Miscellaneous Forest Types

Eastern or Southern Live Oak

Live oak forests, woodlands and savannas in southeastern North America- There are various species (and lower taxa) of Quercus across North America that retain their leaves more-or-less yearlong (or through the cool-season) so as to be described as "evergreen" or "live" oaks. Evergreen or live oaks of one species or another range from the Atlantic Coast at Virginia southward, then along the Gulf Coast, and sporadically westward to the Coast Range of California and Oregon. The evergreen oak of eastern (southeastern) and Gulf coastal North American is the eastern live oak, Virginia live oak, or, simply, live oak (Quercus virginiana). Various forms and varieties of Q. virginiana have been recognized in the region of the Eastern Deciduous Forest of North America. The most distinctive of these varieties of the Eastern Deciduous Forest is Q. virginiana var. maritima, dwarf live oak, which often exist more as a shrub and thus forms shrubland or scrub rather than forest. The live oak of eastern North America growing farther west along the Gulf Coast, and often extending inland for considerable distance, has had (still has) a confusing if not arbitrary taxonomic treatment. Traditionally the live oak species ranging from Virginia and the Carolinas westward to eastern and central Texas and northward to localized pockets in Oklahoma and Arkansas was regarded as Q. virginiana including the more western and northward live oaks designated as Q. virginiana var. fusiformis (Muller, 1951). More recent taxonomists in Texas (Correll and Johnston, 1979; Diggs et al, 1999) elevated Q. virginiana var fusiformis to the species status as Q. fusiformis accompanied by such common names as plateau live oak, escarpment live oak, scrub live oak, and west Texas live oak. Q. fusiformis (= Q. virginiana var. fusiformis) varies in size from shrubs to large trees and in morphology from arboreus (true tree) to scrub form. Typically this taxon spreads by rhizomes and/or root sprouts (often these asexual reproductive organs are referred to as root stocks). Diggs et al. (1999, p.716) stated that it was possible that Q. fusiformis was ... "only a more xeric and cold tolerant subsepecies of the more widespreat Q. virginiana". That has been the traditional interpretation of this taxon, and it was the one followed herein. Forest and range ecologists have been much more reluctant to distinguis between Q. virginiana and Q. fusiformis than have taxonomists. The following treatment of live oak (either the classic mesic, eastern Q. virginiana or the xeric [less mesic], western Q. virginiana var. fusiformis) dealt with the tree form of this species either as woodland (= open forest) made up of individual trees (genetic individuals; different genotypes) or as smaller mottes (Texan usage for "grove") in which many to most tree trunks were shoots of the same tree. In the latter case single or distinct tree trunks are individual or separate shoots of the same genetic plant originating from a common rootstock derived from one acorn (single trunks are modules of the same tree distinguished as the same genotype or genetic individual). It is nigh on to impossible to determine if live oak trunks in a motte or closely spaced live oaks in a larger forest or woodland community are several different trees (unique genotypes; each tree from an embryo) or different tree trunks of the same tree (tree trunks from the same embryo; all shoots derived from one acorn). Usually there are some of both sources, but it is generally accepted by ecologists and plant physiologists that most trunks in a live oak motte are of the same tree meaning that a live oak motte is a clonal plant as in the case of quaking aspen (Populus tremuloides). Also, it must be assumed that trees and shrubs in mottes and many of those in the larger woodland tracts have root-grafted so that the local community does have some features of the Clementsian "super organism". The term woodland rather than forest was used for live oak-dominated range communities because this designation seemed more appropriate (a more apt description) for the more open canopy and, im most instances, only slightly interlocking crowns of mature trees coupled with well-developed herbaceous and shrub-small tree layers in understories. Woodland more than forest seemed to the author to connote more clearly this idea, but at the same time it was possible that the converse was true and that forest was the more relevant or accurate descriptive term. The designation of forest was used by Brown et al. (1998. p. 41) for the Floridian Evergreen (Hammock) Forest biotic community (see below). The Live Oak forest cover (89) of the Society of American Foresters (Eyre, 1980) was described as bottomland forest, but the description for this type clearly included a denser and more species-diverse plant community than the more open vegetational architecture of the Florida hammock and western Gulf of Mexico live oak motte and larger woody communities dominated by live oak.

1. Live oak hammock (hammock is Indian for "shady place")- This is a unique land form and range vegetation type restricted to the southeastern portion of North America, especially Florida. Hammocks are characterized by being fairly level to gently rolling and higher than surrounding land. Yet they are poorly drained and fertile (soils are high in humus) with hardwood communities that are typically dominated by live oak canopy and an understory of cabbage palm (Sabal palmetto), skunk cabbage (Symplocarpus foetidus), carpetgrass (Axonopus affinis), and panicgrasses (Panicum spp.). FRES No. 16 (Oak-Gum-Cypress Ecosystem). There is no Kuchler equivalent for communities this small; the Kuchler system lumped this type in with some other larger unit of which K-81 (Live Oak-Sea Oats) seemed most fitting. SRM 817 (Oak Hammocks). Mixed Hardwood Series 124.1j1 of Floridian Evergreen (Hammock) Forest biotic community 124,1j of Brown et al. (1998). Southern Coastal Plain- Central Florida Ridges and Uplands Ecoregion, 75c (Griffith et al., undated).

2. Interior of a Florida live oak hammock with tell-tale raised yet wet ground—Mid-canopy shrubs include wax myrtle (Myrica cerifera) and American beautyberry (Callicarpa americana) with sparsely scattered saw palmetto (Serenoa repens).FRES No. 16 (Oak-Gum-Cypress Ecosystem). Subunit of K-81 (LiveOak-Sea Oats). SRM 817 Oak Hammocks). Mixed Hardwood Series 124.1j1 in Floridian Evergreen (Hammock) Forest biotic community 124.1j of Brown et al. (1998). Southern Coastal Plain- Central Florida Ridges and Uplands Ecoregion, 75c (Griffith et al., undated).

3. A woodland on the coastal sand prairie- A live oak woodland had developed on the Kenedy sand prairie (Johnston, 1963, p. 460), one form or subtype of Gulf coastal tallgrass prairie inland from the coast but within the Texas Coastal Prairies and Marshes Area. Seen from perspective of Landscape Ecology this range vegetation (these units of hardwood evergreen woodland), ecosystems, and/or landscape elements could be interpreted as patches of live oak woodland in a matrix of seacoast bluestem (Andropogon littoralis= Schizachyrium scoparium var. littorale)- sacahuiste or Gulf cordgrass (Spartina spartae) coastal prairie. From the classical viewpoint of savanna as put forth by Dyksterhuis (1957) this overall grassland-woodland landscape could be seen as a savanna in which units or assemblies of woody vegetation ranging from smaller mottes (Texan for "groves") to extensive "woods" of several hundred acres are the physiogonomic equivalent of vegetation consisting of individual to "a few" trees and/or shrubs isolated on grassland. In other words, this would be a savannah in which the woody elements are larger and include more woody plants (eg. groves) rather than the typical situation of scattered individual trees and/or shrubs. This would not necessarily imply that such a structural or physiogonomic savannah was a successional or genetic (as to origin) savanna, this latter of which is the usual definition or, at least, connotation of an ecotone (a transition) from herbaceous to woody vegetation. Interpretation of the woody vegetational units perhaps would hinge on whether mottes or woodlands were actually one up to "just a few" genetic plants whose individual stems (trunks or boles) were repeating clonal units or, alternatively, if these groves or woodlands were composed of many genetic individuals (each or most trees as denoted by a single tree trunks were derived from one embryo, that is, one acorn). Range vegetation shown here was part of what has long been known as the Wild Horse Desert part of the Rio Grande Plains. This native grazing land is a slightly rolling or hummock aeolian plain of sand entitled the "Kenedy loose sand prairie" that consist of different range plant communities as "a tight mosaic of vegetation types..." (Johnston, 1963, p. 460). The uncertain successional status of large live oak mottes like the one shown here that develop on loose sand uplands was mentioned briefly in the caption of the immediately succeeding slide. The present photograph illustrated the spatial arrangement of sandy sachuiste prairie and mottes dominated by live oak that attest to the "very complex mosaic of vegetation types" (Johnston, 1963, p. 460) of the Wild Horse Desert on the Rio Grande Plains. Norias Division, King Ranch, Kenedy County, Texas. February, late hibernal or early vernal aspect.

