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Forest Range Types of Eastern North America
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The fundamental and practical distinction between coniferous and deciduous forests is useful (and was used herein), but precise, non-arbitrary "lines" are impossible when presenting and discussing forest range types in the eastern half of the continent. This is especially the case when climax or potential natural vegetation is used as the basis for forest types (ie. when cover types, or the more specific management cover types, are discussed as being more or less synonymous with permanent forest types). As discussed in detail below, the epic work of Lucy Braun (1950) is still the definitive basis for the ecological discussion and classification of those North American forests which extend from the Atlantic Coast to slightly beyond the Missouri and Mississippi River drainages. Braun (1950) included all the coniferous forests (forest types, regions, etc.)-- the generic "southeastern pine region"--as part of her one Deciduous Forest Formation. The forest range types included in the following section include coniferous, deciduous, and mixed coniferous-deciduous forests. This is confusing but unavoidable given the nature of the vegetation and the standard understanding (the Braun interpretation) of ecological relations and classification of this forest vegetation. Most of the southeastern pine types presented are management cover types maintained silviculturally as more economically valuable coniferous forests rather than as the climax mixed hardwood-pine forest types. In other words, efforts were made to fit the Society of American Foresters (1980) cover types with the climax types of Braun (1950) and the potential natural vegetation units of Kuchler (1966). The major forest communities or forest zones of eastern North America are broad or wide in their spatial patterns unlike the narrow zonation characteristic of the forests of western North America. The “young” mountains of the western part of the continent are taller (in fact, still getting taller) and as a result have more elevation-based zonation of vegetation than do the geologically older and more eroded (lower) eastern mountains such as the Applachians or Ozarks. So too, are the soils of the Atlantic Coast more zonal (ie. major soil units are larger or broader in spational dimension like those of the vast continental interior whereas soils of the Rocky Mountains and the Pacific Slope ranges are more of the intrazonal spatial scale. See for illustration the national soil map of dominant soil orders and suborders (Soil Survey Staff, 1998). Vankat (1979, p. 137) wrote that relief within the eastern deciduous forest “is quite variable” yet earlier Vankat (1979, p. 41) had also correctly noted that “low hills “ were characteristic of much of this deciduous forest region. Again, contrast this with the extreme physiography of the Rockys or Sierra Nevada-Cascade Ranges. |
| The classic and still-definitive work on forests of eastern North America
(approximately east of the 98th meridian) is the life’s work
of Dr. Lucy Barun (1950). Braun interpreted this entire vegetation as
one great forest formation existing as a mosaic of forest regions which
in turn were made up of community units that she labeled variously as
belts, areas, districts, sections, divisions, etc.
“The Deciduous Forest Formation of eastern North America is a complex vegetation unit most conspicuously characterized by the prevalence of the deciduous habit of most of its woody constituents. This gives to it a certain uniformity of phsiognomy, with alternating summer green and winter leafless aspects. Evergreen species, both broad-leaved and needle-leaved, occur in the arboreal and shrub layers, patticularly in seral stages and in marginal and transitional areas. They are not, however, entirely lacking even in some centrally loocated climax communities” (Braun, 1950, p. 31). “The Deciduous Forest Formation is made up of a number of climax associations differing from one another in floristic compositon, in physiogonomy, and in genesis or historical origin. While the delimitation of associations may be made on a basis of dominant species, and it is from these that the climax is named, dominants alone fo not suffice for the recognition of these units. … Although the delimitation in space of an association is difficult, if not impossible, it is entirely possible to recognize and to map forest regions which are characterized by the prevalence of specific climax types, or by mosaics of types. These regions are natural entities, generally with readily observable natural boundaries based on vegetational features. … Forest regions must not be confused with climax associations. Even though a region is named for the climax association normally developing within it, it should not be assumed that the region is coextensive with the area where that climax can develop. Each of the several climaxes, although characterizing a specific region, nevertheless occurs in other regions.” (Braun, 1950, p. 33-34). Braun (1950, ps. 35-37) listed nine forest regions making up the Deciduous Forest Formation of eastern North America: 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. |
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Historical
Footnote and Editorial
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| 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”. |
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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). |
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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). |
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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. |
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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? |
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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. |
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| 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. |
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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. |
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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. |
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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. |
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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). |
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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). |
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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). |
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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). |
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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). |
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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). |
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White Oak (Quercus alba)-Black
Oak (Q. velutina)-Northern Red Oak (Q. rubra) Forests
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Mixed white-black-northern red oaks forests probably constitute the "umbrella" or general forest type of which forests dominated singularly by white, black and northern red oaks are affilitated with each of these three being a forest cover type--frequently subclimax or climax--itself on certain forest sites. According to cover type descriptions by the Society of American Foresters (Eyre, 1980, ps. 41-44) the white oak- black oak-northern red oak cover type (52) is clearly the most extensive or widely distributed forest type among these various cover types or forms of the oak-hickory association. Another rational perspective of SAF forest cover type 52 is that it is a transition zone among white oak (SAF 53), black oak (SAF 110), and northern red oak (SAF 55) types. In the western portion of the Ozark Plateau upland forests of the white oak-black oak-northern red oak cover type were usually found by this author to have developed on predominately north and east slopes with better drained soils resulting in overall moderately mesic to somewhat dry habitats. This corresponded to the dry-mesic chert forest of Nelson (2005, ps. 125-130). The Society of American Foresters (Eyre, 1980, ps. 41-42) listed white oak as the first of three dominants for this forest cover type and indicated that white oak occurrs across the moisture gradient from dry to moist. This followed the conclusion of Braun (1950, ps. 167- 168) that white oak was more common on north slopes, but that it was a major species on "many slope forests" where black oak was often "its principal associate" except on more mesic habitats where northern red oak held this status. Sugar maple is also a component species on slope forests, especially on limestone soils, "where it may be associated with white oak, red oak, chinquapin oak, basswood, and hickory" (Braun, 1950, p. 168). Where sugar maple is a dominant rather than associate species the forest vegetation is recognized as another form of Ozark Plateau forest (Braun , 1950, ps. 168-169) and a variant or form of the sugar maple cover type and/or the sugar maple-basswood cover type (Eyre, 1980, ps. 29-30, 31-32, respectively). These latter hardwood forests were interpreted as component communities--in effect, edapho-topographic climaxes--of the general oak-hickory association as was explained by Braun (1950, p. 33-34) and quoted in the introduction of this chapter above. The sugar maple types of the Ozark Plateau were treated later in this chapter of Range Types because they are climax forest types (associes vs. associations in the Clementsian model) of the oak-hickory association in Ozark Mountains forests. It was necessary to mention and draw attention (briefly) to Ozark sugar maple forests at this juncture because these cover types blend into the white oak-black oak-northern red oak type (SAF 52) and the northern red oak cover type (SAF 55); that is, there is a continuum among these various cover types, each of which can be climax on specifie forest range sites. |
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15. The mixed oak type above a hollow- Local second-growth stand of the white oak-black oak-northern red oak forest type in the western Ozark Plateau. This forest range vegetation grew on a predominately east slope. It was a more mesic forest site with co-dominants being white and northern red oaks. Woody plants from left to right in fore- and midground: sugar maple saplings with northern red and white oaks behind these; then from right of center (still left to right line-up) white oak, northern red oak, white oak, two northern red oaks, and, the shrubat far right, flowering dogwood (Cornus florida). Saplings behind these trees and shrubs were mostly bitternut or pignut hickory that was generally the associate species. Summer grape (Vitis aestivalis) was present with some growing among the sugar maple saplings (to confuse viewers trying to distinguish plant species). No herbaceous layer in this local stand. Instead there was a leaf layer acting as mulch that formed the ground layer of this forest vegetation. Ozarkers are fond of specifying that the hills are not as high as the "hollers" are deep. The composite mix of dominant trees just enumerated "ringed" the head of a hollow in the Springfield Plateau portion of the Ozark Plateau. This forest had no large animals in it except free-ranging white-tailed deer (Odocileus virginianus). Currently, deer populations were below carrying capacity of this forest range. Approximately 50-60 years ago this had been legal open range resulting in severe overgrazing and overbrowsing by livestock, especially hogs. Fisher Flats, Ottawa County, Oklahoma. June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005). |
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16. Mixing around the edge- A second-growth stand of white, black , and northern red oak with a diverse understorey formed a composite example of this forest cover type at the head of a hollow on a predominately east slope in the western Ozark Plateau (Springfield Plateau). The three Quercus species were "about evenly divided" as tri-dominants. The associate species was pignut, bitternut, or (sometimes) white hickory. The principal shrub was flowering dogwood. The largest tree (left margin, foreground) was white oak as was the tree at far right. Center (somewhat right of center) tree was a black oak. There was some regeneration of all oak and hickory species in spite of considerable shade. It was concluded that for this was the climax vegetation for this forest range as Eyre (1980, p. 42) concluded for the Ozark Highlands. This local second-growth stand of mixed oak was, like the stand presented in the last slide, also at the head of a hollow. Likewise grazing/browsing by large animals was limited to that by white-tail deer. Fisher Flats, Ottawa County, Oklahoma. June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005). |
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17. Under the oaks- A photo-quadrant that featured local herbaceous constituents of the east slope, second-growth stand white oak-black oak-northern red oak forest described in the immediately preceding caption. As in the instance of that stand there was regeneration of all major tree species (the three oaks and bitternut hickory). Again, flowering dogwood was the dominant shrub. The dominant herbaceous species was big bluestem (Andropogon gerardii). In this photograph the most abundant forb was the introduced perennial legume serecia lespedeza (Lespedeza cuneata). Shrubs comprised a pronounced and varied layer including smooth shumac (Rhus glabra) poison oak or ivy (R. radicans= Toxicodendron radicans), summer grape, Virginian creeper (Parthenocissus quinquefolia), greenbriar (Smilax spp.), and flowering dogwood. The various species of woody vines formed a "unifying" layer from ground layer to canopy of tallest trees. Fisher Flats, Ottawa County, Oklahoma. June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005). |
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18. Holler away- Two photographs of a second-growth mixed oak-sugar maple forests at head of another hollow in the Springfield Plateau. The forest range vegetation introduced here and described in the next series of slides was in the general vicinity of the head-of-hollow white oak-black oak-northern red oak-bitternut hickory forest presented immediately above. This was a north versus the predominately east slope of the preceding stands. Composition of the adult and regenerating tree layers (including sugar maple) was the same as east slope stands except that bitternut or pignut hickory was barely present (largely absent) from this north slope forest. The most pronounced differences between the north and east slope forests were in the herbaceous and shrub layers. The dominant herbaceous species on this north slope was the native perennial legume, tick clover (Desmodium nudiflorum), whereas the dominant herb on the slightly drier east slope was big bluestem. Flowering dogwood was common to both east and north slope forests, but on north slope stands the associate shrub was highbush or squaw huckleberry or deerberry (Vaccinium staminium) followed by eastern or common redbud (Cercis canadensis). Young individuals of sassafras (small sapling-size), although a tree species at maturity, comprised another constitutent of the shrub layer. Two photographs showing the same local stand of mature trees of a second-growth mixed oak-sugar maple, Ozark Hihglands forest on a north slope (mesic) with very limited understorey (mostly shed leaf ground layer). The first of these two photographs (vertical) presented a textbook line-up of the species composition of this forest cover type. Tree species from left to right: white oak, northern red oak, white oak, northern red oak , and two white oaks. The second photograph presented most of the same trees in a horizontal plane (left to right): white oak, northern red oak, white oak, northern red oak, and white oak. In the shade of this dense stand there was little if any understorey except for a ground layer of shed leaves with an occasional plant of Desmodium nuiflorum, another unidentified (pre-bloom) Desmodium species, and sugar maple seedling. This tract of forest had not been grazed/browsed by livestock for decades (probably about a half century). From the frontier era up to that time forests in this area had been subjected to free-ranging livestock (including swine) under legal open range status with abuse of forest resources being "the norm". Spring burning of these woods was also a common practice until about the same period. Subsequently the only ungulates-- large vertebrates at all for that matter--in this forest were free-ranging white-tailed deer which ranged throughout this and adjacent forests. At time of these photographs deer numbers had not reached the stage of overpopulation, not yet anyway. Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005). |
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19. A hollow cast of charaacters- At the head of a hollow on a north slope in the Springfield Plateau of the Ozark Highlands a second-growth, hardwood forest of white oak-black oak-northern red oak (tri-dominants) with sugar maple as the associate species had developed. This local stand provided a good example of this climax forest cover type and its range vegetation. Tree species in this photograph (left to right) were a mid-size white oak and two young pole-size northern red oaks. Between these three trees and in mid foreground were two small, crooked-trunk saplings of flowering dogwood. Eastern redbud was present though scarce. Scattered small saplings of sassafras were also present as in the woody layer that was dominated by shrubs (again, mostly flowering dogwood). Major herbaceous species were two species of tick clover (Desmodium nuiflorum and another Desmodium species in pre-bloom stage that could not be positively identified). Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005). |
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20. Principal players- Good species composition "quadrant" showing climax species of: 1) dominant trees, dominant shrub, and dominant forbs which also portrayed layers of vegetation and general forest structure. Plants in a left to right line in foreground of slide were: leaves of black oak (left margin), two black oaks, bent sapling of sugar maple (foreground), and white oak (extreme right margin). Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005). |
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21. Developing forest with the understorey coming first (or forest vegetation on the edge)- An edge of the north slope white oak-black oak-northern red oak forest featured here provided an opportunity to document a successional stage of forest development. All three oak species and sugar maple had ample regeneration at the gap-like edge of this mesic Ozark Hihglands second-growth forest. Summer grape was also well-represented by young and conspicuous plants as was highbush or squaw huckleberry (short woody plants with large linear shade leaves). Some sassafras was present as large seedlings. The major herbaceous species was one of the rosette panicgrasses (one of the species in Dichanthelium subgenus), which in the post-spring bloom and grain-shatter stages could not be identified positively, but seemed most likely to be western panicgrass or western rosettegrass (Panicum lanuginosum= Dichanthelium acuminatum var. implicatum= Dichanthelium lanuginosum). Members of the Dichanthelium subgenus produce winter foliage in the form of a rosette (hence rosette Panicum spp.) from which infloresecences emerge to typically bloom in mid to late apring and then again in autumn. This rosette panicgrass species presented here had formed an extensive colony at edge of the north slope mixed oak-sugar maple forest. Poverty oatgrass (Danthonia spicata) was also abundant-- by standards of the shade cast by this forest even at its outer edge. Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005). |
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22. Highbush huckleberry, squaw huckleberry, or deerberry (Vaccinium staminium)- On oak-hickory forest range of the Ozark Hihglands where this shrub occurs it is a defining species because highbush huckleberry has nothing even approaching the commonness and widespread feature of other shrubs likc flowering dogwood, blackberries, smooth sumac, poison oak or ivy, and Virginia creeper. Highbush huckleberry is usually restricted to the more mesic, yet well-lite forest sites. This specimen was growing at the edge of the north slope, white oak-black oak-northern red oak-sugar maple forest shown in the four immediately preceding photographs. Seedlings in lower corner of foreground were black oak and blackjack oak (Quercus marilandica). Ottawa County, Oklahoma. July (estival aspect); immature fruit stage of phenology. |
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23. Under uneven-aged management- General view of an upland white oak-northern red oak second-growth forest in Boston Mountains. Shortleaf pine (Pinus echinata) is a very infrequent component of this forest range community, but not "anywhere near" associate species status. As such, this forest tract was presented as another example of the mixed oak type in which white oak ranks the edge as the "first-among-equals" dominant. Red maple was an important member of the understorey with its ultimate staus yet to be determined in this forest (see below). Shrubs included flowering dogwood, redbud, buckbrush or coralberry, poison oak, poison ivy, lowbush huckleberry (Vaccinium vacilians), and various kinds of blackberry. Grasses and forbs were in short supply, but some species were noted in the next caption.. Most of the understorey consisted of various layers made up of the age classes of the co-dominant oak species and red maple. Ozark National Forest, Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004) |
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24. White oak-northern red oak forest under uneven-aged management- Appearance "closer-up" of a second-growth upland mixed oak forest under selective harvest. Species composition and structure in the interior of this forest range was emphasized in this photograph. White oak was the most common (major) dominant, but northern red oak gave it a good run for the money. In this view there was tremendous regeneration of bittrnut or pignut hickory (Carya cordiformis as well as northern red oak (center foreground). Red maple at seedling and small sapling age classes was extremely abundant with its successional and domiance status unknown. Most of the understorey was regeneration of oaks.bitternut or pignut hickory, and red maple with the various layers made up of the different age classes of these hardwood trees. Shrub species included flowering dogwood, redbud, buckbrush or coralberry, poison oak and poison ivy, Viraginia creeper, and various kinds of blackberry. Not much by way of grasses and forbs. Virginia and silky wildrye (Elymus virginicus and E. villosus, respective) were the main grasses, but they were sparse and widely scattered. Mayapple, one of the more characteristic herbaceeous dicots throughout the Ozark Plateau, was the most conspicuous forb. Clearly this was was a browse range with white-tail deer the game species of choice (the range animal managed for). Obviously the main commodity from this forest tract was saw timber. It was instructive that wood from this forest included a major representative of the white oak group (Leucobalanus subgenus) and the red oak group (Erythrobalanus subgenus). The red oaks are especially prized for fuurniture manufacture whereas the white oaks have traditionally been used as rougher construction materials as well as cooperage, wooden containers (eg. barrels) made up of a round head (made of wood) on both ends and a body of one to many staves (wooden; long, narrow, and curved one-piece units) held together by hoops (usually metal). Historically one of the most important uses of oak was for timbers (by definition at minimum 5 inches by five inches), especially railroad ties (=crossties) which are eight inches by eight inches and eight feet long. Hickory is prized for tool handles because it is so resistant to shock, breakage, and deformity (tough wood fibers of hickory are very resilant or plastic), is much easier on workers' hands than metal or fiberglass, and aesthetically pleasing (at least to those with good taste and not obsessed with synthetics). Ozark National Forest, Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004) |
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25. Interesting interloper in the understorey- Large seedling of red maple (Acer rubra) in understorey of an upland white oak-northern red oak second-growth forest with uneven-aged management. This species was very abundant in this developing forest, but the ultimate successional status and role of this species in an mixed oak-hickory was unknown. Red maple is a fairly palatable browse for white-tail deer. All species of livestock will browse on hardwoods to some degree or the other. They are especially fond of mast, including acorns and hickolry nuts. Hogs generally browse less than other kinds of livestock, but their disturbance of soil (rooting and wallowing) and feeding on roots makes swine the most potentially destructive of all livestock in hardwood forests, perhaps especially the oak-hickory forest range types. Ozark National Forest, Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004) |
| Note on location: An example of the white oak-black oak northern red oak forest in which black oak was a minor component (generally not present) was presented below at end of the section devoted to the variant forest cover type of American beech (Fagus grandiflora)-white oak-hickory. |
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Black Oak (Quercus velutina)
Forests
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In western parts of the Ozark Highlands, such as the Springfield Plateau, black oak-dominated forests are some of the most common remaining forest tracts as they are found on some of the driest, shallowest, rockiest soils and, hence, generally the harshiest of remaining forest sites. The more mesic forest sites with their deeper, less stoney soils (ie. the less marginal, more-or-less arable land) naturally developed old-growth forest that were dominated by northern red and white oaks (with black oak as an associate species). Most of these more productive mesophytic forests were converted to hill-side pastures, strawberry patches, fruit orchards, and even field or vegetable crops decades ago. Some of the cropland derived by clearing the original (post-Indian) white oak and northern red oak forests was retired from intensive cropping years later as a result of socio-economic developments (ie. loss of small, inefficient farms). Some of the old-fields (abandoned farmland) on these almost subsistence-level (and hobby) farms have gone back through secondary plant succession to the stage that they revegetated to seral or subclimax black oak forest. Also, some old-growth (climax) black oak forest have recovered and are routinely logged (usually high-graded or clearcut) at, say, quarter to half century intervals. Some examples of these second-growth black oak forests and secondary plant succession on them following disturbance was presented in the following section. |
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26. Dormancy afforded a good "look-see"- With leaves on the ground instead of in the canopy an all-in-one view was provided of structure, arrangement, species composition, and lumber crop of a climax black oak-northern red oak forest full of mature trees.Black and white hickory (Carya texana and C. tometosa, respectively) were associate tree species. There were some post oaks, but this was clearly a forest site for the black oak species with black oak the dominant and northern red oak the associate species. There was also mentionable cover of black cherry, which probably indicated infrequent surface fires in this stand. Grasses were few in understorey, but dominant was broomsedge bluestem. There had been a history of cattle grazing in this stand, but it was generally light (mostly breechy ones looking for better pasture and finding worse than they left). Canopy was too dense for much herbaceous understorey other than for that of early season species like Mayapple (Podophyllum peltatum). Many of the larger trees in this stand were over-ripe and dying or even dead. For this forest site and this stocking rate these trees were probably of their maximum size. There were several snags. However most regeneration was hickory. This was a second-growth forest, yet species composition and structure of forest was climax. This forest approached old-growth status. Ottawa County, Oklahoma. FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak). |
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27. Now a summer view- With leaves back up in the canopy a vastly different perspective was afforded of the same climax black oak-northern red oak of mature trees as introduced immediately above. There was ample regeneration of hardwoods, but hickory predominated. Ottawa County, Oklahoma. FRES No. 15 (Oak-Hickory Forest Ecosystem).
Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black
Oak). |
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28. Interior of the climax black oak-northern red oak forest presented in the immediately preceding two sets of two slides each. Abundant reproduction of hickory so that this stand was becoming a hickory phase or variant of the black oak cover type. Ottawa County, Oklahoma. FRES No. 15 (Oak-Hickory Forest Ecosystem).
Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black
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Forest remains: harvest of the black and northern red oaks-black and mockernut hickories forest- The tract of upland black and northern red oaks-black and white or mockernut hickories forest shown immediately above was highgrade-logged and basically clearcut in winter (and on the sly) 15 years after the preceding photographs were taken. The cutover land was re-photographed in July of the second spring-summer growing season post-harvest. Redevelopment of forest vegetation on this improperly logged, degraded tract was a combination of typical old-field plant succession beginning with pioneer or colonizing species including giant ragweed (Ambrosia trifida), horseweed or mare's tail (Conyza canadensis), hairy crabgrass (Digitaria sanginualis), and the naturalized annual, beefsteak plant (Perilla frutescens) along with climax dominants that were regenerating both sexually and asexually. Most asexual reproduction was in black oak (and some northern red oak) by coppicing from mid-size stumps. The other three species besides climax dominants that had appreciable cover were black cherry (Prunus serotina) and sycamore (Platanus occidentalis). Trees with noticeably less cover were , Kentucky coffee tree (Gymnocladus dioica), American elm (Ulmus americana), black walnut (Juglans nigra), and eastern red cedar (Juniperus virginiana). Surprisingly, the typical pioneer tree species, sycamore had less cover than would be expected judging from recovering vegetation on other severely disturbed sites in the immediate vicinity. Other tree species commonly found as pioneer or early seral plants included sassafras (Sassafras albidum) and persimmon (Diospyros virginiana). These two species were also less abundant than in the typical situation observed for disturbance in this vicinity. This cutover land had not been denuded to the extreme old-field state. Most regeneration of dominant climax trees was of the hickory species present as saplings before logging (described and ahown above). Most reproduction of oak (especially black oak) at this point in forest recovery was by stump-sprouting. There were few oak or hickory seedlings at this stage of secondary succession.Most seedlings were sycamore, black cherry, American elm, and sassafras. Some smaller though sexually mature trees that were left standing in the clearcut as well as uncut adult trees on the perimeter of the forest (see below) served as seed trees. Otherwise, re-establishment of oaks and hickories depended on coppicing and/or the soil seed bank. Shrubs were very important in this recovering vegetation. Far and away
the most important shrubs were blackberry (probably This remarkably diverse recovery vegetation also included numerous species of forbs, both native and naturalized. The latter included the biennial, flannel mullein (Verbascum thapsus), and introduced legumes, both perennials such as white clover (Trifolium repens) and red clover (T. pretense) and annuals like Korean or Japanese lespedeza (Lespedeza striata). Other forbs included numerous natives like biennial evening primrose (Oenothera biennis); pokeweed (Phytolacca americana); and various composites, especially giant ragweed (Ambrosia trifida), common ragweed (A. artemisiifolia), Carolina elephant foot (Elephantopus carolinianus), Baldwin ironweed (Vernonia baldwinii), black-eyed susan (Rudbeckia hirta), Canada wild lettuce (Lactuca canadensis), and wingstem crown-beard (Verbesina alternifolia). The pioneering composite, horseweed or mare's tail was widespread and locally dominant although generally not in dominating proportions that might have been expected (and, probably, present in the first season following harvest). There was also some immature smartweed or knotweed plants (Polygonum sp.) that could not be identified as to species along with toothed spurge (Euphorbia dentata). Pokeweed was the most widespread and overall most important forb, but giant ragweed was a close second forming dense stands from which almost all plants of other species were excluded. The only common grass was hairy crabgrass. The perennial grasses were represented almost exclusively by broomsedge bluestem (Andropogon virginicus). Poverty oatgrass (Danthonia spicata) already present in the understorey persisted by larger stumps where it was not "overrun by pioneer species. There was also an occasional plant of purpletop (Tridens flavesus) next to shelter (eg. uncut tree).There were also some plants of green and/or yellow foxtail (Setaria viridis, S. glauca, respectively) which could not be distinguished at vegetative stages present at time of photographs. Likewise there were incidental plants of Carex and Cyperus species. White-tailed deer where the only species of large vertebrate that had access to this was black oak-northern red oak forest which had not been grazed/browsed by cattle or hogs for several decades. In the climax oak-hickory forest that approached old-growth state there were very few plants available for grazing and/or browsing in the understorey. These were limited to leaves and buds on regenerating oak and hickory species, flowering dogwood, poison oak/ivy, and incidental plants (trace amounts in absolute and relative cover) of blackberry, poverty oakgrass, and sedge. This tract of black oak-dominated upland Ozark Plateau forest was clearly transitory forest range. Once (after) canopy cover reached almost 100% almost no light that could penetrate throught the dense foliage to reach the ground (soil) surface. After leaves were fully grown in the forest canopy each spring light could only reach to the level delineated by lower leaves on shrubs like flowering dogwood and saplings of oak, hickory, black cherry, etc. In this climax oak-hickory forest with mature ("over-ripe") trees spring forbs like mayapple (Podophyllum peltatum) were not present except in natural forest gaps and at forest edges. Organization Note: a black oak-pignut or mockernut climax forest that served as permanent forest range was presented and described in the Use and Abuse of Oak-Hickory Forests section of Oak-Hickory Forests-II. Harvest of this forest was an example that socioeconomic factors often override biological ones, and that many if not most human endeavors (not excluding silvicultural operations) involve human emotions including greed and jealosy. Logging of this oak-hickory forest was a case of timber theft. A thieving son snuck in and stole this standing timber from his aged father who for romatic and aesthetic reasons wanted the forest with its many mature trees left as it was. The conniving son left uncut the trees around the edges of this forest to hide what he was doing inside the tract. The timber-buyer left some of the smaller trees inside the perimeter that he felt were not worth felling (probably to the chagrin of the greedy son who worked up all the slash to sell as fuelwood). Not only was this dysfunctional family relations, but it was also improper forest practices: cut-and-run, sloppy, (and illegal) logging of the worst form, the kind that gives the forest products industry a bad reputation. Thief of "free grass" and "timber for the taking" was standard fare in early history of use and abuse of forest and rangeland resources. The important thing for rangemen and foresters to bear in mind is that these resources are, to large degree, renewable (even with improper harvest methods and other forms of abuse). The following series of slides was of the black and northern red oak-black and/or Texas and mockernut hickories upland forest in the second growing season following highgrade logging that was, in effect, land-clearing. |
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29. Missing trees, stolen timber- The second-growth black oak-dominated (northern red oak, associate) forest shown and described here was secretly clearcut with a few smaller trees left around the perimeter so that a son could hide the timber theft from his aged father. The crime scene and second-year successional vegetation was presented in two photographs that gave a general view of the cleared forest. The wasted sound logs in the first of these slides added resource waste and abuse to the weed patch effect. In both slides remaining trees (a young northern red oak in the first; black and northern red oaks, Texas or Ozark and mockernut hickories in the second) "looked on" cutover land supporting a pioneer stage of recovering forest vegetation dominated by such colonizing species as the annual composite, giant ragweed, along with common ragweed, hairy crabgrass and such disturbance-loving perennials as pokeweed, black-eyed susan, Baldwin ironweed, wingstem crown-beard , Carolina elephant's foot. Foreign weeds that benefitted from this disturbance included flannel mullein and beefstake plant. Common shrubs included several species of blackberry, summer and fox grape, and smooth sumac. This stage of secondary plant succession on a black oak-dominated Ozark Highlands upland forest was about half-way into the second warm-growning season following clearcutting of a second-growth forest that approached old-growth status. Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak). |
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30. Into the weeds (and the ticks and chiggers)- Two close-in views of seral vegetation on a forest site that two winters or one an a half warm-growing seasons before had been a climax black oak-northern red oak (dominant and associate species, respectively) upland forest that was harvested by clearcutting (in effect a land-clearing operation). The most abundant herbaceous species were giant and common ragweeds. Other composite forbs included black-eyed susan, common horseweed or mare's tail, an Carolina elephant's foot. Pokeweed and flannel mullein represented native perennial and naturalized Eurasian biennial forbs, respectively. Passionflower (Passiflora incarnata), a perennial herbaceous vine or twining forb, was conspicuous along right margin (about half-way up) in the first slide. The only grass with cover and density worthy of note in the seral range vegetation presented in these two photographs was hairy crabgrass. Shrubs included blackberry, smooth sumac, and summer and fox grape. Tree species present as pre-existing (present before logging), stump sprouts, or seedlings included black oak, northern red oak, post oak, Texas or Ozark hickory, mockernut hickory, black cherry, American elm, red elm, sassafras, and persimmon. Young trees (either too small to make a saw log and/or serving as a cover to conceal the crime sceene from outside the clearcut) included all of the oak and hickory species as well as a few black cherry. Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak). |
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31. Edge effect, a benefit of forest harvest- Two more general views of a clearcut in a black oak-dominated climax forest in the Springfield Plateau. These two photographs high-lighted the effect of edge, the union of standing trees at perimeter of the now clearcut forest and seral range vegetation developing on the clearcut through secondary plant successtion midway through the second warm-growing season post-harvest. Some of the more abundant and conspicuous herbaceous species on this clearing included native forbs common to disturbed habitats: smartweed, Baldwin ironweed, pokeweed, giant ragweed, common ragweed. Most of these were of little or, at least, very limited forage value. The forb of most forage value was red clover, a naturalized, Eurasian, perennial legume (barely visible in the first photograph as pink clusters). How such abundant cover of this valuable forage plant developed by the second growing following logging remained a mystery. Farmers in this local vicinity have overseeded permanent pastures to red clover. Clearcutting had converted a climax forest with limited understorey (most of that regenerating hardwoods, especially hickory species) into a cutover pasture or range that, though a far cry from standards of high-quality tame pasture, provided valuable forage plants for livestock and wildlife including an introduced, perennial legume; a palatable, annual grass, and forbs of diverse palatability). This was transitory forest range. The second of these slides featured the numerous woody species that invaded the clearing the first growing season following clearcutting. Most conspicuous were two-year-old seedlings of sycamore (center of photograph). There were stump sprouts and seedlings of black oak, Ozark and mockernut hickories and black cherry. Most of the latter were seedlings or pre-established saplings. Prominent forbs in successional range vegetation shown in the second slide included Baldwin ironweed, giant ragweed, and common ragweed. Young vines of summer and fox grape trailed and wound their way around newly established large patches of blackberry out of which grew black cherry, sassafras, and elm (both American and red). In this arrangement of seral vegetation there were edges within edges:. edges where blackberry patches met weed (eg. colonizing composites, pokeweed, flannel mullein) patches within the edge that was the perimeter of the climax black oak-red northern oak forest. This was shown in more detail in the next set of two slides. In Nature any disturbance--no matter how traumatic or disturbing of existing species, communities, ecosystems, landscapes, etc--is a boon to some other species, communities, ecosystems, and so on. Clearcutting the climax, near old-growth black oak-northern red oak forest that occupied this site was an extreme perturbation, a drastic alternation of that forest ecosystem, that had dire consequences on many living things ranging from old trees and their dependent fungi species, squirrels, pileated woodpeckers (Dryocopus pileatus), and humans due to dissolutionous family relations between a father and a theiving son. That action, that extreme ecological disturbance, also created greately improved habitat for white-tailed deer and northern bobwhite (Colinus virginianus). One of numerous reasons why populations of bobwhite quail have declined throughout much of the Ozark Plateau Region is that their prime habitat of old-fields, recovering cleared forests, and small farm fields had revegetated back (had secondary plant succession progress closer toward) the pre-white man forest. Advanced seral, subclimax, and climax stage oak-hickory forests provide marginal to poor habitat for bobwhite. "Setting back" forest vegetation to pioneer and other early seral stages was (is) of immense benefit to seral species like bobwhite quail and white-tailed deer. When this climax forest (with its high proportion of rotten trees that were "overripe" for quality hardwood lumber) was logged pileated woodpeckers that lived off of insects that thrived on decaying wood as well as cavity nesting mammals including squirrels, 'possums (Didelphis virginiana), and coons (Procyon lotor) came out "loosers", at least in the short run. Critical parts of their habitat were eliminated. Perhaps this was the situation for wild turkey (Meleagris gallopavo) and raptors like great horned owls (Bubo virginianus) and red-tailed hawks (Buteo jamaciensis), all species that commonly lived in the black oak-northern red oak forest with its old age, over-mature trees. However, turkey and coon soon benefitted from increased production of blackberries, pokeberries, etc. Reduction of canopy cover facilitated predation by raptors although owls are still going to have to find dense tree canopy to escape their arch enemy, the annoying, sleep-depriving crow (Corvus brachyrhynchos). The 'possum will miss her favorite hollow tree, but see what a crop of persimmons will be produced for her descendants in a few years. The edge where uncut, timber theft-blocking trees meet and merge with the new weed and brush patch provided a new corridor as well as ample cover for new feed sources for various species of animals and new growing environments for plants. Use of natural resources always producers "winners"and "loosers". Wise use to conservation prophets like Gifford Pinchot consisted of finding that "blend" which, in the spirit of Jeremy Bentham's utilitarianism, "provided the greatest good to the greatest number for the longest period of time". As was shown in later descriptions and discussions of clearcutting this climax black oak-forest, it was quite likely that in the "long run" Pinchot's standard for wise use was fulfilled on this land, in spite of the villainous crime committed by a treasonous son against the rest of his family. Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak). |
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32. A closer look at edges and species make-up of seral vegetation- Two photographs showing seral forest range vegetation about half-way through its second warm-growing season growing adjacent to adult black and northern red oak trees that were left uncut to hid from view a forest clearcut of a climax black oak-dominated upland forest in the western Ozark Plateau. The plant communites that developed along the uncut forest perimeter occured spatially as "rows" (narrow zones) of woody plants (both tree saplings and shrubs) while there were larger patches of "weedy" colonizing species interior to the "belts" of wody vegetation. In the first photograph a narrow belt of adult black and northern red oaks was in the background while the foreground was an area dominated by giant and common ragweed. The bulk of the recovering forest vegetation (midground of photograph) was a blackberry patch with numerous saplings of black cherry, American and red elms, and sassafras (less persimmon) along with fox and summer grapes. The second photograph also showed a "weed patch" that was almost exclusively giant ragweed with some common ragweed and mare's tail orhorseweed (foreground) contiguous with a zone or "natural row" of blackberry patch with tree saplings and shrubs the major ones of which were smooth sumac, the two grape speceis, black cherry, American elm with some climax hardwoods (more hickories than black or red oaks). Such strips of seral woody vegetation are almost impenetrable to humans as the blackberry picking photographer attested. This is bobwhite habitat par excellence. Even pointers experience some problems getting through such brushy barriers. Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak). |
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33. Sere colonizers, both pioneer newcomers and climax holdovers- Two close-up "photo-plots" of seral range vegetation on a climax black oak-dominated upland forest site in wesstern Ozark Highlands that had been clearcut two winters previously. Recovering forest vegetation in these photographs was about half-way through the second warm-growing season after logging. The plant community of this pioneer or initial seral stage was a combination of colonizing species (giant ragweed, common ragweed, mare's tail or horseweed, hairy crabgrass, and beefsteak plant) native perennial composite forbs (Baldwin ironweed and Carolina elephant's foot), biennial forbs (flannel mullein and biennial evening primrose), native shrubs (blackberry, smooth sumac, summer and fox grape, Virginia creeper), colonizing trees or generally less tolerant tree species (sycamore, black cherry, sassafras, American and red elm), and seedlings and stump srpouts of climax trees (black and red oaks, Ozark or Texas and mockernut hickories). An amazing array of plant species in small local habitats. Seral range vegetation shown in the first slide consisted of a weed patch dominated by giant ragweed with Canada wild lettuce, biennial evening primerose, and greeen and/or yellow foxtail mixed with woody species ranging from black oak and hickory seedlings and stump sprouts to summer grape that had developed in front of an edge of woody vegetation made up of taller shrubs and young tree saplings. Shrubs included blackberry, smooth sumac, fox and summer grapes while saplings ranged from black oaks and two hickory species to American and red elms, black cherry, and sassafras. This was another example of the edge where different plant communities meet producing the edge effect that is so important for certain species of wildlife. Northern bobwhite quail was a species of great local importance that benefits immensely from this kind of seral range vegetation.. The second slide was of seral forest range vegetation dominated by forbs and seedlings and stump sprouts mostly of black oak and the two hickory species. Conspicuous forbs included giant ragweed, horseweed or mare's tail (many of which had upper parts of shoots grazed off by white-tailed deer), Canada wild lettuce (also grazed by deer though less so than mare's tail), toothed spurge, and biennial evening primrose. Red clover was also present but less conspicuous in spite of its showy inflorescneces than the taller-growing "weedy" colonizers. Tha annual grasses, airy crabgrass and green or yellow foxtail, were the representatives of the Gramineae in this photograph There was considerable cover of young Virginia creeper. The tree trunk in center midground was a black oak. Sprouts surrounding this black oak were mostly Ozark and mockernut hickories. Also in this slide was a hollow butt portion (roughly one ana a half foot diameter) of a black oak log that was later used for fuelwood. Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak). |
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34. Forgiving oaks- Four examples of stump sprouting or coppicing in black oak on a black oak-northern red oak upland forest in the western Ozark Highlands (Springfield Plateau section) midway through the second growing season for hardwood species following clearcutting. Coppice was defined by the Society of American Foresters (Helms, 1998) as: "1. the production of new stems from the stump or roots 2. to cut the main stem (particularly of broadleaved species) at the base or to injure the roots to stimulate the production of new shoots for regneration 3. a plant derived by coppicing 4. any shoot arising from an adenvtitious or dormant bud near the base of a woody plant that has been cut back". All four meanings were appropriate for the morphological/physiological phenomenon presented in the four black oak stumps and their second-season suckers. Helms (1998) also provided: "stool- a living stump (capable of) producing sprouts". Coppice shoots or stools (and related forms of vegetative reproduction including water shoots, suckers, and sprouts) are, of course, clones, ramets, modules or modular units of the parent plant, genet, entire vegetative plant, respectively. A simplified explanation is that stump sprouts are natural grafts or naturally formed scions arising from the stock (the stump or root). Forest regeneration by coppicing is a major silvicultural system (or component treatment thereof) for certain hardwood species. Black oak has not generally or typically been viewed as a coppice species or black oak lumber as a coppice crop to the extent as for some other species (eg. black cherry), but on the clearcut described in the above captions a high proportion of the felled black oak did coppice (sprout back from living stumps). In fact, this is the usual regeneration response of black oak, at least of the younger trees. Stump sprouting in black oak was described in Silvics of Trees of the United States (Fowells, 1965, p. 560; Burns and Honkala, 1990, Vol. 2.p. 747). The latter reported that roughly 95% of black oak regeneration in clearcutting was by sprouts from either stumps or "advance reprodution" (new sprouts from dormant buds near juincture of shoot and root). An interesting inverse relationship exist between stump size and successful stump sprouting. Shoots (sprouts= stools) from black oaks that were larger at harves grow faster than those of smaller cut trees; however, stump sprouting was inversely related to stump size, tree age, and forest site with larger stumps and those of older trees (often "same difference") having reduced sprout perduction. In other words stumps of bigger trees (ie. larger diameter stumps) and, hence, usually older trees are less apt to sprout, but if they do sprout these suckers (stump shoots) grow faster (have more rapid grow rates) than suckers from smaller stumps. This is a common--if not nearly universal--physiological response in hardwood species. Black oak stump-sprouts less readily than norther red oak which, as on this oak-hickory forest, is commonly associated with black oak. Fowels (1965, p. 591) cited research that reported over two-thirds of black oak reproduction in the Missouri Ozarks was "of sprout origin". Nonetheless, coppicing is the best possible means to replace 1) "original" (genetically identical) trees and shrubs and 2) climax woody species. In this regard, sawing logs amounts to the same thing as mowing shoots of perennial forage species for hay. With properly harvest the species composition of the tree component of the clearcut forest is essentially the same as it was before logging. In the instance of the upland Ozark Highlands black oak-dominated forest that served as the example for this lesson there was a cruel irony to the clearcutting operation in which a theiving son stole timber from his trusting father. The criminal act of forest harvest actually benefitted the black oak-northern red oak forest. The felled trees were still of an age and/or size that a high proportion of them coppiced (stump-sprouted) resulting in regeneration of both 1) the climax tree species and 2) reproduction of the same genetic individuals (ie. restoration of the identical harvested trees). Obviously the shoots (trunks= boles) of the mature (actually, over-mature) trees will not be restored because they were hauled of to the sawmill for pallats and framing lumber, but genetically these identical trees will regrow (grow back) so as to be be replaced as they were before forest harvest. There will be the same original root systems and basal trunks, the stumps, but morphologically new trunks. Based on canopy cover the proportion of black and northern red oaks in the recovering (seral) forest was less than in the climax forest, but climax tree species (and the same individual trees) were in the seral forest vegetation from the initial (pioneer) stage of plant succession (forest restoration). With asexual reproduction (coppicing) climax trees were present in the initial plant community following clearcutting rather than entering the forest sere later in the sequence of secondary plant succession as would be necessary if climax tree establishment depended solely of sexual reproduction (production of seedlings). Net result will be (barring other or continued severe forest disturbances) a more rapid return to the terminal stage of plant succession (ie. a shorter time interval to replace the climax black oak-northern red oak forest). Ironically, if the near old-growth black and red oaks (many of them already "overripe" with trunks partly hollow with heart rot) had remained standing for more years most of them would have been too old to coppice. Thus regeneration of the climax oak species almost assuredly would have been much lower and slower. Sexual reproduction (seedlings from acorns) of black and red oak is much less likely (much lower probability of successful tree establishment) than is asexual reproduction by coppicing. Furthermore, recall (from photographs and descriptions of the unlogged, nearly old-growth forest) that most tree reproduction in this climax black oak-dominated forest was of the associates, Ozark hickory (a small tree at maturity) and, secondly, mockernut hickory. In this forest of mature oak trees and in absence of natural disturbances like windthrow (blowdown) and fire the climax oak species were, through a combination of natural death due to old age and lower rates of regeneration, being replaced by replaced by hickory species. Clearcutting this black oak-dominated forest resulted in 1) regeneration of the climax oaks along with that of the associate hickory species, 2) increased plant biodiversity due to a combination of regeneration of climax species and colonization by pioneer ("weedy") species, 3) increased forage and browse production for livestock and deer, 4) improved habitat for bobwhite quail, and 5) caused loss of habitat for plant and animal species dependant on climax forest vegetation. |
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35. Black oak forest in an Ozark spring- Interior view of dry chert upland forest dominated by black oak with post oak, red oak, blackjack oak, and black hickory as associates. There are two obvious shrub layers: 1) an upper one dominated by flowering dogwood (State Tree of Missouri; conspicuous here), redbud (State Tree of Oklahoma), and shadbush= eastern serviceberry (Amelanchier arborea) and 2) a lower one dominated by buckbrush= coral berry (Symphoricarpos orbiculatus) and blackberry. Wild grape and Virginia creeper (Parthenocissus quinquefolia) form an aboreal shrub layer while poison ivy (Rhus radicans = Toxicodendron radicans) occurs in both shrub layers ranging from lianas extending to tops of trees to non-climbing thickets.The herb layer is usually limited to early spring species that complete their annual cycle before greening of the forest canopy. Mayapple (Podophyllum peltatum) is the conspicuous dominant herb. Head of hollow on a chert upland. April, early vernal aspect. Ottawa County, Oklahoma. FRES No. 15 (Oak-Hickory Forest Ecosystem). Black oak and red oak form of K-91 (Oak-Hickory Forest). SAF 110 (Black Oak).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" ! |
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36. 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). |
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37. 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. |
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38. 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. |
