Oak-Hickory Forests-IIB

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Forest Range Types of Eastern North America

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

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

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

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

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

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

Ottawa County, Oklahoma. Hiebernal aspect, December. One of numerous forms or variants of wet bottomland forest based on classification of natural communities by Missouri Natural Areas Committee (1987). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). Combination of SAF 93 (Sugarberry-American elm-Green Ash ) and SAF 94 (Sycamore-Sweetgum-American Elm). Too small for Kuchler units. Mixed Hardwood Series in Southeastern Swamp and Riparian Forest biotic community of Brown et al. (1998). Wet Bottomland Forest (Nelson, 2005, ps.153-157). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

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

Ottawa County, Oklahoma. Estival aspect, July. Designation of wet bottomland forest was determined from classification by the Missouri Natural Areas Committee (1987). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). Combination of SAF 93 (Sugarberry-American Elm-Green Ash) and SAF 94 (Sycamore-Sweetgum-American Elm). No Kuchler unit at this small scale. Mixed Hardwood Series in Southeastern Swamp and Riparian Forest biotic community of Brown et al. (1998). Wet Bottomland Forest (Nelson, 2005, ps.153-157). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

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

Ottawa County, Oklahoma. Estival aspect, July. FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). Combination SAF 93 (Sugarberry-American Elm-Green Ash) X SAF 94 (Sycamore-Sweetgum-American Elm). Mixed Hardwood Series in Southeastern Swamp and Riparian Forest biotic community of Brown et al. (1998).Wet Bottomland Forest (Nelson, 2005, ps.153-157). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

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

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

94. Ice ain't nice- Severe damage from an ice storm to trees in a green ash-sycamore wet bottomland forest in the Ozark Plateau. A late autumn precipitation event resulted when warm, moist air from the Gulf of Mexico rose over cold, dry Arctic air (a Norther) resulting in the "winter mix" of, in order, rain, freezing rain, sleet, and snow that accumulated as ice to depths of one to four inches. The heavy accumulations of ice added so much additional weight to trees, power lines, roofs, etc. that numerous of these objects crashed to the ground. Such precipitation events, known as ice storms, can cause some of the most incredible damage in Nature to trees. Locally, extreme damage to trees and shrubs may rival that from tornadoes and hurricanes. Area of impact is frequently larger than that from tornadoes. The ice storm responsible for the damage shown here and in subsequent slides cut a swath about three hundred miles long and a one hundred and fifty miles wide.

The wet bottomlad forest shown here was the green ash-sycamore community introduced above that developed around a natural spring. Most of the downed crowns and individual limbs at this site were from sycamore, the locally dominant tree, but Shummard oak, black oak, and green ash also suffered major damage.

Ottawa County, Oklahoma. Hiebernal aspect, December. One of numerous forms or variants of wet bottomland forest based on classification of natural communities by Missouri Natural Areas Committee (1987). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). Combination of SAF 93 (Sugarberry-American elm-Green Ash ) and SAF 94 (Sycamore-Sweetgum-American Elm). Too small for Kuchler units. Mixed Hardwood Series in Southeastern Swamp and Riparian Forest biotic community of Brown et al. (1998). Wet Bottomland Forest (Nelson, 2005, ps.153-157). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

95. The double-edged sword of water- A hollow and nearly-dead-but-still-standing sycamore was toppled by heavy ice accumulations resulting from late autumn ice storm caused by the atmospheric phenomenon of overriding (warm, moist air rising above cold, dry, heavy air resulting in mixed precipitation of rain, freezing rain, sleet, and snow). This old sycamore and its cohorts, including its side-by-side "sister" tree, became established and survived as facultative phreatophytes along a natural spring in the western edge of the Ozark (= Springfield) Plateau. The ice-downed sycamore was the left sycamore (leftmost, conspicuous, white-barked trunk in left midground) shown in the first slide above that introduced this local green ash-sycamore wet bottomland forest.

Ponded water from the spring was visible in the left midground of the first of these two slides. Ironically it was liquid water that permitted establishment and continued survival of sycamore and it was solid (frozen water in form of ice) that finally ended the life of the old, toppled sycamore. Details of the hollow trunk and its downfall were covered in the immediately following set of photographs and caption.

From perspective of plant succession and development of this wet bottomland forest the most obvious change in this forest community was the on-going replacement of sycamore by western hackberry. The characteristic netted or deeply furrowed bark of the latter species was distinctive on the trunk of the pole-size tree in left foreground of the first photograph and in the three centermost saplings (included the leftmost one on which the sycamore had fallen and bent). The sapling on the right of the first slide and in left midground of second slide was a sapling of Shummard oak (the same sapling in both photographs incidentially). These species dynamics were already in play several years before the old sycamore fell. They will certainly be accelerated thereafter. This pattern of species replacement (the succession of plant species) was taking place at the edge of the wet bottomland forest. In closer proximity to the pool of spring water sycamore and green ash were still the species having greatest recruitment and, thus, were an edaphic climax (in polyclimax theory), postclimax (in monoclimax theory), or a climax that developed along a gradient in which soil moisture was determinative (in climax pattern theory).

The lack of sycamore reproduction and, instead, reproduction of western hackberry was consistent with changes in the hackberry-American elm-bitternut hickory floodplain forest along Modoc Creek described in detail above. These two forest range communities were about 350 yards apart.

Ottawa County, Oklahoma. Hiebernal aspect, December. One of numerous forms or variants of wet bottomland forest based on classification of natural communities by Missouri Natural Areas Committee (1987). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). Combination of SAF 93 (Sugarberry-American elm-Green Ash ) and SAF 94 (Sycamore-Sweetgum-American Elm). Too small for Kuchler units. Mixed Hardwood Series in Southeastern Swamp and Riparian Forest biotic community of Brown et al. (1998). Wet Bottomland Forest (Nelson, 2005, ps.153-157). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

96. Hollow-grounded- Details of the hollow and almost-dead-though-still-standing sycamore shown above that was felled by heavy accumulation of ice in late autumn ice storm. Hollow trees (trees with hollow trunks), contrary to the erroneous popular view, are usually about as strong as trees with solid trunks. Not quite, but close. This is the same basic physics as the fact that under certain kinds of strain a steel or iron pipe is quite strong compared to a solid piece of steel or iron of the same metal and diameter. This is why shafts of bird feathers, the hollow culms of many grasses (think bamboo), and hollow airplane wings (used as fuel tanks with different degrees of fullness) are very strong, especially when compared to their weight. In the same manner hollow trees can stand much strain and stress. In the case of this sycamore the added weight of accumulated ice (about two inches) and accompanied with gusty winds was more than the trunk that was 9/10ths or more hollow could bear up under. In this same ice storm there were large, completely intact trees (including sycamore, American elm, and hackberry) that also crashed. It was no wonder that this over-ripe oldster ended its life cycle thusly.

There was a small, but very deep fire scar on the base of the fallen sycamore that appeared to the point of entry of decay agents (fungal mycelium were present) that permitted decomposition of the dead wood (heartwood) inside the trunk. The lower end of this fire scar was visible in the second of these photographs (at extreme left margin of the photograph about 3/8ths of the vertical distance from bottom of slide). Referral to any freshman Botany text will remind viewers that the layers tissue of the tree trunk moving from interior to exterior were pith (very little of that); xylem of heartwood; followed consecutively by living xylem, cambium, and live phloem; and, finally, bark. It was the heartwood, which in a tree of this age comprised vast bulk of the trunk, that was susceptible to decay by microorganisms and fungii. In fact, in this tree there was remarkably little sapwood and bark to keep this old fellow upright. It was a testament to the strength of hollow trees that this more-dead-than-alive, rotten sycamore stood as long as it did and that it took such extreme stress to bring it down.

Even down this sycamore remained a critical component in the forest range ecosystem of which it was part for decades. As this trunk continues to rot it will release nutrients stored in wood to enter the biogeochemical cycles of carbon, nitrogen, etc. It hollow will undoubtedly serve as a den for coon, possum, feral house cat, etc.

Ottawa County, Oklahoma. December.

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

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

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

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

100. Mature fruit of sycamore- Fruit of sycamore is a dry aggregate of achenes. The second of these slides showed numerous of these achenes close up. The native hardwood with the largest trunk diameter and second overall in size begins it growth as a tree from this small dry fruits. Another of God's miracles in the woods.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Ottawa County, Oklahoma. August.

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

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

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

Permanent spring, Ottawa County, Oklahoma. April.

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

115. Bottomland forest of western hackberry, sugarberry (Celtis laevigata), and American elm (Ulmus americana) with an understory almost exclusively of broad-leaf wood oats (Uniola latifolia= Chasmanthium latifolium) with scattered solidary plants of the shade-tolerant composite, elephant's foot (Elepohantopus carolinianus). Beautiful plant community and an example of permanent deciduous forest range in contrast to the typical transitory feature of these dense hardwood forest types.

Ottawa County, Oklahoma. September, late estival aspect. No obvious FRES or Kuchler designations as this is a smaller unit. Neither was this relatively unique community described by Braun (1950) though it corresponds to general creek and river bottom vegetation in the Ozark Mountains. SAF Cover Type 93 (Sugarberry-American Elm-Green Ash). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

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

117.Winter in the Bosque bottoms- Hardwood bottomland forest dominated by netleaf hackberry, pecan, and American elm with dominant varying from location to location. In this view the dominant was netleaf hackberry with the four foremost trees all of that species. Most trees in background were pecan (largest trees) and American elm.

The dead herbage in foreground was broadleaf woodoats, the dominant warm-season herbaceous species. The tallest dead shoots were of frostweed (Eupatorium ). The dominant cool-season herbaceous species was Canada or nodding wildrye which was featured in the next photograph. There was also abundant Texas wintergrass (Stipa leucotricha). There were scattered individuals of naturalized, annual, Eurasian grasses including rescuegrass (Bromus unioloides= B. catharticus), Japanese brome or Japanese chess (B.japonicus), cheatgrass (B. tectorum), and wild oats (Avena fatua). There were also winter, annual forbs such as shepherd's purse (Capsella bursa-pastoris) that was common at time of this photograph.

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

118. Still winter in the Bosque bottoms- This photograph was taken within a few minutes of the preceding slide, but here the dominant, cool-season herbaceous species, Canada or nodding wildrye, was featured. Almost all of green bunchgrass in foreground was Canada wildrye though some of the green clumps were Texas wintergrass. The biggest tree (leaning right) was pecan (Carya illinoinenisis). The three smaller trees far to left of this pecan were American elm. Most of the trees behind the elms were netleaf hackberry. The dominant warm-season herbaceous species was broadleaf woodoats which was featured in the immediately preceding slide.

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

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

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

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

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

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

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

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

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

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

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

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

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

125. Fruit in the forest- Terminal end of a short shoot of native (hard-shell) pecan in the Grand Prairie of northcentral Texas. End of a short shoot on a pecan in the small bottomland forest range site along Richardson's Creek.

Hunewell Ranch, Tarleton State University. Erath County, Texas. October; nearly fruit-ripe stage.

Use and Abuse of Oak-Hickory Forests

The following section dealt with management (proper and improper) of oak-hickory forests. An example of natural revegetation (reforestation) of a clearcut climax black oak-dominated forest was included above under the section, Black Oak Forest. Discussion of the even-aged (clearcutting) silvicultural method and secondary plant succession of a clearcut black oak forest was placed there because that tract of climax oak-hickory forest had been used as an example of the black oak forest cover type. Secondary succession on the clearcut was part of the forest range vegetation of that cover (dominance) type.

Presentations and discussions in this section were more generic. Examples and principles taught from them were, regardless of the dominance type, applicable across forest range cover types in the oak-hickory forest (Oak-Hickory Region)

126. Interior of an upland mixed oak (white, red, post, black)-hickory forest that has extremely lush and diverse shrub and herb layers. Understory plants include tick clover, blackberry, grape vine, Virginia creeper, poison oak (Rhus toxicodendron= Toxicodendron toxicarium), bluestem, and panic grasses. Second growth forest with excellent regeneration of climax tree species, especially hickories. Note that this excellent botanical diversity and forest regeneration is occurring under light or conservative grazing/browsing by cattle (Herefords are visible in center).

Adair County, Oklahoma. May, late vernal aspect. FRES No. 15 (Oak-Hickory Ecosystem). K-91 (Oak-Hickory Forest). SAF 52 (White Oak-Black Oak-Northern Red Oak) and SAF 110 (Black Oak) combination, dry-mesic chert forest. Oak-Hickory Series of Brown et al. (1998). Ozark Highlands- Dissected Springfield Plateau-Elk River Hills Ecoregion, 39b (Woods et al., 2005).

127. Uneven-aged management in upland oak-hickory forest- Selective cutting (= selective felling) in an Ozark Plateau chert forest dominated by black oak which was harvested for veneer to be used in making church furniture. Trees were felled in winter and this is the scene three months after logging. Most harvested trees were 70-90 years old. Note total absence of any soil disturbance or damage to young trees. Logs were carried— not drug —out by a rubber-tired skidder. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 52 (White Oak-Black Oak-Northern Red Oak) tending toward SAF 110 (Black Oak). Oak-Hickory Series of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

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

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

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

Ottawa County, Oklahoma. August. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 52 (White Oak-Black Oak-Northern Red Oak) tending toward SAF 110 (Black Oak). Oak-Hickory Series of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

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

Ottawa County, Oklahoma. Estival aspect, September. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 52 (White Oak-Black Oak-Northern Red Oak) tending toward SAF 110 (Black Oak). Oak-Hickory Series of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

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

Ottawa County, Oklahoma. Estival aspect, September. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 52 (White Oak-Black Oak-Northern Red Oak) tending toward SAF 110 (Black Oak). Oak-Hickory Series of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

Organization Note: a black oak-dominated (northern red oak,associate species) climax forest before and after clearcutting was included under Black Oak Forest in the first part of Oak-Hickory Forests-I. That arrangement was used for consistency. Photographs of the climax (essentially old-growth) black oak-northern red oak forest were taken 15 years before the clearcutting operation and had been used in Range Types of North America prior to forest harvest. Changes in range vegetation that occurred with forest recovery (secondary plant succession) following clearcutting were included with descriptions and photographs of the climax black oak-dominated forest. That closed canopy upland Ozark Highlands forest was transitory forest range. For all practical purposes a browsable/grazable understory existed in that oak-hickory forest only at seral stages and not in the climax forest.

131. Greed and amateurs in the Ozarks- A second-growth, black oak-dominated forest in the western Ozark Plateau was high-graded for furniture lumber and pallat material (and the felled trees were immature for those uses) by a group of half-assed, clueless buffons armed with barely running chainsaws. Any black oak having the diameter of a soup bowl and maybe one 2X4 in it) was cut while blackjack and post oaks of twice the size were left standing (although many of these were "barked" and otherwise scarred by fun-loving boys driving make-shift skidders through the woods). The result was near-devastation of the forest, but Mother Nature is forgiving (especially in a 40-42 inch precipitation zone).

Appearance of this degraded black-oak Ozark Highlands forest in the first third of the third growing season post-high grading was presented in this and the remainder of the following set of slides. These photographs documented the remarkable--and the remarkably rapid--recovery of this black oak forest and chronicled the resilence of this forest cover type and ecosystem in the Ozarks. This series of slides provided a photographic chronicle of secondary plant succession in the Interior Highlands of the oak-hickory forest region (Braun, 1950,ps. 164-173).

This photograph presented the general outer appearance of the high-graded, black-oak-dominated forest early in the third growing season following amateruish logging. The largest tree along the left margin was a second-growth blackjack oak. The foremost tree on the right was a second-growth post oak. Both of these trees--immature though they were for logs--were larger and older than most of the high-graded black oaks. God's mercy and resilence of this forest vegetation was readily apparent with amble regeneration of black oak along with a typical pattern of secondary plant succession that was reset to approximately mid-sere. The dominant shrub in this "photo-plot" was any of several species of blackberry (Rubus spp.) followed by smooth sumac (Rhus glabra var. glabra), winged sumac (R. copallina var latifolia), poison ivy (Toxicodendron radicans), and wild grapes (Vitis spp.). The dominant herbaceous species was broomsedge bluestem, but there was considerable cover of silky wildrye (Elymus villosus), Virginia wildrye (E. virginica), purpletop (Tridens flavus), beaked panicgrass (Panicum anceps), and occasional large clumps of big bluestem. There were some sedges (Carex spp., Cyperus spp.) present at pre-flowering stages.

Newton County, Missouri. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Chapman et al., 2002).

132. Recovery in spite of greed and amateurs- Two overall composite shots of vegetation recovering via secondary plant succession on a high-graded, black oak forest in the western Ozark Plateau early in the third warm growing season following destructive logging by a bunch of amateur woodcutters. There was almost unbelieveable regeneration of black oak along with that of tree species including Texas or Ozark hickory (Carya texana), blackjack and post oaks, black cherry (Prunus serotina), sassafras (Sassaras albidum), and American elm (Ulmus americana). Shrubs (for which high-grading was an manmade windfall) in the recovering forest vegetation included blackberry species, wild grape, smooth sumac, winged or shining sumac, and Virginia creeper (Parthenocissus quinqueolia). Dominant herbaceous species varied locally and included broomsedge bluestem, silky wildrye, Virginia wildrye, purpletop, beaked panicgrass, naturalized tall fescue (Festuca arundinacea), and big bluestem. Major forbs in the range vegetation presented in this photograph included woodland or hairy sunflower (Helianthus hirsutus), and slender mountain mint (Pycnanthemum tenuifolium).

In the forest range vegetation shown in these two photographs black oak had more overall reproduction than any other tree species, except at isolated spots where local dominants varied from black cherry to post oak to eastern red cedar (Juniperus virginiana). Overall the second most regenerate tree was Texas, black, or Ozark hickory.

Newton County, Missouri. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Chapman et al., 2002).

133. Third growing season- Revegetation of a high-graded, second-growth black oak-dominated forest in western Ozark Plateau in first third or so of third growing season following high-grading. Black and a few choice post oak logs were taken leaving blackjack oak, Texas or black hickory, elm, black cherry, and eastern red cedar. Fortunately, from an economic standpoint as well as recovery of forest vegetation most tree regeneration had been of black oak, the climax dominant, although locally black cherry, sassafras, post oak, and American elm occasionally exceeded black oak reproduction.

The local forest range vegetation presented here was a forest gap where a few "half-grown" black oaks had been felled for small logs out of a local stand that also included Texas or Ozark hickory and post oak. The gap had become a blackberry patch that was surrounded by uncut small trees. The perimeter of the gap also included, smooth sumac, winged sumac, summer grape (Vitis aestivalis), poison oak, and Virginia creeper. Herbaceous species of the gap included grasses such as big bluestem, purpletop, beaked panicgrass, broomsedge bluestem, and a few spots of southern crabgrass (Digitaria sanguinalis), each of which was locally dominant. In this photograph purpletop was the local dominant and it was featured in center foreground. Important forbs included hairy or woodland sunflower, slender mountain mint, giant ragweed (Ambrosia trifida) and tick clover (Desmodium spp.).

Newton County, Missouri. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Chapman et al., 2002).

134. Plenty of new trees and grass- Recovery via secondary plant succession of a high-graded black oak-dominated Ozark forest was shown in this photo-quadrant which offered several lessons for the student of range vegetation. In about the first third of the third warm growing season following improper logging native grasses and forbs had made tremendous recovery along with regeneration of tree species. At this local scale big bluestem was the dominant climax grass with a nice clump growing in front of a nice black oak sapling both of which were already growing at time of high-grade logging. Both of these plants were released by removal of overshadowing crowns of black oak.