4. Live oak motte turned woodland- Exterior view of a live oak woodland or forest (larger than typical live oak motte) situated within seacoast-sacahuista tallgrass coastal prairie. The dominant herbaceous plant growing at perimeter of live oak woodland was Gulf cordgrass or sacahuista. Other tree species present--at rare to trace amounts of canopy cover--werehoney mesquite (Prosopis glandulosa) and common hackberry or sugarberry (Celtis laevigata). Even though crowns of trees produced a fairly closed canopy with considerable interlocking of branches there was a well-developed (e) herbaceous understorey (often consisting of two to three layers) as well as a second (lower) woody layer or understorey of shrubs and small (immature) trees. Areal extent (acreage) of this live oak-dominated range community was considerably larger than typical live oak mottes and therefore was viewed more as a live oak woodland or forest rather than as a grove of trees within or on a prairie. Perhaps this distinction was arbitrary or even incorrect but, as in the case for fire behavior, at some point size (spatial scale) becomes an ecologically critical feature. Impacts and role of fire would be a case in point. Prairie fires could easily burn under or scorch crowns of small mottes whereas with expansive woodland areas there would places where fire could not reach (ie. as size of live oak-dominated stands increase in area there is increased probability that larger proportions of stands will be unaffected by fire). The situation would be similar for dispersal of plant propagules from outside live oak-dominated stands as well as for penetration of light from edges (vs. through canopy) of live oak stands. Successional status of mottes on coastal (and sone iinterior tallgrass and mixed) prairies apparently has not been completely established. Johnston (1963) described, quantified, and discussed changes in range vegetation in the much of the area of the Rio Grande Plains, including that part known as the Wild Horse Desert. Based on personal accounts, including that of a longtime ranchman on the Norias Division of King Ranch, Johnston (1963, p. 464) concluded that live oak had so increased on the sandy seacoast bluestem-sacahuiste coastal prairie that previously separate live oak mottes had coalesced and become interconnected. Causes of the shift from more herbaceous to woody vegetation included the "usual suspects" of overgrazing, seed dispersion by livestock, and reduction or cessation of fire were proposed for the increased in woody vegetation, including that of live oak mottes. Notwithstanding brush invasion due to influences of white man, live oak mottes (of much smaller acreage) are native, potential natural, or climax range vegetation. The range vegetation of both live oak mottes and surrounding Gulf cordgrass or sacahuista sand prairie were samples of two climax (= potential natural) plant communities. Live oak mottes had expanded into range that was formerly "Kenedy loose sand prairie" (Johnston, 1963, p. 460) so that the overall vegetation of this natural grazing ground had departed from climax conditions of the virgin range. Yet species composition, structure, etc. of the two range communities was representative of that of each climax community. A description and discussion of the general or overall range vegetation over much of the Rio Grande Plains was provided by Fulbright in Shiflet (1994) as SRM 719 (Mesquite-Live Oak-Seacoast Bluestem). The Society for Range Management did not offer a separate designation or treatment of live oak motte as a distinct range type. Likewise, the Live Oak forest cover type of the Society of American Foresters (Eyre, 1980) did not include the live oak motte form of the Gulf Coast prairies. Norias Division, King Ranch, Kenedy County, Texas. February, later hibernal or early vernal aspect. FRES No. 16 (Oak-Gum-Cypress Forest and Woodland Ecosystem). Variant of K- 81 (Live Oak-Sea Oats), Variant of SAF 89 (Live Oak). Alternatively and based on geographic location: FRES No. 32 (Texas Savanna), K-55 (Mesquite-Live Oak Savanna). Live Oak variant of SRM 719 (Mesquite-Live Oak-Seacoast Bluestem). Western Gulf Coastal Plain- Coastal Sand Plain Ecoregion 34d (Griffith et al, 2004).

5. Into the woods we go- Inside a live oak motte with a well-developed (and heavily utilized) herbaceous understorey and a shrub/immature tree layer. Almost all tree trunks where those of live oak (clonal trunks or those of individual genotypes was not determined), but there were a few honey mesquite and sugarberry trees (at least nine out of ten trunks were live oak). There were immature trees of three of these species. Most common (abundant) shrubs were Hercules club that also "goes by" tickle-tongue, pricklyash, pepperbark, and toothache tree (Zanthoxylum clava-herculis), lime pricklyash or colima (Z. fagara), and mustang grape (Vitis mustangensis). The herbaceous layer was made up almost exclusively of numerous grasses including: Gulf cordgrass or sacahuista, seacoast bluestem, tumble windmill grass (Chloris verticillata), and hooded windmillgrass (C. cucullata), red lovegrass (Eragrostis secundiflora ssp. oxylepis),Gulf dune paspalum (Paspalum monostachyum), Canada wildrye (Elymus canadensis), white tridens (Tridens albescens), Texas tridens (T. texanus), redtop panicgrass or thatchgrass (Panicum rigidulum), and Ghiesbreght panicgrass (P. ghiesbreghtii) as well as naturalized King Ranch bluestem (Andropogon ischaemum= Bothriochlor ischaemum) and Guineagrass (Panicum maximum). There were no obvious or conspicuous forbs other than an Aster sp ("go figure"). Norias Division, King Ranch, Kenedy County, Texas. February, late hibernal or early vernal aspect. FRES No. 16 (Oak-Gum-Cypress Woodland and Forest Ecosystem). Variant of K-81 (Live Oak-Sea Oats). Variant of SAF 89 (Live Oak). Alternatively and based on geographic location: FRES No. 32 (Texas Savanna), K-55 (Mesquite-Live Oak Savanna). Live Oak variant of SRM 719 (Mesquite-Live Oak-Seacoast Bluestem). Western Gulf Coastal Plain-Coastal Sand Plain Ecoregion 34d (Griffith et al., 2004).

6. Inside the live oaks- Another view of the interior of an upland (loose sand) live oak woodland or an extensive motte dominated by live oak. Tree cover and density was almost exclusively live oak (of both mature and immature tree trunks), but there were occasional honey mesquite, sugarberry or common hackberry, and least abuncant huisache (Acacia farnesiana= A. smallii). These three species were also of both mature and immature ages. Most abundant shrubs were Hercules-club or tickle-tongue (all leafless short shrubs in foreground), lime pricklyash, and mustang grape. Grasses included the dominants, Gulf cordgrass or sacahuiste and seacoast bluestem plus tumble windmillgrass, hooded windmillgrass, red lovegrass, Gulf dune paspalum, white tridens, Texas tridens, Canada wildrye, redtop pancigrass or thatchgrass, and Ghiesbreght panicgrass plus the naturalized King Ranch bluestem and Guineagrass. Strangely, no forbs other than a species of Aster were obvious. Norias Division, King Ranch, Kenedy County, Texas. February, late hibernal or early venal aspect. FRES No. 16 (Oak-Gum-Cypress Forest and Woodland Ecosytem). Variant of K-81 (Live Oak-Sea Oats). Variant of SAF 89 (Live Oak). Alternatively and based on geographic location: FRES No. 32 (Texas Savanna), K-55 (Mesquite-Live Oak Savanna). Live Oak variant of SRM 719 (Mesquite- Live Oak-Seacoast Bluestem). Western Gulf Coastal Plain- Coastal Sand Plain Ecorgeion 34d (Griffith et al., 2004).

7. King Ranch? Ya gotta be kiddin'!- Interior of live oak woodland that developed on a surrounding Western Gulf Coast tallgrass prairie. Morphology of trees, including features of branches and crowns, and general architecture (structure and arrangement of vegetation) of a live oak motte of such area (relative spatial scale) and development as to be a woodland. Trees were exclusively live oak except for occasional (rare) mesquite, Texas hackberry or sugarberry, and, least of all, huisache. There was a prominent herbaceous layer composed of coastal prairie grasses and a woody understorey of shrubs like Hercules club, lime pricklyash, and mustang grape. Grass species included isolated individuals of the dominants, Gulf cordgrass and seacoast bluestem, as well as common and hooded windmillgrasses, Gulf dune paspalum, red lovegrass, white and Texas tridens, Canada wildrye, redtop panicgrass or thatchgrass, and Ghiesbreght panicgrass, plus naturalized King Ranch bluestem and Guineagrass. A quick search for forbs was fruitless. The woodland range shown here does not purport with images of ranches in the popular imagination or perception or as shown in Hollywood Westerns, but there are numerous such live oak mottes and woodlands on large ranches in south Texas,especially those along the Gulf Coast. This scene was on the Norias Division of King Ranch. Live oak woodland range like this furnishes outstanding habitat for the Rio Grande turkey (Meleagris gallopavo intermedia) as well as providing shade for cattle and horses like the renowed Santa Gertrudis and sorrel Quarter Horses of the famous King Ranch. Kenedy County, Texas. February, later hibernal or early vernal aspect. FRES No. (Oak-Gum-Cypress Woodland and Forest Ecosystem). Variant of K-89 (Live Oak). Variant of SAF 89 (Live Oak). Alternatively and based on geographic location: FRES No. 32 (Texas Savanna), K-55 (Mesquite-Live Oak Savanna). Live Oak variant of SRM 719 (Mesquite-Live Oak-Seacoast Bluestem). Western Gulf Coastal Plain-Coastal Sand Plain Ecoregion 34d (Griffith et al, 2004).

8. Crowding an ole patriarch- An old-growth speciment live oak with a unnaturally high (man-caused increase in) cover of understorey trees and shrubs. The well-developed lower tree and shrub layer was clearly a recent (hence almost assuredly anthropogenic) woody invasion. Great spreading limbs of the "big momma" or "gran'paw" live oak were undeniable proof that this ancient tree developed in the open (ie. in absence of crowding). The small-sized (relatively young) understorey trees and shrubs attested to recent establishment of these individuals suggesting absence of fire (at least low fire frequency), perhaps combined with overgrazing or periodic overuse that reduced grass cover. Most of the leafed-out shrubs were honey mesquite (eg. larger shrubs at far-right foreground with trunks inclined to the right) but there were also some huisache. Texas sugarberry or common hackberry was also present though largely dormant. The multi-stemmed shrub with smooth, light-grey bark was lime pricklyash with an accompaning and readily identified mustang grape with its serpentine single-stem covered by dark, deeply furrowed bark. These shrubs were the subject of the next photograph. Norias Division, King Ranch, Kenedy County, Texas. February, later hibernal or early vernal aspect. FRES No.16 (Oak-Gum-Cypress). K-89 (Live Oak). Variant of SAF 89 (Live Oak). Alternatively and based on geographic location: FRES No. 32 (Texas Savanna), K-55 (Mesquite-Live Oak Savanna). Live Oak variant of SRM 719 (Mesquite-Live Oak-Seacoast Bluestem). Western Gulf Coastal Plain-Coastal Sand Plain Ecoregion 34d (Griffith et al., 2004).