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39. "In the Good Ole Summertime" but still "sick"- Sleek cattle and green leaves do not change the fact that this should-be or one-time forest is a degraded plant community (again, notheing but a "stomp lot"). What should be an oak-hickory forest with miscellaneous hardwoods such as black walnut, black cherry, and Kentucky coffeetree become a degraded pasture of mostly Eurasian annual grasses and a few mature trees of climax species from the previous forest stand. Cattle through overgrazing and overbrowsing prevented regeneration of the hardwood trees and also precluded light surface fires that would have killed these non-sprouting junipers and benefitted former climax understorey grasses. Yes, it is true that a climax or late seral stage oak-hickory forest like the stand that was about 150 yards down the road from this "mess" (and that was used as the control plot to present this lesson) would have little grazable/browsable understorey. A comparison of that late successional stage of forest vegetation with the "cow pasture" shown here would suggest to the neophyte that there is more "cow feed" on this degraded former forest. That is not true, not the case at all. The near-climax forest of the control plot is ready for logging. Following harvest of oak, hickory, and walnut logs, native grasses (big bluestem, Indiangrass, and purpletop are the main ones) and numerous shrubs (including blackberry, sumac, buckbrush, and wild plum) as well as regenerated hardwoods (mostly seedlings with some stump sprouts) will soon become re-established and provide range forage and browse for livestock and wildlife (notably white-tailed deer and bobwhite quail). Most importanlty from a forest perspective is the fact that the wood crop (hardwood logs are the agricultural commodity) is a source of revenue along with cattle (feeder calves and cull cows) and wildlife (either as recreational products or a sources of income from egress fees). Over the longterm, a properly managed oak-hickory forest will generate more revenue and produce more resources and commodities than this degraded barnyard with shade trees. Even the latter will eventually die to be replaced by juniper which will be a fire hazard by that time. Of course, agricultural producers are the world's greatest and most sustained optimists. In that spirit one can always hope that the eastern red cedar will escape crown fires and live long enough to grow into lumber that will fetch a fair price and that can be made into fragrant cedar chests to sell to tourists in flea markets. If this owner wanted nothing but pasture for these cows and calves the proper farming practice would be to plant this land to introduced (agronomic) pasture grasses like bermudagrass (Cynodon dactylodon) or tall fescue (Festuca arundinacea), both of which are well-adapted to these shallow rocky Ozark hills. The owner could then properly manage this tame pasture for economical production of beef cattle. Instead and as it was this landowner had nothing but "bragging rights" to running some cows and a lot less income than if he had wisely managed his forest, range, and livestock resources. This joker had not done justice to the revered title of "hillbilly" (just plain "hick" about covered it). Ottawa County, Oklahoma. June. |
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40. 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). |
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Northern Red Oak (Quercus rubra)
Forests
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Oak-hickory forest dominated by northern red oak, northern red oak cover type 55 (Eyre, 1980, ps. 43-44), tend to develop on some of the most mesic forest sites and, therefore, to be some of the most productive forests throughout theoak-hickory association. Unfortunately for these forests (and the foresters and rangemen who would manage them) many were converted into farmland and, later, unceasing concentrations of highways, locations for houses (in all price ranges), and the final consequences of unabated, never though-through, run-amuck suburban sprawl. From a more positive perspective some northern red oak forests remain and some of the degraded (by farming and improper logging, grazing, burning, etc.) tracts have recovered to considerable degree. Perhaps most impressive and inspiring have been efforts made by some states, local governments, and non-government organizations that resulted in preservation of remaining relicts of natural vegetation including northern red oak forests. Missouri has been one of the leaders in this marvelous development in preservation of its natural heritage. Nelson (1987, 2005) described many of these natural communities and listed various units of land and water set aside by Missouri and other cooperating organizations. These included various oak-hickory forests, woodlands, and savannahs in which northern red oak is a constitutent species. |
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41. Stand of young northern red oak- On a north slope of a hollow in the Springfield Plateau of the Ozark Highlands a second-growth of northern red oak was developing to the species composition and vegetational structure of a climax red oak-dominated forest. Sugar maple, also young trees, was the associate species. A good example of young sugar maple was the rightmost tree (small pole or large sapling). The major shrub was flowering dogwood which comprised most of the shrub layer although it was accompanied by its usual shrub associate, eastern redbud. Numerous shrub-sized or small tree-class sassafras were also part of this intermediate woody plant layer. A lower woody layer was made up of regenerating hardwood tree species includnig white oak as well as sugar maple and northern red oak. An herbaceous layer was also present. It was dominated by tick clover. Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005). |
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42. Recovery of northern red oak-A second growth forest dominated by young trees of northern red oak with sugar maple as the associate species on a mesic north slope in the western part of the Ozark Highlands (Springfield Plateau). A few older and larger trees of white oak were present locally in this tract (two larger trees at far right, the rear one of which was dead with exfoliating bark) and there was regeneration of white oak (seedling age class). Most regeneration of all younger age classes was northern red oak with sugar maple (eg. large saplings to front and left of the two large white oaks) being second in this regard. In other words it seemed obvious that white oak was seral to northern red oak and sugar maple. The major shrub was flowering dogwood which was growing all around the foremost northern red oak.in center foreground Tick clover dominated the sparsely populated but continuous herbaceous layer. Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005). |
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43. Restoration of northern red oak and sugar maple- Interior of a second-growth stand of northern red oak with sugar maple as associate species on a moist north slope in the Springfield Plateau portion of the Ozark Plateau (= Ozark Highlands). The shrub layer was dominated by flowering dogwood with eastern redbud and young sassafras saplings functioning as associate shrub species. Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005). |
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44. Tick clover in the understorey- Two species of tick clover or tick trefoil dominated the herbaceous layer of a second-growth northern red oak-dominated (sugar maple was associate) forest on a moist north slope in the Springfield Plateau of Ozark Hihglands. This photo-plot provided a close-up, detailed view of the herbaceous zone of the understorey of this forest that was recovering from logging several decades previously. One species of tick clover was was in full bloom and handily identifed as naked-flowered ticktrefoil (Desmodium nudiflorum) whereas the other species was obviously a late summer- or autumn-blooming species whose identity could only be guessed from its round leaves. The latter was perhaps smooth tick trefoil or beggar's lice (D. marilandicum) which is common on wooded slopes and uplands throughout the Ozark Highlands. Also present at herbaceous level was s seedling of summer grape and a larger seedling of flowering dogwood. Species composition of this forest range vegetation was that of (at least, closely approaching) the climax northern red oak and sugar maple associate forest. The Society of American Foresters description of the northern red oak cover type (Eyre, 1980, p. 44) explained that this forest range type was "probably subclimax", especially on north and east slopes. One species of Desmodium back) s Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005). |
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45. Seeing red in the western Ozarks- Stand of second-growth northern red oak with various age/size classes ranging from large seedling to small sapling to large pole. There was a dead sapling of white oak, but otherwise no evidence of this often associated climax species. Trees in background included a high proportion of black oak along with northern red oak. A shrub-dominated lower woody layer was dominated by flowering dogwood with some scattered individuals of eastern redbud and large seedings and small saplings of sassafras. The major herbaceous species was the papilionaceous legume, naked-flowered tick trefoil. Individuals of another species of tick trefoil or beggar's lice was present which could not be positively identified and may have been smooth tick trefoil. Ottawa County, Oklahoma. July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 55 (Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005). |
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White Oak (Quercus alba)
Forests
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| 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). |
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| 46. 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). |
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47. 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). |
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48. 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). |
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49. 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). |
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50. White oak-black oak-northern red oak forest- White oak clearly dominated this north slope of a western Ozark Plateau forest, but it was not cover or density domiance to the degree that white oak dominated the north slope shown in the preceding set of slides (Roaring River State Park, White River Hills Ecoregion). Development and species diversity of two prominent woody layers below canopy layer was greater and different from that of a more overwhelming dominance by white oak as described for the immediately preceding Ozark Plateau forest.. This north slope and forest hollow was substantially more mesic and had less light exposure than the preceding forest. Flowering dogwood was the dominant of the taller shrub layer. A lower and more diverse shrub layer included wild hydrangia, blackberry, poison oak, grape, Virginia creeper, and lowbush huckleberry. The herbaceous layer(s) was not as diverse as the white oak-dominated north slope forest described above, but the grass component (mostly downy, silky, or hairy wildrye) was much more productive and of far greater foliar cover on this mixed oak Ozark forest. Most common forb was pokeweed (which is more commonly a forest forb on locally disturbed areas). Flag Springs State Park, McDonald County, Missouri. June, late vernal aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Most plausible Society of American Foreters (Eyre, 1980) cover type was SAF 52 (White Oak-Black Oak-Northern Red Oak) and this was predominant cover type of this locality: white oak was not as much as obvious dominant and defining species as was case for SAF 53 (White Oak) which this closely resembled. Oak-Hickory Series of Brown et al. (1998). Dry-Mesic Chert Forest (Nelson 2005, ps. 125-130). Ozark Highlands- Elk River Hills Ecoregion, 39b (Chapman et al., 2002). |
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51. 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). |
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52. 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). |
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53. 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. |
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Northern Red Oak(Quercus rubra)-White
Oak (Q. alba) Forests
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Throughout large areas of the Ozark Plateau there are upland forests with various proportions of red, white, and black oak all of which are dominants (ie. forests of co- or multi-dominants). This is in contrast to other forest cover types in which any one of these three species is the sole dominant. Forest cover types of these three single dominants (SAF 53, SAF 55, SAF 110) were treated immediated above. In the following section the forest cover type with various combinations of these three dominants (White Oak-Black Oak-Northern Red Oak, SAF 52) was covered. In this author's observation, over much of the Ozark Highlands the more mesic white oak and northern red oak tend to be frequently associated with each other whereas the less mesic black oak tends to dominate the more marginal forest sites (stonier, more shallow soil; south slopes), and as the sole dominant tree--at least, the oak--species. All four of these cover types (SAF 52, SAF 53, SAF 55, SAF 110) frequently develop in close proximity with each other. There is commonly conterminous occurrence among the four forest types as recognized by the Society of American Foresters (Eyre, 1980). Making distinctions among these can be somewhat arbitrary. An example of this situation was given above in the White Oak Type (SAF 52) that tended to be an Ozark forest co-dominated by white oak and northern red oak. The current section presented a climax forest (in species composition though not at old-growth state) in which northern red oak and white oak were obviously co-dominant and black oak was not present, period. This example was on a north slope above a deep hollow. |
| 11455 and 11456. |
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Mixed oak Ozark forest-Local stands of white oak-northern red oak as a second-growth forest populated by young adult trees just "shy" of being prime logs for lumber or timber. There was regeneration of both of these tree species adequate to maintain them as the co-dominants of this dry-mesic, chert, upland forest (Nelson, 2010, p. 125-130) in the Springfield Plateau of the Ozark Highlands physiographic province. There was a limited understorey beneath the complete canopy closure of this climax (though not at old-growth state) forest. In addition to sexual regeneration of the two dominants there were seedlings and small saplings of sassafras, a seral (often, pioneer) tree species and pignut hickory (Carya cordiformis), a climax tree of more mesic habitats. Examples of sassafras seedlings and a large sapling of pignut hickory were visible in foreground of the second photograph. The major low shrub in these two stands was buckbrush or coralberry (Symporicarpus orbiculatus) as seen in immediate foreground of the first slide.For all practical purposes there was not an herbaceous layer in this vegetation, the heavy accumulation of leaf litter serving as a mulch that prevented emergence of grasses, grasslike plants, and forbs. (Herbaceous plants were present in some portions of this forest as can be seen in accompanying photographs.) This forest vegetation was growing on the upper perimeter of a hollow, a local feature of the landform form of some ancient mountains like the Ozarks. Hollow as applied to highly eroded mountains was defined in the caption following the next one (two captions from now). Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005). |
| 11457 and 11458. |
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Ozark mixed-oak forest nearing its prime- Textbook example of a white oak-northern red oak forest forest of the dry-mesic, chert upland form (Nelson, 2010, p. 125-130). The second of these two photographs was a closer-in view of vegetation in right midground of the first photograph. This was a second-growth forest of adult trees that had grown to about their maximum size before senescence and interior decay (trunk hollowing) set in. This mixed-oak forest had the species composition (and was approaching the structure and physiogonomy) of the climax stage for this forest site. Most of the adult trees in these two views had DBHs in excess of 18 to 20 inches with some of the larger northern red oak exceeding two foot. Even the largest trees had intact crowns with few broken limbs that could allow entry of decay causing organisms. There was sexual reproduction amble for maintenance of white and northern red oak as co-dominants of the climax forest. There were trees of these two species in various age classes. A few of the large saplings of both oak species had died, presumedly due to less tolerance to conditions of this climax forest (ie. competition for light, soil moisture, root space, or whatever variables). This phenomenon was represented by a small snag of northern red oak bearing a conspicuous broken bole in far-right of first slide and, more prominently, in the second slide (growing among several larger trees). There was, however, a root sprout or sucker of this particular tree at its base bearing the immense shade leaves typical of young oaks growing in the forest understorey. Shade leaves are leaves produced in microhabitats of shade (of varying degrees of darkness or shadows). Shade leaves are much larger yet thinner (reduced leaf thickness) so that they are able to utilize larger proportions of the light available to them. Also growing among the adult oak trees--right in the thick of them nonetheless--was a small pole of pingut hickory, one of the most tolerant and mesic habitat-adapted Carya species in this forest cover type. This young hickory was prominent in the second slide. There were numerous larger seedlings and saplings of red maple (Acer rubra) scattered throughout this forest. A red maple seedling was featured in the immediate center foreground of the first slide. The most conspicuous shrubs in these two "photoplots" was Virginia creeper (Parthoenocissus quinquefolia) and buckbrush or coralberry, bogth of which were visible in immediate right foreground of both slides. There were no herbaceous species, hence no herbaceous layer, in samples of forest vegetation seen in these two "photoplots". Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005). |
| 11459 and 11460. |
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On one edge of a hollow- A large white oak (left) and northern red oak (right) growing at the outermost edge of a hollow in the Springfield section of the Ozark Plateau in a dry-mesic, chert upland forest (Nelson, 2010, p. 125-130). The large seedling to small sapling growing "smack dab" between these two adult trees was red maple, a tree species that was just making its appearance in this second-growth climax (or approaching climax) forest. The leader with narrower serrate leaves at left margin was the shrub or small tree species, hop hornbeam (Ostrya virginiana) which is one of the most tolerant of all North American hardwoods. Hop hornbeam produces wood surpassed in hardness only by that of flowering dogwood (Kurz, 2004, p. 160). The lower shrub layer was dominated by seedlings of climax co-dominants, white and northern red oak, along with buckbrush or coralberry and Virginia creeper. The thick leaf mulch and dense forest canopy prevented development of an herbaceous layer in most parts of this forest.(A few exceptional local areas of herbaceous growth were presented below.) Uppermost outer edge of the hollow was at the right third or so in these two photographs. The term hollow is one which many readers may be either unfamilar with or have a misunderstanding of. In its Glossary of Geology (Gary et al., 1972) the American Geological Institute defined hollow in part as "a low tract of land surrounded by hills or mountains; a small, sheltered valley or basin, especially in a rugged area". OK, but more specifically the term hollow (often pronounced "holler") is used by hill folk in southeastern and southcentral North America to connote a deep, almost gourge-like (albeit smaller) depression in the land thereby giving the impression of a small or local valley nestled within mountains. In ancient mountains like the Ozark Plateau, the mountains of which have been eroded down to their roots, the hollows are actually deep indentions in the general land surface rather than being of the "typical elevation" with mountain peaks having high elevations. This is more of a perception than a definition, but it most accurately captures the ambiance of the landscape (and hillbillies like this author). Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005). |
| 11461. |
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Approaching another edge- Straight-on view on upper "lip"or "brim", (the edge) of the hollow that was introduced in the immediately preceding two-slide set. In the western Ozark Mountains (Springfield scetion of Ozark Plateau; Springfiield Plateau) a climax mixed oak-hickory forest (dominated by white oak and northern red oak) of the dry-mesic, chert upland form (Nelson, 2010, p. 125-130) had developed around and in the interior of a deep and relatively large hollow. The large tree in left foreground and the tree behind and closest to right of it were white oak. The tree in center midground and the larger tree at right midground with a lower leaning limb off right side of its trunk were northern red oak. The small pole-sized (or sapling-size) tree to the left and slightly in front of the lower, right-leaning limb was northern red oak.There were widely scattered shrubs and shrub-like oak saplings at mid-height throughout this forest range. One of these (ascending trunk in right front of the leftmost and largest white oak) was a hop hornbeam. There were several red maple of shrub-size to the right of the large leftmost white oak. The lowest layer of this forest plant community was also made up of hardwood trees--as seedlings of both white and northern red oaks--but probably even more so of red maple seedlings as, for example, in lower right corner. Buckbrush or coralberry and Virginia creeper were common (though with sparse density and cover) lower-growing shrubs. The grape vine on the northern red oak could not identified to species. Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005). |
| 11462 and 11463. |
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On the cusp of an Ozark hollow- Details of some of the forest vegetation introduced in the preceding slide were presented in these two slides that were taken a few steps off the rim and down into the hollow. The largest and foremost tree with a lower, ascending limb off right side of its trunk served as a "landmark" in reference to this immediately preceding photograph. This (and all the large trees, young to middle-aged adults) were northern red oak. The shrub at far left margin (and with the trunk of a northern red oak serving as a backdrop in the second slide) was hop hornbeam, one of the most tolerant of all forest shrubs in North America. The low shrublike, sapling in center midground (to right of large, lower leaning-limb northern red oak) and several others behind it were all red maple. Sapling in right foreground in the first of these two slides was white oak. There were some adult white oak in background. All age classes of both climax Quercus species were represented although some saplings of these oaks had died (for whatever reasons; perhaps from competition). Ground-level, the lowest layer of vegetation, was comprised of seedlings of both of these oak species plus some Virginia creeper, and buckbrush or coralberry as well as a few plants of Christmas fern (Polystichum acrostichoides) and some planrts of a Carex species (unidentifiable in its strictly vegetative state). Thus, in restricted microhabitats there was something of an herbaceous layer although it was sporadic. Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005). |
| 11464 and 11465.. |
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Growing on a cusp- Composition, structure, and partial physiogonomic perspective of an upland, climax white oak-northern red oak forest that developed at the head of an Ozark hollow. This second-growth forest was approaching the ultimte structure and physiogonomy of an old-growth forest (the state of a virgin climax forest). The largest trees were roughly middle-aged adults that in a few decades would be approaching their maximum size. These trees were thus nearing their "prime" from standpoints of quality lumber or timber products and still-rapid rates of growth. Species or botanical composition was clearly that of the climax forest vegetation. Old-growth structure had not been reached in this stand. The three foremost (center foreground) trees in the first slide were northern red oak, the center and smallest (foremost tree) of which had died recently as evidenced by peeling bark at base of trunk and still attached dead leaves on lower branches.The adult tree in midground of both slides was a white oak. In the second photograph this white oak was positioned to the left of the big northern red oak in right midground. Three small saplings and one considerably larger sapling to the close-left of this white oak were red maple as was the large sapling situated between the two northern red oaks. The conspicuous (in both of these slides) liana that had "clum to the top" of the left adult northern red oak was some grape that could not be identified from the lowest leaves which were at least 20 foot off the ground. This plant was likely summer grape (Vitis aestivalis). Most of the lower or ground layer of forest vegetation was seedlings of white oak and northern red oak. Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005). |
| 11466. |
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On around the "holler"- Another view of the large northern red oak (left margin) and white oak (as viewed here it positioned to immediate right of the northern red oak) and "attendant" plants seen in right midground of the immediately preceding slide. One more northern red oak and white oak were in the background (farther down in the hollow). The sapling in right foreground and another (and smaller) sapling at close-right to the foremost white oak were red maples. Most of the plant growth in the lowest--the ground-level--layer of vegetation was made up of seedlings of the two oak species. There were even some seedlings of grape (unidentified Vitis species). Most of the ground layer was Virginia creeper. Buckbrush or coralberry was the next most common shrub. One plant of Christmas fern was to to close-right of the foremost northern red oak. Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005). |
| 11467 and 11468. |
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Looking down deep- Head of a hollow in the Springfield Plateau section of Ozark Mountains on which a climax white oak-northern red oak forest had developed. These two more detailed ("closer-up") photographs presented the lower layers (understorey) of this forest range so as to show what few herbaceous species grew in the surface (ground-level) layer of vegetation. These two "photoplots" were of the same hollow shown previously except that in this local area there were a few plants of some caric sedge (Carex sp.), which in the strictly vegetative state could not be identified, and also of Christmas fern. A conspicuous plant of Christmas fern was featured prominently in center foreground of second slide. There were also several Christmas fern plants along with some of the caric sedge close to the trunk of the trunk of the northern red oak in the second slide. The photographer could not find any grass in this dry-mesic, chert, upland forest (Nelson, 2010, p. 125-130). The high tree density resulted in what, for all intents and purposes, was complete canopy closure (ie. 100% cover by tree crowns). The two largest trunks (the only one in second slide) were northern red oak. The tree in center midground of both slides (and to left-rear of the big northern red oak in second slide) was white oak. Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005). |
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General note: students should take note of (and make a mental note as to) general density or "spacing" of trees growing on the upper edge of the Ozark Plateau hollow featured above. Density is the number of individual organisms present in a specific area of space (land). An application of density in Forestry is the concept of stocking. In silvicultural usage stocking refers to occupancy of forest growing space relative to some preestablished standard. A closely related term used in Forestry is stand density which is either 1) a quantiative measure of stocking shown as number of trees, basal area, or volume of trees per unit of land area or 2) a measure of the degree of tree crowding in forests and shown as some unit relative to growing space (Helms, 1998). The area of land per tree or relative proportion of sky (or canopy) occupied per tree was comparatively large in the white oak-northern red oak forest shown and described immediately above. Trees were not crowded. Instead they were of such density or stocking ("spacing") that crowns of individual trees were large yet without producing lower limbs in the fashion of shade trees. These trees grew as unbranched single trunks that were producing good sawlogs yet with abundant canopy so that they had relatively rapid growth. With complete canopy closure the lower forest layers were limited to plants that have relatively high levels of tolerance. This included species rated as Tolerant to Very Tolerant (hop hornbeam, red maple, climax oak species, Virginia creeper, and among herbaceous species, Christmas fern). In most soils of the Ozark Plateau rapid tree growth and production of large trees having high-quality wood requires low stocking or, in laymen's terms, wide spacing of individual trees. |
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American Beech (Fagus
grandiflora)-White Oak-Hickory
