Miscellaneous Forest Types- IB

Riverarine Forests (Including Gallery Forests) of Humid to Semiarid Zones

Riverarine (of or relating to rivers) forests are a varied group of forests (including woodlands, tree-dominated communities in which crowns of trees do not come into contact) developing along and stretching for different widths away from channels of larger streams. Some riverarine forests and woodlands extend across broad floodplains for considerable distances from river channels whereas other riverarine forests form much narrower communities that are confined closer to the main stream channel. Gallery or fringing forests, a designation more commonly used with tropical forests, refers to forest that develop along the corridors of streams from the stream bank (riparian zone) and extending outward to where bottomland or floodplain woody vegetation contacts grassland and/or savannah (Allaby, 1998). The term thus includes both the riparian zone and the outermost or perimeter zone(s) of forest (or woodland) than is (are) directly influenced by the stream. In perception gallery forests are tree-dominated plant communities that are relatively narrow being limited to habitats that are influenced directly by the river.

In this section gallery forests from the contintental interior of North American grasslands were described and used as representative examples of this general category or group of forest range vegetation.

Niobrara River

The Niobrara River has one of the most botanically and structurally diverse gallery forest in the Western Range Region. The Niobrara River was designated as the Niobrara National Sceinci River (at least 200 miles of it was) in 1991, but much remains to be done with regard to specific management if it is indeed to be saved and not turned into an irrigation ditch like most of the mid-sized rivers of the North American heartland.

The Niobrara is a braided stream, "a stream that divides into or follows an interlacing or tangled network of several small branching and reuniting shallow channels separated from each other by ephemeral branch islands or cannel bars, resembling in plan the strands of a complex braid" (Wilson and Moore (1998). The definitive work on Niobrara (including geological, biological, and cultural aspects) and was that of Johnsgard (2004).

120. Forest lined- Niobrara River with its floristically rich and structurally diverse gallery forest. There were some open areas of tallgrass prairie, but most of this riverarine range vegetation was comprised of tree-shrub-dominated plant communities. Riverarine vegetation of this gallery forest was descripbed in captions below. Forest Descriptions of the forests and adjoining grasslands along the Niobrara River included Barker and Whitman (1989, p. 18-20), Kantak (1995) and Johnsgard (2004, ps. 45-47), the latter of whom drew much from Kantak (1995).

The hardwood zone of this gallery forest would be of the Elm-Ash or Elm-Ash-Basswood (Plains Haradwood Type) described by Barker and Whitman (1989, p.19).

Cherry County, Nebraska. Late June (estival aspect). Range vegetation descriptions given in following captions.

121. A diverse lot- Gallery forest along the Niobrara River. Vegetation of this gallery forest extended from the riparian zone of sandbar willow (Salix exigua), peachleaf willow (S. amygdaloides), red osier dogwood (Cornus stolonifera), and some chokecherry (Prunus virginiana) through a middle zone of green ash (Fraxinus pennsylvanica), American elm (Ulmus americana), box elder (Acer negundo), basswood or American linden (Tilia americana), eastern cottonwood, and bur oak (Quercus macrocarpa) and hackberry (Celtis occidentalis) to the highest and outermost forest zone dominated by ponderosa pine and eastern red cedar (Juniperus virginiana). Obviously there was considerable overlap of many of these species especially in the broad mid-zone to lower parts of the outermost forest edge. Shrubs (in addition to those listed for the riparian zone) included riverbank grape (Vitis riparia), Virginia creeper (Parthenocissis quinquefolia), thicket creeper (P. vitacea) and poison ivy (Toxicodendron radicans= Rhus radicans ).

There were very few forbs in this dense forest, at least at this late spring-early summer season. Most grass was along the moist edge of the bank and consisted of naturalized mooth brome. There was some bottomland switchgrass and Canada wildrye. Also some caric sedges (Carex spp.).

Cherry County, Nebraska. Late June (estival aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest) for the central hardwood zone while the lower or riparian zone could be interpreted as K-89, Northern Flood Plain (Populus-Salix-Ulmus). SAF 235 (Cottonwood-Willow) for riparain zone; SAF 93 Sugarberry (Hackberry variant)-American Elm-Green Ash for central zone; SAF 237 (Interior Ponderosa Pine) for driest, upper zone.Not a relevant unit for most of this forest in Brown et al. (1998).Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001).

122. Braided and lined- This view spanning the Niobrara River showed the stream channel feature of this braided stream. Braided streams have usually been interpreted as having loads of sediment that exceed their capacity to carry them (Wilson and Moore, 1998). The result is deposition of these overloads in the network of small, shallow interchannels and channel bars. Range vegetation along this stretch of the Niobrara River was described in the next two sets of slides.

Cherry County, Nebraska. Late June.

123. Farthest edge of sandhill gallery forest- At uppermost (least mesic) zone of the gallery forest hat developed along the Niobrara River ponderosa pine and easter red cedar co-cominated the vegetation. This was where the forest contacted tallgrass prairie of upland switchgrass, big bluestem, little bluestem, sideoats grama, and blue grama with some cheatgrass. The most abundant (about the only) shrub was smooth sumac (Rhus glabra). Ponderosa pine had invaded the grassland here at the dege of these two types of range vegetation. This may have been due to greater tolerance of this species to fire.

Cherry County, Nebraska. Late June. Forest community was 21 (Pondaerosa Pine Forest and Woodland Ecosystem). K-15 (Eastern Ponderosa Pine Forest with Eastern Red Cedar do-dominant). SAF 237 (Interior Ponderosa Pine) In Brown et al. (1998, p. 37): eastern form of Yellow Pine Series, 122. 62) Grassland was FRES No. 39 (Prairie). K-67 (Nebraska Sand Hills Praire) or, perhaps, more precisely at this location, K-66 (Bluestem Prairie). Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001).

124. Forest meets prairie in the Nebraska Sandhills-Panarama of uppermost zone of gallery forest above Niobrara River coming into contact with tallgrass prairie. At higher (and drier) portions of this riverarine forest some scattered boxelder and bur oak joined co-dominant ponderosa pine and eastern red cedar to make a "last stand" at edge of the regional climax of tallgrass prairie. The prairie was dominated by upland switchgrass with big bluestem, little bluestem, and Canada wildrye associated major species.

Some Kansas Rivers

In the following section five examples of forests were described that developed along rivers or tributries of rivers in northcentral Kansas. This sample included three gallery forests and two bottomland forests that developed on backwaters of river tributaries. These rivers were in the extreme eastern portion of the Great Plains physiographic province, the Plains Border (Fenneman, 1931, ps. 25-27). There has been some confusion and controversy in the literature concerning the proper physiographic treatment of this area; specifically whether it is part of the Great Plains or the Central Lowlands physiographic province. Frye and Swineford (1949) concluded that Fenneman was incorrect as to the western boundary of the Plains Border, but that his delineation of the eastern boundary (at the western edge of the Flint Hills) was correct. Most precisely, this is the Smoky Hills Upland adjacent to the Flint Hills Upland to its east.

In these forests eastern cottonwood was one of the dominant and defininng species, but cottonwood was a seral and not a climax species. This was in contrast to forests treated above in which eastern cottonwood was the climax dominant. Iin SAF 63 (Cottonwood) forest cover type of southern and eastern bottomlands (Eyre, 1980, ps. 62-63) cottonwood (of several species) was interpreted as a temporary pioneer that was replaced by the next seral stage made up of pecan, sugarberry and/or hackberry, green ash, American elm, sycamore, boxelder, and silver maple with the sere terminating in a silver maple-American elm-boxelder climax. In SAF 235 (Cottonwood-Willow) of western lowlands--as from the Great Plains westward--(Eyre, 1980, p. 113) cottonwood (again of several species) comprises an edaphic/topographic climax ("a climatic anamaly") or as "postclimax type", "a subclimax stage of the hydrosere" (Eyre, 1980, p. 113). This latter interpretation was that of the classic Clementsian monoclimax theory. Eyre (1980, p. 113) specified that in the eastern Great Plains SAF 235 "merges with the cottonwood type [SAF 63] or silver maple-American elm". Thus, it was not clear as to which SAF forest cover type (63 or 235) and, hence, successional status the cottonwood-containing forests presented below were best described by.

125. Along and around the bend- Gallaery forest along the Republican River (in early morning light). This was a botanically rich forest from standpoint of tree species which included eastern cottonwood, hackberry or western hackberry, green ash, boxelder, American elm, black walnut, northern catalpa (Catalpa speciosa), bois d'arc or Osage orange or, even, hedge apple (Maclura pomifera), black locust (Robinia pseudoacacia), and red mulberry (Morus rubra). Catalpa and bois d'arc in this area of unknown origin. There is some evidence that neither of these species is native this far north and west and instead owes its existence here to introduction by white man. If this was the case they are still naturalized and part of the forest range vegetation. Red mulberry was present as an understorey tree (the only tree species that at maturity remained in the lower tree layer). There was some regeneration of all tree species, including eastern cottonwood.

There were relative few shrubs in this forest other than buckbrush or coaralberry (Symphoricarpos orbiculatus), riverbank grape (Vitis riparia), and Virginia creeper. Grasses were also scarce. The grass species was Virginia wildrye. Naturalized smooth brome and Japanese chess were present though more at forest edges.

In some parts of this forest there were two rather distinct zones running somewhat paralleal to the river. The zone closest to the stream (more common on river bank) included boxelder and had relatively less cover of eastern cottonwood. The zone farther from the stream consisted of more cottonwood and included bois d'arc, black walnut, and catalpa. Green ash, American elm, and hackberry were fairly evenly distributed except they were less common on the river bank.

Brown coloration of water was due to heavy loads of sediment from heavy late spring rains and flooding.

Clound County, Kansas. Late June (estival aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-89 (Northern Floodplain Forest). Combination of SAF 63 (Cottonwood) and SAF 93 ([Hack]berry-American Elm-Green Ash). In Brown et al. (1998, p. 43): for the eastern cottonwood-black willow zone was either Plains and Great Basin Riparian Deciduous Forest 222.2, Cottonwood-Willow Series 222.21 or Southwestern Riparian Deciduous Forest and Woodland 223.2, Cottonwood-Willow Series 223.21 and for the green ash-hackberry-northern catalpa-bois d'arc-box elder-black walnut mixture would have to be Southwestern Riparian Deciduous Foreest and Woodland 223.2, Mixed Hardwood Series 223.22. Central Great Plains-Smokey Hills Ecoregion, 27a (Chapman et al., 2001).

126. On the edge- Outer margins of a gallery forest along the Republican River contacted tallgrass prairie on the eastern border of the Great Plains. These two "photo-plots" showed the "advancing front" of tree species as far as they could extend (or, at least, had extended) to the prairie. These were small (sapling to small-pole size) eastern cottonwood and hackberry (some of the taller poles) along with some black locust. The latter were present as the lower (more of shrub size) and darker-green plants at right midground in these photographs. A northern catalpa was visible at far left margin in the first photograph.

The dominant prairie species on these more mesic habitats was bottomland switchgrass. Other prairie grassses included big bluestem, prairie cordgrass (Spartina pectinata), eastern gamagrass (Tripsacum dactyloides), prairie dropseed (Sporobolus heterolepis) , western wheatgrass (Agropyron smithii), and inland saltgrass (Distichlis stricta). Naturalized smooth bromegrass was present, but it was most common in locally disturbed areas (eg. spots impacted by flooding). The major forb was Illinois bundleflower (Desmanthus illinoinsis).

127. Points of contact- Two "photo-plots" furnished a distant and a close-in perspective of the contact zone between a gallery forest and tallgrass prairie on the eastern boundary of the Great Plains. The largest trees (treeline in background) were eastern cottonwood followed by hackberry and some green ash.Tree species in midground of both photographs were mostly saplings and small pole-size eastern cottonwood with some black locust. Locust were readily distinguished by the smaller, darker-green leaves and loer height compared to cottonwood (center to right margin of first slide; center to left margin of second slide).

Dominant grass was bottomland switchgrass. Other grasses included prairie cordgrass, big bluestem, eastern gamagrass, prairie dropseed , western wheatgrass, and inland saltgrass and the naturalized perennial, smooth bromegrass, and the Mediterranean annual, Japanese brome. These introduced Eurasian grasses were most common on disturbed areas, due to flooding and past highway construction. Illinois bundleflower was the most common forb.

128. Frontier sentinels- On the edges of gallery forest and adjoining tallgrass prairie representative plant species held their "last bit of ground" like soldiers on a military frontier. Trees were mostly eastern cottonwood with some hackberry as the local associate tree species. A large bois d'arc (right center background or treeline) represented its species.

It was noted above that bois d'arc or Osage Orange is probably not native this far north and west, but was likely introduced by white settlers who planted this thorn-beraing, close-growing species as a hedge, a living fence, to enclose livestock. This was undoubtedly an Anglo tradition transported from the British Isles. Hedge rows did not work (at least not well) and livestock, especially swine, strayed and ranged freely. When barbed wire was invented and adopted immediately by sod-busters trying to fence out cattle to protect their crops it was soon learned that bois d'arc made the most rot-resistant, some of the strongest (and, sometimes, the crookedest), and easiest to use post (they did not have to be split for example). Not only was Osage orange or bois d'arc excellent for hunting bows (reflected in these two common names), but its wood has the greatest longevity when in contact with soil of about any tree in North America (certainly any on the prairie and especially when compared to cottonwood or hackberry).

The small rise in the foreground had been disturbed by human activity (part of a staging area for road construction equipment) which permitted invasion by the rhizomatous, Eurasian (and now naturalized) perennial smooth bromegrass. The native, wide-leafed, rhizomatous eastern gamagrass had established a "roothold" in this bromegrass colony following the anthropogenic disturbance therby attesting to the competitive ability of this "ice cream species".

129. Deep inside- Interior of a gallery forest on a tributary of the Smoky Hill River. Largest trees (abackground of al three photographs) were eastern cottonwood, but smaller trees (saplings and seedlings) were green ash and hackberry indicating that these were replacing cottonwood which had regeneration only in certain local spots. Red mulberry was the dominant of a lower tree layer (the saplings in foreground in all photographs) and the associate tree species of this gallery forest. Redbud was well-represented as a tall shrub species. Buckbrush and wild or Missouri gooseberry (Ribes missouriense), foreground of the second and third photographs, comprised a lower shrub layer. There were almost no herbaceous species in the understorey within the forest interion. Instead grasses and forbs made an herbaceous layer only at edges of this gallery forest. Virginia wildrye, climax decreaser, was the dominant herbaceous plant. Species of the forb in foreground of econd photograph could not be identified in its early vegetative stage.

McPherson County, Kansas. Late June (estival aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-89 (Northern Floodplain Forest). Combination of SAF 63 (Cottonwood) and SAF 93 ([Hack]berry-American Elm-Green Ash). In Brown et al. (1998, p. 43): was either Plains and Great Basin Riparian Deciduous Forest 222.2, Cottonwood-Willow Series 222.21 or Southwestern Riparian Deciduous Forest and Woodland 223.2, Cottonwood-Willow Series 223.21. Central Great Plains-Smokey Hills Ecoregion, 27a (Chapman et al., 2001).

130. Shaded red- Shade leaves of red mulberry (Morus rubra) growing beneath sugarberry or smooth hackberry and hackberry or western hackberry.

Ottawa County, Oklahoma. July.

131. Straight (but not narrow) and muddy- Gallery forest along the straight channel of the Salina River in northcentrtal Kansas at eastern border of Great Plains. Rivers have a river of channel pattern, "the configuration in plan view of a limited reach of a river channel as seen from an airplane" (Wilson and Moore (1998). River patterns include meandering, braided, sinuous, and relatively straight. The Niobrara River covered above was an example of a braided river. The Salina River provided an example of the relatively straight pattern, perhaps the least common of these major recognized river patterns. Although this photograph was not taken from the air and it did not show an extremely long stretch of the river it did provide a limited example of the relatively straight river channel.

This gallery forest was dominated by eastern cottonwood and black willow (Salix nigra). There were younger, smaller trees of green ash and hackberry and less abundance of box elder and American elm. Presence of fewer and smaller individuals of these late seral or climax tree species together with limited regeneration of cottonwood and willow indicated clearly that this forest was undergoing plant succession toward the terminal Ash-Elm-Hackberry forest.

There were only a few shrub species (excepting willow) and even fewer herbaceous species and cover beneath the crowns of eastern cottonwood and the seedlings and saplings of green ash, the dominant tree species of the lower tree layer. Herbaceous cover was limite mostly to edges of the forest or local areas disturbed by flood water.

Saline County, Kansas. Late June (estival aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-89 (Northern Floodplain Forest). Combination of SAF 63 (Cottonwood) and SAF 93 ([Hack]berry-American Elm-Green Ash). In Brown et al. (1998, p. 43): was either Plains and Great Basin Riparian Deciduous Forest 222.2, Cottonwood-Willow Series 222.21 or Southwestern Riparian Deciduous Forest and Woodland 223.2, Cottonwood-Willow Series 223.21. Central Great Plains-Smokey Hills Ecoregion, 27a (Chapman et al., 2001).

132. Forest-lined Salina- Oblique view of a gallery forest dominated by eastern cottonwood and black willow that developed alone the Salina River. Almost of the large trees were cottonwood while black willow (eg smaller tree or large shrub arched over the river in foreground) formed a second or lower tree layer. The area in the foreground was on the edege of oan pening in the forest. Part of this had been disturbed by flooding that removed some soil and deposited alluvium to replace it (or on a spaace nearby) while other parts had been spared and supported dense colonies of climax Virginia wildrye. The flood-perturbed area had been populated by herbaceous pioneer species including giant ragweed and annual sunflower (Helianthus anuus).

Local spots that were still recovering from disturbance where giant ragweed and annual sunflower made dense stands.

133. Salicaceous sistern- Eastern cottonwood and black willow co-dominated a gallery forest along the Salina River on the eastern border of the Great Plains. These two species are Intolerant species (Wenger, 1980, p.3) that pioneer such river habitats as gravel bars and mud flats along channels. Most members of the Salicaceae (willow family) have the successional role of pioneers or early seral species. These species sometimes persist as old individuals into advanced successional stages, perhaps a few of the more favored ones survive to climax. Cottonwood and willlow clearly do not dominate advanced or climax stages. Unlike gallery forests described above that were composed of a variety of woody species, the fringing forest shown here remained dominated by salicaceous species clearly indicating the early successiona status of this forest range community.

The foreground had colonies of Viirginia wildrye, a climax festucoid grass, but it also had local stands of giant ragweed and dense stands of the Eurasian smooth brome on local spots that had been disturbed by action of flood water.

134. Old and young; seral and climax- "Photo-plots" of a stand of gallery forest along the Salina River featuring internal structure and species composition. The large, adult trees were all eastern cottonwood. These trees were still in the "prime" being remarkably free of storm damage, dying crowns, etc. Growing beneath these mature eastern cottonwood were green ash of various age/size classes frnginf from seedling through small-pole size. Pole-size ash were most prominently featured in the third of these three slides although one pole was also in the first and, though much less, in the second slide (far left, lower corner). There was very little regeneration of eastern cottonwood in this forest which highlighted the seral nature of this currently dominant tree species and the more successionally advanced feature of green ash. There were also some small saplings of hackberry in lower layers and a few young box elder (eg. center foreground of second slide), but green ash was the clear "dominant-to-be" of this forest vegetation.

There were some secondary shoots (sprouts) from debarked parts of lower cottonwood trunks. These small sites of injury appeared to have been caused by logs and other debris carried and shoved into trees by force of flood water. This photographer did not observe any feeding activity by beaver (Castor canadensis) any of which would most likely have induced resprouting of cottonwood.

Shrubs were not present (at least not in the area presented here). Herbaceous plants were primarily limited to the climax grass, Wirginia wildrye, although the naturalized agronomic species, smooth bromegrass, was present in localized colonies (especially in disturbed areas next to the adjoining hay field of smooth brome and where flooding had previously denuded parts of the river bank).

135. Relict stand- Photographic dendrogram of a climax bottomland forest of green ash-hackberry-eastern cottonwood on backwater of the Solomon River in central Kansas. Green ash was the dominant species while hackberry was the associate and eastern cottonwood a persistent species (since pioneering the site). Black walnut was present as a minor but indicative species. This was a remnant of old-growth forest on private property that had been spared clearing apparently because it was not cost-effective to drain this small a parcel of wetland. The adjoining private property was also in a climax green ash-hackberry forest but this tract had been taken by the state of Kansas through eminent domain and made into a rest area-roadside park. The small tract of old-growth green ash-hackberry-eastern cottonwood shown here and the contiguous tract dominated by green ash constituted a single example of this range cover type.

The lower layers of this stand (not discernible in this photograph) included a lower shrub layer dominated by buckbrush and an herbaceous layer dominated by Virginia wildrye and various forbs. Where edges of the herbaceous layer(s) of this forest vegetation came into contact with a field of smooth brome there was some invasion of this agronomic grass into the forest stand, but in general this introduced forage species could not compete with the native Virginia wildrye. Details of this forest range vegetation were covered below.

This photograph presented the physiogonomy and general architecture of the green ash-hackberry bottomland forest type. Number of dead limbs in crowns of green ash was typical of old-growth (and "over-ripe") trees.

Ottawa County, Kansas. Late June (estival aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-89 (Northern Floodplain Forest). SAF 93 ([Hack]berry-American Elm-Green Ash). Southwestern Riparian Deciduous Foreest and Woodland 223.2, Mixed Hardwood Series 223.22 (no Mixed Hardwood Series in Plains and Great Basin Riparian Deciduous Forest 222.2). Central Great Plains-Smokey Hills Ecoregion, 27a (Chapman et al., 2001).

136. Dead and dying, but stiill capable of regeneration- Od-growth stand of green ash, a high proportion of which were dead or dying, on backwater wetland of the Solomon River. This was an example of trees that were overmature, "a tree or even-aged stand that has reached that stage of development when it is declining in vigor and health and reaching the end of it natural life apan" or "a tree or even-aged stand that has begun to lessen in commerical value because of size, age, decay, or other factors--note the term has little applicability to uneven-aged stands, which consist of trees of diverse ages and stages of development " (Helms, 1998). The first of these definitions would be most relevant to the stand presented here dimply because this forest was not being used as a commercial forest. Also, overmature was appplied to individual trees (as in both definitions) not the stand which had much regeneration of green ash and hackberry, the climax dominants.

All living female trees (ash species are dioecious) in this stand, including those "more dead than alive", had produced abundant crops of smaras (the dry, indehiscent, one-seeded, winged fruit of ash). In addition, there were different age/size classes of progeny ranging from seedling to saplings; however there were few pole-size ash. There were individual hackberry (associate tree species) of pole-size, but fewer seedlings and saplings of this clearly secondary tree species. in this observer's experience ash species produce fruit sporadically with nothing like the regularity of such hardwoods as oak, hickory (eg. pecan), hackberry, cottonwood, willow, elm, walnut, bois d'arc all of which were associated with green ash in this region. As such, sexual reproduction in ash is more uncertain and episodic in ash (again, in this author's observation) than in associated angiosperms. Saplings (younger trees generally) readily sprout from stumps of green ash (Burns and Hankala, 1990). This silvic feature permits asexual (vegetative) reproduction following fire, flood, windthrow, beaver feeding, etc.

The dominant shrub was buckbrush or coralberry. The understorey dominant was the climax grass, Virginia wildrye. Lower layers of this stand, including reneneration of climax trees, was treated later in this section.

The foprest vegetation shown here was on one edge of the forest tract of which one part was presented in the preceding photograph and other local stands that were described later. This green ash-hackberry-eastern cottonwood (this latter an uncommon though persistent pioneer) had been privately owned and "let be" presumedly because it was such a small tract of forested wetland that it would not be profitable to clear and plant it to smooth brome, the surrounding field crop (ie. beneficial neglect). Kansas condemned the property and built a rest area on part of the land along the highway. The Department of Transportation then commenced close, repeated mowing (shredders or rotary mowers were used) of the back portion of the rest area that was considerable distance from restrooms. Apparently this was a rest area for travelers' dogs the excreta of which fertilized an oversized yard of annual grasses. Seedlings of green ash, hickberry, cottonwood, and black walnut were killed by mowing as were native perennial species such as Virginia wildrye, the herbaceous dominant of the natural forest. Such waste of tax dollars prevented any forest regeneration and left an overgrazed yard that resembled a city lot.

The back part of the rest area was not being used except as an unhealthy disposal for dog dung. "Maintenance" mowing was mostly make-work for government employees. One would think that Kansas, one of the states with the smallest percentage of land in forests, would appreciate the beauty and, especially in this case, watershed value of a native forest. Instead, mindless, make-work mowing had reduced a recently diverse natural forest fo a few dead and dying over-ripe trees surrounded by manicured crabgrass. "Go figure", but pay attention and watch where you step. .

137. Synopsis of a stand- Interior of the climax stand of green ash-western hackberry that was introduced in the immediately preceding photograph. Emphasis in this "photo-plot" was on structure and general species composition deep inside this bottomland forest range that had developed on a backwater drainage of the Solomon River.Most of the adult trees were green ash, almost all females of which bore heavy crops of ripening fruit, with some of western hackberry which was the associate species. The clear dominant of the herbaceous part of the understorey was Virginia wildrye. At edges of this wetland forest that contacted a field of smooth brome there was some invasion of this naturalized agronomic grass "just inside the line" of the green ash-dominated stand. Nodding foxtail or nodding bristlegrass (Setaria faberii), a naturalized native of east Asia, was also present near margins of this forest. There were fewer, though notable, individuals of orchardgrass (Dactylis glomerata), another Eurasian agronomic grass, that grew to typical size in this forest interior. Forbs included pokeweed or pokeberry (a local dominant in more sunlite spots), poison hemlock (Conium maculatum), a deadly poisonous and naturalized exotic, elderberry, and curly dock (Rumex crispus), another naturalized--and sometimes poisonous--Eurasian weed, and, remarkably, numerous individuals of giant ragweed. (More discussion on forbs followed in captions below.)

There was abundant regeneration of both green ash and hackberry. A nice example of hackberry reproduction was the small sapling in far left margin of this photograph. A plentiful crop of ash seedlings and small saplings was obvious in right foreground. Regeneration of climax tree species was discussed in the immediately succeding caption. All larger trees in this view were green ash except for one hackberry in right midground.

138. Lower layers- Details of the understorey of a climax green ash (dominant)-hackberry (associate) bottomland forest on a drainage area of the Solomon River in central Kansas. These two photographs also featured (like the preceding photograph) th interior of this forest range vegetation. The obvious herbaceous dominant was the decreaser grass, Virginia wildrye. There was abundant regeneration of both green ash hackberry. As a general rule all of the North American ash species are interpreted as Intermediate in tolerance rating (Wenger, 1980, p. 3). Burns and Hankala (1990) specified that green ash "... varies from intolerant to moderately tolerant to shade in the northern part of its range" whereas "[i]n In the southern part of its range, green ash would be considered tolerant when young and moderately tolerant as it grows older". Burns and Hankala (1990) noted further that on some forest sites green ash responds as a pioneer (eg. as on alluvium) along with cottonwood and lack willow (25). In sum, the prolific regeneration of green ash in this stand was consistent with that of a climax species for this forest range site. Hackberry " is intermediate to tolerant in its ability to withstand shade" with a successional status extending to subclimax though "...its successional position is difficult to determine" (Burns and Hankala, 1990).