Also present prior to (as well as after) logging was the eastern red cedar (behind and to right of the black oak sapling and big bluestem) and two black cherry saplings (crooked trunks at left midground; somewhat in line but slightly to farther back that the easter red cedar). Obviously, a prescribed burn (or a wild fire) was needed to improve this forest range and facilitate secondary plant succession because a fire would kill the invasive and nonsprouting eastern red cedar as well as "stimulate" the big bluestem to produce sexual shoots (ie. increase seed production of the potnetial natural climax grass for this forest cover type on this range/forest site).

Most tree regeneration in thios photo-plot was Texas or black hickory.

Newton County, Missouri. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Chapman et al., 2002).

135. Another patch in another gap- In a forest gap produced by removal of all trees from a local stand of black oaks a small-scale but diverse forest plant community developed consisting of blackberry, smooth and winged sumacs, Virginia and silky wildryes , broomsedge bluestem, purpletop, beaked panicgrass, hairy or woodland sunflower with black cherry as the main regenerating tree species. There was not much reproduction of black oak in this local gap.

Newton County, Missouri. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Chapman et al., 2002).

136. Next generation of black oak- Outstanding regeneration of black oak, the climax dominant tree species, on a high-graded oak-hickory forest in the western Ozark Plateau about one-third into the third growing season following the most recent forest harvest. There was also considerable reproduction of black cherry. Winged and smooth sumac, blackberry, and wild grape were the main shrubs in this local stand of forest vegetation. Herbaceous species included grasses (broomsedge bluestem, purpletop, beaked panicgrass, big bluestem, naturalized tall fescue), sedges (both Carex and Cyperus species), and tick trefoil (Desmodium rigidum), the forb (a legume) with the most cover.

.Newton County, Missouri. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Chapman et al., 2002).

137. Released- Recovering range vegetation in a microhabitat by stump of a black oak roughly one-third through the third warm growing season following high-grading of a second-growth black oak-dominated Ozark forest. There was no sprouting from this stump, but there was tremendous sexual reproduction of black oak, the climax dominant tree species in all directions of the mmediate microenvironment of the stump. There was also a limited selection of shrub species including blackberry and poison ivy. The local dominant herbaceous species was woodland sunflower.

The conspicuous heart rot in this rather small stump was typical of many harvested black oaks. Cause of this rot was unknown. There was not a fire scar on this stump. From a wood production perspective it was good practice to rid the forest of such trees as the logs from such trees are "pert nar" worthless, being limited to wood chip products or cheap pallats. This is particularily the case when there is so much regeneration of the climax dominant and economically most valuable tree species. Unfortunately, much of the log removal by "hatchet-happy" skidder drivers left scars on trunks of many remaining trees which provided modes of entry for future causes of heart rot and economic losses in the next wood crop.

Newton County, Missouri. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Chapman et al., 2002).

138. Baby boom- Prolific sexual reproduction of black oak, the climax dominant, in a high-graded oak-hickory in the western Ozark Plateau roughly on-third through the third warm growing season following half-assed forest harvest. The second most successful reproducing (also sexual reproduction) was black, Texas, or Ozark hickory. There was also regeneration of black cherry from seed. Other regenerating woody species included summer grape and Virginia creeper. The clumps of green grass at right side of stump were poverty oatgrass (Danthonia spicata). The stump was, of course, black oak. Tan-colored grass culms were broomsedge bluestem. The main forb was woodland or hairy sunflower. The trunk of the sapling (behind and slightly left of the stump) was flowering dogwood.

Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

Moderate Intensity, Spring Surface Fire in Oak-Hickory Forest

The following photographs and capitons provided an example of the effects of a spring wildfire of moderate intensity in the western boundary of the Oak-Hickory Forest Region. The forest was a second-growth, climax, black oak-pignut or bitternut hickory upland forest that was uneven-aged and approaching old-growth age and structure. This was a dry-mesic chert forest (Nelson, 2005, ps. ) in the Springfield Plateau section of the Ozark Uplands.The wildfire occurred in late March or early April during which period all hardwood species were still dormant. Eastern red cedar was the only tree species with leaves during this early spring period. Photographs were taken in late May and early June (approximately eight to ten weeks following the spring wildfire).

The results of this wildfire appeared to be those resulting from natural or anthropogenic ignition during a typical moist early spring (based on the author's experience with this forest range vegetation over a span of a half century). The wildfire was not of the intensity, season, etc. that would induce major disruption or drastic alteration of the oak-hickory forest ecosystem. It was not a wildfire of such severity as to cause major retrogression of the forest plant community nor, therefore, to induce secondary plant succession at lower stages along the forest sere. Even though the surface fire was a cool burn with fire intensity being relatively low for this forest range type, fuel (consisting mostly of shed leaves) had accumulated to considerable depth and extent due to prolongued absence of fire and, in addition, was quite dry such that the wildfire burned long enough to remove essentially all of the leaf/litter layer and exposed the extremely stoney surface of the soil on this upland chert forest. The surface forest fire usually left only incidental visible burn marks (eg. blackened bark), but it generated sufficient heat and/or persisted long enough to topkill plants of tree species ranging in age/size classes from one- or two-year old seedlings up to saplings of over three to four inches DBH.

White-tailed deer, squirrels, cottontail rabbits (Sylvilagus floridanus), and perhaps species of birds that might eat buds, leaves, bark as well as mast were the only grazing/browsing animals in this forest. No livestock had ranged in this forest for at least the last three-fourths of a century-- if ever (other than strays). The rough terrain of this forest tract precluded fencing. One side of this private property included bluffs above a creek. Bluffs were steep and treacherous, but there were local strips of land free enough of limestone edges that served as natural game trails through which livestock could stray. It was not economically (and probably not even physically) feasible to fence this steep upland forest. As such livestock had been excluded from this forest range which was then accessable only to wildlife species.

This second-growth forest had the species composition, structure, etc. of a climax forest with many old-age, hollow trees that formed a closed canopy. Even with heavy canopy cover the forest vegetation included a well-developed understorey with two to four layers (varying by local stands) that provided herbage and browse. This was permanent forest range in contrast to the transitory forest range of the black oak-dominated upland forest that was clearcut and described above.

139. Freshened by fire- Interior of a dry-mesic chert foret (Nelson, 2005, ps. 125-130) in the western Ozark Highlands approximately eight to ten weeks following a cool, surface fire (accidental wildfire). This was a black oak-pignut or bitternut hickory climax upland forest with well-developed layers of forest vegetation including two herbaceous layers and one (sometimes two) woody layer(s) that comprised a grazable/browsable understorey.

Other tree species included northern red oak that dominated a few local stands (these were conterminous with and ecotonal to a sugar maple-pignut hickory climax forest), post oak, chinkapin oak (Quercus muehlenbergii), sugar maple (also ecotonal to the adjoining sugar maple-pignut hickory forest), American or white elm (Ulmus ameicana), a few red or slippery elm (U. rubra), black cherry, and white ash (Fraxinus americana). Probably all age/size classes of these tree species from seedling to large sapling were present in the understorey or lower tree layer if forest gaps were included. Most regeneration of tree species was mockernut hickory and/or black oak, which in conjuction with dominance of canopy cover and total accumulation phytomass, indicatdd that these were co-dominants of this forest that was interpreted as climax vegetation.

Shrub species included the dominant flowering dogwood plus eastern redbud, shadbush or eastern serviceberry (Amelanchier arborea), and buckthorn (Rhamnus lanceolata var. glabrata) in the interior of the forest. Common woody vines included summer grape (Vitis aestivalis), Virginia creeper, and greenbriar, known also as bullbriar or catbriar, (Smilax bona-nox). Poison ivy/poison oak was present, but it was much less common than in other oak-hickory forests in this general area. At forest edge (just out of camera range and adjoining a limestone glade of big bluestem-dominated tallgrass prairie) major shrubs included smooth and skunkbush sumac.

The main herbaceous species in interior of this black oak-pignut hickory forest was pointed-leaf, sticky, or cluster-leaf ticktrefoil or, also, tick trefoil, or tick clover (Desmodium glutinosum), but there were local habitats where pokeweed, carrion-flower (Smilax herbacea), woodland sunflower (Helianthus hirsutus), white vervain (Verbena urticifolia), Indian physic or American ipecac (Gillenia stipulata), and small skullcap (Scutellaria parvula) were locally dominant or, at least, common. There were also other Desmodium species, but these were not blooming and could not be identified positively at species level. Bigbract or large-bracted tick trefoil (D. cuspidatum) was tentatively identified based on vegetative characteristics. Grasses were absent from this understorey except at forest edges--ecotonal to an adjoining glade--were big bluestem, Canada or nodding wildrye, silky wildrye, purpletop, and three-flower (ed) melic (Melica nitens) were abundant.

Tree species in the first photograph included post oak (foremost tree at left with fire-charred/blackened basal trunk), a minor component species, northern red oak (big tree at right background behind five or six saplings) associate species, black oak (large tree behind and to right of the fire-charred post oak; rightmost foreground tree behind and to immediate left of dead foreground sapling), co-dominant species; chinkapin oak, and , pignut or mocker nut hickory (largest tree; right of a waist-high rotten snag; center), co-dominant species; sugar maple (tree closest on the right to the big pignut hickory), indicator species and transitional to adjoining sugar maple-mockernut hickory forest; and Anerican elm (many of the saplings including the one in left midground to right of fire-charred post oak). Also in the first of these two slides was summer grape.

The second of these two photographs was a closer-in view of the center of forest introduced in the first slide. Two large black oak(left and right margins) and pignut or mockernut hickory (the same big tree at right of a rotten snag now barley visible) represented the two co-dominants of this dry-mesic chert forest. The sugar maple (tree to right of the previously noted pignut hickory), was an indicator of the somewhat mesic nature of this forest range site. Also in this second slide were many plants of both clusterleaf ticktrefoil and what was tentatively identified as bigbract ticktrefoil. Species in the herbaceous and low woody layer were presented and described in detail in subsequent slides.

Most regeneration presented in these two slides was pignut hickory, but black oak was well-represented. Successional status of American elm and black cherry on this forest site was unknown, but these are generally regarded and treated as seral species in climax oak-hickory forests.

Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

140. Layers in a recently burnt Ozarks oak-hickory forest- This second-growth black oak-pignut or bitternut hickory upland Ozark Plateau forest of climax composition and structure was burned by wildfire (a cool, surface fire) about eight to ten weeks prior to time of photograph (late March or early April when all hardwood species were dormant and only eastern red cedar had leaves). This slide presented the wildfire-modified structure of this forest range vegetation.

Although the surface fire was a cool burn that only occasionally left visible burn marks (eg. blackened bark) it generated sufficient heat or persisted long enough to topkill plants of tree species ranging in age/size classes from one- or two-year old seedlings up to saplings of over three to four inches DBH. Fire intensity was generally low by standards of this forest type, but fuel (primarily several years accumulation of shed leaves) was dry enough that the wildfire burned of a duration sufficient to consume almost all of the leaf layer thereby exposing the rocky soil surface.

A high proportion of oak and hickory seedlings or small saplings ranging in size up to about one or more inches in basal diameter and roughly eight to less than ten feet maximum height were topkilled, but almost all of these promptly resprouted. By contrast, saplings of American elm and black cherry up to the approximate four inch DBH-size and some up to 18 or more feet in height were topkilled. Almost all of these resprouted from the basal trunk (= "stump") or roots. The glaring difference in tolerance (survivability) to fire exhibited by climax oak and hickory species versus American elm and black cherry was in the larger size-saplings (over three to four times greater height and stem diameter) of the latter that were topkilled. Sprouting appeared to be the same among hardwood tree species because apparently about 100% of topkilled individuals of all hardwood trees resprouted. The difference was in how much more effective even a cool surface fire was in total shoot kill of hardwood trees of species other than oak and hickory. Simply put, in the same forest and when standing side-by-side much larger trees of the nonclimax, associate species were topkilled (ie. only smaller shoots like seedlings and young saplings of the dominant oaks and hickories were killed whereas shoots up to pole-size of black cherry and elm died).

Stark differences between dominant oaks and hickory versus black cherry and elms in response to fire of the same intensity, frequency, at same season (same time, same dormant stage of phenology) was treated in separate photographs later.

The response of co-dominant black oak at large sapling size was shown by the individual in left foreground. Lower limbs appeared to have been killed as they had blackened bark and did not have leaves. Higher limbs and the crown were unscathed and in full-leaf about two months following a fire that occurred dormancy in early spring. Presence of a few persistent leaves from last year on some of these lower limbs was proof that they had been alive going into winter dormancy (unless they died the previous summer which seemed unlikely). The most obvious response of this black oak to the cool wildfire was basal sprouting. The fire had been hot or lasted long enough to induce "stump-suckering" (production of long shoots from the basal trunk), but it had not--up to time of photograph--damaged upper limbs and branches in the crown.

The snag of an American elm of pole size was also representative of this species in the black oak-mockernut hickory climax forest. American elm demonstrated high sexual reproduction with many seedlings and saplings. Even when these were topkilled by cool, surface fire they promptly resprouted. Resprouts from crowns of American elm saplings were conspicuous as, for example, new leaves of one such elm sapling directly in front of the pole-size snag (at first glance it appeared that the sprouts were from the snag, but this was only an illusion produced by camera angle from which the photograph was taken). American (and a few slippery) elms grew throughout the understory of this oak-hickory forest, but there were no individuals of these species larger than small poles. Evidentally, elms had (could) not compete effectively with larger trees of oaks, hickories, and sugar maples all of which had regenerated--and were continuing to regenerate--under their own shade. Such tolerance was indicative of climax species on this forest range type. Elms were major constitutent species in a lower woody layer dominated by shrubs like flowering dogwood, eastern redbud, lanceleaf buckthorn, and shadbush, but elms could not compete with climax oak and hickory dominants nor live long enough to enter the crown canopy. American and red or slippery elm were lasting components of the shrub layer.

Other plant species in the lower layers included Virginia creper, summer grape (both seedlings and fire-induced, basal sprouts), carrion-flower, and ticktrefoils.

Mature trees in right foreground were black oaks having DBHs of 24 to 25 inches. The large tree in left background was a pignut or mockernut hickory of 23 inches DBH.

Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

141. Details of the layers and responses to wildfire- A canopy-to-ground "photoplot" in a recently burnt (eight to ten weeks prior to time of photograph) black oak-bittternut hickory climax forest forest revealed varying responses of different plant species to a cool, surface fire modifications in forest structure. The relatively cool fire that occurred during dormancy of hardwood species (late March or early April) topkilled and stimulated basal trunk ("stump")-sprouting of smaller saplings of trees along with sprouting from live shoots of some shrubs in the lower layers of range vegetation. Responses among angiosperms differed greatly, but this photographer could not find a single individual plant of any shrub species that had been topkilled. This included the woody vines of summer grape like those shown reaching into the canopy of the large black oak in left foreground. This was also the responses of lanceleaf buckthorn and shadbush. An individual of shadbush was immediately behind the large black oak (its trunk bent to the left from behind this black oak. This individual shrub was about as large as any the photographer ever found in the this vicinity. It suffered no detectable injury (did not even have a smoked basal bole) yet it had sprouted profusely from its root crown portion. Flowering dogwood apparently was affected the least of any woody species of similar size. Some smaller plants of flowering dogwood did sprout whereas some larger individuals of this species did not sprout.

This was a dry-mesic chert forest (Nelson, 2005, ps. ) in the Springfield Plateau of the Ozark Highlands that had been burned by wildfire eight to ten weeks previously.

Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

142. New sprouts in old shrubs- A cool, surface wildfire burned (late March or early April) through the dry-mesic, upland, chert Ozark forest dominated by black oak and pignut or bitternut hickory about eight to ten weeks prior to the stage of growth and recovery of forest understorey shown in these two slides. These two slides presented details of the interior of this post-burn forest following the first fire to have taken place here in 40 to 50 years. The first slife featured a large tree in right foreground that was a black oak which two vines of summer grape continued to use as a trellis following the fire. These two shoots of summer grape had responded to the wildfire by sending up new shoots from their bases even though these shoots were still alive and leafed out (clear to the top of the black oak canopy).

In center foreground of the first photograph a nice shadbush or eastern serviceberry had also responded to the fire by basal sprouting even though its crown had also leafed-out and was apparently unaffected by the fire. The full-grown flowering dogweed behind the shadbush appeared to have been completely unaffected by the fire as it had not even sent up new shoots from the trunk base (ie. no stump sprouts). This is about as large as (the maximum size reached by) individuals of flowering dogwood and shadbush growing in upland forests in this portion of the Springfield Plateau, Ozark Mountains. Did the fire induce stump-sprouting in some individuals that will prove beneficial to these old-age (as indicated by large size) shrubs? Or was as-yet unseen damage done to these shrubs such as these two examples of two species? Proliferation of new shoots from the basal trunk cannot be viewed as neutral--even if sprouting and fire are natural phenomena.

The second photograph presented the sprouting shadbush shown in center foreground of the first slide accompanied by the woody vines of summer grape. The shoots of neither of these shrubs revealed any signs of fire-damage all of which having leafed out and growing elongating leaders. The only signs of fire on shoots of these plants (besides smoked bark) was their basal sprouting response. All basal sprouts of the shadbush had been browsed by white-tailed deer so that the terminal bud of each had been removed. None of the newly sprouted shoots of summer grape had been browsed.

Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

143. Top to bottom in a fired upland forest- Forest opening (gap) in front of a small, local stand of black oak in a dry-mesic chert upland forest in the Ozark Highlands eight to ten weeks after a cool, surface fire burned through this forest range vegetation while all woody angiosperm species were still dormant (late March or early April). The large black oak had a DBH of 24-25 inches and a severely damaged crown so that this individual was long past its "prime" as a tree. Several of the larger black and northern red oaks in this climax (though second-growth) forest had moderate to severe crown damage meaning that these individuals were long past the rapid growth stage in their life cycle. Some larger trees were still-living (barely alive) snags with their crowns completely gone. If this forest was being managed for a wood crop over-ripe trees like these were "long over-due" for harvest. They probably had such heart-rot that their wood was unfit for lumber and useful only for fuelwood. Alternatively, they were extremely valuable for cavity nesting animals.

Much of the green (the vascular plant) surface layer in the gap and up to the trunks of mature black oaks was made up of seedlings and both small and larger saplings of pignut hickory and black oak (with slightly more hickory). All of these had resprouted since the wildfire (otherwise there would have been six to eight foot saplings in front of the big black oak).Other lower-growing species included pointed-leaf or cluster-leaf ticktrefoil and, probably, bigbract or large-bracted ticktrefoil (though this latter species was in pre-bloom stage and could not be identified with 100% accuracy), carrion-flower, Virginia creeper, summer grape, and seedlings of American elm.

Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

144. Regeneration of the climax- In a local stand of black oak in a black oak-bitternut hickory forest there was ample reproduction of both of these climax co-dominants following a wildfire (in late March or early April; a cool, surface fire). These two photographs showed survival of both 1) year-old seedlings and 2 of older, topkilled saplings which had respounded by resprouting. In the foreground of both photographs there was a single trunk of a mature black oak. In left midground of the first slide a mature northern red oak represented this more mesic of the two oak species. Both slides showed the abundant sexual reproduction of oak and mockernut hickory as well as that of American elm along with much cover of pointed-leaf or cluster-leaf ticktrefoil, white vervain, summer grape, and Virginia creeper. Some small trees of sugar maple were growing in the more densely shaded background (which was too dark to photograph and reveal anything other than shadows).

A high proportion of the saplings in foreground of both slides was American elm. All of these saplings that had been alive at time of the wildfire had been topkilled and resprouted by time of photographing; however, many of the elm saplings had already died (presumedly via competition with adult oaks and hickories as Dutch elm disease had not reached this tract of forest).

Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

145. Fire in the woods: past and present- Eight to ten weeks after a wildfire (cool, surface fire even though it was an accidental burn) burned (late March or early April) through this climax black oak-pignut hickory forest the spring vegetation on this forest range showed evidence of the recent fire as well as previous fires. The large fire scar at base of the center (largest) tree was unequivacol evidence of past fire(s). This tree was a pignut or bitternut hickory with a DBH of 23 inches. The tree to the left and the smaller trunk to the right of the fire-scarred hickory were black oaks.The shoot of the smaller black oak (the one on the right) showed no affect of the wildfire other than prolific and rapid growth of new basal long shoots. The leaning (to the left) tree behind the hickory and black oaks was a northern red oak. The dead sapling behind and to right of the bitternut hickory was also a northern red oak which had been taken out by extreme competition from adult trees or disease or...

There was much regeneration of the oaks and bittternut hickory along with that of American elm and summer grape. The woody shoot of an adult summer grape was growing between the hickory and left-front black oak. Much of the lower layer of prominent green leaves consisted of pointed-leaf or cluster-leaf ticktrefoil, the most abundant herbaceous species in this tract of forest.

Follow-up timber tale: the year after this wild fire and the set of slides for this section lightening killed the large pignut hickory (basal fire scar and 23 inch DBH) shown here. It was not just top-killed, but total death including of basal stump and root system. An account of vegetation dynamics in the forest gap created by opening in the canopy with loss of this tree was provided below following treatment of this wild fire.

Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

146. Burnt but still coming (or blessing of fire in an oak-hickory forest)- Synopsis shot of the understorey as well as adult trees in a black oak-bitternut hickory dominated upland forest in the western Ozark Highlands about eight to ten weeks following a cool, surface burn (wildfire) that moved through this forest vegetation while all hardwood species were still in dormancy (late March or early April). Eastern red cedar was the only woody species that had green leaves and was capable of photosynthesis at time of the wildfire.

Adult or, even, mature trees in midground were northern red oak (two dark trunks at left) and chinkapin oak (gray or pale-colored trunk at right). The partial trunk of a sapling at right corner foreground was also a northern red oak. Sapling in left corner foreground was an American elm which was not topkilled by the wildfire. The topdead sapling that had rapid growth of basal sprouts (long shoots) was a slippery or red elm. There were also large seedlings or small saplings of black cherry and sassafras that had been topkilled and quickly sprouted back. Shrubs included the dominant flowering dogwood plus eastern redbud and lanceleaf buckthorn.

Herbaceous species included the dominant pointed-leaf or cluster-leaf ticktrefoil and what was tentatively identified as bigbracted ticktrefoil. This local habitat was deeper in the forest and more densely shaded so that Desmodium species were the only herb of any consequence.

Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

147 Adult fire sprouts- Shoots sprouted from a burnt out stump of black oak decades ago and grew into four mid-size tree trunks. These black oak shoots were approaching adult size in these two photographs. In the first slide there was a fifth tree trunk of black oak (to left and rear of the four stump sprouts) that is a separate tree, a genetically distinct (genotype) tree. The four trunks of black oak that sprouted from the stump of the previous black oak that was killed by the fire are the same tree (same genotype). Each of these four tree trunks is a clone, module, or ramet (synonyms) of the original genetic individual just as the fifth black oak trunk is that of a separate, genetically unique tree. In sum, these five tree trunks (boles) are of two trees one (the older/larger) of which had four boles and one that had a single bole like a "normal" tree. The second slide showed basal portions of the stump shoots, including the hole by basees of the four boles which was left when remains of the burned off stump rotted away, proving their common point of origin by asexual reproduction which in this case was suckering, sprouting, or coppicing.

The four boles of one tree are the progeny of fire-induced coppicing. Coppicing in black oak was introduced, described, and references cited in regard thereto above in the section, Black Oak Forests. Examples of two-year-old stump suckers (long shoots that arose from the root crown and/or cut sutmp) were presented in that portion of the Oak-Hickory Forests chapter.

These two photographs were taken about eight to ten weeks after a cool, surface fire (accidental cause; late March or early April) burned off most of the understorey of a second-growth black oak-mockernut hickory upland forest in the western Ozark Highlands (Springfield Plateau section of the Ozark Plateau or Mountains). Recently resprouted yearling and two-year old seedlings and small saplings included those of mockernut hickory (the co-dominant with most regeneration overall), summer grape, Virginian creeper, pointed-leaf or cluster-leaf ticktrefoil, white vervain, small skullcap, and even a young plant of some species of wild lettuce (Lactuca sp.).

Ottawa County, Oklahoma. June (vernal aspect).

148. Sweet root sprouts- Base of a sugar maple in a dry-mesic chert upland forest (Nelson, 2005, ps. 125-130) dominated by black oak and bitternut hickory eight to ten weeks following a cool, surface wildfire (back in late March or early April). Sugar maple was one of several other hardwood tree species including chinkapin, post, and northern red oak along with black cherry, American and slippery elm, and white ash. Major shrubs included flowering dogwood, eastern redbud, shadbush or eastern serviceberry, and lanceleaf buckthorn. The main herbaceous plants were ticktrefoil (Desmodium spp.), the major species of which was . All of these species have the capacity to sprout from stumps, roots, rhizomes, root crowns, etc. In fact the only species in this forest lacking the sprouting response was eastern red cedar. The wildfire topkilled many individual plants that ranged in size from only inches-high seedlings of oaks and hickories to 25 foot tall black cherry with four inch-diameter trunks. The great majority of these topkilled plants--except for those of eastern red cedar--resprouted almost "immediately".

The small, new shoots of sugar maple shown in this slide were root suckers or root sprouts in cotrast to stem or stump sprouts of most of the other resprouting species (and individuals thereof) in this forest. Root suckering is apparently relatively rare in sugar maple, but it obviously took place in this instance. These young, living (this year's) shoots of root origin from the parent plant were themselves fire-induced resprouts from previous season's shoots thar arose from roots in absence of fire. In other words, the root shoots from this sugar maple had arisen as clones or modules in previous growing seasons. These ramets or clones of root origin were topkilled by the surface forest fire and current growth (the only living root shoots) were resprouts of fire-killed root suckers (fire-caused resprouts of normal root sprouts). Both normal (the original) root shoots and regrown root shoots following death of original root suckers were products of asexual reproduction.

Original shoots coming off of roots of the genetically unique mother tree (clones or ramets of the genet) was an example of natural cloning the same as was coppicing in black oak described above. In the case of sugar maple root shoots this is apparently one adaptation (survival feature) of the Very Tolerant sugar maple. Rather than relying strictly on food storage in a maple seed and gambling on risky, high-mortality rate sexual regeneration (reproduction) sugar maple can use photosynthate directly to reproduce asexually with fewer risk and lower mortality rate among its progeny. In this way sugar maple remained a constitutent species in the black oak-mockernut hickory climax forest.

Ottawa County, Oklahoma. June (vernal aspect).

149 Rejuvenated the base (after it killed the top): an example of a tree species with low adult fire-tolerance- Understorey of a black oak-bitternut hickory upland forest in the western Ozark Plateau about eight to ten weeks following a cool, surface wildfire in late March or early April. This was a dry-mesic, chert forest (Nelson, 2005, ps. ) that developed adjacent to a sugar maple-mockernut hickory climax north slope forest. A black cherry of pole size (about 25 feet tall with a trunk diameter over four inches) was completely topkilled, but sprouted long shoots from the root crown of basal shoot (center midground). Oaks and hickories less than half this size were unscathed, but black cherry and elm species showed much less tolerance to fire than did climax hardwood species. Seedlings and small saplings of both American and slippery elms were abundant in this portion of the forest understorey.

Shrubs visible in this "photo-quadrant" included flowering dogwood (small. resprouted seedling in right foreground), Virginia creeper, and summer grape. About the only grass present in this forest (other than at forest edge adjacent to a prairie glade) was hairy woodland brome (Bromus purgans= B. pubescens) seen here at early growth stages.

For a comparison of the higher fire-tolerance of black oak, co-dominant of this climax-composition upland forest, in contrast to high fire-susceptability of black cherry, local associate species, the next slide in this series was offered. This black cherry and the black oaks grew within about 20 to 30 steps of each other.

Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

150. Springing from and staying on newly fired ground: plus an example of a tree species with relatively high adult fire-tolerance- Interior of a western Ozark Highlands upland forest dominated by black oak and bitternut hickory eight to ten weeks following a surface wildfire in late March or early April. Many (in fact, most) individual plants of several species were topkilled even though this was a relatively cool fire.

This photograph featured several black oaks of large sapling to small pole size that were largely unaffected by the wildfire. Most such young trees lost lower limbs, but blackoak with trunk diameters greater than approximately three inches (perhaps slightly smaller) were neither topkilled nor induced to produce long shoots (= resprouts= suckers) from basal trunks or root crowns. This lack of sprouting response was evident in the two small pole-size black oaks in center of this slide. Neither of these (or other nearby) black oaks sent up new basal shoots even though the fire at this local environment burned long and/or hot enough to remove all leaf and litter material from the ground surface and burn much of the bark on trunk bases.

This relatively high fire-tolerance in black oak contrasted sharply with that of black cherry. By way of comparison/contrast readers were encouraged to "roll up" to the immediately preceding slide in which a black cherry much larger than (at least half again as large as) these black oaks was completely topkilled and promptly sprouted from the shoot base and/or root crown. All of these sapling or pole-size trees were growing within 20 to 30 steps of each other.

There were many resprouted plants were of numerous species the most common or abundant of which included seedlings and small saplings of both co-dominants, summer grape, Virginia creeper, ticktrefoil (Desmodium spp) most of which was pointed-leaf or cluster-leaf ticktrefoil, and carrior flower. Viewers should take note that the young, previous seasons' shoots of summer grape that had twined up into the foremost (foreground) black oak were killed by the surface wildfire and had resprouted from their root crown(s). As was the case with trees and other shrubs, the larger the shoot the greater the probability that the shoot would less or little affected by fire (other things being equal, of course).

Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

151. What fire wrought in the woods- Two views of an extremely instructive "photoplot" of the understorey in the interior of dry-mesic, chert, upland forest (Nelson, 2005) eight to ten weeks after a cool, surface wildfire (in late March or early April) burnt off much of the forest understorey. This climax-composition forest range was dominated by black oak-bitternut hickory (locally accompanied by northern red and chinkapin oaks, American and slippery or red elm, sugar maple,and white ash) with two species-rich shrub layers and one or more herbaceous layers. The dominant shrub was flowering dogwood with local associate shrub species varying from lanceleaf buckthorn, shadbush, eastern dogwood, Virginia creeper to summer grape.

The most obvious shrub effects due to forest fire in this forest involved the dual impacts or interaction of defoliation, first by the surface fire followed by white-tailed deer browsing. Deer must have either "camped out"on this black oak-mockernut hickory forest range or else included it as part of their daily feeding route. There were few sprouts on any individual plant of most of the shrub species that had not been browsed by deer. The one exception was the dominant shrub overall: flowering dogwood. There were several (a fairly high percentage) of dogwood plants on which some or all fire-induced sprouts were unbrowsed by deer. By contrast there were almost no fire sprouts on any plant of lanceleaf buckthorn that had not been browsed, and browsed heavily. Deer-browsing on shadbush and summer grape was intermediate between that of buckthorn and dogwood.

This author/photographer could not find a single sprout of black, northern red, chinkapin, or post oak or of bitternut hickory that had been browsed. The same feeding situation obtained for sassafras. Sprouts of the dominant oaks and hickory might as well as been steel posts or creosoted power poles as regards deer feeding. Sassafras would have been the equivalent of asbestos. The most highly preferred tree species for browse was elm, most of which were American or white elm. All plants of Ulmus spp. that sprouted following fire were browsed; usually there were few, if any, shoots of any elm that were unbrowsed. Black cherry was also highly prefered based on proportion of shoots browsed and relative degree of use on these shoots. White ash was the least abundant tree species in this forest being mostly a transitional and only occasional species from an adjoining bluff forest community of sugar maple, mockernut hickory, and white ash.White ash received very little browse use by deer, but relative rarity of this species might have been a factor in this regard (availability being one component of browsing selectivity).

Browsing selectivity on this site provided a textbook case. At left midground in the first of these two slides and featured exclusively in the second of these slides (front and center) there was a small pole-size sassafras that was topkilled and had resprouted. Less than halfway through the first spring-summer growing season this sassafras had new sprouts over five feet tall. These were untouched by deer (apparently the flavor of sassafras shoots to deer is not on par with that of the roots to human taste). There were numerous basal shoots (fire-caused resprouts) of lanceleaf buckthorn (eg. behind and to right of the sassafras) and American elm (eg. right foreground). In contrast to the untouched sassafras, buckthorn and American elm received moderate to heavy browsing by white-teiled deer. Note the almost extreme degree of use on the lanceleaf buckthorn. This feeding pattern undoubtedly reflected deer preferences and relative palatability of browse species to deer, given the same availability of sassafras, lanceleaf buckthorn, and Americn elm (ie. they stood side-by-side and/or back-to-back).

Note: another textbook example of browsing selectivity was presented later (below) in this series of slides on the climax black oak-mockernut hickory forest between hickory and lanceleaf buckthorn.

Most of the understorey in this "photo-plot" was pointed-leaf or cluster-leaf ticktrefoil. Contrary to common sense and human experience this nodulated (papiolionaceous), herbaceous legume received relatively scant utilization by white-tailed deer. There were a few plants of eastern redbud on this local site and these also received fairly limited utilization (given the heavy browsing of deer on accompanying species).

Organization Note: Examples and details of pointed-leaf or cluster-leaf ticktrefoil (known also as sticky tick-clover, sticky tick-trefoil) were presented above in the section entitled Sugar Maple Forests. Forest plants covered under Sugar Maple Forests included many of the same species found in the range vegetation of this black oak-pignut hickory forest. These two examples of those two forest types were contiguous with each other and even included a transition zone between them.

Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

152. More released understorey and resprouting hardwoods- Another "photo-plot" of the second-growth (but climax-composition) upland, chert, Ozark Plateau forest range dominated by black oak and bitternut hickory eight to ten weeks after a cool, surface wildfire (late March or early April) spread through the understorey layers of this vegetation. Pointed-leaf or cluster-leaf ticktrefoil was the dominant herbaceous species at this location. Also locally (in the forest vegetation shown in this slide) low-growing or trailing shrubs, including Virginian creeper and summer grape, made up a lower woody layer while a taller or mid-height woody layer was comprised of lanceleaf buckthorn (three small, single-bole shrubs in center fore-to midground), eastern redbud, and flowering dogwood. The foremost (and smaller buckthorn) was topkilled and resprouted vigerously, but in an exceptional example these new shoot had received no browsing by white-tailed deer.

Large trees in background were mockernut or bitternut hickory (right margin) and black oak (left side).

Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

153. Fire affects: some sprouted; some did not- Understorey in interior of a black oak-bitternut hickory upland chert forest in the Ozark Highlands about eight to ten weeks following a cool, surface burn (wildfire in late March or early April). This was a second-growth forest range with the species composition and structure of this climax forest type (for this forest site). Other important tree species included northern red, chinkapin, and post oaks along with sugar maple, black cherry, American and slippery or red elm, and white ash.

Major shrub species were present in this view featuring the understorey. Flowering dogwood was the dominant shrub species overall although locally shadbush or eastern serviceberry, eastern redbud, and lanceleaf buckthorn were dominant. Summer grape and Virginia creeper were dominant and almost universally present woody vines which following the wildfire resprouted to form a ground surface or lower woody layer.On this local site, resprouted seedlings and small saplings of American elm were associated with sprouts of these species of woody vines (lianas). (Fire-induced sprouts of elm seedlings and saplings were abundant throughout the field of view in this photograph.)

In the forest range vegetation presented here a seedling of white ash, a relatively rare species in this forest tract, had resprouted (lower left foreground). An example of a resprouted, young flowering dogwood (probably a two- or three-year-old seedling) was in right foreground.

The large sapling or small pole in center midground was a black oak that was unfazed by the wildfire and, hence, did not growh stump suckers (fire-induced long shoots).

Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

154. Pyrogenic variety- "Photoquadrant" of the lower layer of the understorey of the second-growth, upland black oak-bitternut hickory forest with species composition and structure of climax vegetation at about eight to ten weeks following a cool, surface burn (wildfire in late March or early April). Main purpose of this photograph was to present the various herbaceous and woody species comprising the lower zone (vertically) that was typical of the outer zone (vs. interior) of this forest range vegetation. Herbceous species included carrion-flower, ticktrefoil (mostly pointed-leaf or cluster-leaf ticktrefoil with some large-bracted or big-bract ticktrefoil), and a few plants of hairy woodland brome. Other herbaceous range plants that were in this lower layer as well as a higher herbaceous layer included white vervain. small skullcap, pokeweed, and woodland sunflower.

Resprouts of shrubs and woody vines were also part of this lower layer (as well as the vertical layer or zone immediately above). Some of those species were greenbriar (known also as catbriar or bullbriar), Virginia creeper, summer grape, skunkbush sumac as well as seedlings and resprouts of saplings of the oaks, especially the co-dominant black oak, and mockernut hickory, the other co-dominant tree species.

Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

155. Two two-time loosers (but still in the woods)- Two saplings of American elm that were topkilled by a fairly cool, surface wildfire (late March or early April), promptly resprouted, and then were browsed by white-tailed deer. These small trees were photographed approximately eight to ten weeks post-wildfire. In that interim period these two examples of fire-induced shoot/stump sprouts (long shoots) of American elm had received moderate to moderately heavy degrees of use by deer. Note that in both of these plants (examples) there few, if any, shoots that had some browsing .

These plants were in the upper shrub layer of range vegetation in a second-growth, upland, chert forest that had the species composition and structure of climax forest of this cover type and forest site. Black oak and pignut or bitternut hickory were co-dominants with other major tree species being northern red and chinkapin oaks, sugar maple, black cherry, American or white elm, red or slippery elm, and white ash. Major shrubs included flowering dogwood, overall major species, eastern redbud, shadbush, lanceleaf buckthorn, Virginia creeper, and summer grape. Sassafras was locally important,but sporadic in its distribution in this forest tract. At forest edges smooth and skunbush sumacs were important. The most important herbaceous species was pointed-leaf or cluster-leaf ticktrefoil. Grasses were non-existent except for very infrequent individuals of hairy woodland brome.

American elm and red elm were two of the more palatable and important browse plants on this oak-hickory forest range.

Ottawa County, Oklahoma. Early June (vernal aspect); less than midway through the first growing season following topkilling by wildfire.

156. Was loosing really winning or, alternatively, is there more waiting to be lost? - A third small tree of American elm served as another example of defoliation (and comparative degree of use) of resprouts of American elm by white-tailed deer on a climax, black oak-bitternut hickory upland forest in the Spring Plateau section of the Ozark Plateau about eight to ten weeks following a cool, surface burn (wildfire in late March or early April). The wildfire topkilled this young American elm (stem diameter was less than one inch) which quickly responded by prolific production of long shoots from root crown or base of bole and, which in turn, were just about as quickly browsed (moderate to heavy degree of use) by deer.