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Presented in this section was an example of a post-climax forest community defined/distinguished by presence of American beech (Fagus grandiflora) as a dominant species within portions of the oak-hickory association. This forest range vegetation developed on atypically mesic, yet well-drained land in the Ozark Plateau. American beech was either the major dominant or co-dominant species along with white oak, pignut or bitternut hickolry (Carya cordiformis) and/or shagbark hickory (C. ovata ). The habitat for this beech-oak-hickory forest in the Ozarks Region is generally restricted to deeper drainages such corridors along tributaries to rivers such as the Buffalo and White Rivers of northern Arkansas. This deciduous forest range community is the potential natural vegetation that could be interpreted as a variant of the White Oak forest cover type (SAF 53). Alternatively, this forest veagetation could be regarded as an "island" or isolated southern extension, and in variant form, of the Beech-Sugar Maple forest cover type (SAF 60). Sugar maple was abundant in the example forest described herein, but oaks, including northern red and white oaks, and hickories were far more abundant and common than sugar maple. This was consistent with the Society of American Foresters description of a variant form of Beech-Sugar Maple (SAF 60) in which this cover type "grades into numerous related types according to degree of disturbance and changes in site". In the Midwest and southern Appalachians this forest vegetation "is commonly associated with uplant oaks" such as white oak, black oak, and northern red oak (Eyre, 1980, p. 34). The forest portrayed below had developed on a soil (Arkana-Moka complex, 20-40% slopes) of generally less than a yard in depth overlying level-bedded limestone. Surface soil layers were generally very cherty and very stoney silt loams that overlaid clay to clay loam subsurface material. Soil was moderate in fertility and neutral to slightly acid or slightly alkaline in reaction (Soil Conservation Service, 1988, ps. 15-16). This forest had been logged--and it was high-grade logging at that--approximately 40-42 years prior to time of photographing and descriping it. Fire scars--some of them very deep and large--on almost all of the larger (hence, presumedly older) beech trees attested to the impact of hot fires on the present forest. Such fires selectively damages smooth-barked beech and sugar maple more than oak and hickory species. This selective defoliation was in all likelihood a major factor that contributed to relatively more oak and hickory and less beech that might otherwise have existed. It was also likely that secondary plant succession was on-going with the present state of forest community development at an advanced seral stage yet still below the ultimate state of development (eg. present forest vegetation was subclimax or preclimax). |
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This beech-white oak forest developed on mesic lowland along Clark Creek which forms a gouge-like tributary to Buffalo River. This was not a floodplain forest nor, strictly speaking, was it a bottomland forest. Neither was this forest range community riparian vegetation. In drier times of the year there is little water in the channel of Clark Creek because the water goes belowground through a fissure. The moist and fertile (though boulder-strewn) land of this drainage produced this unique forest that also included sweetgum, pignut (=bitternut) and mockernut hickories, basswood, sycamore, red mulberry, black walnut, red maple (and less sugar maple), northern red oak, sassafras, and both cucumber tree (Magnolia acuminata) and umbrella magnolia (M. tripetala). Shrubs included redbud, dogwood, spicebush, wild hydrangea, pawpaw, Carolina buckthorn (Rhamnus caroliniana), and American bladdernut. Remarkably little of Rubus species. Forbs included wild ginger, Solomon's seal, shooting star, great lobelia or blue cardinal flower, round-lobed liverleaf (Hepatica nobilis), pokeweed, trout lily, tall anemone (Anemone virginiana), jack-in-the-pulpit, green dragon, yellow-flower leaf-cup or bearsfoot (Polymnia uvedalia), columbine, bloodroot, trillium, sand phlox (Phlox bifida), and several kinds of ferns including Christmans, maidenhair, and broad beech fern (Thelypteris hexagonoptera= Dryopteris hexagonoptera= Phegopteris hexagonoptera). Main grass overall was Canada or hairy, wood brome (Bromus purgans= B. pubescens) along with bottlebrush (Hystrix patula). Also present but less common generally was broadleaf woodoats (Uniola latifolia) and Virginia wildrye (Elymus virginicus). Braun (1950, ps. 33-34) clarified that a climax association that characterizes a forest region can also develop in other forest regions (which are characterized by other forest associations).In the Clementsian model used by Braun (1950, ps. 10-13) this beech-white oak-hickory forest could be interpreted as an example of a variant of the climax beech-maple association (that characterized the beech-maple forest region), but which occurred in the oak-hickory forest region or perhaps of the "transition from the oak-hickory to the oak-pine region" (Braun, 1950, p. 278). Perhaps most precisely by Braun (1950, ps. 170 and 172) interpretation this forest vegetation was an isolated community (an ecological island so to speak) of the Mixed Mesophytic Region or, at least, of the "typically mixed mesophytic aspect". In fact, it seemed likely that Braun described this exact tract of forest (which was in Lost Valley of Buffalo National River--previously Lost Valley State Park--near Ponca, Arkansas) decades earlier (Braun, 1950, p. 172). This conclusion could not be stated with 100% certainity, but the evidence was very strong that such was the case. Interpretation of this forest vegetation based on designations and descriptions of forest cover types by the Society of American Foresters (Eyre, 1980) was explained above. This forest dominated by American beech-white oak (pignut hickory should probably be added for some portions of this climax vegetation) was a mesic lowland forest that had developed along a drainage that furnished a habitat substantially more moderate and more favorable for mesic species than that of adjoining environments. This should not, however, be confused with either 1) a bottomland or floodplain forest or 2) riparian vegetation. Clark Creek was not a perennial stream and it did not sustain moist banks for extended periods of time. Rather it was an intermittent stream the flow of which seeps through fissures and moves underground for much of the growing season. Flooding is also of such short and infrequent (even if violent) occurrence that vegetation along stream banks does not differ in meaningful composition or structure from that distant from the stream. In fact it is probably because this was not a floodplain that enabled American beech to grow here. It is textbook knowledge that beech (one of the most tolerant of North American hardwoods trees) does not thrive on either drier soils or those that are poorly drained (Harlow et al., 1979, 284-285). Beech has been found to be one of the most sensitive species to flooding (Burns and Honkala, 1990, p. 329). |
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Technical note: the photographs presented below were taken in early summer (mid-July) from mid-day to late afternoon (approximately 1230 to 1630 hours Central Standard Time). There was not so much as a wisp of a cloud in the entire sky. Thus shade and shadows represeted the minimum degree of darkness and the maximum quantity of light present in this subclimax to near-climax deciduous forestl. Neither conditions of direct overhead sun nor those of oblique angle of sun (again, both with 100% clear sky) resulted in optimum lighting conditions for photography. It bore repeating, however, that photographs were representative of the maximum light that could penetrate canopy (crown cover). Said another way, photograhic portrayl accurately represented conditions of maximum light intensity available to the understorey under full summer forest canopy. The phenomenon of sunflecks was shown to ideal advantage under these light conditions. Low degree of cloud cover will sometimes diffuse light so as to produce uniforn distribution of light conditions (intensity, quality, etc.) which can reduce or eliminate shade. This, however, also reduces brightness (intensity) of available light necessitating lower f-stops and/or slower shutter speed which in turn results in less depth-of-field (hence less detail) of subject matter, especially of plant parts. Under conditions where cloud cover results in diffusion and more uniform light distribution and the consequence of reducing or eliminating shade there is also reduction or elimination of spaces that have full-sun (maximum birghtness). With such conditions of uniform distribution and reduced intensity of light in forests there is little or no opportunity to present examples of the sunflect phenomenon that is essential for photosynthesis below the forest crown canopy and consequent development of understorey layers, the grazable/browsable zone of forest range. It is textbook knowledge as well as folk wisdom that when forests dominated by American beech are in in full-leaf they are still some of--if not the--darkest of all forests in North America. In this photographers' experience only north-slope forests dominated by the large-leafed bur oak have darker (dimmer) light conditions in the understorey. Slides of leaves and canopy of American beech and forest range communities dominated by this species portrayed the basis of overall dim light and sunfleck patterns characteristic of beech-dominated deciduous forests in the Ozark Plateau. Viewers should bear in mind that low visibility situations seen in most of the photographs presented below were representative of typical light conditions (ie. what they would encounter if they were present) in the dark, often dank, forest dominated by American beech. |
| Treatment of the American beech-white oak-hickory mesic forest began with overall or general external views of the forest and progressed toward the forest interior and understorey ending with photographs which presented details of important range plants and range microsites. Two different locations and distinct forest sites were presented. Both of these forest communities were distinguished by having American beech as a dominant along with oak and hickory species. The first forest tract featured was a forest community dominated by American beech on a north to northeast slope above Boxley Valley (Newton County, Arkansas). The second tract of forest was larger and was a much more botanically and structurally diverse forest community that had developed along the drainage of Clark Creek known as Lost Valley (Newton County, Arkansas). These forest range communities were in the Boston Mountains portion or section of the Ozark Plateaus physiographic province. The Boston Mountains exist as a dissected plateau that was peneplaned earlier than the rest of the province so that this is the oldest part of the Ozark Plateaus (Fenneman, 1938, ps. 655-660). Soils of the Boston Mountains formed from various parent materials with hill crests most commonly being of sandstone while lower strata are generally of chert/limestone. |
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54. Trees over all the hills and in the "hollers"- Exterior view in Ozark Plateau-Boston Mountains of oak-hickory forest with American beech, sugar and red maples, sycamore, black locust (Robinia pseudoacadia), shortleaf pine (Pinus echinata), and sweet gum (Liquidambar styraciflua) as species that were local associates to white and northern red oaks, the general dominants. This general perspective presented the hills and hollows topography typical of the deeply dissected plateaus of the Ozark Plateau Region. July. Buffalo National River, Newton County, Arkansas. |
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55. About as dense and dark as it gets- High noon in early summer in the interior of an oak-hickory forest in which American beech and sugar maple are the major associate species. In the first of these three photographs pignut hickory dominated a stand of mature trees on a north slope. The second photograph featured a second-growth stand dominated by American beech with pignut hickory as associate species on a northeast slope. The third slide showed dominance of beech in this stand (pignut hickory represented by one tree in right background). The second and third slides presented the very liited understorey which was comprised mostly of regenerating beech. This demonstrated the Very Tolerant tolerance rating of this species. In fact, presence of several age classes of beech attested to this feature. Extent of shade on the forest floor at mid-day (1230 Central Standard Time) in early summer portrayed (clearly though darkly) the phenomenon of sunflecks and and the resultant extremely sparse layer of range vegetation on this shaded forest floor. July. Buffalo National River, Newton County, Arkansas. |
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56. American beech, and nothin' else- A northeast slope above floodplain of the Buffalo River was naturally stocked by the single species, American beech. This was an example of a forest consociation. In the first of these two slides there was only one representative of another tree species. Can you find it? Hint: it is a pignut hickory. Note the different age classes of beech present in these two "photo-plolts". Viewers should also take not of relative area of the forest floor as to: 1) proportion of shade and 2) absence of understorey species other than beech. These photographs showed the maximum surface area of the ground receiving sunlight in early summer because pictures were taken at mid-day (1230-1245 hours Central Standard Time) when shadows cast by trees was at a minimum. The portion of shaded vs. sunlite ground surface also presented the concept of constanting changing patches of light caused by changing position of Earth relative to its sun, cloudiness (not shown on this cloudless day), and wind-caused leaf movement. The scant plant cover and density at lower layers of this forest vegetation was largely due to the dense shade cast by beech (the specific epithet, grandifolia, was chosen most appropriately). It was likely however, that grazing/browsing by livestock in the not-too-distant past was also a factor in lack of understorey development. This steep hillside was now part of Buffalo National River, but it had previously privately owned land that was located immediately above an old barn (of sorts) and "cowpens". Any halfway knowledgable forest or range "detective" would have to assume that even a slope of this steepness had received relatively heavy stocking of cattle. Also, this area was some of the last legal open range (land open to the public to run livestock on and therefore subjected to free-roaming livestock) which included free-ranging, privately owned hogs. Swine are potentially the single most destructive species of range livestock on deciduous forest such as oak-hickory and oak-pine cover types. There was no denying the high probability that cattle and hogs had likely impacted this tract of forest in recent history of, say, 15-25 years ago. Nonetheless, much of the absence of understorey development was due to conditions of exteme shade. July. Buffalo National River, Newton County, Arkansas. |
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57. Sign post (or "fools names in public places")- The trunk of an old American beech bore names and other graffiti left by vulgar, clueless scribes (probably urban and rural) who could not resist the temptation to use beech's characteristically smooth bark as sylvan space on which to "leave their mark" The mark left by literal sons of bitches (Canis familaris, C. latrans) quickly dries and, if not too frequently left, leaves little impact on the face of Mother Nature. By contrast, the heavy hand of vain-glorious, ego-driven, illbred, human SOBs bears a longer-lasting imprint of visitors to public property who violate the first rule of the woods, Leave a Clean Camp. Besides, unlike Daniel Boone, these self-centered wood carvers kilt no b'ars under thishere beech. Any claw marks left by the once native black bear (Ursus americanus) would by contrast be most welcome in this designated wilderness administered by the National Park Service. Desecrationof public property aside, the identifying smooth gray bark and dense shade cast by American beech was presented in this Kodak moment on a steep north slope dominated by beech in the Ozark Plateau. Another feature of American beech is root sprouting, development of secondary shoots from roots of mature and even uninjured beech trees. Most of the beech shoots in foreground of this photograph arose as asexual (vegetative) reproduction rather than as sexual reproduction from this tree, the bark of which that bore inscriptions from self-appointed, wood-carver scribes. More examples of root suckering (sprouting) were presented farther below. July. Buffalo National River, Newton County, Arkansas. |
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58. A steep, beech-shaded slope- This steep, north slope was dominated by American beech underwhich shade-adapted plant species comprised some layers of forest vegetation. Root sprouts or secondary shoots (introduced in the preceding slide) and beech seedlings made beech the most abundant plant species at various micro-locations, but there were a few other understorey species including poison ivy, Virginia creeper, Christmas fern (Polystichum acrostichoides), and rolled sedge (Carex convoluta). The mature tree and two smaller trees closest to it on the right were beech as were most of the seedlings and small saplings, almost all of the latter of which were root sprouts of the adult tree. July. Buffalo National River, Newton County, Arkansas. |
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59. Hard to see but instructive- Trunk of the largest American beech shown in the preceding photograph was shown in more detail in these two slides. One distinctive feature of beech--especially older, larger trees--is the self-exposed roots which in normal growth commonly protrude above the ground surface.This growth characteristic results in a fluted basal tree trunk. Also presented in these photographs were the lower layers of forest vegetation--sparse though they were--that developed in the dense shade of a stand of beech. Viewers were again reminded that this photograph was taken at noon time (1230-1245 hours Central Standard Time) when light penetration to the forest floor was theoretically greatest overall (on average). Some plants of rolled caric sedge were visible in the second slide. The phenomenon of sunflecks was presented here,, but shown to better advantage in the next slide. Another phenomenon displayed in the first of these two slides was that of geotropism, the directional movement (and subsequent growth) of a plant to gravity. This old beech was leaning so as to remain upright against the pull of gravity as exerted on this steep hillside (ie. a plant response to the stimulus of gravitational pull). There was also greater accretion of wood in the trunk on its uphill side in order to balance the entire shoot (bole and crown) of this beech which was large by standards of the Ozark Plateau, a marginal habitat for one of the most mesic of North American hardwood (angiosperm) tree species. July. Buffalo National River, Newton County, Arkansas. |
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60. Light-grabbin' grasslike plant- A specimen of rolled sedge (Carex convoluta) at base of American beech on a shaded, steep north slope. This plant was growing close to the base of trunk and exposed roots of the American beech presented in the two photographs immediately above.This species is obviously one of the best adapted of sciophytes (= skiophytes; "shade-loving" plants). July. Buffalo National River, Newton County, Arkansas. |
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61. Details of a shade-lover- Sexual shoots with fruit of rolled caric sedge. These shoots were on the same plant shown in the previous photograph. It was one of the specimens growing at base of the large and leaning American beech featured above. July. Buffalo National River, Newton County, Arkansas. |
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62. Mixed mesophytic forest in Ozark Plateaus Region (Boston Mountains)- General and exterior view of the "mixed mesophytic aspect" (Braun, 1950, p. 170) of the oak-hickory association. It was explained in the introduction to forest range vegetation featured in this section that it was one of the more mesic (and restricted) forms of oak-hickory forest that develops in the Ozark Mountains. Dominance of American beech along with the typical dominant white and northern red oaks plus pignut (=bitternut) hickory in association with sweet gum, sugar maple plus occasional sycamore along with two Magnolia species as smaller (subcanopy) though defining trees described the specieas composition of this unusual climax forest that extended from bottomland to sloping uplands. This stream-drainage forest was in a deep hollow that eminated from tall sandstone bluffs (complete with waterfall and small cave) to terminate on the floodplain of the Buffalo River. The sandstone bluffs are visible in right background of this photograph. The striking white bark (left midground) was that of a beech. Other dominant tree species were white oak, northern red oak, and pignut hickory. Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Mixed mesophytic form of the oak-hickory forest with American beech as defining/distinguishing species and in association with sugar and red maple designated this as a isolated unit of a variant of SAF 60 (Beech-Sugar Maple) rather than as a variant of SAF 52 (White Oak-Black Oak-Northern Red Oak). Likewise, isolated unit of Beech-Maple Series 122.13, Northeastern Deciduous Forest 122.1 of Brown et al., 1998, p. 37). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004). |
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63. Pristine Arkansas- Mixed mesophytic form (Braun, 1950, p. 170-172) of oak-hickory forest. American beech as a local dominant along with sugar maple, sweet gum, and Magnolia species joined the area dominants, white oak, northern red oak, and pignut hickory, to produce a variant of Ozark Plateau deciduous forest that was interpreted as an isolated unit of the beech-maple association. The largest tree in these two slides was a white oak. The next largest tree and with large shade leaves (to left and rear of big white oak) was sugar maple. Most of the canopy in these photographs was American beech. There were small caves or cavelike recessions sunk into the sandstone bluffs behind trees. This forest community developed within the hollow created by the drainage of Clark Creek. Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Mixed mesophytic form of the oak-hickory forest with American beech as defining/distinguishing species and in association with sugar and red maple designated this as a isolated unit of a variant of SAF 60 (Beech-Sugar Maple) rather than as a variant of SAF 52 (White Oak-Black Oak-Northern Red Oak). Likewise, isolated unit of Beech-Maple Series 122.13, Northeastern Deciduous Forest 122.1 of Brown et al., 1998, p. 37). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004). |
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64. Boston Mountains beech- Local stand of American beech with sweet gum as the associate in the overall oak-hickory forest of the Ozark Plateau region. Although white and northern red oak along with pignut hickory were area dominants none of these species was represented in this local stand that had developed along sides of Clark Creek. In fact all regeneration (of which there was an abundance) was beech. This was both sexual reproduction from beech nuts and secondary shoots (suckers) from roots of established trees. Root suckering in beech was discussed below when forest gaps and patch dynamics were covered. Sweet gum is a seral species and its presence in this Clark Creek forest was due to persistence of trees that established about 40 years earlier following clearcutting of this forest tract. Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. |
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65. Things got hot (but they're cool now)- The personality of this beech attested to hot fire(s) in the past. This scar extended over eight feet up the bole of this second-growth specimen and penetrated through to the heartwood. Most trees survive this invasion (at least initially) unless such injury allows entry of pathogens that may later kill the tree, directly or indirectly. Heartwood and much of the sapwood is dead tissue anyway so that loss of such wood has no physiological impact on the plant. Instead the tree is weakened by loss of some support provided by the trunk. This frequently results in toppling of the tree when it is subjected to high winds: blowdown when the trunk is snapped, or, even, twisted off. Examples of such blowdown or windthrow (and the consequent forest gaps) were presented later in this section. For now this beeech was still a "member in good standing" of this forest community. Much of the beech reproduction (which made up most of the regeneration shown here) was asexual in form of root sprouting (suckering). Throughout this section emphasis was frequently placed on the phenomenon of secondary shoot production by root sprouting in American beech. Some of the smaller trees presented here were seedlings showing that both sexual and asexual reproduction was successful in this forest. This was in spite--or perhaps because--of fire which may have initiated beech suckering. Fire(s) that burnt this beech and its neighbors undoubtedly had influences on neighboring plants, for good and bad depending on species. Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. |
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66. Memories of hotter times- An American beech sports a long and deep fire scar that bears testimony of a once-upon-a-hot-time in these Boston Mountains woods. Like the example presented in the preceding photograph, the fire scar of this second beech had burnt through to the heartwood along entire height of the pyric injury. In fact, in both of these cases flames inside the trunk actually burned higher up inside the bole than on the outside. Also, these trees were healing around the injured area so that the scar was simultaneously "growing" as the tree grew in heighrt and girth and also healing over as replacement growth enveloped the injury site. The thin bark and big, protruding roots of beech make it one of the most susceptible hardwoods to fire (Fowells, 1965, p. 176; Burns and Honkala, 1990, p. 329). Thin bark also makes beech prone to sunscald, especially when trees in dense forest (hence protected from damage from sun's rays) are suddently exposed to brighter or even full-sun following logging. This species is also vulnerable to mechanical injury from logging and prunibng as well as damage by insects with piercing-sucking mouthparts (bugs) and diseases in general.(Burns and Honkala, 1990, p. 329). As demonstrated by this and the preceding example some individuals of American beech can live with extensive fire injury as long as additional strrains from wind and gravity and stresses do not exceed physical strength and disease pathogens cannot gain entry through a compromised physical barrier of bark and associated tissues. It was not known if fire was a factor involved in root suckering. Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. |
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67. Innards of a botanical "island"- Composition and structure of an Ozark forest dominated by second-growth American beech. Isolated forest communities of beech-sugar maple occassionally develop in more favorable habitats of the regional oak-hickory forest (ie. isolated communities of the beech-sugar maple association within the surrounding oak-hickory association).Forest vegetation in these two photographs represented the interior of an American beech-dominated variant of the white oak-northern red oak-pignut hickory community of more mesic environments such as this one in the Boston Mountains of the Ozark Plateau region. The two photographs were horizontal and vertical views of the same stand to better represent species make-up and forest structure. While beech requires more mesic forest sites than its co-dominants in this forest (white oak, northern red oak, pignut hickory) it also require better drained soils. Beech is one of the most flood intolerant species of all North American trees, especially flooding durnig the growing season (Burns and honkala, 1990, p. 329). Almost all of tree regeneration in this stand was beech and not that of white oak, the major co-dominant, or sweet gum, a locally abundant associate that is a seral species sometimes persisting into advanced successional stages. This was also the case for some of the other stands presented below. This forest understorey was comprised of both herbaceous and woody species, including beech seedlings and root sprouts as discussed above. The most abundant, though certainly not of great density or foliar cover, herbaceous species overall was hairy wood brome. In locally miacrosites the herb with most cover was broad beech fern. mmon. Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Mixed mesophytic form of the oak-hickory forest with American beech as defining/distinguishing species and in association with sugar and red maple designated this as a isolated unit of a variant of SAF 60 (Beech-Sugar Maple) rather than as a variant of SAF 52 (White Oak-Black Oak-Northern Red Oak). Likewise, isolated unit of Beech-Maple Series 122.13, Northeastern Deciduous Forest 122.1 of Brown et al., 1998, p. 37). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004). |
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68. Renewable resources- That forests are some of man's most readily renewed natural resources was showed in this scene of a 40 to 45 year-old second growth forest of white oak and American beech that had regenerated along Clark Creek following high-grade logging less than a half century earlier. This was also a good example of the species composition and community structure of this forest. Tree species in the foreground were (left to right): beech, white oak( middle and largest trunk), and sweet gum. Most tree reproduction was beech, and of several age classes, the same as the situation presented in the preceding forest stand. Shrubs included redbud, pawpaw, spicebush, flowering dogwood, and American bladdernut. Herbaceous species ranged from hairy wood brome to broad beech fern. Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Mixed mesophytic form of the oak-hickory forest with American beech as defining/distinguishing species and in association with sugar and red maple designated this as a isolated unit of a variant of SAF 60 (Beech-Sugar Maple) rather than as a variant of SAF 52 (White Oak-Black Oak-Northern Red Oak). Likewise, isolated unit of Beech-Maple Series 122.13, Northeastern Deciduous Forest 122.1 of Brown et al., 1998, p. 37). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004). |
