Oak-Hickory Forest

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Forest Range Types of Eastern North America

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

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

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

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

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

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

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

                1. Mixed Mesophytic Forest Region,

                2. Western Mesophytic Forest Region,

                3. Oak-Hickory Forest Region,

                4. Oak-Chestnut Forest Region,

                5. Oak Pine Forest Region,

                6. Southeastern Evergreen Forest Region,

                7. Beech-Maple Forest Region,

                8. Maple-Basswood Forest Region, and

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

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

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

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

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

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

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

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

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

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

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

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

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

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).

15. 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. There was also mentionable cover of black cherry, which probably indicated infrequent surface fires in this stand. Grasses were few in understorey, but dominant was broomsedge bluestem. There had been a history of cattle grazing in this stand, but it was generally light (mostly breechy ones looking for better pasture and finding worse than they left). Canopy was too dense for much herbaceous understorey other than for that of early season species like Mayapple (Podophyllum peltatum).

Many of the larger trees in this stand were over-ripe and dying or even dead. There were several snags. However most regeneration was hickory.

Ottawa County, Oklahoma. FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak). Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).


16. 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).


17. 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).

18. The black oak cover type- 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 (Symphoricarposorbiculatus) and blackberry. Wild grape and Virginia creeper (Parthenocissus quinquefolia) form an aboreal shrub layer while poison ivy (Rhus radicans = Toxicodendron radicans) occurs in both shrub layers ranging from lianas extending to tops of trees to non-climbing thickets.The herb layer is usually limited to early spring species that complete their annual cycle before greening of the forest canopy. Mayapple (Podophyllum peltatum) is the conspicuous dominant herb. Head of hollow on a chert upland. April, early vernal aspect. Ottawa County, Oklahoma. FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak). Oak-Hickory Series 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" !


19. 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).


20. 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.


21. 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.


22. "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.


23. Some respectible-size hardwoods on a rock pile- A climax oak-hickory forest with black oak (two foremost trees on the left; two left foreground trees) the most common tree species along with black hickory and bitternut hickory (Carya cordiformis) and even one blackjack oak (rightmost larger tree). The large tree with the high scar (tree behind foremost tree on left) was a bitternut hickory with a DBH of 24 inches. Not much herbaceous understorey but big bluestem and broomsedge were main species. Flowering dogwood (left margin; just coming into bloom) was major species of the upper shrub layer. A second or lower shrub layer consisted of buckbrush, blackberry, and Virginia creeper, this latter of which covered much of the ground surface and also reached up into tree crowns so as to be in both shrub layers. Grape (right foreground) also extended in both shrub layers.

Very marginal land (Land Capability Unit #8).

April, early vernal aspect. Ottawa County, Oklahoma. FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak). Oak-Hickory Series 122.11, Northeastern Deciduous Forest 122.1 of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

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

24. 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).


25. 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).


26. 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).


27. 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).


28. White oak-black oak-noethern red oak forest- White oak clearly dominated this north slope of a western Ozark Plateau forest, but it was not cover or density domiance to the degree that white oak dominated the north slope shown in the preceding set of slides (Roaring River State Park, White River Hills Ecoregion). Development and species diversity of two prominent woody layers below canopy layer was greater and different from that of a more overwhelming dominance by white oak as described for the immediately preceding Ozark Plateau forest.. This north slope and forest hollow was substantially more mesic and had less light exposure than the preceding forest.

Flowering dogwood was the dominant of the taller shrub layer. A lower and more diverse shrub layer included wild hydrangia, blackberry, poison oak, grape, Virginia creeper, and lowbush huckleberry. The herbaceous layer(s) was not as diverse as the white oak-dominated north slope forest described above, but the grass component (mostly downy, silky, or hairy wildrye) was much more productive and of far greater foliar cover on this mixed oak Ozark forest. Most common forb was pokeweed (which is more commonly a forest forb on locally disturbed areas).

Flag Springs State Park, McDonald County, Missouri. June, late vernal aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Most plausible Society of American Foreters (Eyre, 1980) cover type was SAF 52 (White Oak-Black Oak-Northern Red Oak) and this was predominant cover type of this locality: white oak was not as much as obvious dominant and defining species as was case for SAF 53 (White Oak) which this closely resembled. Oak-Hickory Series of Brown et al. (1998). Dry-Mesic Chert Forest (Nelson 2005, ps. 125-130). Ozark Highlands- Elk River Hills Ecoregion, 39b (Chapman et al., 2002).


29. 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).


30. 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).


31. 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.

Water Oak (Quercus nigra) Forest

The first example of the water oak (Quercus nigra) forest cover type shown immediately below was in a commercial forest in the Big Thicket section of the Texas Pineywoods. This water oak forest vegetation was adjacent to loblolly pine stands and a forest dominated by loblolly pine, water oak, American holly both with a lower woody layer comprised primarily of yaupon or, often called, yaupon holly (Ilex vomitoria).

Water oak has been regarded as Intolerant as to tolerance and as a subclimax species that is quite susceptible to fire damage (Fowells, 1965, p. 630; Burns and Honkala, 1990, Vol. 2, p. 703). Thus while light surface fires tend to maintain pines like the associated loblolly pine, major fire damage as with crown fires would select for regeneration of water oak. In absence of fire plant succession would progress to a climax of hardwoods, which in the Big Thicket would commonly be American beech, southern magnolia, American holly, and climax oaks such as white oak.


32. Water oaks in the Pineywoods- Exterior view of a local stand of water oak growing on a flatland forest site that frequently ponded water. Loblolly pine were growing around perimeter of the water oak stand. Yaupon grew as widely scattered individuals while most of the ground layer was oak leaves with scattered plants of longleaf woodoats (Uniola sessifolia), cottongrass bulrush (Scirpus cyperinus), and green flat sedge (Cyperus virens). These species (from this locale) were featured below under the loblolly pine-water oak-American holly form or subtype of loblolly pine-hardwoods forest. The largest--and also the most scarce-- herbaceouss pecies was bentawn plumegrass (Erianthus contortus) which was also featured below.

Liberty County, Texas. February, late hibernal aspect. This is a component or subtype of the general hardwood-pine southern forest forest that has one of the southeastern yellow pines a dominant or, sometimes, an associate species with oaks, hickories, or even beech as the more common climatic dominant (in contrast to a fire-determined dominant). Overall this forest range vegetation would have to be included in FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem) with K-101 (Oak-Hickory-Pine Forest) being the Kuchler equivalent listed thereunder. SAF 88 (Willow Oak-Water Oak-Diamondleaf [Laurel] Oak). Oak-Pine Series of Brown et al (1998) (but should be for Southeastern Deciduous and Evergreen Forest biotic community). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

Inside the water oaks- Interior of the local stand of water oak presented immediately above. This was a local consociation of Quercus nigra with a "broken" (widely scattered) population of yaupon holly and local herbaceous cover composed variously of longleaf woodoats, cottongrass or woolgrass bulrush, green flat sedge, and panicgrasses (Panicum spp.). This isolated water oak stand was adjacent to a mixed forest of loblolly pine, wter oak, and American holly (covered below).

Liberty County, Texas. February, late hibernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem) with K-101 (Oak-Hickory-Pine Forest) being the Kuchler equivalent listed thereunder. SAF 88 (Willow Oak-Water Oak-Diamondleaf [Laurel] Oak). Oak-Pine Series of Brown et al (1998) (but should be for Southeastern Deciduous and Evergreen Forest biotic community). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004)..


33. One of the more common forms or manifestations of oak forest in the Pineywoods of Texas and Louisiana is the Palmetto-Oak Flats (Ajilvsgi, 1979, ps. 12-13) or, when expressed as to topographic-edaphic rather than botanical features, Clayey Wet Upland Depressions (Diggs et al., 2006, ps. 97-98). Ajilvsgi (1979, p. 12) described overcup oak and laurel oak as dominants whereas Diggs et al., (2006, p. 98) emphasized willow oak and green ash (Fraxinus pennsylvanica) as major plants of the larger tree species. The Society of American Foresters (Eyre, 1980, p. 63) described the willow oak-water oak-diamondleaf or laurel oak type (SRM 88) as developing on a topographic-soil moisture gradient intermediate between the swamp chestnut oak-cherrybark oak type (SRM 91) and the overcup oak-water hickory type (SRM 96) with dominance of SRM 88 tending to change to non-oak hard spceies like green ash under heavy logging or high-grading.