9. Prickly and twisted invaders- Close-up of lime pricklyash and mustang grape that had invaded the undersrtorey of a live oak motte on the sandy plains of the Wild Horse Desert in Rio Grande Plains. Age of these shrubs was not determined, but they were relatively young having not yet grown to height of first horizonal limb of an old live oak under the canopy of which these plants were growing. (This was presented clearly in the immediately preceding photograph.) Successional status of these invading shrubs was unclear, but their relative youth in comparison to the obvious age of the large live oak indicated that lime pricklyash, mustang grape (as well as mesquite and huisache) had established after the live oak had reached adult size. The terms invasion, invaders, and invading shrubs could have one or two meanings in this usage, only one of which was certain. The certain meaning or usage of invasion referred to movement of plants from one area into another and subsequent successful establishment of these new plants in their environment. This is the classic original meaning of invasion in the Clementsian usage (Weaver and Clements, 1938, ps. 131-132144, 148, 166) and, hence, as used by most foresters and rangemen (at least those of the Anglo-American school of "dynamic plant ecology"). From this original meaning a star student of Weaver, E.J. Dyksterhuis adapted invasion and, especially, invader to a second meaning (an extension or outgrowth of the original meaning of Clements). This second usage was applied to plant species that continued to invade (Clementsian meaning) on habitats that had been disturbed (denuded was Clements' term) or that were continuing to be disturbed, especially by overgrazing (Dyksterhuis, 1949). As applied to this live oak woodland Gulf cordgrass has to re-establish itself on (ie. to invade) a previously denuded (as by overgrazing, plowing, drought) range. Gulf cordgrass has to invade (ie. complete or effect an invasion) but because this species declines with disturbances (specifically abusive or improper grazing) it is a decreaser not an invader in usage as applied to plant succession and range condition/trend (Dyksterhuis, 1949). Decreaser corresponds to member of the climax plant community. On the live oak-dominated range type/site featured here lime pricklyash, mesquite, and huisache also invaded when they successfully established. Up to this point invasion has the same meaning as successful establishment of a decreaser such as Gulf cordgrass, but because these particular woody range plants have continuing or on-going invasion under disturbances such as overgrazing or, as is also likely, underburning these shrub species are invaders in contrast to decreasers like Gulf cordgrass. The prevailing judgment of rangemen with regard to such abundant establishment (invasion) of these woody species on the live oak woodland and sacahuista sand prairie range types is that this unnatural excessive invasion is symptomatic of disturbance (perhaps cessation of fire, improper grazing management, or drought) so that these shrubs are invaders in the meaning of Dyksterhuis, 1949). This is an invasion of brush, of woody invaders as per the Dykstehuis (1949) model and therefore noxious range plants. This brush invasion is different from the general (the Clementsian) invasion. Invasion by Dyksterhuis (1949) invaders is range deterioration through the process of retrogression (retrograde movement from climax vegetation). Continuing invasion by species like lime pricklyash, honey mesquite, huisache, and, probably, mustang grape resulting in development of a lower woody understorey constituted brush invasion (changes in the plant community by retrogression) and commensurate loss of economically valuable and ecologically natural forage. The real question is, "How much of the cover of live oak is brush invasion"? Or same thing with reversed emphasis, "How much of the uniting (the coalescing) of live oak mottes into extensive live oak woodland is climax (potential natural) vegetation"? Norias Division, King Ranch, Kenedy County, Texas. February.

10. Ranchman's touch- Naturalized Guineagrass (Panicum maximum) formed the complete understorey of a motte of mature live oak. On this large live oak motte or woodland on the King Ranch, the understorey was such an exclusive single species- stand of Guineagrass that woody species as well as other grasses were absent. This isolated tract was subject only to infrequent defoliation. There are various "versions" of how the introduced Gunieagrass (native of Central and South America) naturalized and spread throughout the Coastal Prairies and Marshes and eastern parts of the Rio Grande Plains. One of the more popular explanations is that Guineagrass was spread by Hurricane Beluah from seed nurseries and experimental plantings at branch stations of the Texas Agricultural Experiment Station. Others insist that Gunieagrass spread from some of the larger ranches in the region, especially King Ranch, that had planted it as the latest "miracle grass". Cypher (1995, ps. 77, 92, 94, 107, 183) described planting of Guineagrass on King Ranch under direction of Bob Kleberg. However most of these plantings and established pasturess of Guineagrass described by Cypher (1995) were King Ranch holdings in Cuba and South America. As late as the early 1970s Guineagrass was probably not established in any part of Texas (Gould, 1975, p. 469). This introduced forage grass is a "new comer" compared to Johnsongrass, bermudagrass, King Ranch bluestem, and buffelgrass. Norias Division, King Ranch, Kenedy County, Texas. February, later hibernal or early vernal aspect. Understorey so modified that FRES, Kuchler, SAF, and SRM designations would be meaningless. Western Gulf Coastal Plain-Coastal Sand Plain Ecoregion 34d (Griffith et al., 2004).

11. Leader of the western taxon of eastern or southern live oak (Quercus virginiana var. fusiformis= Q. fusiformis)- Catkins and new leaves have emerged on this live oak on the Grand Prairie of northcentral Texas. Shedding of last year's leaves and emergence of this year's leaves typically occurs synchronously causing some lay observers to state that "the new leaves are phushing off the old leaves". "Leaving out" usually also occurs with flowering. It's a busy (and critical) time in the life of a live oak. Erath County, Texas. March, immediate pre-anthesis to anthesis stage.

12. Flushing in eastern or southern live oak- The synchronous emergence of new leaves (from preformed buds of previous season) and shedding of old or current year's leaves in trees (such as the plateau form of southern live oak shown here) is known as flushing. These two views of a leader of southern live oak illustrated this phenological phenomenon. Erath County, Texas. March.

13. Off with the old on with the new- Tip of branch in plateau form of southern live oak on which current season's or old leaves (large leaves of dark green color) persisted even as new leaves emerge accompanied by catkins. This phenomenon is designated as flushing. Erath County, Texas. March, early catkin (pre-anthesis) stage of phenology.

14. A live oak's life in the fall- Summer-hardened leaves provided a backdrop for this southern live oak acorn. Southern (eastern) live oak is a white oak (Leucobalanus subgenus), the species of which produce an annual (vs. a biennial) acorn crop. Acorns are a rich and major carbonaceous (energy) concentrate on numerous ranges and range types. Acorns are often extremely important components of the diets of many species of range animals ranging from insects through birds and mammals, the latter often including man (American Indians especially relied heavily on various kinds of acorns.). Excessive intake of acorns (as well as buds, catkins, and leaders) can result in toxicity. Older leaves are not toxic. The poisonous principle is a group of tannins known as gallotannins. Energy stores of acorns generally offset adverse impacts of poisoning for wildlife such as upland game birds, especially wild turkey and white-tailed deer (Odocoileus virginianus). Grand Prairie, Erath County, Texas. October, fruit-ripe.

Various Other Southern Forest Types

15 .Two distinct plant communities comprising locally restricted vegetation types: 1) a swamp of water oak with bald cypress as an associate and 2) a maidencane (Panicum hemitomon) marsh designated as a lowlands range site. These two types together constitute a flat woods pond. FRES No. 16 (Oak-Gum-Cypress Ecosystem) and corresponding K-101 (Southern Flood-Plain Forest) and FRES No.41 (Wet Grasslands Ecosystem) with with no Kuchler units small enough to pick up the maidencane type. Maidencane would be included with Kuchler-83 (Everglades) in Florida. The maidencane marsh type is SRM 819. Mixed Hardwood Series in Southeastern Swamp and Riparian Forest biotic community and Maidencane Series (if and when such is recognized) Series in Southeastern Interior Marshland biotic community, respectively, of Brown et al. (1998). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

16. Edge of two wetland range communities- Boundary between the water oak-bald cypress swamp and maidencane swamp introduced in the preceding slide. The swamp portion of this flatwoods pond was FRES No. 16 (Oak-Gum-Cypress Ecosystem) and corresponding K-101 (Southern Flood-Plain Forest) while maidencane marsh was FRES No.41 (Wet Grasslands Ecosystem) with with no Kuchler units small enough for this region so that instead maidencane would be included with Kuchler-83 (Everglades) in Florida. Maidencane rangeland cover type was SRM 819 (Freshwater Marsh and Ponds). Swamp with water oak dominant and bald cypress the associate species comprised a combination or "hybrid" of SAF ted

17. Maidencane in the spring- Hardin County, Texas. May, late vernal aspect. Maidencane Series (if and when such is recognized) in Southeastern Marshland biotic community, respectively, of Brown et al. (1998). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

18. Bay-Gall Bog or Titi in the Texas Big Thicket- This vegetation is the most impenetrable “jungle” or “tangle’ in the Big Thicket. The local mound-and-intermound relief creates a bog ecosystem. The soil series of the mound microrelief (on the mound) has the spodosol soil series Babco. This is currently the only spodosol mapped in Texas. The dominant plants are red bay (Persea borbonia) and sweet bay or swamp bay (Magnolia virginiana) among the hardwood trees and shortleaf and loblolly pine from the conifers. Gall, swamp cyrilla or, by the Indian name, titi (Cyrilla raecmiflora) is the dominant species of the shrub layer along with gallberry (Ilex coriacea; not to be  confused with the preceding gall), bull-briar (Smilax bona-nox), saw-brier (S. glauca), buttonbush (Cephalanthus occidentalis), and wax myrtle (Myrica cerifera) dominate the shrub layer. Completing this “tangle” is the herbaceous understory often dominated by rather rank-growing ferns.