All of the trees in the first photograph were green ash. A nice pole-size hackberry--complete with warty bark--was featured in the second photograph.

Dominance of the herbaceous component of this climax forest by Virginia wildrye was obvious in both of these slides. Numerous specimens of giant ragweed and pokeweed were conspicouos in the second slide. Presence of giant ragweed, an annual pioneer on distrubed sites, in a dense colony of Virginia wildrye, a climax, cool-season, perennial grass seemed unusual until it was realized that such a combination might facilitate efficient use of resources in lower layers of this wet forest. Also in the second "photo-plot" was tall or pasture thistle (Cirsum altissimum), a native, biennial composite. Perhaps, biennialism in conjuction with warm- and cool-season and annualism and perennialism allowed even more explotiation of limited resources. Shrubs were lacking in this stand even though buckbrush was found in other parts of this forest tract.

139. Structure of a stand- Paired photographs (the first presented a taller and more distant view vs. a shorter and closer view in the second) the outer margin of a bottomland forest stand dominated by green ash with hackberry the associate species. Emphasized was stand structure and species composition (a lot of biodiversity) at edge of this forest community in order to compliment description of the forest interior treated in the last two photo-captions.

This forest tract came into contact with a field of smooth brome into which it was slowly invading. At this contact there were some localized stands of both cheatgrass and Japanese chess, two naturalized Mediterranean grasses. Smooth brome had invaded the herbaceous layer of the forest at some areas of contact between these two plant stands. Orchardgrass, another exotic agronomic grass, was also present in trace amounts in this understorey. Somewhat more common, though nowhere near abundant. was the Eurasian weed, curly dock. With even less cover was poison hemlock (right margin of seccond slide) Elderberry (middle of midground in both slides; more obvious in second slide between ash trunks) as was pokeweed, giant ragweed, and tall thistle.

All adult trees were green ash. Absence of shrubs was notable.

140. Composition of a stand- Another view of the forest vegetation presented in the preceding paired photographs. This perspective was diagonal to the view presented in the second of the preceding pair of slides (use forked trunks in left margin of both as a bearing). Forb species were more prominent in this diagonal perspective especially curly dock, giant ragweed, pokeweed, and tall thistle. Virginia wildrye was everywhere.

Shrub species were almost non-existant in this portion of the stand of green ash-dominated wet forest. Buckbrush was common locally in this forest tract though not in this more pristine stand.

141. Layered sequence- A triololgy of ever closer views of the layers in a stand of green ash on a drainage of the Solomon River at extreme eastern border of the Great Plains. This was at the outer margin of the stand of mixed age green ash introduced above. Species included a mixed assemblage of cheatgrass and Japanese chess (light brown straw in foreground of first two slides), smooth brome (foreground) poison hemlock (left foreground of first two slides), elderberry (to left of biggest trunk in all slides), pokeweed, giant ragweed, and Virginia wildrye, the overall dominant herbaceous species.

An interesting phenomenon was shown with the dead shoot of a sapling of red mulberry (right foreground of the three slides). An extremely light surface fire had burned a short distance into this forest in the previous year with such low intensity that it left little evidence of its passing. It was, however, enough to topkill the red mulberry whidh had already resprouted . Examples of red mulberry as a lower layer tree were presented above.

142. Gnarled nurse and straight suckling- Textbook example of the nurse plant phenomenon. An old-growth eastern cottonwood that had pioneered a backwater area of the Solomon River provided a nurturing microhabitat for a western hackberry that had grown to small-pole size. On this wet forest site hackberry and green ash were dominants of the potential natural vegetation in which some eastern cottonwood persisted at least to subclimax stage) lo. This was the outermost edge of this forest where it contacted a field of smooth brome. Recent road construction (including a rest area) had set the forest sere back to bare soil so that the herbaceous plants in foreground were annual pioneers like giant ragweed, horseweed or marestail (Conyza canadensis), lambsquarter (Chenopodium album), hairy crabgrass (Digitaria sanguinalis), and annual sunflower. Future visits should reveal a nice crop of eastrn cottonwood seedlings. The cycle goes on..

143. Hackberry on the (sort of) prairie- Forest dominated by hackberry or western hackberry on an upper drainage of the Solomon River in central Kansas at edge of what remained of a a wet prairie. Featured here was hysiogonomy and general crownline of a consociation of Celtis occidentalis (accompanied by a few scattered American elm, green ash, eastern cottonwood, bois d'arc, and northern catalpa) that formed along an upland drainage at edge of wet prairie. This prairie had been highly modified by human action when used as a staging area for highway construction. The original cover of bottomland switchgrass and prairie cordgrass had been greatly reduced by movement of heavy roaad equipment several years earlier. Smooth brome and reed canarygrass (Phlaris arundinacea) had replaced native prairie plants (at least for a time) on this wet prairie that was sustained by a small ephemeral stream that branched off of the the main channel of this small upstream river tributary. Shallow channel of this ephemeral branch was at center-right. The larger channel of the tributary was lined by a narrow forest that was mostly a stand of hackberry made of numerous age/size classes.

Various shrub species in this hackberry-dominated forest formed irregular layers in the vegetation. Shrubs included chokecherry (Prunus virginiana), the tallest shrub, buckbrush, the most common shrub and that formed a low shrub layer, and redroot (Ceanothus ovatus) at outer margins of the forest. The main herbaceous species was Virginia wildrye. This forest had a dense enough canopy that neither smooth brome nor reed canarygrass had invaded the forest understorey.

Tree species were slowly expanding out into the disturbed prairie at points of contact between forest and grassland. The tree seedlings out in the middle of the damaged wet prairie were bois d'are or Osage orange. It was explained above that bois d'arc is probably not native in this region but instead naturalized after having introduced by white settlers for use (largely unsuccessful) as a hedge or living fence. Invasion of Osage orange into the degraded grassland was an example of brush invasion on an old field or "go-back land".

Cloud County, Kansas. Late June (estival aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-89 (Northern Floodplain Forest). SAF 93 ([Hack]berry-American Elm-Green Ash). Southwestern Riparian Deciduous Foreest and Woodland 223.2, Mixed Hardwood Series 223.22 (no Mixed Hardwood Series in Plains and Great Basin Riparian Deciduous Forest 222.2). Central Great Plains-Smokey Hills Ecoregion, 27a (Chapman et al., 2001).

144. Great Plains conglomeration; woody expansion- Contact bewteen a hihgly man-modified wet prairie (formerly dominated by bottomland switchgrass and prairie cordgrass but largely converted to a stand of reed canarygrass with scattered colonies of smooth bromegrass) and a hackberry-dominated gallary forest. The prairie had been used during dry periods as a staging area for highway construction (heavy equipment parking area) with such destructive consequences that these two perennial Eurasian grasses successfully invaded the denuded grassland. Woody plants were also encroaching from edge of he adjacent forest into the former tallgrass paraire. This included some hackberry but mostly the native shrub, redroot. Chokecherry and buckbrush were also moving into the disturbed grassland. At other spots bois d'arc (probably not native to this area) had invaded (and was continuing to do so) the denuded area that was in effect an old field. The forest had not been impacted (at least not directly) by movement of heavy equipment.

The obviously old-growth hackberry was surrounded by various age/size classes of its race ranging from pole-sized trees of roughly half-grown adult dimensions (left of the patriarch tree) through large saplings (right side of patriarch) down to seedlings. There were some widely scattered small trees of American or white elm, green ash, northern catalpa, and even eastern cottonwood, the latter limited to open spots beneath the othrwise dense forest canopy. Shrub species on the forward margin of this forest were mostly redroot followed by buckbrush and chokecherry.

FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-89 (Northern Floodplain Forest). SAF 93 ([Hack]berry-American Elm-Green Ash). Southwestern Riparian Deciduous Foreest and Woodland 223.2, Mixed Hardwood Series 223.22 (no Mixed Hardwood Series in Plains and Great Basin Riparian Deciduous Forest 222.2). Central Great Plains-Smokey Hills Ecoregion, 27a (Chapman et al., 2001).

145. Gallary forest on fringe of Great Plains- Paired views of a hackberry-dominated fringing forest that developed along a tributary of tthe Solomon River at the eastern border of the Great Plains. First photogrpah presented the physiononomy and exterior structure of this wet-mesic forest. Second photograph showed species composition and layering of this forest where its outermost edge contacted a degraded tallgrass prairie.

This forest was a consociation of western hackberry (with incidental trees of American elm, grreen ash, and eastern cottonwood) which had the essential features of a climax forest. For all practical purposes all age classes of hackberry were presnnt ranging from the knotty old-growth specimen (left margin of both photographs) through pole-size, large and small saplings down to seedlings. The broad, oval-leafed shrubs at the most exterior edge of the woody range plant community, which included the hardwood forest plus its edge,(midground of first slide; foreground of second slide) was redroot. Accompanying this major shrub was chokecherry and buckbrush, the latter more common farther back and in interior of forest.

Grasses at this point of contact between deciduous forest and degraded tallgrass parairie were reed canarygrass and smooth brome, both introduced (from Eurasia) agronomic forage species. Reed canarygrass was more abundant ("choked out" even smooth brome) in wetter spots. There was also some now-dead straw of the Eurasian annual brome, Japanese chess. A few never-say-die, hangers-on of switchgrass and big bluestem were noted for the record.

146. Young trees along an old river- Sycamores ranging in age/size classes from sapling through pole to young adult had pioneered the banks of the Marais des Cygnes River in the Osage Cuestas section (of the Central Lowlands physiographic province) in eastern Kansas. Some disturbance(s), such as flooding or human dredging, in the "not-too-far-back" had denuded this section of the river channel enough to permit invasion of the "new land" by the Intolerant sycamore. The associate tree species on this "run" of river was honey locust, an example of which was at left margin just in front of young sycamore.

Present (though not visible in this slide) in more interior parts of this bottomland forest was American elm, silver maple (Acer saccharinum), and a few relict giant eastern cottonwood whichwere not reproducing. The lower layers of this floodplain forest were limited due to extremely dense shade. Widely scattered understorey species included woody vines or creepers which ascended into the forest canopy. These included poison oak-ivy, Virginia creeper, and riverbank grape (Vitis riparia). There were local small patches of buckbrush or coralberry (Symphoricarpos orbiculatus) The main understorey herbaceous species was giant ragweed on freshly flood water-scoured "new ground" and pokeberry or pokeweed on land with some established plants. Farther away from the dense tree canopy the main herbaceous species was poison or spotted hemlock (Conium maculatum) as was shown below.

Brown (sediment-laden) water of this river was testament to recent heavy rains. Such heavy loads of sediment are what made bottomlands so fertile. This provided students with an example of riparian vegetation, the plant community that grows closest to the stream (ie. on the stream bank, less-frequently flooded parts of the river bed, and immediately adjacent floodplain).

Osage county, Kansas. Mid-June (late spring). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-89 (Northern Floodplain Forest). SAF 94 (Sycamore-Sweetgum-American Elm). Not a good unit in Brown et al. !1998, p. 43), but Warm Temperte Swamp and Riparian Forests (223), Southwestern Riparian Deciduous Forest and Woodland (223.2), Mixed Broadleaf Series (223.22) came closest. Western Corn Belt Plains, Loess and Glacial Drift Plains Ecoregion 47i (Chapman et al., 2001).

147. Rich array along another river- The Delaware River running brown with sediment through the glaciated land of northeastern Kansas during spring flood time. There was quite a "cast of character species" aligned along the fertile bank of this shallow stream. Tree species included sycamore, black willow, black walnut, basswood or Ameriocan linden, American or white elm, silppery or red elm, silver maple, boxelder, honey locust, and green ash BUT no cottonwood in this streatch of river forest. The extra-dense shade of this diverse-species forest had very limited lower layers of vegetation except for small openings (as were a tree had died liaving a forest gap). These opening, like newly deposited mud along the banks and higher bed portions of the river, supported mostly giant ragweed and naturalized smooth brome. Woody vines such as poison oak/ivy, Virginia creeper, and riverbank grape grew up into the forest canopy as if to unit the various layers of this forest range.

The range plant community seen here was riparian vegetation being that which grew on the river bank and immediate floodplain.

Lower seral stages resulting from past disturbances supported a species-rich understorey as well as pioneer or colonizing tree species. An example of this stage of species diversity was presented in the slides immediately following this caption.

Brown County, Kansas. Mid-June (late spring). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-89 (Northern Floodplain Forest). SAF 94 (Sycamore-Sweetgum-American Elm). Not a good unit in Brown et al. !1998, p. 43), but Warm Temperte Swamp and Riparian Forests (223), Southwestern Riparian Deciduous Forest and Woodland (223.2), Mixed Broadleaf Series (223.22) came closest. Western Corn Belt Plains, Loess and Glacial Drift Plains Ecoregion 47i (Chapman et al., 2001).

148. Silver stream star- Silver maple (Acer saccharinum) is one of the "also-ran-species" of tree in riverine forest of the humid to subhumid zones. It was not a dominant or even associate (other than in microsites) tree in bottomland forests in more western portions of the Central Lowlands province, but it was consistently present species. For example, silver maple was one of the trees in floodplains of the Delaware and Marais des Cygnes Rivers of eastern Kansas. Thus, the appropriateness of its inclusion here.

Newton County, Missouri.. Late April.

149. Silver samaras and leaves- Leaves and fruit along with a typical twig of silver maple. The fruit of Acer species has been interpreted variously as either a schizocarp or samara.. Smith (1977, p. 165) offered the options, "... as either two samaras, separating at maturity, or a samaroid schizocarp, separating into two-winged mericarps at maturity". Smith defined samara as a dry fruit type that is indehiscent with a single seed and an attached conspicuous wing-like appendage (ie.a winged achene) and schizocarp as a dry indehiscent fruit originating from a syncarpous gynoecium splitting at maturity into one-seeded closed segments called mericarps (Smith, 1977, ps. 66, 307).

Newton County, Missouri.. Late April.

150. Graduating class of the first seral school- Farther back from the bank of the Delaware River in northeastern Kansas trees of pioneering eastern cottonwood and sycamore had reached size classes from large sapling yp through young adulthood on the outermost edge of the riparian zone. The author played amateur archeologist (crime scene investigator might be more apt) and determined that in the not-too-distant past highway construction crews had useed this higher part of the river's immediate floodplain that was close to a major highway as staging ground for major construction of bridge repair and road resurfacing. The devestated bottomland forest was undergoing secondary plant succession. Within a quarter century or less of construction-and-distruction action this young stand of two pioneering hardwood tree species had reforested the denuded floodplain.

Forest succession was on-going with young boxelder (Acer negundo) coming in as the replacement (of sycamore and eastern cottonwood) dominant species in the next seral stage on this bottomland forest range.

Even at this early seral there was diverse understorey of both shrub aand herbaceous layers. The dominant shrub was rough-leaf dogwood (Cornus drummondii) plants of which were larger than smaller seedlings of boxelder which remained decades away from wresting dominance away from eastern cottonwood and sycamore. Decreaser (climax) grasses had already returned with the dominant being a bottomland ecotype of switchgrass (visible as tall, bleached shoots with grainless panicles still otherwise intact). There was also at least one species of rosette panicgrass (Panicum of the Dichenthelium subgenus= Dichenthelium) still without vernal sexual shoots so as to make positive identifiction impossible. Naturalized smooth bromegrass was everywhere as to be expected in this close proximity to a state highway in northern Kansas. The most common forb was Indian hemp (Apocynum cannabinum) of the dogbane family (Apocynaceae).

Nemaha County, Kansas. Mid-June (late spring). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-89 (Northern Floodplain Forest). SAF 94 (Sycamore-Sweetgum-American Elm). Not a good unit in Brown et al. !1998, p. 43), but Warm Temperte Swamp and Riparian Forests (223), Southwestern Riparian Deciduous Forest and Woodland (223.2), Mixed Broadleaf Series (223.22) came closest. Western Corn Belt Plains, Loess and Glacial Drift Plains Ecoregion 47i (Chapman et al., 2001).

151. Deleware River's sylvanian outpost- At the outermost margin of bottomland forest that had developed along the heartland stream a more mesic forest community consisted of three dominant tree species: green ash (crown with leaves; tall, senescing tree at left in first slide and tallest tree to immediate left of big-girthed tree in second slide), bur oak (the largest tree; right-leaning with conspicuous rotting lower limb in both slides), and hackberry (multi-limbed tree to left of big bur oak in first slide). At this greater distance from the river and on slightly higher ground (ie. beyond the riparian zone) the forest cover type changed from a variant or form of the sycamore-cottonwood belt (SAF cover type 94) to a variant of the sugar[hackberry]berry-elm-green ash cover type (SAF 93). Bur oak as a constituent in this forest vegetation was a spatial relict or an ecological "sentry" from the adjoining or "close-by" bur oak forest cover type (SAF 42). From that perspective this forest plant community was an ecotone between SAF 93 and SAF 42. Most regeneration was of green ash and hackberry with only a few seedlings of bur oak. This was consistent with bur oak as a sort of outlier or "drifted" relict from the neighboring bur oak forest.

Bois d'arc or hedge apple was invading the extreme border of this forest (eg. saplings conspicuous beneath the ancient bur oak) where the forest community joined a degraded tallgrass prairie. Invasion by bois d'arc was the direct result of European man, the race that introduced this species from Oklahoma, Missouri, Texas, and Arkansas for attempted use as a living fence against free-ranging cattle (hence the common name of "hedge" throughout Kansas). While Indians did make their bows from this wood (hence the Anglized form from French, bois d'arc) the tribes in what became Kansas got this wood through trade with southern tribes. The white man brought bois d'arc to Kansas where it became a major pest (as well as the source of the most durable wood for fence post in North America).

There was a rich understorey as the tree layer was composed of more widely spaced and fewer species than in the densely shaded riparian zone forest of the same river as detailed above. The most abundant herbaceous species was poison or spotted hemlock followed by the naturalized, Eurasian perennial grass, smooth brome both of which were obvious in both of these slides. Other understorey species (woody and herbaceous species) were presented in the next slide.

Brown County, Kansas. Mid-June (late spring). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-89 (Northern Floodplain Forest). SAF 93 (Hackberry-American Elm-Green Ash). Not a good unit in Brown et al. !1998, p. 43), but Warm Temperte Swamp and Riparian Forests (223), Southwestern Riparian Deciduous Forest and Woodland (223.2), Mixed Broadleaf Series (223.22) came closest. Western Corn Belt Plains- Loess and Glacial Drift Plains Ecoregion 47i (Chapman et al., 2001).

152. A treasure chest or Pandora's box (depending on one's point of view)- The botanically diverse understorey of a green ash-hackberry-bur oak bottomland forest that developed on the floodplain of the Deleware River in the Dissected Till Plains (glaciated portion) of northeastern Kansas. Major herbaceous species in this "photoquadrant" included yellow giant hyssop (Agastache nepetoides), lopseed (Phryma leptostachya), stinging nettle (Urtica dioica), some species of smartweed or knotweed (Polygonum sp.), and pokeberry. Most abundant woody species included bois d'arc and riverbank grape, but green ash and hackberry dominated with exclusive cover by these two dominant trees to far rear of camera range.

This zone of an array of species was farther in the forest than the outermost zone which was presented in the two preceding photographs and that was dominated by poison hemlock and smooth bromegrass.

Brown County, Kansas. Mid-June (late spring). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-89 (Northern Floodplain Forest). SAF 93 (Hackberry-American Elm-Green Ash). Not a good unit in Brown et al. !1998, p. 43), but Warm Temperte Swamp and Riparian Forests (223), Southwestern Riparian Deciduous Forest and Woodland (223.2), Mixed Broadleaf Series (223.22) came closest. Western Corn Belt Plains, Loess and Glacial Drift Plains Ecoregion 47i (Chapman et al., 2001).

153. Mill Creek changes- Esterior of a bottomland forest that developed along Mill Creek which is a smaller stream in the Smoky Hills of the eastern Great Plains or, more accurately, the margin of Great Plains and Central Lowlands physiographic privinces. Physiogonomy as well as general structure and species composition was preented in this overall view.

This floodplain forest was another case in which mature and senescing trees species of pioneer (colonizing) and/or earlier seral stages that were currently dominant were in the successional process of being replaced (through different rates of regeneration) by successionally more advanced--greater tolerance for shade and competition--species of trees. Eastern cottonwood and associated tree species like black walnut and red mulberry were either not regenerating (eastern cottonwood and black walnut) or reproducing at rates that appeared to be inadequate to maintain current proportions in the tree population (red mulberry) while hackberry and American elm were asecnding the throne to climax vegetation. There were some large seedlings or very small saplings ofboxelder and green ash, but these (especially green ash) were proportionately much less than those of hackberry and American elm. In terms of Society of American Foresters forest cover types (Syre, 1980) Cottonwood (SAF 63), "a temporary, pioneer type" (Syre, 1980, p. 62) was being replaced through plant succession by the Hackberry-American Elm-Green Ash (SAF 93) which "tends to be long term in the successional scale" (Syre, 1980, p. 65).

Other trees species present as adults in this bottomland forest (though not distinguishable in this photograph) included boxelder and honey locust. There was considerable regeneration of boxelder (though less than hackberry and american elm) through sexual reproduction, but the author found no reproduction of honey locust.

As this sucessional drama unfolded mature trees of lower seral stages and less tolerance persisted while climax and/or advanced seral-stage species invaded the sere so that there was increasing shade (ie. decreasing light reaching lower strata of the forest range) and a declining understorey, at least of herbaceous species. the most abundant of which was pokeberry (foreground of this photograph) that survived in small or local openings and along the forest edge. The second most common herb varied locally including wood nettle (Laportea canadensis), Virginia wildrye (Elymus virginiana), fowl mannagrass (Glyceria striata), to a Carex species which could not be identified at this time due to absence of inflorescence and fruit.

There was a shrub layer composed primarily of woody vines, mostly Virginia creeper and bristly greenbrier (Smilax hispida). Riverbank grape and Missouri gooseberry (Ribes missouriensis) were also present but quite localized in contrast to the two more common lianas.

Trees in this slide included the climax hackberry (left and center left), eastern cottonwood (three trunks at center-right) and red mulberry (dying at right and slightly foreward of other trees).

Along Mill Creek, Washington County, Kansas. Mid-June (late spring). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-89 (Northern Floodplain Forest). Ultimately, SAF 93 (Hackberry-American Elm-Green Ash); currently, SAF 63 (Cottonwood). Not a good unit in Brown et al. !1998, p. 43), but Warm Temperte Swamp and Riparian Forests (223), Southwestern Riparian Deciduous Forest and Woodland (223.2), Mixed Broadleaf Series (223.22) came closest. Central Great Plains- Smoky Hills Ecoregion 27a (Chapman et al., 2001).

154. Partiarchs present but being phased out- Vegetational dynamics in a bottomland hardwood forest in Kansas' Smoky Hills. The first of these "photographic dendrograms" presented mature eastern cottonwood (two largest trunks) and black walnut (immediately behind the cottonwoods) with no reproduction and hackberry (young trees at far left and far right) and American or white elm (many seedlings and saplings) in foreground as the young trees of the next successional stage which is climax. The second "photographic dendrogram" showed younger (and smaller) trees of American elm (left-most trunk which had a geniculate bend) and hackberry (center trunk with less prominent bends) replacing the pioneering (though persistent) eastern cottonwood (biggest trunk). In this second photograph the conspicuous leaves in the foreground that dominated the lowest layer of forest vegetation were those of Virginia creeper. Bristly greenbrier was the other major woody vine in this forest range.

In various forest cover types certain tree species which are pioneers--the first to colonize the "new land" created by disturbance--persist into the climax or, at least, subclimax through old age over which time span they grow to large size. Eastern cottonwood is one such species. Black walnut and red mulberry were two other such species in this forest although the successional state of red mulberry was not known.

155. Interior of a Smoky Hills, mixed hardwood, bottomland forest- Species composition and structure of a floodplain forest range going through plant succession from a forest of eastern cottonwood as dominant and black walnut as associate, a seral (maybe up to subclimax) stage, toward a climax hackberry-American elm forest. Cottonwood and black walnut persisted into the climax as mature to over-mature (senescing) trees, but tree regeneration (represented by numerous age classes) was hackberry, American elm, boxelder, and green ash in that relative order. Seedlings and saplings of these Tolerant tree species contributed at least two lower woody layers. Young trees were prominent in all of these photographs attesting to the success of regeneratioin of Tolerant trees.

Red mulberry was also present as mature and senescing trees without much regeneration. Successional status of red mulberry was not clearcut. Burns and Honkala (1990) noted that red mulberry is a component of forest cover types Cottonwood (SAF 63) and Hackberry-American Elm-Green Ash (SAF 93) and is regarded as Tolerant as to tolerance class "... as it often grows as an understory tree". A good-sized red mulberry with a prominent knothole was in left midground of the second of these two photographs. Senescing ("past-their-prime") cottonwood were featured in both of these photographs. The first slide featured (left to right) a mature black walnut, two old eastern cottonwood, and a young American elm. The second slide featured the rightmost cottonwood shown in the first slide at closer camera range with a pile of debris at base of its trunk. The forest floor was littered with downed limbs and branches of "over-ripe" cottonwood.

Shrubs in these photographs included such lianas as riverbank grape, Virginia creeper, bristly greenbrier, and Missouri gooseberry. Herbaceous species present in the understorey included pokeweed or pokeberry, wood nettle, Virginia wildrye, fowl mannagrass, and a species of caric sedge.

An ecological mystery- In this forest, as well as several others presented in this section, there were not mature trees of hackberry and American elm, the dominant and climax tree species. Instead, in each of these bottomland forests within which hackberry or sugarberry and American elm, slippery elm, or cedar elm were obviously the regenerating climax species there were only young trees of these species. In these same forests there were large (in some cases immense) trees of species that were pioneers and, in come cases, up to subclimax status. It was not known why this age/size class distribution existed--and so consistently--in so many forests that were at (or approaching) climax. To be specific, it could not be determined why there were only young trees of climax species. There was an obvious explanation as to why there were only mature trees of lower successional states (eg. pioneer stage) in these climax (at least subclimax) forests: young trees (especially seedlings and saplings) of pioneer and other lower seral communities were small plants of Intolerant species that could not regenerate (survive to become established) in the shaded and root-competitive habitat created by their parents and species of more advanced plant succession. What was not known was why trees of Tolerant, and usually climax, species were still immature. Another way of stating this was, why were there not fully grown trees of climax species present in forests that had climax composition, structure, and physiogonomy?

The partial answer is that these forest communities were second-growth forests that had developed to the stage of having the botanical composition, vegetational layers, and outer appearance of climax forests, but they had not yet reached the fully developed state of old-growth climax forests. These second-growth forests were still young enough in successional time frame that they had not developed to the old-growth stage or form that has age/size classes of ancient and senescing trees of the climax tree species. The question remained, why were there not climax forests at old-growth state of development? The most logical answer to this lies in time of settlement, both settlement by European white man as well as US government-forced relocation of American Indians. Prior to Anglo-American influence (ie. before whites dispossed the Indians) the aboriginal tribes for the most part had not felled and cleared large areas of forest. Most impact of Indians on forests--bottomland or upland--was periodic fire; use of acorns, walnuts, pecans, etc. for food staples, and killing of browsing forest animals like deer for meat. Europeans by contrast cleared much larger areas of primival forest for farmland, timber sales, and fuelwood. Even forests that were not cleared outright where highly modified by action of Europeans that ranged from damming of rivers themselves or streams draining into them (ie. multipurpose dams), channalization and dredging (for flood control, navigation, "beautification"), and grazing by livestock (consumption of mast, especially by swine, was especially harmful for tree recruitment).