The photographs shown below were of a water oak-willow oak forest with a lower shrub layer made up almost exclusively of dwarf palmetto and a herbaceous layer(s) of sedges, rushes, bulrushes, and panicoid grasses. Views of the Oak-Palmetto Flats in these slides were presented so as to view this forest range vegetation going from exterior to deep interior as if the viewer were traveling to and then into it.


34. Coming onto the Oak-Palmetto Flats- Exterior view of an example of the willow oak-water oak-diamondleaf (laurel) oak type showing physiogonomy and overall species composition of this form of Pineywoods. Dominant species of this stand was water oak with willow (locally known as "pin" oak). Laurel oak was a distant third Quercus species. Blackgum or black tupelo (Nyssa sylvatica) was another associate tree species. The largest tree with the horizontal upper limb and fire-scarred basal trunk was an ancient water oak readily idetified by the sporadically scattered, prominent "warts" of bark. Loblolly pine was represented by one conspicuous tree in center midground. There were other infrequent loblolly pine throughout. Young trees grouped at right foreground were a mixture of water and willow oak and very black tupelo. Dwarf palmetto made up a lower shrub layer. Grassses and grasslike plants comprised one or two (rarely three) herbaceous layers in the forest understorey. Herbaceous plants were most common around perimeter of the forest vegetation. Individuals of broomsedge bluestem (Andropogon virginicus) were prominent in foreground understorey.

Hardin County, Texas. February, hibernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem) with K-101 (Oak-Hickory-Pine Forest) being the Kuchler equivalent listed thereunder. SAF 88 (Willow Oak-Water Oak-Diamondleaf [Laurel] Oak). Oak-Pine Series of Brown et al (1998) (but should be for Southeastern Deciduous and Evergreen Forest biotic community). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).


35. Edge of an Oak-Palmetto Flats forest range- Around perimeter of a stand of water and willow oak with dwarf palmetto were various local assemblages of herbaceous plants. The latter included cottongrass bulrush and miscellaneous sedges, both Carex and Cyperus species (eg. green flat sedge [C. virens]), along with panicgrasses, especially beaked panicgrass (Panicum anceps); paspalums like brownseed paspalum (Paspalum plicatulum), and both broomsedge and bushy bluestem. These latter two species are invaders. These same species also formed herbaceous strata beneath the oaks and pines though with less continuous cover and smaller plants, conditions likely resultant from fairly dense shade. Water and willow oaks are Intolerant species.

Hardin County, Texas. February, hibernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem) with K-101 (Oak-Hickory-Pine Forest) being the Kuchler equivalent listed thereunder. SAF 88 (Willow Oak-Water Oak-Diamondleaf [Laurel] Oak). Oak-Pine Series of Brown et al (1998) (but should be for Southeastern Deciduous and Evergreen Forest biotic community). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).


36. Into the Pineywoods flats we go- These three photographs were a pictorial "walk to the woods", a sequence of slides showing the range vegetation of a water oak-willow oak- loblolly pine-palmetto-herbaceous plants Pineywoods flats. Continually closer-in views allowed presentation of the herbaceous layer(s) of native vegetation that was better developed at outer edge of the forest stand. Some of the common herbaceous species of this vegetation were presented below in the section devoted to the loblolly pine cover type, specifically the loblolly pine-water oak-American holly form or subtype thereof. The smaller trees in foreground with unshed lower limbs (most of them still alive but senescing) were willow oak. Locals hereabouts apply the otherwise confusing and nonstandardized common name of "pin oak" to Quercus phellos. "Pin" in several oak species refers to any of the lower, usually dead, unshed limbs (ie. dying or dead limbs on species that do self-pruning, but instead become well-seasoned or preserved and, hence, persistent on the lower bole). There were a few scattered woody vines, the only one of which the author-photographer identified was rattan (= Alabama supplejack).

Once inside the Pineywoods flats the interior of the water oak-willow oak-dwarf palmetto-herbaceous range community revealed a "closer-in" view of plant species composition and the lower woody layer of palmetto and the local vertical zone of herbaceous species. Largest trunk was that of a young to mid-age water oak with bark characteristic of an immature tree. At this stage of maturity bark of water oak and willow oak is so similar as to be indistinguishable, thereby making reliance on leaves and buds necessary for definitive indetification. "Warty" bark on older water oak bark was just forming on this straight-trunked specimen, but some smaller water oaks had larger "warts".

Grass shoot (visible in both photographs) in front of this water oak was broomsedge bluestem, a common invader of Oak-Palmetto, which was common and conspicuous throughout this oak flats stand. Almost all herbaceous species were grasses or grass-like plants and, as this was dead of winter and this range had been grazed so that most species had to be identified by vegetative features, most herbs could not be identified by the author who was a "stranger to these parts". The tallest green herb was cottongrass bulrush (shown and described briefly below). There were no prominent forbs in this forest range vegetation. Dwarf palmetto comprised a single-species, lower, woody layer.

Hardin County, Texas. February, hibernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem) with K-101 (Oak-Hickory-Pine Forest) being the Kuchler equivalent listed thereunder. SAF 88 (Willow Oak-Water Oak-Diamondleaf [Laurel] Oak). Oak-Pine Series of Brown et al (1998) (but should be for Southeastern Deciduous and Evergreen Forest biotic community). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).


37. "Up-and-dicular" perspective of a Oak-Palmetto Flats- Structure and species composition of the water oak-willow oak-dominated Pineywoods flatwoods described under horizontal photographs above. Architecture of this stand was displayed to better advantage in these two photographs. Most hardwood trees were water oak and willow oak of sapling to small pole size. Those with persistent lower limbs were willow oak. There was an "occasional" black tupelo (also of sapling-pole size).

Cover and density of palmetto was shown to good advantage in the first of these two slides while the frequent openings within the palmetto that were populated by grasses and grasslike plants were evident in the second slide. Tree in left foreground with live lower limb was willow oak.

This stand was obviously a second-growth forest. A cohort of sapling to small pole size oaks had developed beneath larger, established (older) but very widely scattered, mature oaks of both species. Structure and, especially, botanical composition of this stand was typical of climax water oak-willow oak-laurel oak-palmetto vegetation. Both willow oak and water oak are classified as Intolerant and recruitment of these species had been possible under a mostly open sky (sparse canopy of oak and loblolly pine). Natural thinning of oaks had already commenced as evidenced by the dead toppled pole (visible in both photographs). This will undoubtedly continue resulting in more dead younger oaks and fewer, though larger, trees (fewer boles but more board foot/acre) and eventually greater oak crown cover (increased--though by no means closed--tree canopy).

The apparent dominant herbaceous species was cottongrass bulrush. Numerous individuals of broomsedge bluestem were conspicuous with their tannish yellow shoots dispersed among bulrush and other grasslike plants.

Hardin County, Texas. February, hibernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem) with K-101 (Oak-Hickory-Pine Forest) being the Kuchler equivalent listed thereunder. SAF 88 (Willow Oak-Water Oak-Diamondleaf [Laurel] Oak). Oak-Pine Series of Brown et al (1998) (but should be for Southeastern Deciduous and Evergreen Forest biotic community). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).


38. Closing, composite shot of Pineywoods Oak-Palmetto Flats- All-in-one shot of species composition and structure (architeture) the water oak-willow oak-loblolly pine-palmetto-cottongrass bulrush-broomsedge bluestem community featured above. All of these species except loblolly pine, which dominated (generally and/or locally) their respecive layers of vegetation, were visible (if not obvious). In addition, rattan (= Alabamas supplejack) was featured prominently growing up trunks of oaks in left midground. Almost all oak trunks of any age are hosts to various crustose lichen, at least on north and east exposure.