The largest trunk (in center) is a loblolly pine, the trunk immediately behind and to the right of it is a water oak, the two trees immediately behind and to the right of the water oak are sweet bay magnolias, and the left foreground tree is a red bay. Most of the shrubs in the foreground understory are swamp cyrilla or titi. Hardin County, Texas. May. There is no specific FRES or Kuchler for this local community that grows within the FRES No. 13 (Loblolly Pine-Shortleaf Pine forest Ecosystem). Mixed Hardwood Series in Southeastern Swamp anbd Riparian Forest biotic community of Brown et al. (1998). South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

19. Interior of a Texas Big Thicket Bay-Gall Bog- Detail of the shrub layer described in the preceding slide caption. Note the seedling or young tree stage of loblolly pine in the foreground and the adult loblolly pines in background indicating that this is the dominant conifer for this unique local community. Hardin County, Texas. May.

20. The floor or herbaceous layer of a Bay-Gall Bog dominated by ferns. Over 20 species of ferns are native to the Big Thicket and there are another four or five species that may have naturalized here. The ferns are growing on a mound of Babco soil. Hardin County, Texas. May.

21. Profile of Babco soil (the only spodosol mapped in Texas)- Spodosols comprise the  soil order characterized by having  a light gray eluvial horizon over a reddish aluminum- and/or iorn-enriched horizon. They typically occur in humid areas.  The Babco pH ranges from 3.1 to 3.6. Hardin County, Texas. May.

22. Sandjack= bluejack oak (Quercus incana)-sandhill bluestem scrub type— The bluestem is a taxonomic complex of little bluestem, including the taxa often shown as Andropogon divergens or Schizachyrium scoparium var. divergens, and slender bluestem (Andropogon tener= Schizachyriumtenerum). A few post oaks are associates of bluejack oak. Composites and various prickly pears (Opunia spp.) are scattered throughout the bunchgrass sward. An aeolian ("blowsand") ridge community. Beech Creek Unit, Big Thicket National Preserve, Hardin County, Texas. May. FRES No. 14 (Oak-Hickory Forest Ecosystem). A variant of K-72 (Oak Savanna). One of the many forms of Southern Scrub Oak, a variant of SAF 72 (Southern Scrub Oak). A Scrub Oak Series of Brown et al. (1998), but one was not shown for this region. Sandy upland variant of South Central Plains- Flatwoods Ecoregion, 35f (Griffith et al., 2004).

Sundry Bottomland Forest Types

23 .Bottomland (floodplain) gallery oak-hickory forest— a "finger" of the eastern deciduous forest projects into the climatic or regional climax tallgrass paririe here in the Cherokee Prairie in the Osage Plains division of the Central Lowlands physiographic province. This gallery forest community is classified by the Missouri Natural Areas Committee (1987) as wet-mesic bottomland forest. It is dominated by pin oak (Quercus palustris) represented here by the largest tree with the light-colored trunk (center). Associated species also visible include: western hackberry (Celtis occidentalis), black cherry (Prunus serotina), persimmon (Diospyros virginiana), bois d'arc, red mulberry (Morus rubra), and silver maple (Acer saccharinum). Dominant shrub is Missouri gooseberry (Ribes missouriense). Herb layer is absent. Missouri State Prairie Park, Barton County, Missouri. June, early estival aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). One riparian form or part of K-73 (Mosaic of Bluestem Prairie [K-66] and Oak-Hickory Forest [K-91]). Variant of SAF 65 (Pin Oak-Sweetgum). Central Irregular Plains- Cherokee Plains Ecoregion, 40d (Chapman et al., 2002).

24. Bottomland forest of western hackberry, sugarberry (Celtis laevigata), and American elm (Ulmus americana) with an understory almost exclusively of broad-leaf wood oats (Uniola latifolia= Chasmanthium latifolium) with scattered solitary plants of the shade-tolerant composite, elephant's foot (Elepohantopus carolinianus). Beautiful plant community and an example of permanent deciduous forest range in contrast to the typical transitory feature of these dense hardwood forest types. Ottawa County, Oklahoma. September, late estival aspect. No obvious FRES or Kuchler designations as this is a smaller unit. Neither was this relatively unique community described by Braun (1950) though it corresponds to general creek and river bottom vegetation in the Ozark Mountains. SAF Cover Type 93 (Sugarberry-American Elm-Green Ash). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al., 2005).

Pecan (Carya illinoinenisis)

Pecan has been regarded as the largest of North American hickories. This species frequently forms extensive bottonland forests along major rivers (floodplains in general). Such floodplain forests usually hve an understorey that is either herbaceous or a combination of herbaceous-woody layers that is amenable to proper--usually, light--grazing, commonly as permanent forest range on a sustained yield basis. Some examples of these pecan bottom ranges were presented below.

25. Bottomland forest (actually more of a woodland physiogonomy as crowns overlap but slightly) of pecan (Carya illinoinenisis) with an understory devoid of shrub layers and with the herb layer dominated by Canada wildrye with frostweed (Verbesina virginica) an important associate. Indiangrass and eastern gamagrass add a tallgrass element. In the upper two post oaks (left center) entered this otherwise single-species stand or pecan consociation. In the second photograph two large pecans (only their lower trunks showing) shaded a carpet of the two cool-season perennial native grasses, Canada wildrye Texas wintergrass, while warm-season perennials like little bluestem, Indiangrass, and perennial dropseeds "waited in the wings" for their time on the shaded stage. This tree-dominated vegetation was an irregular gallery forest along the South Llano River. The pecan is the stately State Tree of Texas and the community shown here is known as "pecan bottoms" by locals who frequent it come nut season. Another beautiful permanent deciduous forest range type. Kimble County, Texas. June, estival aspect. An "island" or isolated part of FRES No. 15 (Oak-Hickory Forest Ecosystem) and K-91 (Oak-Hickory Forest). No SAF designation readily fit this range vegetation. This forest range appeared similar to the Cottonwood and Live oak Bottomland Types (SAF 63 and SAF 89), but most likely a Variant SAF 94 (Sycamore-Sweetgum-American Elm) that had the former SAF designation of Sycamore-Pecan-American Elm. Edwards Plateau- Edwards Plateau Woodland Ecoregion, 30a (Griffith eat al., 2004).

26. Texas-size- Though it was hard to show with all the shade, several widely scattered giant pecans formed a pecan bottom much like those that existed in Texas river bottoms before European man presumptuously assumed that he could improve things. For big trees like our "centerfold hero" here stocking rate has be relatively low. Prof. Benton Storey (Texas A&M Universty) felt that production of big pecan trees and high nut yields were best obtained with only one tree per acre. Pecan bottoms would thus be savannahs or, at most, woodlands because their crowns would not contact each other at such stocking. Studded T steel post serving as stakes for pecan seedlings and saplings and the top of one post by the pecan-picker's "centerfold" provided scale for size comparison. This part of the understorey of this river bottom woodland had been mowed (perhaps in anticipation of nut season). Kimble County, Texas. June, estival aspect. An "island" or isolated part of FRES No. 15 (Oak-Hickory Forest Ecosystem) and K-91 (Oak-Hickory Forest). No SAF designation readily fit this range vegetation. This forest range appeared similar to the Cottonwood and Live oak Bottomland Types (SAF 63 and SAF 89), but most likely a Variant SAF 94 (Sycamore-Sweetgum-American Elm) that had the former SAF designation of Sycamore-Pecan-American Elm. Edwards Plateau- Edwards Plateau Woodland Ecoregion, 30a (Griffith eat al., 2004).

27. East Texas pecan bottoms- Bottomland mixed forest in the Texas Pineywoods dominated by pecan with hackberry and water oak as associates. Foremost tree at left margin was a large water oak with a single bole of high-quality timber. The large tree at right-of-center with numerous major limbs and scaley, brown bark was pecan as were most of the trees (pole-size) in background. The smallest of three trunks in foreground (in front of and aligned along left edge of pecan was sugarberry (Celtis laevigata). Flowering dogwood (center green shrub) formed an interupted upper shrub layer. The invasive alien Japanese honeysuckle (Lonicera japonica), seen here as the green plant in left to center foreground, comprised a spotty though dense lower shrub layer. The exotic and now widely naturalized Japanese honeysuckle (a horticultural escape "gone wild") is a major noxious plant problem in forest understories, especially in pecan bottoms. There were only a few grasses (panicgrasses were most obvious), grasslike plants (limited to Carex spp.), or forbs in the understorey of this bottomland forest. The alien L. janpoica appeared to have crowded out herbaceous plants.Species of green briar and grape along with rattan vine or Alabama supplejacck formed a "jungle" of vines among the smaller pecans in bacdground. Local mowing at edge of this forest near a campground had apparently effectively controlled woody vines in this sample of lowland Pineywoods forest. Davy Crockett National Forest, Houston County, Texas. March, vernal aspect (post dogwood blooming stage ). FRES No. 15 (Oak-Hickory Forest Ecosystem). K-91 (Oak-Hickory Forest). Variant SAF 94 (Sycamore-Sweetgum-American Elm) that had former SAF designation of Sycamore-Pecan-American Elm. Closest biotic community designation of Brown et al. (1998, ps. 37, 38) was Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1, but there should have been a Oak-Hickory Series for Southeastern Deciduous and Evergreen Forest 123.. South Central Plains- Southern Tertiary Uplands Ecoregion, 35e (Griffith et al., 2004).