Such forest modification varied from minor disturbances to complete destruction with more time required for recovery of forests from the more drastic degrees of denudation. An example of forest destruction and forest recovery by secondary plant succession would be clearing of forest for cropland and subsequent natural reforestation of old fields (abandoned farmland). In most instances (apparently in all cases presented here) there was simply not enough time for complete recovery of second-growth forests to the ultimate old-growth form of climax forests. Bottomland forests in Kansas provided a convenient example. The earliest direct affect of white man on forests in what became the state of Kansas can be said to have begun in about 1830 when, under authority of the Indian Removal Act, Indian tribes from the Great Lakes Region were forceably moved to the land that became Kansas three decades later. Kansas Territory was created in 1854 at which time white settlement legally began. Statehood was granted to Kansas in 1860. The Homestead Act of 1862 probably contributed as much if not more than any other factor to European settlement of the Jayhawker State.

By the early Nineteenth Century forests on land that that was to become Kansas were subject to human impacts which were of different degrees and kinds than those these forests had been subject to for several thousand years prior to that period (not to mention millenia of pre-human time under which these plant and animal communities evolved). From the earliest period in which European influence could have impacted Kansas forests until the time of photographs presented here was approximately 180 years. Most forests were probably influenced--directly or indirectly--by Europeans for considerably less time than this. Less than two centuries was, more than likely, not an adequate length of time for drastically altered forests to redevelop to old-growth forests.

Another possibility as to why forests shown and described here had not developed into old-growth forests would be just the opposite of human destruction and subsequent forest recovery. That alternative possibility was that actions by European man contributed to increased rather than decreased area in forests; that instead of forest redevelopment and reformation following deforestation (ie. reforestation) this was actually afforestation, establishment of forests on land where the preceding vegetation was not forest (Helms, 1998). Such a conclusion was reached by Abrams (1986) from a study of gallery forests on the Konza Prairie, a long-term ecological research unit, in northeast Kansas. Comparsions of notes on forests in land survey records (original reports by surveyers of the General Land Office of the United States Department of Interior) to area in present-day forests indicated that there had been "dramatic expansion of gallery forests along the stream channels..." from 1859-1939 and again periodically up until 1978 (Abrams, 1986). "Increases in woody vegetation were attriburted to decreased fire intensity and frequency since European settlement" (Abrams, 1986). In these gallery forests establishment of hackberry and elms occurred from ten to thirty years after recruitment of bur oak and chinquapin oak. Abrams (1986) predicted that members of Ulmaceae would eventually replace those of Fagaceae as overstorey dominants with final outcome of elm dominance depending on role of Dutch Elm Disease.

If the afforestation alternative is closer to correct than that of natural reforestation, the absence of mature and senescing trees of more tolerant species (hackberry, elms, boxelder, green ash) would be explained by the same insufficient period of time for plant succession and maturation/senescence of trees of climax dominant species. Either way, the operative process is plant succession with the same ultimate forest community being a climax of hackberry, elm, and other more tolerant species with pioneer tree species (eg. eastern cottonwood, black walnut, honey locust) surviving as aged and dying members in the old-growth forest.

156. Young forest around an old forest on tallgrass prairie- Outer view of a forest of black locust and black walnut with Kentucky coffeetree as associate species around the margin of a climax forest of green ash (most abundant)-hackberry-elm (both American and slippery)-eastern cottonwood (persistent pioneer) along a stream in a Smoky Hills tallgrass prairie. The blooming shrub was roughleaf dogwood. Most common tallgrass species were bottomland ecotypes of switchgrass (represented by the tall, bleached-out straw), Indiangrass, and big bluestem. Also present was the naturalized smooth bromegrass.

The forest on the outer edge and the forest in interior of this tree-dominated tract were two different forest communities with the exterior community of black locust, black walnut, and Kentucky coffeetree being a seral stage of the climax forest of green ash, hackberry, and elm forest.

Washington County, Kansas. Mid-June (late spring). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-89 (Northern Floodplain Forest). Ultimately, SAF 93 (Hackberry-American Elm-Green Ash). Not a good unit in Brown et al. !1998, p. 43), but Warm Temperte Swamp and Riparian Forests (223), Southwestern Riparian Deciduous Forest and Woodland (223.2), Mixed Broadleaf Series (223.22) came closest. Central Great Plains- Smoky Hills Ecoregion 27a (Chapman et al., 2001).

157. Close and poisonous quarters- Stand of poison or spotted hemlock (Conium maculatum) growing along the Marais des Cygnes River. The river vegetation (both riparian and that away from channel bank and still on the floodplain) of this river provided an opportunity to study bottomland forest range in the Osage Questas province of the Central Lowlands. This forest vegetation was similar to that of other rivers in this region that were covered herein.

One unique opportunity afforded by bottomland of the Marais des Cygnes River was a large stand of poison hemlock. This introduced Eruopean forb has naturalized throughout much of North America. Poison or spotted hemlock is typically a biennial (less commonly a short-lived p;erennial) that contains at least five similar alkaloids that are structurally related to nicotine with similar physiological functions (Kingsbury, 1964, ps. 379-383). All species of livestock and humans have been poisoned by this member of the Unbelliferae, carrot or parsley family. This is the species that yielded the execution extraction of which Socrates was forced to drink. In the words of the American Association of University Professors, "Academic freedom is not free".

Numerous poisonous plant bulletins and sudry publications have discussed poison or spotted hemlock, Stephens (1980, ps. 80-82) being a good one for the Jayhawker State. The definitive authority for North America is Burrows and Tyrl (2001, ps. 54-57).

Osage County, Kansas. Mid-June.

158. Shoots poison- Several shoots of poison hemlock in the stand introduced immediately above. This species is easily confused with its relative--and not infrequently its neighbor--water hemlock (Cicuta maculata) upon initial sightings of either species by neophytes. They are readily distinguished by knowledgable rangemen (or for that matter by anyone who will take a second look). Leaves of poison or spotted hemlock appear fern-like being of a triangular-shape and three to four times pinnately compound (dissected) whereas water hemlock leaves have larger leaflets and are only one to three times pinnately compound (Stephens, 190, ps. 79-80). One does have to look somewhat closely to distinguish between these two. They have almost identical species ranges. Despite common names water hemlock is the more deadly poisonous of these two species.

This colony of poison hemlock was particularily dense with large plants growing, as they were, on the floodplain of the Marais des Cygnes River in the Osage Questas portion of the Tallgrass Prairie Region.. Osage County, Kansas. Mid-June, full-bloom stage.

159. Umbels in an umbel- Secondary units (umbels) of the overall umbel inflorescence of poison hemlock (first two slides) followed by a closer-yet view of one unit (sub-umbel) of a secondary umbel unit on the same poison hemlock plant. Umbel inflorescences of the Umbelliferae are compound arrangements of small flowers or "umbels in larger umbels in larger yet umbels".

Enjoy the photographic subject, study this naturalized range forb, and wash your hands after being around these lethally toxic poisonous plants.

Osage County, Kansas. Mid-June, full-bloom stage.

Bosque River- A Texas Example of a Subhumid Zone River

A tract of relict bottomland mixed hardwood forest on the floodplain of Bosque River in northcentral Texas provided a Godsent, nearly pristine example of one of the most widely distributed forest range types in southcentral North America. This was a climax sugarberry (Celtis laevigata)-cedar elm (Ulmus crassifolia)-pecan (Carya illinoinensis) forest with a Canada wildrye (Elymus canadensis)-broadleaf woodoats (Uniola latifolia) understorey.

This range plant community was presented in four "mini-segments" each showing and describing this forest range vegetation at a seasons of the year typical for this mild contintental climate in a subhumid precipitation zone. First, however, was a short--though somewhat dramatic--section showing what is meant by floodplain forest.

Students who desired a detailed description of the forest range vegetation presented below were referred to Rosiere et al. (2013).

160. "How high's the water?" (from a song by Johnny Cash)- Flash flood on Bosque River after cresting about 18 hours earlier and continuing to recede.

"A flood is the inundation by a body of water on land that is normall dry...Flooding is a normal part of the hydrologic cycle" (Newton, 2003, ps. 120-121).A flsash flood was defined by the American Geological Institute (Gary et al., 1972 Wilson and Moore, 1998) as "a local and sudden flood or torrent or relatively great volume and short duration, overflowing a stream channel in a usually dry valley (as in a semiarid areea), carrying an immense loas of of mud and rock fragments, and generally resulting from a rare and brief but heavy rainfall over a relatively small area having steep slopes. It may also be caused by ice jams, [heavy or rapid snowmelt], and by dam failure.". Flash flood is distinguished from other forms of flooding that may take longer to crest and, then, to recede. "Flash" means rapid-- both beginning and ending.

Note from the above definitions and from this author's introduction of floodplain forests that flooding is a natural phenomenon. These two "snapshots in the life of a river" illustrated the life-giving, absolutely essential provess of flooding to development and survival of a bottomland or floodplain forest.

The first of these two slides was of the main channel of Bosque River. The second slide was at confluence of the main channel (left) with Dry Branch (right and center), a tributary of Bosque River.

Viewers should see the beauty of the muddy water. No, that "dirty" water "ain't" for drinking; its for "eating"--of soil minerals carried in the "liquid life" by forest plants. Alluvium is as essential as the water itself for life of the floodplain forest range. Alluvium is " a general term for clay, silt, sand, gravel, or similar unconsolidated detrital material, deposited during comparatively recent geologic time by a stream or other body of running water as sorted ro semisorted sediment in the bed of the streanm or on its flood plain or delta...especially such a deposit of fine-grained texture (silt or slity clay) deposited during time of flood" (Wilson and Moore, 1998). Also: "Sediments deposited by running waterof streams and rivers. It may occur on terraces well above present streams, on the presernt flood plains or deltas, or as a fan at thebasse of a slope" (Soil Science Society of America, 2001).

Similarly students should take note of all that conspicuous debris (eg. rotten logs, uprooted trees), carried by flood water is essential to the forest as which is future plant nutrients. Such debris may function as local dams that create essential habitat for plants and animals of the forest community. In its hydrological meaning the American Geological Institute (Wilson and Moore, 1998) defined derbis from a geological view as:"Any surfical accumulation of loose material detached from rock masses by chemical and mechanical means, as by decay and disintegrtion. It consist of rock fragments, soil material, and sometimes organic matter". In a more ecological context, debris was defined by Wilson and Moore (1998) as: "Solid waste from natural and man-made sources deposited indiscriminately on land and water".

Bosque River, Erath County, Texas. Late January.

Spring-Bosque River

Spring (General)- The following few sets of photographs introduced and presented an overall perspective of the climax sugarberry-cedar elm-pecan forest range in spring.

Transition Winter to Spring (Late Winter/Early Spring Stage)-Bosque River

Climax associate (and a shade tree without superior)- An estraordinarily fine example of American elm growing on a tributary (Dry Branch) of a small river (Bosque River) at perimeter of a sugarberry-scedar elm-pecan bottomland forest in the West Cross timbers in northcentral Texas. This tree had probably not reached its maximum size for this habitat and its genetic potential, but it was a large specimen and it showed to admantage the general habit or morphology characteristic of a species capable of producing extremely trees by angiosperm standards. This tree had quite a fruit (samara) yield, but most of these dry fruits had been killed by an extremely low-temperature (10 degrees Fahrenheit) a week prior to taking of these slides. A good number (even though the percentage was low) of the samaras on this old parent tree had survived. Natural selection insured tht a fairly hard freeze (not to mention a light frost) would not kill all of progeny of this native tree species.

American elm was an associate species of this mixed hardwood floodplain forest whereas cedar elm was one of three tri-dominant tree species (second behind sugarberry and ahead of pecan). Cedar elm drops its fruit in autumn so distinguishing between these two Ulmus species was a snap in this forest. It is the plant architecture (gross form or morphology) of American elm coupled with its vast, sprawling, hold-tight root system (that makes this species almost impossible for wind to uproot) that endeared this native tree as the ultimate shade tree in much of North America. It is also a rapid-growing tree and one that is generally tolerant to shade and competition.

The conspicuous herbaceous understorey of this forest range consisted almost exclusively of Canada wildrye, broadleaf woodoats, and hairy woodland brome (in that relative order). The dominant shrub in this climax vegetation was fiddleleaf green-brier. In the two views seen here there were a number of plants of elderberry and trumpet creeper.

There were numerous entries to burrows of bank-dwelling beaver in the soil of the foresst range seen here which made for careful human travel over this land.

Floodplain, Bosque River, Erath County, Texas. Early March; hibernal aspect, flowering stage of phenology. FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash) and/or SAF 94 (Sycamore-Sweetgum-American Elm). Absence of sycamore and sweetgum and presence of sugarberry suggested that SAF 93 was more appropraite; however, the former cover type designation for SAF 94 included pecan so choice was unclear. Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

Architeccture of a climax associate- Three progressively closer camera distance views of a large American elm that was growing on the upper bank of a tributary (Dry Branch) of a small river (Bosque River) in the West Cross Timbers in northcentral Texas. Many adult trees of American elm are characterized by widely spreading crowns that develop from massive limbs some of which may grow to diameters rivaling that of the the trees' trunk. It is this growth habit that makes American elm such a wonderful shade tree. American elms have wide-reaching root systems with long lateral roots to match their spreading crowns. This is why wind storms do not uproot American elms in the fashion of tree species that have a smaller peg-root system. A larger, stronger tornado will break off American elms, but cyclonic winds wil not usually uproot the well-rooted American elm. Branches may be torn from American elms, but they are promptly replaced by these raipdly growing beautiful trees.

Many mighty American elm were killed with introduction of the alien Dutch elm disease, which is caused by sac fungus of any of three ascomycetes (ascomycete microfungi) species especially Ophiostoma novo-ulmi that is spread especially effectively by the (also introduced) pest, European elm bark beetle (Scolytus multistriatus), the American elm bark beetle (Hylurgopinus rufipes) and banded elm bark beetle (Scolytus schveyrewi), formerly placed in its own family,Scolytidae but now interpreted as being in family Curculionidae, subfamily Scolytinae). Extirpation of the magnificant American elm from eastern North America by this combination of fungus and vector ranked with loss of American chestnut (Castanea dentata). Such loss amounted to an ecological disaster and inexcusable destruction. For whatever reason, numerous adult (and comparatively large) trees of American elm in northcentral Texas have not been killed by Dutch elm disease. That includes a number of American elm along the Bosque River such as the fine specimen shared here.

Other tree species in the far background of these photographs ranged from sugarberry, cedr elm, pecan, black walnut, and bois d'arc. The main shrub in this floodplain mixed hardwood forest range was fiddleleaf or saw green-brier, which was the dominant shrub overall and grew throughout this climax vegetation. Other shrubs within views of these slides were trumpet creeper, poison ivy, and in the patch of plants at right, rusty blackhaw haw (Viburnum rufidulum). The herbaceous layer was dominated by Canada or nodding wildrye, broadleaf woodoats, and hairy woodland brome (in that relative order).

Floodplain, Bosque River, Erath County, Texas. Early March; hibernal aspec, flowering phenology in American elm. FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash) and/or SAF 94 (Sycamore-Sweetgum-American Elm). Absence of sycamore and sweetgum and presence of sugarberry suggested that SAF 93 was more appropraite; however, the former cover type designation for SAF 94 included pecan so choice was unclear. Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

Spring (late winter) view of on-going forest succession- Physiogonomy, structure, and composition of a mixed hardwood bottomland forest along the outer bank of a small of a river in the West Cross timbers of Texas. Crowns of two adult trees of eastern cottonwood, pioneers members of this forest, stand above "later arrivals" of higher successional status. Younger member tree species seen here were immature trees (large sapling or small pole size) of American elm and sugarberry along with, though at smaller crown cover, cedar elm, pecan and bois d'arc or Osage orange. There was also some cover of common elderberry, a major shrub in this floodplain forest. Dead stalks in the immediate foreground were of giant ragweed and Johnsongrass. There was young growth of Canada wildrye and broadleaf woodoats.

Bank of Bosque River, Erath County, Texas. Mid-March; late winter (late bloom/early fruit phenology). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash) and/or SAF 94 (Sycamore-Sweetgum-American Elm). Absence of sycamore and sweetgum and presence of sugarberry suggested that SAF 93 was more appropraite; however, the former cover type designation for SAF 94 included pecan so choice was unclear. Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

Closer spring (late winter) view of forest succession- On the outer bank of a river in the West CrossTimbers of Texas young trees of American elm, sugarberry, cedar elm, pecan, and bois d'arc or Osage orange (in that relative order) had become established under the open canopy dominated by pioneer eastern cottonwoods that were now at full maturity (though still in their "prime of life"). There were several plants of mustang grape that grew from ground level to tops of younger, smaller treees. Green foliage in the understorey was herbage of Canada or nodding wildrye, broadleaf woodoats, Japanese chess and cheatgrass. The first two of these grasses were climax (decreaser species) herbaceous co-dominants. There were some dead shoots of giant ragweed and Johnsongrass in the midground along an opening on the inner river bank.

166. On a bank of the Bosque- View of a climax bottomland forest on a north bank (so south slope) of the Bosque River. All the large crowns were of pecan. The small tree in right foreground was American elm. On portions of the river bank were periodic ,recurrent running water scoured the bank giant ragweed dominated the vegetation (almost to exclusion of any other plant species). Cool-season Canada wildrye and warm-season broadleaf woodoats were the dominant herbaceous species of the forest understorey in their respective seasons. Canada wildrye dominant in mid spring.

Erath County, Texas. Early May (vernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash) and/or SAF 94 (Sycamore-Sweetgum-American Elm). Absence of sycamore and sweetgum and presence of sugarberry suggested that SAF 93 was more appropraite; however, the former cover type designation for SAF 94 included pecan so choice was unclear.Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

167. Vernal verdure: springtime in the Bosque bottoms)- Four views of an elm (cedar and American)-pecan-sugarberry climax lowland forest in northcentral Texas in spring. This forest had developed was in the bottomland of the Bosque River, a typically perennial stream. The dominant species in most stands was cedar elm with pecan being the associate to lesser co-dominant species. Sugarberry or southern hackberry (Celtis laevigata) and American elm were other major species. Some younger individuals of bur oak (Quercus macrocarpa) were present suggesting that this was also a climax species that was returning on this previously human-disturbed forest. Eastern cottonwood was present as a few widely scattered trees. At upper elevational zone of the forest there were a few old-growth live oak (Quercus virginiana= Q. virginiana var. fusiformis= Q. fusiformis) of possibly hybrid of these taxa). Shrubs included mustang grape, smooth sumac (Rhus glabra), common greenbriar, Virniginia creeper (Parthenocissus quinquefolia), and trumpet creeper (Campsis radicans).

The dominant herbaceous speceis in this vernal aspect was Canada wildrye. Broadleaf woodoats was the warm-season dominant. Jonsongrass (Sorghum halepense), a naturalized native of Africa, was dominant in spots during summer, but it was an overall dominant at any time. The same situation applied to the naturalized annual bromes, cheatgrass (Bromus tectorum), rescuegrass (B. unioloides= B. catharticus= B. wildenowii), and Japanese chess (B. japonicus). There were a few rare-as-hen's-teeth shoots of the native giant cane (Arundinaria gigantea) suggesting that this extremely palatable bamboo had been nearly grazed out in the distant past. The dominant cool-season forb was the naturalized Eurasian umbel, hedge parsley (Torilis nodosa). Dominant warm-season forb was frostweed or white crownbeard (Verbesina virginica). Elderberry was present as large, isolated plants (Sambucus canadensis).

All of the larger trees in the first of these four photographs were cedar elm while the herbaceous understorey was overwhelmingly composed of Canada wildrye. The second photograph featured a bur oak sapling (center midground) surrounded by a dense and verdent stand of Canada wildrye with a young pecan of pole-size and a pecan sapling were at left.

The third photograph presented a classic composition view of this forest range vegetation: cedar elm (three large trees in center; one on left has dual or split trunk), pecans (far left), sugarberry (background to right of two-trunk cedar elm), and American elm (small tree to left and behind the rightmost cedar elm and sapling in right-corner foreground). In this second slide broadleaf woodoats was a distant second major grass to Canada wildrye. Elderberry and giant ragweed were conspicuous in midground.

The fourth photograph had quite a diversity of species including smooth sumac and common greenbriar as the major shrubs along with frostweed, the dominant forb at peak standing crop in autumn, and hedge parsely, a common forb in the vernal aspect. Grasses in the fourth slide included Canada wildrye, the spring dominant herbaceous species, broadleaf woodoats, generally dominant warm-season grass (mature panicles avisible in foreground), Johnsongrass, a warm-season dominant in small spots (it was conspicuous in foreground), and three Eurasian annual bromes, rescuegrass, cheatgrass, and Japanese chess. Trees included cedar elm, sugarberry, pecan, and bur oak (these four were present in background) and American elm (pole-size tree in right foreground). Larger tree behind and to left of this American elm was pecan. Greenbriar climbing on the American elm with smooth sumac to right and slightly in front of this elm. A mustang grape was in the large pecan behind the american elm in right foreground.

Spring (Development)- The following sets of photographs featured vernal development of range vegetation on the sugarberry-cedar elm-pecan climax forest from onset of spring through "green-up" to conclusion of spring season. In most of the plant species comprising this forest range community a major portion of vegetative growth is completed prior to beginning of summer (estival solstice). Summer in this southerly latitude and the subhumid precipitation zone is usually a time of water stress in all except the most moisture-rich years which, of course, are much more rare than periodic dry spells and outright drought. Rainfall during July, August, and September can be adequate to abundant, but the high temperatures and frequent low humidity "suck" moisture from the sandy soil of this bottomland environment resulting in a growth "holding pattern " (ie. a period of reduced or "suspended", as in halted, plant growth). Thus summer for most species, especially herbaceous ones, is probably best descriped as a phenomenon of quasi-summer domrancy. A combination of shortening photoperiod, cooling temperatures, and generally higher soil moisture by early autumn hastens or permits flowering and fruit production in range plant species and completion of their annual cycle.

168. Afternoon sun on new shoots- The warm, bright glow of afternoon light fostered early spring growth of leaves and culms of broadleaf woodoats and Canada wildrye on the floor of a climax sugarberry-cedar elm-pecan bottomland forest that developed on the floodplain of the Bosque River. (The oblique light also highlighted the developing herbaceous layer of this three-layered forest range vegetation. Woodoats was the predominant grass in these two photographic views though Canada wildrye, the overall, slightly-rated-the-edge "number one" herbaceous species, had a respectable showing here.

The two tree trunks were young sugarberry, the dominant tree species of this range plant community.

These two photographs were from the same camera angle only at closer distance in the second slide (ie. same trees and the same clump of broadleaf woodoats in front of the foremost sugarberry trunk). The dead grass straw in center background of both photographs was that of last year's woodoats.

Erath County, Texas. Late March (early spring); second leaf-beginning culm elongation stage of phenology. FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash) and/or SAF 94 (Sycamore-Sweetgum-American Elm). Absence of sycamore and sweetgum and presence of sugarberry suggested that SAF 93 was more appropraite; however, the former cover type designation for SAF 94 included pecan so choice was unclear.Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

169. On locally disturbed ground- Stand of giant ragweed, an annual composite of great dimensions, with some annual ryegrass (Lolium multiflorum) on a flood wter-scoured area on the upper bank of Bosque River. Denuded local habitats (microhabitats) like this were commonly dominated by this native pioneering species along the river in a climax sugarberry-cedar elm-pecan bottomland forest. Perennial forbs were more common in undisturbed parts of this forest's understorey.

The boundary of the local disturbance (denuded area) created by flood water was very discrete or sharp as shown in the first of these photographs. Above (higher up on the river bank) the grass species were Canada wildrye and broadleaf woodoats, the dominant herbaceous plants of this forest range. The trees in the first slide were an American or white elm (Ulmus americana) in the foreground with sap oozing from a wound. Green briar was climbing this "bleeding" trunk" The dramatically leaning tree behind the elm was pecan. It was shown below that leaning pecans were a common feature of this forest with many such trees reaching immense proportions and surviving to old age.

The first of these two slides was taken from the top of the river bank down into the river channel to give a general view of the giant ragweed stand in relation to the rest of the bank that was not disturbed (denuded by flowing water). The second photograph of this set presented the stand of giant ragweed at closer camera range, but from an angle starting at the river bed and zooming up the river bank. In this second slide the American elm and pecan were in left background. The mostly dead tree center and left background in first and second photographsrespectively, was red mulberry (Morus rubra).

Erath County, Texas. Mid-May (vernal aspect at mid- to late spring). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash) and/or SAF 94 (Sycamore-Sweetgum-American Elm). Absence of sycamore and sweetgum and presence of sugarberry suggested that SAF 93 was more appropraite; however, the former cover type designation for SAF 94 included pecan so choice was unclear.Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

170. On settled ground- Mid-morning sunlight filtered through the lightest of cloud cover permitted full depth-of-field views in these two and several succeeding photographs of the range plant community of a bottomland climax sugrberry-cedar elm-pecan forest with a remarkably well-developed herbaceous layer. Several species of shrubs, including lianas, formed a sporadic or erratic lower woody layer between that of trees and herbaceous plants. Almost all of the grass herbage, which was the vast proportion of herbaceous biomass, was of the two native, cool-season, perennial bunchgrasses, Canada wildrye and broadleaf woodoats. There was minor cover and biomass of nimblewill (Muhlenbergia schreberi), a native, warm-season perennial. The two, naturalized, Eurasian, annual grasses, cheatgrass and Japanese chess, were present as widely scattered plants at trace amounts. The most common (and conspicuous) forb was frostweed or white crownbeard followed--"at a distance" by pigeonberry (Rivina humilis). Both of these are native warm-season species. Another common and well-distributed forb was hedge parsley, a naturalized, Eurasian, annual umbel.

Foremost tree in both slides was sugarberry, the most important of three dominant tree species. Large trees in mid- to background were pecan (the third-ranking dominant tree) which had formed a grove of old-growth specimens, some of which were massive and, undoubtedly,.quite old. Sugarberry was in the process of replacing pecan through proportionately greater rates of reproduction.

171. Placid views of a "settled community"- Range vegetation of a climax sugarberry-cedar elm-pecan bottomland forest on the upper (an older) floodplain of Bosque River. Most of the lush herbaceous standing crop was grass herbage comprised of Canada wildrye and broadleaf woodoats which were native, cool-season species having cespitose habit. Nimblewill, a native, warm-season perennial, was also present. There were a few plants of cheatgrass and Japanese brome, annual Eurasian grasses. Frostweed was the most abundant forb with pigeonberry coming in as a distant second. Both of these are native warm-season species. Hedge parsley, a naturalized, Eurasian, annual umbel, was another commonly seen forb.

These two photographs showed the inner edge of a stand of young sugarberry which had grown at the exterior of a grove of old-growth pecan. This grove was featured in the last two-slide set immediately above and the next two-slide set immediately below.