Hardin County, Texas. February, hibernal aspect. FRES No. 13 (Loblolly-Shortleaf Pine Ecosystem) with K-101 (Oak-Hickory-Pine Forest) being the Kuchler equivalent listed thereunder. SAF 88 (Willow Oak-Water Oak-Diamondleaf [Laurel] Oak). Oak-Pine Series of Brown et al (1998) (but should be for Southeastern Deciduous and Evergreen Forest biotic community). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

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

39. Dwarf palmetto, blue palmetto, swamp palmetto, dwarf palm, blue palm, etc. (Sabal minor)- Large, mature swamp palmetto with previous season's floral stalk and spent inflorescence. This true palm is most commonly acaulescent (lacking a trunk or bole) though sometimes there are individuals that have a single, short woody stalk which would "pass for" a trunk. The shoot or stem does not branch and is characterized as woody or pithy in nature.

The speciment portrayed here was growing in the water oak-willow oak-lobollly pine-palmetto-cottongrass bulrush-broomsedge stand featured above. Hardin County, Texas, February.

Savanna of Oak-Hickory Forest and Tallgrass Prairie
The tallgrass prairie-hardwood forest savanna or transition was given a chapter of its own in this publication entitled, Tallgrass Savvanna, plus a section on Prairie Peninsulac in the chapter, Tallgrass Prairie (Interior). One photograph and caption was included at this point to highlight locations for these important deciduous forest and eastern prairie grasslands.
40. Oak-hickory forest and savanna— This is the widespread transition zone or intermediate community type between actual oak-hickory tallgrass prairie understory savanna and the western edge of the oak-hickory forest proper.Post oak and black hickory dominate the tree layer while black and red oak are associated species. Understory is primarily nodding or Canada wildrye and purpletop (Tridens flavus). Virginia creeper covers much of the understory and climbs into tree tops. Blackberry, buckbrush, Mayapple, and black-eyed susan dominate various microsites depending on cover of tree canopy. Chert savanna-chert forest composite type with fire determining which form prevails. June, late vernal aspect. FRES No. 15 (Oak-Hickory Ecosystem). K-73 (Mosaic of Bluestem Prairie [K-66] and Oak-Hickory Forest [K-91]). SAF 52 (White Oak-Black Oak-Northern Red Oak) but with black hickory more than black oak; SRM 801 (Dry Savanna). Oak-H:ickory Series of Brown et al. (1998).
Dominant, Associate, Widespread, and Just Plain Interesting Plants
of the Central and Southern Forests
41. Composite view of Oak-Hickory Forest- Tree on left is pignut or bitternut hickory, the forked trunks with burl on left trunk is black cherry, a young pignut hickory to its immediate right, and the two larger white-barked trunks in background visible between the hickory and cherry are post oaks. The barely detectable short shrub dominating the understory is buckbrush which in this winter season has shed its fruit. Hibernal aspect . Ottawa County, Oklahoma. Hibernal aspect, Late December. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 40 (Post Oak-Blackjack Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

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


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

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

Newton County, Missouri. October.



45. Shadbush or eastern or downy serviceberry (Amelanchier arborea)- Another shrub of the oak-hickory-- and general eastern deciduous-- forest is downy serviceberry. The common name of shadbush comers from the hill folk who noted that blooming of this species often coincided with the spring runs of shad (Alosa spp.). Downy serviceberry is often confused with flowering dogwood because the flowering periods of these two species often overlap and that of one species may precede or lag behind the other depending on conditions in any one spring. (Dogwood gets all the credit by the way.)

Serviceberry is one of the many members of the rose family which is the single most important family of range browse plants in North America. Amelanchier species are in the Pomoideae, the pome fruits subfamily, of Rosaceae. Shadbush undoubtedly provides some browse and the fruits are eaten by birds and furbearers, but it is not common enough to be a major feed plant.

The bluffs of Modoc Creek, Ottawa County, Oklahoma. April.

46. Downy serviceberry in full bloom- In the opinion of your author when this species is flowering it is often the most striking and showy shrub in the oak-hickory forest. Much of the enjoyment of the spring blooming in the hills is due to serviceberry and not flowering dogwood, but a human population of predominantly city dudes does not know the difference and just calls everything other than redbud a "dogwood". Ottawa County, Oklahoma. April.
47. Flowering shoot tip of downy serviceberry- The inflorescence and two newly emergent leaves of shadbush or eastern serviceberry. Ottawa County, Oklahoma. April.

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

Springfield Plateau section of Ozark Plateau; base of limestone hill in Newton County, Missouri. June.

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

50. Catkins (male flowers) and mature fall fruit of common hazelnut (Corylus americana)- Another (but usually sparsely populated) shrub of the oak-hickory forest is the hazelnut. This nut was highly prized by the early American backwoodsmen, the proper historic name for the first wave of frontiersmen who moved westward beyond the Fall Line into the dark, forbidding, and mysterious deciduous forest of eastern North America. For millinia prior to the first footprint of man (God only knows what shade of skin or species of Homo it was) hazelnut was feed for wildlife ranging from wild turkey to furbearer to native ruminant.

Male flowers of this monecious species are borne in catkins that are some of the first flowers in the late winter or early spring in the oak-hickory forest. The specimen shown here is from an individual of the variety C. americana var. indehiscens (formerly interpreted as C. cornuta) delineated as a separate taxon by the united fruit bracts that make the fruit to appear unopened. Fruits appear singularly up to as many as four in a cluster like the one pictured here.

The family to which this little nutbearer belongs is another source of controversy. Some treat it as a member of the birch family (Betulaceae) with separate tribes of Betuleae and Coryleae while other workers place it in it's own hazel family (Corylaceae).

On second terrace of Modoc Creek in Ozark Plateau. Ottawa County, Oklahoma. Catkins, February; fruit, October.

51. Pawpaw (Asimina triloba)- Pawpaw or custard apple is another (though less common) shrub of the eastern deciduous forest, in particluar the oak-hickory forest. As is the case for many of the understorey plants, especially forbs, pawpaw reproduces asexually by suckering from extensive creeping rootstocks. In fact, this is usually the main mode of reproduction there being springs when pawpaw does not bloom (and fruit-set is much less frequent than flowering). On bottomland forests pawpaw often forms estensive colonies with broad, shiny leaves. Ottawa County, Oklahoma. August.
52. Immature fruits of pawpaw- When pawpaw does bloom and set fruit, which is less frequent than non-fruiting years, it produces this backwoods delicacy for 'possums (Didelphis virginiana), coons (Pyocyon lotor), and coon- and 'possum-hunting hillbillies. The green color of the thin fruit skin indicated that this fruit was still immature. Ripe fruit turns yellow and finally brown when over-ripe. Ottawa County, Oklahoma. August.

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

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


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

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


55. Missouri gooseberry (Ribes missouriense)- This shrub does not approach the abundance of Rubus, spp., Linderia benzoin, or even Vaccinium spp., but it is a climax understorey shrub in the oak-hickory forest types. In these forests gooseberry sometimes grows in colonies infrequently starting new plants from tips of shoots that come in contact with soil. More commonly Missouri gooseberry occurs as a solitary or, sometimes, as a few plants that form large clumps. This specimen was unusually fruitful.

Ottawa County, Oklahoma. Estival aspect, July.