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69. A unique blend in Boston Mountains- A western mesophytic form of the oak-hickory forest association developed in a deep, narrow hollow that formed around Clark Creek which drained into Buffalo River. The forest stand shown here was on the terrace of this intermittant stream. The four young trees featured in the foreground were (left to right): sycamore beech, beech, and white oak. Beech dominated smaller/younger age classes as well as that of the largest trees (the canopy layer of this second-growth forest). Undrstorey species were primarily spicebush (a shrub) and pokeweed or pokeberry (rank-growing perennial forb). This was another two-slide set presenting horizontal and vertical perspectives to present a greater sample of the understorey and structure of the forest stand up into the canopy. The herbaceous species with most foliar cover was pokeweed or pokeberry. There was some local spot dominance of the irregular herbaceous layer by broad beech fern. Most abundant shrub was spicebush. Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Mixed mesophytic form of the oak-hickory forest with American beech as defining/distinguishing species and in association with sugar and red maple designated this as a isolated unit of a variant of SAF 60 (Beech-Sugar Maple) rather than as a variant of SAF 52 (White Oak-Black Oak-Northern Red Oak). Likewise, isolated unit of Beech-Maple Series 122.13, Northeastern Deciduous Forest 122.1 of Brown et al., 1998, p. 37). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004). |
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70. "Would you believe that this is Arkansas?"- "Shore as shootin'" this north slope along the deep drainage (a small hollow) of Clark Creek in the Boston Mountains supported a beech-dominated mixed mesophytic aspect (Braun, 1950, p. 170) of oak-hickory forest. In fact, more precisely this was an isolated representative community of the beech-maple forest association that developed within the overall oak-hickory association. However, sugar maple was quite limited whereas this was not the case for beech which dominated overwhelmingly the tree seeding and small sapling layer of (tree regeneration in) this forest community. Likewise, American beech dominated the layer of largest trees which were second-growth individuals less than 50 years old, this location having clearcut about 40 to 42 (maybe 45) years previously. In this mixed mesophytic aspect forest white oak was the close second co-dominant species to beech. Ultimate triumph of beech as the ultimate climax dominant tree of this forest sere was already evident at this relatively short time following severe forest disturbance. Sweet gum is a seral species on this forest site that persist into climax or, at least, advanced seral stages. A representative individual of sweet gum was present in midground of this photo-quadrant (behind and to right of the ceenter and foremost beaech tree). Shrubs in understorey included spicebush (typically the most common), pawpaw, redbud, flowering dogwood, and wild hydrangea (typically the least common). Forbs were essentially absent at this estival aspect of this lowland (creek drainage) forest, but wild ginger was the most abundant herbaceous species in forest range vegetation presented in these two photographs. The dark-brown piece of wood in the first of these photographs (near right margin; in front of a beech) was a fallen limb (species was not discernable) that apparently died while part of the forest canopy. Such fallen dead wood was a frequent phenomenon in this dense, mesic forest even though most of the largest trees were young second-growth. Dark woody vines in the second of these photographs (at least three were visible) represented numerous species of grape (Vitis sp?) that are common throughout the entire Ozark Plateau Region. The forest composition and structure presented in these two slides was representative of that that developed from the foot almost to the top of north slopes. Beech was replaced to some extent by white oak and pignut hickory in upper parts of both north and south slopes (beginning lower on south slopes of course). These two photographs were of the beech-white oak stand that developed in a secondary drainage into Clark Creek, an intermittant stream. Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Mixed mesophytic form of the oak-hickory forest with American beech as defining/distinguishing species and in association with sugar and red maple designated this as a isolated unit of a variant of SAF 60 (Beech-Sugar Maple) rather than as a variant of SAF 52 (White Oak-Black Oak-Northern Red Oak). Likewise, isolated unit of Beech-Maple Series 122.13, Northeastern Deciduous Forest 122.1 of Brown et al., 1998, p. 37). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004). |
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71. Composite shot of an American beech-white oak bottomland forest- A hardwood forest of mixed mesophytic form in the Boston Mountains included beech as a dominant (along with white oak, northern red oak, and pignut hickory which are the typical dominants of more mesic forest environments in the oak-hickory association). This beech-oak-hickory forest range developed in a small hollow through which a stream (Clark Creek) flowed intermittantly. Specifically the flow of water typically became subterranean during drier seasons (winter and mid through late summer, especially).Thus this forest range community was neither riparian vegetation nor a floodplain forest. Rather this was a very mesic (and periodically a wet) forest due to the sheltered environment of a deep, narrow hollow yet one with well-drained soil (very cherty/very stoney silt loams overlying a subsoil of clay to clay loam) and slightly acid to slightly alkaline generally fertile soil. It was this combination of edaphic features together with protection from more severe physical condition afforded by the deeply dissected topography that permitted dominance by American beech as well as typical dominant species of oak and hickory. Most of the plants in lower layers seen in this photo-quadrant were seedlings of beech, first or most common, and white oak, second. Most numerous and with greatest cover in the lowest herbaceous layer was wild ginger, the overall most abundant forb in this mesic, stream-drainage forest at least in the estival aspect. The most obvious shrub in this slide was some species of grape that was well-represented by conspicuous lianas. Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Mixed mesophytic form of the oak-hickory forest with American beech as defining/distinguishing species and in association with sugar and red maple designated this as a isolated unit of a variant of SAF 60 (Beech-Sugar Maple) rather than as a variant of SAF 52 (White Oak-Black Oak-Northern Red Oak). Likewise, isolated unit of Beech-Maple Series 122.13, Northeastern Deciduous Forest 122.1 of Brown et al., 1998, p. 37). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004). |
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72. Arkansas as the first Arkansawyers knew it- American beech and white oak were co-dominants here on the upper portion of a north slope that was topped with bluffs and aligned parallel to an intermittant stream (Clark Creek. This co-dominant beech-white oak mixture was typical of most photographic samples of this mixed mesophytic aspect forest that developed in hollows of the deeply dissected Boston Mountains. Higher up near top of the bluffs sugar maple and wild hydrangea comprised a different forest community. Much of the lower woody layers of the forest vegetation seen here was regeneration of the two dominant tree species, but spicebush, pawpaw, Caroling buckthorn, and American bladdernut plentiful so that shrub layers were well-repesented. Cucumber tree and umbrella magnolia were sparsely present and along with the more common flowering dogwood comprised an irregular taller shrub or even lower tree layer. Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Mixed mesophytic form of the oak-hickory forest with American beech as defining/distinguishing species and in association with sugar and red maple designated this as a isolated unit of a variant of SAF 60 (Beech-Sugar Maple) rather than as a variant of SAF 52 (White Oak-Black Oak-Northern Red Oak). Likewise, isolated unit of Beech-Maple Series 122.13, Northeastern Deciduous Forest 122.1 of Brown et al., 1998, p. 37). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004). |
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73. A little "beech" of northern deciduous forest in the Ozark Region- In the Boston Mountains the mixed mesophytic aspect (Braun, 1950, p. 170) of the regional climax oak-hickory forest was dominated at low to above mid-level elevations of moist hollows by American beech. Some larger second-growth beech trees were featured. in this photograph to illustrate the interior of what is almost certainly either a post-climax form or, alternatively, an edahic or physiographic climax of the oak-hickory association in the Ozark Plateau. Forest vegetation like this is more like what would be expected in New England than in the southcentral forest region of North America. It was remarked variously throughtout this chapter of the Oak-Hickory Forest that this anomalous (and, again, post-climax or edaphic/topographic climax) vegetation defied ready and unambiguous classification as to forest cover type. Given the widely scattered presence of sugar and red maple (mostly at higher elevations and in shelter of bluffs, especially of the former) this could be interpreted as isolated units of the Beech-Maple Association within the surrounding and otherwise unbroken Oak-Hickory Association. This general relation of what could be seen as "outliers" of other forest associations within masses of the regional association was explained by Braun (1950, ps. 33-34). It seemed plausible to the present author that this beech-dominant (or co-dominant, as with white oak) forest was a physiographic or an edaphic climax as suggested by Braun (1950, p. 34). Braun's recognition of such climaxes was her incorporation of Arthur Tansley's polyclimax concept into her general reliance on Clements' monoclimax concept which, of course, would view this beech-dominated forest in the Oak-Hickory Region as post-climax. Whatever this forest vegetation was quite atypical (more mesic for one thing) for Ozark forests. Most regeneration portrayed here was beech which showed to good advantage the rating of Very Tolerant for this species. Shrubs in this forest habitat within a deep, narrow hollow included spicebush, pawpaw, American bladdernut, flowering dogwood, and some Carolina buckthorn. The most common herbaceous species was wild ginger. Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Mixed mesophytic form of the oak-hickory forest with American beech as defining/distinguishing species and in association with sugar and red maple designated this as a isolated unit of a variant of SAF 60 (Beech-Sugar Maple) rather than as a variant of SAF 52 (White Oak-Black Oak-Northern Red Oak). Likewise, isolated unit of Beech-Maple Series 122.13, Northeastern Deciduous Forest 122.1 of Brown et al., 1998, p. 37). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004). |
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74. Second-garowth stand of American beech- A spot in a mixed mesophytic aspect forest generally dominated by American beech and white oak in which beech was represented by various age classes which illustrated the Very Tolerant feature of this species and provided an example of a consociation. The small, dead limb in mid foreground had fallen from the largest beech providing an example of the self-prunning feature of this species. There were also grape vines growing up into crowns of beech in the background which should not be confused with self-prunned, hanging branches. The understorey in this scene was rather sparse with broad beech fern being the most common herbaceous species. Beech regeneration made up most of the lower layers of vegetation. Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. |
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75. Some more light; some more biodiversity- Paired (vertical and horizontal) photo-plots showing understorey vegetation by a large second-growth American beech where there was a lower density of beech trees, saplings, and seedlings than was typical for the mixed mesophytic forest of which this local stand was part. The narrow terrace of an intermittant stream (Clark Creek) in the bottom of a hollow provided an unusually moist and shelted environment in which a "vegetational island" of beech forest had developed in which white oak, northern red oak, and pignut hickory varied locally in dominance with American beech. Sweet gum, sugar maple, sycamore, basswood, umbrella magnolia, cucumber tree, and mockernut hickory were some of the other larger woody species growing in this forest range vegetation. White oak, sugar maple, and sweet gum (this latter a seral species persisting into claimx or near-climax stages) were local associates growing immediately behind this matriarch beech. There was regeneration of beech, sugar maple, and white oak though primarily of beech, the reproduction of which was represented by several age classes. The presence of beech and sugar maple (both Very Tolerant species) in age classes ranging from young adult to seedling suggested to this worker that the mesic forest which had developed in this extraordinarily favorable forest site was a local variant of the Beech-Sugar Maple forest within the surrounding regional climax of the Oak-Hickory Association. Shrubs present in these two slides included spicebush and, in background, American bladdernut and pawpaw.Herbaceous species (both as to kinds as well as density and cover) were considerably more abundant in the more open space associated with this large beech. This microhabitat provided an example of ground cover by hairy wood (= Canada, woodland) brome and broad beech fern, the two species having greatest foliar cover, along with wild ginger, round-lobed liverleaf, bottlebrush, Virginia creeper, and poison ivy. Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Mixed mesophytic form of the oak-hickory forest with American beech as defining/distinguishing species and in association with sugar and red maple designated this as a isolated unit of a variant of SAF 60 (Beech-Sugar Maple) rather than as a variant of SAF 52 (White Oak-Black Oak-Northern Red Oak). Likewise, isolated unit of Beech-Maple Series 122.13, Northeastern Deciduous Forest 122.1 of Brown et al., 1998, p. 37). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004). |
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76. Beech fern on bottom of beech forest- In spite of generally shaded conditions in the beech-white oak mixed mesophytic Boston Mountains the Almighty provided a small shaft of light in late afternoon that permitted sharing of this pteridophytic forb with loyal viewers. This broad beech fern (Thelypteris hexagonoptera= Dryopteris hexagonoptera= Phegopteris hexagonoptera) was one of a number of its kin that locally dominated--frequently to exclusion of all other herbs--the herbaceous layer in this post-climax or physiographic/edaphic forest. This specimen was growing at base of the large American beech featured in the two-slide set shown immediately above. The fern was accompanied by round-lobed liverleaf which was barely visible to the right. Lost Valley, Buffalo National River. Newton County, Arkansas. |
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77. What a little light can do- Another set of paired photo-quadrants to present progressively closer-in views of understorey vegetation at base of another large American beech on the terraace of Clark Creek. This was another example of a floristically diverse local assemblage of woody and herbaceous species that had developed near a mature beech tree in one of the numerous hollows in the Boston Mountains. The species composition of understorey layers growing by this tree was similar to, yet distinctly different from, that living by the beech in the three preceding slides. In this local forest stand the more common understorey species were California or hairy wood brome, jack-in-the-pulpit, poison ivy, and Virginia creeper. Unlike the species mix of lower vegetational layer by the beech in the previous photo-quadrants this microsite had numerous seedlings of pignut or bitternut hickory, another defining climax tree species of more mesic forests of the Ozark Plateau. It was reported above that the Boston Mountains have been regarded as being one of the more deeply dissected plateaus in the Ozark Plateaus Region (Fenneman, 1938, p. 655-660). These geologic dissections have formed hollows that in this part of the Boston Mountains furnish more protected and moist environments that support forest vegetation dominated and/or characterized by more mesic species that are typical of deciduous forests which are climax at more northerly latitudes. It iseemed plausible to this observer the combination of sandstone and limestone parent materials formed soils with extra favorable combinations of pH, fertility, water-holding capacity, drainage, and depth that were uniquely conducive to growth and survival of more demanding, persnickety species like the vulnerable beech which requires growing conditions seldom found in the Ozark Mountain Region. In turn, beech provides a sheltering microclimate for understorey species that is also uncommon in the Ozarks. Larger beech trees typically had an understorey immediately beneath them that had less cover of woody plants, including, in some instances, fewer seedlings and saplings of beech (at least unless fire or some injury resulted in root suckering). Herbaceous species like broad beech fern, Canada or woodland brome, and jack-in-the-pulpit benefitted from this set of conditions. Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). Mixed mesophytic form of the oak-hickory forest with American beech as defining/distinguishing species and in association with sugar and red maple designated this as a isolated unit of a variant of SAF 60 (Beech-Sugar Maple) rather than as a variant of SAF 52 (White Oak-Black Oak-Northern Red Oak). Likewise, isolated unit of Beech-Maple Series 122.13, Northeastern Deciduous Forest 122.1 of Brown et al., 1998, p. 37). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004). |
| A wind storm approximately 7-10 weeks prior to time of this Clark Creek
series of slides blew down (some uprooted; others with hollow trunks [probably
due to fire injury] twisted off) some of the largest (presumedly oldest)
trees of American beech and pignut hickory, two climax tree species. Forest
gaps created by local blowdown provided photographic examples of patch
dynamics. Three examples (adult trees) of beech and one of pignut hickory
were presented in the following sequence. Interestingly, the species most
apt to benefit from death of beech and hickory was beech and hickory,
respectively. Each of these climax, tolerant tree species had already
reproduced either adequately sexually (in case of hickory) with enough
seedlings and pole-size trees to dominate the gap or (in instance of beech)
abundantly, both sexually and asexually (clonal shoots from trunk and/or
root) to monopolize the newly created patch. Thus there will be very little
secondary plant succession in gaps. In case of beech, recruitment will
be partly by new shoots of the previous genet (same genotype). This will
be equivalent of new shoots regrown from rootcrowns, stolons, rhizomes
of perennial grasses in a pasture or hayfield.
It was once felt "that beech root suckers do not development into
desirable trees" although some root sprouts could grow independently
of the parent tree (Fowells (1965, ps.174-175). This was accepted as
textbook knowledge (Harlow et al., 1979, p. 284). Subsequent studies
revealed that reproduction of this climax species "is almost exclusively
by suckering in the 'beech gaps'" in northeastern forest. Suckering
is also important-if not essential-in more northern and western portions
of this species' range (Burns and Honkala, 1990, ps. 327-328; original
studies cited and summarized). An example of this marginal extent of
American beech would be the Ozark Plateau in western Arkansas. It is
now realized that "[r]oot sprouts can develop into desirable trees"
(Burns and Honkala, 1990, p. 328). It is still accepted that stump sprouts
from other than smaller beech trees (less than four inches in diameter)
remain alive for only short periods These two tolerant species were not only capable of reproducing in their own shade (generic, poetic description of any climax plant species), but also in patches of sunlite habitats formed by their removal. This undoubtedly explained to a large extent why this second-growth forest had returned to climax species composition in less than half a century following a logging operation that amounted-more-or less-to clcarcutting. Soil seedbank was also a factor especially in case of white oak. An old folk proverb used as a play on words by Mr. Shakespeare taught its own lesson in Forest Ecology: "Ill blows the wind that profits nobody. (W. Shakespeare. 1591. Henry VI, Pt. 3 ii. v. 55]. Blowdown or windthrow of individual large, adult trees created gaps that could be exploited by other plants in lower strata of range vegetation which could then use light previously captured or excluded from lower forest layers by crowns of mature trees. In other words the philosophical saying, "Its an ill wind that blows no man good" is an ecological misnomer. Ain't no such wind.. In the mesic beech-white oak-pignut hickory forest featured here (and in all other forests), as in a Florida or Texas city following a hurricane, devastating winds always bring some good fortune, some ecological "profit", to those who can benefit from the misfortune of others and, in doing so, benefit the community. A tour of four such local scenes of distruction and reconstruction followed in Arkansas' Boston Mountains mesophytic beech forest. |
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78. Snapped right off- This large second-growth American beech provided the first of four examples of a forest gap and the pattern of patch dynamics in a mixed mesophytic forest in athe usually sheltered habitat of Clark Creek hollow. This and the next example of a forest gap were formed by a combination of 1) weakened trunks caused by fire damage and subsequent wood rot caused by organisms that gained entry through the fire scar and 2) strong winds. These two beech were broken off at the location of fire and rot damage. Wind was a necessary but not a sufficient factor for blowdown and gap creation. Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. |
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79. Broke on both ends- The second of three examples of blowdown of individual beech trees along the drainage of Clark Creek in the Boston Mountains of northwestern Arkansas. The same, spring wind storm and previous fire(s)-induced damage to the tree trunk that provided the preceding example of beech blowdown was responsible for breakage shown here. Both trees broke off in high winds at the fire scars, the site of fire damage and resultant wood rot, proving in these two cases that fire and wind were both necessary physical factors for tree loss. There were numerous other adult beech trees in this forest that had fire scars, some of which were much more extensive than the two examples shown here. These other trees which had considerably greater fire damage (two of which were presented earlier in describing this forest) were not broken off (or otherwise topppled) by these winds because they were farther down along the drainage of the stream and in locations that were more sheltered from wind.force. The second of the two photographs of this second patch presented a local stand of white oak that were immediate beneficiaries of increased light made available due to beech blowdown. However these white oak were almost as tall as the downed beech so it was not clear how much the"lucky ones" of the main co-dominant species would benefit from light aspects of the newly created gap. There would certainly be more growing space at the edge of this gap along with additional water and soil mineral nutrietns. Edaphic resources may or may not have been limiting. One obvious thing was that the white oak were "damn lucky" that the main limbs of the beech crown broke off due to their own weight on the way down and before reaching the imperiled white oak. (second photograph). "A miss is as good as a mile". Reminded the author of some "close calls" in his own life. Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. |
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80. Uprooted beech- A third example of a wind-downed adult American beech in the drainage (hollow) of Clark Creek in the Boston Mountains. This beech, unlike the two preceding examples, fell by being uprooted rather than by having its trunk twisted or snapped off at fire scars and site of rotten wood. In this third example wind without fire brought down an adult beech. This does not mean, however, that wind alone was a sufficient condition for gap creation. Rather it was a combination of saturated (or nearly so) soil and strong winds that uprooted this large tree. Water that had sustained this tree for decades and enabled it to reach maturity contributed to its demise. While the mechanism of patch formation was different the resul, tree blowdown, was the same. Likewise, the general pattern of forest dynamics was the same. The same species of climax dominants--mostly American beech and, secondly, white oak--immediately filled the void formed by removal of light-intercepting crowns of adult trees of these same species. In this forest gap there were age classes of American beech extending from seedlings to larger saplings (two in right foreground of first photograph). There were numerous secondary shootrs arising from close-to-the surface roots of the downed beech that were not pulled up when the tree was uprooted. There was nearly zero herbaceous vegetation in this newly created (about two month-old) forest gap. It was not determined at this time if there were stunted individuals of herbaceous species (say, perhaps, hairy wood [Canada] brome) that might be released later with more light. No such plants were noticed. Also, it was not known if annual species would colonize the gap in the next growing season. There was a small stand of young, pole-size white oak at far edge of the gap that were "all set" to benefit from the additional resources made available with loss of this large American beech. Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. |
| Clarifying note: All three of the beech trees blown down by wind (two broken off at fire scars, one uprooted farom wet soil) were growing on fairly level ground so that slope and gravitational force were not factors in these cases of windthrow. |
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81. Too close for comfort- Several beeches barely missed being brought down when this huge pignut hickory was bested by spring winds. How many gusts or even gales over course of its life had whistled through the crown of this monsterous member of a usually sheltered mesic forest that for decades had called the hollow of Clark Creek its home? This massive old hickory finally met its Waterloo when a combination of strong winds and saturated soil on a north slope was too much stress-and-strain for its massive trunk and crown that finally bite the ground that had sustained it through many previous storms. What successional change did this forest gap, this hole in the upper canopy, initiate? Simply put: none. Beech, the most mesic and finicky of the climax tree species was the one with the most young adult trees to get immediate benefits of added light, space, etc. (first of these two slides; background of second slide). Adults of bitternut hickory were also poised to reap any rewards of increased light, spece, soil nutrients, etc. ((tree in middle foreground of second slide). There were also saplings and seedlings of pignut hickory that could benefit from light now capable of reaching to lower strata of the previously closed canopy. Also, bittternut or pignut hickory does develop root sprouts and its seedlings have a high shade tolerance (Burns and Honkala, 1990). Yes, gaps in forest canopies can permit winds to "scoop" down into the remaining forest vegetation resulting in more windthrow of shallow-rooted trees which were able tor remain upright in large part due to the unbroken surface afforded by continuous canoy cover (the outer most layer of "skin" of the forest). Such is of little importance from the standpoint of forest succession when most of the woody layers are dominated by various age classes of the climax rtree species (beech, white oak, and pignut hickory in this forest). Any subsequent windthrow, and gaps created by wind disturbance, will amount to replacement of one age class of the climax dominants by another age class of the same and/or other climax dominants. But the forest will miss this large and longtime resident of the community. Even trees do not live forever, at least not the same trunk and crown. Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. |
| "The rich get richer and the poor get poorer"- This popularized
(and perhaps cynical) folk wisdom has been used for centuries. It became
part of popular culture in 1921 when song writers Gus Kahn and Raymond
Eagan incorporated the already well-known sentiment in the song, Ain't
We Got Fun: "There's nothing surer: The rich get rich and the
poor get—children". In the case of patch dynamics when
and where American beech, the climax dominant of the mixed mesophytic
aspect forest (Braun, 1950, p. 170), was "overthrown" by windthrow
the vacated "successional throne" was immediately ascended and
reclaimed by children of the rich, progeny of
the climax beech. There was very little ecological "presidential
succession" by other (presumedly seral) plant species. Instead root
suckers (= sprouts; secondary shoots arising from roots close to the soil
surface) emerged when released from suppression by apical dominance of
the standing adult tree. There was also some sexual reproduction from
beech nuts, but this was considerably less than the axexual or vegetative
regeneration from root sprouting.