The first of these two slides was shot under a full-sun sky which showed the phenomenon of sunflecks which are periodic areas or flickering spots of light where "holes" in the crown canopy permitted penetration of light rays to the forest floor or herbaceous stratum. The second slide of these slides--like the two preceding slides and the second in the next two-slide set-- were taken under very thin cloud cover which diffused the light uniformly thereby eliminating sunflecks (and shadows) to show consistent color as well as plants consistently (ie. no plants were partially hidden by shade).

Sunflecks are an extremely important factor in conduction of photosynthesis (hence, survival) of plants in the understorey of forests. One of the more widely cited sources on sunflecks is that of Chazdon and Pearcy (1991).

172. Vernal vestiture- The ultimate green of spring in the vegetation of a climax sugarberry-cedar elm-pecan bottomland forest range that developed on the upper (an older) floodplain of Bosque River. Most of the herbaceous layer was made up of Canada wildrye and braodleaf woodoats which are native, cool-season bunchgrasses. Forbs included frostweed, pigeonberry, and hedge parsley.

The first of these two photographs was taken under a full-sun sky with some plants, including pecan trunks, in direct light and other plants in shade. The second photograph (taken with a mere few minutes before the first photograph) was with complete, though thin, cloud cover so that under diffuse light there was minimum shade and therefore fewer plants in the darker light of shadows. The full-sun shot (first slide) demonstrated the phenomenon of sunflecks, short exposures to sunlight when branches and leaves in the forest canopy are parted to permited to flickering "patches" of light to reach lower layers of forest vegetation. Sunflecks have been shown to be essential for survival of plants in lower strata of forest vegetation (Chazdon and Pearcy, 1991). Ultimately these flickering or short bursts of light are key to production of shrub and herbaceous biomass and, thus, of browse and forage for range animals.

The second (vertical) photograph was a closer camera range of the leaning pecans in central midground of the first (horizontal) photograph. A fairly high proportion of pecan, even at old-growth stage, have grown into a drastically leaning position. This showed the past influence of events that caused the young tree at some stage(s) of development to bend and then remain in that distorted position. This phenomenon was succinctly described by the old adage, "As the twig is bent so grows the tree". As applied to teaching the words of Alexander Pope (1734, ps. 149-150) in Epistle to Cobham were repeated here: 'Tis education forms the common mind, just as the twig is bent, the tree's inclined". Anyway, on this forest range the ancient deformed trees were botanical history writ in the living plant.

Also noticable in the second photograph was a rotting log which was a huge limb that fell from the larger (foremost) pecan which had several shed limbs (and rotting stubs high up on the woefully bent trunk. The ole bugger could tell its share of storm stories, drought tales, and so forth. Again, the life of a tree writ in its injuries. Of such is life-- in tree or man.

173. Handsome grove- A stand of old-growth pecan on the upper bank of Bosque River with an herbaceous understorey dominated by Canada wildrye and broadleaf woodoats. Forest and, sometimes, more like woodland , groves of pecan such as this are almost always limited to bottomland, especially floodplain, sites. Hence the folk term designation of "pecan bottoms". For whatever reasons, pecan trees in groves developing on these bottomland forest sites typically grow and develop into leaning, bent, twisted, and other misshapen forms frequently with limbs almost as massive as the trunks from which they arise. Loss of these heavy limbs from wind storms, gravity, and aging (the natural pattern and process of shedding older plant parts) results in burls and decaying appendage stumps often culminating in hollows and cavities which add "charaacter" to the already picturesque patriarchs.

Large size, old age, and distorted form notwithstanding, in many instances old-growth pecans do not sexually reproduce at rates sufficient to maintain pecan at these greater proportions as forest development progresses to climax. The old-growth individuals persist into a sugarberry-cedar elm-dominated climax, but more frequently than not smaller, younger pecans are not part (at least not a dominant part) of climax forests. That sugarberry and elms (both American and cedar elm) eventually succeed pecan as the more important dominants was represented in these slides by smaller sugarberry and cedar elm trees in the background and, most conspicuously, by sugarberry leaves at left margin in the second slide.

Nonetheless, pecan does persist as the third dominant of climax bottomland forest as young trees as well as senescing, old-growth trees. Next slide, please...

Erath County, Texas. Late April (vernal aspect at early to mid-spring). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash) and/or SAF 94 (Sycamore-Sweetgum-American Elm). Absence of sycamore and sweetgum and presence of sugarberry suggested that SAF 93 was more appropraite; however, the former cover type designation for SAF 94 included pecan so choice was unclear.Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

174. Pecan grove just above the river- A local grove of pecan growing on the upper bank of Bosque River furnished viewers with a second--though less photogenic than the immediately preceding--example of this form of range plant community within the mixed hardwood bottomland forest climax. Species composition and general structure of this example was about the same as that of the preceding pecan grove with two notable exceptions. First and most importantly, the forest range of this grove had numerous pecan seedlings (in foregrounds of both slides) coming in to maintain pecan as a dominant tree species of the climax forest that will eventually become dominated primarily by sugarberry and cedar elm. Commonly, pecan persist into the climax forest range mostly, even exclusively, as ancient, senescing, old-growth trees the huge crowns of which occupy as much of the canopy as sugarberry and cedar elm trees which are more numerous but of younger age and smaller size (and disproportionately less crown cover).

In this pecan grove, however, there was more than enough regenertion of pecan at the seedling stage to insure maintenance of pecan even as younger trees into climax. That is, barring some event, agent, or condition that takes out the seedlings as, for instance, a surface fire. Even then though (and assuming that fire killed existing seedlings) more seedlings could replace those there were lost. In fact, a surface fire could in theory remove enough leaf litter, grass straw, and other forms of organic mater that gemnination and emergence of pecan would be enhanced and result in as many, if not more, seedlings than before the burn. In absence of experimentation this remained unknown though such an outcome ranged from possible to inevitable.

The second difference between this pecan grove and the one presented immediately above was presence of snags in this pecan stand. The second of these two slides was a nearer view a snag and fallen limb seen in the background of the first photograph. In this pecan grove, pecan spanned the range of age/size classes from small seedling (again in foreground of both photographs) to snag. The extremes (= the end points) of the age/size spectrum were present though not with all classes. Saplings and pole sizes were not represented. This pecan grove existed as an even-aged stand with the present old-growth generation in the process of being replaced by the seedling genertion with intermediate ages and sizes absent.

Lower layers of this bottomland forest range included an herbaceous understorey dominated by Canada wildrye and broadleaf woodoats and a component of lianas--including mustang grape, common green-briar, poison ivy, coral honeysuckle, and, the alien, Japanese honeysuckle--that extended from soil to canopy.

Special emphasis was put on the horrid invasion of Japanese honeysuckle. This exotic shrub (introduced as a horticultural ornamental) is found in almost all bottomland forest of this or related cover types from northcentral Texas through the Pineywoods. Japanese honeysuckle unquestionably threatens forest integrity.

175. Natural pasture in bottomland forest- Floodplain of Bosque River (stream channel in background with both river banks visible) featuring the herbaceous understorey dominated--nearly exclusively--by Canada or nodding wildrye and broadleaf woodoats. In this vernal society the herbaceous layer was roughly 60:40 percent Canada wildrye and broadleaf woodoats. Other (and only incidental) grasses included three species of naturalized annual bromes: Japanese chess, downy brome or cheatgrass, and rescuegrass (first two, Eurasian; latter, South American). Forbs were very sparse yet the native, perennial dominants were indicator species (indicative of fertile, low-lying range sites). Frostweed or white crownbeard was the overall dominant forb with pigeonberry and pokeberry or pokeweed "swapping off" as the infrequent dominant and, forestwide, the native associate species. In this vernal society the most abundant native annual forb was cleavers or bedstraw (Galium aparine) while the most widespread naturalized forb was the Eurasian annual, hedge parsley (Torilis nodosa). Giant ragweed was generally the dominant forb in estival and autumnal societies, especially on locally disturbed microsites, but frostweed, pigeonberry, and pokeberry were local dominants at these seasons.

Dominant climax trees were sugarberry, cedar elm, and pecan (in that order) with pecan persisting as a climax dominant primarily as old-growth (and obviously senescing) individuals. There was, however, regeneration of pecan at all age/size classes at rates apparently adequate for maintenance of this species to the state of potential natural vegetation. (This phenomenon was shown and explained above as well as below.). The two associate tree species on this portion of the floodplain were in the Moraceae: 1) red mulberry (Morus rubra) and 2) bois d'arc, Osage orange, or hedge (horse) apple (Maclura pomifera). These two species are native to this region where they are typically smaller, even understorey, trees though with occasional trees of relatively large size (see below). Several species of trees and shrubs that were found elsewhere on this floodplain (black walnut and rusty black-haw for respective example species) were absent from scenes seen here and in the immediately suceeding two-slide set. By contrast, lianas like mustang grape, common green-briar, and trumpet creeper were fairly common.

All of the trees in the background of the first photograph were sugarberry except for one bois d'arc. In the second slide the two trees in left background were red mulberry and sugarberry while the foremost tree in left midground was also sugarberry. The snag in right midground was a pecan as were all of the large trees in the background of this second photograph. The smaller tree that was fully leafed-out was American elm. The two rightmost trees (actually there are three, but only two are visible) in the second slide (bois d'arc and sugarberry) were featured in a slide-caption unit below (ie. stay tuned).

176. Closer look at the forest forage crop- Two "photoquadrants" that stressed the herbaceous understorey of a climax sugrberry-cedar elm-pecan bottomland forest that had developed on the floodplain of Bosque River. The beautiful green of Canada or nodding wildrye and broadleaf woodoats (roughly a 60:40 percent ratio) in early spring growth "dominated" the vernal society of this lowland forest range. Other herbaceous species were largely incidental other than at local scale where disturbances such as flood-scouring of stream sides and banks created favorable microhabitats for "weedy" (pioneer) species like giant ragweed, pokeberry, and annual bromes (Japanese chess, downy brome or cheatgrass, rescuegrass). Other grass species including Johnsongrass and annual ryegrass were even less common and were typically restricted to disturbed "mini-areas".

Most of the trees in these two slides were sugarberry. The leafed-out sapling was American elm. Another American elm that had "leafed" ahead of other trees was shown in the last slide of the above set. These two trees were typical specimens which showed that American elm is one of the first trees to leaf-out in this area.

The second slide depicted the woody species diversity of this forest range. The three foremost trees in the second photograph were sugarberry, the foremost one had a nice shoot of bedstraw growing and lying against its trunk. The largest tree was a red mulberry in center midground. One of the two trees to right of this red mulberry was a bois d'arc with its side-by-side companion a sugarberry. A mustang grape arose in front of and later "clumb up" the red mulberry. Common green-briar grew beside (slightly to left of) the mustang grape.

177. Dead and dying- Broken-top snag of sugarberry with a dying (senescing) sugarberry to its right-rear (left-center of slide) and old and much larger pecan in background. Downed limb was off the left-center (of slide) dying sugarberry. Herbaceous layer was co-dominated by Canada or nodding wildrye and broadleaf woodoats. There was considerable cover of Japanese honeysuckle (Lonicera japonica), a horribly invasive exotic, in bckground. This range plant community was on the upper bank of Bosue River.

The big lesson in forest succession and tree silvics was that on sugarberry-cedar elm-pecan bottomland forest sugarberry never attains the great size (mass, trunk diamteter, crown spread, limb size, or tree height) of pecan even though sugarberry is the "number one" dominant of this forest cover type. Sugarberry is a species of smaller trees. Adults of sugarberry do not grow to the larger size of pecan or, even, cedar elm. It is as simple as that. Whether this is due to shorter life span (longevity), genetic basis to shorter stature and generally smaller adult size, or some other factor(s) was not known. The explanation is somewhere in the silvics of sugarberry. Certainly sugarberry appears later in succession (development of the forest vegetation) whereas pecan is a pioneer (colonizing) species or, at very least, appears at an early seral stage and then persist as massive trees into the climax forest. From this perspective sugarberry is a shorter-lived species on the forest sere and the much greater size of pecan can be explained.

Erath County, Texas. Mid-April; vernal aspect and dead to late senescing stage of life cycle of Celtis laevigata.

178. Now a look at the crop of trees- On the outer bank and immediate (adjoining) floodplain of Bosque River a climax sugarberry-cedar elm-pecan bottomland forest had developed with an herbaceous layer dominated by Canada wildrye and broadleaf woodoats across which were scattered individual trees and local aggregations or asemblages of trees of such dispersion as to form an open canopy forest or, even, a woodland-like structure. In this portion of the mixed hardwood forest range, crowns of widely sacttered trees usually did not come into contact except where trees grew close together in a locally clumped pattern. Some of these local "clumps" were of the same species (most often same-species 'bunches" were of sugarberry) whereas other such localized aggregations were composed of several tree species.

One of these milti-species tree groups was featured in these two slides that served as "photodendrograms". This group of three trees of as many species included (left to right): bois d'arc or Osage orange, sugarberry, and pecan. Bois d'arc and red mulberry are both in Moraceae or mulberry family, native to this region, and widespread in this bottomland forest growing from stream banks to outer edges of the floodplain. Presence of sugarberry in this local assemblage was representative of its role as the number-one dominant of this cover type. A pecan sapling in this tree group was especially noteworthy because it demonstrated (and depicted in a successionally symbolic way) that while pecan persist as a dominant into the climax forest mostly as long-lived, senescing, old-growth trees there are enough young trees so that all age/size classes of pecan grow in the climax vegetation. (Seedlings and smaller saplings of pecan were shown in other photographs in this section.) Two examples of old-growth pecan were the two trees in far-distant right background of the first photograph.

The presence of two native Moraceae species (bois d'arc and red mulberry) suggested a biotic affinity to river bottom forests far to the north. It was reported elsewhere in this publication (including above in the current chapter) that these same two species of the mulberry family were common in floodplain forest in central Kansas and eastern Nebraska. It has been generally accepted that bois d'arc was native only in what are now the states of Texas, Arkansas, and Oklahoma with this species having naturalized as far north as present-day South Dakota due to introduction by the whiteman for use as hedgerows (living fences which did not work well) and shelterbelts (McGregor et al., 1977). Even though Osage orange, hedge, or bois d'arc was not native to river bottom forests as far north as Kansas and Nebraska, naturalization and extensive presence of this species there now still shows a similarity of these riverine habitats (ie. an abiotic affinity if not also a biotic one). It also showed that human impact has increased similarity among bottomland hardwood forests. "Is man part of nature or not" (Tansley, 1935).

Besides nodding or Canada wildrye and broadleaf woodoats, the herbaceous co-dominants of this forest range, other (and mostly incidental) herbs in the range vegetation shown here included bedstraw or cleavers, hedge parsley, downy brome or cheatgrass, Japanese chess, and rescuegrass. Frostweed or white crownbeard was the major or dominant (most abundant; greatest foliar cover) forb, but it was nowhere near a dominant species of this forest range community.

179. Gnawed on- Beaver (Castor canadensis) had been at one time in the recent past been important range mammals in a climax sugarberry-cedar elm-pecan bottomland forest on the floodplain of Bosque River in the West Cross Timbers of northcentral Texas. These were bank beaver (those that live in borrows dug into stream banks) rather than lodge beaver (those that damn a stream and create a pond in which they errect a lodge of logs felled by them). At the time of these photographs and description of this forest range apparently there were no beaver living in this area of the forest. Specifically, scientists studying this forest could not find any evidence of current beaver residence such as recent feeding or tracks.

Not too far back (roughly five to ten years previously), however, beaver had fed on bark of sugarberry, cedar elm, and red mulberry. The telltale, gnaw scars of bark-feeding by beaver at base of tree trunks of these species was indisputable. Examples of standout, healing wounds from feeding beaver were included in this discussion of the sugarberry-elm-pecan cover type because defoliation by beaver can be a major determinant of species composition, structure, successional development, and even function of forest range.

The first of these four photographs presented a sugarberry (right) and a large red mulberry (left) browsed by beaver on the outer bank of Bosque River. Green growth of the current season (early spring) and dead stalks of last year's growth of Canada wildrye and broadleaf woodoats, herbaceous dominants of this forest range, comprised almost all of the understorey. Other herbs were visible and described in the remaining three photographs.

The second and third slides were detailed views of the scar left by beaver-browsing on the red mulberry shown at the right of the first photograph. Beaver had not only eaten the bark of this tree but also had gnawed into the cambium layer (maybe even deeper) as well. Chips spat out by beaver are larger than wood shreads from a properly sharpened chainsaw, but such shreads rot relatively quickly. Absence of shreaded wood, chips, amall bits of browsed bark, etc. indicated that these browse scars had been made several years ago (time enough for complete rotting of beaver leavings). In addition to co-dominant nodding or Canada wildrye and broadleaf woodoats,other visible herbaceous species included bedstraw or cleavers, giant ragweed, and hedge parsley.

The fourth slide was a close-in view of the sugarberry shown at right in the first slide. Bark of this tree was more noticably covering the beaver feeding scar than was the case for the red mulberry. Trumpet creeper was the liana shoot on the right side of the sugarberry trunk. Herbs at base of this trunk included Canada wildrye, broadleaf woodoats, hedge parsley, bedstraw, and henbit (Lamium amplexicaule).

The author was able to find only one stump and one part of the trunk of a sapling as evidence of beaver feeding on trees smaller than adult-sized trees as shown here. Simply put, there was no evidence of any beaver presence at time of these photographs. It was not known what factor(s) was responsible for absence of beaver in years following a time period of considerable feeding on large trees by beaver.

Also unknown was what, if any, impacts beaver browsing might have had composition, structure, development, and function of the climax sugarberry-cedar elm-pecan floodplain forest. Donkor (2007) studied beaver browsing in boreal forests and concluded that these largest of North American rodents influenced forest species composition through their selective browsing. All agents of defoliation, including manmade implements, defoliate selectively (= exhibit selectivity). On this bottomland hardwood forest, beaver browsed on certain tree species and not on others. Specifically, there were no feeding scars on any bois d'arc trees in this forest yet a number of red mulberry bore such gnaw scars, including one large red mulberry that had beeen killed from complete girdling by beaver. Both of these species are members of the mulberry family. Likewise, there was a much lower percentage of browsing on cedar elm than on sugarberry. Finally, the author could not find any evidence of beaver browsing on pecan. The dominant tree species that had the greatest number of individual trees had received the most beaver browsing (on a propotional or relative basis) and the "least" of the three dominant tree species that had the smallest number of trees (trunks) had zero browsing by beaver. Again, it was not known whether such selective browsing had any impact on this forest and, in fact, there was only limiited evidence (one dead adult tree and one dead sapling) that browsing by beaver had any lasting influence on individual trees.

That Celtis species are a major feed source for beaver (a long-know fact) was indicated by the common name of "beaverwood" for C. occidentalis (Small, 1933, p. 442).

Erath County, Texas. Late April.

180. Return of beaver- After an absence of several years from the climax sugarberry-cedar elm-pecan bottomland forest along this stretch of Bosque River (see immediately above) beaver "returned with a vengende" and commenced feeding on bark of sugarberry and, especially, cedar elm. Two comparatively large cedar elm, both of which had been girdled by beaver, were presented in this slide-caption set. These trees will likely be carcasses with any survival of these two trees dependent on coppice shoots (resprouts or new shoots) arising from the lower trunk base of roots.

Meanwhile taps for the trees. it was shown in various sections of Range Types of North America (see chapter, Miscellaneous Forests- II also) tha North American beaver (Castor canadensis) can have major, even profound, impacts of forests. Note robust plants of inland or broadleaf woodoats at base of girdled elms.

Bank, Bosque River, Erath County, Texas. Late November; late autumn.

181. Art work spelled death to trees- Details of bark-feeding by beaver on trunks of cedar elm. Tooth marks (by incisors) left by bark-browsing beavers were the "calling card" of death to these two tree trunks. Beaver had cut through the cambium or (deeper) thereby stopping sap flow through cambium or xylem hence stopping life of these trunks. Will trees resprout?

Stay tuned.

Bank, Bosque River, Erath County, Texas. Late November; late autumn.

182. Dam construction by the original hydrologic engineers- Low dam built by beaver across the main channel of Bosque River at it's confluence with a local ephemeral tributary. (The tributary was at left in both of these photographs.) During a dry autumn and winter (exceptional drought on Palmer scale) beaver constructed this dam of loose sticks (twigs up to large branches and saplings) that fell from pecan, American and cedar elms, sugarberry, eastern cottonwood, and red mulberry plus some woody shoots felled by beaver for feed. Rocks were also placed by beaver in the dam (second slide). These were selected from among those already present in the stream channel. Leaves and other debris, including algae from a "bloom" fed by runoff of commercial fertilizer applied along the banks by grounds workers of Stephenville Texas) quickly lodged in spaces among the beaver-laid material. This resulted in a relatively "water-tight" dam.

The spillway of the beaver dam was at the low bank side (left terminus as seen in the first of these two slides).

Herbage on banks of the river and tributary was primarily biomass of broadleaf woodoats and Canada or nodding wildrye. Large trees in background were old-growth pecan.

Erath County, Texas. Mid-March.

183. Details of dam construction- Materials and their placement in a beaver dam built across main channel of Bosque River at it's point of convergence with a major ephemeral tributary. Very little of the woody material was felled by beaver and instead had fallen (was shed) from standing trees or washed into the stream. A small portion of woody material in the dam had been felled and stripped of bark as browse by beaver. Wood (and some rocks) were gathered and maneuvered into place "busy beavers". The bale of straw on the upper (higher) bank (right side of dam in second slide) served as the "corner stone" of the dam. Street construction contractors, Jay Mills Contracting, (webmaster@ci.stephenville.tx.us), improperly anchored this bale on an upstream bank to prevent soil erosion. A rain storm of moderate intensity swept the bale (along with tons of soil) into Bosque River.

Beaver began builting their dam off of this bale and extended the structure out from the bale. (Note that sticks were aligned across--in direction of width of--the channel immediately above the bale versus aligned with length--up and down--the channel in rest of dam.) Obviously beaver are opportunistic builders.

Erath County, Texas. Mid-March.

184. A nice find- Local stand of hairy wood(land) brome of Canada bromegrass (Bromus pubescens= B. purgans) on floor of a climax bottomland sugarberry-cedar elm-pecan forest. Hairy wood brome is extremely uncomon in the Western Cross Timbers where this lovely stand was found on restricted area above Dry Branch that drained into Bosque River. In fact, it was only in relatively recent times that rangemen were aware of existence of this species in the Cross Timbers and Prairies vegetational or land resource area of Texas. Bromus pubescens was not shown as being here in either Gould (1962) or Hignight (1988). This author was estatic when he happened on such a marvelous stand. So much so he had to share it with other grass lovers.

It was emphasized throughout this treatment of the sugarberry-cedar elm-pecan type that Canada wildrye and broadleaf woodoats were co-dominant herbaceous species of this range plant community. For whatever reason this local stand of woodland bromegrass was a nearly exclusive population (consociation) in which common green-briar was about the only other species present. At the margins of this hairy wood brome stand there was a mingling of this native perennial with plants of Canada wildrye and broadleaf woodoats, but woodland brome was in essence segregated from other herbaceous species.

The tree trunk at far left of this slide was a large cedar elm. Sugarberry, pecan, red mulberry, and black walnut were present in distant background, but in the local stand of forest range vegetation presented here common green-brair (locally abundant) was about the only other plant species present.

Erath County, Texas. Mid-May (vernal aspect at mid- to late spring), hairy wood brome was in soft- to medium-dough stage. FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash) and/or SAF 94 (Sycamore-Sweetgum-American Elm). Absence of sycamore and sweetgum and presence of sugarberry suggested that SAF 93 was more appropraite; however, the former cover type designation for SAF 94 included pecan so choice was unclear.Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

185. A hairy stand- A local population of hairy woodland or Canada brome in the herbaceous layer of a climax sugarberry-cedar elm-pecan bottomland forest that developed along Dry Branch, an ephemeral sstream draining into Bosque River. Both of these "photoplots" were taken at the more upslope edge of the grass stand. The large tree trunk in the second "plot" was of a pecan on which a robust plant of Virginia creeper had found a ready source of support. There were also several individuals of common green-briar. Other plants species were absent which was a notable feature of this local population of woodland bromegrass.

Bromus pubescens (B. purgans) is quite rare in this region. It was only recently officially "discovered" in the Texas' Cross Timbers and Central Prairies vegetational area (Diggs et al., 1999). Its presence attested to the pristine state of this relict tract of mixed hardwood floodplain forest.

Detailed showings of hairy wood brome were presented below with other important plant species of this forest range.

Erath County, Texas. Mid-May (vernal aspect at mid- to late spring), hairy wood brome was in soft- to medium-dough stage.

186. More dead than alive, but alive- Three views of an ancient pecan that is dead except for a narrow band of cambium that supplies water and mineral nutrients to and translocates photosynthate (manufactured food) from two, remaining, small branches in what was left of a once large crown. This is the "bristlecone pine equivalent" of a pecan. This decript, "damn near-dead" pecan is not only still part of this bottomland forest, but it also provides unique habitats for organisms that may not be supplied otherwise. For example, the hollow cavities (which comprise most of the remaining parts of this decaying carcass) afford resting and nesting sanctuary for furbearers, squirrels, and cavity nesting birds. The dead, rotting wood provides nutirients and growing space for various fungii, algae, and mosses. The remaining trunk and large limbs serve as a source of space and physical support for woody climbers which on this tree included trumpet creeper, common green-briar, poison ivy, and coral honeysuckle (Lonicera sempervirens).

Standing dead (or almost-dead) trees are called snags,"a standing, generally unmerchantable dead tree from which the leaves and most of the branches have fallen"; standing trunks that are rotting or, more specifically, are "composed primarily of wood in advanced stages of decay and deterioration"are called soft snags (Helms, 1998). Another definition of snag by the Society of American Foresters (Helms, 1998): "a standing section of the stem of a tree, broken off usually below the crown". Snags are viewed as a form of a forest's biological legacy (see below).

Foliage of narrow leaves at base of pecan was that of Canada wildrye and broadleaf woodoats, co-dominants of the herbaceous understorey.

Erath County, Texas. Late April. (vernal aspect at early tomid-spring). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash) and/or SAF 94 (Sycamore-Sweetgum-American Elm). Absence of sycamore and sweetgum and presence of sugarberry suggested that SAF 93 was more appropraite; however, the former cover type designation for SAF 94 included pecan so choice was unclear.Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

187. Dying trallis- Another ancient pecan, which like the one presented in the last three-slide series, had senesced almost to the point of death. This "over-ripe","way-past" mature tree persisted in the same mixed hardwood bottomland forest of Bosque River. This "just hanging on" tree was another soft snag (see immediately above) with just a few remaining branches that were supplied by a ribbon of live cambium. This snag supported one more species of liana than the one viewed above with mustang grape (the woody climber growing from the forest floor in far lower left foreground) along with trumpet creeper (largest liana directly on the trunk), coral honeysuckle (visible as leaves at base of trunk), poison ivy (on center of trunk), green briar (also on center of trunk).

Old-growth support other growth- An old-growth pecan (one of three climax tri-dominant species) in a mixed hardwood bottomland forest with the native woody vine, coral honeysuckle (Lonicera sempervirens) growing on trunk and lower limbs of this specimen of the State Tree of Texas.