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

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

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

59. Highbush huckleberry, squaw huckleberry, or deer berry (Vaccinium stamineum). Ottawa County, Oklahoma. June.
60. Poverty oatgrass (Danthonia spicata)- This cool-season bunchgrass often forms rather extensive carpet-like stands made up of these green clumps in the understorey of oak-hickory forests. This occurs mostly on the drier habitats like the steeper of south slopes and the more shallow, stoney, and acidic soils where the larger grasses (eg. the bluestems, panicgrasses, beakgrain, and woodreed grass), forbs, and shrubs like blackberry (Rubus spp.) are absent or stunted. Poverty oatgrass is also more common on "edges" between deciduous forest and tallgrass prairie where the more drought- tolerant trees predominate. For example, in the habitat shown here oak leaves in the background belonged to blackjack (Quercus marilandica). Ottawa County, Oklahoma. June.
61. Panicle of stout woodreed or woodreed grass- This festucoid grass (tribe Aveneae) is a major provider of nutritious forage in the understories of carefully managed (ie.conservatively stocked) oak-hickory forests. It is a major species only on a local basis because years of overgrazing have greatly reduced it’s abundance and even occurrence. Woodreed is adapted to diverse sites within the oak-hickory forest range type thriving on both bottomland and stream bank habitats as well as shallow upland soils. Bank of Modoc Creek, Ottawa County, Oklahoma, September.
62. Stout woodreed is a perennial grass that has an obvious prolonged growing season and pronounced allocation of resources. In the Ozark Plateau flowering occurs in late summer or early fall (August to October), but new basal shoots appear soon after completion of the annual life cycle (death) of current-season shoots. Thus there are usually green shoots in some phenological stage yearlong, but the growth and development of these shoots is extremely slow. In this photograph young shoots (tillers: intravaginated shoots) of two to four inches in length are growing among last year’s tillers which reached their maximum mature height of three to four feet. Shallow savanna range site (a shallow upland soil overlying a solid layer of chert). Ottawa County, Oklahoma, December.

63. Close-up of the woodreed tillers of the previous slide- Allocation of resources so as to maintain some live tillers yearlong is likely an adaptation to the dark shade existing from spring through autum due to  dense canopy of the forest during this frost-free period.(mid-April to mid-October in the Ozarks). Woodreed requires a full year under these conditions of limited light to fix enough carbon to complete the annual life cycle and store root reserves to initiate next year’s growth. Any plant living in habitats with such a low light intensity and a protracted regime of photosynthesis and growth will obviously be quite vulnerable to defoliation and require careful grazing management.    

Not only is conservative stocking important for regeneration of shrubs and lumber trees but it is also essential to prevent overgrazing of grass species growing in the dimlite, stressful understory. Given the shallowness of soils on some sites in deciduous forest range in conjunction with the commonness of periodic summer drought it is likely that grasses like stout woodreed and beakgrain suffer from the stress of water shortage. This would be “true in spades” for these summer-flowering festucoid grasses which must compete with the more heat- and drought- tolerant dominant panicoid grasses like big bluestem and beaked panicgrass (Panicum anceps). Combinations of dense shade, shallow soils, drought, competition, etc. make for a harsh environment even in humid precipitation zones. Excerbation of this condition with overgrazing or improper season of use is one reason why deciduous forest ranges have become deteriorated.

Furthermore (and contrary to popular misconception) these deciduous woods ranges were the last open range (the true meaning of open to the public, a grazing commons, and not just absence of fences) in the United States. The actual Public Domain ranges of the Intermountain West were closed and came under some scientific regulation by the Grazing Service (later, Bureau of Land Management) with attempted enforcement of the Taylor Grazing Act in the 1930s. By contrast, state laws officially closing the range (passage of mandatory “fence-in” laws) in Missouri, Arkansas, Oklahoma, and Texas were not passed until the 1950s. Free-ranging, acorn-eating, razorback, rooter hogs could be found sleeping and rooting on county courthouse squares through the first half of the 1950s throughout portions of the Ozark, Boston, Quachita Mountains in Oklahoma, Missouri, and Arkansas and in the Big Thicket of Texas. The account for the latter can be found in Sitton (1995, ps. 194-273 passim). Rowley (1985, ps.239-241) noted that control of livestock grazing on National Forest “… in the South [Region 8] remained an unfinished business in the early 1960s…”.and that in “… the 1950s range continued to be the southern region’s ‘ bastard child’. Closing the southern ranges had always been a battle in intself (King, 1982) because (and this is another scientific, economic fact that is contrary to popular, and incorrect, opinion) livestock raising was far more important economically and culturally than it was credited with. This was true even for the antebellum South when the planter class was at its zenith (McDonald and McWhiney, 1975). It is no wonder why professionally trained foresters still harbor strong prejudices against even proper (sustainable) livestock grazinig in the eastern deciduous forests.

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

66. Nimblewill (Muhlenbergia schreberi)- This eragrostoid grass (Eragrosteae tribe) is another widely distributed grass in understories of oak-hickory forest ranges. Nimblewill grows on a diversity of habitats ranging from fairly dense, mesic woods to open prairies on shallow soils and as a component of stremside vegetation to a common species on abandoned fields (= go-back land) or even old city lots and schoolyards. It is generally accepted that nimblewill is probably the best-adapted of any muhly in the southern oak-hickory forests and savannahs, including adaptation to disturbances like overgrazing and trampling. Nonetheless, nimblewill was interpreted by Tyrl et al., 2002, p. 109) as "... characteristic of the mid to late stages of plant succession".

Although nimblewill is neither rhizomatous nor stoloniferous it frequently grows as localized colonies especially in forest clearings, lanes, old barnyards, etc. This habit is due to adventituous rooting at nodes of the low, decumbent tillers from which short-branched, largely unspread panicles arise. Nimblewill has a unique feature (unique among North American Muhlenbergia species) in that morphology or growth form differs greatly from the early season, short plants with broad leaf blades to the late-season form with ascending to even sweeping shoots.

Nimblewill specimens presented here were growing in shade of post and blackjack oaks at extreme western edge of Ozark Plateau where relatively large-sized tallgrass prairies and oak-hickory forests developed in a complex vegetational mosaic. This patchwork of range vegetation was part of the southern extension of the prairie peninsula.

Ottawa County, Oklahoma. December: hibernal aspect, dormancy.


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

Ottawa County, Oklahoma. December: hibernal aspect, dormancy.


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

In bottomland forest of Modoc Creek, Ottawa County, Oklahoma. April.


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

First terrace of Modoc Creek, Ottawa County, Oklahoma. May.

70. Detail view of green dragon- Leaves and the spadix and spath of the "flower" of green dragon were presented here. Modoc Creek, Ottawa County, Oklahoma. May.
71. Deer's tongue, adder's tongue, or dog-tooth violet (Erythronium albidum)- This is a forest lily that emerges early in the spring through the deep layer of shed oak and hickory leaves on the forest floor. It then promptly flowers before the overstorey trees grow their annual canopy that affectively blocks most light from reaching the ground. Hill-tromping hillbillies enjoy this delightful forb as a harbinger of spring. Ottawa County, Oklahoma. March.
72. Wake robin (Trillium sessile)- This is another member of the Liliaceae that emerges early in the spring when it can get enough light to complete it's annual cycle of life. Wake robbin grows best and to it's largest size on rich, moist soil. Ottawa County, Oklahoma. April.
73. May apple or mandrake (Podophyllum peltatum)- May apple is probably the most widespread and, as it grows in sizeable colonies from extensive rootstocks and sports large, spreading, uniquely shaped leaves, the most conspicuous forb of oak-hickory forests. The fruit is edible, but the foliage does not appear to be grazed by vertebrates. This mandrake should not be confused with the other species of that name that is native to the Mediterranean Region (the mandrake of the Holy Bible). This forb is often the locally dominant species of the upper herbaceous layer of the multi-layered deciduous forest. Ottawa County, Oklahoma. May.

74. Dutchman's Breeches (Dicentra cucullaria)- The range of this forest forb extends from Quebec westward to Oregon and Washington and south to Missouri and Oklahoma. As seen from this photograph it "does it's thing" early in the spring before the trees leaf out and exclude light from the forest floor. Even then, as also clear from this slide, light is limited by trunks and larger limbs.

This species was discussed in Notes on Western Range Forbs (Dayton, 1960, p. 220-222) where it was noted that the bulbs are poisonous to cattle when they are pulled from the soil and eaten along with the shoot. Burrows and Tyrl (2001, ps.706-709) provided a recent synthesis. They noted that Dicentra species contain several isoquinoline alkaloids that affect the nervous system and cause trembling and stggering, but they added that the plants were so uncommon as pose no serious problem, and besides animals usually completely recovered.

