest in the western edge of the Ozark (Springfield) Plateau in southwestern Missouri. Internal structure and composition of this white oak form of the Oak-Hickory Association (Clements, 1920; Weaver and Clements, 1938) or Oak-Hickory Region (Braun, 1950) was shown in these three "phototransects".
All the mature trees in these views were white oaks most of which were at the senescent stage of their life cycles (the stage that precedes death) or, else, near death. Two years of Moderate Drought followed by one year of Severe Drought on this cherty soil habitat might have been just a little too much for these oldsters. These trees were clearly approaching end of their life cycle; they did have (more-or-less) the size or aged shape of the typical (and few) old-growth specimens that can still be found in this area.
An interesting--if not unusual--phenomenon in this climax (and approaching old-growth stage) forest was that most--though not all--regeneration was not of white oak or its second-place co-dominant, northern red oak, but of bitternut or pignut hickory and American elm. This development took place during prolonged drought and it remained to be seen if these two slightly more mesic species would remain the new dominants. The obvious ecological question raised by this development was if the white oak-northern red oak forest was climax, or subclimax with climax being a more bottomland form with bitternut hickory and American elm the true climax co-dominants.
Given a historical fire regime in this forest (and this region generally) it was possible that the white oak or white oak-northern red oak forest was a pyric (fire) climax whereas the bitternut hickory-American elm was the climatic climax or the actual, true climax for this forest site. Traditionally, white oak has been interpreted as the climax cover type for moderately dry to mesic forest sites in the Ozark Highlands (Eyre, 1980, p. 43).
The understorey consisted of two woody layers with a 1) lower woody layer of blackberry, buckbrush or coralberry, Virginia creeper, and poison ivy, the latter two of which were woody climbers that grew up into tree crowns and 2) a middle or mid-layer of red mulberry and redbud. There was not an herbaceous layer, but silky wildrye and hairy caric sedge were the widely scattered by main herbaceous. There were a few sparsely scattered colonies of Mayapple.
McDonald County, Missouri. Early June; late vernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF53 (White Oak). Cold Temperate Forest and Woodland 122, Northeastern Deciduous Forest 122.1, Oak-Hickory Series 122.11 of Brown et al. (1998, p.37). Ozark Highlands- Springfield Plateau ecoregion 39a (Chapman et al., 2002).
99. Is what grows, what goes?- Two views "deep inside" a mesic upland forest of the Oak-Hickory Association (Clements, 1920; Weaver and Clements, 1938) or Oak-Hickory Region (Braun, 1950) in the far-western portion of the Ozark (Springfield) Plateau. This forest was the White Oak or a variant White Oak-Northern Red Oak cover type (Eyre, 1980) of the westernmost association of the Eastern Deciduous Forest Formation (Braun, 1950).
The large trunk in the left foreground of the first slide was white oak whereas that of the second slide was northern red oak. Trees in the background of both slides were mostly white oak with northern red oak well-represented. There were no trees of black oak (Quercus velutina) in this forest tract.
The woody understory consisted of two layerrs, the upper of which was dominated by red mulberry and redbud while the lower woody layer consisted of blackberry, buckbrush or coralberry, Virginia creeper and poison ivy (these latter two are lianas that routinely "clumb to the top" of adult oak trees). There was not an herbaceous layer, but there were scattered plants of silky wildrye and fuzzy wuzzy or hairy caric sedge.
This was a second-growth forest, but as was described above the number, proportion, and distribution of aged (senescing) and dead or dying adult white and northern red oaks indicated that this was an old-growth forest or, at least, one that was approaching the stage of an old-growth forest. Throughout this part of the Ozark Plateau your author has observed numerous oak trees of various species that died of old age in protected areas such as in home sites and inaccessible areas where log removal was impossible (logging was not feasible). Only a few of these cherished or inaccessable trees of northern red oak or white oak species that had lived to the old-growth stage and died of natural causes were much larger than the specimens found in this forest. Based on this empherical evidence it seemed most likely that adult white oak and northrn red oak in this upland, chert forest site had reached the end of their natural lives and attained their maximum size for the habitat in which they grew. It seemed an inevitable conclusion that these trees were old-growth individuals and the forest they reigned was an old-growth forest (though, being second-growth, not a virgin forest).
Two years of Moderate Drought and the current year of Severe Drought (Palmer Index) could have hastened senescence or death of these adult white oaks and northern red oaks. Contrary to this possibility was the fact that there was more reproduction of the more mesic-adapted bitternut hickory and American or white elm than of the co-dominant white oak and northern red oak. Time (at the scale of plant succession) will reveal if the two oak species would retain dominance or if forest succession was moving toward a bitternut hickory-American elm dominated forest more typical of bottomland forests in this region. Recurrent fire in this Ozark Plateau forest and general area (a fire scar in white oak was presented below) left open the possibility that the white oak-northern red oak forest was a fire type or subclimax to a climax (climatic climax) forest dominated by bitternut hickory and American elm that is sometimes found in bottomland or lower slope habitats in this part of the Ozark Highlands. (Examples of this latter forest cover type were presented in Range Types of North America in the Forest chapters, Southern and Central Forests and Oak-Hickory Forests-II.)
McDonald County, Missouri. Early June; late vernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF53 (White Oak). Cold Temperate Forest and Woodland 122, Northeastern Deciduous Forest 122.1, Oak-Hickory Series 122.11 of Brown et al. (1998, p.37). Ozark Highlands- Springfield Plateau ecoregion 39a (Chapman et al., 2002).
100. Internal organization with possible changes ahead- The interior of a white oak-northern red oak (the latter a lesser co-dominant, but more than an associate) mesic upland forest in the Spring Plateau of southwestern Missouri. There was a two-layer woody understorey: 1) intermediate woody layer dominated by redbud and red mulberry and 2) lower woody layer of blackberry, buckbrush or coralberry, Virginia creeper, and poison ivy with the two lianas covering much of the ground surface yet climbing into adult trees. Herbaceous species were largely limited to silky wildrye and hairy caric sedge. There were a few local colonies of Mayapple.
The interesting ecological lesson shown in these two 'photoplots" was that most regeneration was of bitternut or pignut hickory (eg. small sapling in cneter of both slides) and American elm. There was some regeneration fo the co-dominant white and northern red oaks, but the greatest number of smaller trees (seedling, small sapling, large sapling) was of bitternut hickory and American elm. This development was conspicuous after two years of Moderate Drought followed by the current year of Severe drought (Palmer Scale). Furthrmore, most of the adult white and northrn red oaks were either at senescent stage or, else dead or dying. This end of life stage could have been hastened by the drought. Persistence of bitternut hickory and American elm (to perhaps become the climax dominants) remained to be followed on the sere of this forest range. White oak was regarded as climax on moderately dry to moist forest sites in the Ozark Plateau (Eyre, 1980, p. 43).
Evidence of past and not recent fire in this oak-hickory forest (see presentation and discussion of a fire scar in white oak below) raised the possibility that white oak-northern red oak was either subclimax or a fire climax while the climax (= climatic climax) was the slightly more mesic species mix of bitternut hickory and American elm. Development of root sprouts (genotypic offshoots or clones arising from roots) is often common with bitternut hickory (Burns and Honkala, 1990) though there were only a few adult bitternut hickory in this upland forest.
Even though adult white and northern red oaks had not achieved the size and typical shape of old-growth trees sometimes found in this area, it was felt that adult trees in this forest tract had reached (or nearly so) their maximum development and survived to typical longevity or life span (based on tree size, shape, soundless of trunk and limb, etc.). Thus it seemed that this forest vegetation was the climax (or subclimax, fire climax?) for this forest site and that the forest, even though it was second-growth, approached the state of old-growth forest.
McDonald County, Missouri. Early June; late vernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF53 (White Oak). Cold Temperate Forest and Woodland 122, Northeastern Deciduous Forest 122.1, Oak-Hickory Series 122.11 of Brown et al. (1998, p.37). Ozark Highlands- Springfield Plateau ecoregion 39a (Chapman et al., 2002).
101. Different views, different light; same plants- Three slightly different views (different camera angle and overcast versus full-sun sky) of the interior of a white oak-northern red oak upland second-growth forest in the far-western Ozark Highlands. The co-dominant northern red oak was represented by the largest trunk with prominent butt-swell in center of the first slide and at far left (near margin) of the second and third slides. The co-dominant white oak was represented here by several trees (largely unidentifiable) in the background, large tree along right margin in second photograph, and the dual trunk (or two close-growing trunks) just to the right of the largest trunk that was the northern red oak noted in the preceding sentence.
The second woody layer or intermediate woody understorey included red mulberry and redbud . There was also a lower (essentially ground-level) layer of woody species that included buckbrush, Virginia creeper, poison ivy and blackberry, both primocanes and florocanes. Given that these woody vine species (Virginia creeper and poison ivy) can growth to tops of tallest trees these woody trailers occupied the three woody layers.
The major herbaceous species were silky wildrye and hairy or fuzzy wuzzy caric-sedge.
In the local areas shown here there was more regeneration of bitternut or pignut hickory and American elm than of the co-comnant white oak and northern red oak. This phenomenon in relation to forest succession was discussed above. White oak was interpreted as climax for the Ozark Plateau on moderately druy to moist sites (Eyre, 1980, p. 43).
McDonald County, Missouri. Early June; late vernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF53 (White Oak). Cold Temperate Forest and Woodland 122, Northeastern Deciduous Forest 122.1, Oak-Hickory Series 122.11 of Brown et al. (1998, p.37). Dry-Mesic Chert Forest (Nelson, 2010). Ozark Highlands- Springfield Plateau ecoregion 39a (Chapman et al., 2002).
102. Young whippersnapper among oldsters- A small-size sapling of bitternut or pignut hickory with large shade leaves in the understorey of a second-growth, upland oak-hickory forest in the western Ozark (Springfield) Plateau. It could be determined if this this shoot was of seed origin or if it arose vegetatively from roots (ie. was an offshoot of an older tree of the same genotype). This shoot was in the understorey of a generally dense-canopy white oak-northrn red oak forest.