Outer bank of Bosque River, Erath County, Texas. Mid-April; peak bloom of honeysuckle.

No quite as old as the river, but ...- An ancient black walnut with only one living branch (and it was a small one) growing as part of a mixed hardwood forest along Bosque River in north central Texas. The biblically based expression "old as Methuselah" was certainly appropriate in this instance. The small amount of remaining living tissue was characteristic of other trees such as the conifers, bristlecone pines like those in the White Mountains of California (Pinus longaeva), that survive for many years with only one or two strips of living tissue while the rest of the tree slowly rots decay. In this case of a stream bank in a moist river bottom in the subhumid zone wood decay was much considerably more rapid. The only part left living on this battered old black walnut was a small branch near the crotch of the trunk where the first limb (now mostly rotted away) branched off (visible in both slides). And, guess what, the one, small, remaining shoot was blooming. It served as an example of the flower cluster, the catkin, of this species (see below).

The green foliage at base of this snag included Canada or nodding wildrye and broadleaf woodoats (climax co-dominant herbaceous species of this rforest range), naturalized annual ryegrass (Lolium multiflorum), and the native shrub elderberry (Sambucus canadensis).

Like ninety year church parishioners, the ancient "more-dead-than-alive" (senescing) trees are still part of the forest community, the range ecosystem. This ole snag of a still- (though barely) living tree served as a trellis for the following species of lianas (woody vines): mustang grape, trumpet creeper, and poison ivy.

South bank of Bosque River, Erath County, Texas. Mid-April; blooming of black walnut. FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash) and/or SAF 94 (Sycamore-Sweetgum-American Elm). Absence of sycamore and sweetgum and presence of sugarberry suggested that SAF 93 was more appropraite; however, the former cover type designation for SAF 94 included pecan so choice was unclear. Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

Methuselah blooming- One of the few remaining live branches on the more-dead-than-alive, ancient black walnut introduced immediately above, and it was in early stages of sexual reproduction. With backcrop of an azure-blue sky (right after a late norther) this sexual leader might well have been this ancient tree's last attempt at fruit production. These catkins of the monoecious black walnut were all males (closer views immeidately below).

South bank of Bosque River, Erath County, Texas. Mid-April; blooming of black walnut.

Methuselah details- View of several catkins and a few new leaves (first slide) and close-up of one of these catkins (second slide) on the leader of the ancient black walnut presented above. Black walnut flowers are monoecious; these long, hanging catkins were males. All flowers on this leader were male.

South bank of Bosque River, Erath County, Texas. Mid-April; blooming of black walnut.

Start of a black walnut- Young seedling of black walnut that was growing in the Springfield Plateau of northeastern Oklahoma. Seedlings such as this specimen grow very rapidly due to considerable storage of nutrients in the richly flavorable (to humans and squirrels) and extremely hard-shelled nut.

Ottawa County, Oklahoma. Mid-June; early seedling stage of phenology.

How it starts in the soil- Three views at progrsssively closer camera distance of the basal part of a recently germinated black walnut seedling (the one presented in the immediately preceding photograph) in the western Ozark (Springfield) Plateau orf northeastern Oklahoma. This is near the far western edge of the oak-hickory forest (as distinguished from the Cross Timbers form oak-hickory-tallgrass savanna).

The "meat" of the delicious black walnut fruit is primarily the two cotyledons which serve as the stored nutrients for the seedling during germination and emergence.

Ottawa County, Oklahoma. Mid-June; early seedling stage of pnehology.

188. Last act- A pecan that is still part of the forest in its final stage performance as a rotting log. A once-mighty member of the forest canopy as gone back to rest on the forest floor until it once again becomes part of the soil that sustained its life as a member of the governing guild of dominant trees. Now the old cellulose carcass is home to wood roaches, rodents, salamanders, spiders, and centipedes. The decomposing remains of trees (eg. rotting logs), along with those of shrubs, grass, forbs, etc., provide their final ecological role or service as habitat for animals and organisms like fungus, water storage, soil protection, organic matter to improve soil properties, and, ultimately, to return mineral nutrients that it borrowed during its life as part of the forest ecosystem. As with man; so too, trees. "... for dust thou art, and unto dust thou shalt return" (Genesis 3:19).

Frequently, some senior faculty members are characterized as "dead wood" because by measurements of publications and new courses developed they are less productive than when they were "young Turks" still aspiring for their final promotion. Such tenured university professors who have ceased to be highly productive in research output and new course offerings have been described in the lumber or orchard metaphor as "over-ripe" (senescent trees with decaying wood from broken limbs) and past their prime for lumber. Some may even be "hollow trees" taking up space in the forest where younger, fast-growing young trees could be standing using limited resources to produce "new wood" for the "next crop". What such a slanted and uninformed perspective of "dead wood" overlooks (or conveniently ignores) is that these "dead snags" or "downed timber" are also part of the forest. They are "senior statesmen" of the faculty who have matured to that stage of the academic life cycle where they are more involved in developing ("mentoring") graduate students and assistant professors. Or, alternatively (perhaps simultaneously), these "standing dead trees" are doing disproportionately more service on committees (tenure, curriculum, accredation, etc.) or even tutoring or advising more undergraduates, all aactivities which they had to kept to minimums when they were still in the stage of their careers known as "Publish or Perish", the Academy's equivalent of Darwin's struggle for existance. And, finally, when the last of the "old trunk" falls to the forest floor at retirement or death (which ever comes first) the "farewell party" might well be about as educational as entertaing. If the "old tree" left behind any artifacts like books, notes, antique office supplies, a charred pipe (from days when universities were still humane), or interesting memorabilia graduate students, fellow professors, secretaries, or even janitors might "scrounge" the remains for useful items. In ecological terms, recycling nutrients.

Common green-briar. poison ivy/oak were slowly twining up and over the remains of the former forest elder. A species of bracket fungus ws growing on the rotting wood of these big limbs. Canada wildrye, broadleaf woodoats, and hairy wood brome were native perennial grasses that were growing around this big detritis.Giant ragweed and hedge parsley were the principal forbs.

189. Reading plant succession- Sugarberry sapling (rightmost tree trunk; small with warty bark) established by three trunks (roughly 14 inch DBH) of young adult pecan and a fairly large mustang grape (darkest and smallest trunk) left no doubt as to thel fact that hackberry is a successionally more advanced tree species than pecan which it will ultimately supplant on this sere. This was a second-growth mixed hardwood forest on a wet (semi-swamp) habitat. The land shown here was a southeastern slope from which water seeped for protracted periods during wetter and/or cooler weather (autumn through spring except during dry winters). The major plant species on much of the ground layer was common greenbriar (Smilax bona-nox) which with muitang grape grew into the canopy layer to unite all the layers of this forest vegetation.

This young, developing forest had been severely or radically invaded by two exotic shrubs brought in by whiteman for ornamental yard species: 1) Japanese honeysuckly (Lonicera japonica) and 2) callery, the Bradford cultivar, pear (Pyrus calleryana). These horribly invasive species were slowly destroying this seral forest by converting it to a brush patch of species that should have never been allowed to enter North American soil. Invasion by alien (exotic) woody species such as these is one of the gravest threats to forests in the Southeastern Region of North America.

There was no herbaceous understorey--in fact, few herbaceous plants ata all--in this dense and heavily invaded/degraded seral forest.

Erath County, Texas. Mid-March (hibernial aspect; late winter). Early leaf stage of common greenbriar. FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash) and/or SAF 94 (Sycamore-Sweetgum-American Elm). Absence of sycamore and sweetgum and presence of sugarberry suggested that SAF 93 was more appropraite; however, the former cover type designation for SAF 94 included pecan so choice was unclear.Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

Summer- Bosque River

Summer (General)- The following few sets of photographs introduced and presented an overall perspective of the climax sugarberry-cedar elm-pecan forest range in late summer and at peak standing crop.

190. Summer verdure above the Bosque- Deep shade and rich stains of green might give the appearance of coolness, but this local community of a climax, mixed hardwood (sugrberry-cedar elm-pecan), botttomland forest was anything but cool (temperature-wise). The ambient temperature was near (or above) the "century mark" on a blistering late-summer day along Bosque River in the West Cross Timbers of northcentral Texas.

The tree with its large lower limb leaning over the channel of Bosque River was an American elm. The tree to immediate left of the American elm was a cedar elm, one of the three climax dominant tree species of this forest. Trees in the distant background were old-growth pecans growing as a local single-species grove. The herbaceous understorey was co-dominated by Canada wildrye and broadleaf woodoats, the latter of which predominated in the live, summer vegetation (estival society). The yellow composite was giant goldenrod (Solidago gigantea). Most abundant shrubs were woody vines, primarily common green-briar and trumpet creeper. These were the only shrubs that were within the two "photoplots" presented here. Other shrub species--both native and exotic--were presented in many of the other slides herein.

There were two non-native woody species that had invaded local areas of this floodplain forest. Japanese honeysuckle, an aggressive exotic invader, was extremely abundant to the left of both photographs. Japanese honeysuckle has become one of the most ecosystem-threatening exotic species of forest throughout the Southeast and Southcentral Forest Regions (Kellison et al, 1998; Miller, 2003).Another aggressively invasive, alien shrub in this Bosque River forest was common or Chinese privet (Lignustrum sinense). Both of these introduced (originally for ornamental landscaping) woody species have naturalized to considerable degree, especially japanese honeysuckle, and pose major threats to composition, structure, and function of forest ecosystems (Miller, 2003). Kellison et al. (1998) reported that river terraces and levees of southern floodplain forests of North America almost always had sizable cover and proportions of Japanese honeysuckle.

Erath County, Texas. Mid-September (late estival aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash) and/or SAF 94 (Sycamore-Sweetgum-American Elm). Absence of sycamore and sweetgum and presence of sugarberry suggested that SAF 93 was more appropraite; however, the former cover type designation for SAF 94 included pecan so choice was unclear. Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

191. Stifling summer and stagnant water- On north bank of a muddy, almost-still-watered Bosque River a pecan (leftmost; largest tree) and an American elm (right teee) grew as members of a climax, mixed hardwood floodplain forest range. Cedar elm was one of three dominant tree species (sugarberry and pecan being the other two), American or white elm was locally well-represented as in this "snapshot" of the river bottom forest vegetation. Canada wildrye--visible as dead straw of this cool-seson C3 festucoid grass--was the overall co-dominant herbaceous species of the forest understorey. Other distinguishable herbaceous species were giant ragweed and frostweed or white crownbeard. Common green-briar was the major woody species in this "photographic dendrogram", but Japanese honeysuckle was also present at disturbing extent foliar cover and proportion of forest species composition (eg. the green leaves and brown stems in front of pecan).

This and the other photographs labeled as mid-September were taken on a sultry summer afternoon with the thermometer's mercury "flirting" back and forth with 100 degrees Fahrenheit, and no wind with roughly 60% relative humidity. Not a photographer's favorite atmospheric conditions, but the Nikon fm did not seem to mind. Forest, woodland, and even tree-savanna vegetation blocks much to most wind movement, especially in the interior of such plant communities, as well as increasing humidity through transpiration. Comfort benefits to animals afforded from shade provided by trees is thus somewhat negated by reduction of cooling wind flow and increased humidity (water content) of air. Roots, fallen logs, and crowns (in the air or on the ground) all contribution to rduction of falling raindrop impact and, to lesser degree, with snow meltwater. Trees are especially effective in protection of watersheds (= water catchments).

Pecan Bottoms- The next five slide-caption sets featured two pecan groves as local communities within a climax sugarberry-cedar elm-pecan floodplain forest. These are forest stands, within which pecan is overwhelming--if not the sole--dominant tree, characteristically have a poorly developed woody (shrub) underlayer and, instead, a well-developed herbaceous layer. The predominant growth or life form of shrubs are lianas (woody vines) such as mustang grape, trumpet creper, common green-briar, poison ivy, and/or Virginia creeper. These local populations of pecan--most trees of which are near-mature (approaching maturity), mature, or post-mature (senescing: slowly dying)--with a very sparse layer of shrubs and a lush herbaceous layer are known in Texas parlance as pecan bottoms. Most such forest or woodland assemblages have developed on bottomland sites such as floodplains so pecan bottoms is an apt and very descriptive name (as, for that matter, are most of the colorful folk names applied to plant species and vegetation).

192. Inside a pecan grove in summer- Interior of a local population of old-growth pecans having the size and shape of ancient trees of this species. These old-growth specimens were senescing and the very low rate of pecan recruitment (very few seedlings) was not adequate to maintain the single-species dominance of this local community (sub-community) of a climax sugarberry-cedar elm-pecan bottomland forest. Instead, sugarberry, the overall "number one" dominant tree species, was slowly succeeding to eventually become the local dominant (ie. replacing pecan as the canopy dominant). Leaves of sugarberry were distinguishable at right margin of this slide. Pecan, a pioneer tree species, is of such longevity as to persist into the climax forest, but this is often as aged and senescing (slowly dying after reaching maturity) trees. Such pecans are gaints; they are also dying at completion of their life cycles).

The herbaceous layer was dominated by Canada wildrye (most of the straw or dead shoots). Broadleaf woodoats, overall co-dominant, climax, herbaceous species, was present but not well-represented in this local assemblage of a mixed-hardwood floodplain forest. Most of the green shoots scattered among dead shoots (straw0 of Canada wildrye were those of two herbaceous composites: giant ragweed, native annual) and frostweed or white crownbeard (native perennial). There were also a number of green shoots of common green-briar.

193. Changes in the grove- Interior of another pecan grove within a climax sugarberry-cedar elm-pecan bottomland forest. Featured in this "photographic dendrogram" was a mature (a quite old) pecan on the trunk and into the crown of which numerous species of woody vine had grown. These lianas included mustang grape (the species most obvious in this photograph), trumpet creeper, poison ivy, and common green-briar. There were a number (at least six) of other mature pecans visible (in the left background) in this grove, but this picturesque stand of mature pecan trees was clearly on its way out. All of the saplings (and, not visible here, tree seedlings) in this forest community were sugarberry (four saplings to right of and behind the center-pieced pecan) and cedar elm (distant background). Judging from other local stands within and detailed study of this floodplain forest it was apparent that long-lived pecan would persist into the climax forest, but strictly speaking pecan is subclimax whereas sugarberry and cedar elm are the climax dominants that are capable of regenerating at the terminal stage of forest succession. Pecan is an Intolerant species whereas sugarberry has traditionally been classified as Tolerant and cedar elm, though less studied, appears to be Intermediate in regards to shade, competition, etc. (Burns and Honkala, 1990).

The dominant herbaceous species of this forest range was Canada widlrye which is obvious in this photograph as all the amber-colored straw (dead shoots). Broadleaf woodoats, overall the herbaceous co-dominant, was less abundant here. The major forb in this photographic sample was pigeonberry while frostweed or white crownbeard, the overall dominant forb, was second most abundant.

194. Clumb on and burnt out- Deep inside a pecan grove of a mixed-hardwood bottomland forest an old pecan boasted two fire scars and two companion vines, a mustang grape (the larger vine stem) and a trumpet creeper, that used the pecan as a source of support and outlet to sunlight. These two lianas were shown in more detail in the second of these slides. Trumpet creeper was the woody vine aerial adventituous roots clinging to the bole of the pecan. Other woody vines common in the local community seen here were common green-briar and poison oak/ivy. All five trunks of adult trees in foreground extending to mid-ground (one to right and slightly behind the fire-scarred pecan, one seen between these two, and the two larger trunks at distant right) were pecan. The few saplings visible were all sugarberry, the ultimate climax dominant tree species.

Tolerance ratings of sugarberry, cedar elm, and pecan (the three climax dominants of this forest type) are Tolerant, Intermediate, and Intolerant, respectively (Burns and Honkala, 1990). Pecan is a pioneer or colonizing species that requires (or essentially so) bare, mineral soil for germination/emergence (Burns and Honkala, 1990). Probably pecan is more precisely seen as a subclimax species, but it is a comparatively long-lived tree that persist into the climax largely as a function of this longevity. Many or, even, most old-age pecans in the climax canopy are post-mature and senescing whereas sugarberry and cedar elm occupying the canopy are younger and generally much smaller trees. This was especially the case for sugarberry in this floodplain forest. The largest sugarberry on this forest range was a snag that was presented below.

In this view of a local pecan grove within a climax sugarberry-cedar elm-pecan forest conspicuous herbaceous species included Canada wildrye, co-dominant herb (amber-color straw); broadleaf woodoats, co-dominant herb (not obvious green shoots); frostweed or white crownbeard, the major (and most conspicuous) forb species; pigeonberry.

The "other side" of these two fire scars was shown in the next slide...

195. Living chimney- Heat and flames that entered the pecan featured in the preceding two-slide set exited its trunk through a large wound created when and where the lowest limb of this tree was torn off by some accident and agent known but to God. Draft created by the fire burning up from the lower trunk resulted in enhanced combustion of wood, created a large area of burn-out, and resulted in this huge, higher fire scar. This phenomenon is known as a fire chimney. Most of the fire-consumed wood was heartwood and dead or dying sapwood. Most of the tree's live tissue was undamaged (or minimally harmed) so the tree lived on more or less unscathed. Hollow cylinders (things like hollow trees, pipe, grass straws) are almost as strong as solid cylinders so this fire-hollowed tree was not adversely affected, at least not overly so. The fire scar was not life-threatening to the pecan. It was interesting, somewhat unusual, and source of a good object lesson.

Erath County, Texas. Floodplain of Bosque River. Mid-September.

196. Going back to the ground- A nearly gone snag of pecan standing in a climax sugarberry-cedar elm-pecan floodplain forest along Bosque River in northcentral Texas. It was explained variously throughout this treatment of mixed hardwood forest that pecan is a pioneer of early--typically the first--seral stage of forest succession that as a result of long into the forest climax, typically as very large (and very old) trees. Many of these ancient pecans are senescing and others are more dead than alive. (An example of one such "Methuselah pecan" was shown above under the section entitled, Spring- Bosque River.) Still other pecans are dead and in various stages of decay and final incorporation into the Bunyun (series) soil of this bottomland forest. (Other examples of this rotting-and-return phenomenon were also included at various points in this discussion, the Spring-Bosque River section being one of these.)

This rotting pecan snag was all that remained of a once mighty river-bottom monarch. The crown and lower limbs of the "wooden skeleton" crashed to earth and, being more in contact with moist soil, decomposed a faster rates than the snag that was not in such contact but stood in the drier microclimate of the atmosphere. Eventually though, enough water accumulated in the snag's dead wood that the last remains of this forest king were going to the trees final resting place.

Range plant species within this photographic view included cedar elm (immediately behind pecan snag and 9n badkground), pecan (in background), the invasive Japanese honeysuckle (right foreground behind grass shoots), coral honeysuckle ( in front of snag), redbud (large seedling in left margin), American elm (leaves, extreme left-rear margin), red mulberry (upper center across to upper right margin), broadleaf woodoats (green, broad-leafed grass in immediate foreground), Canada wildrye (dead straw among woodoats shoots), and common green-briar (climbing the mulberry above the snag).

197. Gone to grass- "...for dust thou art, and unto dust shalt thou return" (Genesis 3:19). Jehovah's admission to Adam upon expulsion from the Garden of Eden could be applied to this old pecan tree even though it "played by the rules". Those are the rules, however. Ashes to ashes and dust to dust. Atmospheric carbon into plant organic matter into soil organic matter back into atmospheric carbon then again into plant organic matter. The endless biogeochemical cycle of carbon. Ecosystem structure and function. Such was the slow and inevitable drama of nutrient cycling in this ecosystem of a climax sugarberry-cedar elm-pecan bottomland forest along Bosque River.

Students should note that the remains--what little remains--of this pecan is still part of this mixed-hardwood forest range ecosystem. Organic matter in the rotting wood of this decomposing pecan trunk served as mulch to retain water and release it to broadleaf woodoats (green shoots) and Canada wildrye (dead, amber straw), the co-dominant climax grasses of the understorey of this forest range community.

Wooy plants in midground (to the right behind the rotting wood) were a small chittamwood (Buemlia), two young cedar elm, a dead sapling of American elm, and a live sapling of pecan, this latter being the rightmost sapling (directly behind the dead shoot of Canada wildrye). Even though pecan is a pioneer species there are sometime saplings and even eedlingr beneath mature and dying pecans. This pecan sapling was one such. Pecan was remaining as a member of the climax vegetation of this bottomland forest range. Pecan was growing side-by-side with cedar elm, the second of the three climax tree species (sugrberry being the other, and the most tolerant, of the three).

198. Biggest of its kind (and the deadiest)- The snag of this long-dead sugarberry was the largest trunk of its species in the climax sugarberry-cedar elm-pecan forest on the floodplain of Bosque River. None of the adult sugarberry trees in this forest even approached the size (and, most likely not the age) of the largest pecan. Reasons for this were not known. The right-leaning snag behind the sugarberry snag was a pecan that was unusually small for dead adult trees of this species in this forest.

Dead--both standing and fallen--mature trees are one of the structural and demographical characteristics of old-growth forests. Presence of mature dead trees (those that aged to the natural end of their life cycles) was a diagnostic feature that this mixed-hardwood forest was at or near the old-growth stage of vegetation development.

These two views of the same photographic sample of the climax-composition forest Young trees (eg. saplings along left margin in first slide; behind sugarberry snag in second slide) were mostly sugrberry and cedar elm though there were some young pecan trees (in background of both slides). The dominant--almost exclusive--shrub was common green-briar. There was some Japanese honeysuckle (indistinguishable in background). Most of the herbage was dead material (straw of dead shoots) of Canada or nodding wildrye, co-dominant with broadleaf woodoats which were a minor component in the field of view of these two photographs. Green herbage was primarily that of frostweed or white crownbeard and giant ragweed, both of which are native .composites of perennial and annual life cycle, respectively.

Closer examination of the snag (both photographs) revealed that the characteristic short knobs on the trunk of sugarberry extends behind the bark to the actual wood of the bole.

A botanical diversion- Plateau or escarpment live oak or, traditionally, southern live oak (Quercus virginiana= Q. virginiana var. fusiformis= Q. fusiformis) is, depending on which authorities are consulted, either a more xeric, western form, variety, or subspecies of southern live oak or, alternatively, a completely different species (Diggs et al, 1999, p. 716). The definitive work of Muller (1961) established for most students in the Lone Star State that except for a very part of southeast Texas the proper species designation for this live oak is Q. fusiformis.

In the East (Lower) Cross Timbers and West (Upper) Cross Timbers of north Texas this oak is limited to calcareous soils, primarily limestone outcrops. Given the relative sparcity of such edaphic environments in the sandy soils of the Cross Timbers, this evergreen oak is of quite limited distribution in this forest range cover type. Francaviglia (2000, ps. 43) reported that in the Crosss Timbers plateau live oak grew on well-drained but mesic soils avoiding both extremes of wetness and dryness. Francaviglia (2000, ps. 83) quoted journal entries of explorers that mentioned live oak growing on the higher ground of such rivers as the Brzos, Colorado, and Trinity.

The short portion immediately below funrished a quick gaze at an example of plateau live oak on a local habitat (= microsite) of a climax, mixed-hardwood bottomland forest (sugarberry-cedar elm-pecan cover type) that had developed along Bosque River in the West Cross Timbers of northcentral Texas. Plateau live oak of this example were growing on calcareous (limestone-underlaid) soil at a higher elevation of this bottomland habitat. This local, live oak specimen was old-growth and of immense size (and undoubtedly of comparatively great age) consisting of either one, two or three trees (DNA studies were not performed= genotype[s] were not determined). There were three large trunks of such similar size that each trunk appeared to be a genetically individual tree.

It was explained in the Eastern, Southern (Plateau) Live Oak section at beginning of this chapter, Miscellaneous Forest Types, that plateau live oak abundantly reproduces both sexually (acorns) and asexually (root sprouts). With this combined regeneration it is next to impossible to determine if closely growing "trees" are actually separate trees (each trunk is a unique genotype) or simply clones or modules of the same genotype that arose as root offshoots (so-called root sprouts or sucker shoots). Such clones or ramets are actually branches (limbs, is more precise usage) arising off of woody rhizomes commony called "rootstocks". The only ways that sexual or asexual status (originating from acorn or rootstock, respectively) of a given "tree" (shoot) can be determined is by either excavation to trace origin of the shoot to a rhizome or, sometimes, "stump" (basal part of an older trunk) or to determine genetics through DNA (Deoxyribonuclic acid) analysis. Neither of these conclusive (and intensive) methods was used in this instance so number of actual trees per se (genetically unique or individual tree shoots) was not known. This knowledge may or may not have been ecologically relevant to this short lesson.

199. Livin' on the edge- Plateau live oak, a minor species of a climax sugarberry-cedar elm-pecan bottomland forest, growing on one of the highest and driest ridges on the floodplain of Bosque River. This specimen consisted of three large trunks that forked off into big limbs and their various branches forming discontinuous crowns. It was not determined (see immediately preceding paragraph of introduction) if these trunks and crowns (shoots) were one, two, or three individual trees (genotypes). They appeared to be three genetically distinct trees given their similarity in size, but at this close distance they had undoubtedly root-grafted (a typical characteristic of this species) and functioned as one tree. Periodic high yields of acorns assured genetic diversity and potential for dispersal at greater distance from the parent tree(s).

Plateau live oak is only an incidental tree species of the mixed hardwood bottomland forest type that develops on less water-rich environments throughout much of the general Southern Region. In fact, it is often present as incidental and isolated individual trees. Thouth sparse in abundance these "far-and-few-between" trees are often massive in size, extensive in canopy cover, "long on age", and, in general, far more important in the forest ecosystem than suggested by their low rate of occurrence (abundance). Acorns are one of the few natural concentrate feeds (a feature held in common with pecan and walnut) in such Celtis-Ulmus-dominated bottomland forest ranges. These nuts greatly complement and/or supplement ungulate diets that otherwise consist overwhelmingly of forage from Canada wildrye and broadleaf woodoats, C3 grasses that are the herbaceous co-dominants of this range cover type. Plateau live oak is a white oak (Leucobalanus subgenus) and hence produces an acorn crop each year (acorns mature over course of one growing season in contrast to two years in case of the red oaks, Erythrobalanus).

Erath County, Texas. Mid-September (late estival aspect). A subunit or stand component of the general units of a climax sugarberry-cedar elm-pecan bottomland forest cover type. FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash) and/or SAF 94 (Sycamore-Sweetgum-American Elm). Absence of sycamore and sweetgum and presence of sugarberry suggested that SAF 93 was more appropraite; however, the former cover type designation for SAF 94 included pecan so choice was unclear. Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

200. Some big ole good 'uns- Three trunks of plateau live oak growing on a high rise of a bank of Bosque River in a climax sugrberry-cedar elm-pecan bottomland forest in the West Cross Timbers of northcentral Texas. It was explained above that it could not be determined without DNA analysis or, less definitively, by excavation whether these trunks were three separate trees (three genotypes), trunks of the same tree (a single genotype), or two genetically distinct trees. It was even possible that the two, closest adjoining trunks had originated as two seedlings or, more precisely, two plumules from the same acorn. This phenomenon of dual plumules arising from the same acorn was shown in the first section of this chapter, Eastern or Southern (Plateau) Live Oak. The actual genetics of these three trunks and crowns (shoots) is, in effect, irrelevant from standpoint of this discussion of forest range vegetation. Given the common characteristic of plateau live oak to root graft especially among close-growing individuals it was almost certain that these three trunks functioned as a single tree regardless of genotype(s).