Your author noted that this is another application of the Cardinal Principles of Range Management, Proper Season of Use in this case. When forbs like the ones shown in this section are growing there is very little valuable forage in the oak-hickory forest because what grasses and valuable forbs (eg. legumes) do grow have not produced feed. Ergo: stay the heck off the oak-hickory forest ranges at this time. Perhaps of even more importance is the fact that twigs and buds are very susceptible to browsing injury at this season and growth such that regeneration of hardwood species is easily adversely affected. The wood products from these oak-hickory forests are a greater source of revenue than is the 10-30 acres per AUM valuation. It does not require a post doc to figure this out folks.

Base of limestone bluff along Modoc Creek in Springfield Plateau section of Ozark Plateau; Ottawa County, Oklahoma. March.

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

76. Bloodroot (Sanguinaria canadensis)- Here is yet another picturesque range forb from the early spring floor of the oak-hickory forest. And it comes with another colorful common name. This one derived from the red- or orange-colored exudate from the rhizomes that was used by backwoodsmen as a dye to add a little color to their drab garb of buckskins and hunting shirts and gingham or flour sack dresses.

Bloodroot is a flowering neighbor of it's pantalooned "cousin" just presented. Both bloodroot and Dutchman's breeches bloom at about the same time and in close proximity to one another. Bloodroot seems to grow slightly higher on hillsides, especially those adjoining streams, than does Dutchman's breeches. Yes, these two species are "kissin' cousins" of a sort. Some taxonomists placed Dicentra species in the Fumariaceae (fumatory family) while others interpreted Dicentra as belonging to the subfamily Fumarioideae in the Papaveraceae (poppy family). Bloodroot has commonly been interpreted as a member of the Papaveroideae subfamily of Papaveraceae.

On a limestone bluff alongside Lost Creek in the Springfield Plateau section of the Ozark Plateau physiographic provinece. Ottawa County, Oklahoma. March.


77. One shoot (from a rhizome) of bloodroot at peak bloom and accompanied by a detailed view of it's inflorescence- Take note of the fruit immediately above the leaf. Fruit of bloodroot is a one-locular (locule= cavity of, in this case, the ovary) capsule. This fruit is very similar, obviously, to the capsule of poppy.

Bloodroot is one of the first native forbs to flower in the spring on the leaf-covered floor of oak-hickory forests. To be able to live through another hard winter and share the joy of the bloodroot in bloom is one of Nature's blessings to the hill-billy.

Bluff above Lost Creek, Ottawa County, Oklahoma. March.

78. Solomon's seal (Polygonatum commutatum) - This and the next species are members of the asparagus tribe of the lilly family (Liliaceae) are found in the vast deciduous forest region of eastern North America. The pictured specimens of these two species were growing in the oak-hickory forest association in the Ozark Plateau. They were conspicuous plants and though of no commercial value often elicit favorable responses from people visiting the "woods", in particular native plant and wild flower buffs. Young spring shoots of Polygonatum and Smilacina species were eaten by Indians and backwoodsmen. The range of P. communatum is from New England westward to Oklahoma. Ottawa County, Oklahoma. May.
79. Flowers and leaves of Solomon's seal- The bell-shaped flowers and rounded to broadly acute leaves were obvious between views in these two photographs. Polygonatum species are strongly rhizomatous. The origin of the common name was from the seal-resembling scars on the rhizomes. Ottawa County, Oklahoma. May.
80. False Solomon's seal or false spikenard (Smilacina racemosa)- This forest forb was growing on a moist north slope on top of a bluff in an oak-hickory forest in which sugar maple (Acer saccharum) was the dominant. The understorey woody (= shrub) layer was dominated by flowering dogwood a trunk of which was pictured along the right margin of the photograph. Forbs such as this have little or no feed value although Dayton (1960, p. 23) reported that deer eat the berries of Smilacina species. Their main practical vlaue in Forestry and Range Management is as biotic diversity. Professionals in these natural resource fields are frequently called upon to provide names for conspicuous plant species and questioning laymen are always impressed when rangemen and foresters can spout back the name. This is more so the case for those plants that have little economic value because it shows that professional resource managers know even the minor species, those that are not major lumber or forage and browse plants. Above Modoc Creek, Ottawa County, Oklahoma. May.
81. Inflorescence of false Solomon's seal or Solomon's plumes- This bright flower cluster was on a specimen growing above Modoc Creek in Ottawa County, Oklahoma. May.

Pterophyte is the term for spore-bearing vascular plants. These are "intermediate" (in evolutionary development, taxonomy, etc.) between the traditional units of Byrophyta and the Spermatophyta (seed-bearing plants, usually a taxon at the division level). Traditionally the pterophyte taxonomic level (division) has been called Pteridophyta. All three of these traditional units or taxa were included in the Embryophyta (subkingdom of plants in which the zygote develops into a multicellular embryo while enclosed within the female sex organ or within the embryo sac). In the traditional taxonomic hierarchy the vascular plants (those possessing xylem and phloem including both Pteridophyta and Spermatophyta) were distinguished from the Bryophyta by being placed in the Tracheophyta (the taxon, variously a division or superdivision, of vascular plants). In other words, Tracheophyta minus Pteridophyta leaves Spermatophyta (Gymnospermae and Angiospermae). The Pteridophyta included the club mosses (Lycopocineae), horse-tails or scouring rushes (Equisetineae), and the ferns (Filicineae). Some taxonomic schemes had a taxon designated Pteropsida that grouped (largely on basis of complex, relatively large leaves) the ferns, gymnosperms, and angiosperms.

Spore-bearing plants from the thallophytes through and including the pteriophytes have been called cryptogams (pteridophyts are vascular crypotgams). Plants have thus traditionally been grouped taxonomically in various arrangements depending on what features were of concern for different purposes. In effect, this is a form of taxonomic bilingualism. It is desirable, even essential, in instances where several features are of interest simultaneously. This is often the case when studying vegetation and, even more so, when applying this study to production agriculture. Those individuals are uninformed who view terms and taxa like the ones just discussed as meaningless or useless because they are archaic or, even, obsolete (two different things). If these individuals arrogantly persist in viewing such words as meaningless these folks are stupid (incapable of learning).

There are several taxa of pteridophytes in range and forest vegetation. Pteridophytes are especially common in the various deciduous forest cover types. A few were selected for inclusion in this section devoted to the oak-hickory forest.


82. Colony of smooth scouring rush (Equisetum laevigatum)- Equisetum is, as obvious from the prefix, the genus of pteridophytes given the common name of horsetail. Many of the Equisetum species do not have the namesake horsetail arrangement of primitive leaves encircyling the shoot. Those Equisteum species whose leaves are reduced to rudiments were given the common name of scouring rush, a common name which appeared to aptly describe even those having a "horsetail". In fact, given that most Equisetum species in many locales are "bald" tails, scouring rush is often the more apt common name (genus designation notwithstanding).

Lack of well-developed leaves that have a typical leaf form was a criterion used in classifying ferns as more advanced and closer to spermatophytes than are the club mosses and scouring rushes (ie. ferns are the least primitive of the vascular cryptogams).

Within the oak-hickory forest many of the vascular cryptogams grow best on the more moist habitats.This colony of smooth rush was growing along a high bank of Lost Creek flowing through the extreme western edge of the Ozark Plateau section of the oak-hickory forest. Ottawa County, Oklahoma. July.


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

Specimen from the stand shown immediately above. Bank of Lost Creek, Ottawa County, Oklahoma. July.


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

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

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

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

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

Newton County, Missouri. May.

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

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

Ottawa County, Oklahoma. January.