In this second-growth forest there was more regeneration of bitternut hickory and, in some portions of this forest, white or Americn elm than of the current do-dominant white oak and northern red oak. American elm is usually rated as Intrmedaite in tolerance so it is probably not much more likely to reproduce in this forest than are the dominant oak species and the asociate bitternut hickory.
Tolerance of bitternut or pignut hickory varies from Intolerant to Intermediate, but its tendency to sprout prolificly gives it a competitive advantage over some other tree species such that bitternut hickory is capable of growing beneath moderately dense forest canopies. , especially on sites dry enough to restrict reproduction of more tolerant, but more fire- or drought-sensitive speciesfs
McDonald County, Missouri. Early June; late vernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF53 (White Oak). Cold Temperate Forest and Woodland 122, Northeastern Deciduous Forest 122.1, Oak-Hickory Series 122.11 of Brown et al. (1998, p.37). Dry-Mesic Chert Forest (Nelson, 2010). Ozark Highlands- Springfield Plateau ecoregion 39a (Chapman et al., 2002).
103. Why it' s called "oak-hickory"- Seedlings of white oak and bitternut or pignut hickory on the floor of a white oak-northrn red oak dominated upland forest in the far-western Ozark (Springfield) Plateau. In the first of these two slides seedlings of these two species were growing around a fire-scared white oak. The second slide presented two seedlings of white oak in front of some seedlings of bitternut pignut hickory.
Bitternut or pignut hickory is a deep-rooted, relatively wind-firm tree that sprouts readily such that it is adapted to certain forest disturbances, but this hickory is not fire-tolerant. Bitternut hickory is susceptible to fire at all stages of its life cycle from seedling to adult (Burns and Honkala, 1990). By contrast white oak is regarded as moderately resistant to fire according to online publications of the US Forest Service. The thick, corky bark of white oak has more insulating value than the smoother, tighter bark of such species as bitternut or pignut hickory.
One of the major reasons why white oak, a species of Intermediate tolerance, is a dominant forest tree species that can persist into and form a climax is that all stages of the white oak life cycle can survive under a forest canopy for for upwards of a century (Burns and Honkala, 1990).
McDonald County, Missouri. Early June; late vernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF53 (White Oak). Cold Temperate Forest and Woodland 122, Northeastern Deciduous Forest 122.1, Oak-Hickory Series 122.11 of Brown et al. (1998, p.37). Dry-Mesic Chert Forest (Nelson, 2010). Ozark Highlands- Springfield Plateau ecoregion 39a (Chapman et al., 2002).
104. Scar tells a tale; a hole teaches an Ecology lesson- Old fire scar on the basal trunk of a white oak in a second-growth oak-hickory forest in the Ozark Plateau. A surface fire that burned decades earlier injured the base of this white oak when it was a much younger tree. Some bark, cambium, and sapwood was killed by the fire followed by decay and loss of wood that left a this hole. Living tree tissue has gradually been closing around around the damaged area, buth a fire scar remained as this hole.
Decay of heartwood from fire injury is a major source of white oak loss (Burns and Honkala, 1990).
Some tree species are more tolerant of fire than other species. The native oaks are some of the more fire-tolerant tree species for a number of reasons. While most of these fire adaptations are probably still unknown one of the obvious factors is that oak wood, especially some of the white oak species (Leucobalanus subgenus of Quercus) like post and white oak, are more resistant to decay.
Fire scars like the one seen here were an indication that the white oak-red oak upland forest described in this section had a history of fire. Occurrence of fire was a likely reason that oak species rather than more tolerant species like American elm dominated this forest. It is likely that forests like this white oak-northern red oak dominated community was something of a fire type. There were indications that the whit oak-northern red oak was subclimax compared to American elm and pignut hickory.
McDonald County, Missouri. Early June; late vernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF53 (White Oak). Cold Temperate Forest and Woodland 122, Northeastern Deciduous Forest 122.1, Oak-Hickory Series 122.11 of Brown et al. (1998, p.37). Dry-Mesic Chert Forest (Nelson, 2010). Ozark Highlands- Springfield Plateau ecoregion 39a (Chapman et al., 2002).
The following sequence of slides and caption sets were devoted to one of the most legendary, well-known, and beloved of all the oak species: North American white oak (Quercus alba). White oaks are awe-inspiring trees of things of inspiration due to their often picturesque shape, immense size, long life, and overall beauty from bark to leaf. White oak is the State Tree of Connecticut, Illinois, and Maryland. In Silvics of North America (Burns and Honkala, 1990) Rogers said it well: "white oak is an outstanding tree among all trees".
White oak was a dominant of certain forest cover types (eg. SAF 52, SRM 53) in the virgin forests of the Ozark Plateau. Small fortunes were made from the sale of white oak, primarily for railroad ties and cooperage, mostly staves for whiskey barrels. In this way, white oak both helped build the infrastructure and bind solidity of the American nation as well as being central to one of its most persistent sins. Financial empires both large and small were built on the explotiation of Ozark forests, especially those in which white oak was a major species. For example, the largest landowner in the author's boyhood Ozark community made his fortune selling white oak for barrel staves. "Cut-and-run" logging was the order of the day. Some of these cut-over forest recovered and are managed using the science of proper-use forestry. Many more acres were converted to farmland (muck of which is now in permanent pasture of introduced forage species) or, ultimately, small residential areas (crabgrass disclimaxes).
Silvics of white oak was presented in Burns and Honkala (1990). Therein, white oak was rated as Intermediate in tolerance with trees of this species being able to respond well to release following disturbance such as logging or windthrow through both 1) persistence via the die-back mechanism (tree shoots can completely die yet resprout from stump and/or roots) and 2) seedling establishment. White oak is climax in the Ozark-Boston Mountains on other than mesic cove sites. White oak is a dominant in forests due to its persistence in understory as a more tolerant young tree, die-back ability, rapid growth following release, and tree longevity (Burns and Honkala, 1990). On many second-growth forests, most white oak is of resprout origin resulting in nearly "pure" stands of white oak with even-aged harvest being about the only silvicultural method capable of maintining such stands (Burns and Honkala, 1990). White oak does produce abundant acorn crops with subsequent ample sexual regeneration. This mast is readily eaten by livestock as well as most wildlife species.
105. Double-barreled- In this example of white oak (Quercus alba) bole and bark it was not known if the two trunks were two distinct trees (two genotypes) or two remaining stump sprouts (coppice shoots) following injury or felling of a previous tree shoot. White oak coppices prolifically from young stumps, but fails to resprout from stumps of older, larger trees with thicker bark (Burns and Honkala, 1990). These weer relatively young trees with trunks far from their potential maximum size.
McDonald County, Missouri. Early June.
106. Characateristic crown- Lower crown and upper bole of a young white oak in the Springfield Plateau of southwest Missouri.
McDonald County, Missouri. Early June.
107. Oak leaf cluster- The Oak Leaf Cluster is a military medal signifying vraiously bravery, loyalty, and distinguished service. The relevance or symbolic significance attested to the esteem with which oak trees were held. The fact that oak leaves symbolize such virtues as fidelity, courage, and "service above and beyond the call of duty" reflects oaks (Quercus spp.) as forest kings. White oak is one of the paramount species in this regard.
In less melodramatic contexts white oak leaves are symmetrical and moderately to deeply lobed and of compartively large size.
McDonald County, Missouri. Early June.
108. Galled in the woods- Leaves of white oak with conspicuous insect galls on these otherwise characteristic organs of photosynthesis. In writing the silvics in (Burns and Honkala, 1990) Rogers noted that damage from gall-forming insects was generally are "of minor importance".
McDonald County, Missouri. Early June.
109. Future prince white oak- Seedling of white oak in a white oak-northern red oak upland forest in Ozark (Springfield) Plateau of southwestern Missouri. Seeedlings of white oak develop long taproots giving them drought tolerance, but as these baby white oaks grown older the taproot atrophied and replaced by the peg root system characteristic of many hardwoods, including the oaks. As white oaks grow and age they develop large lateral roots so as to maintain a deep-root system (by oak standards) throughout life (Burns and Honkala, 1990).
This baby tree was growing in a year that went from Severe to Extreme Drought (Palmer Index). The two previous two warm-growing seasons had also been times of Extreme Drought. This worsening drought situation had apparently caused (or contributed to) death of neighboring adult white and northern red oaks.
McDonald County, Missouri. Early June.
110. A hairy one on oak leaves- Local group (an aggregrate, as in aggregated or clumped dispersion) of plants of hairy leaf caric sedge (Carex hircutella= C. complana var. hircutella) in understrey of a second-growth northern red oak-white oak forest that had the composition and structure of a climax forest of this cover type. This member of the subgenus Eu-Carex is a rather uncommon or infrequent species and it occurred in widely speced, extremely localized groups within this forest community. Hairy leaf caric sedge was, however, the dominant herbaceous species in an otherwise absent herbaceous layer.
McDonald County, Missouri. Early June; immediate post anthesis stage.
111. At bottom- Basal shoots at soil surface of hairy leaf caric sedge on floor of a northern red oak-white oak Ozark Plateau forest. This was the dominant herbaceous species in a second-growth forest at climax composition and structure. Hairy leaf caric sedge was very sparsely populated in this forest that was generally lacking an herbaceous layer at a stage approaching old-growth.
No livestock were on this forest range, but it abounded in white-tailed deer. There was no evidence of grazing on herbage of any plants of hairyleaf caric sedge in this forest.
McDonald County, Missouri. Early June.
112. Spikelets near the floor- Inflorescences with prominent spikelets that bear early stage achenes in hairy leaf caric sedge on floor of a northern red oak-white oak second-growth climax forest. The fruit type in Carex species is an achene which differs from a grass caryopsis in having the seed attached loosely (or not at all) to the fruit wall and with embryo embedded deeper in the interior of the seed instead of peripherally as in the grass seed.