In the instance of blowdown of pignut or bitternut hickory, a co-dominant climax tree species of upper slopes, adult beech trees that were already established-- and luckily missed the crashing corpse of Carya cordiformis--enabled, even strengthened, continued dominance by the most mesic of the climax species. There were also numerous seedlings and saplings of pignut hickory to insure reclaiming of the climax crown by this regionally dominant species. Any benefit to pioneer and other seral species was short-lived. Some more lyrics from Ain't We Got Fun attested to this ecological fact in the mesic Boston Mountains forest: "There's nothing surer: The rich get rich and the poor get laid off". On forest range that is either the most mesophytic variant of oak-hickory forest or an isolated and variant community of beech-sugar maple forest, creation of gaps by natural (and, apparently, human) disturbance does not cause denudation and retrogression that results in secondary plant succession. Instead asexual and sexual offspring of the climax trees "fill in the gaps". Patch dynamics is overwhelmingly replacement of adults of the climax tree species by clones and new genotypes of these same mature shoots. Wind and chainsaws merely mow off adult standing crop (both biomass and necromass) which subsequently regrows by resprouts and new trees of the climax species. Plant succession essentially does not exist in patches (including clearcuts) created in this forest dominance type. |
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82. Even unto death- (or still a part of the forest)- Log of a downed American beech providing a protective and moist, rich microsite for spotted touch-me-not or jewelweed (Impatiens capensis). A big beech fell across a walking ttrail and a trail crew with the National Park Service bucked it up and rolled the rounds off the trail into this neat pile which provided the improved habitat for this forest forb (one of the more mesic forb species in the Ozark Plateau Region). It was also habitat for some species of shelf fungi as its decaying wood provided a substrate for this species that is a decomposer in this forest ecosystem..This shot also showed the role of ecosystem reducers. Even at end of its life cycle the woody carcass of a tree, especially a big one that takes longer to rot, remains a part of the forest ecosystem and continues to provide "goods and services" in the forest range. Not only did the rotting beech log furnish habitat for the producers and reducers shown in theis photograph it also undoubted provided shelter and a food source for soil-swelling consumers such as earthworms and arthropods like centipeds. Nutrient recycling, soil protection (erosion risks are high on the steep slopes above streams like Clark Creek), and water retention (the decaying wood holds and slowly releases water back into the hydrologic cycle) are all important roles served by dead wood on forest range. Benefits of dead and downed wood to forest watersheds were illustrated in this final stage of a forest tree. Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect.. |
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83. One of the few grasses in a beech-dominated forest- Bottlebrush (Hystrix patula) is one of very few grass species that can survive in the dense shade of an Ozark Plateau forest dominated by American beech. Canada (= hairy wood brome), most common, broadleaf woodoats (Uniola latifolia), and Virginia wildrye were the only other species of grass this rangeman could find in this dark forest. Divine Providence furnished a rare and very fleeting shaft of "heavenly sunlight" that enabled this hapless photographer to bring his loyal viewers this gramineous or graminaceous (both adjectives are correct) delight. Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. |
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| 84. A study in American beech (Fagus grandifolia)- A mature beech at old-age including some sucker shoots arising from surface roots. Leaves and the smooth, gray bark of beech were also presented in this composite shot. Buffalo National River. Newton County, Arkansas. July. |
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85. Shady details- Close-up photographs of basal trunk of the American beech presented in the preceding photograph. The photograph presented details of beech bark and (in front of trunk) a secondary shoot arising from one of the upper roots that was naturally protruding above ground surface. Second photograph showed this entire root sucker. The thin, smooth bark of American beech makes it very susceptible to fire injury and sunscald (Burns and Honkala, 1990, p. 329). Sunscald refers to the condition of "localized injury to bark and cambium caused by a sudden increase in exposure of a stem or branch to intense sunlight (insolation) and high temperatures" (Helms, 1998). Certain smooth, thin-barked tree, including both beech and sugar maple, are also susceptible to winter sunscale which can develop on the side of trunks and limbs that are exposed to warmer temperatures and brighter light of winter afternoons. Buffalo National River. Newton County, Arkansas. July. |
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86. Leaves of American beech #1-Leaves and twig on the root sucker of beech introduced in the preceding two slides. Northeast slope above Boxley Valley in Boston Mountains. Phenomena of tolerance, shade leaves, and sunflecks was explained in the two-slide set presented immediately below. Buffalo National River. Newton County, Arkansas. July. |
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87. Leaves of American beech #2- Another (the second) example of a beech root sucker. This specimen was growing in the terrace (broad stream bank) of Clark Creek that drains into Buffalo River. Both this and the previous example of beech leaves also showed the extremely shady conditions produced by the dense foliage of American beech and high this Very Tolerant species can grow and have successful vegetative (asexual) reproduction in the dim microenvironment beneath its own dense foliage. These two photographs (and the two photographs shown immediately above) were taken using natural light at mid-day (1245 hours Central Standard Time). This photographer had to wait until a shaft of light from the appropriate direction struck the leaves of these root sprouts. This was for a brief period lasting no more than a quarter hour. Most likely there were other such brief periods when the orientation of Earth to Sun permitted streams of light to strike these leaves. This is the phenomenon of sunflecks, several examples of which were presented in this section. Obviously these-more-often-shaded-than-sunstruck beech leaves were, on average, more sinks than sources of photosynthate. These lower leaves almost assuredly had to be supported by leaves in the crown which functioned as photosynthate sources for these energy and nutrient-dependant organs. Leaves on these beech root sprouts (secondary shoots arising from near-surface or even protruding roots) were example of shade leaves, those which due to their position in shaded environments are larger and thinner than co-hort sun leaves that are in environments which receive full-sun light (even if on the same tree). Buffalo National River. Newton County, Arkansas. July. |
| Final footnote: presence of white oak as co-dominant with American beech (rather or more than with sugar maple, basswood, pignut hickory being co-dominants) probably reflected spatial contact with the climax white oak-northern red oak upland (south-slope; ridge top) forest contiguous with the local mesic environment afforded by banks of Clark Creek and north-slope limestone bluffs aligned with this stream. It was not clear if white oak trees would live to such age as to remain a component of ultimate climax-as in old-growth-forest or if white oak was of high-seral status. If this latter was the situation then this mixed mesophytic aspect (Braun, 1950, p. 170) forest could be viewed as subclimax with even greater dominance by beech as the topographic or edaphic climax (polyclimax model) or post-climax (monoclimax model). It did not appear to have structure (and perhaps not function) of true virgin (=old-growth) forest for this forest site and cover type, but there was not an old-growth stand of this (or similar) forest cover type with which to compare the forest discussed above.. |
| The next portion of this section described an upland white oak-northern red oak forest that was contiguous with the beech-white oak-hickory forest treated immediately above. This oak forest range community was similar to (and could be considered as a vaiant of) the white oak-black oak-northern red oak cover type dealt with earlier in this chapter. Near absence of black oak, one of the most widely distributed Quercus species throughout the Ozarks Plateau Region, combined with the conterminousness of this forest tract with that of the beech-white oak forest in the hollow (drainage) of Clark Creek just covered made inclusion of this white oak-northern red oak forest here, rather than above, a logical arrangement. |
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88. Rainbow of hardwoods- Mesic upland mixed oak forest with a wide array of woody angiosperms. An upland white oak-northern red oak forest immediately above the mixed mesophytic beech-dominated forest that had developed along Clark Creek provided a rather stark contrast (over a remarkably short distance) of two forest range types in which oak species were climx dominants to one extent or another. In this upland oak forest the species composition of shrub layers was in rather striking contrast to that of the more mesic forest deep in a stream drainage. From one perspective this difference was as dramatic as absence of adult beech from the upland forest (next paragraph). Differences between these two adjacent forest cover types--again,one a lowland and the other an upland community-- were in both kinds of shrubs and structure of the shrub strata. Mesic shrub species of pawpaw, wild hydrangea, and American bladdernut were absent from the upland mixed oak forest. Instead the defining shrub, lowbush huckleberry (Vaccinum vacillans), was widespread in the upland and absent (or nearly so) from the lowland beech-dominated forest. Also more common in the upland mixed oak forest, and essentially absent from the mixed mesophytic beech forest, was sassafras and buckbrush or coralberry. Also present in the upland and absent (at least rare) in the lowland forest was eastern red cedar (Juniperus virginiana). Absence--undoubtedly exclusion (one way or the other)--of fire clearly accounted for presence, and increasing abundance, of this nonsprouting juniper. Sweet gum was not found on the upland forest whereas it was an indicator tree species in the mexic beech-dominated forest below. One of the most telling signs (though what it told was not clear at this point in development of vegetation) was absence of large adult beech from the upland. Beech was preent in the upland white oak-northern red oak forest but only as widely scattered seedlings and small saplings. It was not knownif these would survive to adulthood. Flowering dogwood was the most abundant of the taller shrub species (foremost trunk; midground in first slide) followed--as distant runner-up--by red maple. As already noted, lowbush huckleberry was the most common of lower-growing shrub species. Sassafras was the most conspicuous shrub in the second slide (left foreground with single bole). Sassafras is a species that occurs as both shrub and tree, and as small to mid-size tree. Conspicuous specimens (tall, straight, clear trunks) of white oak and northern red oak were present in both of these photographs. The second photograph was a zoomed-in view of the two largest representatives of these species introduced in the first photograph. This proved to be an advantageous arrangement to present the two subgenera of Quercus. The two large trees in midground of the first photograph and then featured in the foreground of the second photograph were white oak (left) and northern red oak (right) representing Leucobalanus and Erythrobalanus, respectively. At immediate left side of the featured white oak in second slide was a healthy local stand of Canada or hairy, wood brome. Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004). |
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89. About all the principal acators on stage- Composition of mesic upland forest dominated by white oak and northern red oak. Largest tree in center was white oak. To its immediate right (partially concealed by trunk) was the multi-stemed shrub of red maple. A number of individuals of northern red oak were prominent in right fore- and midground. A small sapling of American beech with large shade leaves represented this species at left foreground. Time alone would tell if beech succeeded to a climax dominant on this upland forest site, or even if scattered smaller plants survived to adulthood. Shrubs present in this photo-plot included flowering dogwood, sassafras, and red maple at shrub size and with multi-trunks (just noted). Low-growing shrub in fore- to midground was lowbush huckleberry which was featured at a closer-in distance in foreground of the next slide. Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004). |
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90. All major layers present and accounted for- View of mesic upland white oak-northern red oak forest with a wide selection of tree and shrub species representing all major strata of this forest range community. Lowbush huckleberry, the dominant low shrub species in this tract was photographed in foreground to emphasize abundance and importance of this species. Flowering dogwood, red maple, and sassafras as shrubs were plentiful. Redbud was also present but to extent of the just-listed species. The shrub at far left was cucumber tree (Magnolia acuminata). Pole-size tree at far right was northern red oak. All-in-all, an interesting aggregation of range plants. Lost Valley, Buffalo National River. Newton County, Arkansas. July, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak). Oak-Hickory Series of Brown et al. (1998). Boston Mountains- Upper Boston Mountains 38a (Woods et al., 2004). |
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Bur Oak (Quercus macrocarpa)
Forest
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Bur oak as a forest cover type (SAF 42; Eyre, 1980, ps. 39-40, 236) is the most northward and westward of the eastern and central continental tree oaks. Bur oak comprises the only oak forest cover type in more northern portions of the Central Lowlands and Great Plains physiographic provinces. Bur oak is extremely tolerant of drought and fire and forms forests, groves and savannahs within (and within) the central grasslands of North America from the eastern edge of the Prairie Peninsula (Transeau, 1935) through to the Black Hills (Eyre, 1980, p. 39). Bur oak is to this vast region what post oak and blackjack oak are in the south (eg. Cross Timbers). These two forest range types meet near northern reaaches of the Flint Hills Region. Presented in this section was an example of the bur oak forest cover type (SAF 42) found on a north slope and outward to a ridge top in the Nebraska Sandhills. This was about as dry (least mesic) an environment as bur oak is adapted to, at least to the extent of forming forest communities. This example was a forest and not a woodland or savanna although at its outer edges it did form small groves with tree density and understorey more suggestive, or even typical, of woodland. This contiguous bur oak-dominated community (both forest and grove phases, or two communities if so interpreted) was the climax (ptential natural) range vegetation on this sere except where frequent, close, mechanical mowing had converted the native herbaceous understorey to a manmade one made up of Eurasian perennial grass species adapted to such intensity and frequency of defoliation. Examples of both native and naturalized herbaceous layers were presented following a brief section that introduced the foliage and fruit features of bur oak. This relict stand of bur oak forest had developed on what was at this time the Valentine, Nebraska city park (and almost any city park says it all). This forest range vegetation occupied both upland, including and especially a mesic north-slope, and bottomland site. Vegetation on all but the north-slope was highly modified by yard-mowing, manicure-it urban man (even in this over-whelmingly rural region). The natural herbaceous understorey of the upland phase was Canada wildrye (Elymus canadensis) except on north-slope upland forest range where the dominant herb was—as on the bottomland phase—long-beak(ed) or Sprengel’s sedge (Carex sprengelii). It was likely that there was also some bristleleaf or ebony sedge (C. eburnea) was growing in association with long-beak sedge, but C. eburnea could not be positively identified given absence of inflorescences. Under frequent, close mowing Kentucky bluegrass (Poa pratensis) dominated the lowland understorey. With less frequent mowing smooth brome (Bromus inermis) dominated both bottomland and upland phases. Both of these introduced Eurasian grasses have naturalized widely and under a farmer (vs. forester or rangeman) frame of mind, and management consistent therewith, the native grasses and sedges were crowded out by aggressive, highly competitive agronomic forages. The timeless story since Cain and Abel. The farmer (cornhusker in the Cornhusker State) again won out over the pastoralist. |
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91. Big tree on the prairie- Beautiful and massive specimen of bur oak that made its home in the floodplain of the Bosque River in the Western Cross Timbers of northcentral Texas. Bur oak is obviously a minor member--especially when compared to the dominant post and blackjack oaks--of the scattered forest and woodland communities in southern portions of the tallgrass prairie region. In more northernly parts of this vegetational realm bur oaks replace blackjack and even post oak as the dominant oak. In these northern parts of the Prairie Peninsula and tallgrass-true prairie region bur oak would be second--if that--only to eastern cottonwood as a dominant tree both on grassland and isolated forest communities such as gallary forests. Bur oak is the most widely distributed of all the oak species that have the role of hardwood dominant across the North American central prairies (McGregor et al., 1977, ps. 39-41). Post oak and blackjack oak barely extend northward to Iowa whereas bur oak extends to Ontario. Bur oak forms forests, woodlands, and groves (distinguished by tree density, extent of canopy cover, and size of stand resulting in varying degrees of understorey development) in more northern areas of North America as in the Nebraska Sandhills, an example of which was used for photographs and corresponding descriptions and explanations in this section. Hamilton County, Texas. July. |
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92. Bur(r) or mossy cup oak (Quercus macrocarpa)- Young leaves and catkins of the dominant oak of the northern tallgrass, true, and mixed prairies northward from central Kansas. Bur oak can be interpreted as the northern ecological equivalent of post and blackjack oaks as the aboreal dominant of the grassland-deciduous forest ecotone (Vankat, 1979, p. 221). This species produces the largest acorns of any oak in North America and it is the oak of the famous oak groves and savannas of the northern grasslands (Peattie, 1938). Seedlings rapidly send their tap roots deep into the fertile prairie soils and become quickly established after germination (Weaver, 1968, ps. 135-139). This genetic adaptation to drought combined with the species’ thick fire resistant bark (Allen, 1967, p. 15) make it admirably suited to drought- and fire-prone prairies. It’s range extends far south of it’s region of dominance into central Texas where it sometimes dominates bottomland savannas. Hamilton County, Texas, April. |
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93. Leaves and fruit of buroak- Burr oak (either one or two "rs" are used) is regarded as having the largest acorn of any Quercus species in North America. Like other white oak species buroak bears fruit every year (versus a two-year cycle) in the red or black oaks. The combination of large acorns and production each year makes this species one of the most valuable producers of mast in the eastern deciduous forest (ie. the eastern part of North America). Burroak is also known as mossy-cup oak, both common names in reference to the conspicuous tapered tips of the scales of the cup which present a fringe-like appearance. The geographic range of. Q. macrocarpa extends from eastern Canada (New Brunswick and Quebec) to Texas. Hamilton County, Texas. September. |
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94. Unusual leaf arrangement in bur oak- Bur oak has a unique arrangement of leaves along its leaders. Leaves grow in dense clusters or whorls at intervals along young branches (leaders) with one of these clusters (complete with the current year's acorns) on each of several internodes extending back toward the trunk until older internodes cease to produce such clusters. Two such leaf and acorn clusters on one leader were shown in this photograph. These were obviously the two youngest internodes which showed the last two year's growth of this shoot (one internode being produced each growing season). Erath County, Texas. Late September; approaching fruit-ripe stage of phenology. |
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95.. Clusters of leaves (and acorns) in bur oak- Two clusters or whorls of leaves and acorns in bur oak. The first photograph presented a whorl from a lengthwise view whereas the second photograph presented more of an oblique and semiend-on view of the second whorl. Bur oak concentrates its leaves in large clusters infrequently along its branches (one cluster per internode) whereas most oak species produce individual leaves and some secondary (usually short) shoots off of young branches (= leaders or shoots) at more frequent intervals along their internodes. Erath County, Texas. Late September; approaching fruit-ripe stage of phenology. |
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96. Clustered at the end- End-on views of two different clusters or whorls (at progressively closer distance) of leaves in bur oak. This is a unique and eye-catching arrange of foliage and fruit in a Quercus species especially well-adapted to drought and fire. Bur oaks grow at variuos densities ranging from isolated individual trees to small groves to dense forests (see rest of section immediately below). Function of this unique arrangement (pattern) of leaves in bur oak was unknown to this author. Erath County, Texas. Late September; approaching fruit-ripe stage of phenology. |
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Technical Note: The following sequence of photographs of the Bur Oak forest cover type (SAF 42) was taken in early summer (late-June). Two aspects were presented and described: 1) north-slope and 2) hilltop. These represented two distinct aspect-based bur oak forest communities even though both were: 1) dominated by bur oak, 2) the same cover (dominance) type, and 3) upland forest. Both aspect-determined bur oak communities were photographed at the same time (about 0730 to 100 hours Central Standard Time). This time of day was most advantageous for pphotographing the north-slope forest which, having a general northeast orientation (hillside alignment), received most light in the understorey (maximum land surface area exposed to full-sun lighting; greatest understorey coverage of greatest light intensity) in early to mid-morning. Obviously time of daylight hours (excepting very early morning or late evening) was largely irrelevant for photographing vegetation of hilltop forest. Relatively high density of adult bur oaks on the north-slope forest coupled with dense individual crowns of this large-leafed species resulted in one of the darkest understorey habitats ever seen by this photographer. Most of the understorey area (square footage, yardage, meterage, etc.) of the north-slope forest range received somewhere between half to three-fourths of an hour of direct light daily. During the remainder of daylight time plants--especially herbaceous species--were in relatively dense shade and obtained only indirect light. Herbaceous species in bur oak forest have to be some of the best adapted and most extreme examples of sciophytes (skiophyes) or sciaphlic (skiaphilic) plants, "shade-loving" or plants that have evolved to shade (shady habitats), in the eastern deciduous forest. Viewers should realize that the shady (generally dark; poorly lite) images presented below, while of relatively poor viewing quality (compared to range types in other biomes), "caught" exactly normal light conditions as they existed, and at their brightest (time of day in which most light penetrated forest crown canopy to reach ground level). No artificial lighting was used. In other words, intensity and quality of light seen in photographs were those that the viewer himself would have experienced, the images that would have traveled through his cones, rods, and optic nerve, had he moved through this forest environment.One can but marvel at the adaptation of the herbaceous plants in bur oak forest range. |
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97. Sylvan scene in Nebraska Sandhills (Are you kiddin"?)- Send this as a postcard from the Nebraska Sandhills and those who deal in sterotypes will asert, "You're kidding me". Nope, the real thing.. Bur oak Sandhill forest on a north slope in early summer. Most of the trees in these two "dendrographic" photographs were bur oak (including the largest tree and the pole-size sampling in left front of the "big 'un"), but white or American elm (Ulmus americana) and green ash (Fraxinus pennsylvanica) were associates that were present (as in background of these two slides). (Incidentially the commerative specific epitthet pennsylvanica means literally "Penn's woods" [ in honor of the Quaker founder, William Penn] ; " sylva from the Latin and Greek roots is in reference to "wood" or "forest"). This was an old-growth stand of bur oak with the climax herbaceous species still dominating the understorey. This was clearly virgin range vegetation. The graceful herbaceous understorey in these two "photo-plots" was almost exclusively made up of long-beak or Sprengel's sedge (Carex sprengelii). Canada wildrye (Elymus canadensis) was the associate herbaceous species overall though very little of this grass was present in forest range vegetation presented here. Eastern red cedar (Juniperus virginiana) had in absence of fire begun to encroach as an invader into this sylvan Eden. Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Kuchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001). |
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98. Appreciative of what it receives- Close-up "photo-quadrant" of local sward of long-beak or Sprengel's sedge presented in the first photograph of bur oak forest treated in this series. This is the very fleeting maximum intensity of light that these shoots will receive.Short duration of light from early morning rays on a north slope in Nebraska Sandhills. Cherry County, Nebraska. Late June. |
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99. Forest and sunflecks- Nebraska Sandhills bur oak forest and shade-adapted herbaceous species. Light conditions of early morning, the major part of day during which light could reach ground level of this north-slope forest, provided a textbook illustration of the sunfleck phenomenon. In dense-canopy plant communities like forest (and even grassland swards) sunlight penetrates through foliage at varying lengths of time. The shortest of these light periods or bouts (time lengths or durations of light at whatever light intensity) which can last only a few seconds up to several minutes are known as sunflecks (Smith et al., 1989; Chazdon and Pearcy, 1991). The converse of sunflecks are shadeflecks which are shade fluctuations due to cloud cover between shaded spots and the sun. In vegetation, such as that of a bur oak forest featured here, wind movement of leaves, travel of Earth relative to sun, and any cloud cover are responsible for the ever-changing state of light that penetrates to lower strata of the range plant community. The general condition or phenomenon of changing or dynamic light as to duration and related intensity or "fluctuation in irradiance" as it was described by Chazadon and Pearcy (1991, p. 760) is an all-important abiotic factor in growth and survival of plants in lower layers of range vegetation-- the understorey of this bur oak forest. Light dynamics interacts with other factors (abiotic and biotic) such as defoliation, precipitation, and, as in this north-slope forest, aspect. Dense shade produced by the large leaves and close interlocking crowns of bur oak on the aspect most shielded from sunlight resulted in severe conditions of light deficiency related stress for understorey plants in this bur oak forest. Range plants in lower layers of this north-slope bur oak-dominated forest community were sciophytes (sciophytes), "shade-loving" species of the highest adaptation. There were a few, though extremely patchy or irregular, layers of woody plants below the forest canopy. These vegetational strata were made up primarily of regenerating bur oak (seedlings, saplings, and poles), American elm, green ash, and, least of all, eastern red cedar. This photographer did not see any shrub species. The herbaceous layer (which was also the ground layer) of this forest range was dominated mostly by long-beak or Sprengel's sedge with Virginia wildrye the associate species. The major--and about the only--forb was anise root (Osmorhiza longistylis). This virgin range vegetation was a fine representation of old-growth bur oak forest (even if there were too many shadows to show it to best advantage). Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Kuchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001). |
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100. A study in shadows- Closer-in photograph of the two big bur oaks and associated herbaceous layer introduced in the last slide (second slide of the preceding pair of photographs). This will be about all--and clearly the most (greatest intensity and longest duration of)--light that the understorey of this north slope bur oak forest will receive, and this was close to longest day of the year. Early morning and late evening were the major (about the only) times that sunlight could fall on the ground at this extreme north aspect. The phenomenon of sunflecks as explained in the preceding caption was obvious. Understorey was herbaceous, except for basal sprouts of bur oak (see below), and consisted primarily of long-beak sedge with some Canada wildrye. The only forb of note was anise root (Osmorhiza longistylis). This was virgin vegetation in structure and composition and in all layers; a good representation of climax bur oak forest range. Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Kuchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001). |
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101 About as bright as it ever gets- The slant of earlier morning light on this north slope is just about as much photosynthetically active radiation as the understorey of this north-slope, Nebraska Sandhills bur oak forest ever gets. These paired photographs presented an example of light dynamics in the understorey layer of this densely shaded forest range. The second of these two photographs was at a closer distance to trees, but the major difference in pattern of light and shade on the forest floor was due to changing radiation that occurred with passage of time. These photo-dynamics resulted from 1) variation due to the diurnal (day-and-night) cycle of light and 2) annual cycle due to travel of Earth on its yearly orbit around the sun. Sunflecks lasting only a few minutes or, perhaps more commonly, mere seconds (or fractions of seconds) are largely due to wind movement of leaves in the forest canopy. This rapid photo-dynamics--very short-term spots of alternating sun and shade--could not be captured in photographs or sequences photographs, but the prevalence of light dynamics, hence the concept of sunflecks, was represented photographically. Trees and saplings in these slides were all bur oak. The herbaceous layer was mostly long-beak sedge with some Canada wildrye and, even less, anise root. Old-growth bur oak forest serving as a reminder of what a virgin forested range was like in a most unlikely grassland region. Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Kuchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001). |