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

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


Bryophytes are those range and forest nonvascular plants of the general groups of mosses, hornworts, and liverworts. (Recall that "plants" was used herein as in the older or more traditional usage of the two kingdoms of organisms that the author viewed as more practical for discussion of vegetation.). More specifically bryophytes are members of the Bryophyta (a division of plants in the Plantae); in short, they are nonvascular spore-producing plants. Bryophytes have alternation of generations (gametophyte and sporophyte), but in contrast to the vascular plants the dominant and conspicuous generation of bryophytes (ie. the so-called plant) is the gametophyte (gametophytic generation). While bryophytes lack differentiation into true roots they have root-like structures called rhizoids which anchor the gametophytes to their substrate (bryophyte rhizoids do not function in absorption).

Bryophytes, like thallophytes (thallus-- the type of plant body that is undifferentiated into root and shoot-- plants such as algae, bacteria, and fungi), are primarily important in Range Management and Forestry as reducers or decomposers and therefore in processes like soil formation and plant succession (ie. the Clementsian process of reaction now more commonly known as facilitation). This was discussed above immediately before presentation of fungus species.

Raven et al. (1992, ps. 298-316) provided an excellent introductory discussion of the Bryophyta. Shaw and Goffinet (2000) wrote a comprehensive and the recent classic text on the Bryophyta for the "really serious" student.


90. Gametophye of a "true moss" (order of Bryales), Musci- "Moss" is one of the most confusing and most misleading names in all of Botany. "Moss" is applied to every sort of "plant" from reindeer moss (a lichen eaten by caribou and reindeer) to Spanish moss (a monocotyledonous epiphyte often eaten by native and domestic ruminants). Obviously such "mosses" are range plants by definition because they are native plants that serve as feed for grazing/browsing animals. Absent this axiomatic and self-evident definition, these and other "plants" growing on range are still range plants even if they function in less conspicuous roles such as decomposition, soil formation, nutrient cycling, and plant succession. True mossses provided such examples of these roles or functions in forest and range ecosystems.

"True mosses" has been the designation traditionally used for the one (of three) class in the division of Bryophyta know as Musci. In some of the more recent works (eg. Raven et al., 1992, p. 308) Bryophyta division still consist of three classes one of which is that of the true mosses but known as Bryidae (instead of Musci). The Musci or Bryidae bryophytes are partially saprophytic being dependent on decaying organic mattter. The specimen photographed here (the species of which was not identified ) was attached to a piece of chert that had a deep facet which had acumulated rotting oak leaves. Oak-hickory forest in Ozark Plateau. Ottawa County, Oklahoma. January.


91. Gametophytic and sporophytic generations of a species of true moss (Funaria hygrometrica), Musci- In these two macrolense shots the leafy gameotphyte is visible at the base of the moss while the mature sporophytes are very prominent as the apex of the moss body. These shown here were female sporophytes known as archegonia (singlular is archegonium) or archegonial heads which consist of a capsule (= sporangium; plural is sporangia) that are borne on a seta (stalk). The first of these two slides presented a view from the top to show the overall appearance of moss in the sporophytic stage.

The second slide was a side-view of the moss which showed more clearly both the leafy gametophytic generation and the stalked capsule of the sporophytic generation. The covering of the capsule is the calyptra the sharp, pointed tip of which is the operculum. This is the lid of the capsule which when shed opens up the capsule, the inside of which contains spores which are released upon this opening. While these are not vascular plants and are not differentiated into roots, stems, and leaves there is some differentiation of tissues. This occurs in the stalk as well as in the leafy gametophyte.

At the base of this sectional sample of moss are the rhizoids which are the root-like structures (more like root-hairs actually) of the gametophytic generation that function only to anchor or hold the plants (absorption of water and mineral nutrients occurs directly through the gametophyte).

Oak-hickory forest in Ozark Plateau. Ottawa County, Oklahoma. March.


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

Newton County, Missouri. April.

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

The relatively high primary productivity of the oak-hickory forest results in heavy accumulations of detritis ("yields" of necromass) that are rich substrata for many kinds of reducers (= decomposers) among forest and range plants. (The author acknowledged the value and legitimacy of the five kingdom taxonomic system, but herein used "plants" in the traditional botanical usage for simplicity, utility, and practical application.) Botanical organisms functioning as reducers extend from bacteria, algae, fungi, lichens, and bryophytes to vascular plants. A sample of these was included below beginning with the fungi. Not all fungi are saprophytes. Some fungi are saprohytic, of course, but others form mycorrhiza or the mycorrhizal symbiosis with roots of vascular plants while others are parasitic (on vascular plants, insects, or even othr fungi). Some fungus species are a combination of saprophyte and parasite (examples were presented below). Likewise, not all saprophytes are fungi or bryophytes. An example of a saprophytic vascular plant (a dicotyledon) was presented below following presentation of several fungal species.

Fungi are one of the three major groups of organisms traditionally regarded as plants (or plantlike taxa) that lack vascular tissue (ie. simplistically defined as nonvascular, undifferentiated lower plants). These "plants" (plantlike organisms) were historically regarded as thallus plants or thallophytes, members of the Thallophyta (usually listed as a subkingdom). Thallophytes are those organisms having plant bodies not differentiated into roots and shoots, lacking vascular tissue, and having gametes enclosed only by a cell wall such that their zygotes do not develop into embryos while inside the female sex organ (Wilson et al., 1971, ps. 447-451). With acceptance of the five kingdom classification system of organisms proposed by such stellar scientists as Robert H. Whittaker the older or traditional two kingdom scheme fell into disuse. While this development was generally an improvement (especially for the teaching of Biology to beginners) some of the taxonomic groupings (eg. thallophyte) still make sense especially for applied use in Agriculture. The author of this web publication periodically resorted to groups like the Thallophyta for practical use (ie. ease of teaching cencepts in conservation, agricultural production, etc.).

Thallophytes and, as discussed below, bryophytes are essential to development of vegetation and soils. These lower plants typically form one or more layers of vegetation. Also, they are usually pioneer species. These primitive plants are among the first organisms to grow on the raw parent material of a sere such that they improve the habitat for higher plants of later seral communities. This was what Clements termed reaction, one of the processes in development of vegetation (Clements, 1916a, 79-96 passim; Weaver and Clements, 1938, ps. 234-241). Years later this process was termed facilitation (Connell and Slatyer, 1977) and in time became known as the facilitation model of plant succession (Begon et al., 1990, ps. 632-633, 635, 641-642). Clements' specific view (almost as much philosophical as theoretical) was that "[e]ach climax formation had its individual or ontogenetic development"... such that "it shows a phylogenetic development from a preceding climax or community". "If the phylogeny of the community comprises the same general process as that of the species, it should be recapitulated by the ontogeny as seen in the sere" (Clements, 1916a, ps. 344-345).

An assumed phylogeny beginning with lower plants such as algae, liverworts and mosses, lichens, etc. (as determined from the fossil record) implied-- at least to Clements-- that development of vegetation on current seres began with these thallophytes and bryophytes. This recapitulation component of Clements' grand theory of plant succession was probably the least understood part of his complicated model of vegetation development, but undoubtedly all students of plant succession have observed that some of the earliest species to pioneer an area (especially a prisere, a fresh or newly created bare area of parent material or the area having to undergo primary succession) are the primitive plant forms. Whatever processes and paths of development are involved, the thallophytes and bryophytes do facilitate development of range and forest plant communities by the processes of plant succession. It was for this reason that a few of these species were included in this publication on range and forest cover types.

There are numerous ouytstanding field guides to the fungi (ie. mushrooms) of North America including Krieger (1967), Orr and Orr (1979), Lincoff (1981), Pacioni (1981), McKnight and McKnight (1987), Metzler and Metzler (1992), and the massive Arora (1986). Mycology is the study of fungi. There are numerous outstanding texts and references for the fungi. Pritchard and Bradt (1984) was recommended for readability yet thoroughness. Carlile and Watkinson (1994) was a comprehensive text that covered basic biology, including ecology, evolution, genetics, etc., but for overall, concise reference (especially for beginners) the basic Botany textbook of Raven et al. (1992, ps. 208-243) seemed easiest to use.


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

Banks of Modoc Creek, Ottawa County, Oklahoma. April.