Plateau live oak is not common in the deep sandy soil of the Cross Timbers. This oak species is generally less common than post oak (Quercus stellata) or, in some areas, even bur oak (Q. macrocarpa) in the Cross Timbers. Plateau live oak is limited to higher calcium-containing, better-drained soils so that this oak species is of limited distribution in true Crosss Timbers (versus adjoining Grand or Fort Worth Prairie) vegetation. Plateau live oak is typically restricted to calcareous-soil, often those underlaid with calcium parent material such as caliche (encrusted calcium carbonate). Such was the case for the few--though giant--plateau live oak growing in the climax mixed hardwood forest that developed on the floodplain of Bosque River such as the specimen presented here.

In spite of limited occurrence within floodplain forests, such as the sugarberry-cedar elm-pecan cover type featured here, these sparsely distributed live oaks have an impact noticeably greater than what would seem apparent from their comparative scarcity. This is true even for forest physiogonomy because plateau live oak is the only tree species that retains green leaves throughout the otherwise winter period of tree dormancy. Plateau live oak is a species in the white oak subgenus (Leucobalanus) of Quercus. Leucobalanus oaks bear acorn crops annually in contrast to biennial crops of the red oaks (Erythrobalanus species). In size of both trunk and canopy, adult plateau live oaks are rivaled in this floodplain forest community only by mature pecan. Species composition (relative proportion) of this live oak species in the forest canopy, the dominant layer of forest range, is much greater than suggested general frequency or occurrence in the forest community simply because live oak crowns are so large, and again these are these are the only criwns with leaves to conduct photosynthesis during late autumn through early spring.

The evergreen canopy of plateau live oak does usually exclude grasses and forbs from the lower layer(s) of this forest range type. Co-dominant grasses, Canada wildrye and broadleaf woodoats (both C3, cool-season species), along with sand lovegrass, purpletop, and sideoats grama (C4, warm-season grasses) are locally abundant in the semi-shaded habitat beneath huge plateau live oak. Green shoots of broadleaf woodoats and the straw-colored dead shoots of dormant Cnada wildrye were obvious in both of these photographs as well as the immediately preceding slide. Other common herbaceous species were frostweed or white crownbeard and pigeonberry. Shrubs in these two "photogrphic dendrograms" included mustang grape (see the next two photographs), common green-briar, Virgina creeper, and poison oak/ivy. The plant of mustang grape seen in these two photographs was featured in the two slides/caption immediately following this caption.

Other tree species present in forest vegetation presented in these two views included sugarberry and cedar elm, the two (of three) dominant tree species of the forest climax responsible for most regenertion at terminus of forest succession. For instance, sugarberry was represented by the sapling to the immediate left of the three trunks in the first slide an the same sapling seen in the "gun sight" of the two large trunks (between these boles) in the second slide. There were two mid-size pecan (the third dominant tree species) in background of the second of these photogrpahs.

Dead and slowly rotting lower limbs on these live oak trunks were visible in these two and the preceding photograph. As a general rule, once live oaks start to lose teir lowest large limbs, such as seen here, trees had about reached their maximum size and state of maturity and have started to senes (rate of tissue death exceeds that of tissue growth; no net replacement or accumulation of new woody material; necromass formation exceeds biomass accretion). Nonetheless, these trees (or shoots of the same tree as the case may be) had many decades of life left baring natural accidents or "hatchet-happy" human action.

Redirection: Another example of plateau live oak on the floodplain of Bosque River (roughly 1/5th mile downstream from the specimen just shown) was presented above in the first section of this chapter.

201. Drapped grape-Relatively large mustang grape (Vitis mustangensis) on a higher ridge above Bosque River and climbing up into plateau live oak, cedar elm, and pecan in a mixed hardwood bottomland forest. Saplings of sugarberry, the first-among-equals dominant tree species of the climax, were present but too small to serve as much support for this large and, undoubtedly, quite old liana. Mustang grape is by far the largest woody vine in the climax sugarberry-cedar elm-pecan forest. The tree at far left margin of first photograph was a mid-size (and still young). The two center (and also mid-size and young) trees in center of first photograph were cedar elm. These same cedar elms were shown in right midground of the second slide along with a small pecan (centermost of three trees in this second photograph) that was not discernable in the first photograph.

There was also considerable cover of Virginia creeper (eg. on trunk of pecan at left margin of first slide) and common green-briar in the forest vegetation presented in these two views. Herbaceous species included Canada wildrye (visible as dead, light-amber shoots), white crownbeard or frostweed, pigeonberry, and broadleaf woodoats (background of second slide). Canada or nodding wildrye and broadleaf woodoats were co-dominant herbaceous species of this forest range.

This was the same plant of mustang grape that was included in the two immediately preceding photographs that featured an example of plateau live oak. That live oak was at right margin of the first of these two slides and in distant background of the second slide.

Autumn- Bosque River

Autumn (General and Development)- The following sets of photographs presented a general or overall perspective of the climax sugarberry-cedar elm-pecan forest range in autumn. Autumn is the season of "fulfillment" for most plant species (and cetainly of the range plant community). Autumn is also the period of peak standing crop for all woody and almost all herbaceous species. The dominant plant species of the herbaceous layer are the two native perennial grasses, Canada wildrye and broadleaf woodoats. Although both of these bunchgrasses are C3, cool-season species, the point of peak plant development varies from late summer through autumn (exact period for this point in the annual growth or life cycle depends on precipitation during this time frame). Broadleaf woodoats typically do not set and shatter ripe grain until late autumn because this species basically requires both cool- and warm-seasons of growth to complete its annual production cycle. Canada wildrye sets and shatters some ripe grain by early to mid-summer, but most florets are retained through at least mid-autumn. Furthermore, the rank and grain straw-like shoots of these grasses do not undergo much decomposition until late winter or early spring (although leaves fall by early winter). In consequence, the maximum herbage of grass is in autumn which conicides with that of the forbs (both annual and perennial) which reach peak fruit development and ultimate plant size in autumn also.

202. Synopsis shot upstream- On a tributary of Bosque River about one-third mile above the bottomland climax (sugarberry-pecan-cedar elm) forest covered in preceding photographs a seral forest community had developed (apparently following destruction--including use of bulldozers--of the pre-existing forest). Tallest, largest trees in background were eastern cottonwood, a pioneering tree species. Trees (or large, single-bole shrubs) along left margin were black willow. The smaller tree with conspicuous yellow leaves was a young green ash which signaled onset of late succession as green ash is a climax indicator species though not a dominant or even associate member of the potential natural vegetation of this variant of hardwood bottomland forest.

Giant ragweed was the major herbaceous species (even on land that had not been recently disturbed) signaling that advanced community development was still a way off. Johnsongrass was the most abundant grass, but Indiangrass was well-represented which indicated that this vegetation had developed considerably beyond early seral stages. Texas nightshade (Solanum triquetrum) was common and still at peak bloom.

Erath County, Texas. October, peak of warm-growing season (onset of leaf fall in deciduous woody species, fruit ripening in herbs ranging from annual forbs to warm-seasson perennial grasses).

203. Edge of autumn and forest- Exterior of a mixed hardwood bottomland forest in early autumn aspect. At this outermost edge of a climax sugarberry-cedar elm-pecan forest two cedar elms flanked a red mulberry (one elm to left and right of the mulberry in center foreground) on the bank of Bosque River. Herbaceous species dominated the understorey. Canada wildrye and broadleaf woodoats were the overall dominant herbaceous plants, but the most conspicuous herbs in the foreground of these two photographs were the annual forbs, giant ragweed and lambsquarters. Giant ragweed was the dominant native forb on local disturbed areas such as patches of bare soil formed by flood waters. Lambsquarters, a naturalized Eurasian chenopod, was more restricted in occurrence. Other major forbs included frostweed or white crownbeard, a perennial composite tha was the climax dominant forb of the herbaceous layer; pokeberry (Phytolacca americana) and pigeonberry (Rivina humilis) of the Phytolaccaceae (poke family), and red sprangletop (Leptochloa filiformis), a native annual grass commonly growing into robust plants. All of these herbaceous species were growing in the immediate vicinity of vegetation presented in these two slides.

Other woody species included a number of woody climbers including common green-briar, the most widely distributed liana in this forest, and trumpet creeper, a specimen of which was growing on the left side of the left cedar elm. Other woody species that were in background of these photographs and indistinguishable were not listed in this caption.

Erath County, Texas. Late October (early autumnal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash) and/or SAF 94 (Sycamore-Sweetgum-American Elm). Absence of sycamore and sweetgum and presence of sugarberry suggested that SAF 93 was more appropraite; however, the former cover type designation for SAF 94 included pecan so choice was unclear.Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

204. Autumn air- A return to the banks of Bosque River at end of autumn (termination of warm-growing season) and just prior to onset of the cool-growing season brought forth these and many scenes of river bottom mixed hardwood forest vegetation and the range resources afforded thereby. Two old-age pecan (one on either side of the Bosque) and the snag of a sugarberry that was once a flooplain patriarch.

At the terminus of the warm-growing season the understorey dominant was broadleaf woodoats, the major perennial grass at this point of the annual cycle, described variously throughout this section. Dead shoots of the now-dormant Cqanada or nodding wildrye, dominant cool-season herbaceous species stood beside broadleaf woodoats bearing evidence of the partitioning of plant-growth resources in this diverse range ecosystem. Two composites species comprised most of the forb component of the herbaceous layers: 1) giant ragweed, an annual capable of exploiting "scours" along the band that were typically created by the rushing water of spring floods and 2) frostwed or white crownbeard, a perennial that monopolizes stable environments which afford habitats for climax vegetation. The green bush in right background (beneath horizontal right limb of bigger pecan) was the naturalized citrus species called bitter orange (Citrus trifoliata= Poncirus trifoliata).

Erath County, Texas. Early November (autumnal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash) and/or SAF 94 (Sycamore-Sweetgum-American Elm). Absence of sycamore and sweetgum and presence of sugarberry suggested that SAF 93 was more appropraite; however, the former cover type designation for SAF 94 included pecan so choice was unclear.Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

205. Farther down banks of Bsoque River- Downstream from the stand of range vegetation just presented was still more floristic diversity. The first of these two photographs presented in left foreground (left to right) a tree each of Osage orange or bois d'arc, pecan, and sugarberry. Trees in background were almost all pecans. Broadleaf woodoats dominated the understorey through the local range plant community shown in this first slide.

The second slide presented another "photo-sample' of species composition of this sugarberry-pecan-cedar elmbottomland forest. In it all discernable trees (including the largest in center midground) were sugarberry with two exceptions: 1) the small sapling behind and to right of this sugarberry which was a bois d'arce and 2) the pole to right of the sugarberry which was a young pecan. Broadleaf woodoats which was the dominant, warm-season, herbaceous species and frostweed or white crownbeard which was the dominant warm-season forb comprised the great bulk of the understorey. There were a number of dead though still standing shoots of Canada wild rye which was the dominant cool-season herbaceous species.

Greater details of the herbaceous layer of this forest range vegetation were presented in the next two-slide set.

206. A study of the herbaceous zone- Two "photoquadrants" of the herbaceous understorey of a climax sugarberry, pecan-cedar elm climax bottomland forest range along the Bosque River in autumn (and surrering through a severe drought). First photograph presented some green broadleaf woodoats, the dominant warm-season herbaceous species, surrounded by dead and broken-over shoots of dormant Canada or nodding wildrye, the dominant cool-season herbaceous species "escourting" a young pole-size sugarberry, the potential dominant tree of this climax range vegetation.Dead shoot with withered brown leaves in right corner was that of pigeon-berry (Rivina humilis), a sporadically abundant forb of the Phytolaccaceae (pokeweed family).

Second photograph was a all-in-one sample of species composition and structure of the herbaceous layer. Dead, broken shoots were of dormant Canada wildrye; most of the green shoots were broadleaf woodoats; and tall, still-green, wide-leafed forb was frostweed. Wildrye was the dominant, cool-season herb while woodoats and frostweed were overall dominant, warm-season herbaceous species and dominant warm-season forb, respectively.

207. Afternoon autumn light- Clear sunlight shinning behind a representative sample of a climax sugarberry-pecan-cedar elm climax bottomland forest gave viewers this relatively rare glimse of productive and pristine range at peak standing crop at end of the annual plant-growing cycle. The herbaceous understorey was dominated overwhelmingly by broadleaf woodoats accompanied by frostweed or white crownbeard, the dominant forb and overall associate species of the herbaceous layer.

The overall dominant tree was sugarberry, a specimen of which was in left background. Cedar elm (a specimen was in right background), though sometimes locally dominant, was, along with pecan, one of the associate species.

208. Samples of the more complete structure- The preceding slides showing the autumnal aspect of a climax sugarberry-pecan-cedar elm bottomland forest that developed along Bosque River showed range vegetation consisting of tree species (though of several age classes of the various species) and an herbaceous understorey. Some local stands of this forest range vegetation also had a shrub component. In some local assemblages of this range plant community the shrubs consisted mostly of lainas or lanes (climbing woody species). These two photographs--the first, a more distant view and the second, a closer-in view--gave readers an idea of this mixed hardwood forest vegetation when species of woody climbers were part of its structure.

Range vegetation in the first photograph included a young adult of cedar elm (left margin) and a larger specimen of the same in center background. The tree in far right background was a sugarberry. The second photograph featured a young adult of sugarberry (left foreground) and a diverse herbaceous layer consisting of broadleaf woodoats (at peak standing crop and with ripening grain), dormant (dead stalks) of Canada wildrye, and a few frostweed plants. In the background (so as to show height to which climbing shrubs can ascend into trees) common greenbriar (Smilax bona-nox), poison oak or ivy (Rhus radicans= Toxicodendron radicans) and trumpet creeper (Campsis radicans) were represented. The latter was conspicuously climbing a tree trunk in right background.

209. The next generation- On the south bank of Bosque River young trees foretold (barring unforeseen catastrophic phenomenon) of the future composition of this local stand of a sugarberry-pecan-cedar elm climax. A sapling of cedar elm (left) and one of bur oak (right) stood surrounded by a herbaceous layer composed of Canada wildrye, pigeon-berry, giant ragweed, and Texas nightshade (Solanum triquetrum).

Erath County, Texas. Early November (autumnal aspect).

210. Turning colors on the north bank- A young green ash was turning a golden yellow from top of crown downward while a towering eastern cottonwood flanking the ash on the far left with still all-green leaves hearlded the onset of autumn on an upper north bank of Bosque river. Tall, dead stalks to immediate left of the green ash were of giant ragweed, a rank-growing, opportunistic, annual composite representing r-selected species that grew on locally disturbed parts of the bank that had been scoured by flood waters.

211. Sample on the slope- A part of the north bank of Bosque River furnished an example of forest range vegetation that developes on a prominent south slope along watercourses in the Western Cross Timbers. Tree species in this local assemblage included sugarberry, cedar elm, American elm, pecan, bois d'arc or Osage orange with shrubs that included trumpet creeper, poison oak or ivy, and southern or rusty black-haw or, sometimes written as, blackhaw (Virburnum rufidulum) plus an herbaceous layer that included broadleaf woodoats, Canada wildrye, frostweed, and giant ragweed.

212. Sugarberry everywhere no matter what the associate- Two different local stands of a climax mixed bottomland hardwood forest demonstrated that sugarberry was the overall--and the ultimate--dominant of this forest cover type regardless of size of its associates. The first photograph was of an immense cedar elm that dwarfed a pole-sized sugarberry to its left. The second photograph also included immediate left foreground surrounded by young pecan, and a "hemongous" shoot of mustang grape (Vitis mustangensis) in immediate right foreground. The consistency to which sugarberry was "always present" and at various age classes showed it to be the overall dominant of this example of a sugarberry-pecan-cedar elm-American elm-eastern cotttonwood-bois d'arc cast of arboreal characters. In the examples of this climax range vegetation the understorey dominant was Canada wildrye which was dormant and represented only by dead shoots at end of this year's plant-growing cycle.

213. General structure, internal arrangement, and species composition- General views of the interior of a climax mixed hardwood (sugarberry, pecan, cedar elm) bottomland forest above (on upper bank of) Bosque River. In the first photograph a local stand of pecan (two trees at left) and cedar elm (three trees at right) had a diverse understorey conisting of a lower shrub layer represented by a colony of poison oak ot ivy (distinguishable as by its bright yellow leaves) and an herbaceous layer dominated here by Canada or nodding wildrye with frostweed or white crownbeard as the major forb.

The second photograph featured another stand or grove of thisbottomland mixed hardwood forest. In this local range plant community there was an old-growth pecan (center background; four main limbs arising from the short trunk) with a younger pecan to its left (largely hidden by saplings of sugarberry, pecan, and cedar elm) and flanked by two young adult sugarberry (to sides and in front of the picturesque pecan). In front and to left of all these (and with an identifying broken limb) was a bois d'arc or Osage orange. Presence of bois d'arc and red mulberry clearly showed the ecological relatedness of this bottomland forest that developed along a southern river with bottomland forests that formed along rivers far to the north (compare to rivers in northern Kansas as presented above). The herbaceous layer in this grove consisted mostly of forbs with white four o'clock and frostweed being the most and second most abundant, respectively, species.

214. Structure of the understorey- Layers and species composition of understorey of a sugarberry- pecan-cedar elm bottomland forest. This "photoplot" was of the lower layers of forest range vegetation that was below the rightmost tree (with forked trunk) in the first slide in the two-slide set immediately preceding this photograph. A lower shrub layer was dominated locally by poison ivy or oak (birhgt yellow leaves in foreground). Mustang grape, represented by a large individual in right background, represented the arboreal shrub layer (trumpet creeper and climbing plants of poison ivy were other species of woody climber that were in this bottomland forest though not represented in this slide). Frostweed or crownbeard the overall dominant forb of the herbaceous layer was represented by a large plant in immediate center foreground). The locally dominant herbaceous species was Canada or nodding wildrye. This cool-seson festucoid grass was dormant at end of the end of the current, annual plant-growing cycle, but much straw from its shoots bespoke to the cover and biomass that had been produced.

Erath County, Texas. Early November (autumnal aspect).

Travel-log note: The next two (2) sets of slides and captions presented a short walk up to, through, and out of a grove of a climax sugarberry-pecan-cedar elm bottomland forest that was dominated by old-growth pecans. After emerging from this first grove a second pecan-dominated grove of this forest cover type was traversed and described in two (2) sets of slides. Then there was a brief interlude at a single photostation (one slide) showing sugarberry replacing pecan as the dominant climax tree of this forest community before a third pecan-dominated stand was entered and described using three (3) sets of slides only to be greeted by a grove of young sugarberry that, as presented in two (2) sets of slides, again showed pecan ultimately being replaced over the long course of plant succession by sugarberry, the most-dominant of terminal (climax) tree species. Enjoy the short Texas autumn and its lesson in vegetation development in a climax bottomland forest of mixed hardwoods.

215. Aproaching a local old-growth grove- Two species "standing guard" on the perimeter of this local pecan stand in the first slide and from the vantage point on left flank in the second slide prevealed much about this forest range vegetation. The tree in the right foreground of first slide and left foreground of second slide was red mulberry. Individual plants (rather than groups) of this species were distributed throughout this bottomland forest that had developed along Bosque River, including an ephemeral stream draining into th river (presented below). Presence of red mulberry and also Osage orange or bois d'arc (both species members of Moraceae) that was widely distributed throughout this mixed hardwood forest showed the ecological and floristic affinity of this rcverine vegetation to that of rivers as far north as northern Kansas (see again above). The tree to left of red mulberry in first slde and to right and behind this red mulberry in the second slide was a young adult sugarberry which, as shown throughout this treatment of Bosque river forest, was the potential number one dominant tree species of this climax plant community. (These same two trees were also present in the first of two photographs in the next slide set to serve as "landmarks" for viewers.)

The three shrubs in center midground of the second photograph shown here were lanceleaf buckthorn (largest shrub) and two immature plants of chittamwood or gum elastic (Bumelia lanuginosa var. oblongifolia). Chittamwood is widely distribute throughout the West Cross Timbers and Prairies vegetational area, but lanceleaf buckthorn is, in this author's observation, quite limited in distribution, usually being restricted to more mesic habitats. Accompanying these shrubs were saplings of pecan and cedar elm attesting to some regeneration of these tree species that were typically secondary dominants to sugarberry in climax stands..

The herbaceous layer consisted of Canada wildrye (the dominant cool-season herbaceous species), broadleaf woodoats (dominant warm-season herbaceous species), frostweed or white crownbeard (typically the most important forb), and pigeon-berry (a locally dominant forb; usually associate forb to frostweed).

216. Inside a local grove of a pecan bottoms- Several old-growth pecan (only three such trees were visible in these photographs) formed nucleus of a local stand of climax sugarberry-pecan-cedar elm bottomland forest that had developed along the upper bank of Bosque River. For purposes of orientation and continuity the first photograph included the red mulberry and accompanying young adult sugarberry that were featured in the two immediately preceding photogarphs. Also still visible in this first slide (behind mulberry and sugarberry in right background) were shrubs of chittamwood and red or rusty or southern blackhaw along with saplings of pecan and cedar elm.

The three large pecans (right midground and background of first slide; obvious in second slide) were representative of this species, the mature individuals of which were the largest trees in this bottomland forest though generally second to sugarberry in total number of plants, bole density, and crown cover. Furthermore (and most telling from standpoint of succession), sugarberry regeneration far exceeded that of pecan and cedar elm combined (more on this below). Pecan is regarded as the largest and fastest growing of the North American hickories, but also the least tolerant along with having the lightest, weakest wood of the Carya species (Harlow et al., 1979, p. 259).

All of these characteristics were evident--at least as evident as can be shown in still photography--in the scenes presented in these and subsequent slides. Lower rates of regeneration and tree recruitment as compared to sugarberry was just noted and shown, both here and in subsequent slides. The comparatively large (often, immense) size, including diamters of trunk and major limbs of pecan were by old-growth specimens shown here as well as below. Considerably less visible in these photographs was the breakage of limbs (of large dimensions) that littered the ground beneath most of these huge and over-mature pecans. Pecan bottoms abound in downed limbs and branches and potential "widow makers". Do not tarry under old pecan trees in windy weather!

The herbaceous layer was described with several slides above and below and most commonly consisted of broadleaf woodoats (warm-season dominant), Canada or nodding wildrye (cool-season dominant), frostweed or white crownbeard, and pigeon-berry. Locally giant ragweed was the dominant herbaceous species, and which with having the largest shoots of any herb was infrequently the sole herbaceous species. Such single-species herbaceous stands were usually confined to stream banks where flood waters had scoured away existing litter and vegetation thereby forming microsite-sized "old fields" for this pioneering, r-selected native weed (an annual composite). Beneath huge pecans in this grove such an environment (a natural "old field") existed with local exclusive populations of the rank-growing giant ragweed. Individuals of giant ragweed were conspccuously smaller than plants growing as weeds in crop fields, fence rows, etc. Such was likely response to conditions of lower light intensity and duration under canopies of pecan and associated trees. All of these phenomena were featured in the second of these two slides where several plants of giant ragweed were growing in front of the backdrop of a large pecan trunk. This area had not been grazed by livestock, including free-ranging feral swine, so animal disturbance was not a factor in giant ragweed habitat.

Conspicuously absent was any regeneration (replacement or recruitment) of pecan beneath canopy of established (and "way past" mature) pecan that were potentially parent trees.

217. At the edge of a second pecan-dominated grove- In distance of a short walk from the grove of old-growth pecan described in the preceding two sets of slides was another local assemblage of the climax sugarberry-pecan-cedar elm forest on the upper bank of Bosque River. Here too, pecan provided all of the large (and, presumedly, old) trees; however the other two dominant tree species were represented (only sugarberry was visible and those being in background of both of these photographs).

Unlike the stand described immediately above this pecan grove had some pecan reproduction, both sexual and asexual (although mostly asexual). The young sapling to immediate left of the foreground pecan (both slides) might (with some good fortune) replace the large, in-its-prime pecan. Details of asexual reproduction via stump suckers or sprouts from leaning pecan in background was shown in detail in the next two-slide set.

The understorey of this pecan-dominated grove was mostly that of the herbaceous layer including broadleaf woodoats (dominant warm-season herb), Canada or nodding wildrye (dominant cool-season herb), frostweed or white crownbeard, pigeon-berry (Rivina humilis), and--as shown below--local populations of nimblewill (Muhlenbergia schreberi). In vernal societies hedge parsley (Torillis arvensis), an annual Eurasian weed (Umbelliferae), is a locally dominant forb in the understorey of this bottomland forest.

Shrubs, other than occasional and incidental shoots of poison oak (nonclimbing form), were absent from this local grove with one important exception. That exceptional species was the naturalized and highly invasive Japanese honeysuckle (Lonicera japonica). This horticultural introduction (from Asia) has naturalized over a large region of North America and become one of the more aggresive exotic invaders of forest communities, often threatening survival of native species and some functions of forest ecosystems. Japanese honeysuckle became particularily widespread in central and east Texas where it is frequently the botanical equivalent of feral hogs. Samples of natural vegetation of the sugarberry-pecan-cedar elm climax presented throughout this section were more-or-less free of Japanese honeysuckle except at local scale. For instance, the twinning shoot entangling the pecan sapling at left of mature tree was that of Japanese honeysuckle.

218. Points of interest inside a second pecan stand- Interior of the grove dominated by old-growth pecan introduced in the preceding two-slide set. The herbaceous layer, which made up almost of the understorey in this old-growth grove, consisted of broadleaf woodoats (beautifully gracing foreground of the first of these three slides), Canada or nodding wildrye, frostweed or white crownbeard, pigeon-berry, and, locally, nimblewill (an eragrostoid grass).

One of the major points of emphsis in this set of photographs was regeneration, especially asexual reproduction, of pecan. Pecan can, under the so-called "right conditions", produce shoots off of stumps (stump sprouts, water shoots) that develop into adult trees. In fact, some of the commercially important cultivars of pecan can be established from grafts on genetically selected root stocks (Burns and Honkala, 1990). Production of suckers or sprouts from base of a trunk of a mature and not old-aged pecan was visible in all three of these photographs.

The second and third photographs provided closer views of this phenomenon. There were two distinct suckers or sprouts arising from an injury at the trunk base of the leaning pecan. These sprouts were each an example of an epicormic shoot or epicormic sprout, "a shoot arising spontaneously from an anventitious or dormant bud on the stem or branch of a woody plant often following exposure to increased light levels or fire" (Helms, 1998). Cause of injury was unknown, but did not appear to be a fire scar. (There was no evidence of fire in this bottomland forest.) Viability of either or both of these "stump" (basal shoot) sprouts was yet to be proven. Apical dominance of the clonal parent might function to retard rapid development of these sprouts. Such morphological development was an obvious ecophysiological "bet hedging" by this tree. If something happened to the parent tree (shoot) these secondary shoots would be available to maintain the genet (genetic individual; genotype of this plant). The second and third slides presented these two epicormic shoots from opposite sides.