McDonald County, Missouri. Early June; immediate post anthesis stage.
Presented in this section was an example of a post-climax forest community defined/distinguished by presence of American beech (Fagus grandiflora) as a dominant species within portions of the oak-hickory association. This forest range vegetation developed on atypically mesic, yet well-drained land in the Ozark Plateau. American beech was either the major dominant or co-dominant species along with white oak, pignut or bitternut hickolry (Carya cordiformis) and/or shagbark hickory (C. ovata ). The habitat for this beech-oak-hickory forest in the Ozarks Region is generally restricted to deeper drainages such corridors along tributaries to rivers such as the Buffalo and White Rivers of northern Arkansas. This deciduous forest range community is the potential natural vegetation that could be interpreted as a variant of the White Oak forest cover type (SAF 53). Alternatively, this forest veagetation could be regarded as an "island" or isolated southern extension, and in variant form, of the Beech-Sugar Maple forest cover type (SAF 60). Sugar maple was abundant in the example forest described herein, but oaks, including northern red and white oaks, and hickories were far more abundant and common than sugar maple. This was consistent with the Society of American Foresters description of a variant form of Beech-Sugar Maple (SAF 60) in which this cover type "grades into numerous related types according to degree of disturbance and changes in site". In the Midwest and southern Appalachians this forest vegetation "is commonly associated with uplant oaks" such as white oak, black oak, and northern red oak (Eyre, 1980, p. 34).
The forest portrayed below had developed on a soil (Arkana-Moka complex, 20-40% slopes) of generally less than a yard in depth overlying level-bedded limestone. Surface soil layers were generally very cherty and very stoney silt loams that overlaid clay to clay loam subsurface material. Soil was moderate in fertility and neutral to slightly acid or slightly alkaline in reaction (Soil Conservation Service, 1988, ps. 15-16). This forest had been logged--and it was high-grade logging at that--approximately 40-42 years prior to time of photographing and descriping it. Fire scars--some of them very deep and large--on almost all of the larger (hence, presumedly older) beech trees attested to the impact of hot fires on the present forest. Such fires selectively damages smooth-barked beech and sugar maple more than oak and hickory species. This selective defoliation was in all likelihood a major factor that contributed to relatively more oak and hickory and less beech that might otherwise have existed. It was also likely that secondary plant succession was on-going with the present state of forest community development at an advanced seral stage yet still below the ultimate state of development (eg. present forest vegetation was subclimax or preclimax).
This beech-white oak forest developed on mesic lowland along Clark Creek which forms a gouge-like tributary to Buffalo River. This was not a floodplain forest nor, strictly speaking, was it a bottomland forest. Neither was this forest range community riparian vegetation. In drier times of the year there is little water in the channel of Clark Creek because the water goes belowground through a fissure. The moist and fertile (though boulder-strewn) land of this drainage produced this unique forest that also included sweetgum, pignut (=bitternut) and mockernut hickories, basswood, sycamore, red mulberry, black walnut, red maple (and less sugar maple), northern red oak, sassafras, and both cucumber tree (Magnolia acuminata) and umbrella magnolia (M. tripetala). Shrubs included redbud, dogwood, spicebush, wild hydrangea, pawpaw, Carolina buckthorn (Rhamnus caroliniana), and American bladdernut. Remarkably little of Rubus species. Forbs included wild ginger, Solomon's seal, shooting star, great lobelia or blue cardinal flower, round-lobed liverleaf (Hepatica nobilis), pokeweed, trout lily, tall anemone (Anemone virginiana), jack-in-the-pulpit, green dragon, yellow-flower leaf-cup or bearsfoot (Polymnia uvedalia), columbine, bloodroot, trillium, sand phlox (Phlox bifida), and several kinds of ferns including Christmans, maidenhair, and broad beech fern (Thelypteris hexagonoptera= Dryopteris hexagonoptera= Phegopteris hexagonoptera). Main grass overall was Canada or hairy, wood brome (Bromus purgans= B. pubescens) along with bottlebrush (Hystrix patula). Also present but less common generally was broadleaf woodoats (Uniola latifolia) and Virginia wildrye (Elymus virginicus).
Braun (1950, ps. 33-34) clarified that a climax association that characterizes a forest region can also develop in other forest regions (which are characterized by other forest associations).In the Clementsian model used by Braun (1950, ps. 10-13) this beech-white oak-hickory forest could be interpreted as an example of a variant of the climax beech-maple association (that characterized the beech-maple forest region), but which occurred in the oak-hickory forest region or perhaps of the "transition from the oak-hickory to the oak-pine region" (Braun, 1950, p. 278). Perhaps most precisely by Braun (1950, ps. 170 and 172) interpretation this forest vegetation was an isolated community (an ecological island so to speak) of the Mixed Mesophytic Region or, at least, of the "typically mixed mesophytic aspect". In fact, it seemed likely that Braun described this exact tract of forest (which was in Lost Valley of Buffalo National River--previously Lost Valley State Park--near Ponca, Arkansas) decades earlier (Braun, 1950, p. 172). This conclusion could not be stated with 100% certainity, but the evidence was very strong that such was the case. Interpretation of this forest vegetation based on designations and descriptions of forest cover types by the Society of American Foresters (Eyre, 1980) was explained above.
This forest dominated by American beech-white oak (pignut hickory should probably be added for some portions of this climax vegetation) was a mesic lowland forest that had developed along a drainage that furnished a habitat substantially more moderate and more favorable for mesic species than that of adjoining environments. This should not, however, be confused with either 1) a bottomland or floodplain forest or 2) riparian vegetation. Clark Creek was not a perennial stream and it did not sustain moist banks for extended periods of time. Rather it was an intermittent stream the flow of which seeps through fissures and moves underground for much of the growing season. Flooding is also of such short and infrequent (even if violent) occurrence that vegetation along stream banks does not differ in meaningful composition or structure from that distant from the stream. In fact it is probably because this was not a floodplain that enabled American beech to grow here. It is textbook knowledge that beech (one of the most tolerant of North American hardwoods trees) does not thrive on either drier soils or those that are poorly drained (Harlow et al., 1979, 284-285). Beech has been found to be one of the most sensitive species to flooding (Burns and Honkala, 1990, p. 329).
Technical note: the photographs presented below were taken in early summer (mid-July) from mid-day to late afternoon (approximately 1230 to 1630 hours Central Standard Time). There was not so much as a wisp of a cloud in the entire sky. Thus shade and shadows represeted the minimum degree of darkness and the maximum quantity of light present in this subclimax to near-climax deciduous forestl. Neither conditions of direct overhead sun nor those of oblique angle of sun (again, both with 100% clear sky) resulted in optimum lighting conditions for photography. It bore repeating, however, that photographs were representative of the maximum light that could penetrate canopy (crown cover). Said another way, photograhic portrayl accurately represented conditions of maximum light intensity available to the understorey under full summer forest canopy. The phenomenon of sunflecks was shown to ideal advantage under these light conditions.
Low degree of cloud cover will sometimes diffuse light so as to produce uniforn distribution of light conditions (intensity, quality, etc.) which can reduce or eliminate shade. This, however, also reduces brightness (intensity) of available light necessitating lower f-stops and/or slower shutter speed which in turn results in less depth-of-field (hence less detail) of subject matter, especially of plant parts. Under conditions where cloud cover results in diffusion and more uniform light distribution and the consequence of reducing or eliminating shade there is also reduction or elimination of spaces that have full-sun (maximum birghtness). With such conditions of uniform distribution and reduced intensity of light in forests there is little or no opportunity to present examples of the sunflect phenomenon that is essential for photosynthesis below the forest crown canopy and consequent development of understorey layers, the grazable/browsable zone of forest range.
It is textbook knowledge as well as folk wisdom that when forests dominated by American beech are in in full-leaf they are still some of--if not the--darkest of all forests in North America. In this photographers' experience only north-slope forests dominated by the large-leafed bur oak have darker (dimmer) light conditions in the understorey. Slides of leaves and canopy of American beech and forest range communities dominated by this species portrayed the basis of overall dim light and sunfleck patterns characteristic of beech-dominated deciduous forests in the Ozark Plateau. Viewers should bear in mind that low visibility situations seen in most of the photographs presented below were representative of typical light conditions (ie. what they would encounter if they were present) in the dark, often dank, forest dominated by American beech.
Treatment of the American beech-white oak-hickory mesic forest began with overall or general external views of the forest and progressed toward the forest interior and understorey ending with photographs which presented details of important range plants and range microsites. Two different locations and distinct forest sites were presented. Both of these forest communities were distinguished by having American beech as a dominant along with oak and hickory species. The first forest tract featured was a forest community dominated by American beech on a north to northeast slope above Boxley Valley (Newton County, Arkansas). The second tract of forest was larger and was a much more botanically and structurally diverse forest community that had developed along the drainage of Clark Creek known as Lost Valley (Newton County, Arkansas). These forest range communities were in the Boston Mountains portion or section of the Ozark Plateaus physiographic province. The Boston Mountains exist as a dissected plateau that was peneplaned earlier than the rest of the province so that this is the oldest part of the Ozark Plateaus (Fenneman, 1938, ps. 655-660). Soils of the Boston Mountains formed from various parent materials with hill crests most commonly being of sandstone while lower strata are generally of chert/limestone.
113. Trees over all the hills and in the "hollers"- Exterior view in Ozark Plateau-Boston Mountains of oak-hickory forest with American beech, sugar and red maples, sycamore, black locust (Robinia pseudoacadia), shortleaf pine (Pinus echinata), and sweet gum (Liquidambar styraciflua) as species that were local associates to white and northern red oaks, the general dominants.
This general perspective presented the hills and hollows topography typical of the deeply dissected plateaus of the Ozark Plateau Region.
July. Buffalo National River, Newton County, Arkansas.