95. Ink cap (Coprinus radians)- On this rotting log covered with decomposing oak leaves is the complete body of a fungus in the group, basidiomycetes (Basidiomycotina is a subdivision of the higher, fleshy, or true fungi), more specifically the Hymenomycetes class therein. The species shown here is one of the best or standard textbook examples of the true fungi, those species that produce fleshy fruit-bodies know generically as mushrooms and/or toadstools. The two general parts of a mushroom are 1) the filaments of hyphae which form a network known collectively as the mycelium or spawn and 2) the fruiting body, often called the carpophore, which bears the reproductive structures that produce spores. The fruit-body is the obvious, often conspicuous, part of the true or higher fungi (ie. the "mushroom") which forms from the mycelium. The latter part of the true fungi are not differentiated into roots and shoots and are typically not seen by human eyes. The only part of fungi usually visible (again, to humans) is the fruit-body, the mushroom. If some of the mycelium (hyphae filaments) remain attached to the carpophore when someone picks or knocks over the mushroom he would naturally think of the mycelium as the "roots" which is obviously incorrect.

C. radians is unusual in having bright orange hyphae threads and, when these are accompanied by the carpophore (as shown here), students have the rare opportunity to see the whole "plant" (entire fungus body). In this slide there is one live or "fresh" and one dead (withered) carpophore. Some fungal species have a carpophore or mushroom that includes a prominent "cap" as its hymenium, the spore-bearing surface of the Ascoymetes and Basidiomycetes. In C. radians the spore-bearing underside of the cap is dark-colored hence the common name of ink cap (next slide). Obviously this fungus is a saprophyte. It was shown to offer students a classic example of a saprophyte or saprophytic "plant" which is one of the major categories within the reducer or decomposer group of organisms in forest and range ecosystems.

First terrace of Modoc Creek, Ottawa County, Oklahoma. April.


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

Bank of Modoc Creek, Ottawa County, Oklahoma. April.

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

98. Wood ear or tree ear (Auriculularia auricula) on dead elm (Ulmus americana) limb- In the first slide, almost indistinguishable on this dead limb, are dry fruit-bodies (accompanied by equally dried-out lichens) of a species of the Hymenomycetes (class) of the Basidomycetes (subdivision) known by the colorful (and most appropriate) common name of wood ear.

In the second slide, which had been taken 24 hours earlier when the fruit-bodies (the "mushrooms") were conspicuous, wood ear carpophores (and accompanying lichens) had swollen from a recent rain.

This saprophytic species is but one of many fungi that quickly decompose the vast detritis or necromass (dead organic matter that was previously biomass) produced in the oak-hickory forest. This species of Hymenomycetes was growing in the "dead of winter", a common season for appearance of many carpophores due to abundance of water in detritis as a result of low evaporation rates.

Ottawa County, Oklahoma. December.

99. Wood or tree ear (Auriculularia auricula) capophores fully hydrated- Close-up shot of wood ear immediately after a warm winter rain. Ottawa County, Oklahoma. December.
100. Wood ear fruit-body drying out- Appearance of the member of Hymenomycetes known as wood ear about six or seven hours after cessation of winter rain. The capophore was quickly returning to the more common dehydrated state as shown above. Ottawa County, Oklahoma. December.

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

Newton County, Missouri. December.


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

Oak-hickory in Ozark Plateau. Newton County, Missouri. December.


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

Oak-hickory forest in Ozark Plateau. Ottawa County, Oklahoma. April.


104. Oak conk- The basidiocarp of a woody shelf fungus showing the underside with the layer of tissue from which the spores are released from the numerous spore-producing tubes. The typical cracked cap of the polypores (family: Polyporaceae) was also clear in this photograph. This was one of numerous carpophores on the trunk of the old black oak shown in the preceding slide.The shelf- or bracket-like basidiocarps of this and related species are just as woody as the substrata they grow on.

Polypores have generally been viewed as more saprophytic than parasitic, but most species function in both capaacities. Arora (1986, p. 549) pointed out that the mushrooms of this group (Polyporaceae and related genera) "are absolutely indispensable to the forests of this world" in their function as decomposers. These are the major "wood-rotting fungi" causing 90% of the rotten-wood damage to standing timber in North America. This fact could be viewed as indicating a major forest pathology crisis, and at one time foresters who interpreted Forestry as mere tree farming (and thus sought to eliminate all "enemies of the forest") may have held such views. That was before application of the ecosystem concept brought back a more balanced perspective or maybe just some old-fashioned woods wisdom in new terms for original principles. Today's foresters and rangemen appreciate the essential role of decomposers in reduction of detritis (plus the ecological role of detritis itself), cycling and availability of nutrients, and provision of food for consumers (both vertebrates and invertebrates). Arora (1986, p. 549) stated the case for the polypore fungi and put the role of decomposition in proper perspective. "Without them there would be no logging industy in the first place: every cut stump, felled log, and lopped-off limb would be indefinitely on the forest floor, the woods would quickly become impenetrable, and the new trees would have neither room nor nutrients to grow".

Oak-hickory forest in Ozark Plateau. Ottawa County, Oklahoma. April.


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

Ottawa County, Oklahoma. October.


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

Ottawa County, Oklahoma. June.

107. Pinesap- Close-up photograph of the vascular saprophyte presented in the preceding slide. This specimen had less pubescence than is typical for this species. It was growing on the organic matter of rotting oak and hickory leaves in the western part of the Ozark Plateau. Ottawa County, Oklahoma. June.
108. Interior of an upland mixed oak (white, red, post, black)-hickory forest that has extremely lush and diverse shrub and herb layers. Understory plants include tick clover, blackberry, grape vine, Virginia creeper, poison oak (Rhus toxicodendron= Toxicodendron toxicarium), bluestem, and panic grasses. Second growth forest with excellent regeneration of climax tree species, especially hickories. Note that this excellent botanical diversity and forest regeneration is occurring under light or conservative grazing/browsing by cattle (Herefords are visible in center). Adair County, Oklahoma. May, late vernal aspect. FRES No. 15 (Oak-Hickory Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak) and SAF 110 (Black Oak) combination, dry-mesic chert forest. Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Dissected Springfield Plateau-Elk River Hills Ecoregion, 39b (Woods et al., 2005).

109. Uneven-aged management in upland oak-hickory forest- Selective cutting (= selective felling) in an Ozark Plateau chert forest dominated by black oak which was harvested for veneer to be used in making church furniture. Trees were felled in winter and this is the scene three months after logging. Most harvested trees were 70-90 years old. Note total absence of any soil disturbance or damage to young trees. Logs were carried— not drug —out by a rubber-tired skidder. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 52 (White Oak-Black Oak-Northern Red Oak) tending toward SAF 110 (Black Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

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

“Forest Management- the practical application of biological, physical, quantitative, managerial, economic, social, and policy principles to the regeneration, management, utilization and conservation of forests to meet specified goals and objectives while maintaining the productivity of the forest” (Helms, 1998).