There were also some saplings in this stand (behind vegetation featured here) that had arisen by sexual reproduction (ie from the nut).

Another notable feature of some pecans is the leaning, bent, twisted, warped, or otherwise deformed shape of their trunks. This may be of practical importance only or, at least, mostly from the perspective of saw timber, of which pecan is not particularly important or valuable. It is, however, a common enough characteristic of this species to be remarked upon. Causes of the various individual deformed trunks of pecans growing in this forest were unknown. Such were noted and the old advantage involked, "As the twig is bent so grows the tree".

219. True tales told by plants- A sugarberry seedling, small sugarberry sapling, and a cedar elm seedling were growing growing by a large, mature pecan.The cedar elm seedling was in immediate center foreground, sugarberry seedling was to left of pecan, and sugarberry sapling was immediately in front of pecan. In addition, there were two small pole-sized sugarberry (one to left and opposite; one to left and rear of big pecan). There was no such reproduction of pecan.

Any beginning student of ecology could interpret this lesson: the tree species that achieve dominance in the future composition of this forest was sugarberry or, even, cedar elm and not pecan. This pattern of successional replacement was repeated throughout this climax sugarberry-pecan-cedar elm bottomland forest (see even more below). It was shown above as well as in subsequent slides that most of the outstandingly large trees growing on this forest range were pecan. Yet, in spite of some regeneration pecan was being replaced by sugarberry (with both cedar and American elm holding their own).

Explanation for the presence of huge pecan and smaller (by inference, much younger) sugarberry was not known by this author or living locals. Certainly sugarberry does not grow to the relatively enormous dimensions of pecan, the largest hickory in North America, but this fact did not explain absence of sugarberry of comparable (relative to pecan) size and age. One could speculate that perhaps larger sugarberry had been cut for fuel wood while leaving the beloved, nut-bearing pecans. Such a speculative proposal would not explain presence of cedar and American elms of size and age roughly comparable to the larger-growing pecan. Nor would this or related conjectures explain why numbers and density of sugarberry was increasing while that of pecan was static or slightly declining.

A much more rational explanation was to be found in silvics of these species, especially in regards the phenomenon of tolerance. Tolerance rating of sugarberry [hackberry, in general] varies (Wenger, 1984, p. 3) as to forest site (site-specific), but it is considerable higher or greater in this feature than the fairly intolerant pecan (Burns and Honkala, 1990). Pecan is basically Intolerant ((Wenger, 1984, p. 3) though less so than cottonwood and willow (ie. the only associated species more Intolerant than pecan are pioneers like cottonwood). Burns and Honkala (1990) regarded pecan as subclimax. By contrast, sugarberry is relatively tolerant to intermediate (though again this is site-specific) and is capable of establishing in the forest understorey (Burns and Honkala, 1990). Finally, sugarberry is a relatively short-lived tree (about 150 years is maximum) in contrast to the fairly long-lived pecan. A combination of these two silvic features rationally explained size, successional replacement, and forest composition described above.

This one photograph provided pictoral evidence of the same.

The understorey shown in this local forest assemblage included both woody and herbaceous species. The latter included broadleaf woodoats, Canada wildrye, frostweed, white four o'clock, and nimblewill. The major shrubs were common greenbriar and the invasinve alien, Japanese honeysuckle.

220. Entering a third grove with old-growth pecan- General panned view of another stand of sugarberry-pecan-cedar elm bottomland forest that developed on the broad, upper bank of Bosque River in the West Cross Timbers of northcentral Texas. From this distance most of the major range plant species of this forest vegetation were present.Most of the largest trees in background were pecan, but there was limited regeneration of pecan and instead almost all of the mid-size and smaller trees were sugarberry and cedar elm. Mustang grape supplied a conspicuous arboreal shrub component (visible in background) this diverse forest range community though common greenbriar was far more abundant (and annoying to the author). Chittamwood and southern or rusty blackhaw provided a taller shrub element while poison oak or ivy contributed most of the lower shrub layer.

Herbaceous species included broadleaf woodoats (warm-season dominant), Canada wildrye (cool-season dominant), frostweed or white crownbeard, and pigeon-berry as the major grasses and forbs. There were small local populations of nimblewill, a widely distributed eragrostoid grass of eastern deciduous forests.

A "sliver" of the trunk of a sugarberry was in estreme right margin symbolizing successional replacement of the generally subclimax pecan by the more tolerant and climax sugarberry.

221. More true tales told by plants- Young adult trees of the Tolerant- to Intermediate-rated sugarberry (Wenger, 1984, p.3; Burns and Honkala, 1990).were crowding this old-growth representative of the largely Intolerant-rated pecan (Wenger, 1984, p. 3) at the edge of another pecan-dominated grove of climax sugarberry-pecan-cedar elm bottomland forest. (See preceding captions for explanation of size, age, age class, etc. relations of these tree species based on the phenomenon of tolerance.) This grove on upper bank of Bosque River in northcentral Texas appeared to have been spared major disturbances so that plant succession had progressed to the climax stage (or near it). The subclimax pecan (Burns and Honkala, 1990) was being succeeded by the generally short-lived but climax sugarberry along with cedar and American elms which are Intermediate in tolerance rating (Wenger, 1984, p.3; Burns and Honkala, 1990, ps. 808-811).

Sugarberry regeneration was visible asa young tree (left foreground of first slide) and sapling (at left and behind small sugarberry tree in first slide and at left margin of the second slide). Cedar elm reproduction was also evident at seedling stage as, for example, seedlings to front and left of the featured old-growth pecan (most noticable in lower left corner of second slide).

Technical note: the Epson Perfection 700 slide scanner is far from perfect. The one used in production of this publication managed to "naturally" crop almost every slide (among several thousand scanned) including removal of most of the sapling and seedlings just referred to. This author recommends that readers not purchase this shoddy equipment. On the scanner purchased by this photographer, two (so far) of the little plastic tabs that retain slides in the slide holder had to be glued on to prevent breaking off. Epson makes inferior equipment using the cheapest of material-- and, in experience of this author, most Epson service representatives are arrogant and of no assistance to the customer. Avoid Epson scanners!

The other major lesson taught by these two photographs is that of the sapling with compound leaves growing in front of the immense, old-growth pecan. (This sapling was also visible--though barely perhaps--in the immediately preceding and succeeding slides.) At first glance this plant appeared to be a hearty pecan that would more than likely replace the large, still-in-its-prime pecan. Unfortunately (for the pecan race) this was not the case. Instead this sapling, which at first "tricked" this author-photographer, was tree-of-heaven or Chinese sumac (Ailanthus altissima), an alien shrub (horticultural introduction) that was a horribly invasive brush species in this locality. Pecan had not reproduced in this local habitat (microsite) and instead had an noxious exotic plant invade its immediate vicinity. Fortunately (for the pecan species) tree-of-heaven (a member of Anacardaceae) has--much like sumac--a shallow root system that develops aroung woody rhizomes so that smaller plants are easily pulled from the soil. The author made sure that this sapling of tree-of-heaven was ripped out as soon as these photographs were taken. The lesson of invasive alien plants--even in pristine climax vegetation--was recorded and shared with students (ya'll take every opportunity to destroy such exotic invaders).

222. Leaving the third grove and entering a new successional order- Upon leaving the third local stand of climax sugarberry-pecan-cedar elm bottomland forest described in this section (background) another forest stand was entered. This latter stand was composed mostly of young adult sugarberry. It was uphill from the grove dominated by old-growth pecan. This local difference in relief might have been a factor involved in differences of dominant tree species; however, species composition of shrubs, grasses, and forbs was obviously the same on both these local microsites. Major grasses were broadleaf woodoats (the dominant, warm-season herbaceous species), Canada or nodding wildrye (the dominant, cool-season herbaceous species), and nimblewill a (locally abundant but otherwise minor grass species). Major herbaceous species in this local "photo-plot" were Canada wildrye, which was easily distinguished by its dead shoots, broadlaeaf woodoats, pigeon-berry, and giant ragweed (annual pioneer composite species) in that approximate order of dominance. Pigeon-berry and giiant ragweed grew in dense--sometimes exclusive--populations so as to be local (microsite) dominants. Pigeon-berry, obvious as wilted green plants growing in groups in the foreground, was the local dominant over much of the area presented here.

Leading edge of the sugarberry-dominated grove was represented by two sugarberry at left foreground. Three (3) photographs of this sugarberry stand were shown and described in the next three-slide "installment" immediately below.

223. The next reign and final order of successiona- A stand of sugarberry had developed on bottomland that was slightly uphill from and adjacent to the last of three groves dominated by old-growth pecan (background of first photograph). These sugarberry were adult trees though young and still relatively small as compared to potential mature size. Tolerance of sugarberry varies though it has generally been interpreted as having a rating of Intermediate while pecan, the least tolerant of the hickories (Harlow et al., 1979, p. p. 259), was rated as Intolerant (Wenger, 1984, p. 3; Burns and Honkala, 1990).

All trees of pole-size or larger shown in the second and third of these three photographs ("photoplots" in deeep interior of the sugarberry stand) were sugarberry. No pecan were present: there had been no regeneration of pecan i n this large local stand within the climax sugarberry-pecan-cedar elm bottomland forest that developed on the upper bank of Bosque River. There was regeneration of cedar elm as represented by a large seedling or small sapling in center midground of the first and third photograph and in immediate near-center foreground of the second photograph.

Small saplings with compound leaves (and closely resembling pecan unless buds be examined) that were visible in all three photographs, especially conspicuous in left foreground of second slide, were tree-of-heaven, a dreadfully invasive exotic.

Native shrubs included chittamwood, an example of which was the lower left corner of the second slide, mustang grape, such as the one in left background of first slide, common greenbriar, one of which was climbing a tree-of-heaven in the first slide, and poison oak or ivy. The dominant herbaceous species in understorey of this stand was Canada or nodding wildrye. It was conspicuous as the straw or amber-colored dead shoots topped by spike inflorescences. Broadleaf woodoats, the dominant warm-season herbaceous species throughout this forest, was present but not abundant in this local sugarberry stand. Frostweed or white crownbeard and pigeon-berry were the major forbs. Several plants of both species were prominent in foreground of the first and third photographs.

224. New look of the final order of development- Sugarberry trees that were probably approaching maximum size and age for this site grew on the uphill perimenter of a bottomland forest stand comprised almost entirely of this single species. There had been some regeneration of cedar elm (as shown in the preceding set of slides) in the interior of this local stand (eg. small trees in right-center background), but pecan was absent at any and all age classes.

The forest understorey shown in this "photoplot" consisted of herbaceaous species except for an occasional plant of the exotic and highly invasive Japanese honeysuckle. Major plant species in this herbaceous layer were Canada or nodding wildrye, the locally dominant grass, pigeon-berry, and frostweed or white crownbeard in that approximate order of general abundance and cover.

225. Jack be nimble, Jack be quick, Jack jump over the log- Local colony of nimblewill (Muhlenbergia schreberi) growing at base of an old-growth pecan and beside a fallen limb of the same in a climax bottomland forest of sugarberry, pecan and cedar elm dominants on upper bank of Bosque River in the West Cross Timbers of northcentral Texas.

Also present within and around margins of this local population of colony of nimblewill were seedlings of cedar elm, senescing shoots of pigeon-berry, some Canada or nodding wildrye, and a twisting shoot of the alien invader, Japanese honeysuckle. Although it was not the major lesson of this slide caption it should be noted that there was not any reproduction or the Intolerant pecan (Wenger 1984, p. 3) , but rather regeneration of the Intermediate-tolerance cedar elm (Burns and Honkala, 1990).

Nimblewill is one of the more widely distributed grasses in the eastern deciduous forest. Its range extends from Maine and Manitoba south to Florida and the other Gulf States westward to the Plains States. Although nimblewill is a weedy invader of horticultural turfs it is probably best regarded as an increaser on forest ranges when compared to decreasers like broadleaf woodoats and nodding wildrye.

Nimblewill is frequently stoloniferous with adventituous rooting from lower nodes. Thus it was impossible to determine from field observation if population shown here was of one or several clonal plants (genetic individuals; genotypes) or if included numerous genetically unique individuals.

Erath County, Texas. Early November; grain-ripening stage of phenology.

226. Nimble shoots- Shoots of nimblewill in a local population growing by an old-growth pecan in a climax sugarberry-pecan-cedar elm bottomland forest on upper bank of Bosque River in northcentral Texas.

Erath County, Texas. Early November; grain ripening stage of phenology.

227. Cool-season and warm-season citizens- Local assemblage (or a "photosample") of the herbaceous layer of a climax sugarberry-pecan-cedar elm bottomland forest that developed on upper bank of Bosque River in West Cross Timbers of northcentral Texas. First photograph presented the full floristic array of major herbaceous species in this understorey: Canada or nodding wildrye (dominant cool-season herb and most abundant one at this botanical party), broadleaf woodoats (dominant warm-season herb), frostweed or white crownbeard, and pigeon-berry. White crownberad (the wilted, green-leaved forb) and dead shoots of the dormant Canada wildrye shared the spotlight in the second photograph.

Erath County, Texas. Early November.

228. Telling old trunks teach ecophysiological lessons- Snag of a former monarch pecan and past-its-prime mature and senescing pecan stand as remnants at outer part of a grove dominated by old-growth pecan. Smaller tree in distand background (appearing to be between the two pecan) was a bois d'arc or Osage orange. A young adult sugarberry was behind the bois d'arc. Understorey consisted of a shrub layer consisting mostly of common greenbriar and poison oak or ivy (nonclimbing form) and and herbaceous layer dominated by broadleaf woodoats (in warm-growing season) and Canada wildrye (in cool-growing seson) with the major forbs being frostweed or white crownbeard, giant ragweed, and/or pigeon-berry depending on local habitat (microsite).

Perhaps the most striking feature in this scene was the shedding (loss) of dead limbs from the snag and off of the live but over-mature pecan. The ground beneath the snag and extending partly under crown of the live pecan was covered to considerable depth by the litter of spent limbs some of which were of enormous size. Hickory species are renowned for the strength and resilence of their wood (best of all wood for tool handles for instance), but pecan is the least in these wood qualities. Pecan wood is relatively weak and fallen timber from crowns of dead trees are a common characteristic. Obviously all wood rots sooner or later, and snags (trunks of dead trees with missing limbs and/or branches) represent stages in decomposition of standing dead trees. This state of tree breakdown is more prominent in large trees that have big, spreading limbs like pecan. There is also loss of lower limbs from pecan when these older and, often, larger units progress through senescence and, eventually, death. Such shedding is a dramatic version of self-pruning. Another cause of limb loss is breakage due to wind, gravity, ice coating, and so on. These sorts of injury (defoliation) affect healthy limbs, branches, and twigs. The live pecan had examples of both natural prunning via senescence (big knot on left side of trunk below the four large limbs) and defoliation by storm injury (outer portion of lower left limb that had extended beyond snag). A striking example of release from apical dominance was the nearly vertical branch that had developed below the lost (storm-torn) portion of the lower limb. Old trees tell such grand stories if one takes the time to experience them.

229. On the margins- A small motte of southern live oak (Quercus virginiana= Q. fusiformis= Q. virginiana var. fusiformis) growing on the outermost edge of a climax sugarberry-pecan-cedar elm bottomland forest on upper bank of Bosque River in West Cross Timbers. Growing beside the motte was an old-growth specimen of pecan (left and slightly to rear of live oak in first slide) and an individual of bois d'arc or Osage orange (right and to rear of live oak in second slide). Presence of bois d'arc as well as numerous trees of red mulberry (presented both above and below thes slides and catpion) showed floristic and ecological affinity with river bottom forest as far north as northern Kansas (recall from above). Also growing to right of the live oak motte were four adult pecans of intermediate size and, by extension, age.

Motte is a somewhat distinctive as used in Texas (and perhaps adjacent areas). Scifres (1980, p. 322) defined motte as "a distinct clump, usually circular, of woody plants in open grassland or shrubland". The term is applied especially to clumps (ranging in size from a few square yards to several acres) of live oak where there are numerous oak shoots-- some of which are of different trees (unique genotypes) while others are clonal sprouts from woody rhizomes or "rootstocks from established trees--that form a distinctive group and with other plant species growing therein. Field observation is usually inadequate in determining whether any given shoot is clonal (of asexual origin from roots or rhizomes) or a unique genotype (of sexual origin from acorns). Cover provided by live oak functions as the nurse plant phenomenon such that certain species are more common, larger, vigerous, etc. in and/or around edges of live oak mottes than in adjacent, surrounding habitats with different groups of plants.

In the forest vegetation presented here species composition of the understorey did appear to be different beneath live oak than outside the motte. This understorey was a combination of low shrubs, woody climbers, and herbaceous species yet it had fewer plant species than in most other parts of this forest. The two most abundant species were Canada wildrye and common greenbriar. Poison oak or ivy of the nonclimbing form was the second most common shrub. There was some frostweed, but considerably less than throughout the vegetation as a whole.

In this general region of northcentral Texas live oak is almost invaribly restricted to calcareous soils and therefore is restricted to tallgrass-dominated grasslands like those of the Grand Prairie and does not grow in the Cross Timbers savanna. These two general range types have developed in an intricate mosaic of range plant communities with diverse ecotonal vegetation among these. Live oak frequently grows in such ecotones, especially as mottes.

Mottes of live oak were treated as a separate section in the Live Oak chapter.

The forest range vegetation shown here was ecotonal (of a transitional nature) between Grand Prairie grassland that was just beyond this forest and the West Cross Timbers bottomland forest that developed on the upper bank of Bosque River. This bottomland forest was primarily (or overall) a sugarberry-pecan-cedar elm climax as was described above. Along the perimeter of this climax forest vegetation there was a slightly different floristic "flavor" with relatively less sugarberry and more plants of incidental or minor species including black walnut (Juglans nigra) and boxelder (Acer negundo).

Erath County, Texas. Early November (autumnal aspect). Ecotonal range vegetation of the following designations. FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash) and/or SAF 94 (Sycamore-Sweetgum-American Elm). Absence of sycamore and sweetgum and presence of sugarberry suggested that SAF 93 was more appropraite; however, the former cover type designation for SAF 94 included pecan so choice was unclear.Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

Travelog Note: An ephemeral (dry most of the year) stream that drained into Bosque River provided a tall, broad bank that provided an environment for forest vegetation that was somewhat different and more distinctive than that which had developed along the main channel of Bosque River. The Soil Survey for Erath County, Texas (Soil Conservation Service, 19) designated this usually dry tributary of the Bosque as Dry Branch. That designation was used and the dynamic vegetation associated with that tributary was presented in the description of bottomland forest range that followed.

230. A sylvan lineup- Mature trees of cedar elm, black walnut, and pecan (left to right) grew on the outermost edge of the tall, steep bank of Dry Branch that drained into Bosque River in the West Cross Timbers of northcentral Texas. A large sapling or small pole of sugarberry growing just off of this bank's edge was shown in the second of these three slides. Most of the huge trees on the opposite upper bank (background of all slides) were pecan though there were also some cedar elm, black walnut, and boxelder associated with these old-growth pecan. Comparatively large red mulberry grew on sides of the proximate (near) bank just below the edge of it. The major shrubs were common greenbriar, poison oak or ivy, southern or rusty blackhaw, and eastern redbud. Other less abundant shrubs were mustang grape, Virginia creeper, and trumpet creeper. Most tree seedlings and saplings were of pecan. These were so abundant that replacement of mature pecan was assured barring catastrophic disturbance or devestation of the forest as for instance--and abhor the thought--for commencial development by modern, urban man.

The dominant herbaceous species, which was also the dominant of the understorey in toto, was broadleaf woodoats, though in very local areas Canada or nodding wildrye warrented this distinction. These were the major warm-season and cool-season herbaceous species, respectively. Forbs, though widespread and abundant thoroughout the rest of this bottomland forest, were for all practical purposes absent from this range vegetation.

231. Autumn in the air; colors all around- Interior of a climax sugarberry-pecan-cedar elm bottomland forest in West Cross Timbers. This grove was on the upper bank of Dry Branch, an ephemeral stream draining into Bosque River. Large tree on the opposite bank of Dry Branch (left background) was a massive old-growth pecan. Foremost and rightmost tree was a cedar elm whereas the tree to its left (right-of-center midground) was a yound adult pecan. Leftward leaning tree in background was a black walnut. Shrub species included common greenbriar, the dominant shrub, redbud, southern blackhaw, mustang grape, poison oak (ivy), and trumpet creeper

Overall understorey dominant was broadleaf woodoats with Canada or nodding wildrye as associate. Most common forb was white crownbeard or frostweed.

232. Beauty in structure (or "Oh shucks, its jist plain purty")- Textbook example of a climax, mixed hardwood, bottomland forest that developed along Dry Branch, the channel of an ephemeral stream, that drained into Bosque River in northcentral Texas. The first of these two "photoplots" panned across both banks of Dry Branch while the second was a nested "photoplot" that provided greater detail of a key portion of the first or larger "photoplot". In the panning, across-Dry Branch view the large tree (left background) leaning over the channel was a black walnut (more discussion of it farther below) whereas the largest tree (with woody vine) and two trees behind it (on the right channel bank in midground) were pecan. In the second slide, the nested "photoplot", the large pecan with its liana (a mustang grape) that was introduced in the panned "photoplot" was shown at shorter focal length and from a different angle. This second "photoplot" also included a pole-sized black walnut and generally showed more clearly (at closer distance) the various major species of this amazing vegetation.

The herbaceous layer, the more prominent part of the understorey of this forest community, was dominated by broadleaf woodoats (this was also the major warm-seson herbaceous species). Canada wildrye (major cool-season herbaceous species throughout most of this forest range) was present only in small to trace amounts on upper banks of this ephemeral stream channel. Major shrubs included common greenbriar (most abundant and general dominant of the shrub layer), poison oak or ivy, mustang grape (including the prominent example already noted), and eastern redbud. The latter was exemplified by the small specimen growing beside the largest pecan (to left of pecan in both slides). There was much reproduction of pecan and less of black walnut. Large seedlings and small saplings of pecan were visible behind and to right of largest pecan (right midground of first slide and right foreground in second slide).

The immense tree with the symetrical forked trunk across Dry Branch in center background of the second slide (and also in the first photograph of the very next two-slide set) was an old-growth pecan.

233. Vegetation fit for a nut-lover- On the level top of the upper bank of Dry Branch that drained into Bosque River this local assemblage included black walnut and pecan with both species represented by numerous age classes. Regeneration of these nut-bearing hardwoods was guaranteeded other than by the intervention of Man--the Manipulator of Ecosystems; the one entrusted by his Creator to cherish creation--who alone holds the power to destroy, in human time scale, such a remarkable forest plant community.

These two "photoquadrats", with a nested arrangement (explained later in this caption), were subplots of the larger plot presented in the immediately preceding two-slide set that panned across and over Dry Branch to include both of its banks. Many of the same "landmark" plants in those "photoplots" were included in these two photographs.

Two pole-size black walnut (foreground of first photograph) and several black walnut seedlings joined pecan which was one of three dominant tree species of this bottomland forest and the major tree species along this ephemeral stream. The second photograph was a closer-in view of part of the range vegetation introduced in the first photograph, specifically the mature (larger) and younger (smaller) pecans in right midground.

Broadleaf woodoats was the dominant of the herbaceous layer which was the prominent part of the understorey. Canada wildrye, the domiant cool-season herbaceous species throughout much of this forest range, was present only in trace amounts in the vegetation shown and described in these two "nested photoquadrants". Likewise, forbs were essentially absent. There was a rich array of shrubs including common greenbriar, the most abundant shrub, mustang grape (a specimen of which was to the immediate left of, slightly behind, and climbing the larger pecan in the second slide), poison oak or ivy, trumpet creeper, and eastern redbud (one plant of this species was growing to the immediate left of the larger pecan in the second slide).

234. Above Dry Branch- An ephemeral stream shown as Dry Branch in the Soil Survey, Erath County, Texas (Soil Conservation Service, 197) that drains into Bosque River had cut a deeply incised channel in the alluvial Bunyon soil on which a climax mixed hardwood (sugarberry-pecan-cedar elm climax) bottomland forest had developed. Along this this streatch of Dry Branch several mature to senescing trees of red mulberry (Morus rubra) sported characteristically large leaves which had turned a bright yellow that shone colorfully on an autumn morning. The two largest trees in center foreground of these two slides represented this species, the presence of which in this climax community showed the affinity of this river bottom forest to those in as far north as northern Kansas (see above).

Immense trees (though baredly visible) in background were pecan. Dead grass shoots were those of dormant nodding or Canada wildrye, the dominant cool-season herbaceous species of this forest range.

235. Looking down on Dry Branch- Still more of the remarkable biodiversity and structure of a climax mixed hardwood bottomland forest in the West Cross Timbers of northcentral Texas. A young, pecan (left trunk) with an attending common greenbriar climbing its straight bole and a black walnut (big tree at right) stood above seedlings of sugarberry (lower left corner) in the first of these two photographs.

The second photograph presented another perspective of the black walnut introduced in the first slide and revealed that it was noticeable leaning (a geotropism to its position on the steep, sandy bank of this ephemeral stream. Sapling in center midground of this second photograph was a boxelder (Acer negundo). A specimen of eastern redbud was to immediate right of this boxelder. The golder leaves of poison oak and some everpresent woodoats in the foreground added to the understorey of this diverse bottomland forest range.

236. Down in Dry Branch- The herbaceous layer of range vegetation in a mixed hardwood forest that developed along Bosque River. This ephemeral stream was designated Dry Branch in the Soil Survey of Erath County, Texas (Soil Conservation Service, 1973). Tree species along this stream included pecan (the huge tree on the upper left bank in first photograph), sugarberry and cedar elm (mostly seedlings and saplings), red mulberry (young adult and over-mature, senescing trees), black walnut, and a sapling of boxelder. Obviously the dominant herbaceous species on banks of Dry Branch was broadleaf woodoats with giant ragweed, the major forb, colonizing local microsites washed bare by flood waters that infrequently flash down this ephemeral stream.

237. Lining Dry Branch- Scenes of the remarkable (and beautiful) range afforded by a climax mixed hardwood (sugarberry-pecan-cedar elm) forest along a dry branch leading into Bosque River. The first of these three photographs provided a more distant view of the forest range vegetation in general. The second photograph gave a perspective looking down Dry Branch toward Bosque River. This slide featured a large black walnut (foremost trunk) and a pecan (lighter barked tree behind and to left of the walnut). A sapling of boxelder was to immediate left of this pecan. The third photograph gave an upstream view of Dry Branch with a huge forked trunk pecan (right background) with a pole-sized boxelder to its left. The herbaceous layer shown in all three slides was dominated by broadleaf woodoats with the annual composite, giant ragweed (an r-selected species), the associate herbaceous species and the major forb along banks of Dry Branch and Bosque River.

Tree species above Dry Branch included sugarberry, cedar and American elms, pecan, red mulberry, bois d'arc or Osage orange, and eastern cottonwood. Shrubs ranged from southern black haw to poison oak or poison ivy to woody climbers like trumpet creeper and mustang grape. Broadleaf woodoats and Canada wildrye mace up most of the herbaceous layer.