This example served to teach another lesson in management of ranges and ask an interesting question. The observant viewer will have noted that there was one (and only one) large, long shoot that received no defoliation by deer. The developing terminal bud with its growth-regulating apical meristem on this one shoot was still intact. If this one "Lone Ranger" long shoot continued to escape defoliation it would (presumedly) quickly replace the fire-killed young bole. The replacement shoot could then continue to grow and develop toward maturity; die in competition with climax oaks, hickories, and sugar maple; succumb to Dutch elm disease; or live to a ripe old age and return to the earth from which it grew. Any of these (or other) possible outcomes would depend on continued escape of this intact shoot from subsequent deer defoliation. If deer eventually browse this shoot (remove the terminal bud and apical meristem) then the fire-killed and subsequently browsed elm will likely remain a hedged bush rather than develop into a tree.

Whether a continually browsed (hedged) shrub or, alternatively, a tree capable of producing wood or habitat for birds, squirrels, and furbearers is the management goal will determine whether either final browsing outcome is desirable or undesirable. If a continual game browse range is desired then the hedged shrub would be preferred. If a fully developed, sexually mature elm tree is desired in order to propagate other elms and with the full-grown tree ultimately serving as a source of wood then escape of the remaining shoot would be preferred outcome.

If (for sake of this argument) the "sole, surviving heir" capable of developing into a replacement tree does survive it is possible (on the surface it is logical or rational to assume) that browsing by deer was a net benefit to this individual (unique genetotype of) American elm. This is plausible because browsing could have equated to "pruning by Mother Nature" in a way that facilitated more rapid growth, development, and sexual maturity of the remaing shoot. The remaining single shoot would have less stress from competition by the population--most likely an overpopulation-- of heterophyllous or long shoots (=suckers= sprouts). This in turn would increase the likelihood of survival by the tree (the original genetic individual) to maturity and successful sexual reproduction. It was conceivable that deer browsing amounted to beneficial or, perhaps, even essential pruning.

On the other hand, it was also quite likely that deer would "make another call" of this recovering American elm and remove the last intact shoot thereby reducing the former single-stem sapling to a continually cropped, circular hedge (ie. a persistent shrub with axesual reproduction the only alternative to regeneration). Time will tell. Another excuse to visit the woods.

Ottawa County, Oklahoma. Early June (vernal aspect); less than midway through the first growing season following topkilling by wildfire.

157. Leaves still at the top, but now at the bottom too- Two plants of lanceleaf buckthorn that featured selectivity by two defoliators in tandem. In a climax black oak-bitternut hickory upland forest in the Springfield Plateau section of the Ozark Plateau (Highlands) a cool surface burn (wildfire in late March or early April) induced basal stem or trunk ("stump")-sprouting of numerous species many of which were subsequently browsed (sometimes heavily) by free-ranging white-tailed deer.

In the first example (slide) a nice, single-trunk lanceleaf buckthorn exhibited no signs of the wildfire other than profuse sprouting from the trunk base or, perhaps, the root crown. In the second exampe (slide) a much smaller, two-trunked lanceleaf buckthorn had prominently charred and heat-cracked bark on its boles beside numerous long shoots (=suckers=sprouts) that were induced by the fire. This smaller buckthorn was, however, also not topkilled. It, too, showed no signs of fire-damage other than burnt bark and basal sprouting.

The second of these photographs furnished another lesson in browsing selectivity. A resprouted seedling of mockernut or bitternut hickory with zero defoliation (no deer-browsing) was growing beside the browsed lanceleaf buckthorn (hickory was in front and to the left of the buckthorn). It was mentioned previously that the author could not find any seedlings, fire-induced resprouts, or saplings of oak and hickory species, climax tree species, whereas buckthorn, apparently a climax shrub species, had been consistently consumed by deer (frequently at fairly high degrees of use). In the concept of selective browsing the hardwood tree species had low or poor palatability for deer and, for their part, deer demonstrated their preference for buckthorn and their lack of preference for mockernut hickory. Students should pay attention to the fact that, by definition, palatability is a characteristic or feature of plants while preference is a feature of animals. Availability of a plant species, a dual function of presence of the plant species at time and place that animals could utilize it, was obviously not a factor in the total combination of the phenomenon called selection. Selection or selectivity= palatability + preference + availability.

Note: another textbook example of browsing selectivity by deer was presented earlier (above) in this series of slides on climax black oak-mockernut hickory forest between sassafras and both lanceleaf buckthorn and American elm.

Ottawa County, Oklahoma. Early June (vernal aspect); less than midway through the first growing season following topkilling by wildfire.

158. When selectivity by one defoliator is enough- Two general views of the climax, upland black oak-bitternut hickory forest in the Springfield Plateau section of the Ozark Mountains described under several photographs presented above. These two slides were taken approximately eight to ten weeks following a cool, surface fire (anthropogenic wildfire; late March or early April). These photographs presented the general structure, including layering, and overall species composition of the forest range. Spermatophytes (seed-bearing plants) in this second-growth, climax-composition and structure forest were limited to angiosperms (fruit-bearing or flowering plants) except for one species of gymnosperm (spermatophytes bearing naked seeds or seeds not encased in fruits): eastern red cedar (juniper). Eastern red cedar was the only woody species with leaves present at the time of the wildfire. This conifer was also unique on this forest range site in being the only species consistently incapable of producing resprouts (long or heterophyllous shoots) from trunk or roots (root crown). Thus topkilling of eastern red cedar was synonymous with killing eastern red cedar.

Eastern red cedar grew beneath the nearly closed canopy of oak species (black, northern red, chinkapin, and post oaks), mockernut hickory, sugar maple, black cherry, and an occasional white ash and elm (American and red) and in (as a constitutent of ) a higher shrub layer made up of flowering dogwood, shadbush or eastrn serviceberry, lanceleaf buckthorn, and eastern redbub (and with summer grape and Virginia creeper) growing from ground to canopy layer. In this oak-hickory forest that had not burned in several decades (probably not in the last three or four) eastern red cedar had begun to invade.

The wildfire that rejuvenated numerous plants of various species and merely set back younger plants of other species was death to eastern red cedar. The wildfire was not hot enough to even consume juniper needles, but it was hot enough to give "complete" or "total" kill (vs. only topkill) of this nonsprouting juniper even if heat was just enough to scorch leaves. (And the dead coniferous invaders in this climax hardwood forest were "purtier than a Christmas tree" to foresters and rangemen.)

More discission on invasion and control of eastern red cedar in oak-hickory forests was provided in the next series of slides.

Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

159. Four years after- The lower woody layer and herbaceous layer(s) of a climax dry-mesic upland forest in the western Ozark Plateau four years following a lightening-set surface fire that burned theough this vegetation (in mid-spring). Plants present included seedlings and saplings (of various ages) of climax tree species including black oak, pignut hickory, white ash, American elm, and post oak as well as shoots of the shrubs redbud, flowering dogwood, shadbush, and summer grape. Many of the shoots of these shrub species were rootcrown or stump sprouts that resulted from either topkill or damage-induced growth due to the wild fire.

Herbaceous species, all of which were unaffected by the fire, included hairy wood brome (Bromus purgans= B. pubescens), pointed-leaf, sticky, or cluster-leaf tick-trefoil (Desmodium glutitinosum), carrion-flower (Smilax herbacea), black-eyed susan (Rudbeckia hirta), and heartleaf skullcap (Scutellia ovata).

Ottawa County, Oklahoma. Late July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

160. Long look for some long plants- Plants of bur-reed caric sedge (Carex sparganoides) in herbaceous layer of understorey in upland climax black oak-bitternut hickory) forest in Ozark (Springfield) Plateau. Plants of bur-reed caric sedge grew together in local populations of sparse cover. This cover was comprised of comparatively large, weak-stemmed individuals having a resultant prostrate growth-form. Such plants are easily overlooked by the casual observer. Rangemen with disciplined concentration and keen eyes should have no trouble in detecting these plants. The first of these two slides presented a typical rangeman's view from standing position. The second photograph presented two (2) typical plants of bur-reed sdege from a bent-over or mid-height distance.

Bur-reed caric sedge was the major grasslike plant in the rock-strewn soil of this comparatively xeric forest range community. The dominant graminoid of the herbaceous layer was hairy wood brome. The dominant herbaceous species was pointed-leaf, sticky, or cluster-leaf tick trefoil. Other associated forbs included black-eyed susan (Rudbeckia hirta) and heartleaf skullcap (Scutellia ovata) which were featured below.

In an unusual growing season (bitter-cold, wet winter followed by wet, warm spring and then unusually hot, dry summer with exceptional drought) bur-reed caric sedge grew larger and produced more fruit than was typical for this forest range site.Such circumstances provided fine photographic fodder and an educational opportunity for students of forest range vegetation.

This was a dry-mesic, chert forest by Missouri standards of forest community classification (Nelson, 2005, ps. 125-130).

Ottawa County, Oklahoma. Mid-July; fruit-ripe phenological stage.

161. Long caric shoots in a dry forest- Closer looks at bur-reed caric sedge in understorey of a black oak-pignut hickory upland forest. In contrast to an upright and a bent-over or leaning view (two immediately preceding slides) the current two perspectives (camera distances) presented rangemen with squatting-distance views of bur-reed sedge on a chert outcrop, dry-mesic Ozark Highlands forest.

The broad leaves of neighboring plants were American elm, which was locally an associate species to the co-dominant black oak and pignut or bitternut hickory whose leaves made almost all of the litter layer beneath the sedge.

Ottawa County, Oklahoma. Mid-July; fruit-ripe phenological stage.

162. Shoots and spikes under oaks and hickories- Basal shoots and a spike of ripening achenes (first slide) and shoots with leaves along with a spike of maturing achenes (second slide) of bur-reed cric sedge in the herbaceous layer of a black oak-pignut hickory upland chert forest in western Ozark (Springfield) Plateau. Some senescing sedge leaves were prominent in both of these photographs.

Ottawa County, Oklahoma. Mid-July; fruit-ripe phenological stage.

Ripening burrs on a drooping reed- Spikelets of achenes on spikes of bur-reed caric sedge on floor of an oak oak-pignut hickory chert upland forest in the western Ozark (Springfield) Plateau. Further details of the floral units were given in the immediately succeeding slide-caption set in which ripe achenes were presented.

Ottawa County, Early June; soft-dough stage of phenology.

163. Burred tops- Four different views of spikes of bur-reed caric sedge that developed in the herbaceous layer of a black oak-pignut hickory climax forest. Many authorities describe/define the inflorescence (whole flower/fruit cluster) type of Carex species as a spike with individual groups being spikelets and each unit of the spikelet being a floret. C. sparaganioides spikelets are androgynous meaning that staminate florets are above lower pistillate florets, that is female florets subtend the male floral units (Steyermark, 1963, ps. 311, 335; Kaul et al., 2006, p. 379-380). Distinct spacing (well-delinated spaces) among spikelets is a readily seen feature in this Carex species (Steyermark, 1963, ps. 333, 335). The rich brown to golden color of florets (spikelets) is another charactristic of this species (Fernald, 1950, p. 308).

Interestingly, the same figure in both Fernald (1950, p. 308) and Steyermark (1963, p. 333) did not show the prominent often--but not always present--bract at base or beneath lower spiklets that are present as seen here, in photographs "on the web", and illustrated in Flora of North America (Flora of North America Editorial Coommittee, 2003, p. 296).

These specimens were in a dry-mesic chert upland forest (Nelson, 2005, ps. 125-130), and bur-reed sedge was the major grasslike plant species in this Ozark Highland forest range.

Ottawa County, Oklahoma. Mid-July; fruit-ripe phenological stage.

164. Two forbs on the floor- Black-eyed susan (Rudbeckia hirta), left, and heartleaf skullcap (Scutellia ovata), right, growing of the floor of a climax black oak-pignut hickory four years following a lightening-ignited surface fire four years ago. Heartleaf skullcap was a dominant forb and black-eyed was an associate forb in this forest both before and after the fire (fire aspects had no observable impact on either forb species.

Ottawa County, Oklahoma. Late July (estival aspect).

In the heart (and bright light) of the forest- Two plants of heart-leaf skullcap of the floor of an upland, dry-mesic black oak-pignut hickory climax forest in the Ozark (Springfield) Plateau during an Extreme Drought (Palmer Index) These drought-defying rascals had grown to heights over over two and a half feet (the author has seen some over four feet tall In a "good year", meaning with wet, cool growing conditions) season and had put out large inflorescences (see upper shoot with inflorescence of another plant in the next, subsequent slide; immediately below).

Most forbs in climax oak-hickory forests grow either early in the spring before trees are fully leafed-out to produce nearly complete canopy cover or they grow at edges of the forest where they can get adequate light. By contrast, heartleaf skullcap grows deep in these forest and blooms lataer in the warm-growing under a closed canopy. The biggest and more vigerous specimens of this species, however, occur in parts of the forest floor where the crown canopy is either "thinner" or where the land surface gets longer periods of direct light (as in light specks) and/or more indirect sunlight.

Ottawa County, Oklahoma. Late June; late-bloom phenological stage.

Eat your heart out- Upper part of shoot, including inflorescence and terminal leaves, of heart-leaf skullcap in a dry-mesic, upland, black oak-pignut hickory climax forest in the western Ozark (Springfield) Plateau. This tough character was blooming during an Extreme Drought and following two preceding years of Severe Drought.

Plus, this battle-bruished plant had grown to produce this level of production with noticable leaf damage from feeding insects (note partly riddled leaves). From this author's observations, insects were the main herbivores that fed on heart-leaf skullcap. The photographer found no evidence of feeding on heart-leaf skullcap by rodents, rabbits, or white-tailed deer, all of which were plentiful on this forest range which was not grazed by livestock.

Ottawa County, Oklahoma. Late June; late-bloom phenological stage.

The plants of heart-leaf skullcap shown in the three slides of the immediately preceding slide-caption sets bloomed in a different year from plants presented in the three slide-caption sets immediately below. The latter plants photographed in late May were in earlier stages of bloom progression than the plants presented above that were blooming in Late June.

Note that in the Late June plants individual flowers in bloom were near the top of the inflorescence (flower cluster) whereas individual blooming flowers in the Late May plants were near bottom to midway up in inflorescences. Blooming in Scutellia species is indeterminate: flowering progresses upward from bottom to top of inflorescence and outward from flowers closer to rachis to those at tips of branches of flower clusters.

165. Heart of the forest- Heart-leaf skullcap (Scutellaria ovata), a dominant forb, in the herbaceous understorey of a climax black oak-pignut hickory five years after a lightening-ignited surface fire burnt across this forest range. Although there is much variation in morphological features of this species (Steyermark, 1963, ps. 1271-1272), heart-leaf skullcap is one of the largest-growing Scutelleria species in the oak-hickory forests of the Ozark Highlands. This dominant forb grew in association with such herbaceous species as bur-reed caric sedge, hairy wood brome (Bromus purgans= B. pubescens), sticky (= pointed-leaf, or cluster-leaf) tick-trefoil or tick clover (Desmodium glutinosum), and carrion-flower (Smilax herbacea).

Plants and plant parts shown in this series of photographs were growing and flowering on extremely rocky, shallow soil in the second consecutive summer of Extreme to Extraordinary Drought (Palmer Scale). Heart-leaf skullcap is reasonably drought-tolerant. (most native species are).

Ottawa County, Oklahoma. Late May; peak-bloom phenological stage.

166. Leave your heart in the Ozarks- Two views of upper shoots (of two plants) featuring the inflorescences and leaf of heart-leaf skullcap on a western Ozark (Springfield) Plateau climax upland forest, a dry-mesic, chert forest (Nelson, 2005, ps. 125-130) dominated by black oak and pignut hickory.

Ottawa County, Oklahoma. Late May; peak yet early bloom phenological stage.

167. Flowers after your heart- Inflorescences of heart-leaf skullcap--presented so as to feature individual flowers--on plants growing in the haerbaceous understorey of a climax black oak-pignut hickory upland chert forest five years following a lightening-ingited surface fire.

Heart-leaf skullcap can--in wet years--be one of the largest forbs in the interior of oak-hickory or, conversely, in dry years it is a stunted little runt, but it is almost always bright and flashy flower.

Ottawa County, Oklahoma. Late May; peak-bloom--though early bloom--phenological stage.

Side-view of heart-seeking flowers- View of flower cluster of heart-leaf skullcap as seen from the side. Th short bristles of pubescence were also visible in this slide. It was explained above that flowering pattern (progrsssion) in heart-leaf skullcap is indeterminate.

Ottawa County, Oklahoma. Late June; peak-bloom phenological stage.

Readers could handily see from these photogrphs that heart-leaf skullcap makes a nearly ideal species for folks desiring native plants in their yards and gardens. As increasing water shortages increase acceptability--not to mention desirability--of natural landscaping there will be on-going, trial-and-error experiments to find native plants best adapted for such uses. Heart-leaf skullcap is a a "real looker" in this regard. It is a comparatively large forest forb with pretty, eye-catching leaves to accompany its beautiful flowers. What is more, based on observations made on the climax upland oak-hickory forest being described here, white-tail deer do not graze this species. This woods-watcher did not find any evidence of defoliation of heart-leaf skullcap even though a small herd of white-tail deer had been attracted to and grazed some of the other herbaceous species on this previously burnt-over forest range. Finally, it was noted above that this species is quite drought-tolerant.

168. Nimble on the floor- Single plant of nimblewill (Muhlenbergia schreberi) on the floor of a dry-mesic, chert outcrop upland black oak-pignut hickory dominated forest. in the Ozark (Springfield) Plateau. This forest had burnt off (spring-season surface fire) four years prior to this growth of nimblewill. There were a few scattered plants of nimblewill in the herbaceous zone of this forest, but the dominant--though sparsely scattered--grass was hairy woodland brome.

Nimblewill is a native perennial that is probably better descriped as a bunchgrass; however nimbleweill grows adventituous roots and forms modules (clonal units) along nodes of the decumbent shoots such that it has a semi-sodforming habit (Tyrl et al., 2008, p. 136-137). In this manner nimblewill frequently grows as "dense colonies" (Steyermark, 1963, p. 160). Nimblewill has a unique habit or growth form in that early season shoots are characteristically lower-growing with wider leaf blades whereas by late late-season shoots take on a more up-right form with ascending to arching culms from which contracted panicles emerge (Hitchcock and Chase, 1950, p. 397).

Nimblewill is a common forest grass in forests of the Ozark Highland Region, and undoubtedly is a member of the climax plant community, but it is generally an increaser or even invader (when present at higher cover and density) particularily on local denuded (disturbed) areas. In facat, nimblewill is so adapted to and readily populates disturbed patches that it is a weed in lawns where domesticated turfgrasses like the bluegrasses (Poa species)or bentgrasses (Agrostis species) are preferred (Fernald, 1950, p. 169). In the author's experience and observations in the Ozark Plateau nimblewill is readily eaten by all species of livestock.

Ottawa County, Oklahoma. Late July; advanced stage of plant growth, pre-bloom.

 

169. Sex at the end (and the beginning of winter)- Acrocarpis moss (Weissia viridula= W. controversa) of the leaf-littered winter understorey of a black oak-bitternut or pignut hickory upland chert forest. Acrocarpis mosses comprise one of two major moss taxa (pleurocarp mosses being the other) in which archegonia, female sex organs (and, with sexual development, their capsules), grow at terminal ends of the stemlike "shoots" of mosses.

In mosses (the bryophytes, non-vascular spore-bearing plants, in general) the generations are the opposite of the tracheophytes (vascular plants) in that it is the gametophytic generation is haploid and both photosynthetic and absorptive of soil nutrients whereas the sprophytic generation is the diploid genertion that is dependent on the rhizoids (root-like orgns absorptive organs) of the gametophytic generation for nutrients (Cronquist, 1971, 283-284).

The plant shown here is the gametophyte that is not yet bearing the capsules of the sporophyte (sporophytic generation).