114. About as dense and dark as it gets- High noon in early summer in the interior of an oak-hickory forest in which American beech and sugar maple are the major associate species. In the first of these three photographs pignut hickory dominated a stand of mature trees on a north slope. The second photograph featured a second-growth stand dominated by American beech with pignut hickory as associate species on a northeast slope. The third slide showed dominance of beech in this stand (pignut hickory represented by one tree in right background). The second and third slides presented the very liited understorey which was comprised mostly of regenerating beech. This demonstrated the Very Tolerant tolerance rating of this species. In fact, presence of several age classes of beech attested to this feature.
Extent of shade on the forest floor at mid-day (1230 Central Standard Time) in early summer portrayed (clearly though darkly) the phenomenon of sunflecks and and the resultant extremely sparse layer of range vegetation on this shaded forest floor.
July. Buffalo National River, Newton County, Arkansas.
115. American beech, and nothin' else- A northeast slope above floodplain of the Buffalo River was naturally stocked by the single species, American beech. This was an example of a forest consociation. In the first of these two slides there was only one representative of another tree species. Can you find it? Hint: it is a pignut hickory. Note the different age classes of beech present in these two "photo-plolts". Viewers should also take not of relative area of the forest floor as to: 1) proportion of shade and 2) absence of understorey species other than beech. These photographs showed the maximum surface area of the ground receiving sunlight in early summer because pictures were taken at mid-day (1230-1245 hours Central Standard Time) when shadows cast by trees was at a minimum. The portion of shaded vs. sunlite ground surface also presented the concept of constanting changing patches of light caused by changing position of Earth relative to its sun, cloudiness (not shown on this cloudless day), and wind-caused leaf movement.
The scant plant cover and density at lower layers of this forest vegetation was largely due to the dense shade cast by beech (the specific epithet, grandifolia, was chosen most appropriately). It was likely however, that grazing/browsing by livestock in the not-too-distant past was also a factor in lack of understorey development. This steep hillside was now part of Buffalo National River, but it had previously privately owned land that was located immediately above an old barn (of sorts) and "cowpens". Any halfway knowledgable forest or range "detective" would have to assume that even a slope of this steepness had received relatively heavy stocking of cattle. Also, this area was some of the last legal open range (land open to the public to run livestock on and therefore subjected to free-roaming livestock) which included free-ranging, privately owned hogs. Swine are potentially the single most destructive species of range livestock on deciduous forest such as oak-hickory and oak-pine cover types. There was no denying the high probability that cattle and hogs had likely impacted this tract of forest in recent history of, say, 15-25 years ago. Nonetheless, much of the absence of understorey development was due to conditions of exteme shade.
July. Buffalo National River, Newton County, Arkansas.
116. Sign post (or "fools names in public places")- The trunk of an old American beech bore names and other graffiti left by vulgar, clueless scribes (probably urban and rural) who could not resist the temptation to use beech's characteristically smooth bark as sylvan space on which to "leave their mark" The mark left by literal sons of bitches (Canis familaris, C. latrans) quickly dries and, if not too frequently left, leaves little impact on the face of Mother Nature. By contrast, the heavy hand of vain-glorious, ego-driven, illbred, human SOBs bears a longer-lasting imprint of visitors to public property who violate the first rule of the woods, Leave a Clean Camp. Besides, unlike Daniel Boone, these self-centered wood carvers kilt no b'ars under thishere beech. Any claw marks left by the once native black bear (Ursus americanus) would by contrast be most welcome in this designated wilderness administered by the National Park Service.
Desecrationof public property aside, the identifying smooth gray bark and dense shade cast by American beech was presented in this Kodak moment on a steep north slope dominated by beech in the Ozark Plateau.
Another feature of American beech is root sprouting, development of secondary shoots from roots of mature and even uninjured beech trees. Most of the beech shoots in foreground of this photograph arose as asexual (vegetative) reproduction rather than as sexual reproduction from this tree, the bark of which that bore inscriptions from self-appointed, wood-carver scribes. More examples of root suckering (sprouting) were presented farther below.
July. Buffalo National River, Newton County, Arkansas.
117. A steep, beech-shaded slope- This steep, north slope was dominated by American beech underwhich shade-adapted plant species comprised some layers of forest vegetation. Root sprouts or secondary shoots (introduced in the preceding slide) and beech seedlings made beech the most abundant plant species at various micro-locations, but there were a few other understorey species including poison ivy, Virginia creeper, Christmas fern (Polystichum acrostichoides), and rolled sedge (Carex convoluta). The mature tree and two smaller trees closest to it on the right were beech as were most of the seedlings and small saplings, almost all of the latter of which were root sprouts of the adult tree.
July. Buffalo National River, Newton County, Arkansas.
118. Hard to see but instructive- Trunk of the largest American beech shown in the preceding photograph was shown in more detail in these two slides. One distinctive feature of beech--especially older, larger trees--is the self-exposed roots which in normal growth commonly protrude above the ground surface.This growth characteristic results in a fluted basal tree trunk.
Also presented in these photographs were the lower layers of forest vegetation--sparse though they were--that developed in the dense shade of a stand of beech. Viewers were again reminded that this photograph was taken at noon time (1230-1245 hours Central Standard Time) when light penetration to the forest floor was theoretically greatest overall (on average). Some plants of rolled caric sedge were visible in the second slide.
The phenomenon of sunflecks was presented here,, but shown to better advantage in the next slide. Another phenomenon displayed in the first of these two slides was that of geotropism, the directional movement (and subsequent growth) of a plant to gravity. This old beech was leaning so as to remain upright against the pull of gravity as exerted on this steep hillside (ie. a plant response to the stimulus of gravitational pull). There was also greater accretion of wood in the trunk on its uphill side in order to balance the entire shoot (bole and crown) of this beech which was large by standards of the Ozark Plateau, a marginal habitat for one of the most mesic of North American hardwood (angiosperm) tree species.
July. Buffalo National River, Newton County, Arkansas.
119. Light-grabbin' grasslike plant- A specimen of rolled sedge (Carex convoluta) at base of American beech on a shaded, steep north slope. This plant was growing close to the base of trunk and exposed roots of the American beech presented in the two photographs immediately above.This species is obviously one of the best adapted of sciophytes (= skiophytes; "shade-loving" plants).
July. Buffalo National River, Newton County, Arkansas.
120. Details of a shade-lover- Sexual shoots with fruit of rolled caric sedge. These shoots were on the same plant shown in the previous photograph. It was one of the specimens growing at base of the large and leaning American beech featured above.
July. Buffalo National River, Newton County, Arkansas.
121. Mixed mesophytic forest in Ozark Plateaus Region (Boston Mountains)- General and exterior view of the "mixed mesophytic aspect" (Braun, 1950, p. 170) of the oak-hickory association. It was explained in the introduction to forest range vegetation featured in this section that it was one of the more mesic (and restricted) forms of oak-hickory forest that develops in the Ozark Mountains. Dominance of American beech along with the typical dominant white and northern red oaks plus pignut (=bitternut) hickory in association with sweet gum, sugar maple plus occasional sycamore along with two Magnolia species as smaller (subcanopy) though defining trees described the specieas composition of this unusual climax forest that extended from bottomland to sloping uplands. This stream-drainage forest was in a deep hollow that eminated from tall sandstone bluffs (complete with waterfall and small cave) to terminate on the floodplain of the Buffalo River. The sandstone bluffs are visible in right background of this photograph.
The striking white bark (left midground) was that of a beech. Other dominant tree species were white oak, northern red oak, and pignut hickory.
Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Mixed mesophytic form of the oak-hickory forest with American beech as defining/distinguishing species and in association with sugar and red maple designated this as a isolated unit of a variant of SAF 60 (Beech-Sugar Maple) rather than as a variant of SAF 52 (White Oak-Black Oak-Northern Red Oak). Likewise, isolated unit of Beech-Maple Series 122.13, Northeastern Deciduous Forest 122.1 of Brown et al., 1998, p. 37). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004).
122. Pristine Arkansas- Mixed mesophytic form (Braun, 1950, p. 170-172) of oak-hickory forest. American beech as a local dominant along with sugar maple, sweet gum, and Magnolia species joined the area dominants, white oak, northern red oak, and pignut hickory, to produce a variant of Ozark Plateau deciduous forest that was interpreted as an isolated unit of the beech-maple association. The largest tree in these two slides was a white oak. The next largest tree and with large shade leaves (to left and rear of big white oak) was sugar maple. Most of the canopy in these photographs was American beech. There were small caves or cavelike recessions sunk into the sandstone bluffs behind trees.
This forest community developed within the hollow created by the drainage of Clark Creek.
Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Mixed mesophytic form of the oak-hickory forest with American beech as defining/distinguishing species and in association with sugar and red maple designated this as a isolated unit of a variant of SAF 60 (Beech-Sugar Maple) rather than as a variant of SAF 52 (White Oak-Black Oak-Northern Red Oak). Likewise, isolated unit of Beech-Maple Series 122.13, Northeastern Deciduous Forest 122.1 of Brown et al., 1998, p. 37). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004).
123. Boston Mountains beech- Local stand of American beech with sweet gum as the associate in the overall oak-hickory forest of the Ozark Plateau region. Although white and northern red oak along with pignut hickory were area dominants none of these species was represented in this local stand that had developed along sides of Clark Creek. In fact all regeneration (of which there was an abundance) was beech. This was both sexual reproduction from beech nuts and secondary shoots (suckers) from roots of established trees. Root suckering in beech was discussed below when forest gaps and patch dynamics were covered. Sweet gum is a seral species and its presence in this Clark Creek forest was due to persistence of trees that established about 40 years earlier following clearcutting of this forest tract.
Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect.
124. Things got hot (but they're cool now)- The personality of this beech attested to hot fire(s) in the past. This scar extended over eight feet up the bole of this second-growth specimen and penetrated through to the heartwood. Most trees survive this invasion (at least initially) unless such injury allows entry of pathogens that may later kill the tree, directly or indirectly. Heartwood and much of the sapwood is dead tissue anyway so that loss of such wood has no physiological impact on the plant. Instead the tree is weakened by loss of some support provided by the trunk. This frequently results in toppling of the tree when it is subjected to high winds: blowdown when the trunk is snapped, or, even, twisted off. Examples of such blowdown or windthrow (and the consequent forest gaps) were presented later in this section.