28. Another example of a pecan bottom- This example of bottomland forest dominated by pecan was along the small slow-moving Alarm Creek on extremely fertile alluvium. The four big trunks in right foreground and center midground were pecan. Other trees included both post and blackjack oaks. Hackberry was also well-distributed. Greenbriar was the dominant shrub. There was a well-developd herbaceous layer of Canada wildrye, little bluestem, and sand lovergrass (Eragrostis trichoides). Small isolated patches of giant cane (Arundinaria gigantea) grew along banks of the creek. Erath County, Texas. April. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-75 (Cross Timbers). Variant SAF 94 (Sycamore-Sweetgum-American Elm) that had former SAF designation of Sycamore-Pecan-American Elm. (Griffith et al., 2004). Closest biotic community designation of Brown et al. (1998, ps. 37, 38) was Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1, or more descriptively perhaps, a transition of this Oak-Hickory Series and Bluestem "Tall-grass" Series, 142.11, Plains Grassland 142.1 Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

29. Young pecans and tallgrasses in summer- Part of the floodplain of the Bosque River in the Western Cross Timbers supported this stand of "half-growed" young pecans with bur oaks of the approximate same age cohort as associate tree species. Understorey was locally dominated by Canada wildrye, especially prominent in the second of these photographs where that cool-season member of the Hordeae or Tritaceae tribe was taller than the top wire of the fence enclosing this nice sample of "pecan bottoms". The associate herbaceous species varied locally from such species as the native and-should-have-been-one-of-the- climax -dominants Indiangrass to naturalized Johnsongrass. Other grasses included big, little, and silver bluestems; Texas wintergrass, several perennial dropseeds, sideoats grama (Bouteloua curtipendula) tumble windmillgrass (Chloris verticillata), and tumblegrass (Schedonnardus paniculatus). Forbs were varied but, as to be expected, composites "ruled" with prominent and common species including western ragweed (Ambrosia psilostachya), Baldwin ironweed (Vernonia baldwinii), iand frostweed or white crownbeard (Verbesina virginica). The major shrub was common greenbriar or catbriar (Smilax rotundifolia). Other shrubs were trumpet creeper (Campsis radicans) and poison oak. Hamilton County, Texas. June. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-75 (Cross Timbers). Variant SAF 94 (Sycamore-Sweetgum-American Elm) that had former SAF designation of Sycamore-Pecan-American Elm. (Griffith et al., 2004). Closest biotic community designation of Brown et al. (1998, ps. 37, 38) was Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1, or more descriptively perhaps, a transition of this Oak-Hickory Series and Bluestem "Tall-grass" Series, 142.11, Plains Grassland 142.1 Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

30. Pecan timber- The forest form of pecan bottoms was represented by this stand of midddle-aged trees on the first terrace of a small creek floodplain in the patchwork of range plant communities where there is an intermixing of Western Cross Timbers and Grand Prairie in northcentral Texas. All trees were pecan whose tall straight trunks supported such woody vines as mustang grape, fiddleleaf or fringed greenbriar, and Alabama supplejack. Dominant understorey shrub was Carolina buckthorn. Most woody ground cover was the invasive, alien Japanese honeysuckle (Lonicera japonica), a treadful, horrid, etc. (adjectives fall short of vile description) woody weed. Major grasses included Canada wildrye, purpletop, Johnsongrass, and Texas wintergrass in that order. Major forbs were Baldwin ironweed (Vernonia baldwinii), and Indian plantain (Cacalia plantaginea). Erath County, Texas. April, early to mid-vernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-75 (Cross Timbers). Variant SAF 94 (Sycamore-Sweetgum-American Elm) that had former SAF designation of Sycamore-Pecan-American Elm. (Griffith et al., 2004). Closest biotic community designation of Brown et al. (1998, ps. 37, 38) was Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1, or more descriptively perhaps, a transition of this Oak-Hickory Series and Bluestem "Tall-grass" Series, 142.11, Plains Grassland 142.1 Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

31. Bottomland pecan forest- Higher stocking of pecan trees on bottomland produces a forest in contrast to the savanna oar open woodland form of this range type as shown above. All trees in these two photographs were pecan. Dense stocking resulted in tall, relatively straight boles in marked contradistinction from the widely spreading crowns of pecans with more spacing among trees. Pecans with stocking rates represented by forest vegetation presented in these two photographs produce wood and not fruit. This is the forest community of a lumberman not an orchardman, and the form of vegetation that develops in absence of fire. There was a well-developed woody vine layer that extended from ground level to tops of forest canopy which was comprised of mustang grape, rattanvine or Alabama supplejack, and fiddleleaf greenbriar. Carolina buckthorn grew as a smaller tree or larger shrub in the lower woody layer. The herbaceous layers included tallgrass species like Canada wildrye, purpletop, and the naturalized Johnsongrass along with composite forbs the two most common of which were Baldwin ironweed and Indian plantain. Much of the lower layer was composed of the invsive (and apparantely naturalized) Japanese honeysuckle. Erath County, Texas. April, early to mid-vernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-75 (Cross Timbers). Variant SAF 94 (Sycamore-Sweetgum-American Elm) that had former SAF designation of Sycamore-Pecan-American Elm. (Griffith et al., 2004). Closest biotic community designation of Brown et al. (1998, ps. 37, 38) was Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1, or more descriptively perhaps, a transition of this Oak-Hickory Series and Bluestem "Tall-grass" Series, 142.11, Plains Grassland 142.1 Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

32. Woody understorey of pecan bottom- From ground level and extending upward just shy of tree branching the range vegetation of a bottomland pecan forest was comprised of three woody vine species (mustang grape, fiddleleaf greenbriar, and Alabama supplejack) and the shrub or small tree, Carolina buckthorn. The wsidespread exotic invader, Japanese honeysuckle, was absent from this photoplot. The herbaceous portion of this forest vegetation visible in the background was presented in the next photograph. Erath County, Texas. April, early to mid-vernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-75 (Cross Timbers). Variant SAF 94 (Sycamore-Sweetgum-American Elm) that had former SAF designation of Sycamore-Pecan-American Elm. (Griffith et al., 2004). Closest biotic community designation of Brown et al. (1998, ps. 37, 38) was Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1, or more descriptively perhaps, a transition of this Oak-Hickory Series and Bluestem "Tall-grass" Series, 142.11, Plains Grassland 142.1 Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

33. Herbaceous layer (s) of a bottomland pecan forest- In addition to the woody vines of mustang grape, fiddleleaf greenbrier, and Alabama subblejack or rattanvine in right foreground and background of this photoquadrant important forbs of the herbaceous component included Indian plantain (left corner of foreground) and scattered smaller, apparently shade-stunted Baldwin ironweed. Many of the small green shoots with forb-like leaves were new sprouts of greenbrier and supplejack. Erath County, Texas. April, early to mid-vernal aspect. FRES No. 15 (Oak-Hickory Forest Ecosystem). K-75 (Cross Timbers). Variant SAF 94 (Sycamore-Sweetgum-American Elm) that had former SAF designation of Sycamore-Pecan-American Elm. (Griffith et al., 2004). Closest biotic community designation of Brown et al. (1998, ps. 37, 38) was Oak-Hickory Series 122.11 of Northeastern Deciduous Forest 122.1, or more descriptively perhaps, a transition of this Oak-Hickory Series and Bluestem "Tall-grass" Series, 142.11, Plains Grassland 142.1 Cross Timbers-Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

Eastern Cottonwood (Populus deltoides)

Eastern cottonwood is a fast-growing, deciduous species (Salicaceae) capable of maturing into a large tree. In fact, it has one of the most rapid growth rates of North American hardwoods having been described by Cooper and Haverbeke (in Burns and Honkala, 1990, vol. 2, p.530) as "... the fastest-growing commercial forest species in North America". This species has historically been subdivided into the two taxonomic varieties of eastern cottonwood (P. deltoides var deltoides) and plains cottonwood (P. deltoides var. occidentalis) for more central and western forms, respectively (Fowells, 1965, ps. 514- 522; Burns and Honkala, 1990, vol. 2, ps.530-543). In addition, various other workers re-evaluated the taxonomy of the tree or, sometimes, shrub known as Rio Grande cottonwood as P. deltoides ssp. wislizenii (Powell, 1988, p. 87-88). This taxon was formerly interpreted as a variety of Fremont cottonwood (P. fremontii var. wislizenii) or, in the "bible" of Texas flora (Correll and Johnston, 1979, p. 455), as a separate species (P. wislizenii). Along watercourses Rio Grande cottonwood grows into large trees. Eastern cottonwood is usually the dominant--sometimes, the exclusive (or nearly so)--tree species of bottomland forests. Many of these forests have a grazable understorey so as to constitute forest range. However, grazing must be carefully managed or the highly palatable foliage of young cottonwoods will be so heavily defoliated, even by range animals that are more grazers than browsers (eg. beef cattle), that this tree species will be greatly reduced or even eliminated to point of eventual loss of the forest. In the past, many eastern cottonwood-dominated forests were destroyed by confining livestock on them in what were in essence dry lots. At one time this was especially common for swine operations (feeder pig and/or market hog) where mature cottonwood trees were used as improvised or natural shade and the former forestland was simply a barnyard or pig sty. Examples of properly managed, lightly grazed cottonwood forests were presented below, along with an example of overbrowsing by beef cattle.