The first of these two slides presented an overall view of the aboveground moss plant growing atop a chert rock surrounded by black oak and bitternut hickory leaves on the winter-aspect surface soil of a second-growth forest (at its old-growth climax state) in the western Ozark (Springfield) Plateau. This was the same forest-- an example of Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130)-- that the vascular plants shown above were growing in.

The second slide provided a close-up (macrolens) view of the stemlike structures of the gametophyte that was shown "in full" in the first slide.

Good summary treatment of the mosses was that of Wilson et al. (1971, ps. 595-600).

Ottawa County, Oklahoma. Early January; later gametophytic stage of phenology.

The next set of slides and captions were of a second-growth, chert, upland oak-hickory forest dominated by (in approximate general order): black oak, post oak, blackjack oak, northern red oak (rarely), and Texas or Ozark hickory. Oldest/largest trees in this mixed oak-hickory forest were of mid-age (in contrast to the mostly mature trees of the black oak-bitternut or pignut hickory described immediately above). Both of these tracts of oak-hickory forest had burnt off at about the same time (late March or early April) in separate wildfires that were of about the same overall fire behavior and impact on forest vegetation. Both wildfires were generally cool surface fires that did not leave fire scars on trees that had mature bark, but that did topkill sprouting species and kill "outright" nonsprouting eastern red cedar.

The black oak-post oak-blackjack oak-Texas or Ozark hickory forest had not been grazed/browsed by livestock for about three decades. Prior to that time beef cattle had accesses to this second-growth forest range which, in earlier stages of forest recovery following logging had substantial quantities of herbage, browse, and mast. White-tailed deer ranged freely through this forest.

This tract of second-growth forest was best described when using existing cover types of the Society of American Foresters (Eyre, 1980) as a combination ("hybrid") of Post Oak-Blackjack Oak (SAF 40) and Black Oak (SAF 110). Black oak was generally the most abundant tree that commanded the most cover of the forest canopy, but post and blackjack oaks were almost as common and locally dominant (as was black oak on other local habitats). Both SAF 40 and SAF 110 have been regarded as climax or at least subclimax in the Ozark Highlands (Eyre, 1980, ps. 39, 43) where, incidentially, these types (and combinations thereof) might be edaphic, topographic, or pyric climaxes (forest types).

169. Fire at the edge- Appearance of the edge between a second-growth mixed oak (black, post, and blackjack oaks)-Ozark or Texas hickory upland forest and a remnant of degraded tallgrass prairie about eight to ten weeks following a cool, surface burn (wildfire; late March or early April). Black cherry is a limited component species of this mixed oak-hickory forest (SAF 40 X SAF 110) which has traditionally been interpreted as climax vegetation throughout much of the Ozark Plateau Region. Some plants of eastern red cedar (juniper) had invaded this forest (a major topic below).

Flowering dogwood and eastern redbud were co-dominants of the upper shrub layer. Various species of woody vines (lianas) extended from the ground layer to crown canopy: Virginia creeper, poison oak.ivy, greenbriar (mostly Smilax bona-nox), and both summer and fox grapes. Herbaceous species in the interior of this oak-hickory forest were very limited (mostly to ticktrefoils and caric sedges) so that an herbaceous layer was not present except for local colonies of Mayapple. Herbaceous species at edges of this forest and an adjoining grassland of former tallgrass parairie that was presently invaded by introduced Eurasian species, pioneer or colonizing annuals, and afew perennial broadleaf weeds. Forest-grassland edge plant communities were presented and discussed in the immediately following two photographs and caption.

There were two obvious effects of the recent wildfire on range vegetation of this oak hickory forest. The most general effect was reduction in cover of the multi-layered understorey. The visual effect of this reduction in understorey cover was due to topkilling of many (most) smaller, individual plants of shrubs and seedlings/saplings of the tree species. Essentially as many of these plants as were topkilled quickly sprouted back from basal meristem of shoots ("stumps") or root crows (ie. even those plants which had all of their aboveground, perennial biomass killed initiated new growth from perenniating buds at or below ground level).

The second and, to the eye of this photographer-rangeman, most striking or immediately visible outcome of this fire was the completely scorched (and thus "completely dead") eastern red cedar (juniper) that had invaded this ecotonal forest range site to which juniper is not native. Eastern red cedar is native to this general area (Springfield Plateau section of the Ozark Plateau or Ozark Highlands), but this conifer is not native to, not a natural component species of, oak-hickory forests, tallgrass prairie, of savannahs (ecotones) where the first two of these general range types merge or overlap.

In the Ozark Highlands and parts of the Central Lowlands (eg. Cherokee Prairie) eastern red cedar naturally exists (is a natural component of the native climax vegetation) only in habitats (eg. range and forest sites) naturally secure from fire. Such habitats include environments with shallow, stoney (low water-holding capacity) soils where plant cover is sparse and, therefore, fuel is inadequate for fires (fire regimes) that would kill the extremely fire-sensitive juniper. Such harsh, water-limited environments have fires of such limited intensity, frequency etc. that some (generally only a few) eastern red cedar survive to ages that enable them to grow large enouth to escape future infrequent, low-intensity fires.

Invasion of eastern red cedar in to climax or subclimax oak-hickory forests and tallgrass prairies is a sure sign of mismanagement of forests and grasslands (sloppy husbandry of the range, poor stewardship of the land). So much so in fact, that absence (or presence) of eastern red cedar is a "litmus test" for proper (or improper) management of forests and ranges. Death of eastern red cedar on this forest range was celebrated--Hip! Hip!, Hip! Hurray!--in more detail below.

These two photographs displayed the physiogonomy, general structure, and conspicuous species (eg. dominants) at edge of this Ozarks oak-hickory forest and tallgrass prairie in part of the Prairie Peninsula first described by Transeau (1935). The large tree in this two-slide sequence was blackjack oak. The sapling to left of this blackjack was black oak. The topkilled, smaller sapling to left of the black oak (larger) sapling and most of of the other saplings of various ages/sizes under and surrounding the black oak sapling and the adult blackjackwere saplings of Carya texana known variously as black, Ozarks, or Texas hickory. This is one of the smaller Carya species in this region. It is also one of the best adapted tree species for shallow, infertile soils, frequent drought conditions, tolerance to defoliation, and rapid regeneration on disturbed habitats. The fire-scorched eastern red cedar was obvious.

Low-stature (at time of photographs) plant species in foreground of the edge included tree seedlings, shrubs, grasses, grasslike plants, and forbs. This layer of the vegetational edge and these plants were shown and described in detail in the immediately following two-slide set. The most conspicuous species in these slides were the two forbs that were the major herbaceous species: 1) giant ragweed, the overall most abundant (apparent relative cover, biomass, density) species and 2) woodland sunflower, second most common (second greatest apparent relative cover, biomass, and density). The most common grass in edge vegetation was broomsedge bluestem, but big bluestem was well-represented (see slide set immediately below). Naturalized Eurasian grasses in this ecotonal range vegetation included tall fescue (Festuca arundinacea), orchardgrass (Dactylis glomerata), perennial ryegrass (Lolium perenne) and common bermudagrass (Cynodon dactylon). Two naturalized Eurasian legumes included white clover (Trifolium repens) and red clover (T. pratense). These naturalized introduced grasses and legumes grew at outer-most edge of the transition zone. They did not grow in either the extreme "dripline edge" or the interior of the mixed oak-Ozark hickory forest.

Ottawa County, Oklahoma. Late May (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 40 (Post Oak-Blackjack Oak) X SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry- Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

170. Details of the fired edge- Two "photo-plots" featuring species composition and general biodiversity of the outer zone of the edge of range vegetation at ecotone (transition) between mixed oak (black, post, blackjack oaks)-Ozark or black hickory, upland, chert forest and degraded tallgrass prairie about eight to ten weeks after a cool, surface fire burned through dormant range vegetation (man-caused wildfire; late March or early April; dry fuel). This was that part of the edge vegetation adjoining the grassland so that herbaceous species made up a plurality of this zone of the ecotonal range plant community.

Forbs dominated this local range vegetation. Major species (in approximate overall order of species makeup based on estimated cover and/or biomass): giant ragweed, woodland sunflower, black-eyed susan (Rudbeckia hirta), lopseed (Phryma leptostachya), Carolina elephant's foot, annual ragweed, and pointed-leaf or cluster-leaf ticktrefoil. Lopseed and elephant's foot were the most abundant forbs in the range vegetation that was recorded in the first of these two slides. The second of these slides displayed species biodiversity of this edge range vegetation. Species visible in this second photograph included saplings of tree (eg. the resprouted black cherry in upper right-hand corner), shrubs (summer grape, Virginia creeper) and forbs (giant ragweed, annual ragweed, woodland sunflower, lopseed, elephant's foot, ) in addition to the conspicuous clumb of big bluestem (left center , and

Ottawa County, Oklahoma. Late May (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 40 (Post Oak-Blackjack Oak) X SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

171. Cleaned up underneath (for now)- Interior of a second-growth, chert upland, black oak-post oak-blackjack oak-Ozark hickory forest approximately eight to ten weeks after a cool, surface wildfire (late March or early April; hardwood species dormant). There were two conspicuous effects of this wildfire. Most glaring and of most immediate importance was the easy and effective removal of eastern red cedar (an invader on this range site) from this Ozark mixed oak-hickory forest. Secondly--and of most moment from the management standpoint of browse production on this forest range--was proliferation (asexual reproduction) of long or heterophyllous shoots (=suckers= sprouts) from roots or trunk bases ("stumps") of topkilled seedlings and saplings of oaks, black or Ozark hickory, black cherry, and both American and red elms. The latter three species are not climax dominants of this forest type nor are they as tolerant of fire as the climax oak and hickory species. The fire selectively reduced these seral tree species more (to a greater relative degree) than climax plant species.

Even a cool, early spring fire when the dominant trees and shrubs (all were angiosperm species) were dormant resulted in a burn of sufficient heat and duration that seedlings and small saplings of hardwood species were topkilled. Killed "completely" (root as well as shoot) were most trees (up to large sapling size) of eastern red cedar, a non-sprouting conifer, that had invaded this forest range. Eastern red cedar was the only woody species in this forest that was "leafed out" at time of the early spring wildfire. It was also the only woody species that does not resprout (produce long shoots, suckers, sprouts). Killed crown= killed tree.

This wildfire--a convenience fire that burned out of control--"cleaned up the woods" (the initial purpose in parlance of the hillbilly who set it) and in the process rejuvenated browse species, reduced cover of seral species or, same thing, facilitated plant succession, and eliminated eastern red cedar, an invader (invasive species) that was a weed tree.

Smaller saplings in center midground and larger trees in right background were black oak; larger trees in left foreground were post oak. The largest and most common forb in the understorey in this scene was pokeweed. Most common plant species in lower layer (just above soil surface) was resprouting Virginia creeper. There was also extensive cover of summer grape and buckbrush or coralberry. Black cherry was the tree species with most seedlings in this lower layer of forest vegetation shown from this view (photo-frame).

Ottawa County, Oklahoma. Late May (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 40 (Post Oak-Blackjack Oak) X SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

172. Timber Stand Improvement- View inside a second-growth mixed oak (black, post, and blackjack oaks)-Ozark hickory on a chert upland in the Springfield Plateau section of the Ozark Highlands approximately eight to ten weeks following a cool, surface wildfire (late March or early April; eastern red cedar was the only woody species that was not dormant). Heat and/or length of burning time were adequate to topkill hardwood seedlings and small saplings and kill both root and shoot of most plants of the non-sprouting coniferous cedar (juniper). Since eastern red cedar does not sprout from roots, bases or sides of trunks, or branches (does not develop new shoots from existing woody tissue) death of existing live wood equates to death of the plant. If fire (or anything else) kills the shoot it kills the plant. Simply put: scorch the crown; kill the cedar. Over and done with.

The early spring, relatively low-intensity wildfire effectively ridded this oak-hickory forest of juniper. This included the largest plants in the forest. Saplings of eastern red cedar larger than ten feet in height and three inches in bole diameter were killed. It was all over except for the celebrating.

Fire--in this case, wildfire-- resulted in improvement in this forest stand. Fire eliminated a native tree species that was not a component species of the potential natural (climax) vegetation of this forest type. Fire, a natural factor in this forest range ecosystem (and an abiotic influence under which it evolved) removed an invasive plant species. Fire was God's and the Indians' form of brush control. In fact, fire of sufficient frequency serves as brush prevention, the preferred level of noxious plant management for forest or range. Fire is a tool in range and forest improvement.

For generations, Timber Stand Improvement was such a common and widely used term in the foresters lexicon that it was customarily shown as TSI. The Society of American Foresters (1Ford-Robertson, 1971) defined TSI as a "a loose term comprising all intermediate cuttings made to improve the composition, constitution, condition and increment of a timber stand". At time of this writing the Society of American Foresters (Helms, 1998) had decided that TSI was obsolete and used "stand improvement- an intermediate treatment made to improve the composition, strucrture, condition, health, and growth of even- or uneven-aged stands" with intermediate reatment defined as "any treatment or tending designed to enhance growth, quality, vigor, and compostion of the stand after establishment or regeneration and prior to final harvest".

Aside from the opinion of the current author-photographer that forest health is a substandard term, the concept of TSI or stand improvement (or whatever term will be used when the next SAF forestry glossary comes out) was appropriate for the effects of wildfire in this forest. Increased production of browse and herbage and elimination of eastern red cedar from this forest range were improvements by whatever term or definition was used.

The two large trees in right foreground and the one to their left in center midground were black oak, overall dominant tree species of this tract of mixed oak-Ozark or Texas hickory forest. Much of the lowest layer of vascular plants was seedlings of tree species, including black cherry, and American and red elms, though most sexual reproduction was of hickory and oaks. There was somewhat more regeneration of hickory than of the oak species. Sprouts (fire-induced shoots) of Virginia creeper, summer grape, and buckbrush or coralberry were common throughout the understorey. The grass highlighted against a dark background in the lower right corner was thin or thinseed paspalum (Paspalum ciliatifolium var. muhlenbergii=P. muehlenbergii= P. setaceum var. muehlenbergii, as now generally accepted to be a variety of P. setaceum the latter was most current at time this description was written). This grass grew only sporadically at forest edge. It could not survive in the forest interior.

Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 40 (Post Oak-Blackjack Oak) X SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

173. More Timber Stand Improvement (a dead cedar is a good cedar)- Species composition and structure of a chert, upland mixed oak-Ozark or black hickory forest in the western Ozark Highlands as apparent eight to ten weeks after burning by an early spring wildfire (cool, surface fire; late March or early April). This second-growth forest was dominated (other than at local scale= localized stands) by black oak, but post and blackjack oaks and black or Texas hickory were general associates while black cherry and, less frequently, American and slippery or red elm were local associates. Eastern red cedar (juniper) was an invader and a weed tree.

Weed tree- "1. any tree of a species having little or no economic value on the site in quetion 2. a tree of little or no economic value, more particularly when competing with one or more desirable timber trees at any stage of development and therefore due for elimination from the crop" (Helms, 1998).

It should be emphasized that for many weed species there is an ecological basis in addition to the primary economic criterion for correct designation of weed or pest. This is clearly the case for eastern red cedar as exemplified on the oak-hickory forest described here. Some weeds (a la eastern red cedar) are alien species (or, at least, species that are not indigenous to the site they have invaded), that can migrate and become established due to disturbance (human-induced or otherwise). Disturbance certainly includes lack of fire in native plant communities in which fire was a natural abiotic factor under which natural (climax) vegetation developed and ecosystems evolved.

On most oak-hickory forests eastern red cedar is a species that is not a natural constitutent of this climax vegetation. Rather this conifer is an invader, an invasive species, that can only persist on such forest range in continued absence of fire. Eastern red cedar is a weed tree--both ecologically and economically--that becomes established under exclusion of fire. Conseversely, fire prevents invasion of this woody weed and/or controls it once it becomes established (if the trees are not too large to crown-scorch).

Even though the spring burn whose impacts were described here was a cool, surface fire (and all hardwood species were still in dormancy) there was sufficient heat generated or long enough duration of burning to kill almost all of the invading eastern red cedar. The surface fire did not consume the winter-dessicated needles of this juniper species nor did the fire blacken much of the bark on trunk bases, but a mere scorching of the crowns was sufficient to kill even the largest individual junipers (biggest trees were saplings about ten feet tall with stems of about three-inch diameter).

Numerous seedlings and saplings of oaks and hickory were topkilled as were those of the less fire-tolerant black cherry and elms which lost shoots of saplings larger than those of the climax dominants. For all practical purposes, all topkilled saplings and seedlings of hardwood trees promptly sprouted back (produced long shoots from roots or trunk bases) so that while the fire killed from one to several seasons of aboveground perennial growth it did not kill these plants. No flowering dogwood and eastern redbud were topkilled except for a few seedlings and very small saplings. Likewise, none of the larger plants of these two shrub species produced basal shoots (=sprouts= suckers).

Although this wildfire burned within a week of (and less than a mile from) the fire in the black oak-mockernut hickory upland forest described in the preceding slide series, the fire in the mixed oak-black hickory forest described here was a substantially cooler fire. It appeared from investigations subsequent to this fire that it burned downhill (ie. a backing fire) and on a day during which temperatures was considerably lower. Consequently the fire in the mixed oak-Ozark hickory forest had less impact on hardwoods (trees and shrugs). There was little if any difference in impacts on eastern red cedar between the two forests (forest types). It does not take a hot or persistent fire to kill an eastern red cedar. Fortunately, this conifer is a species that like the house cat (Felis silvestris catus=Felis domesticus=Felis catus) is easily killed. Apparently animal taxonomists are in as much disagreement over proper scientific names as agrostologists are of grasses, but the relevant thing is that house cats and eastern red cedar die easily if properly attacked. A good and faithful steward will keep both of these species out of the oak-hickory forest, and kill any he finds that are already there. Death to pests!

In sum, the wildfire selectively eliminated non-sprouting eastern red cedar, reduced the undercanopy of the shrub and lower tree layers, increased browse production, and facilitated plant succession in the forest.

The largest tree trunk (left foreground) and the one to left and mostly obscured by the largest trunk were black oaks. The tree behind these black oaks and left of the left cedar was northern red oak. The two trees in center foreground and midground to the left of the right scorched cedar and the pole-sized tree behind and to right of left scorched cedar were post oaks. The lowest layer of vascular plants (just above ground level) was comprised of seedling and resprouts of tree species, Virginia creeper, buckbrush or coralberry, and summer grape.

Ottawa County, Oklahoma. Early June (vernal aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 40 (Post Oak-Blackjack Oak) X SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

174. Fire is a selective feeder- "Photo-transect" showing selective defoliation by an early spring wildfire (cool surface fire; late March or early April) in the lower (herbaceous) layer of mixed oak-Texas or black hickory upland forest in the western Ozark Highlands. Shown here was the burn scene at local habitat scale about eight to ten weeks post-fire. In center foreground a caric sedge (Carex sp?) that was either dormant had but few leaves present at time of fire was thriving. All around the sedge were new shoots and leaves of Virginia creeper, and behind the sedge was a dead seedling of eastern red cedar. This young tree had been killed by the cool, surface fire which did not generate enough heat nor send flames high enough to incinerate this cedar (juniper). Instead the crown of this baby tree was merely scorched, but that was enough to kill a small plant of this non-sprouting coniferous species.

In this same forest and from the same fire much larger cedars (saplings with up to three-inch diameter boles) were also killed with no more visible damage than scorched crowns. Examples of this effect were presented in immediately preceding slides and captions. It was conceivable that some smaller, short-stature cedars might have escaped the fire due to passage of flames and/or heat above them. Such was not the case. The flaming angel of death did not pass over smaller junipers. Instead, eastern red cedar is so sensitive to fire affects that a broad range of age/size classes of cedar in this forest were susceptible to fire-kill.