For now this beeech was still a "member in good standing" of this forest community. Much of the beech reproduction (which made up most of the regeneration shown here) was asexual in form of root sprouting (suckering). Throughout this section emphasis was frequently placed on the phenomenon of secondary shoot production by root sprouting in American beech. Some of the smaller trees presented here were seedlings showing that both sexual and asexual reproduction was successful in this forest. This was in spite--or perhaps because--of fire which may have initiated beech suckering.
Fire(s) that burnt this beech and its neighbors undoubtedly had influences on neighboring plants, for good and bad depending on species.
Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect.
125. Memories of hotter times- An American beech sports a long and deep fire scar that bears testimony of a once-upon-a-hot-time in these Boston Mountains woods. Like the example presented in the preceding photograph, the fire scar of this second beech had burnt through to the heartwood along entire height of the pyric injury. In fact, in both of these cases flames inside the trunk actually burned higher up inside the bole than on the outside. Also, these trees were healing around the injured area so that the scar was simultaneously "growing" as the tree grew in height and girth and also healing over as replacement growth enveloped the injury site.
The thin bark and big, protruding roots of beech make it one of the most susceptible hardwoods to fire (Fowells, 1965, p. 176; Burns and Honkala, 1990, p. 329). Thin bark also makes beech prone to sunscald, especially when trees in dense forest (hence protected from damage from sun's rays) are suddently exposed to brighter or even full-sun following logging. This species is also vulnerable to mechanical injury from logging and prunibng as well as damage by insects with piercing-sucking mouthparts (bugs) and diseases in general.(Burns and Honkala, 1990, p. 329).
As demonstrated by this and the preceding example some individuals of American beech can live with extensive fire injury as long as additional strrains from wind and gravity and stresses do not exceed physical strength and disease pathogens cannot gain entry through a compromised physical barrier of bark and associated tissues. It was not known if fire was a factor involved in root suckering.
Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect.
126. Innards of a botanical "island"- Composition and structure of an Ozark forest dominated by second-growth American beech. Isolated forest communities of beech-sugar maple occassionally develop in more favorable habitats of the regional oak-hickory forest (ie. isolated communities of the beech-sugar maple association within the surrounding oak-hickory association).Forest vegetation in these two photographs represented the interior of an American beech-dominated variant of the white oak-northern red oak-pignut hickory community of more mesic environments such as this one in the Boston Mountains of the Ozark Plateau region. The two photographs were horizontal and vertical views of the same stand to better represent species make-up and forest structure.
While beech requires more mesic forest sites than its co-dominants in this forest (white oak, northern red oak, pignut hickory) it also require better drained soils. Beech is one of the most flood intolerant species of all North American trees, especially flooding durnig the growing season (Burns and honkala, 1990, p. 329).
Almost all of tree regeneration in this stand was beech and not that of white oak, the major co-dominant, or sweet gum, a locally abundant associate that is a seral species sometimes persisting into advanced successional stages. This was also the case for some of the other stands presented below.
This forest understorey was comprised of both herbaceous and woody species, including beech seedlings and root sprouts as discussed above. The most abundant, though certainly not of great density or foliar cover, herbaceous species overall was hairy wood brome. In locally miacrosites the herb with most cover was broad beech fern. mmon.
Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Mixed mesophytic form of the oak-hickory forest with American beech as defining/distinguishing species and in association with sugar and red maple designated this as a isolated unit of a variant of SAF 60 (Beech-Sugar Maple) rather than as a variant of SAF 52 (White Oak-Black Oak-Northern Red Oak). Likewise, isolated unit of Beech-Maple Series 122.13, Northeastern Deciduous Forest 122.1 of Brown et al., 1998, p. 37). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004).
127. Renewable resources- That forests are some of man's most readily renewed natural resources was showed in this scene of a 40 to 45 year-old second growth forest of white oak and American beech that had regenerated along Clark Creek following high-grade logging less than a half century earlier.
This was also a good example of the species composition and community structure of this forest. Tree species in the foreground were (left to right): beech, white oak( middle and largest trunk), and sweet gum. Most tree reproduction was beech, and of several age classes, the same as the situation presented in the preceding forest stand. Shrubs included redbud, pawpaw, spicebush, flowering dogwood, and American bladdernut. Herbaceous species ranged from hairy wood brome to broad beech fern.
Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Mixed mesophytic form of the oak-hickory forest with American beech as defining/distinguishing species and in association with sugar and red maple designated this as a isolated unit of a variant of SAF 60 (Beech-Sugar Maple) rather than as a variant of SAF 52 (White Oak-Black Oak-Northern Red Oak). Likewise, isolated unit of Beech-Maple Series 122.13, Northeastern Deciduous Forest 122.1 of Brown et al., 1998, p. 37). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004).
128. A unique blend in Boston Mountains- A western mesophytic form of the oak-hickory forest association developed in a deep, narrow hollow that formed around Clark Creek which drained into Buffalo River. The forest stand shown here was on the terrace of this intermittant stream. The four young trees featured in the foreground were (left to right): sycamore beech, beech, and white oak. Beech dominated smaller/younger age classes as well as that of the largest trees (the canopy layer of this second-growth forest). Undrstorey species were primarily spicebush (a shrub) and pokeweed or pokeberry (rank-growing perennial forb).
This was another two-slide set presenting horizontal and vertical perspectives to present a greater sample of the understorey and structure of the forest stand up into the canopy. The herbaceous species with most foliar cover was pokeweed or pokeberry. There was some local spot dominance of the irregular herbaceous layer by broad beech fern. Most abundant shrub was spicebush.
Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Mixed mesophytic form of the oak-hickory forest with American beech as defining/distinguishing species and in association with sugar and red maple designated this as a isolated unit of a variant of SAF 60 (Beech-Sugar Maple) rather than as a variant of SAF 52 (White Oak-Black Oak-Northern Red Oak). Likewise, isolated unit of Beech-Maple Series 122.13, Northeastern Deciduous Forest 122.1 of Brown et al., 1998, p. 37). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004).
129. "Would you believe that this is Arkansas?"- "Shore as shootin'" this north slope along the deep drainage (a small hollow) of Clark Creek in the Boston Mountains supported a beech-dominated mixed mesophytic aspect (Braun, 1950, p. 170) of oak-hickory forest. In fact, more precisely this was an isolated representative community of the beech-maple forest association that developed within the overall oak-hickory association. However, sugar maple was quite limited whereas this was not the case for beech which dominated overwhelmingly the tree seeding and small sapling layer of (tree regeneration in) this forest community. Likewise, American beech dominated the layer of largest trees which were second-growth individuals less than 50 years old, this location having clearcut about 40 to 42 (maybe 45) years previously. In this mixed mesophytic aspect forest white oak was the close second co-dominant species to beech.
Ultimate triumph of beech as the ultimate climax dominant tree of this forest sere was already evident at this relatively short time following severe forest disturbance. Sweet gum is a seral species on this forest site that persist into climax or, at least, advanced seral stages. A representative individual of sweet gum was present in midground of this photo-quadrant (behind and to right of the ceenter and foremost beaech tree).
Shrubs in understorey included spicebush (typically the most common), pawpaw, redbud, flowering dogwood, and wild hydrangea (typically the least common). Forbs were essentially absent at this estival aspect of this lowland (creek drainage) forest, but wild ginger was the most abundant herbaceous species in forest range vegetation presented in these two photographs. The dark-brown piece of wood in the first of these photographs (near right margin; in front of a beech) was a fallen limb (species was not discernable) that apparently died while part of the forest canopy. Such fallen dead wood was a frequent phenomenon in this dense, mesic forest even though most of the largest trees were young second-growth. Dark woody vines in the second of these photographs (at least three were visible) represented numerous species of grape (Vitis sp?) that are common throughout the entire Ozark Plateau Region.
The forest composition and structure presented in these two slides was representative of that that developed from the foot almost to the top of north slopes. Beech was replaced to some extent by white oak and pignut hickory in upper parts of both north and south slopes (beginning lower on south slopes of course). These two photographs were of the beech-white oak stand that developed in a secondary drainage into Clark Creek, an intermittant stream.
Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Mixed mesophytic form of the oak-hickory forest with American beech as defining/distinguishing species and in association with sugar and red maple designated this as a isolated unit of a variant of SAF 60 (Beech-Sugar Maple) rather than as a variant of SAF 52 (White Oak-Black Oak-Northern Red Oak). Likewise, isolated unit of Beech-Maple Series 122.13, Northeastern Deciduous Forest 122.1 of Brown et al., 1998, p. 37). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004).
130. Composite shot of an American beech-white oak bottomland forest- A hardwood forest of mixed mesophytic form in the Boston Mountains included beech as a dominant (along with white oak, northern red oak, and pignut hickory which are the typical dominants of more mesic forest environments in the oak-hickory association). This beech-oak-hickory forest range developed in a small hollow through which a stream (Clark Creek) flowed intermittantly. Specifically the flow of water typically became subterranean during drier seasons (winter and mid through late summer, especially).Thus this forest range community was neither riparian vegetation nor a floodplain forest. Rather this was a very mesic (and periodically a wet) forest due to the sheltered environment of a deep, narrow hollow yet one with well-drained soil (very cherty/very stoney silt loams overlying a subsoil of clay to clay loam) and slightly acid to slightly alkaline generally fertile soil. It was this combination of edaphic features together with protection from more severe physical condition afforded by the deeply dissected topography that permitted dominance by American beech as well as typical dominant species of oak and hickory.
Most of the plants in lower layers seen in this photo-quadrant were seedlings of beech, first or most common, and white oak, second. Most numerous and with greatest cover in the lowest herbaceous layer was wild ginger, the overall most abundant forb in this mesic, stream-drainage forest at least in the estival aspect.