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102. Shade-adapted range plants- This two-slide set of a north-slope, Nebraska Sandhills bur oak forest featured the 1) herbaceous understorey dominated by long-beak sedge with Canada wildrye as associate species and anise root as principal (essentially sole) forb and 2) base of a mature bur oak with stump sprouts and a sapling. The sapling was at such distance (about a foot and a half) away from the trunk of the adult tree that it was apparently a separate tree derived from an acorn (most likely the adult bur oak it was closest to). Thus this bur oak sapling appeared to be a sexual offspring of the adult oak, a separate and unique genetic individual produced by an adult bur oak through fruit. It was possible that this sapling grew from an acorn produced by a neighboring bur oak and transported next to the big oak shown here by a combination of gravity, wind, and steep slope or by a food-storing squirrel. There was some probability that this sapling was actually a basal sprout, secondary shoot, that arose from the adult tree, but this seemed not to be the case. By contrast, the small stump sprouts to the right of the sapling were asexual offspring from the large bur oak. In this case the new oak shoots (stump sprouts or stump suckers which are secondary shoots) were clones (clonal progeny) of the parent tree. These were genetically identical to the big bur oak from which they arose. Stump sprouts were secondary shoots of the existing bur oak tree the same as new limbs, branches, and buds. Such clonal shoots are often called offshoots. This asexual reproduction is vegetative reproduction (synonyms), human horticultural forms of include grafting and budding. More (and more clear) examples of suckering from healthy, uninjured, mature bur oak was shown below near end of this section. Almost all tree species are capable of both sexual and asexual reproduction, but sexual regeneration is less successful in forest of dense shade unless the species are relative in tolerance (Tolerant or Very Tolerant). Vegetative regeneration is not limited (at least not nearly as much so) because the parent tree can translocate photosynthate and other nutrients to its own shoots (clonal organs). By contrast, sexually produced progeny are their "own plants" and must be able to survive under the shade of their parents at age classes ranging from seedling through sapling to pole-size. Most of these individual smaller trees--at least in species having lower tolerance--die. An example of this reality was the stick (at left of trunk and sapling) which was the remains of a small sapling tht could not survive in the shade of (compete with) the aduld bur oak. Burns and Honkala (1990) reported that bur oak has usually be interpreted as being Intermediate in tolerance rating although certain observers viewed bur oak as more tolerant than other climax oaks such as northern red and white oak. In the Prairie Peninsula Region however bur oak stands have been invaded by other oaks including white oak, and bitternut hickory (Burns and Honkala, 1990). This was far to the east and on more mesic habitats than in the Nebraska Sandhills where the more mesophytic oaks and hickories are absent. These same authors specified that in forests to the east and north bur oak died from suppression (due largely to shading?) after reaching sapling size. That same response was frequent in the north-slope Sandhills bur oak forest described here. Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Kuchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001). |
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103. Dominants of a deep, dark lower layer- Local sunlit patch of the herbaceous layer in a north-slope bur oak forest in Nebraska Sandhills. Plant species seen here were the climax herbaceous species that dominates the understorey of a sandhills bur oak forest..Long-beak or Sprengel's caric sedge (Carex sprengelii) was the dominant--almost exclusive--species with Canada wildrye the associate species. Anise root was the only forb of consequence in the herbaceous layer. These were reported as characteristic (indicator) herbaceous species in bur-oak dominated range vegetation (Barker and Whitman, 1989, ps.17-18). The first of these two photographs was of a single-species stand of long-beak sedge. Second photograph included a shoot of Canada wildrye accompanying the long-beak sedge. Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Kuchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001). |
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104. Contaminated understorey- Sward of herbaceous layer of a north-slope bur oak forest in Nebraska Sandhills consisting of long-beak or Sprengel's sedge and of naturalized Kentucky bluegrass (Poa pratensis). Kentucky bluegrass and smooth bromegrass (Bromus inermis) are two widespread Eurasian, perennial festucoid grasses introduced by the white man that have natrualized widely in North America. These two introduced grasses are commonly used throughout the greater Northern Great Plains Region (as well as areas of adjoining regions) for grazing and, in case of smooth brome, for both pasture and hay. They continue to have ample opportunities for naturalization and establishment across a vast acreage, especially on disturbed sites in humid through semiarid zones, so they are properly regarded as naturalized range plants.. Continued heavy mowing up to edge of the north-slope, old-growth bur oak forest described heein enabled both Kentucky bluegrass (as shown here) and smooth brome (covered below) to invade outer parts of the native herbaceous understorey of this bur oak stand. This invasion allowed displacment, to varying extents, of native graminoid species like long-beak sedge and Canada wildrye. These two agronomic, Eurasian grasses were able to outcompete natives only where there was continued disturbance by repeated, heavy mowing over specific locations or of adjacent areas from which the exotics could spread for short distances into unmowed native sward. The sward shown here was an example of the latter situation. Cherry County, Nebraska. Late June, peak flowering stage of both long-beak sedge and Kentucky bluegrass. |
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105. Long-beak or Sprengel's caric sedge (Carex sprengelii)- Three photographs showing progressively greater detail of sexual shoots on the dominant herbaceous species in the understorey of a north-slope Nebraska Sandhills bur oak forest. Cherry County, Nebraska. Late June, peak flowering stage just after anthesis. |
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106. Broadleafed Lone Ranger- Anise root (Osmorhiza longistylis) in the herbaceous layer of the understorey of a north-slope bur oak forest in Nebraska Sandhills. This was only forb that this photographer could find in this climax forest range. It is a characteristic species; generally a dominant and indicator species in the bur oak-dominated forests of this region (Barker and Whitman, 1989, p.18). Its graminoid neighbors were long-beak caric sedge, the dominant, and Canada wildrye, the associate species. Cherry County, Nebraska. Late June, late pre-bloom phenological stage. |
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107. Invasion at the edge- Lower and outer edge of a north-slope bur oak forest in Nebraska Sandhills with an individual eastern red cedar prominently and happily growing in a mid-layer of vegetation. This immediate local was just above Minnechaduza Creek. This accounted for the lush stand of herbaceous riparian vegetation in the foreground. which "crept in" into this "photo-plot" useed to show size of the cedar relative to typical size of adult bur oak (to immediate right of eastern red cedar). Human suppression of fire had permitted eastern red cedar to establish sporadically both along edge and in interior of the bur oak-dominated forest. Most of the herbaceous riparian vegetation was the introduced Eurasian grass, smooth brome which invaded the bur oak understorey under frequent, heavy mowing. Also present with much cover and density, though lacking height of the bromegrass,was another Eurasian grass, Kentucky bluegrass. Other riparian species were native grasslike plants (Cyperaceae) that were still only in vegetative stages and unidentifiable by this worker. These had, however, been overwhelmed by the introduced grasses. The only native plants that could persist under cover of the agronomic grasses (which were competitive with native herbaceous species only under heavy mowing) were bur oak, green ash, American elm, and eastern red cedar. And mowing wiped out all plants of these tree species except saplings and adult trees. Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Kuchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001). |
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108. Bur oak stand at the edge and changed by man- Part of the perimeter of the north-slope Sandhills bur oak forest adjacent to a field of smooth bromegrass. Extension of close (short-height) mowing, over a number of years, into the edge of the bur oak forest had either directly killed out long-beak sedge, Canada wildrye, and anise root and/or permitted invasion of smooth brome and Kentucky bluegrass that "choked out" (outcompeted, overshadded, etc.) these natives. Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Kuchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001). |
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109. Advance guard of bur oak-dominated vegetation being invaded by enemy legions- Two views of a bur oak forest with herbaceous layer(s) that were highly man-modified. This local small stand of bur oak stood in stark contrast to the virgin climax bur oak forest featured above. These two photographs showed the outer- and lowermost perimeter of the bur oak community covered in this section. The adult bur oaks and a few patches of the natural understorey beneath mature trees had persisted (where oak trunks were too close together to permit entry and operation of rotary mowers [=shreaders]) along this perimenter. Generally, however, continued close mowing of the range vegetation had done two things: 1) prevented regeneration of bur oak by cutting off all bur oak seedlings and 2) largely converted the natural herbaceous understorey to an anthropogenic or manmade (=artificial) layer of smooth brome with local stands of Kentucky bluegrass. Change in species composition of the herbaceous layer(s), replacement of native species with naturalized Eurasian species, was due to either direct killing of long-beak sedge, Canada wildrye, and anise root by defoliation or indirectly by reducing competitiveness of these range species and/or favoring the taller-growing smooth bromegrass along with denser-growing, more cutting-toleraant Kentucky bluegrass. It was explained above that both of these introduced, agronomic forage grasses have naturalized throughout much of the North America range country. Where mowing disturbance (= abnormal defoliation regimen) occurred throughout much of the growing season the understorey was converted from an herbaceous layer of native grasslike, grass, and forb species to one of naturalized agronomic grasses (ie. conversion from native to naturalized range plants). Also, the lower woody layer(s) was totally eliminated because mowing prevented regeneration of all tree species. Beyond doubt or debate, continued close mowing as had been practiced on this forest range for years will eventually eliminate the bur oak forest (as soon as the already old trees die off) and convert the plant community into a manmade grassland of smooth brome and Kentucky bluegrass, a farm field of domestic forages. Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Kuchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001). |
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110. Artificial understorey- Continued close mowing by roatary shredders killed out--directly or indirectly--the climax herbaceous species (long-beak sedge, Canada wildrye, anise root) of a bur oak forest understorey and converted the sward to a single-species layer of smooth brome.A small remnant of smooth bromegrass remained uncut where this beautiful trunk of an old-growth specimen of bur oak prevented mowing (an inexperienced or lazy shredder operator did not get close enough to the oak). The other glaring outcome of mowing was absence of regeneration of bur oak, dominant tree species, and green ash, the associate species. If present mowing practices continue (and they obviously had been in effect for a number of years) there will eventually be no bur oak forest left. Instead, the plant community will be a monoculture of smooth brome (with local spots of Kentucky bluegrass). All native vegetation will be wiped out (ie. the forest range will be no more and a hay field will have replaced it). Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Kuchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001). |
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111. Genuine-article understorey- Two trunks of large bur oak prevented close mowing of the native herbaceous understorey comprised of long-beak or Sprengel's caric sedge, the dominant, and Canada wildrye, the associate species. Survival of these climax species permitted some of the composition and structure of a bur oak forest range to persist. This part of the old-growth bur oak forest that had developed on a north slope in the Nebraska Sandhills was mowed with less frequency than some of the other forest understorey with the result that the climax species composition of the sward (herbaceaous layer) persisted. Tree regeneration (bur oak, green ash, American elm) was still completely prevented as any seedling was mowed off where density of adult trees and saplings did not prevent entry of rotary mowers. Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Kuchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001). |
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112. Nebraska Sandhills bur oak grove- A bur oak grove, the outermost edge of which was shown here, had developed on a sunlit ridge conterminous with the north-slope bur oak forest described immediately above. Both phases--open, fullsun grove and dense, deeply shaded forest--were climax range vegetation and represented the extremes physiographic and structural manisfestations of the bur oak forest cover type. The greatest differences were in the understorey vegetation, as to vegetational structure and species composition of herbaceous and woody (shrub and smalltree) layers. The bur oak grove phase (or community, if the two extremes were to be viewed as two different range plant communities) had no species of woody plants except bur oak (in contrast to American elm, green ash, and eastern red cedar in the north-slope forest). Also absent from the bur oak grove on the drier and more sunlit ridge was long-beak cric sedge which made up most of the herbaceous biomass on the north-slope forest. Instead, Canada wildrye, associate on the north-slope forest, was the overall dominant of the herbaceous layer in the bur oak grove. Giant ragweed (Ambrosia trifida) was a locally dominant herbaceous species and the most abundant forb in the oak grove. Anise root, the only forb, on the north-slope forest was absent in the grove vegetation. This bur oak grove was on the border of a field of the Eurasian perennial grass, smooth brome, which, as shown above, had invaded the understorey of a north-slope bur oak forest under repeaded close, mechanical mowing when the native long-beak sedge, Canada wildrye, and anise root were killed out by such extreme (intense and frequent) defoliation. Where trunks of bur oak were too close together to permit intrusion of this bloody equipment and overmowing the native, cool-season Canaada wildrye dominated the understorey other than in microsites where giant ragweed and bur oak stump sprouts held this honor. Physiogonomy and structure of this Sandhills bur oak grove was presented in this two-slide episode. Details of the herbaceous layer of this grove was presented in the next two-slide episode. Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Kuchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001). |
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113. Edge of a bur oak grove- Lower zone of tree trunks and herbaceous understorey of a Nebraska Sandhills bur oak grove. This stand of climax vegetation was conterminous with the north-slope bur oak forest featured earlier. In contrast to the deeply shaded understorey of the bur oak forest in which long-beak caric sedge was dominant and Canada wildrye was the associate species, the herbaceous layer of the grove phase of the bur oak cover type was dominated by Canada wildrye except in small areas (microhabitats) where giant ragweed held supremancy or where basal sprouts of bur oak overtopped the dominant cool-season, festucoide grass. Canada wildrye was featured in both of these photographs with the local sward in left foreground of first slide shown at close range in the second slide. Some of the dead (light brown or buckskin-colored) shoots of Canada wildrye included some of the previous year's growth (slightly more faded) as well as some of those of the current growing season. Almost all of the current year's shoots of Canada wildrye were still in the boot. It was not known why a few of the current growing season's shoots had already matured, died and gone into dormancy. Did make for a nice contrasting picture with Canada wildrye very conspicuous. Giant ragweed, a warm-season annual composite of mature size that lives up to its common name, was still in early growth and thus not shown in proportion to its yield of biomass at peak standing crop. This herbaceous understorey was the vernal society even though it was summer and bur oak were representative of the estival aspect. Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Kuchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001). |
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114. Parents, progeny, and playmates- Sprouting (suckering), the production of secondary shoots, from base of trunks in non-injured adult bur oak trees. Bur oak in both of these "photo-plots" were on a ridge at edge of a bur oak grove where light could penetrate from all directions throughout most of the day. This was a different condition of radiation than that of the north-slope bur oak forest described at beginning of this section. Differences in understories of the bur oak grove considered here and the bur oak forest described previously were explained in the immediately preceding caption. It was strikingly evident that full-grown (adult) bur oak produced basal sprouts in both deep shade and full sunlight (forest and grove stands, respectively). This was an understandable characteristic when it was born in mind that these secondary shoots were being supported by the parent shoot (ie. sprouts were a photosynthate sink from the parent tree that was the a nutrient (food) source for the basal suckers. (Sprouting frrom stumps and basal trunks of bur oak was considered in more detail in the next succeeding set of photographs.) What was substantially different in understories of bur oak forest versus grove was in herbaceous species and tree species other than bur oak (see again preceding caption). Mature sexual shoots of Canada wildrye, the overall dominant herbaceous species in the bur oak grove, with conspicuous spikes were present in the first of these two p;hotographs. Also prominent in this first slide was the pioneering, annual composite, giant ragweed. The second slide presented a fullsun shot of a basal sprout of bur oak at edge of the grove where frequent mowing had killed out the native Canada wildrye and its replacement with the introduced Eurasian Kentucky bluegrass. Cherry County, Nebraska. Late June, estival aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Kuchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001). |
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115. Living stumps- Two stumps of bur oak with prolific and vigorous sprouting or suckering. These trees were at edge of a bur oak-dominated forest in the Nebraska Sandhills. Stump sprouting is an adaptation to fire, a key abiotic factor of grasslands. Bur oak is one of the most fire-adapted Quercus species in North America with sprouting from stumps or injured tree trunks one of its most important means of regeneration. In fact, bur oak sprouts prolifically even from old trees and those not subjected to injury. This was shown in several preceding photographs for bur oak growing in deep shade and fullsun environments. Bur oak has long been recognized for its "thick fire-resistant bark" (Burns and Honkala, 1990), but prolific sprouting (suckering) is an accompanying adaptation to recurrent fire that was evolved in the generally fire-prone habitat of this species biological range. Burns and Honkala (1990) explained that prolific sprouting was common following burning or cutting of bur oak only up to pole-size with such secondary shoots being of poor quality except those of seedlings. This latter conclusion was confusing to say the least given that shoots of seedlings are not sprouts (secondary shoots) at all but rather primary shoots derived from acorns. These same authors noted that larger trees (mature or full-grown age class?) also produced basal sprouts but that vigor and quality of these sprouts had not been evaluated relative to age, size, etc. of parent trees. The two examples shown here were pictorial evidence that seemed to contradict findings reported in Burns and Honkala (1990). The profuse (and plenty vigorous) stump sprouts shown off here were from mid-size trees (a foot or more in basal diameter). This was considerably smaller than the full-grown bur oaks presented in the immediately preceding set of two photographs, and these appeared vigorous enough too. Cherry County, Nebraska. Late June. |
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116. Fire ran through the bur oaks- At the outer edge of a grove form of a bur oak forest along a small stream through a hot fire with high flames had burnt just two to three prior to time of photograph. This forest range was in the Dissected Till Plains of the Central Lowlands physiographic province of northeast Kansas which is about the southern limit of forest dominated by bur oak. Thee two slides presented an outside-the-forest view to show physiogonomy and structure of the plant community. The lush growth of the herbaceous layer was dominated by seedlings of giant ragweed (Ambrosia trifida) with scattered plants of Canada wildrye, Canada wood nettle (Laportea canadensis), and pokeberry or pokeweed (Phytolacca americana). Giant ragweed is a pioneering annual; in fact, it is the most abundant pioneer or colonizing forb in this area. The hot fire had been a disturbance of such severity as to create an ideal habitat for invasion by giant ragweed which "eclipsed" the other herbaceous species which were perennials. These perennials had been present years before the burn, but they, too, appeared to have benefitted from the fire jucging by growth of other plants of these species in a few isolated spots that did not burn. This phenomenon was explained in captions below. The saplings in foreground of these (and subsequent) photographs were all hackberry and American elm. Almost every one of these saplings (and seedlings that were too small to show up in the slides) were top-killed by the heat of the obviously hot fire which burned with intensity adequate to partially kill lower branches of bur oak. This latter feature of the fire was shown and described in greater detail below. In this region most climax bottomland forests are dominated by various combinations of hackberry, American elm, slippery elm, and green ash (the Tolerant climax species) with old trees of such Intolerant, colonizing species as eastern cottonwood, sycamore, black walnut, and honey locust persisting into climax as associates. This is Society of American Foresters (Eyre, 1980) cover types 93 and 94. These mixed hardwood forests are the climatic climax. By contrast, bur oak forests, Society of American Foresters (Eyre, 1980) cover type 42, are a pyric climax. Differences in alluvial soils of these floodplain forest are not so different as to account for differences in forest range types. In the absence of periodic fire the bur oak forest would eventually proceed through plant succession to a hackberry-American elm climatic climax. That is, it would be the potential natural vegetation along streams in this climatic regime as interpreted from perspective of polyclimax and climax pattern theories. Or, what amounts to the same thing, bottomland forests dominated by hackberry and American elm would be the edaphic climax, the natural termination of such stream bottom seres. Either way, it is only periodic hot fires that maintain bur oak, which is Intermediate in tolerance (Burns and Honkala, 1990), as climax along stream bottoms (ie. bur oak is a fire type, a pyric climax). In monoclimax theory all such forests are postclimax to the climatic (= zonal or regional) climax of tallgrass prairie. Individuals of bur oak would undoubted persist into a hackberry-American elm climax as bur oaks are relatively long-lived. Burns and Honkala, (1990) reported that some bur oaks bear fruit up to ages of 400 years, the longest of any Quercus species in North America. Progression through plant succession from a seral bur oak-dominated forest to a hackberry or hackberry-elm climax forest was the pattern of vegetation development predicted for gallery or floodplain forests in northeast Kansas by Bellah and Hulbert (1974) and Abrams (1986). Abrams (1985) concluded that fire frequency (mean fire interval) bur oak-chinquapin oak gallery forests in the Flint Hills ranged from about eleven to twenty years with an average "somewhere between that range" but with a historic interval of two to three years. Clearly the historic frequency of two to three years would have maintained the oak forests and prevented progression to hackberry or hackberry-elm. It is likely that even an average fire return interval of roughly fifteen years in contemporary time would maintain the bur oak-chinquapin oak forest. Again, bur oak forest (SAF 42) is a fire type or pyric climax. Spring Creek, Marshall County, Kansas. Mid- June, vernal aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Kuchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Western Corn Belt Plains, Loess and Glacial Drift Plains Ecoregion 47i (Chapman et al., 2001). |
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117. Late spring after fire-Apperaance of a bur oak grove two to three months following a hot fire that effectively top-killed seedlings and saplings of hackberry and American elm that had established beneath the bur oak canopy. Fire intensity was adequate to create a disturbance that favored rapid establishment of a dense population of giant ragweed, the most common and prolific pioneering plant in this region of the Dissected Till Plains of northeast Kansas. Rapidly growing seedlings of giant ragweed, an annual composite, accounted for the greatest herbaceous cover, but there was also an abundance of Canada wildrye and pokeweed (though widely scattered with most cover being local) along with local exclusive colonies of Canada wood nettle (see photograph and caption below). At outer margin (just under forest canopy) there were large individuals of what appeared to be--without much familarity of early growth shoots--giant sumpweed (Iva xanthifolia). The first of these two photographs was a closer-in view of the bur oak at center-left in the second photograph of the preceding two-slide set. The trunk of this tree had split almost to ground level yet was still standing and healthy. Heartwood of a tree is dead tissue anyway so as long as the tree was not weakened to point of crashing and did not have heart-rot it remained as healthy is if the trunk was intact. The second of the photographs of this caption presented a summary view showing several top-killed saplings of hackberry and American elm. The rightmost bur oak (center-right) sported an old fire scar and ws in every way healthy from all outward appearances. This fire-scarred tree was featured below. Spring Creek, Marshall County, Kansas. Mid- June, vernal aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Kuchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Western Corn Belt Plains, Loess and Glacial Drift Plains Ecoregion 47i (Chapman et al., 2001). |
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118. Old and the new after a fire- Close-growing and very old bur oaks (first slide) along with seedlings of giant ragweed and this season's shoots of Canada wood neetle, pokeweed, and Canada wildrye as an herbaceous layer plus some saplings of hackberry and American elm (see especially second slide) comprised the range vegetation of a bur oak forest two to three months following a hot surface fire. In the background there were several pole-sized trees and saplings of red mulberry (Morus rubra) that formed an erratic lower tree layer. One of these saplings was shown in center midground of the second slide. The sapling in center foreground of this second photograph was one of the few young hackberries that was not top-killed by the fire. Other saplings of hackberry and American elm that were not topkilled outright had already started to "peel bark" and appearedlikely to die. The hackberry sapling featured here defied the odds and appeared ready--at this juncture anyway--to take its place ultimately as an adult in the grove of bur oak. It was explained in the immediately preceding caption that bur oak is a fire type forest. In absence of periodic fire bur oak--other than as a few persistent old-age trees--would be replaced by hackberry and American elm which would be the climax dominants on this bottomland sere. The latter two are Tolerant species whereas bur oak is Intermediate in tolerance rank (Burns and Honkala, 1990). Of course, fire is as much as a part of climate (or a consequence of climate if one prefers) as drought, wind storms, and floods. More precisely, lightening is as much part of the atmosphee as precipitation and temperature and fires from lightening with fuel made possible by climate are major factors responsible for maintenance of bur oak forests within the surrounding zonal vegetation of tallgrass parairie. Various studies such as those by Kucera (1960) and Bragg and Hulbert (1976) showed that woody plant communities, including forests, expanded in the tallgrass prairie region since European settlement (due in large part to fire suppression). Abrams (1985) estimated that the mean fire interval for bur oak-chinquapin dominated gallery forest in the Flint Hills of Kansas was two to three years and roughly eleven to twenty years with intervention by white settlers and contemporary man. Abrams (1986) documented this phenomenon and predicted that hackberry, elm, and redbud (Cercis canadensis) would gradually replace bur oak and chinquapin oak--at least in absence or reduced frequency of fire--as the climax forests for bottomland forest sites in the Flint Hills and Dissected Till Plains in northeast Kansas. Abrams (1992) issued a later report based on his earlier findings and concluded that for some areas of Kansas the presettlement vegetation of tallgrass prairie had become either a bur oak-hackberry overstorey with a hackberry-elm understorey or a chinquapin oak overstorey with a elm-redbud-chinquapin oak understorey. This analysis struck the current author as an interim report because it seemed likely to him that the ultimate state would be a climax hackberry-elm forest. Spring Creek, Marshall County, Kansas. Mid- June, vernal aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Kuchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Western Corn Belt Plains, Loess and Glacial Drift Plains Ecoregion 47i (Chapman et al., 2001). |
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119. Just another encounter with fire- Outer edge of a bur oak grove two to three months following a surface fire that burnt through this bottomland forest. Judging from fire scars and general understanding of bur oak forest and the natural envioroment of this area it was obvious that this was the most recent of many such fires in these old bur oaks. The fire had been hot enough to create a disturbance so severe that giant ragweed, the most abundant pioneer species in this area, had already established populations that pretty much excluded other herbaceous species except the climax perennials, Canada wildrye and Canada woodnettle (wood nettle). Even these shade-tolerant and already present perennial herbs appeared to have benefitted greatly form the fire. Without this fire a dense layer of bur oak leaves would functioned as mulch and largely smothered most herbaceous plants. Such a layer of large bur oak leaves (= mulch) was present in several small spots that did not burn. Giant ragweed was absent from these and Canada wildrye plants were smaller than those growing on land that had burned. Note that the lowermost branches of bur oak were largely leafless, but with enough leaves to prove that they were still alive (barely) and that heat from the last flames injured (appeared to have more-or-less killed) these smaller, lower limbs. This was an example of fire-pruning, but such defoliation did not necessarily kill these organs. It was likely that new shoots would arise from intercalary meristem on these lower branches. The sapling to right of foremost bur oak was hackberry. It leafed-out following the blaze, but was nonetheless severely injured by the heat from had to be an unusually hot fire. This one of the few hackberry saplings that was not totally top-killed and it had entire strips the total height of the shoot that already had peeling bark. All American elm sapling in the understorey were top-killed. Abrams (1985) studied fire frequency within gallery forests of northeast Kansas (the Fllint Hills) and found a natural fire frequency of two to three years and an extended mean fire interval of roughly fifteen years during time of occupation by European man. Either the natural or anthropogenic fire frequency would probably maintain bur oak groves and prevent development of hackberry-American elm forest on the forest range site featured here. The fire scar on the foremost bur oak was an old wound that was already present and in process of healing before the recent blaze. This "honor scar" was presented and described further in the next slide. Spring Creek, Marshall County, Kansas. Mid- June, vernal aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Kuchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Western Corn Belt Plains, Loess and Glacial Drift Plains Ecoregion 47i (Chapman et al., 2001). |