34. Floodplain forest of plains cottonwood in the old bed of the Republican River- Seen here is a wetland forest site and a wetland range site (background and foreground, respectively). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). Cottonwood form of K-89 (Northern Flood-Plain Forest).The forest site is an eastern cottonwood consociation (SAF Forest Cover Type 63, Cottonwood) with an herbaceous understory of Indiangrass. Populus deltoides Association (if and when recognized), Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). The range site also has Indiangrass as the dominant herb, but Canada wild rye and prairie sandreed are important associated species. October, autumnal aspect. Dundy County, Nebraska. Western High Plains- Flat to Rolling Cropland Ecoregion, 25d (Chapman et al., 2001).

35. Interior of a floodplain or wetland forest on the former bed of the Republican River- View inside the plains cottonwood forest of the preceding slide, but showing local importance of scattered green ash (Fraxinus pennsylvanica). This vegetation is about 60 years old having begun development when the river flooded and changed course. The climax tree species were already established at time of that disturbance, but this perturbation removed most of the woody plants except the larger cottonwoods and ash allowing migration and establishment of prairie grasses as well as sedges and rushes more typical of the numerous marshes in the area. Dundy County, Nebraska. October, autumnal aspect. FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem).Cottonwood form of K-89 (Northern Flood-Plain Forest). SAF 63 (Cottonwood). Populus deltoides Association (if and when recognized), Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). October, autumnal aspect. Western High Plains- Flat to Rolling Cropland Ecoregion, 25d (Chapman et al., 2001).

36. Semiarid gallery forest- Exterior view of a riparian zone forest of Rio Grande cottonwood (Populus deltoides ssp. wislizenii), netleaf hackberry (Celtis reticulata), Mexican walnut (Juglans major), and lanceleaf sumac (Rhus lanceolata) At outer (least mesic) edge of this gallery forest some of the more moisture-loving grasses formed a thick sward. The large cespitose grass was Wright's sacaton (Sporobolus wrightii) the dominant understorey species; Canada wildrye was the major associate species. The most common forb was horehound (Marrubium vulgare), a naturalized Eurasian weed. This range vegetation was in the Trans-Pecos Basin and Range vegetational area, the west Texas portion of the Chihuhuan Desert Region. It was in Chihuhuan Deserts 24 (ecoregion level III), Chihuhuan Montane Woodlands 24d (ecoregion level IV) based on Griffith et al. (2004). It was an example of mesic forest in a semiarid climate. FRES No 17 (Elm-Ash-Cottonwood Forest Ecosystem). No aqppropriate Kuchler unit for southern and.or southwestern cottonwood gallery forest. SAF 63 (Cottonwood). Populus deltoides Association (if and when such is recognized), Cottonwoow-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Trans-Pecos Basin & Range Vegetational Area: Chihuhuan Deserts- Chihuhuan Montane Woodlands Ecoregion 24d (Griffith et al., 2004). June, summer aspect. Calamity Creek, Jeff Davis County, Texas.

37. Great Plains floodplain forest- Plains cottonwood and various species of willow formed a species-rich forest complete with prolific regeneration and a prominent herbaceous understorey. In these two slides the dominant herbaceous plant at local level ws American licorice (Glycyrrhiza lepidota), but grasses, sedges (Carex spp.), spike rush (Elocharis spp.) rushes (Juncus spp), and cattails (Typha domingensis) were the major understorey plants throughout most of the vegetation. Canada wildrye and bottomland switchgrass were most common with Johnsongrass (Sorghum halepense) the associate. Even the annual Eurasian grass, cheatgrass (Bromus tectorum) was common locally. FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). There is no Kuchler unit describing southern or southwestern cottonwood-dominated riparain forests. SAF 63 (Cottonwood). Populus deltoides Association (if and when such is recognized), Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Rolling Red Plains Vegetational Area: Central Great Plains- Red Prairie Ecoregion 27h (Griffith et al., 2004). June, summer aspect. Flood plain, North Fork of Red River. Wheeler County, Texas.

38. Wild or American licorice (Glycyrrhiza lepidota)- Wild licorice is one of the more common legumes along watercourses throughout the Great Plains. It occurs in both the riparian zone and at outer (drier) margins of rivers, creeks, and floodplains. July, full bloom. Custer State Park, Custer County, South Dakota.

39. Healthy regeneration of plains cottonwood and willow- One of the major problems in structure and sustainability of riparian and floodplain forests is reproduction under concentrated livestock grazing. This is especially so in arid and semiarid regions where livestock, especially cattle, tend to "hang in on water". The forest shown here was grazed periodically at light utilization (low degrees of use). There was extremely vigerous reproduction, both sexual and axexual, of both cottonwood and willow species. Generally members of the Salicaceae sprout vigerously particularily under moist soil conditions as on this subirrigated floodplain in the rolling red prairies. Successful reproduction of the salicaceous shrubs is a good indicator of proper management of the grazing resource and range ecosystem. Correct identifiction of willows (Salix spp.) to the correct species is a challenging task. It is often impossible to obtain positive identification without the catkins. There appeared to two and, probably, three species in this bottomland forest. This most likely "mixture" being some combination of black willow (S. nigra), peachleaf williw (S. amygdaloides), coyote willow (S. exigua), or sandbar willow (S. interior). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). No Kuchler unit for southern or southwestern cottonwood and/or willow riparian forest. SAF 63 (Cottonwood). Populus deltoides association (if and when such is recognized), Cottonwood- Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Rolling Red Plains Vegetational Area: Central Great Plains- Red Prairie Ecoregion 27h (Griffith et al., 2004). June, summer aspect. Flood plain, North Fork of Red River. Wheeler County, Texas.

40. Excellent regeneration of plains cottonwood and willow species on a Great Plains bottomland forest- This is a close-up view of outstanding production of the "next generation" of eastern cottonwood (foreground) and willow (background) on a subirrigated river bottom in the Great Plains. Mixed prairie vegetation surrounded this wetland ecosystem.Trees were functioning as phreatophytes. FRES No.17 (Elm-Ash-Cottonwood Forest Ecosystem) .No Kuchler unit for southern or southwestern cottonwood and/or willow riparian forest. SAF 63 (Cottonwood) Populus deltoids association (if and when such is recognized), Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Rolling Red Plains Vegetational Area: Central Great Plains- Red Prairie Ecoregion 27h (Griffith et al., 2005). June, summer aspect. Flood plain, North Fork of Red River. Wheeler County, Texas.

41. Understorey of a cottonwood-willow bottomland forest- A lower woody layer of regenerating plains cottonwood and willow, an herbaceous layer of grasses and grass-like plants, and a multi-storied liana layer of greenbriar (Smilax sp.) and grape (Vitis sp.) added to the mature tree layer such that this range plant community formed a true forest structure on a wide expanse of river floodplain. Understorey graminoids included bottomland switchgrass, Canada wildrye, Johnsongrass, sedges, spike rushes, rushes, and cattail. This was another of the widespread and naturalized Johnsongrass. The most common forb was wild or American licorice. Proper range management with emphasis on light and periodic grazing/browsing (vs. the widespread practice of continuous heavy grazing) permitted perpetuation of this forest. This was an outstanding example of good stewardship and sustainability of range resources. FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). No Kuchler unit for southern and southwestern cottonwood and/or willow gallery forest. SAF 63 (Cottonwood). Populus deltoides association(if and when such is recognized), Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Rolling Red Plains Vegetational Area: Central Great Plains- Red Prairie Ecoregion 27h (Griffith et al., 2004). June, summer aspect. Flood plain, North Fork of Red River. Wheeler County, Texas.