The most obvious shrub in this slide was some species of grape that was well-represented by conspicuous lianas.
Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Mixed mesophytic form of the oak-hickory forest with American beech as defining/distinguishing species and in association with sugar and red maple designated this as a isolated unit of a variant of SAF 60 (Beech-Sugar Maple) rather than as a variant of SAF 52 (White Oak-Black Oak-Northern Red Oak). Likewise, isolated unit of Beech-Maple Series 122.13, Northeastern Deciduous Forest 122.1 of Brown et al., 1998, p. 37). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004).
131. Arkansas as the first Arkansawyers knew it- American beech and white oak were co-dominants here on the upper portion of a north slope that was topped with bluffs and aligned parallel to an intermittant stream (Clark Creek. This co-dominant beech-white oak mixture was typical of most photographic samples of this mixed mesophytic aspect forest that developed in hollows of the deeply dissected Boston Mountains. Higher up near top of the bluffs sugar maple and wild hydrangea comprised a different forest community.
Much of the lower woody layers of the forest vegetation seen here was regeneration of the two dominant tree species, but spicebush, pawpaw, Caroling buckthorn, and American bladdernut plentiful so that shrub layers were well-repesented. Cucumber tree and umbrella magnolia were sparsely present and along with the more common flowering dogwood comprised an irregular taller shrub or even lower tree layer.
Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Mixed mesophytic form of the oak-hickory forest with American beech as defining/distinguishing species and in association with sugar and red maple designated this as a isolated unit of a variant of SAF 60 (Beech-Sugar Maple) rather than as a variant of SAF 52 (White Oak-Black Oak-Northern Red Oak). Likewise, isolated unit of Beech-Maple Series 122.13, Northeastern Deciduous Forest 122.1 of Brown et al., 1998, p. 37). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004).
132. A little "beech" of northern deciduous forest in the Ozark Region- In the Boston Mountains the mixed mesophytic aspect (Braun, 1950, p. 170) of the regional climax oak-hickory forest was dominated at low to above mid-level elevations of moist hollows by American beech. Some larger second-growth beech trees were featured. in this photograph to illustrate the interior of what is almost certainly either a post-climax form or, alternatively, an edahic or physiographic climax of the oak-hickory association in the Ozark Plateau. Forest vegetation like this is more like what would be expected in New England than in the southcentral forest region of North America.
It was remarked variously throughtout this chapter of the Oak-Hickory Forest that this anomalous (and, again, post-climax or edaphic/topographic climax) vegetation defied ready and unambiguous classification as to forest cover type. Given the widely scattered presence of sugar and red maple (mostly at higher elevations and in shelter of bluffs, especially of the former) this could be interpreted as isolated units of the Beech-Maple Association within the surrounding and otherwise unbroken Oak-Hickory Association. This general relation of what could be seen as "outliers" of other forest associations within masses of the regional association was explained by Braun (1950, ps. 33-34). It seemed plausible to the present author that this beech-dominant (or co-dominant, as with white oak) forest was a physiographic or an edaphic climax as suggested by Braun (1950, p. 34). Braun's recognition of such climaxes was her incorporation of Arthur Tansley's polyclimax concept into her general reliance on Clements' monoclimax concept which, of course, would view this beech-dominated forest in the Oak-Hickory Region as post-climax.
Whatever this forest vegetation was quite atypical (more mesic for one thing) for Ozark forests.
Most regeneration portrayed here was beech which showed to good advantage the rating of Very Tolerant for this species. Shrubs in this forest habitat within a deep, narrow hollow included spicebush, pawpaw, American bladdernut, flowering dogwood, and some Carolina buckthorn. The most common herbaceous species was wild ginger.
Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Mixed mesophytic form of the oak-hickory forest with American beech as defining/distinguishing species and in association with sugar and red maple designated this as a isolated unit of a variant of SAF 60 (Beech-Sugar Maple) rather than as a variant of SAF 52 (White Oak-Black Oak-Northern Red Oak). Likewise, isolated unit of Beech-Maple Series 122.13, Northeastern Deciduous Forest 122.1 of Brown et al., 1998, p. 37). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004).
133. Second-garowth stand of American beech- A spot in a mixed mesophytic aspect forest generally dominated by American beech and white oak in which beech was represented by various age classes which illustrated the Very Tolerant feature of this species and provided an example of a consociation. The small, dead limb in mid foreground had fallen from the largest beech providing an example of the self-prunning feature of this species. There were also grape vines growing up into crowns of beech in the background which should not be confused with self-prunned, hanging branches.
The understorey in this scene was rather sparse with broad beech fern being the most common herbaceous species. Beech regeneration made up most of the lower layers of vegetation.
Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect.
134. Some more light; some more biodiversity- Paired (vertical and horizontal) photo-plots showing understorey vegetation by a large second-growth American beech where there was a lower density of beech trees, saplings, and seedlings than was typical for the mixed mesophytic forest of which this local stand was part. The narrow terrace of an intermittant stream (Clark Creek) in the bottom of a hollow provided an unusually moist and shelted environment in which a "vegetational island" of beech forest had developed in which white oak, northern red oak, and pignut hickory varied locally in dominance with American beech. Sweet gum, sugar maple, sycamore, basswood, umbrella magnolia, cucumber tree, and mockernut hickory were some of the other larger woody species growing in this forest range vegetation. White oak, sugar maple, and sweet gum (this latter a seral species persisting into claimx or near-climax stages) were local associates growing immediately behind this matriarch beech. There was regeneration of beech, sugar maple, and white oak though primarily of beech, the reproduction of which was represented by several age classes.
The presence of beech and sugar maple (both Very Tolerant species) in age classes ranging from young adult to seedling suggested to this worker that the mesic forest which had developed in this extraordinarily favorable forest site was a local variant of the Beech-Sugar Maple forest within the surrounding regional climax of the Oak-Hickory Association.
Shrubs present in these two slides included spicebush and, in background, American bladdernut and pawpaw.Herbaceous species (both as to kinds as well as density and cover) were considerably more abundant in the more open space associated with this large beech. This microhabitat provided an example of ground cover by hairy wood (= Canada, woodland) brome and broad beech fern, the two species having greatest foliar cover, along with wild ginger, round-lobed liverleaf, bottlebrush, Virginia creeper, and poison ivy.
Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Mixed mesophytic form of the oak-hickory forest with American beech as defining/distinguishing species and in association with sugar and red maple designated this as a isolated unit of a variant of SAF 60 (Beech-Sugar Maple) rather than as a variant of SAF 52 (White Oak-Black Oak-Northern Red Oak). Likewise, isolated unit of Beech-Maple Series 122.13, Northeastern Deciduous Forest 122.1 of Brown et al., 1998, p. 37). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004).
135. Beech fern on bottom of beech forest- In spite of generally shaded conditions in the beech-white oak mixed mesophytic Boston Mountains the Almighty provided a small shaft of light in late afternoon that permitted sharing of this pteridophytic forb with loyal viewers. This broad beech fern (Thelypteris hexagonoptera= Dryopteris hexagonoptera= Phegopteris hexagonoptera) was one of a number of its kin that locally dominated--frequently to exclusion of all other herbs--the herbaceous layer in this post-climax or physiographic/edaphic forest. This specimen was growing at base of the large American beech featured in the two-slide set shown immediately above. The fern was accompanied by round-lobed liverleaf which was barely visible to the right.
Lost Valley, Buffalo National River. Newton County, Arkansas.
136. What a little light can do- Another set of paired photo-quadrants to present progressively closer-in views of understorey vegetation at base of another large American beech on the terraace of Clark Creek. This was another example of a floristically diverse local assemblage of woody and herbaceous species that had developed near a mature beech tree in one of the numerous hollows in the Boston Mountains. The species composition of understorey layers growing by this tree was similar to, yet distinctly different from, that living by the beech in the three preceding slides. In this local forest stand the more common understorey species were California or hairy wood brome, jack-in-the-pulpit, poison ivy, and Virginia creeper. Unlike the species mix of lower vegetational layer by the beech in the previous photo-quadrants this microsite had numerous seedlings of pignut or bitternut hickory, another defining climax tree species of more mesic forests of the Ozark Plateau.
It was reported above that the Boston Mountains have been regarded as being one of the more deeply dissected plateaus in the Ozark Plateaus Region (Fenneman, 1938, p. 655-660). These geologic dissections have formed hollows that in this part of the Boston Mountains furnish more protected and moist environments that support forest vegetation dominated and/or characterized by more mesic species that are typical of deciduous forests which are climax at more northerly latitudes. It iseemed plausible to this observer the combination of sandstone and limestone parent materials formed soils with extra favorable combinations of pH, fertility, water-holding capacity, drainage, and depth that were uniquely conducive to growth and survival of more demanding, persnickety species like the vulnerable beech which requires growing conditions seldom found in the Ozark Mountain Region.
In turn, beech provides a sheltering microclimate for understorey species that is also uncommon in the Ozarks. Larger beech trees typically had an understorey immediately beneath them that had less cover of woody plants, including, in some instances, fewer seedlings and saplings of beech (at least unless fire or some injury resulted in root suckering). Herbaceous species like broad beech fern, Canada or woodland brome, and jack-in-the-pulpit benefitted from this set of conditions.
Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Mixed mesophytic form of the oak-hickory forest with American beech as defining/distinguishing species and in association with sugar and red maple designated this as a isolated unit of a variant of SAF 60 (Beech-Sugar Maple) rather than as a variant of SAF 52 (White Oak-Black Oak-Northern Red Oak). Likewise, isolated unit of Beech-Maple Series 122.13, Northeastern Deciduous Forest 122.1 of Brown et al., 1998, p. 37). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004).