Ottawa County, Oklahoma. Early June (vernal aspect).

175. More selective feeding by fire- Local scale (microsite, microhabitat, microenvironment) assemblage of plants in a mixed oak-Ozark hickory, upland, chert forest in western Ozark Plateau. Viewers should note that the cool, surface fire was not hot enough to harm moss growing on trunk of post oak, and at time of fire the moss was quite dry (in fact it was the type of moss carried by frontiersmen in their tenderboxes for tender when starting fires with their flints). Such a cool fire was sufficient to readily kill non-sprouting eastern red cedar. The point was made again: eastern red cedar is readily killed by even low-intensity fire.

The mossy tree trunk was post oak. Most of the leaves at or just above ground level were Virginia creeper (five-leaflet leaves) and buckbrush or coralberry or, frequently, coral berry (opposite leaves aligned along stems).

Management implication: eastern red cedar, an invasive weed tree on some range types, is so easily killed by fire that the safest (least-likely to escape) surface fire imaginable is adequate to eliminate most seedlings and small to mid-size saplings of this non-sprouting conifer from oak-hickory forest (and from tallgrass prairie and oak-hickory-tallgrass savanna). Therefore, fire these ranges and fire them frequently so as to improve chances of killing eastern red cedar (cedar trees will be younger and smaller under more frequent burning).

Ottawa County, Oklahoma. Early June (vernal aspect).

Attention: Viewers attention was drawn to the fact that needles of eastern red cedar (juniper) saplings and seedlings were scourched and not consumed by the relatively cool, early spring, surface fire. Readers attention was also directed to the absence of charred bark on juniper boles and branches except for the basal-most portions of trunks. Yet, these non-sprouting junipers were killed--leaves, bole, and root. A surface fire so cool and fast-moving that it left "nary a mark" on seedling or sapling was nonetheless effective in killing easter red cedar up to sapling-size. In the words of Jackie Gleason: "How sweet it is"! Praise the Lord and pass along the drip torches.

Severe Intensity, Summer Surface Fire in Oak-hickory Forest

The following section presented results (including initial disturbance or denudation and early secondary plant succession) of a hot summer wildfire during Severe Drought (Palmer Scale) in a second-growth post-oak-black oak dominated forest in the western edge of the Ozark (Springfield) Plateau. This fire took place on the hot (ambient temperature above 100 degrees Fahrenheit; relative humidity around 30%) afternoon of 21 July. The wildfire resulted from convenience burning (household trash and dead tree leaves in a yard) in defiance of a county burn ban. As could have been expected from this wildfire-waiting-to-happen situation, a small fire that was incompletely confined "got away" and quickly engulfed a neighboring hardwood (generally a post oak-black oak) forest tract. There was a limited understorey to this forest, but accumulated dry leaves in Severe Drought under hot, low-humidity atmospheric conditions provided enough dry fuel that heat from flames was adequate to kill leaves in the tops of fifty to sixty foot oaks as well as tree saplings, shrubs, and herbaceous plants.

The man who started the fire was jailed and faced criminal charges. The results (if any) of this legal action were unknown to this author even though at the time he lived within six miles of the crime. This wildfire was in effect a case of arson (in some legal degree) given that it involved reckless behavior that led directly to property damage and violated an official burn ban. It would have been a misdemeanor not felony arson in the Missouri penal code.

The case of wildfire that resulted from arrognce, stupidity, and carelessness provided a good case study of forest regeneration, forest range management, and secondary plant (forest) succession for students out there in www.rangeland.

176. Same as crowned out- Appearance of a post oak-black oak second-growth forest 46 hours after a hot summer wildfire during Severe Drought killed leaves in crowns of tallest trees (about thirty to forty feet tall) at this local area. Although there were some blackjack oaks and, fewer chinquapin oaks (Quercus muhlenbergia), black hickory was the associate tree while blackberry (Rubus sp.) was the dominant shrub followed by flowering dogwood. In local areas of more open canopy (lower tree density), hackberry (Celtis occidentalis) and American or white elm (Ulmus americana) were co-dominants. There were also some plants of the native, but invasive (for this forest range site) tree species, eastern red cedar and bois d'arc or Osage orange (Maclura pomifera). There were also some trees of common or Virginia persimmon (Diospyros virginiana) and sassafras (Sassafras albidum) in local areas.

Overall this native vegetation was an Ozark Plateau oak-hickory forest, but it was patchwork or mosaic community which appeared to show various levcels of disturbance by the heavy hand of the whiteman.

Tree density was so great that tree crowns prevented all direct light and most brighter indirect light from reaching the lower forest levels. There was very little herbaceous cover under this dim-light condition, but there were a few plants of broomsedge bluestem, silky wildrye, pokeberry (Phytolacca americana), and naturalized sericea lespedeza (Lespedeza cuneata). Earlier in the spring to early summer there were scattered colonies of Mayapple (Podophyllum peltatum), shoots of which had died as plants entered dormancy (probably hastened by drought).

The first slide presented the outer edge or margin of this forest in which chinquapin oak grew. There were also plants of eastern red cedar and bois d'arc or Osage orange established on the perimeter of this forest tract. These two invaders of oak-hickory forest in much of the Ozark Plateau were killed by the intense heat of this wildfire. The nonsprouting eastern red cedar was "killed dead" (death of shoot equals death of plant) whereas bois d'ar was completely top-killed, but likely to resprout (at least weakly).

The second slide showed evidence of the high intensity of the fire by revealing killed green tree leaves and the complete combustion of much fuel (including saplings that burned off at ground level, then fell, and were nearly completely consumed as they lay on the ground). This fire was so hot that lower, sap-filled, live limbs burned up completely or nearly so. This baby did burn!

Newton County, Missouri. Late July. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 40 (Post Oak-Blackjack Oak) X SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

177. Sizzled understorey- Appearance of a post oak-black oak second-growth forest 46 hours after a hot summer wildfire during Severe Drought and with the afternoon air temperature slightly above 100 degrees Fahrenheit (relative humidity around 30%). Featured here was a young eastern red cedar about 12 feet tall and a smaller eastern red cedar (about six feet in height) both of which were killed while scourced needles remained on their branches. The larger (pole-sized) tree to left of the larger eastern red cedar was chinquapin oak and the smaller tree (large sapling) to right of the larger eastern red cedar was post oak. Although leaves on the chinquapin oak were still green they later died. Leaves on the post oak were burned brown. The lower bushes with scourced leaves in the first slide were shoots of buckbrush or coralberry (Symphoricarpos orbiculatus). There were also burnt canes of blackberry, the overall dominant shrub.

Newton County, Missouri. Late July. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 40 (Post Oak-Blackjack Oak) X SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

178. Still smoldering- The interior of second-growth post oak-black oak forest in the ozark (Springfield) Plateau 46 hours after a hot summer wildfire (during Severe Drought) burned the vegetation with such heat that the uppermost leaves in fifty and sixty feet tree were scourched. There were some trees of blackjack oak, black or Texas hickory, and chinquapin oak in this "photoplot" of the burnt-over forest. Also there were some small (six to about 12 feet in height) trees of eastern red cedar.

Ambient temperature at time of the wildfire was slight higher than 100 degrees Fahrenheit and with low relative humidity (30-35%). Heat of the fire itself was extreme for this kind of fuel which burned cleanly as was seen by the presence of white ash on the soil surface.

This fire was so hot that it burned down through basal tree trunks to the upper tree roots which were still burning two days later as seen in this photograph (smoke in background). There was limited understorey, but there were local colonies of blackberry, buckbrush or coralberry as well as larger shrubs of flowering dogwood. Predominant herbaceous species were broomsedge bluestem, silky wildrye ( (Elymus villosus) and the exotic forage legmue, sericea lespedeza. There were also plants of the perennial forb, pokeberry or pokeweed. Leaves of all woody species were scourced and died soon after the fire. (Tree leaves appear green in this slide, but they were already dead.)

Newton County, Missouri. Late July. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 40 (Post Oak-Blackjack Oak) X SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

179. One year after- Edge of a second-growth post oak-black oak forest at edge of a prairie in the western Ozark (Springfield) Plateau one year following a hot summer wildfire during Severe Drought that either top-killed or severely damaged crowns of the tallest trees. These three slides sonstituted a set of "nested photoplots" with the first slide being a general view of the perimeter (margin) of this forest tract while the second and third slide focused in on the "skeleton" of a fire-killed eastern red cedar which was a "landmark" in center background of the first slide. The largest green crown in the first slide (right margin) was a blackjack oak while other green crowns were those of post oak and black oak. There were also some trees of black or Texas hickory. The three principal shrubs were blackberry, flowering dogwood, and buckbrush or coralberry. In some local areas of this forest tract where there was a more open canopy, hackberry and American elm were co-dominant tree species. Likewise, there were "spots" comprised of common or Virginia persimmon and sassafras.

Students should note (in the first slide) that most of the taller trees had leafed out and from a distance appeared to be "unfazed" by the the hot wildfire of one year ago. A closer examination (see below) revealed otherwise. The fairly large eastern red cedar that was the centerpiece of this nested "photoplot" series was clearly dead. Eastern red cedar does not resprout so this tree was dead "forever and ever, amen". On this forest site, eastern red cedar is an invader (invading species) even though it is a native tree species. Fire is the main environmental factor that "purges" (more-or-less) the oak-hickory forests of the Ozark Plateau of this coniferous weed tree.

This forest tract was at the edge of a former tallgrass prairie that had been converted into cropland, the latest crop of which was the agronomic forage grass, tall fescue (Festuca arundinacea), with quite a bit of cover of sericea lespedeza. These two, introduced, herbaceous species had "come back with a vengence" following the hot summer wildfire of one year earlier. This was shown in the first slide. The second and third slides were of vegetation in the oak-hickory forest itself. These two slides featured the herbaceous species of the burnt-over forest (along with the fire-killed eastern red cedar). The major herbaceous plants were two annual composites: 1) prickly lettuce (Lactuca scariola= L. serriola) in subfamily Liguliflorae whose members secrete a milky latex sap and 2) horseweed or mare'stail (Conyza canadensis) in subfamily Tubuliflorae, the larger of the two subfamilies of Compositae (Asteraceae). These two annual composites were clearly r-selected species. In some local microsites (as shown below) there were plants of common or bull thistle (Cirsium vulgare), a naturalized Eurasian biennial composite.

Newton County, Missouri. Late July. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 40 (Post Oak-Blackjack Oak) X SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

180. Sundown on burn pioneers and other fire beneficiaries- Early evening light through a second-growth post oak-black oak forest in the Springfield Plateau revealed the harmful impact on trees and the release of an herbaceous understorey one year after a hot summer wildfire burned through this timber tract during Severe Drought. Atmospheric conditions at time of the wildfire included ambient temperature above 100 degrees Fahrenheit and low relative humidity (roughly 30 to 35%). Other tree species in this forest tract included blackjack oak and black or Texas hickory plus a few local colonies of sassafras and common persimmon. Blackberry was the dominant shrub. It was present as dead, burnt canes from last year's wildfire and resprouts (primocanes; first year, non-flowering shoots ) from the current growing season.

Some trees, including some of the taller (fifty to sixty foot height) ones, had been killed (no stump-sprouting) although most of these larger trees had not yet died even though their crowns had been devestated. The first of these three slides showed the recovering herbaceous layer of this burnt-over forest. Dominant plant species in foreground of this first photograph were silky wildrye (a native, cool-season, perennial, festucoid grass), tall fescue (an introduced, cool-season, perennial, festucoid grass), prickly lettuce, horseweed or mare'stail, and Japanese hedge parsley (Torilis japonica), this latter is a naturalized, annual exotic of the carrot or parsley family (Umbelliferae= Apiaceae).

The second slide showed a local area of the herbaceous understorey that was dominated by Japanese hedge parsley with a large common or bull thistle (Cirsium vulgare), a naturalized, Eurasian, biennial weed at left foreground and a young mare'stail or horseweed in right foreground (all-in-all a classic shot). The young trees (saplings) in this second slide were mostly hackberry and American elm; they were "accompanied" by an estern red cedar. The nonsprouting eastern red cedar was dead, but there were a few "anemic" and already dying basal sprouts of the hackberry and American elm.

The third slide presented the following DEAD (total top-kill with no resprouting) trees: four blackjack oaks (only three trunks clearly visible at left foreground), two post oaks (left midground), and one eastern red cedar (right midground). Crowns of trees in distant background showed tht these woody citizens were "hurtin' for certain". Local co-dominant forbs in foreground of this slide were Japanese hedge parsley, a naturalized, Eurasian annual, and pokeweed or pokeberry, a native, often-pioneering perennial.

These three slides were taken at the perimeter or just inside the margin of this forest tract where there was more light and closer proximity to tame pasture, and consequently, greater development of the herbaceous understorey.

Newton County, Missouri. Late July. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 40 (Post Oak-Blackjack Oak) X SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

181. A dead one deep inside the woods- An eastern red cedar ( between about 16 to 20 feet tall) that was killed (crown death equals complete kill of this nonsprouting conifer) in the interior of a post oak-black oak dominated second-growth forest one year following a hot, summer wildfire during Severe Drought. In this local area there were a lot of resprouting common or Virginia persimmon and sassafras. Other tree species included black or Texas hickory, blackjack oak, chinquapin oak, hackberry, American elm, bois d'arc (in that approximate order of cover deep inside this burnt-over forest tract). Blackberry, flowering dogwood, and buckbrush or coralberry were the dominant shrub species.

There had been very little herbaceous growth in the interior of this second-growth Ozark Plateau forest. One year after the wildfire there was a dense and well-developed herbaceous understorey consisting of mare'stail or horseweed, prickly lettuce, pokeweed or pokeberry, common or bull thistle, Japanese hedge parsley, flannel mullein (Verbascum thapsus) with a few plants of the native perennial grass, broomsedge bluestem (Andropogon virginicus).

Newton County, Missouri. Late July. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 40 (Post Oak-Blackjack Oak) X SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

182. Death and new life deep inside the woods- Interior of a second-growth post oak-black oak forest in the western Springfield Plateau one year after it had been severely burned by a hot summer afternoon wildfire the previous July. Air temperature was slightly above 100 degrees Fahrenheit and relatively humidity was around 30%. This wildfire took place during a Severe Drought.

Both of these slides featured a local stand of post oak that was made up of young, yet large, pole-sized trees of forest (complete canopy versus more open or widely spaced) growth habit (form).

In mid-growing season of the first year following this extremely intense (by oak-hickory forest standards) wildfire none of the post oaks seen here had died, but they were severely injured with most limbs in their crowns being dead or dying. There were some black hickory in with this post oak stand such as the small sapling in left-center foreground of the first slide which had been top-killed but was vigerously sprouting from its basal trunk.

Buckbrush or coralberry was the shrub species with most cover in the interior area seen in these two slides. There was also considerable cover of Virginia creeper (Parthenocissus quinquefolia). Almost all shoots of these two low shrubs (Virginia creeper covers the land surface as well as climbing to tree crowns) had been killed but were readily resprouting in this first full-growing season since the summer fire.

Almost all of the herbaceous growth (primarily a single-storey layer other than locally as shown below) in these two "photoplots" was that of forbs including pokeberry, prickly lettuce, mare'stail or horseweed, Japanese hedge parsley, plus a few plants of giant ragweed (Ambrosia trifida). The two Eurasian weeds known as common or bull thistle and flannel mullein which were locally abundant (even dominant) elsewhere were absent in the views presented in these two slides.

Newton County, Missouri. Late July. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 40 (Post Oak-Blackjack Oak) X SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

183. Happy forbs and a snag that will outlast them- Two views on the perimeter or outer margin of an Ozark Plateau second-growth forest dominated overall by post oak and black oak that had been burned a year earlier in a hot summer (July) wildfire during Severe drought. Locally dominant and associate tree species from blackjack oak, black hickory, chinquapin oak, American elm, hackberry, bois d'arc, common persimmon, and sassafras. There were were local patches of blackberry in this area.

In the local area featured here a charred large sapling of post oak stood "front and center" while a top-killed bois d'arc seedling or small sapling stood to the left of the of the blackened post oak snag. The pioneer forb species surrounding the dead hardwoods included co-dominants horseweed or mare'stail and prickly lettuce with flannel mullein, Japanese hedge parsley, and annual fleabane (Erigerion annuus) being major or locally, associate forb species. Prickly lettuce and flannel mullein are naturalized, Eurasian weeds that pioneer disturbed locations in oak-hickory forest throughout the Ozark Plateau. Prickly lettuce, like horseweed and annual fleabane (native, annual composites), is an annual while flannel mullein is a biennial.

Newton County, Missouri. Late July. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 40 (Post Oak-Blackjack Oak) X SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

184. More fuel meant more dead trees- An open canopy stand (a woodland) of post oak at one end of an Ozark (Springfield) Plateau second-growth hardwood forest co-dominated by post oak and black oak as it appeared one year after a hot, summer (late July with ambient tempaerature above 100 degree Fahrenheit) wildfire during a Severe Drought. Broomsedge bluestem was the dominant understorey species at time of the wildfire. There was quite a fuel load of broomsedge herbage (both accumulated dead and live biomass) for the hot wildfire. Enough heat was generated that the crowns and trunks of the good-sized, young post oaks in this stand trees were killed or severely damaged. At time of these slides it was impossible to know if these post oaks would die, but it appeared most likely that they would succumb eventually either directly from fire injury or indirectly from disease, drought, or some combination of these and other factors.

The understorey remained largely herbaceous, except for blackberry which had resprouted with primocanes, following the wildfire. The pre-fire understorey dominant species of broomsedge bluestem was lost at this point in secondary plant succession. Instead, the post-fire dominat was pokeberry or pokeweed with flannel mullein the overall associate herbaceous species while the next most important species was annual fleabane. Flannel mullein is a naturalized, Eurasian biennal while the pokeberry is a native, pioneering perennial.

The first or vertical slide was taken when a cloud passed in front of the sun whereas the second or horizontal slide was taken a few minutes later when the cloud disippated. Different forb species were more visible under the two light conditions; annual fleabane was better seen with overcast sky while pokeberry was more visible under full sunlight.

Newton County, Missouri. Late July. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 40 (Post Oak-Blackjack Oak) X SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

185. Dead and dying trees the second season- An open canopy stand (a woodland) of post oak at one end of an Ozark (Springfield) Plateau second-growth hardwood forest co-dominated by post oak and black oak as it appeared two years after a hot, summer (late July with ambient tempaerature above 100 degree Fahrenheit) wildfire during a Severe Drought. These two images are of the same views (= same areas) as shown in the two immediately slides of this vegetation last year (the previous summer; almost exactly one yer later).

There were pronounced differences over this one year period in this oak-dominated woodland vegetation. The trees--both living (barely) and dead--were the same individuals except that some more trees had died while other trees had increased size and foliage area of limbs and trunk sprouts (shoots originating from intercalary meristem following heat damage. This can be seen by comparing the first or vertical photographs in this and the preceding slide/caption unit.

The bigger difference, however, was in the understorey of this woodland range vegetation. In the first full spring/summer growing seaason (last year's vegetation; that shown in the two immediately preceding slides) following the hot summer wildfire (two years prior to time of these two slides; first slides in this chapter section devoted to a hot, summer, surface fire) first season post-fire understorey vegetation was dominated by the native, perennial known as pokeberry or pokeweed with flannel mullein was the overall associate understorey species was flannel mullein, a naturalized Eurasian biennial of the snapdragon family. After these two forb species the next most important herbaceous plant was annual fleabane and with mare's tail or horseweed second to fleabane. Both of these forbs are native annual composites. There was slso considerable cover of the exotic annual forb, Japanese hedge parsley.