42. Palatable resprouts- Root sprouts (suckers) from a felled, senescing eastern cottonwood on a prairie in the western Ozark Plateau. The three sets of root shoots presented here (first and second photographs) were two-year-old suckers or heterophyllous shoots produced from lateral roots (third photograph) of a relatively old, "half-rotten" eastern cottonwood after it was felled for fuel wood. The shoots, some of which in these photographs were over eight feet in height, had been completely defoliated (100% of leaves removed) by commercial beef cattle (cows and calves of mixed breeding) in both the first and second years of shoot development. The small leaves present on shoots at time of photographs were current-season's regrowth (early part of current--second--warm growing season). Several (a high proprotion) of cottonwood root suckers were broken off by browsing cattle so that this extreme or severe degree of use included both eating of leaves and upper twigs of current growing season as well as complete breaking off of many root suckers that were the production of two growing seasons. Several of the broken shoots and branches were visible in some of the photographs presented here and immediately below. Cattle defoliation in the first growing season was heavy, but not total as in the second season of growth. In the second growing season many of the first and current year's root shoots were broken off at their point of origin on the lateral roots. This was done by cattle who "rode down" the root suckers in order to reach the uppermost leaves and terminal buds (ie. cattle ate all leaves and much of the upper parts of shoots, terminal twigs, plus broke off entire shoots by the browsing technique of "riding down"). "Riding down" is the feeding behavior whereby browsing animals reach up as high as they can on plants and then commence walking forward so that forage or browse plants bend down beneath the animals' weight and pass progressively beneath the feeding animals' brisket and belly until the animals have fed as far as the plant will bend to within reach of the animals' mouth. In the process, some of the brittle shoots and branches are broken off while the more limber shoots spring back-- minus leaves, fruit, buds, twigs, etc. that were consumed by the animals. In this feeding pattern animals can remove plant material at much greater heights than they could otherwise reach by merely streatching their necks and extending their tongues. Herbaceous plant species growing adjacent to and beneath eastern cottonwood shoots was mostly the introduced pasture grass, tall fescue (Festuca arundinaceae), which has naturalized across most of the greater Ozark Region and had self-seeded onto this former tallgrass prairie. The second-most common herbaceous species on this local habitat was Japanese chess or Japanese brome (Bromus japonicus), a naturalized Eurasian annual grass. It appeared that none of the herbage of these two non-native grasses had been eaten by cattle although considerable quantities of biomass had been trampled. Cattle seemingly had quite a browsing bout on this spot as quantities of dung were left. There was no evidence of browsing by deer, rodents or rabbits. Total herbivory (=herbivorous action) that did and would continue to influence regeneration of eastern cottonwood on this pasture included at least four components: 1) complete removal of leaves from cottonwood shoots, 2) total elimination of some cottonwood shoots by breakage at ground level (ie. total loss of some two-year old trees or two growing seasons of plant regeneration), 3) feeding selectivity that severely injured or setback cottonwood while at the same time having much less defoliation impact on annual and perennial grasses (both non-natives), and 4) deposition of dung with importation of more potential plant nutrients than was recycled or otherwise generated on this local environment. Question to the "village idiot" (so that he can be "above average"): "will grasses or cottonwood derive most benefit for plant growth and/or survival from the combined influences of cattle feeding?" Eastern cottonwood (both eastern and plains varieties), like most other members of the Salicaceae, readily coppices or resprouts following major injury or defoliation. Fowells (1965, ps. 516, 521) and Burns and Honkala (1990, vol. 2, ps. 533, 539) described vegetative reproduction in eastern cottonwood. Eastern cottonwood sprouts vigerously from both stumps and roots. In the pattern of most sprouting species, vegetative (asexual) reproduction declines with increaseing age of trees while vegetative propagation, both natural and atrtificial, is vigerous in young plants. Fowells (1965, p. 516) and Burns and Honkala (1990, vol. 2, ps. 533) cited research which found that eastern cottonwood (P. deltoides var. deltoides) stump-sprouted up to ages of 25 years. Heterophyllous shoots (root sprouts or suckers) shown here were from a much older tree. Over half of the trunk (beyond the center growth rings) of this eastern cottonwood had rotted away so that it was impossible to accurately determine tree age. (Rotting appeared to have begun around an old fire scar.) Recollections by the author and his conversations with elderly observers firmly established age of this felled tree in excess of 80 years. Root suckering began in early spring following sawing of the old tree in the preceding winter. On various occasions this author has observed resprouting by various kinds of cottonwood (Populus spp.) in larger and older tindividual trees following sawing, bulldozing, f lood breakage, etc. Obviously resprouting in cottonwood is not limited to trees that are younger than a quarter century. It was also obvious that regeneration of eastern cottonwood depends upon protection from overbrowsing. It was possible that some of the herbivory described above benefitted (or had potential to benefit) eastern cottonwood. Dunging provided extra plant nutrients, especially nitrogen, available to surviving cottonwood shoots. Dung might also benefit grass proportionately more than cottonwood so that increased competition from grass resulting from soil fertilization was detrimental to cottonwood. It was conceivable that breakage of some cottonwood root sprouts (suckers) had the effect of beneficial prunning which would enable surviving shoots to grow larger faster and become too large for cattle to "ride down" and thereby escape most further browsing impacts (other than continued dunging if and when cattle rested in shade of growing cottonwoods). Overall, however, and up to the point of time recorded here, even the village idiot made 100% of this pop quiz by explaining (in simple talk) that the browsing shown here was excessive and could result in death of resprouts and ultimate loss of eastern cottonwood from this pasture. Ottawa County, Oklahoma. June.

43. Overutilization of palatable eastern cottonwood by cattle- Excessive (extreme or severe) defoliation of two-year old root suckers of eastern cottonwood on a prairie in the Ozark Plateau. Beef cattle (crossbred cows and calves) removed all leaves and many shoot tips (including developing buds) from these heterophyllous shoots by early to mid-spring of the second growing season. This necessitated production and development of a new set of leaves (the leaves present in these and preceding photographs). Numerous branches were broken from shoots that survived "rding down" browsing by cattle, and lots of the two-year old shoots (root suckers) did not survive the aggressive feeding by cattle. This was an example of overuse which, if continued, might well result in killing and loss of eastern cottonwood from this pasture. These slides demonstrated the importance of protection of young eastern cottonwoods that are extremely palatable (even to animal species that are grazers more than browsers) from excessive browsing. Regeneration of eastern or plains cottonwood such as that shown above in a plains cottonwood-willow bottomland forest would not have been possible with continued overuse at the severity shown here and in the three immediately preceding slides. If degree(s) of use showh here continued for just a few more years (number unknown; would vary depending on many factors) the eastern cottonwood will die out on this pasture. The old felled cottonwood tree described in the preceding caption will be dead afterall. The main lesson from these photographs was the concept of proper degree of use (the first and most important of the Four Cardinal Principles of Range Management), often expressed as Proper Use Factors. Secondary lessons from these slides involved range animal behavior, including both the phenomenon of "riding down" and feeding selectivity. A degree of defoliation approaching "anililation" of eastern cottonwood coupled with non-eating of tall fescue and Japanese brome (though with some trampling and covering with dung of these grasses) provided a "textbook example" of feeding selectivity, the threeway interaction of animal preferences for certain plant species, relative palatability of different plant species, and availability of herbage and browse of certain plant species. Ottawa County, Oklahoma. June.

44. It'll do its part and its best, but eastern cottonwood cannot rise from the dead indefinitely- Close-up view of one of the severely browsed shoots that suckered from roots of a felled eastern cottonwood described above. All leaves are regrowth that replaced earlier full-grown leaves that were eaten by browsing beef cattle. These were some shoots that survived extreme degrees of browsing. Many shoots did not survive. (This exterme defoliation might be expected from overpopulations of browsers like deer, but this was from just "plain ole cows".) If the rangeman takes care of the feed resource it will take care of him. If this cattleman wanted natural replacement of a nice shade tree a drastic change of management was in order. A quicky, makeshift exclosure of wovn wire would protect these young shoots until they could grown beyond reach of the cattle. If this cattleman had the goal of elimination of eastern cottonwood from this pasture he had a good start toward that objective. In other words, if eastern cottonwood was regarded as a brush species (a woody noxious plant) and a single-species stand of tall fescue was the desired result then cattle browsing appeared to be a very effective method of biological control to reduce (perhaps totally eliminate) this woody species from this grazing (or haying) unit of land. As it happened such was not the case. This cattle-owner was simply running cattle for tax purposes and social status (ie."dodging taxes" and "bragging rights"). This photographic case study was used to illustrate how excessive degree of use and/or general imporper animal browsing can prevent or greatly reduce regeneration of eastern cottonwood. Degree of use on eastern cottonwood would almost certainly have been much less if cattle-grazing had been restricted to late autumn through winter-early spring when cottonwood was dormant and nutritive value and palatability of the naturalized cool-season grasses was higher. Ottawa County, Oklahoma. June.

45. Standing in river water- Mature Rio Grande cottonwood trees were flooded by water from the Rio Grande River that flowed a short distance from this location. River water rose above the ground level via subsurface flow. Ponded water was visible because manicuring with power tools for a picnic area (in a national park) eliminated the understorey of what was (would have been) a bottomland or gallery forest in the Chihuhuan Desert. The herbaceous layer in this radically modified vegetation consisted almost exclusively of introduced perennial grasses including bermudagrass (Cynodon dactylon), King Ranch bluestem (Andropoogon ischaemum= Bothriochloa ischaemum), and Dallisgrass (Paspalum dilatatum). Natural vegetation of diverse structure, including a shrub layer below the tree layer and the invasive Eurasian salt cedar (Tamarix chinensis), was present in the background. FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). No Kuchler unit for southern or southwestern cottonwood and/or gallery forest.. SAF 63 (Cottonwood). Populus deltoides association (if and when such is recognized), Cottonwood-Willow Series in Southwestern Riparian Deciduous Forest botic community of Brown et al. (1998). Trans-Pecos Basin & Range Vegetational Area: Chihuhuan Deserts- Low Mountains & Bajadas Ecoregion 24c (Griffith et al. 2004). June, summer aspect. Floodplain of Rio Grande (water mostly from Rio Concho), Big Bend National Park, Brewster County, Texas.