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

110. Stump sprouting (= coppicing) of young black oak- These are stump sprouts at the near end of the first growing season post-logging on the black oak-dominated chert upland oak-hickory in the Ozark Highlands seen in the preceding slide. Stump sprouts like these from the root collar or basal trunk arise from dormant buds whereas stool sprouts arise from adventitious buds between bark and wood. The latter are short-lived and of no value for regeneration (Kramer and Kozlowski, 1979, ps. 61, 150-151). Stump sprouting among haradwoods varies due to many factors including species, age of felled tree, and season of felling. Generally, younger trees sprout better than older ones because the bark is thicker in older trees and the dormant bud may not be able to emerge through it.In addition, older trees are more likely to have connections between bud and pith interrupted (Smith, 1986, p. 471). Sprouting is most abundant and vigerous when trees are cut during dormacy (Kramer and Kozlowski, 1979, p. 272). Sprouts that arise at the basal part of a stump are superior to those growing from the top of a stump because they are stronger and less apt to break off (Smith, 1986, p. 472). Cutting low stumps like the ones seen in these two slides is a trademark of good forest harvest practice. It is the mark of sloth and sloppiness to leave high stumps with trees of this size and straight boles with little butt swell. Ottawa County, Oklahoma. August. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 52 (White Oak-Black Oak-Northern Red Oak) tending toward SAF 110 (Black Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
111. Healthy low stump sprouts of a young black oak- Regeneration of certain hardwood species like oak by coppicing is a sound regeneration method. Combined with uneven-aged management it provides a minimum of impact on the forest ecosystem and wildlife habitat. There are times when clearcutting or even-aged management is also beneficial. It may result in faster regeneration, greater wood yield and improve habitat for deer through greater browse production. It can also release climax tallgrass species and produce more forage for livestock and wildlife as shown in the next slide. Ottawa County, Oklahoma. Estival aspect, September. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 52 (White Oak-Black Oak-Northern Red Oak) tending toward SAF 110 (Black Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
112. Release of big bluestem the first summer following selective cutting in an oak-hickory forest- This is a fine colony of big bluestem (appropriately called “timber grass” by local hillbillies) at anthesis at the end of the first growing seasonfollowing the selective harvest (eight months post-logging) seen in the last three slides. This stand of the native decreaser grass was released from growth-limiting shade along a skid trail. Other such examples of release occurred throughout this chert upland black oak-dominated forest in the Ozark Plateau. It is a textbook example of transitory forest range (as soon as the crowns of oak and hickory trees fill in the openings the grass and forbs will again be suppressed). This demonstrates how Forestry and Range Manageament often go hand-in-glove.
Miscellaneous Forest Types Within the General Oak-Hickory Region
Forest regions of Braun (1950, ps. 33-37) were each named for the "climax association normally developing within it", but forest region and forest association are not always coextensive. Also, while "a specific climax association charactrizes a region" there are numerous forest communities within a region that have species compostion, forest structure, and physiogonomy that are more like (and floristically have more in common with) forest associations of other forest regions (Braun, 1950, p. 34). These forest assocations, forest dominance types, and perhaps other hierarchial and classification units of vegetation may be physiographic or edaphic climaxes rather than climatic climaxes whereas other forest vegetation might be postclimax, preclimax, or subclimax (Braun 1950, p. 13). Some of these "outlier" or "island" forest cover types within the Oak-Hickory Forest (specifically the Oak-Hickory Forest Region) and characteristic plant species of them were presented below.
113. Bottomland hardwood forest on north slope in Ozark Plateau- Extremely species-diverse community shown here in early spring. Species include sycamore, extreme right foreground; western hackberry, foremost trunk; black walnut (Juglans nigra), two trees immediately behind preceding foreground trees; chinkapin oak, grey trunk just behind walnuts; sugar maple (Acer saccharum), two black trunks forming V-shape in background and one behind and partly concealed by the sycamore; basswood or American linden (Tilia americana), on upslope to left of maples; and white ash (Fraxinus americana), left-center midslope. Redbud and flowering dogwood are in the shrub layer but the dominant shrub is spicebush (Lindera benzoin) which comprises a lower shrub layer.The herbaceous layer, which is clearly visible in this early spring stage, includes bloodroot (Sanguinaria canadensis), trout lily or yellow dog-tooth violet (Erythronium americanum), broadleaf waterleaf (Hydrophyllum canadense), rue anemone (Anemonella thalictroides), false rue anemone (Isopyrum biternatum), and scattered colonies of the ever-present Mayapple. Following completion of their annual growing season these species are replaced in their location by great Indian plantain (Cacalia muhlenbergii) which then dominates a tall herb layer through mid-summer. This is a unique transect view going from the first terrace of a mid-size creek to the top of limestone bluffs. The upper-most forest is dominated by black and red oaks as shown in preceding slides. Downslope from the upland oak forest is a mid-slope sugar maple-basswood-chinkapin oak zone. The bottomland or floodplain forest is the sycamore-hackberry-black walnut-white ash forest. Thus this transect includes SAF cover types 61(River Birch-Sycamore, variant form) or variant of 94 (Sycamore-Sweetgum-American Elm), 26 (Sugar Maple-Basswood), and 52 ((White Oak-Black Oak-Northern Red Oak), the latter is indistinguishable in background. Ottawa County, Oklahoma (bank and bluffs of Lost Creek). March, early vernal aspect. Mesic bottomland forest according to Missouri Natural Areas Committee (1987). FRES No. 15 (Oak-Hickory Forest). No obvious Kuchler unit. Brown et al (1998) units of Mixed Hardwood Series in Southeastern Swamp and Riparian Forest biotic community. Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

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

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

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

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

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


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

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


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

Ottawa County, Oklahoma. Estival aspect, July. FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). Combination SAF 93 (Sugarberry-American Elm-Green Ash) X SAF 94 (Sycamore-Sweetgum-American Elm). Mixed Hardwood Series in Southeastern Swamp and Riparian Forest biotic community of Brown et al. (1998).


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

118. Leaves of green ash- Ottawa County, Okahoma. July.

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

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


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

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

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

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

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

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

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

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

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

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

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

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


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

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

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

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

Ottawa County, Oklahoma. August.


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

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


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

Permanent spring, Ottawa County, Oklahoma. April.


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


136. Bottomland forest of western hackberry, sugarberry (Celtis laevigata), and American elm (Ulmus americana) with an understory almost exclusively of broad-leaf wood oats (Uniola latifolia= Chasmanthium latifolium) with scattered solidary plants of the shade-tolerant composite, elephant's foot (Elepohantopus carolinianus). Beautiful plant community and an example of permanent deciduous forest range in contrast to the typical transitory feature of these dense hardwood forest types. Ottawa County, Oklahoma. September, late estival aspect. No obvious FRES or Kuchler designations as this is a smaller unit. Neither was this relatively unique community described by Braun (1950) though it corresponds to general creek and river bottom vegetation in the Ozark Mountains. SAF Cover Type 93 (Sugarberry-American Elm-Green Ash). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).
137. Spikelets of wood oats (Uniola latifolia)- The inflorescence of wood oats is one of the most attractive of North American Gramineae. In extensive stands (like the one seen immediately above) wood oats creates a stikingly beautiful aspect. Uniola (= Chasmanthium) species have some of the most laterally compressed spikelets of any grass. Ottawa County, Oklahoma. August.

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

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


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

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

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


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

Davy Crockett National Forest, Houston County, Texas. March, vernal aspect (post dogwood blooming stage ). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). Variant SAF 94 (Sycamore-Sweetgum-American Elm) that had former SAF designation of Sycamore-Pecan-American Elm. Closest biotic community designation of Brown et al. (1998, ps. 37, 38) was Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1, but there should have been a Oak-Hickory Series for Southeastern Deciduous and Evergreen Forest 123.. South Central Plains- Southern Tertiary Uplands Ecoregion, 35e (Griffith et al., 2004).


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

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


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

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


143. White Christmas in the Ozarks- Snowfall (especially of major accumulation like a half foot or more) is rare in the oak-hickory forests of the Ozark Mountains. In fact, it is not unusual for there to be winters without snow and with winter precipitation being limited to rain or the treacherous "winter mix" of rain, freezing rain, sleet, and snow (ice storms). The "winter wonderland" of wet snow and an accumulation of 10 inches shown in these photographs demonstrated the ever-changing weather of the Ozark Region and the necessity of some winter adaptation for it's species. Besides it was "kinda purty".

Post oak and black hickory were dominant tree species with northern red oak and black cherry the major associates of the canopy layer. Understorey was dominated by buckbrush and poison oak; Virginia creeper formed a uniform intermediate layer extending from ground level to tops of trees while wild grapes (Vitis spp.) were local dominants of the intermediate layer. In early spring the herbaceous layer was usually composed exclusively of Mayapple (Podophyllum peltatum) colonies. All plant life had settled in for a short winter nap in this scene which had all but entirely melted within 24 hours of these photographs.

Ottawa County, Oklahoma. Christmas Day, 2002.


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