By partial contrast, the understorey vegetation this summer (almost eaxctly one year later) as presented in these two slides still consisted of much cover of pokeweed and some of the same alien, naturalized forbs, but numerous shrub species--including blackberry, winged sumac, buckbrush (Symphoricarpos orbicularis), and cat-brier (Smilax tamnoides; S. bona-nox was also present in this forest but not in this local vegetation), all of which are native species, had already established themselves as had the native, perennial silky wildrye (Elymus villosus) and exotic, agronomic forage secies, especially tall fescue and sericea lespedeza. There were also plants of wild bergamot (Mondarda fistulosa), a native perennial of the mint family, and individuals of purpletop (Tridens flavus) and broomsedge bluestem two native, perennia grass species. The latter warm-season panicoid grass had been damaged and severely set back by the hot, summer wildfire while purpleto appeared to have been released by fire.

There were some not-very-healthy-looking stump sprouts of post, blackjack, and black oaks. The main stump-sprouting, hardwood species, however, was bois d'arc. Osage orange, or hedge apple (Maclura pomifera) which is native to the Ozark Plateau. In fact, the English word, Ozark supposedly was derived from the French bois d'arc meaning literally "bow wood" (for its use by Indians for their long bows).

Perhaps most interesting was presence of some sprouts (from stumps of small, young trees) along with seedlings of hackberry (Celtis occidentalis) in this second full spring/summer growing season following the hot, summer, surface fire. Hackberry and elms (Ulmus americana and U. rubra) are often climax trees in this area, but it has long been accepted that fire reduced these less fire-tolerant species so that the oak and hickory forests of the Ozark Plateau was at least partly a result of period fire. In this postulated theme, periodic fire eliminated (at least greately reduced) presence hackberry and elm species that were less tolerant of fire than the oak and hickory species. In the instance described herein, hackberry and bois d'arc came back as promptly as the climax oak and hickory species.

It appeared that this woodland community was, in actuality, an oak-hickory-hackberry-elm savanna with a potential understorey dominated by big bluestem (Andropogon gerardii) or, perhaps, co-dominated with silky wildrye, both of which are dexcreasers on this forest range site. The climax dominant big bluestem had been largely grazed out and replaced by broomsedge bluestem (an invader) with numerous woody plants (most of which were increasers or invaders). Silky wildrye was represented by more plants and greater cover than any other native grass species. The cool-season, C3 wildrye was largely or entirely dormant at time of the summer wildfire so it was much less vulnerable to fire damage than were warm-season species like big bluestem and the increaser eragrostoid grass, purpletop.

Newton County, Missouri. Late July. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 40 (Post Oak-Blackjack Oak) X SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

186. Fire-dead post oak in a burnt-over woodland- Medium-sized post oak top-killed by a hot summer (late July) wildfire that burned over a second-growth hardwood forest in the western Ozark Plateau a year earlier. Intense heat was generated by this fire that burned during an afternoon of temperatures in excess of 100 degrees Fahrenheit with relative humidity of about 30%, plus this was during a Severe Drought.

The bark on the trunk base of this post oak split and separated from the cambium.

Major forest plant species in the herbaceous understorey during middle of the warm-growing season included the dominant pokeberry; flannel mullein, associate herbaceous species; Japanese hedge parsley, and annual fleabane. There were primocanes of resprouting blakcberry.

The view in the first slide included fire-killed eastern red cedar and top-killed bois d'arc or osage orange, two native tree species that are invaders on this forest range site.

Newton County, Missouri. Early July.

187. Growing around a snag- Pokeberry and winged sumac co-dominated a local habitat (microsite) around the snag of a young, pole-sized post oak (first, horizonatal slide) while pokeweed dominated the microhabitat aroung the snag of an older, young adult-sized black oak (second, vertical slide) in the third summer (sesond spring-summer growing season) following a hot, summer, surface wildfire during Severe Drought in a post oak-blackjack oak-hackberry-elm-big bluestem woodland or "close" savanna in the western Springfield Plateau. On the outer margin of this tree-grass range vegetation black oak was the associate species to locally dominant species.

These two images were of recovering range vegetation on local areas that were contiguous with each other. Other plant species in this second seral stage-plnat community (the first seral stage of the preceding year included annual and biennial colonizing forbs in addition to the perennial pokeweed) were the following native species (not in any particular order of cover, importance, etc.): the perennial grass known as purpletop (Tridens flavus); blackberry (Rubus sp.); coon or 'possum grape (Amelopsis cordata), which was climbing the post oak trunk in the first slide; an unidentified boneset or thoroughwort (Eupatorium sp.), Baldwin ironweed (Vernonia baldwinii), a few plants of mare'stail or horseweed, an unidentified Erigeron species, an unidentified annual smartweed or knotweed (Polygonum sp.) and rough or Virginia buttonweed (Diodia virginiana). Naturalized annual species in this seral range vegetation included Japanese hedge parsley, hairy crabgrass (Digitaria sanguinaria), a few remaining (and flowering) plants of the biennial flannel mullein, and beefsteak plant (Perilla frutescens). There were some barely-managing-to-survive plants of broomsedge bluestem, but these individuals had clearly been injured by the wildfire/drolught "double-whammy".

It was remarkable that about one-third of the way through the second full-warm growing season (the third partial summer) following the devestating widlfire (that was concurrent with Severe Drought) native perennials had become established. Some plants of these species such as blackberry, 'possum or coon grape, Baldwin ironweed, and purpletop had most likely been present at time of the wildfire, but some other plants also appeared to be post-fire individuals. Most of these species were increasers and not major members of the climax vegetation. Big bluestem was the potential natural (= climax) herbaceous dominant, but this decreaser had been greatly reduced (just about eliminated) by a long history of overgrazing.

Newton County, Missouri. Late July. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 40 (Post Oak-Blackjack Oak) X SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

188. Smoky Bear told us this could happen- Two views of the lower trunk of a snag of a comparatively large, mature, and prime-of-life black oak that had been killed three summers ago by an intensely hot summer wildfire coupled with Severe Drought in the western Ozark (Springfield) Plateau. There were not any sprouts (shoots from intercalary meristem) at any location on this tree adult tree. Bark on this trunk had either already either peeled away (first slide) or was in the process of peeling off (second slide).

Major plant species around this black oak snag included flannel mullein (left front of trunk in first slide), pokeberry (left front of trunk in second slide), blackberry, some species of knotweed or smartweed (immediate foreground, second slide), Baldwin ironweed, poison ivy, rough or Virginia buttonweed, Japanese hedge parsley, and an unidentified boneset or thoroughwort species.

This specimen and several other individuals of black oak (another example was presented in the immediately succeding slide) were more abundant--and larger--at the outer edge or margin of this forest range dominated by post oak and blackjack oak with black or Texas hickory plus younger trees (more successionally advanced?) of hackberry. The author did not find any seedlings of black oak in this recovering (via secondary plant succession) forest or woodland vegetation. There were a few rapidly growing seedlings of persimmon and sassafras and, even fewer, stump (basal trunk) sprouts of top-killed smaller (and younger?) black oak. There was, however, more resprouting of bois d'arc than of black oak. Widespread experience and common observation has been that oaks and hickories are more tolerant of fire than is bois d'arc (and, also, hackberry, American elm, red or slippery elm), but three summers after this wildfire/drought "double whammy" combination there was more sprouting from basal trunks and rootcrowns of bois d'arc than of the supposedly more fire-tolerant post oak, blackjack oak, and black oak.

The long-held view (backed by research findings and field observations) that recurrent fire is a major factor responsible for existence of oak-hickory forests (and reduction, even elimination, of other hardwood tree species) was not supported by empherical findings from the present anecdotal/photographic set of observations. There was one exception to this general finding in the current observational investigation that was consistent with research into impacts of fire on oak-hickory forests: fire took out the non-sprouting eastern red cedar.

Newton County, Missouri. Late July. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 40 (Post Oak-Blackjack Oak) X SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

189. Dead black oak in a fired-through woodland- Young adult black oak top-killed by a hot summer (late July) wildfire that burnt through an Ozark Plateau second-growth, hardwood forest during Severe Drought. Post lak and black oak were co-dominants of this forest that also included black hickory, blackjack oak, chinquapin oak, American elm, hackberry, and smaller amounts of persimmon and sassafras. The major shrub species was blackberry.

This was an extremely hot wildfire (it burned on an afternoon of temperatures above 100 degrees Fahrenheit and relative humidity of about 30% during a Severe Drought) by oak-hickory forest standards. Heat from the wildfire was so intense that it cracked the basal bark of trees and separated the inner bark from the cambium.

This tree grew in a woodland (open tree canopy) rather than a forest (closed canopy) form of the post oak-black oak community. The more widely tree crowns permitted productive growth (high yields or heavy biomass) of broomsedge bluestem that dominated the herbaceous understorey. Broomsedge furnished a heavy load of easily ignited fuel that resulted in such intense heat that oak trees suffered the extent of injury seen here (and in the immediately preceding slide-caption set).

The herbaceous understorey one year after the wildfire consisted of pokeweed, flannel mullein, and annual fleabane rather than the pre-fire dominant of broomsedge bluestem which was demolished when the wildfire burned this panicoid grass during mid-growing season and prolonged drought.

Newton County, Missouri. Early July.

190. The next summer- Another example of what Smoky Bear warned of was seen in these two views. The lower trunk of a fire-killed black oak in the third summer (second full spring/summer growing season) following an extremely hot, summer surface wildfire during Severe Drought. This was the same trunk (seen from a slightly angle) presented in the two immediately preceding slides. The peeling bark (separating from the cambium layer) was obvious in both sets of slides.

A plant of some Aster species was growing to the front (front-center, first slide; slightly to the left front, second slide) of the trunk while a plant of common ragweed (Ambrosia artemisiafolia) accompanied it (right of the common ragweed). In the background were plants of beefsteak, horseweed or mare'stail, and prickly lettuce (Lactuca scariola= L. serriola).

This young adult tree had been growing at the edge of a woodland dominated by post oak and blackjack oak with hackberry and American or white elm and, to lesser degree, red or slippery elm.

The wildfire was human-caused through careless and in defiance of a county wide burn ban. The "firebug" spent a day or two in the county jailhouse. Apparently there was no further punishment for the crime. Successional time will tell the ecological outcome of the crime. For now, all that can be known was that some beautiful, straight-trunked, yound climax trees were destroyed and other plant species responded to their advantage to the fire/drought disturbance. This was a textbook example of Clementsian denudation (Clements, 1916). It was also what Smoky Bear campaigns warned of with the waste caused by wildfire. Ecological results--at least some results--were the opposite of the benefits of prescribed fire or of more natural spring fires. Fire--like axes, plows, ropes, guns, bulldozers, and so on--can be either a creative and life-enhancing tool or a destructive and mere agent of death.

Newton County, Missouri. Late July. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 40 (Post Oak-Blackjack Oak) X SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

191. Fire pioneers and beneficiaries at their peak- The herbaceous understorey of a second-growth hardwood forest (co-dominated by post oak and black oak) at peak standing crop the year following a hot wildfire in late July (an afternoon of temperatures above 100 degrees Fahrenheit and relative humidity of about 30%) during Severe Drought. (Fire was started by a clueless landowner practiceing convenience burning of trash and accumulated tree leaves.)

Species present included the climax decreaser grass, silky wildrye; prickly lettuce (a naturalized Eurasian annual composite); horseweed or mare'stail (a native annual composite), Japanesse hedge parsley (naturalized exotic member of Umbelliferae), longleaf ground-cherry (Physalis longifolia), blackberry, and the woody climber, fiddleleaf green-brier (Smilax bona-nox).

Most of the hebaceous species in this pioneer seral stage of secondary plant succession were at their peak standing crop and in young to maturing fruit phenological stages. This pioneer plant community was at the outer edge (perimeter) of an Ozark Plateau oak-hickory forest.

Newton County, Missouri. Late July. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 40 (Post Oak-Blackjack Oak) X SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

Upland game management note: Devestating though this hot, summer wildfire (coupled with Severe Drought) was to the second-growth oak-hickory forest, fire impacts on vegetation were a boon to northern bobwhite quail (Colinus virginianus) and, even more so to white-tailed deer (Odocoileus virginianus). Habitat for the "king of upland game birds" was greatly improved by "opening up" of the dense forest which permitted growth of herbaceous pioneer species that afford both cover and valuable concentrate feeds for this r-selected, gallinaceous bird. While bobwhite use these pioneeer plant species primarily for cover and fruits like achenes as concentrates, white-tailed deer feed directly on the herbage of prickly lettuce and, less frequently, mare'stail or horseweed.

In this respect, annual composites like prickly lettuce, horseweed, and, even, common thistle that are correctly classified as weeds from an agronomic or horticultural perspective (see for example regarding horseweed and prickly lettuce, Whitson et al., 1992, ps. 122-123, 152-153, respectively) are not weeds from the standpoint of valuable wildlife species such as white-tailed deer and bobwhite quail. In fact, such pioneering forbs are an important component of the habitat of such game animals. These pioneering forbs are crop plants for these wildlife species.

In just a few years secondary plant succession will result in replacement of pioneering "weeds" with plants of higher successional order, but for now the "fire crop" provided improved habitat for bobwhite quail and white-tailed deer.

As a final thought, the irony of this phenomenon was that while the wildfire was a criminal misdemeanor (some degree of arson) that violated several statutes of the state penal code it had some beneficial impacts, especially in the short term. Overall, however, impacts were negative to the timber owner and neutral to the self-recreating forest over the long term, the time scale of plant (forest) succession. Such is Life.

192. Peaked pioneers of a fired forest- Three "photoplots" of pioneer forb species that comprised the herbaceous understorey of a second-growth hardwood (mostly oak-hickory) forest in the Springfield Plateau at peak standing in the first full-growing season following an extremely hot summer (late July) wildfire that burned during Severe Drought the previous year (a week later than one full year after the forest fire).

The first "photoplot" presented a synopsis or overall view of the seral herbaceous layer while the second and third "photoplots" were "subplots" or "nested photoplots" of the first or overall (larger) "camera plot". The co-dominant forbs were 1) the native, annual composite known as horseweed or mare'stail and 2) the introduced and now naturalized, annual composite called prickly lettuce. These two annual forbs are in the Tubuliflorae and Liguliferae subfamilies of Compositae (Asteraceae), respectively. The associate herbaceous species was also a forb, but a native, perennial of family Phytolacaceae known variously as pokeberry, pokeweed, or pokesalad. Pokeberry is unusual in being a pioneering herb that, number one, is a perennial and, number two, one that often persist indefinitely from an immense, fleshy taproot. Other pioneering herbaceous species are transient occupants on the forest sere. In this first slide there were a few plants of Japanese hedge parsley which were conspicuous by their end-of-life, dried, tan-colored shoots as seen in right-of-center foreground.

The second slide featured the two co-dominant composites (horseweed and prickly lettuce) and the associate herbaceous species, pokeberry. Plants of prickly lettuce were shedding their parachute-equipped achenes while the berries of pokeweed varied from shed-fruit, to ripe, to still green-fruit in the same cluster. Fading and drying leaves of pokeberry indicated that these shoots had matured and were preparing to die as this native perennial was slowly approaching the dormant pahse of its annual cycle. Mares'stail or horseweed was in its bolting stage or very early stge of panicle development. This native, annual forb was "a ways away from" peak standing crop, but the composite or sum of all herbaceous species (including silky wildrye as seen above) was the point of peak standing crop or maximum yield of herbage.

The third slide was an uncommon (for this area of the Ozark Plateau) combination of primarily mare'stail or horseweed with lesser cover and density of prickly lettuce as an upper herbaceous layer and with a lower herbaceous layer made up of wild sorrel (Rumex hastatus) that was visible as the dark reddish brown shoot in center foreground.

Newton County, Missouri. Late July. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 40 (Post Oak-Blackjack Oak) X SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

Interesting successional exception: absent from the first seral or pioneer stage of secondary forest succession shown and described above was the naturalized biennial forb, wild carrot or Queen Anne's lace (Daucus carota), This experienced Ozarker woods walker did not spy one plant of wild carrot try as he might. Wild carrot was not present in the first year's pioneering seral community period, and this hillbilly would have found the conspicuously showy umbel of this ancestor of the domestic carrot had it graced this recovering forest range vegetation.

193. Into the second full-growing season- Range vegetation in the interior of a second-growth hardwood (mostly oak-hickory) forest in the Springfield Plateau at early summer in the second spring-summer growing season (third partial summer) after an extremely hot summer (late July) wildfire that burned during Severe Drought. These two images were two slightly different angles (camera positions) of roughly the same forest community with the bowed-to-the-right tree trunk serving as a "landmark". This bowed (and broken off) trunk and the two tree trunks to the left and left-front of it were snags of blackjack oaks killed by the extremely intense heat of the summer wildfire (and, possibly, in combination with drought). None of these trunks had resprouted; an unusual response given the youth and comparatively small size (large saplings to small poles) of these blackjack oaks. Sapling-size snags in the background (especially right background) of the first (vertical) slide were post oaks.

Another interesting development was the emergence of numerous seedlings of hackberry (in foregrounds of both slides) following the wildfire. As a general rule, hackberry (as well as elm species) is much less fire-tolerant than the dominant hardwoods, post and blackjack oaks and black hickory. As such, recurrent fire selects for the oak and hickory species over hackberry and elm so that fire is a major variable in maintenance of oak-hickory forest. In this instance, however, there was apparently more post-fire regeneration of the less-tolerant hardwood species. There were some seedlings of the oaks and, to lesser extent, black or Texas hickory, but there were more seedlings of hackberry. Similarily, a higher proportion of Osage orange or bois d'arc resprouted than of the oak species and black hickory. Obviously, none of the fire-killed eastern red cedar sprouted (= regrew) as this conifer is a non-resprouting species.

Major plant species in the understorey (both herbaceous and woody components) in addition to hackberry seedlings included silky wildrye, pokeweed or pokeberry, Baldwin ironweed, sericea lespedeza, purpletop, and wild bergomont among herbaceous species and cat green-brier, winged sumac, blackberry, poison ivy, Virginia creeper (eg. shoot of a Virginia creeper just above the big burned spot on the foremost blackjack oak), and (as explained in the preceding paragraph) some stump sprouts and seedlings of the two oak species as well as stump shoots of bois d'arc or Osage orange in the woody component.

At this point (this seral stage) in range plant community succession (only one-third of the way through the second complete warm-growing season) the pioneer or first colonizing annual species such as horseweed or mare'stail, prickly lettuce, Polygonum species, and Erigeron species were much diminished in cover and density compared to the previous (last) year which was the first complete spring/summer growing season after the intense wildfire. These annual colonizing or pioneer species dominated their sere for only one year, the year immediately following the extraordinargy summer fire coupled with Severe Drought.

Comparison of this second seral-stage re-development of forest or woodland range vegetation (these two slides) with the first seral (initially colonizing) stage following wildfire (the two slide/caption sets immediately above) provided a dramatically stark contrast in successional plant communities (seral stages) between the first and second spring/summer growing periods. Forest range succession was rapid on this sere (this forest range site). Frederic Clements' "dynamic vegetation" was (in 1950s parlence) "super-sonic" fast in successional time scale.

Newton County, Missouri. Late July. FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 40 (Post Oak-Blackjack Oak) X SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

Forest Gap; Patch Dynamics- Lightening Strike

The following provided an example of the phenomenon known as a disturbance gap or disturbance patch and, hence, patch or gap dynamics as disturbed or destroyed vegettion re-develops via plant succession (usually secondary succession). This is a condition where a local--often a relatively small local--area has partial or complete defoliation due to such things as windthrow; small, spot fire; disease or insect kill, or lightening strike followed by the process of secondary plant succession in that restricted gap or patch. This is known as patch or gap dynamics. An example of the patch dynamics due to lightening strike was included at this point.