Erath County, Texas. Early November (autumnal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash) and/or SAF 94 (Sycamore-Sweetgum-American Elm). Absence of sycamore and sweetgum and presence of sugarberry suggested that SAF 93 was more appropraite; however, the former cover type designation for SAF 94 included pecan so choice was unclear. Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

238. Death before frost- In a large plant of Virginia frostweed or crownbeard (Verbesina virginica) this monarch butterfly (Danaus plexippus) was observed during the annual autumn migration of these butterflies to their wintering habitat in Mexico. Frostweed is a dominant, native forb (though only a minor or, at best, local associate species) in the riverbottom forest of sugarberry, cedr elm, and pecan. The monarch was not in a typical resting position and upon closer examination it was discovered...

Bank of Bosque River, Erath County, Texas.

239. Didn't quite make it to Mexico- Adult, female monarch butterfly captured and killed by a Carolina praying mantis (Stagmomantis carolina) in a large, blooming Virginia crownbeard or frostweed. This drama took place in a bottomland forest of sugarberry, cedar elm, and pecan as the monarch was migrating to her winter home in Mexico. Like el bandito crossing the Brazos at Waco (Billy Walker) this monarch died in a river bottom at the hands of her enemy.

Predation or the predator-prey relationship is one of the important interactions among species. Predation is part and parcel of range ecosystems. Predation takes place along all trophic levels of secondary productivity food chains except the first level or herbivory (unless one interpretes herbivory as a form of predation).

Bank of Bosque River, Erath County, Texas.

240. Chitinous prey and predator- A Carolina praying mantis feeding on a female monarch butterfly that made the fatal mistke of stopping to rest in a large Viriginia frostweed enrout to her winter home in Mexico. This mantid is a native, insectivorous predator that is both beneficial and bane to man as it kills some insects that benefit humans and other insect species that are harmful to Homo sapiens. Either way, the Carolina mantis is a native animal and one that is part of natural range ecoxystems as well as those more modified by human action.

Praying mantis or mantid species are members of the order Orthoptera (family, Mantidae) along with such phytophagous (plant-eating) insects as crickets and grasshoppers as well as cockroaches that are general feeders.

Bank of Bosque River, Erath County, Texas.

241. Corpse: well-used or wasted?- A Carolina mantis killed and fed on this monarch butterfly that was migrating to its wintering ground in Mexico. The praying mantis ate only parts of the head (actually it could not be determined if the head was eaten or if it fell off without being eaten), thorax, and abdomen. Legs and wings of the butterfly were left still attached to the exoskeleton of thorax and abdomen. Is this a typicl feeding pattern or are wings sometimes eaten?

Bank of Bosque River, Erath County, Texas.

Directions to a related range type- A wet (essentially subirrigated throughout much of the year) prairie or prairie savanna dominated by bottomland switchgrass (Panicum virgatum) and Maximillian sunflower (Helianthus maximillianii) with narrowleaf cattail (Typha domingensis), heath aster (Aster ericoides), and pink boneset (Eupatorium incarnatum) as associate herbs and with black willow (Salix nigra) as a woody component had developed immediately above the mixed hardwood forest of Dry Branch, Bosque River. Treatment of this wetland range vegetation was included in a section entitled, "Wetland Odd-Lots" in the chapter, Meadows and Related Marshes, under the heading,Grasslands.

Winter- Bosque River

242. Straw on the floor- Standing dead culms from the previous growing season of Canada wildrye, mostly, and broadleaf woodoats, secondly, in understorey of a climax sugarberry-cedar elm-pecan bottomland forest range. The trees in this photograph were a massive cedar elm (left midground), adult sugarberry (center background), and bois d'arc (right midground). An on-goining Severe Drought (Palmer Index) had hampered phenological development of cool-season species, both annual and perennial.

Erath County, Texas. Early March (late winter; hibernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash) and/or SAF 94 (Sycamore-Sweetgum-American Elm). Absence of sycamore and sweetgum and presence of sugarberry suggested that SAF 93 was more appropraite; however, the former cover type designation for SAF 94 included pecan so choice was unclear. Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

243. Winter in the winter of their lives- Local grove of trees in mixed hardwood forest that developed on the floodplain of Bosque River. Species and age class composition showed successional pattern of climax community composition. Mature pecans (right foreground and mid-background) along with dead pecan (large tree left foreground with peeled bark) and mature, though middle-aged, cedar elm (left margin, foreground) with saplings of sugarberry coming in as the ultimate dominant species of the climax foresst range. The shrub with dark, knobby bark to immediate right of sugarberry sapling was rusty black-haw (blackhaw) which was a locally dominant understorey woody species and one characteristic of the climax range vegetation. Oother major shrubs were trumpet creeper and common green-briar. Specimens of both of these species were growing on the cedar elm at far left.

An herbacaeous component--except for an occasional plant of broadleaf woodlats--was largely absent in this more densely shaded microsite. The land was in death grip of Sevee Drought (as measured on Palmer Drought Severity Index).

Inside the bottom- Interior structure and composition of amixed hardwood(sugarberry-cedar elm-pecan; tri-dominant) bottomland forest on the floodplain of a small river in the Western Cross Timbers of northcentral Texas. These two slides presented the same forest scene (eg. same trees, woody vines) from two slightly different camera angles. The large tree in foreground (left margin and left-center foreground of first and second slide, respectively) was an old-growth (maturity apporaching senexcence) specimen of black walnut with mustange grape climbing it from the right side of its trunk. This black walnut was surrounded (left, right and behind) by smaller (and, more than likely, youngr) by sugarberry, the number one dominant of this variant of a forest range subtype. Fiddleleaf or saw green-brier grew everywhere.

The herbaceous understorey, which in this late winter aspect contained both last year's dead shoots and the current year's immature, green (living) shoots, was made up overwhelmingly by three climax, cool-season, perennial grasses:1) Canada or nodding wildrye, 2) broadleaf woodoats, and 3) hairy woodland brome (in that relative order of density, cover, abundance). These three species are decreasers in this forest range site. There were some forb species in this foresst range ecosystem, but none of these was visible in these two views.

Floodplain, Bosque River, Erath County, Texas. Early March; hibernal aspect. FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash) and/or SAF 94 (Sycamore-Sweetgum-American Elm). Absence of sycamore and sweetgum and presence of sugarberry suggested that SAF 93 was more appropraite; however, the former cover type designation for SAF 94 included pecan so choice was unclear. Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

Inside with the lead actor- Two views of a sugarberry-cedar elm-pecan bottomland forest that developed along a small river in the Western Cross Timbers of northcentral Texas in the late winter season. These two "photo-plots' showed interior composition and structure of this forest range that was at the climax stage of successional development. All tree trunks in thee two slides were of sugarberry, the "number one" (of three) dominant tree species of this potential natural vegetation. Most of these tree trunks (most were large pole size) were commensal hosts to various species of liana (woody vines) including mustang grape, trumpet creeper, poison ivy, and. of course, the everywhere, ever-present fiddleleaf or sqw green-brier, the dominant shrub of this forest community. A limb of black walnut was barely present (an Epson Perfection 700 scanner cut off most of it that was more competely captureds in a 35mm slide) in upper right margin of the first slide.

The well-developed understorey consisted of three native, cool-season, perennial grasses (three decreasers): 1) Canada or nodding wildrye, 2) broadleaf woodoats, and 3) hairy woodland brome (in that relative order of composition based on foliar cover and plant/shoot density). Both last year's dead shoots and this year's immature live shoots of these three grass species made almost all of the herbaceous layer of this climax forest range vegetation. None of the few forb species that were infrequently present in this herbaceous understorey were in the forest vegetation shown here.

Herbage in these two views of the climax forest range had been produced during drought (Severe Drought during most of previous growing season and Extreme Drought so far in current growing season; Palmer Drought Index).

Two other climax woody members - In a sugarberry-cedar elm-pecan bottomland forest on floodplain of a small river in northcentral Texas this old-growth forest tree form of American elm was playing host to the climbing woody shoots of trumpet creeper. The dead herbage in the immediate foreground was mostly that of Canada wildrye with a few shoots of broadleaf woodoats. Most of the younger trees in the background were sugarberry, the ultimate single most important dominant of this climax forest. Logs in midground were the half-rotten trunk and massive limbs of a huge fallen pecan that died naturally at a "ripe old age". Forest vegetation seen here was during Extreme Drought (Palmer Scale).

American elm is a associate of this potential natural forest vegetation. Cedar elm is the number two tri-dominant (behind sugarberry and befgore pecan), and while American elm is only an associate tree species in this climax forest community, both elm species are defining species and warrent inclusion in the title (s) of the Society of American Foresters (Eyre, 1980) forest cover.

At late winter during bad drought- Interior structure and composition of a climax sugarberry-cedar elm-pecan bottomland forest featuring a fire-scared American elm with a small sapling sugarberry to its left. Large trees in distant background at right were cedar elms while trees to left background of the American elm wre sugarberry. The well-developed and productive understorey of this climax forest vegetation was comprised almost entirely of three decreaser grass species: 1) Canada wildrye, 2) broadleaf woodoats, and 3) hairy woodland brome (in that relative order of cover and herbage yield). These three climax grasses are native, perennial, cool-season species although woodoats defies an easy categorization with regard to major season of growth. Herbage of these three grasses included both last season's dead shoots and current growing season's production. This amount of grass growth and production had occurred during Extreme Drought (Palmer Index).

Transition Winter to Spring (Late Winter/Early Spring Stage)-Bosque River

244. Starting to spring forth- A young American elm with late flowers and young fruit (samara) stood out conspicuously (at far left) among pecan (largest trees), sugarberry, and cedar elm which are the climax dominants of this mixed hardwood bottomland forest range. The herbaceous understorey visible on the grass-covered bank was almost exclusively Canada or nodding wildrye and broadleaf woodoats. The strikely bent tree was a pecan.

Erath County, Texas. Late February (late winter; hibernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash) and/or SAF 94 (Sycamore-Sweetgum-American Elm). Absence of sycamore and sweetgum and presence of sugarberry suggested that SAF 93 was more appropraite; however, the former cover type designation for SAF 94 included pecan so choice was unclear. Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

245. When this associate is most obvious- American elm in full fruit (foremost tree in center midground and branches in foreground) on Bosque River floodplain. Flowering and fruit-bearing of American or white elm is one of the earliest harbingers of spring in this mixed hardwood bottomland forest. The green grass was Canada wildrye with some broadleaf woodoats as local associate grass species. In background the largest trees were pecan; smaller trees were sugarberry and cedar elm. American elm is a local associate to the other three tree species in the climax forest.

246. The class of climax in the forest school- Graphic (by standards of vegetational dynamics) example of plant succession in a mixed hardwood floodplain forest playing out along Bosque River. Three young sugarberry (large pole size) coming up beside (under the canopy) of a young adult pecan provided this picturesque example of the dominant climax tree species slowly taking dominance from pecan. Pecan is a pioneer or colonizer species that becomes established quickly on bared mineral soil and then through long life span persist into the climax bottomland forest as a secondary dominant to sugarberry and cedar elm (often with American elm as a climax associate). Eastern cottonwood and sycamore exhibit this same role and pattern of persistence, but only cottonwood is native to this particular forest site and forest cover (dominance) subtype, the sugarberry-cedar elm-pecan bottomland forest.

Green grass was primarily Canada or nodding wildrye and, secondarily, broadleaf woodoats.

247. Growing green in winter- Top-down view of several (at least five) cespitose plants of hairy woodland brome (first slide) and oblique view of a cespitose plant of Canada wildrye (left) and of hairy woodland brome (right) (second slide) in the understorey of a sugarberry-cedar elm-pecan bottomland forest that developed on the floodplain of Bosque River in the West Cross Timbers of northcentral Texas. Both Canada wildrye and hairy woodland brome are native, cool-season, perennial, festucoid (Fescudoideae subfamily) grasses. In the comparatively mild winters of northcentral Texas, plants of these two festucoid species typically develop and maintain lush, rich-green shoots (primarily leaves) throughout winter and complete their annual growth cycle by mid-spring. Some of last year's spent leaves and culms were visible in these two slides, especially the second photograph.

Canada wildrye was the dominant herbaceous species in this mixed hardwood floodplain forest while hairy woodland brome was primarily a major associate species. In local areas (spatial parcels of range larger than microsites and smaller than range sites) hairy woodland bromegrass was the dominant and, sometimes, co-dominant with Canada wildrye or broadleaf woodlats. Overall, broadleaf woodoats was second in general abundance, cover, shoot density, etc. to Canada wildrye with which it was sometimes a co-dominant.

The following photographs were taken on the official vernal equinox (the first day of spring). A drought classified as Severe on the Palmer Drought Severity Index had existed in this locality (Bosque River bottoms, Erath County, Texas) from late autumn or early winter through to time of these photographs. Growth (both phenological development and biomass production) had been retarded, delayed, hampered, etc. to such an extent that they were several weeks "behind schedule". In reality phenological development and herbage production of cool-season species were probably at least a month later than the average progression of these biological events.As such, range vegetation and plant growth/development presented in this section were representative of a typical winter even though the calendar showed beginning of spring.

248. Winding down- Winter's end in a southcentral floodplain forest mixed hardwoods with a well-developed herbaceous understorey only two days before the vernal equinox. Eat your hearts out Minnesotans. Part of a relict stand of climax sugarberry-cedr elm-pecan forest with a well-developed herbaceous layer. Forest vegetation featured here (and in the two subsequent slides) was a grove of old-growth pecan in which most tree recruitment was sugarberry and cedar elm. Pecan were large to immense size adults with largest rrees showing signs of senscence. Smaller trees in left midground and a large majority of those in background were sugarberry, the tree species that appeared to be in process of succeeding to the foremost dominant.

The shrub-sized tree with green leaves was a cedar elm which is typically the first of the three dominant tree species to leaf out or bear fruit (American and slippery or red elm flower and bear fruit prior to leaf formation). The green herbaceous growth was a mixture of Canada wildrye and broadleaf woodoats, the dominants of the herbaceous layer of the understorey. Phenological development of these two perennial, cool-season, C3 grasses wass substantially delayed by Severe Drought (Palmer Drought Severity Index).

Tree species composition was shown and described in the two immediately succeeding two photographs, except that the smaller trees in the left background were not included in these next two slides. Those trees (to left of downed limbs with dried leaves) were sugarberry. Abundance of broken branches littering the forest floor were discussed in the immediately succeeding caption.

Erath County, Texas. Mid-March (late winter; hibernal aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash) and/or SAF 94 (Sycamore-Sweetgum-American Elm). Absence of sycamore and sweetgum and presence of sugarberry suggested that SAF 93 was more appropraite; however, the former cover type designation for SAF 94 included pecan so choice was unclear. Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

249. The Old guard and the New (or a brace of photographs worth a thousand words)- Mid-range and close-in views (first and second photograph, respectively) of the climax vegetation of a mixed hardwoods (sugarberry-cedar elm-pecan) bottomland forest shown in the immediately precding photograph. That slide and the two shown here provided a zooming-in sequence of "photoplots" of this range plant community. These two photographs centered on a grove of old-growth pecan with an understorey of Canada wildrye and broadleaf woodoats (the hrbaceous dominants) with some nimblewill (the associate grass species), pigeon berry, and frostweed. All of these major herbaceous species were dormant except the cool-season dominants. There were also several plants of common bedstraw or cleavers (Gallium aparine) and an unidentified crudifer species (foreground of second photograph).

These two "photoplots" showed the "relay floristics" aspects of species replacement in in final stages (subclimax to climax) of the classic Clementsian model of plant succession. Sugarberry was succeeding (replacing) pecan as the number one dominant tree species. The foremost tree (pole-size bole) in left foreground and the smaller pole to immediate left of the huge pecan were sugarberry. There was considerably less tree recruitment (establishment of new trees) of cedar elm than of sugarberry and least of all for pecan (though there were seedlings of the three dominant tree species). These two slides gave the "picture worth a thousand words".

Moreover, sugarberry and cedar elm were young sub-adult trees whereas pecan were of the life-cycle stages ranging from fully-grown, mature adults to "over-ripe" or post-mature (senescent) to dead or almost dead trees. Viewers should note, for emphasis, the downed and rotting large limbs in these two photographs as well as in the immediately preceding photograph. Some of these downed limbs and branches were due to wind-pruning, but most reflected the senescent (aging and dying) state of the crowns of these old trees. In fact, most of the branches were broken off by wind were already dead and rotting in place or hollow to one degree or the other.

In this bottomland forest pecan persisted into the climax vegetation as large trees that were mature or senescing while recruitment was highest for sugarberry and cedar elm. In this tract of pristine forest sugarberry and cedar elm were the ultimate (climax) dominants while pecan was a persistent member from earlier stages and probably as a subclimax species. Again, however, limited reproduction (mostly sexual) of pecan appeared ample for maintenance of this species (regardless of successional status) in the climax forest. Burns and Honkala (1990) categorized pecan as subclimax. Wenger (1984) and Burns and Honkala (1990) regarded pecan as Intolerant on the tolerance scale. In their first description of North American forest cover types the Society of American Foresters (Eyre, 1954, ps. 30-31) recognized Sycamore-Pecan-American Elm (Type 94) with the three listed species plus boxelder being the dominants with green ash and sugarberry (hackberry) as associates. In Eyre (1954, p. 30) there was also a Sugarberry-American Elm-Green Ash (Type 93) with associate species including cedar elm and pecan. Pecan was in both of these types as either a dominant or associate species. In the revision of North American forest cover types (Eyre, 1980, ps. 65-66) sweetgum replaced pecan in Type 94. For the bottomland forest featured here the earlier type descriptions (Eyre, 1954, ps. 30-31) was more accurate.

A Severe Drought (according to Palmer Drought Severity Index) throughout late autumn and throughout winter had delayed phenologicl advance and biomass production of herbaceous species.

250. Species composition seen in winter- Range vegetation of a mixed hardwood bottomland forest that had developed on the west bank of Dry Branch, an ephemeral stream channel draining into Bosque River in the West Cross Timbers of northcentral Texas. This range plant community was described in considerable detail above, especially under the autumn subsection. The two photographs presented here were overall or summary views with the second slide being farther downstream along Dry Branch from the land shown in the first photograph. Specifically, the three large trees (one black walnut and two cedar elm) in right background of first photograph were the "center of attraction" in the second photograph in which the largest and darkest trunk was black walnut flanked by two cedar elm. The tree behind these three in the second slide was an immense pecan with a remarkable straight trunk. The shrub with green leaves to immediate left of this huge pecan was a southern or rusty blackhaw which ws featured in the immediately succeeding two-slide set. In the first photograph there were four red mulberry trees (the three foremost trees and a small pole between these three and the black walnut and flaniking cedar elm) at different distances on the west bank of Dry Branch.

The herbaceous layer was a "three-way mixture of Canada wildrye, braodleaf woodoats, and hairy woodland brome. The first two were the overall dominant herbaceous species throughout this forest range. In this specific and restricted local habitat hairy wood brome was a third dominant. The dead shoots with curled leaves (eg. lower left corner of first photograph) were plants of broadleaf woodoats. The huge tree in the background of the first of these photographs (on the east bank of Dry Branch and slightly inclined over it) was a pecan. The second (farther back) tree in the first slide that was conspicuously leaning over Dry Branch and the tree (at closer camera range) in the second slide was another black walnut. This author did not find any seedlings, saplings, or poles of black walnut which strongly suggested that the large, mature trees of this species had established at an earlier stage of plant succession and persisted into the climax forest as long-lived seral (perhaps, subclimax) species. On the tolerance scale black walnut was classified as Intolerant (Wenger, 1984; Burns and Honkala, 1990) the same as pecan. On this forest range there was some recruitment of pecan.

251. Early bloomer amongst the oldtimers- A single plant of rusty or southern black-haw (Virburnum rufidulum) at base of a massive pecan on west bank of Dry Branch, an ephemeral stream flowing into Bosque River. These two photographs were "closer-to-it" views of this locale introduced in the background of the immediately preceding photograph. Trunk to immediate right of large pecan was a cedar elm as was trunk of dead tree behind it and pecan. The woody vine on the big pecan was mustang grape (Vitis mustangensis). Common green-briar was also growing up and into the pecan. Most herbaceous frowth was Canada wildrye and broadleaf woodoats with some hairy wood brome. The large leaning (over Dry Branch) tree in left background was a black walnut.

Rusty black-haw was in full bloom (photographs of this plant were presented below with other woody range plants of this climax sugarberry-cedar elm-pecan forest that developed on the Bosque River floodplain.

252. Starting a new tree; restarting old grass- A seedling of sugarberry beside a burnt stick and by a plant of broadleaf woodoats with young leaves on the floor of a climax sugarberry-cedar elm-pecan forest on the Bosque River floodplain in West Cross Timbers. Canada wildrye, the other co-dominant was also present. In this local spot there was more woodoats than wildrye. All three of these species were dominants of this rforest range. Sugarberry was first among the three dominant tree species and these two, native, cool-season bunchgrasses were co-dominants of the herbaceous layer.

This "photoplot" illustrated the concept of microsite (= microenvironment or microclimate). The burnt stick--evidence of not-too-distant past fire on this foresst range--provided some shelter and imporved microhabitat (shade, windbreak, source of moisture release and nutrients) that might have been just enough of an advantage that permitted establishment of this sugarberry seedling. Decaying tree leaves and twigs, grass roots, and other sources of organic matter also furnished nutrients and increased available water to aid in survival (up to now) and, perhaps, eventual growth to maturity of the adult sugarberry.

Erath County, Texas. Mid-March (late winter; hibernal aspect).

253. What is to come- This was perhaps the ultimate "picture worth a thousand words" among this set of photographs. At base of a giant, old-growth pecan was a two-year-old seedling of cedar elm. This baby tree with it glaring new leaves presented a dramatic snapshot of the future forest composition as to dominant forest trees. Cedar elm not pecan was going to be the dominant of this stand unless some drastic disturbance (eg. a hot fire) intervened. Other photographs in this section (eg. the immediately preceding slide) showed regeneration of sugarberry with that species also replacing, to a large degree, pecan which was dominating the forest canopy of this grove at this point in plant succession (advanced stage of vegetation development on the sere).

It was concluded that for this range type, a form or variant of forest cover type SAF 93 (Sugarberry-American Elm-Green Ash) and/or SAF 94 (Sycamore-Sweetgum-American Elm), pecan was the third dominant as a long-lived subclimax tree species that persisted into the climax whereas sugarberry and cedar elm were more adapted to shade and general competition so as to have greater regeneration at climax stage. It was explained in the two immediately following photograph-caption sets that reproduction (primarily sexual reproduction) was substantially greater for sugarberry and cedar elm than pecan, although the latter did have adequate reproduction to maintain its presence as different age classes in the climax forest vegetation.

Erath County, Texas. Mid-March (late winter; hibernal aspect).

254. Heirs to the climax throne: new plants of the dominant trees- A seedling each of sugarberry (left) and cedar elm (right) on the floor of a climax sugarberry-cedar elm-pecan bottomland forest. Pecan is probably best regarded as a subclimax species that persist into the climax vegetation while sugarberry and cedar elm are the more prominent or predominant trees (ie. "true" co-dominants) at end of the sere. The basis (the "litmus test") of climax species is that they replace themselves and are not succeeded by other species on the sere. Climax species are those that reproduce "in their own shade" so to speak and do not "improve" or further the environment of the sere for other more successionally advance species. Climax plant species are the most successionally advanced. They are the species "left standing" at termination of the sere. Sustained self-reproduction is the only criterion for climax. .

Sugarberry and cedar elm are those tree speceis with far greater recruitment (regeneration or reproduction) than pecan even though pecan appeared to be reproducing at rates adequate to maintain (pretty much) its presence in the climax forest community. Pecan regeneration was proportionally so much less than that of sugarberry and cedar that these latter two species were increasing their "controlling share" of canopy cover while pecan barely "held its own". This photograph (along with the next successive slide) showed that successional pattern most poignantly.

Erath County, Texas. Mid-March (late winter; hibernal aspect).

255. The new regime in its infancy- Sexual reproduction of sugarberry and cedar elm, the two major dominant tree speciesof a mixed hardwood floodplain forest with a sporadic shrub and well-defined herbaceous layer. Several seedlings (at least eight) of cedar elm and two seedlings of sugarberry on the leaf litter strewn floor of a climax sugarberry-cedar elm-pecan bottomland forest. It was explained in the immediately preceding caption that there was abundant regeneration of these two species whereas there was much less reproduction of pecan. No pecan seedlings were growing in this "photoplot".

Sugarberry was regarded as Tolerant while pecan was classified as Intolerant on the forest tolerance groupings ((Wenger, 1984; Burns and Honkala, 1990). Burns and Honkala (1990) concluded that there was not sufficient research to definitively categorize cedar elm, but evidence suggested the rating of Intermediate. Reproduction of these three tree species on this bottomland forest generally was in line with these categories. The area of forest floor presented in this photograph was a more sunlite local microenvironment which could explain the greater number of cedar elm seedlings in contrast to fewer of those of sugarberry.

Herbaceous growth in this plot was almost exclusively Canada wildrye and broadleaf woodoats.

Erath County, Texas. Mid-March (late winter; hibernal aspect).

256. Floodplain floor- Floor of a floodplain forest with a canopy dominatd by sugarberry, cedar elm, and pecan and with one (sometimes two) lower woody layers and an herbaceous layer. The co-dominant herbaceous species were Canada wildrye and broadleaf woodoats, but a local dominant (and the overall associate) herbaceous species was hairy woodland brome (Bromus pubescens= B. purgans). These two slides (a set of "nested photo-plot") featured hairy woodland brome as the local dominant (a micro-habitat consociation). There were four or five separate plants (genotypes) in the second slide.

These plants were growing in a Severe to entry level Extreme Drought (Palmer Index).

Floodplain, Bosque River, Erath County, Texas. Early March; winter growth stage of cool-season perennial grass species. FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash) and/or SAF 94 (Sycamore-Sweetgum-American Elm). Absence of sycamore and sweetgum and presence of sugarberry suggested that SAF 93 was more appropraite; however, the former cover type designation for SAF 94 included pecan so choice was unclear. Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

257. Leaf-in-leaf; pioneer and climax dominants- Recently emerged seedlings of giant ragweed growing beside new-shoot growth of Canada wildrye and broadleaf woodoats on a recent, water-scoured disturbance on the upper bank of Bosque river. The two perennial, cool-season grasses are overall dominants of the herbcaceous understorey of the sugarberry-cedar elm-pecan climax bottomland forest that developed on the floodplain of this river. The annual, warm-season composite is a local dominant on disturbed microsites noitable those created by flood waters of the usually slow-moving river. Also in the first of these two "photoquadrants" were some plants of the naturalized, Eurasian species, annual ryegrass (Lolium multiflorum), which thrives with some disturbance. Annual ryegrass was not found with the native perennial grasses (ie. the annual cannot compete with already established perennials) nexcept where disturbances like flood water created local denudation (exposed bare soil).

The dead, partially rotted stalks in the second "photoquadrant" were last year's shoots of giant ragweed. This "new ground" had been made by flood waters in the winter and spirng of the preceding year and plant succession was, one year later, still hospitable to annuals of native and naturalized species. Last year's floods were not catastrophic enough to wash away the long established climax dominant grasses, Canada wildrye and broadleaf woodoats.

Erath County, Texas. Mid-March (late winter; hibernal aspect); seedling stage of giant ragweed, late winter growth of perennial grasses.

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