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120. New beauty in an scar- An old fire scar on an ancient bur oak that was renewed by a surface fire only two to three months prior to time of photograph. The tree had started started healing the wound caused from previous fire(s) and the most recent burn did not harm the new bark tht was slosly growing over the scar. This was a direct view of the scar that was shown from the side in the foremost bur oak featured in the immediately preceding two slides. Forbs growing in front of the oak were young seedlings of giant ragweed, the herbaceous species that most benefitted from the recent fire. This is the "price" ("rent" so to speak) to be paid by the fire-adapted bur oak for living in a pyric habitat in which less fire-adapted and more tolerant hardwoods such as hackberry and elms cannot survive. In absence of periodic fire hackberry, green ash, American elm, and perhaps boxelder would eventually replace bur oak through plant succession. These more tolerant species are the climatic climax whereas bur oak is a pyric climax (ie. bur oak forest is, in effect, a fire type in the overall environment of the Dissected Till Plains or, sometimes, Glacial Drift Plains of northeastern Kansas). Spring Creek, Marshall County, Kansas. Mid- June. |
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121. After the latest in a long history- Inside a bur oak grove through which a hot surface fire burned two or three months before. This bottomland forest range had developed along a small stream at edge of a tallgrass prairie in the Dissected Drift Plains of northeastern Kansas. There were several "featured attractions" in these two photogrphs. The most prominent of these was the large fire scar on one bur oak trunk that extended from the ground to over 15 feet in height. this was an old scar from previous fire(s) that was "freshened up" by the most recent burn. Another, though much smaller, fire scar on another bur oak tunk was presented and discussed in the immediately preceding photograph and caption. The current author interpreted bur oak forests in the Dissected Till Plains as a fire-climax type with bottomland forests on this forest site in absence of recurrent fire developing through plant succession to a climax hackberry-American elm forest. Effects from fire wounds and scars on bur oak are probably not know with any certainity. Entry of disease pathogens (eg. various forms of fungal rot) and insects through tree wounds, including those caused by fire, is always a possibility. One common suspicion--though perhaps without scientific proof--is that fire (heat or charring of wood) has something of a cauterizing effect so that disease entry is less likely with fire wounds/scars than with similar wounds (eg. those ranging from ax blazes to debarking by falling trees). The charred (charcoal) surface of a fire wound, even a fresh one) is a different (drier, perhaps chemically) environment than the moist, peeled-surface wound left by mechanical injury. What is certain from even brief scanning of the literature is that bur oak fire scars as records of fire regimes, especially fire frequency, have been analyzed in considerably more detail than the impacts of fire damage on tree health. It is also certain that fire in bur oak-dominated and influenced communities (forests, open groves, savannahs, even prairies) is a natural part of the habitat of this range vegetation. Furthermore, it follows that bur oak is adapted to fire and, was the species not so adapted to and tolerant of fire to the degree it is, this species would not be a member of these range plant communities. It is axiomatic that the overall impact of fire on bur oak is minimally adversive, neutral at least, and almost assuredly positive for survival in range vegetation of which it is an important member. Peterson and Reich (2001) concluded that on bur oak was a fire-resister, a designation given by Rowe (1983) to shade-intolerant tree species that suffer little or no damage from low-severity fres. Peterson and Reich (2001) explained that "[f]ire rarely killed mature bur oaks, even those in the smaller size classes ...". They (Peterson and Reich, 2001) noted further that even saplings of bur oak grew corky bark of such thickness as to protect the cambium from most fires. Burns and Honkala (1990) cited a volume of literature attesting to the thick fire-resistant bark of bur oak, which along with its general drought-tolerance, accounted for presnece of bur oak on xeric sites as well as mesic ones where bur oak was an associate on sugar maple-American beech forests. Another "featured attraction" in this pair of photographs, especially the second one, was the death (or near death) of shoots of hackberry and American elm saplings standing in stark contrast to the survival of adult trees of bur oak even with fire scars where sizable portions of their trunks were removed. This is a photographic lesson showing that it is primarily periodic fire (with browsing obviously less important) that maintains bur oak groves and forests which would otherwise develop into hackberry-elm forest with only persistent and senescing adult bur oaks (at status of associate species at most). Bur oak forests and woodland as pyric climax (a climax fire type) was discussed in above photo captions (complete with citation of relevant literature) so that further discussion was not deemed necessary or desirable in this current caption. This was an opportune point to acknowledge that in more xeric environments (eg. those to the west the Dissected Till Plains such as the Smoky Hills or Sandhills in subhumid to semiarid precipitation zones) it is probable that limited soil moisture, especially in drought, rather than fire is the primary variable responsible for maintenance of bur oak and restriction of Tolerant tree species like hackberry, elm, box elder, etc. Also featured here was presence of red mulberry (lower branch with large leaves extending downward from upper-left corner of first photograph). Red mulbery was usually observed to be a smaller tree of the second woody (lower tree) layer, but as one of the--if not the single most--consistent tree species in bottomland forests in humid and subhumid zones extending from the Cross Timbers and Central Prairies in Texas, Ozark Mountains in Missouri and Oklahoma, and through to the Great Plains in Nebraska. In southern mixed hardwood forests, such as those in northcentral Texas, red mulberry frequently grew to relatively large size (eg. 16-18 inch DBH) with straight boles. Red mulberry was found as a consistent member of bottomland forests were dominance varied from that by eastern cottonwood, sycamore, pecan, bur oak, elm, green ash, and hackberry or sugarberry and where soil texture ranged from primarily sandy to predominantly clay. In progressing northward, dominance by members of Ulmaceae goes from exclusively sugarberry (Celtis laevigata) as in north Texas to side-by-side co-dominance of sugarberry and hackberry (C. occidentalis) in northern Oklahoma to exclusively hackberry in northern Kansas and Nebraska. Red mulberry grew with an array of dominant tree species. Again this was usually, though not always, as an understorey or lower-height tree species. Species in the herbaceous layer shown in these two photographs included giant ragweed, Canad wood nettle, Canada wildrye, pokeweed, and giant sumpweed in that order based on estimated relative cover. Giant ragweed, an annual composite, was the number one pioneer species of denuded land in this area and far-and-away the dominant of the herbaceous layer except where there were local colonies of Canada woodnettle farther in the interior of this bur oak forest. Canada wildrye, a festucoid grass of the barley or wheat tribe (Hordeae or Tritaceae) that responds to disturbance as a decreasere, was able to "hold its own" against the rapid-growing and rank seedlings of giant ragweed. Giant sumpweed was present as sparse though very conspicuous individuals. Spring Creek, Marshall County, Kansas. Mid- June, vernal aspect. FRES No. 15 (Oak-Hickory Forest and Woodland Ecosystem). No Kuchler unit for bur oak forest; closest was Bur Oak Savanna (unit 81 in Kuchler, 1964, unit 72 in .Garrison et al., 1977, including map). SAF 42 (Bur Oak). In Brown et al. (1998, p. 37) would be Quercus macrocarpa Association 122.11? (say 111) of Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1. Western Corn Belt Plains, Loess and Glacial Drift Plains Ecoregion 47i (Chapman et al., 2001). |
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122. Sting after the burn- Local colony of Canada wood nettle or, sometimes, stinging nettle (Laportea canadensis) in the interior of a bur oak forest in late spring after a fire in late winter or early spring only a few months earlier. This species is widely distributed in moist forest with a species range in North America from Saskachian east to the Maritime Provinces and south to Florida and west to eastern third of Kansas. This member of the spurge family (Euphorbiaceae) packs quite a sting, condiderably more so according to Stephens (1980, p. 21) than the stinging nettles (Urtica species). Wear long pants and move through it gingerly. The author has negotiated many of a patch of Canada wood nettle and never gotten more than a tiny tingle by showing it the respect it deserves. The wood nettle was accompanied by several plants of giant ragweed. Marshall County, Kansas. Mid-June. |
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Water Oak (Quercus nigra)
Forest
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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. |
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123. 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). |
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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).. |
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124. 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. |
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125. 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). |
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126. 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). |
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127. 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). |
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128. "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). |
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129. 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). |
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| 130. 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. |
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| 131. Watery preparation-Outer parts of longer branches of water oak at peak bloom stage. Most catkins were staminate, but both sexes of these unisexual flowers were present. Quercus species are monecious. This tree was preparing for fruit production. Freestone County, Texas. Late March. |
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| 132. Male flowers- Cluster of staminate catkins in water oak. Freestone County, Texas. Late March. |
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| 133. Female flowers- Pistillate catkins of water oak showing a leader with several catkins (first slide) and detailed view of some small clusters of these catkins (second slide). Freestone County, Texas. Late March. |
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| 134. Watery production- Leader of water oak bearing a heavy yield of acorns. Water oak is in the red oak subgenus (Erythrobalanus) the taxon within which two years (growing seasons) are required for growth and maturity of individual acorns. Freestone County, Texas. Early September. |
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135. Water oak production up close- Details of leaves and acorns of water oak. Water oak leaves provide a textbook example of the spatulate (spatula- or spoon-shape) . leaf shape. From the perspective of Range Management and Wildlife Management both leaves and acorns are sources of agricultural production because foliage and fruit are feed for livestock and wildlife (browse and mast, respectively) Freestone County, Texas. Early September. |
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136. 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. |
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Savanna of Oak-Hickory Forest and
Tallgrass Prairie
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| 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. |
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137. 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). |
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Dominant, Associate, Widespread,
and Just Plain Interesting Plants
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of the Central and Southern Forests
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138. 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). |
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Cherries in the woods- Leaves and ripening fruit of black cherry (Prunus serotina). The fruit type of Prunus species is pome. Black cherry is the source of commercial cherry wood, one of the most beautiful of all North Americn hardwoods. Black cherry is not a domiinant or even associate tree species in climax oak-hickory forests, but in absence of fire it increases quickly and can overwhelm the climax dominant and associate oak and hickory species. Ottawa County, Oklahoma. Late June. |
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139. 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). |
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| 140. 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. |
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141. 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. |
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142. 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. |
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| 143. 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. |
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| 144. Flowering shoot tip of downy serviceberry- The inflorescence and two newly emergent leaves of shadbush or eastern serviceberry. Ottawa County, Oklahoma. April. |
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145. 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. |
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| 146. Inflorescence of Goat's Beard- Goats's Beard is a dioecious, but the flower clusters are similar for both sexes. Newton County, Missouri. June. |
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147. 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. |
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| 148. 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. |
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| 149. 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. |
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150. 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 |
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151. 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. |
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152. 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. |
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| 153. 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 |
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| 154. 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 |
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| 155. Lowbush huckleberry (Vaccinium vacillans)- Ericaceae Vaccinioideae Flag Springs State Park, McDonald County, Missouri. June. |
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| 156. Highbush huckleberry, squaw huckleberry, or deer berry (Vaccinium stamineum). Ottawa County, Oklahoma. June. |
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157. Bittersweet- Bittersweet or American bittersweet (Celastrus scandens) is probably the best known member of the Celastraaceae (bittersweet or staff-tree family). This liana (woody vine) can be found on tallgrass prairie, particularily where associated with larger woody plants on which it can twine and in protected habitats (brush fencerows are a bittersweet favorite). More commonly bittersweet grows in open oak-hickory forest and the oak-hickory forest-tallgrass savanna such as in the Prairie Peninsula. The individual shown here (first slide) was growing on a persimmon sapling, and without inflicting damage on its supporting host. Fruit presented in the second slide was immature. The examples presented were obviously female plants of this typically dioecious species. This woody vine was one of the first to be planted as an ornamental pioneers and second-stage farmers because wives of badkwoodsmen could dig them up or plant cuttings "free for the taking" throughout much of eastern North America plus the flaming red or bright orange fruit persist throughout much of the winter to add a touch of color in the bland season. Vines (1960, p. 660) mentioned that many kinds of wildlife, including some of the most valuable upland game and song bird species consumed the brillantly colored fruit which, incidentially, is usually interpreted as a capsule. Steyermark (1963, p. 1010) reported that white-tailed deer readily ate the leaves of this liana. Springfield Plateau portion of Ozark Plateau (Highlands), Ottawa County, Oklahoma. July. |
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158. Bittersweet fruit- Mature fruit of climbing or American bittersweet. Bittersweet fruit is a globose capsular fruit that breaks open as three valves exposing fleshy seeds covered with arils (Steyermark, 1963, p. 1010). This brightly colored covering of the seed, the aril, "an outgrowth of the hilum which takes the form of a partial covering around a seed" (Smith, 1977, ps. 161, 289). This detail was visible in the second of these two slides. Springfield Plateau portion of Ozark Plateau (Highlands), Ottawa County, Oklahoma. October. |
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159. Burning bush (but flaming colors and Moses were nowhere to be found)- Burning bush or eastern wahoo (Euonymus atropurpureus) is another member of the Celastraaceae (bittersweet or staff-tree family) indigenous to the eastern deciduous forest, including the oak-forest forest association at its western "vegetational frontier" adjoining tallgrass prairie. Eastern wahoo is a smaller understorey shrub that is usually shorter and smaller in trunk diameter than more common shrubs like flowering dogwood, redbud, and shadbush. Burning bush does best on more moist and fertile soils like bases of bluffs along creeks and other alluvilal habitats. It is widespread, however, including occurrence on prairies. Steyermark (1963, p. 1008) reported that eastern wahoo was native to every county in Missouri. Vines (1960, p. 661) and Fernald (1950, p. 983) gave the range of this species as extending from Montana eastward to Ontario and southward throughout southeastern North America to the Piney Woods of east Texas and into the central outliers of the deciduous forest in Kansas and Nebrasak. This shrub whould be more widely cultivated as a native ornamental. It is interesting as well as attractively colorful. Western edge of Ozark Plateau (Springfield Plateau section), Ottawa County, Oklahoma. June. |
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160. Tiny floral sparks- The small, purple flowers of burning bush or eastern wahoo. Vines (1960, ps. 661, 1032) described the inflorescence of eastern wahoo as consisting of seven to fifteen flowers, each perfect, borne on slender peduncles in axillary, trichotomous (three-branched or tri-forked) cymes. Western edge of Ozark Plateau (Springfield Plateau section), Ottawa County, Oklahoma. June. |
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161. Nifty fruit, and finally some firey color (but still waiting for Moses)- The brightly colored fruit of Euonymus atropurpureus showed basis of the common name of burning bush. This fruit type is a lobed, loculicidal capsule (Fernald, 1950, p. 983) with a braightly colored arils as was described above for bittersweet, the first member of the staff-tree family presented above. In a good crop year the colorful fruits of this shrub are so abundant that the logical origin of the common name, burning bush, becomes apparent. Wahoo was apparently derived from the language(s) of some tribes of American Indian from the meaning of "arrowwood" after a major aboriginal use of shoots of this shrub. Western edge of Ozark Plateau (Springfield Plateau section), Ottawa County, Oklahoma. October. |
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162. Black oak buddies- Poverty oaktgrass (Danthonia spicata) and big bluestem (Andropogon gerardii) comprised most of the herbaceous layer of a black oak that was properly managed. The owner of this forest range jealously protected his native forage from overgrazing which was the history (and in the not-too-distant-past) of this forest range. Poverty oatgrass( the lighter green foliage with brown [mature] sexual shoots) is a cool-season species that had almost completed its annual growth cycle. Big bluestem (the taller, bigger diameter bunch in left foreground) is a warm-season--and a short shoot--species that was still phenological stage just prior to shoot elongation and preboot stages. Due to the considerably greater mature size of individual plants of big bluestem, this the sole dominant and single most important herbaceous species (obviously a decreaser) of this range type, will yield a disproportionately greater part of total herbage at peak standing crop. The oatgrass had done all it could do, but the big bluestem had just started to come on. Poverty oatgrass has been interpreted variously as an increaser or invader. In experience of this author (who had observed this range and neighboring ones for over 50 years), poverty oatgrass is an increaser and a species that most landowners would be more than happy to see. (It is certainly superior to alien annual grasses and pioneer forbs that characterize most small tracts of black oak forest.) Poverty oatgrass is increaser on range sites where big bluestem, three-flower melic (Melica nitens), Indiangrass (Sorgastrum nutans), Canada or wood brome (Bromus purgans), and various tickclovers (Desmodium spp.) are decreasers; purpletop (Tridens flavus) and rosin weed (Silphium integrifolium) are increasers and invaders include a "zillion" annual and perennial composites. Ottawa County, Olahoma. June. Phenological stage: grain-shatter in poverty oatgrass, pre-shoot elongation in big bluestem. |
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| 163. 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. |
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| 164. 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. |
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| 165. 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. |
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166. 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. |
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| 167. 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. |
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| 168. 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. |
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169. 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. |
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170. 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. |
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171. Local stand of nimblewill- This small colony or population of nimblewill shoots (along with a few shoots of Carex molesta) was growing along the edge of an oak-hickory Ozark forest. This provided an example of typical habit and morphology of this eragrostoid grass. Nimblewill is not a heavy or high herbage-producer, but it is one of the best adapted native grasses to stress caused to a combination of defoliation and shade. Ottawa County, Oklahoma. July; immediate pre-bloom stage of phenology. |
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172. Shooting in the shade- - Shoot of nimblewill growing in its common and, apparently, "prime habitat" in the Ozark Plateau. In shade of post and blackjack oaks, black cherry, and hackberry and amid blackberry canes this nimbleweill plant had produced a robust (for this species) shoot. Shoots of nimblewill are, as revealed here, distinctive among grasses. Ottawa County, Oklahoma. July; immediate pre-bloom stage of phenology. |
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173. Shooting a panicle- Sexual shoot of nimblewill with largely exerted panicle. Typically the panicle of nimblewill remains partly enveloped in the boot (encased in the leaf that subtends the inflorescence) so that branches off of the central floral axis appear appressed and ascending so as to resemble a semi-condensed panicle. Traditionally the fruit of Muhlenbergia species has been interpreted as an achene rather than a caryopsis. Spikelets in this panicle were pre-anthesis. Ottawa County, Oklahoma. July; early summer. |
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174. 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. |
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175. 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. |
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| 176. 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. |
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| 177. 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. |
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178. Trout lily (Erythronium americanum)- This even showier species is more restricted in its environments in the Ozark Plateau oak-hickory forest than is E. albidum. The trout lily featured here was growing on a bench on the north slope of limestone bluffs that supported a sugar maple-bitternut hickory (Acer saccaharum-Carya cordiformis) mesic-limestone forest (Oak-Hickory Forest- II).. On bluffs above Modoc Creek, Ottawa County, Oklahoma.April. |
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| 179. 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. |
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| 180. Walking up on wake robbin- Two succcessively closer views of T. sessile in the western Ozark Highlands. These fine specimens were growing on the boggy soil of a green ash (Fraxinus pennsylvanica)-sycamore (Platanus occidentalis) wet bottomland forest (see Oak-Hickory Forest-II). |
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181. May apple or American 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. |
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182. Harvest time for may apple- Fruit-bearing and senescing shoots of may apple on the floor of an oak-hickory forest in the Ozark Plateau. The annual cycle for these plants was just about over. May apple are among the first perennial plants to green up in the spring as they must accomplish much of their food production and storage in preparation for winter dormancy before the closed-crown canopy of these dense woods precludes much by way of photosynthesis. This becomes even more critical when--on rare occasion (a notable uncommon phenomenon)--may apples produce a bumper crop of berries as was the case shown here. Leaves of Virginia creeper, the dominant herbaceous species in this forest, were visible on the outer edges of these photographs. Ottawa County, Oklahoma. Early July. |
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183. Berry on the bottom- A ripe fruit of may apple still attached to the crotch or point of branching in this species was presented in the first of these three photographs. The second and third slides showed the fruit cut open with seeds remaining attached and then with seed cluster removed and set aside, respectively. May apple fruit is a berry. Though edible, this berry is pretty much tasteless (unless one is extremely hungry-- as in starving). A bona fide hillbilly would eat his specimen. The one who gave you these shots fulfilled this hill culture requirement. Ottawa County, Oklahoma. Early July. |
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184. 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. |
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| 185. 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. |
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186. 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. |
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187. 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. |
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188. Some other seasoning to go with oak and hickory- Wild ginger (Asarum canadense) grows farom New Brunswick westeward to Minnesota and south through Kansas and Oklahoma to Arkansas and Alabama thereby encompassing most of the eastern deciduous forest formation in its range. Its best habitat is rich wooded slopes, especially calcareous soils associated with bluffs like the ones above Lost Creek in the western part of the Ozark Plateau on which this lovely plant was growing. The common name of wild ginger indicated the use that Indians and white frontiersman made of the rhizomes (either fresh or dried) as a seasoning and substitute for the real thing (Steyermark, 1963, p. 572). On east-facing bluffs above Lost Creek, Ottawa County, Oklahoma. April. |
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189. In rocks and earth tones- Wild ginger growing amid rocks on the east side of limestone bluffs above Lost Creek in the western part of the Ozark Plateau (Springfield Plateau section). The purplish-brown color of flowers of this species blend in with fallen leaves and moist soil almost as if camoflage was the objective. Wild ginger propagates asexually from long rhizomes that give off a pungent aroma (Fernald, 1950, p. 562) which as explained in the preceding caption were used as a seasoning. On east-facing bluffs above Lost Creek, Ottawa County, Oklahoma. April, full-bloom stage.. |
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| 190. 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. |
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| 191. 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. |
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| 192. 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. |
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| 193. 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. |
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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. |
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194. 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. |
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195. 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. |
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196. 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. |
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| 197. 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. |
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198. 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. |
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| 199. 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. |
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200. 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. |
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201. Put-together morphology- Details of frond of Christmas fern shown in a three-slide sequence from general morphology of frond blade (first slide)to details of leaflets or secondary leaves (second and third slides) of the frond. Leaflets are alsso known as pinnae (pinna, singluar). Students were referred to Diggs et al. 91999, p. 178) and standard Botany texts such as Weoer et al. (1982, ps. 586-589) for explanations of morphological structure of fern fronds, rhizomes, and roots. Christmas fern is aptly named (in the northern hemisphere) as it thrives in winter when it gets the greatest amount of sunlight with leaves shed from trees and shrubs. Bank of Modoc Creek, Ottawa county, Oklahoma. Late December. |
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202. 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. |
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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. |
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203. 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. |
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204. 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. |
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205. 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. |
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| 206. Close-up of juniper hiar cap moss- This is the gametophyte (gametophytic generation) of this species. Newton County, Missouri. April. |
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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. |
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207. 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. |
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208. 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. |
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209. 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. |
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| 210. 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. |
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211. 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. |
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| 212. 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. |
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| 213. 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. |
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214. 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. |
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