194. Blasted- Gap created in an upland black oak-pignut hickory forest by death of large (23 inch DBH) adult pignut or bitternut hickory due to lightening strike. Lightening knocked off one of two major limbs at base of tree crown and also resulted in total or complete death of the hickory. This was not just top-kill, but also death of rootcrown and roots. Saturated soil was perhaps a contributing factor to such thorough killing of tree through better grounding. Another factor was undoubtedly age given the physiological phenomenon whereby older trees sprout much less readily and vigerously than do younger trees. Amazingly, no other plants were affected by the lightening strike. One large single trunk of a flowereing dogwood was broken off by crash of the falling limb from the pignut hickory; however, this dogwood had alredy been top-killed by the moderate-intensity surface fire and resprouted from its rootcrown two springs earlier so that there was literally zero direct impact from the lightening strike on any plant except the hickory.

This was appearance of the forest gap in the early part of the second summer after death of tree and creation of the gap with initiation of vegetation dynamics. The initial or nearly immediate effect was release of all plants that had previously been shaded (for varying periods of time) by the canopy of the hickory, the holder of the high ground as it were. Tree species were the major beneficiaries of release in the gap. Tree release was as seedlings and saplings that were almost all of climax species including pignut hickory, the major released species, black oak, American elm, white ash, and northern red oak. The one seral tree species released was sassafras (Sassafras albidum). There were no grasses or grasslike plants in the forest gap because the soil surface was almost completely covered by seedlings and saplings of the various tree, shrub, or even forb species. The major forb species was pointed-leaf, sticky, or cluster-leaf ticktrefoil (also written as tick trefoil and known as tick clover). Other--though much less common--forbs were carrion-flower (Smilax herbacea) and pokeberry. Important shrubs in the gap itself included fox grape, flowering dogwood, Virginia creeper, and blackberry.

Pioneer (weedy) plant species were not present in thsi forest gap. The author could not find a single plant of an annual or biennial species. The closest species to early colonizers were pokeberry and sassafras. For all practical purposes secondary plant succession was not taking place. Patch dynamics consisted of establishment, growth, and maturation of young plants of climax species.

Ottawa County, Oklahoma. Late June (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

195. Kaboom!- A large (23 inch DBH) pignut hickory struck down in its prime by lightening that completely killed it (not just top-kill but total death). This hickory had a remarkably straight bole with relatively taper which forked into large limbs from which were the entire crown originated. One of these limbs was knocked off by the lightening strike. Saturated soil resulting recent heavy rains served as thorough grounding for the tree. No other plant was directly affected by the lightening strike. The crashing limb of the hickory broke off a large, dead trunk of flowering dogwood that had been topkilled three years earlier by a wild surface fire of moderate-intensity. The coppice or stump sprouts of this dogwood were unphased by the fallen limb.

The opening in the forest canopy that was created by death of the pignut hickory was a "windfall" for all other plants. These consisted primarily of seedlings and saplings of the climax tree species, especially pignut hickory of which there were more sexual offspring than any other tree. Other climax trees with both saplings and seedlings included white ash and black oak. There were seedlings only of northern red oak and American or white elm of the climax group and also seedlings of the seral (for this forest range site) black cherry (Prunus serotina). There was one already existing sapling of American elm that had been severely scourched by the wild fire and that had partly leafed out, but it was not going to make it (or amount to anything if it did).

Abundant shrub species in this still-young forest gap included fox grape, Virginia creeper, flowering dogwood, and blackberry. The most abundant forb was sticky, pointed-leaf, or cluster-leaf tick clover or tick trefoil, the herbaceous dominant for this forest range.site Other forbs included carrion-flower (an herbaceous green-briar), wingstem crownbeard (Verbesina helianthoides), and pokeberry or pokeweed. Grasses were not present in the thick "jungle" of this forest gap even though hairy mountain brome, Canada wildrye, and woodland bluegrass were scattered variously throughout this upland forest.

Ottawa County, Oklahoma. Late June, first slide; Late July, second slide (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

196. Shattered life and new life- Lightening-shredded trunk (23 inch DBH) of a pignut hickory was at center of a gap in a black oak-pignut hickory climax upland chert forest in the western Springfield Plateau. Complete (crown to root system) kill of this once-mighty hickory set into motion patch dynamics in this forest gap. Change in vegetation was nothing more than release of already present climax species of trees, shrubs, and forbs. The lightening strike affected directly only the one, mature, hickory tree. Neighboring adult black oaks, flowering dogwood, seedlings and saplings of other trees, etc. were unaffected by the lightening bolt, but clearly benefitted from the inpouring of light after the crown of the pignut hickory was reduced to a skeleton of bare branches. In fact, one of two limbs that formed the crown was knocked off by lightening strike.

There were saplings and seedlings of pignut hickory, the species that produced the majority of sexual progeny, along with those of black oak, and white ash. Reproduction of American elm and northern northern red oak, the other climax tree species in the gap, was limited to seedling classes. There was one partly leafed-out sapling of American elm that had been mostly top-killed by a surface wild fire of moderate intensity three springs ago, but it was not going to do anything. There were also some seedlings of wild cherry which is not a climax species for this forest type or site. The most abundant shrubs were fox or winter grape and Virginia creeper. Flowering dogwood was another important shrub many of which consisted of stumm sprouts following the wild fire. The most common and productive (greatest cover and density) forb was sticky, pointed-leaf, or cluster-leaf ticktrefoil (tick clover) which was a climax understorey dominant for this forest and forest site. Other forbs included carrion-flower (an herbaceous green-briar) and wingstem crownbeard, this latter visible in right midground. Grasses, such as hairy mountain brome and Canada wildrye which were present in various local microhabitats throughout this forest, were not in the gap.

The large sapling snag with falling bark and lichen growth to immediate right of the pignut hickory was northern red oak that had been dead for several years prior to lightening strike apparently having lost the war in competition for light (and/or whatever else) with the larger, older hickory. Just no justice in these woods: the nice northern red oak died too soon to grow to greatness. Its younger siblings will have to be the ones that become part of the climax. forest.

Plant succession was not occurring on this forest gap. Development of vegetation consisted of growth, maturation, and reproduction of individual plants of the climax species. Pioneeer and advanced seral plant species (other than a few sassafras plants) were not present.

Ottawa County, Oklahoma. Late July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

197. "One generation passeth away,and another generation cometh..." (Ecclesiastes 1:4)- Several seedlings and one sapling of pignut or bitternut hickory at base of the trunk of an old-growth pignut hickory killed the previous year by lightening. Pignut hckory was the plant species with the greatest reproduction (both asexual and sexual) in the forest gap created by death of this single, adult hickory tree. Other climax tree species that had seedlings in the gap included black oak, northern red oak, american elm, and white ash. Also present (as in this photograph at far left margin) were seedlings of black cherry. The dead sapling at upper right corner was an American or white elm that was killed three years previously by a wild surface fire of moderate intensity. Leaves and woody vine of fox grape were conspicuous at lower right. Leaves of Virginia creeper were visible at left.

Ottawa County, Oklahoma. Late July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

198. Same patch, next year-View of the lightening-created gap in a second-growth, upland, black oak-pignut hickory forest that was introduced above as it appeared in the next (and a drought) year. This was the third year, growing season, and part of summer after lightening blasted a comparatively large (23 inch DBH) pignut hickory and created a microsite of new forest land on which the forest community had to redevelopment. Development of new forest vegetation was primarily one of released existing plants and relatively little or quite limited appearance of typical pioneer or colonizing species. The most abundant of these early seral species was sassafras. By contrast there were several existing plant species that made rapid growth due to reduced competition, greater light, etc. (ie. that were released) with death of the large hickory and creation of an opening in the foret canopy. Most of these released species were climax or subclimax species including: seedlings and saplings of the tree species pignut hickory, black oak, American elm, and white ash; shrub species such as flowering dogwood, eastern redbud, and summer grape; and forbs like sticky tickclover or sticky tick-trefoi and carrion flower.

The shrubs and forbs were present before lightening killed the pignut hickory as were some saplings of the four tree species. In fact, most plants of the tree species had established prior to lightening strike, but there were seedlings that had emerged and established after lightening created the forest gap. None of the shoots of the four tree species were stump sprouts or suckers (ie. release and regeneration of trees resulted from sexual reproduction). By contrast, most (probably not all) of the increased cover and biomass of shrubs and herbaceous species wasasexual reproduction (ie. proliferation of shoots from rootcrowns, rhizomes and so on).

Ottawa County, Oklahoma. Mid- July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

199. The early years have the big changes- The greatest changes on a redeveloping sere (ie. recovey of disturbed or perturbed vegetation following denudation of a parcel of land via secondary plant succession) take place in the earlier or younger stages. This is the case under textbook old-field or cutover forest succession. That general pattern or form of recovery--in other words, classic secondary succession as described by countless workers even before Clements (1916) reviewed this work and gave Ecology what became known as the Clementsian paradigm--is the typical or usual scheme of plant community redevleopment on parcels of land of such size created by farming (fields), intensive logging (clearcut forests), ranching (especially overgrazed ranges), large wildfires, road construction, etc.

Small gaps such as those in forests formed by death of damage to a few trees as caused by localized blowdown, lightening, uprooting, or beaver feeding and similar small areas of denudation or disturbance on shrublands or, even, grassland (eg. a small burnt patch after the flames burned out in short order) differ in at least one aspect of defoliation from these situations. That aspect or facet of defoliation or denudation is area. The impacts of the other defoliation (denudation) facets of intensity, frequency, seasonality, and selectivity are moderated by decreasing size of the affected (denuded) area. This is because existing plant propagules are close at hand as well as the phenomenon by which remaining vegetation lessens impacts of such variables as ambient temperature, wind, precipitation, even humidity by presence of shade, windbreaking, transpired water, etc. This might be partially counteracted by attraction of animal attraction or, alternatively, this might also moderate or even facilitate recovery of vegetation via seed dispersal, addition of soil nutrients via animal excretion, and so forth.

Ottawa County, Oklahoma. Mid-July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

200. Growing on my mamma's grave- Pignut hickory seedlings that are approaching sapling size growing next to the shattered trunk of the parent hickory (23 inch DBH) that was killed by lightening gthree years ago. There were also young trees of this same size (and presumedly roughly same age) of black oak whose parents missed the lightening bolt and were standing on the edge of this forest gap. There were also seedling of similar size/age of white ash the parents of which grew farther away and lower down on an adjoining north slope as well as American elm the fruit-bearing trees of which grew in closer proximity to the three-year-old gap. There were also seedlings/saplings of slightly smaller size (based on general height) of sassafras. There were no adult sassafras in this forest so origin of propagules was unknown: could have soil seedbank, carried in by wind or animals, whatever.

There were young plants (four to six foot shoots) of flowering dogwood and redbud along with stems of Virginia creeper and summer grape. Herbaceous species were sticky tick-clover and carrion flower were presented in accompanying slides.

Ottawa County, Oklahoma. Mid-July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

201. Where forest succcession is just more of the same- Climax dominant tree species immediated reclaimed forest gap created when lightening killed a large (23 inch DBH) pignut hickory. Member species of the climax forest represented by seedlings (some of which approached small sapling size) included pignut hickory (probably the progeny of the dead tree), black oak, American elm, and white ash. There were also seedlings of sassafras, a seral tree species the propagule source of which was undetermined. Climax understorey shrubs were also poresent including redbud, flowring redbud, Virginia creeper, and summer grape.

Herbaceous species were not descernible in this slide, but they were presented elsewhere including the next two-slide set.

Ottawa County, Oklahoma. Mid-July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

202. Rotting corpse and burial attendants- One of two main limbs of a large pignut hickory that fell following death by lightening strike three years ago.Surrounding the limb were two major understorey forbs, sticky tick-clover or sticky tick-trefoil. Also conspicuous were yearling seedlings of pignut hickory and American elm showing that climax tree species were replacing the dead climax co-dominant tree species. Likewise, these two forbs had been present in the climax forest only in a suppressed state. No suppression in this lightening -created gap in the forest canopy. In essence there was very limited secondary plant succession in this small patch. Instead there was enhanced growth of already present climax forbs and seedlings of climax tree species. There was also sexual regeneration by seedlings thqat emerged following gap formation.

The decomposer (reducer) component of this small, desturbance forest ecosystem was represented by numerous fruiting bodies of the shelf-fungus, white cheese polypore (Tyromyces chioncus), growing along the fallen limb of the lightening-killed hickory. a pignut hickory killed three years earlier by lightening.

Ottawa County, Oklahoma. Mid-July (estival aspect). FRES No. 15 (Oak-Hickory Forest Ecosystem), K-91 (Oak-Hickory Forest), SAF 110 (Black Oak). Oak-Hickory Series 122.11 of Brown et al. (1998). Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

203. Being digested by a "cheesy" character- Fruiting bodies of the shelf fungus, white cheese polypore (Tyromyces chioncus), growing on and rotting (digesting) the limb of a pignut hickory that crashed three years ago after the large adult tree was struck by lightening. The gap created in the forest canopy by the lightening strike was treated in detail above (and summarized immediately below). These and and the two immediately preceding slides emphasized to students that decompoesrs are part of range ecosystems. Such fungal reducers can be especially "photogenic" in forest range ecosystems.

Tyromyces chioncus is a member of Polypodaceae (polypore family) within order Polyporales, class Hymenomycetes and in subdivision, Basidiomycotina (Basidiomycetes).

Ottawa County, Oklahoma. Late July.

204. "Cheesy" details- Two carpophores (fruiting bodies) of white cheese polypore (Tyromyces chioneus) on downed and slowly rotting limb of pignut hickory. A closer-in shot of tow of the many such spore-producing structures of this particular organism in a Dry-Mesic, Chert Forest (Nelson, 2005, ps. 125-130) in the western margin of the Ozark Highlands (Springfield Plateau).

The involuntary, hardwood host (source of detritis) of this heterotrophic organism, a fungal saprophyte, was killed three years ago by lightening strike.This served as a reminder to students of Ecology that any disturbance serves as habitat for some living thing. The blast from above that ended the life of this magnificant pignut hickory created a forest gap that released the offspring of the killed hickory (described above) and that created a new forest environment for other species, including white cheese polypore.

Ottawa County, Oklahoma. Late July.

Big lesson from a big boom: The concept, the pehnomenon, that emerged from this example of patch dynamics in a forest gap was that secondary plant succession did not--and, by extension, to similar forest gaps does not--apply at such small spatial scale. Range scientists such as Harold Heady (1975, ps. 10 [Figure 1-3], 12-70 passim) quite properly defined and described the four defoliation aspects or components of: intensity, frequency, season, and selectivity. These aspects of defoliation have been expaned to those of herbivory in general. These four correspond roughly to the Four Cardinal Principles of Range Management: proper degree (intensity) of use, proper season (season and frequency, interaaction thereof) of use, proper distribution of use (interaction of frequency and intensity), and proper kind and class of range animal (selectivity). (See again Heady, 1975, p. 7, Figure1-2).

To these four aspects or components of defoliation fire ecologists more recently added a fifth: size of area or extent (of defoliation) versus intensity (as in degree of) defoliation. By way of example consider a large-scale forest fire covering hundreds of square mile as compared to a quickly extinguished spot fire that grew to only a fewhundred square feet. Opportunity for regeneration by dispersed seed will be greater for smaller areas. Size and severity of modified microclimate and surface for accelerated erosion will be less for smaller areas of disturbance.In final, smaller disturbed patches will recover more quickly--other things being equal--than will larger areas of defoliation disturbance.

Furthermore, other things will not be equal, they will be more severe for larger areas of disturbance (bigger defoliation patches). If the black oak-pignut hickory forest on which the lightening-created gap occurred had been clearcut there would have been local areas of pioneer annuals such as crabgrass and weedy composites (ie. r-selected species). In the lightening-caused patch seem above there were only climax or, at least, high seral stage species and no annuals or biennials (ie. primarily K-selected species).

Response also varies with plant species that comprise the range--in this instance, forest range--community. One of the most important variables in this context is sprouting characteristics of species, especially of those harvested. It was shown above, for example, how readily black oak coppiced (stump-sprouted) following clearcut logging. This was also shown earlier in this chapter for a sugar maple-northern red oak-pignut hickory-white ash-forest which was less than two miles distant from the lightening-induced forest gap. In that climax, mesic (north-slope), sugar maple-dominated forest most of the felled mature trees of sugar maple, American elm, basswood, pignut hickory, and northern red oak coppiced such that the climax tree species were still present in the recovering forest vegetation during the first year following clearcutting. In fact, most of the same genetic individual trees started regrowth and remained in the forest community though, of course, at the "infant" stage. On the clearcut forest there were all of the standard pioneering (weedy) grasses (hairy crabgrass, Japanese brome) , forbs (sow-thistle, mare's tail or horseweed, giant ragweed), and, even, trees (sassafras), but there were also climax, native, herbaceous species such as wildryes, hairy bromegrass, and woodland bluegrass.

Still, size or area of disturbance was a factor in species composition of the disturbed range vegetation. Larger areas of defoliation (eg. clearcut forest) was more apt to have seral plant species and undergo secondary plant succession (at least components thereof) than small localized areas of disturbance characterized by death or defoliation of a single or few individual plants, even when these individuals were comparatively large.

To summarize, changes in vegetation within small disturbed areas (patch dynamics) such as forest gaps are less dramatic and dynamic than changes in plant communities within larger disturbed areas like clearcuts. In extremely small forest gaps, for example, plant succession might not occur because injured or damaged individuals either regrow (eg. coppice shoots from stumps) much like grasse and forb regrow following mowing and/or sexual progeny spring up from the soil seed bank. Instead of the lengthy and complex pattern of secondary--not to mention primary--succession, recovery of range vegetation from denudation might be a simple matter of progression of the same climax species or, even, the same genetic plants through their life cycles. In the case of the forest gap created by death of an adult, lightening-struck pignut hickory, forest recovery was a simplier, shorter-term matter of seedlings of pignut hickory and, to somewhat lesser degree, of other climax tree species growing to adulthood. Instead of forest gaps being "new land" of a denuded area (an old field, "go-back ground", or cutover forest) and beginning plant succession at the initial seral (pioneer) stage, revegetation of the climax community ( black oak-pignut hickory forest in this instance) was a matter of progressing through the life cycle of a new cohort of the sporophytic generation of climax (especially, dominant) plant species.

In essence, if the disturbance gap is small enough plant succession does not occur. Instead, individuals (sexual and/or asexual in origin) of the climax species just grow to mature, adult-sized plants. In such cases, there is little if any change in plant species composition of the disturbed vegetation. Climax species composition remains over the course of recovery. Changes in vegetation on small denudation gaps (patch dynamics) are limited to those of plant community structure (seedling or stump sprouts through old-growth size and form). In such circumstances development is growth, tissue differentiation, maturation, and reproduction of plants (ie.species-specific plant life cycle) not development of vegetation (ie. plant succession).

205. A better kind to bust through- Carrion-flower (Smilax herbacea) is the only Smilax species in this area of the Oak-Hickory Forest Region that is not a horribly thorned, tough liana known variously as green-briar, cat-briar, bristle-briar, etc. S. herbacea is an herbaceous vine without hard, sharp, thorn-like stem appendages. Carrion-flower was one of the more plentiful forbs on the forest gap formed by lightening-induced death of the mature pignut hickory as detailed above. It also provided students with an example of a monocotyledonous forb.

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