A wind storm approximately 7-10 weeks prior to time of this Clark Creek series of slides blew down (some uprooted; others with hollow trunks [probably due to fire injury] twisted off) some of the largest (presumedly oldest) trees of American beech and pignut hickory, two climax tree species. Forest gaps created by local blowdown provided photographic examples of patch dynamics. Three examples (adult trees) of beech and one of pignut hickory were presented in the following sequence. Interestingly, the species most apt to benefit from death of beech and hickory was beech and hickory, respectively. Each of these climax, tolerant tree species had already reproduced either adequately sexually (in case of hickory) with enough seedlings and pole-size trees to dominate the gap or (in instance of beech) abundantly, both sexually and asexually (clonal shoots from trunk and/or root) to monopolize the newly created patch. Thus there will be very little secondary plant succession in gaps. In case of beech, recruitment will be partly by new shoots of the previous genet (same genotype). This will be equivalent of new shoots regrown from rootcrowns, stolons, rhizomes of perennial grasses in a pasture or hayfield.
It was once felt "that beech root suckers do not development into
desirable trees" although some root sprouts could grow independently
of the parent tree (Fowells (1965, ps.174-175). This was accepted as
textbook knowledge (Harlow et al., 1979, p. 284). Subsequent studies
revealed that reproduction of this climax species "is almost exclusively
by suckering in the 'beech gaps'" in northeastern forest. Suckering
is also important-if not essential-in more northern and western portions
of this species' range (Burns and Honkala, 1990, ps. 327-328; original
studies cited and summarized). An example of this marginal extent of
American beech would be the Ozark Plateau in western Arkansas. It is
now realized that "[r]oot sprouts can develop into desirable trees"
(Burns and Honkala, 1990, p. 328). It is still accepted that stump sprouts
from other than smaller beech trees (less than four inches in diameter)
remain alive for only short periods
(Harlow et al., 1979, p. 284).
These two tolerant species were not only capable of reproducing in their own shade (generic, poetic description of any climax plant species), but also in patches of sunlite habitats formed by their removal. This undoubtedly explained to a large extent why this second-growth forest had returned to climax species composition in less than half a century following a logging operation that amounted-more-or less-to clcarcutting. Soil seedbank was also a factor especially in case of white oak.
An old folk proverb used as a play on words by Mr. Shakespeare taught its own lesson in Forest Ecology: "Ill blows the wind that profits nobody. (W. Shakespeare. 1591. Henry VI, Pt. 3 ii. v. 55]. Blowdown or windthrow of individual large, adult trees created gaps that could be exploited by other plants in lower strata of range vegetation which could then use light previously captured or excluded from lower forest layers by crowns of mature trees. In other words the philosophical saying, "Its an ill wind that blows no man good" is an ecological misnomer. Ain't no such wind.. In the mesic beech-white oak-pignut hickory forest featured here (and in all other forests), as in a Florida or Texas city following a hurricane, devastating winds always bring some good fortune, some ecological "profit", to those who can benefit from the misfortune of others and, in doing so, benefit the community. A tour of four such local scenes of distruction and reconstruction followed in Arkansas' Boston Mountains mesophytic beech forest.
137. Snapped right off- This large second-growth American beech provided the first of four examples of a forest gap and the pattern of patch dynamics in a mixed mesophytic forest in athe usually sheltered habitat of Clark Creek hollow. This and the next example of a forest gap were formed by a combination of 1) weakened trunks caused by fire damage and subsequent wood rot caused by organisms that gained entry through the fire scar and 2) strong winds. These two beech were broken off at the location of fire and rot damage. Wind was a necessary but not a sufficient factor for blowdown and gap creation.
Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect.
138. Broke on both ends- The second of three examples of blowdown of individual beech trees along the drainage of Clark Creek in the Boston Mountains of northwestern Arkansas. The same, spring wind storm and previous fire(s)-induced damage to the tree trunk that provided the preceding example of beech blowdown was responsible for breakage shown here. Both trees broke off in high winds at the fire scars, the site of fire damage and resultant wood rot, proving in these two cases that fire and wind were both necessary physical factors for tree loss. There were numerous other adult beech trees in this forest that had fire scars, some of which were much more extensive than the two examples shown here. These other trees which had considerably greater fire damage (two of which were presented earlier in describing this forest) were not broken off (or otherwise topppled) by these winds because they were farther down along the drainage of the stream and in locations that were more sheltered from wind.force.
The second of the two photographs of this second patch presented a local stand of white oak that were immediate beneficiaries of increased light made available due to beech blowdown. However these white oak were almost as tall as the downed beech so it was not clear how much the"lucky ones" of the main co-dominant species would benefit from light aspects of the newly created gap. There would certainly be more growing space at the edge of this gap along with additional water and soil mineral nutrietns. Edaphic resources may or may not have been limiting.
One obvious thing was that the white oak were "damn lucky" that the main limbs of the beech crown broke off due to their own weight on the way down and before reaching the imperiled white oak. (second photograph). "A miss is as good as a mile". Reminded the author of some "close calls" in his own life.
Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect.
139. Uprooted beech- A third example of a wind-downed adult American beech in the drainage (hollow) of Clark Creek in the Boston Mountains. This beech, unlike the two preceding examples, fell by being uprooted rather than by having its trunk twisted or snapped off at fire scars and site of rotten wood. In this third example wind without fire brought down an adult beech. This does not mean, however, that wind alone was a sufficient condition for gap creation. Rather it was a combination of saturated (or nearly so) soil and strong winds that uprooted this large tree. Water that had sustained this tree for decades and enabled it to reach maturity contributed to its demise.
While the mechanism of patch formation was different the resul, tree blowdown, was the same. Likewise, the general pattern of forest dynamics was the same. The same species of climax dominants--mostly American beech and, secondly, white oak--immediately filled the void formed by removal of light-intercepting crowns of adult trees of these same species. In this forest gap there were age classes of American beech extending from seedlings to larger saplings (two in right foreground of first photograph). There were numerous secondary shootrs arising from close-to-the surface roots of the downed beech that were not pulled up when the tree was uprooted.
There was nearly zero herbaceous vegetation in this newly created (about two month-old) forest gap. It was not determined at this time if there were stunted individuals of herbaceous species (say, perhaps, hairy wood [Canada] brome) that might be released later with more light. No such plants were noticed. Also, it was not known if annual species would colonize the gap in the next growing season. There was a small stand of young, pole-size white oak at far edge of the gap that were "all set" to benefit from the additional resources made available with loss of this large American beech.
Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect.
Clarifying note: All three of the beech trees blown down by wind (two broken off at fire scars, one uprooted farom wet soil) were growing on fairly level ground so that slope and gravitational force were not factors in these cases of windthrow.
140. Too close for comfort- Several beeches barely missed being brought down when this huge pignut hickory was bested by spring winds. How many gusts or even gales over course of its life had whistled through the crown of this monsterous member of a usually sheltered mesic forest that for decades had called the hollow of Clark Creek its home? This massive old hickory finally met its Waterloo when a combination of strong winds and saturated soil on a north slope was too much stress-and-strain for its massive trunk and crown that finally bite the ground that had sustained it through many previous storms.
What successional change did this forest gap, this hole in the upper canopy, initiate? Simply put: none. Beech, the most mesic and finicky of the climax tree species was the one with the most young adult trees to get immediate benefits of added light, space, etc. (first of these two slides; background of second slide). Adults of bitternut hickory were also poised to reap any rewards of increased light, spece, soil nutrients, etc. ((tree in middle foreground of second slide). There were also saplings and seedlings of pignut hickory that could benefit from light now capable of reaching to lower strata of the previously closed canopy. Also, bittternut or pignut hickory does develop root sprouts and its seedlings have a high shade tolerance (Burns and Honkala, 1990).
Yes, gaps in forest canopies can permit winds to "scoop" down into the remaining forest vegetation resulting in more windthrow of shallow-rooted trees which were able tor remain upright in large part due to the unbroken surface afforded by continuous canoy cover (the outer most layer of "skin" of the forest). Such is of little importance from the standpoint of forest succession when most of the woody layers are dominated by various age classes of the climax rtree species (beech, white oak, and pignut hickory in this forest). Any subsequent windthrow, and gaps created by wind disturbance, will amount to replacement of one age class of the climax dominants by another age class of the same and/or other climax dominants.
But the forest will miss this large and longtime resident of the community. Even trees do not live forever, at least not the same trunk and crown.
Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect.
"The rich get richer and the poor get poorer"- This popularized (and perhaps cynical) folk wisdom has been used for centuries. It became part of popular culture in 1921 when song writers Gus Kahn and Raymond Eagan incorporated the already well-known sentiment in the song, Ain't We Got Fun: "There's nothing surer: The rich get rich and the poor get—children". In the case of patch dynamics when and where American beech, the climax dominant of the mixed mesophytic aspect forest (Braun, 1950, p. 170), was "overthrown" by windthrow the vacated "successional throne" was immediately ascended and reclaimed by children of the rich, progeny of the climax beech. There was very little ecological "presidential succession" by other (presumedly seral) plant species. Instead root suckers (= sprouts; secondary shoots arising from roots close to the soil surface) emerged when released from suppression by apical dominance of the standing adult tree. There was also some sexual reproduction from beech nuts, but this was considerably less than the axexual or vegetative regeneration from root sprouting.
In the instance of blowdown of pignut or bitternut hickory, a co-dominant climax tree species of upper slopes, adult beech trees that were already established-- and luckily missed the crashing corpse of Carya cordiformis--enabled, even strengthened, continued dominance by the most mesic of the climax species. There were also numerous seedlings and saplings of pignut hickory to insure reclaiming of the climax crown by this regionally dominant species. Any benefit to pioneer and other seral species was short-lived. Some more lyrics from Ain't We Got Fun attested to this ecological fact in the mesic Boston Mountains forest: "There's nothing surer: The rich get rich and the poor get laid off".
On forest range that is either the most mesophytic variant of oak-hickory forest or an isolated and variant community of beech-sugar maple forest, creation of gaps by natural (and, apparently, human) disturbance does not cause denudation and retrogression that results in secondary plant succession. Instead asexual and sexual offspring of the climax trees "fill in the gaps". Patch dynamics is overwhelmingly replacement of adults of the climax tree species by clones and new genotypes of these same mature shoots. Wind and chainsaws merely mow off adult standing crop (both biomass and necromass) which subsequently regrows by resprouts and new trees of the climax species. Plant succession essentially does not exist in patches (including clearcuts) created in this forest dominance type.
141. Even unto death- (or still a part of the forest)- Log of a downed American beech providing a protective and moist, rich microsite for spotted touch-me-not or jewelweed (Impatiens capensis). A big beech fell across a walking ttrail and a trail crew with the National Park Service bucked it up and rolled the rounds off the trail into this neat pile which provided the improved habitat for this forest forb (one of the more mesic forb species in the Ozark Plateau Region). It was also habitat for some species of shelf fungi as its decaying wood provided a substrate for this species that is a decomposer in this forest ecosystem..This shot also showed the role of ecosystem reducers.
Even at end of its life cycle the woody carcass of a tree, especially a big one that takes longer to rot, remains a part of the forest ecosystem and continues to provide "goods and services" in the forest range. Not only did the rotting beech log furnish habitat for the producers and reducers shown in theis photograph it also undoubted provided shelter and a food source for soil-swelling consumers such as earthworms and arthropods like centipeds. Nutrient recycling, soil protection (erosion risks are high on the steep slopes above streams like Clark Creek), and water retention (the decaying wood holds and slowly releases water back into the hydrologic cycle) are all important roles served by dead wood on forest range. Benefits of dead and downed wood to forest watersheds were illustrated in this final stage of a forest tree.
Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect..
142. One of the few grasses in a beech-dominated forest- Bottlebrush (Hystrix patula) is one of very few grass species that can survive in the dense shade of an Ozark Plateau forest dominated by American beech. Canada (= hairy wood brome), most common, broadleaf woodoats (Uniola latifolia), and Virginia wildrye were the only other species of grass this rangeman could find in this dark forest.
Divine Providence furnished a rare and very fleeting shaft of "heavenly sunlight" that enabled this hapless photographer to bring his loyal viewers this gramineous or graminaceous (both adjectives are correct) delight.
Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect.
143. A study in American beech (Fagus grandifolia)- A mature beech at old-age including some sucker shoots arising from surface roots. Leaves and the smooth, gray bark of beech were also presented in this composite shot. Buffalo National River. Newton County, Arkansas. July.
144. Shady details- Close-up photographs of basal trunk of the American beech presented in the preceding photograph. The photograph presented details of beech bark and (in front of trunk) a secondary shoot arising from one of the upper roots that was naturally protruding above ground surface. Second photograph showed this entire root sucker.
The thin, smooth bark of American beech makes it very susceptible to fire injury and sunscald (Burns and Honkala, 1990, p. 329). Sunscald refers to the condition of "localized injury to bark and cambium caused by a sudden increase in exposure of a stem or branch to intense sunlight (insolation) and high temperatures" (Helms, 1998). Certain smooth, thin-barked tree, including both beech and sugar maple, are also susceptible to winter sunscale which can develop on the side of trunks and limbs that are exposed to warmer temperatures and brighter light of winter afternoons.
Buffalo National River. Newton County, Arkansas. July.
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145. Leaves of American beech #1-Leaves and twig on the root sucker of beech introduced in the preceding two slides. Northeast slope above Boxley Valley in Boston Mountains. Phenomena of tolerance, shade leaves, and sunflecks was explained in the two-slide set presented immediately below.
Buffalo National River. Newton County, Arkansas. July.
146. Leaves of American beech #2- Another (the second) example of a beech root sucker. This specimen was growing in the terrace (broad stream bank) of Clark Creek that drains into Buffalo River. Both this and the previous example of beech leaves also showed the extremely shady conditions produced by the dense foliage of American beech and high this Very Tolerant species can grow and have successful vegetative (asexual) reproduction in the dim microenvironment beneath its own dense foliage.
These two photographs (and the two photographs shown immediately above) were taken using natural light at mid-day (1245 hours Central Standard Time). This photographer had to wait until a shaft of light from the appropriate direction struck the leaves of these root sprouts. This was for a brief period lasting no more than a quarter hour. Most likely there were other such brief periods when the orientation of Earth to Sun permitted streams of light to strike these leaves. This is the phenomenon of sunflecks, several examples of which were presented in this section. Obviously these-more-often-shaded-than-sunstruck beech leaves were, on average, more sinks than sources of photosynthate. These lower leaves almost assuredly had to be supported by leaves in the crown which functioned as photosynthate sources for these energy and nutrient-dependant organs.
Leaves on these beech root sprouts (secondary shoots arising from near-surface or even protruding roots) were example of shade leaves, those which due to their position in shaded environments are larger and thinner than co-hort sun leaves that are in environments which receive full-sun light (even if on the same tree).
Buffalo National River. Newton County, Arkansas. July.
Final footnote: presence of white oak as co-dominant with American beech (rather or more than with sugar maple, basswood, pignut hickory being co-dominants) probably reflected spatial contact with the climax white oak-northern red oak upland (south-slope; ridge top) forest contiguous with the local mesic environment afforded by banks of Clark Creek and north-slope limestone bluffs aligned with this stream. It was not clear if white oak trees would live to such age as to remain a component of ultimate climax-as in old-growth-forest or if white oak was of high-seral status. If this latter was the situation then this mixed mesophytic aspect (Braun, 1950, p. 170) forest could be viewed as subclimax with even greater dominance by beech as the topographic or edaphic climax (polyclimax model) or post-climax (monoclimax model). It did not appear to have structure (and perhaps not function) of true virgin (=old-growth) forest for this forest site and cover type, but there was not an old-growth stand of this (or similar) forest cover type with which to compare the forest discussed above..
The next portion of this section described an upland white oak-northern red oak forest that was contiguous with the beech-white oak-hickory forest treated immediately above. This oak forest range community was similar to (and could be considered as a vaiant of) the white oak-black oak-northern red oak cover type dealt with earlier in this chapter. Near absence of black oak, one of the most widely distributed Quercus species throughout the Ozarks Plateau Region, combined with the conterminousness of this forest tract with that of the beech-white oak forest in the hollow (drainage) of Clark Creek just covered made inclusion of this white oak-northern red oak forest here, rather than above, a logical arrangement.
147. Rainbow of hardwoods- Mesic upland mixed oak forest with a wide array of woody angiosperms. An upland white oak-northern red oak forest immediately above the mixed mesophytic beech-dominated forest that had developed along Clark Creek provided a rather stark contrast (over a remarkably short distance) of two forest range types in which oak species were climx dominants to one extent or another.
In this upland oak forest the species composition of shrub layers was in rather striking contrast to that of the more mesic forest deep in a stream drainage. From one perspective this difference was as dramatic as absence of adult beech from the upland forest (next paragraph). Differences between these two adjacent forest cover types--again,one a lowland and the other an upland community-- were in both kinds of shrubs and structure of the shrub strata. Mesic shrub species of pawpaw, wild hydrangea, and American bladdernut were absent from the upland mixed oak forest. Instead the defining shrub, lowbush huckleberry (Vaccinum vacillans), was widespread in the upland and absent (or nearly so) from the lowland beech-dominated forest. Also more common in the upland mixed oak forest, and essentially absent from the mixed mesophytic beech forest, was sassafras and buckbrush or coralberry. Also present in the upland and absent (at least rare) in the lowland forest was eastern red cedar (Juniperus virginiana). Absence--undoubtedly exclusion (one way or the other)--of fire clearly accounted for presence, and increasing abundance, of this nonsprouting juniper. Sweet gum was not found on the upland forest whereas it was an indicator tree species in the mexic beech-dominated forest below.
One of the most telling signs (though what it told was not clear at this point in development of vegetation) was absence of large adult beech from the upland. Beech was preent in the upland white oak-northern red oak forest but only as widely scattered seedlings and small saplings. It was not knownif these would survive to adulthood.
Flowering dogwood was the most abundant of the taller shrub species (foremost trunk; midground in first slide) followed--as distant runner-up--by red maple. As already noted, lowbush huckleberry was the most common of lower-growing shrub species. Sassafras was the most conspicuous shrub in the second slide (left foreground with single bole). Sassafras is a species that occurs as both shrub and tree, and as small to mid-size tree.
Conspicuous specimens (tall, straight, clear trunks) of white oak and northern red oak were present in both of these photographs. The second photograph was a zoomed-in view of the two largest representatives of these species introduced in the first photograph. This proved to be an advantageous arrangement to present the two subgenera of Quercus. The two large trees in midground of the first photograph and then featured in the foreground of the second photograph were white oak (left) and northern red oak (right) representing Leucobalanus and Erythrobalanus, respectively.
At immediate left side of the featured white oak in second slide was a healthy local stand of Canada or hairy, wood brome.
Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004).
148. About all the principal acators on stage- Composition of mesic upland forest dominated by white oak and northern red oak. Largest tree in center was white oak. To its immediate right (partially concealed by trunk) was the multi-stemed shrub of red maple. A number of individuals of northern red oak were prominent in right fore- and midground.
A small sapling of American beech with large shade leaves represented this species at left foreground. Time alone would tell if beech succeeded to a climax dominant on this upland forest site, or even if scattered smaller plants survived to adulthood.
Shrubs present in this photo-plot included flowering dogwood, sassafras, and red maple at shrub size and with multi-trunks (just noted). Low-growing shrub in fore- to midground was lowbush huckleberry which was featured at a closer-in distance in foreground of the next slide.
Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004).
149. All major layers present and accounted for- View of mesic upland white oak-northern red oak forest with a wide selection of tree and shrub species representing all major strata of this forest range community. Lowbush huckleberry, the dominant low shrub species in this tract was photographed in foreground to emphasize abundance and importance of this species. Flowering dogwood, red maple, and sassafras as shrubs were plentiful. Redbud was also present but to extent of the just-listed species. The shrub at far left was cucumber tree (Magnolia acuminata).
Pole-size tree at far right was northern red oak.
All-in-all, an interesting aggregation of range plants.
Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004).
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