46.   Bottomland (floodplain) cottonwood forest- Exterior (physiogonomic) view of a local forest community dominated by narrowleaf and black cottonwood (Populus angustifolia, P. trichocarpa, respectively). A deciduous forest with an herbaceous understory of introduced Eurasian grasses developed along this alluvial bar of the Gallatin River. Gallatin County, Montana. June. FRES No. 17 (Elm-Ash-Cottonwood Ecosystem). K-89 (Northern Flood-Plain Forest). SAF 235 (Cottonwood-Willow). Cottonwood-Willow Series (222.51) in Rocky Mountain Riparian Deciduous Forest biotic community (222.5) of Brown et al. (1998, p. 43). Middle Rockies- Townsend Basin Ecoregion, 17w (Woods et al., 2002).

47.   Interior of a floodplain narrowleaf and black cottonwood forest- The two cottonwood species form an interlocking canopy and primarily constitute a single layer of woody plants, however prolific cottonwood reproduction has formed somewhat of a secondary (lower) woody plant layer which is intermediate to the lower herbaceous layer. This herb layer is almost exclusively introduced Eurasian grasses (orchardgrass, timothy, and smooth bromegrass; in that order of dominance, at least aspect dominance) and the Eurasian forb, burdock  (Arctium minus). Gallatin County, Montana. FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem).  K-89 (Northern Flood-Plain Forest). SAF 235 (Cottonwood-Willow). Cottonwood-Willow Series (222.51) in Rocky Mountain Riparian Deciduous Forest biotic community (222.5) of Brown et al. (1998, p. 43). Middle Rockies- Townsend Basin Ecoregion, 17w (Woods et al., 2002).

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48.   Branches, leaves, and inflorescences of narrowleaf cottonwood- Utah County, Utah. June.

49. Black cottonwood (Poplar trichocarpa)- This specimen of nearly giantic size was an associate to western redcedar (Thuja plicata) at edge of a grove of the latter species. Glacier National Park, Montana. July.

50. Leaves and bark of black cottonwood- This nice specimen was growing on the beautiful campus of Oregon State University. Benton County Oregon.

51. Fruit of black cottonwood- The comose seeds in a ripe, opened capsule of black cottonwood. All members of the Salicaceae are dioecious so that each black cottonwood is either male or female. It is the cotton-like comose fruit that was responsible for the common name of cottonwood, and because only females bear fruit it was female trees responsible for the name (and the debris resulting from the pubescence). Mutnoman County, Oregon. June.

52. Cottonwoods and water- Outer edge or perimeter of a black cottonwood with an herbaceous understorey along the Methow River in the Okanogan Valley at base of the Northern Cascades foothills. Basin wildrye dominated the rangeland just outside of the black cottonwood floodplain forest. Okanogan County, Washington. June (early summer). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem).  K-89 (Northern Flood-Plain Forest). SAF 235 (Cottonwood-Willow). Cascade-Sierran Riparian Deciduous Forest (222.4)- Cottonwood-Willow Series (222.41) of Brown et al. 1998, p. 43). Columbia Plateau- Okanogan Valley Ecoregion, 10m (Environmental Protection Agency, undated).

53. Black cottonwood river bottom forest- Physiogonomy, structure, and species composition of a black cottonwood forest that developed on the floodplain of Methow River. Black cottonwood constituted the sole tree species and tree (upper) layer of this forest range community. Larger saplings and pole-size young trees of black cottonwood made up a sporadic or interrupted lower tree or upper shrub layer. The lowest layer of vascular plants was composed of both herbaceous and shrub species with species composition and structure of this layer varying considerably based on forest gaps from fallen or felled trees, the latter by beaver (Castor canadensis). There were colonies of blue wildrye, the dominant native grass, that ranged from "pure" (ie.single-species) stands to local stands mixed with introduced and naturalized Eurasian perennial grasses including smooth brome, Kentucky bluegrass, reed canarygrass (Phlaris arundinaceae) and intermediate wheatgrass (Agropyron intermedium) and star Solomon's seal, the main range forb. Bush honeysuckle, black twinberry, or black bearberry (Lonicera involucrata) was the dominant understorey shrub throughout much of this understorey although common snowberry (Symphoricarpos alba) was a local dominant in better-lite microsites. Western virgin's bower or western clematis (Clematis ligusticifolia) united the lower and uppermost layer of this forest range vegetation by twining its way to the cottonwood canopy. There were scattered ponderosa pines along outermost edges of the cottonwood forest, but pines did not penetrate into the cottonwood-dominated community where lack of light was a (probably, the) major limiting factor that excluded ponerosa pine. Felling of cottonwood (mostly mid-size trees) by beaver appeared to be the major phenomenon that permitted establishment of ponderosa pine around perimeter of the black cottonwood forest. There was no Douglas-fir, period. The larger log (far left) was a black cottonwood casualty to wind; the smaller log of black cottonwood (center foreground) fell to hungry, industrous beaver. Okanogan County, Washington. June (early summer). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem).  K-89 (Northern Flood-Plain Forest). SAF 235 (Cottonwood-Willow). Cascade-Sierran Riparian Deciduous Forest (222.4)- Cottonwood-Willow Series (222.41) of Brown et al. 1998, p. 43). Columbia Plateau- Okanogan Valley Ecoregion, 10m (Environmental Protection Agency, undated).

54. Into the woods- View from outermost margin of a black cottonwood floodplain forest revealing species composition and structure of a multi-layered, relatively open, single-tree species forest community. Range plant species in this bottomland forest were given in the immediately preceding caption. Besides black cottonwood, plant species distinguishable in the understorey included star Solomon's seal, bush huckleberry or black twinberry, blue wildrye, and smooth brome. Light conditions were those of early morning (about two hours post sunrise) which from that point to mid-morning was the period of most light (greatest intensity) at this eastern edge of the forest. Okanogan County, Washington. June (early summer). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem).  K-89 (Northern Flood-Plain Forest). SAF 235 (Cottonwood-Willow). Cascade-Sierran Riparian Deciduous Forest (222.4)- Cottonwood-Willow Series (222.41) of Brown et al. 1998, p. 43). Columbia Plateau- Okanogan Valley Ecoregion, 10m (Environmental Protection Agency, undated).

55. Morning light beaming through the cottonwoods- Early morning light permitted this perspective on range vegetation of a black cottonwood floodplain forest that had an herbaceous understorey in all but the shadiest interior of the forest. This slide of a black cottonwood forest range along the Methow River provided an example of the density at which adult trees can occur and survive in competition for light. It was not known how many, if any, of these large, individual shoots of black cottonwood were clones. Some of the more closely spaced trunks such as those shown here could have been stump sprouts--and thus ramets or clones--of a preceding tree (which in turn could have been a sprout from a pre-existing trunk). Alternatively, each of the trunks in this local stand could have been a separate tree (ie. each could have been a unique genetic individual). As was shown below, these black cottonwood were producing an abundant seed crop that was readily dispersed (as anyone having experience with cottonwood trees can attest). Black cottonwood also readily regenerates asexually, including coppicing from stumps created by wind breakage, flooding, beaver feeding, and human felling. Unlike most hardwood species in which coppicing decreases with age and size of tree, most cottonwood species, including black cottonwood, sprout prolifically largely irrespective of size and age of trunk. Sprouting from broken or felled trunks often takes place immediately below the site of injury or loss, including several yards above ground when wind or ice "tops" (breaks off the crown). Major range animals in this black cottonwood-dominated range vegetation were mule deer and beaver. Okanogan County, Washington. June (early summer). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem).  K-89 (Northern Flood-Plain Forest). SAF 235 (Cottonwood-Willow). Cascade-Sierran Riparian Deciduous Forest (222.4)- Cottonwood-Willow Series (222.41) of Brown et al. 1998, p. 43). Columbia Plateau- Okanogan Valley Ecoregion, 10m (Environmental Protection Agency, undated).

56. Illuminating scenes- Another view permitted by early morining light revealed the well-developed understorey of a black cottonwood forest range that developed on the floodplaind of the Methow River. The dominant shrub was bush honeysuckle or black twinberry. The major grass was the native blue wildrye, but the naturalized Eurasian perennial grasses smooth brome, Kentucky bluegrass, reed canarygrass, and intermediate wheatgrass were locally dominant to co-dominant with blue wildrye. Small saplings and regrowth sprouts of black cottonwood formed an interrupete intermediate layer between the canopy (crown layer) and the combination woody and herbaceous understorey. Okanogan County, Washington. June (early summer). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem).  K-89 (Northern Flood-Plain Forest). SAF 235 (Cottonwood-Willow). Cascade-Sierran Riparian Deciduous Forest (222.4)- Cottonwood-Willow Series (222.41) of Brown et al. 1998, p. 43). Columbia Plateau- Okanogan Valley Ecoregion, 10m (Environmental Protection Agency, undated).

57. Beaver cotton to cottonwood- Beaver were obviously fond of the inner bark and cambium of black cottonwood in this floodplain forest along the Methow River. This young sapling had been felled and the bark neatly eaten from much of its trunk. The cottonwood had resprouted as is typical of this (and most) species in the Salicaceae. DeBell (in Burns and Honkala, 1990, p.64) explained that black cottonwood develops suckers (heterophyllous shoots) from stumps, roots, and even broken branches that become buried Remarkably, black cottonwood can shed small, leafed shoots which frequently take root. Apparently this is one means of propagation by black cottonwood becomes established on soils of stream bars. Okanogan County, Washington. June (early summer).

58. Bower bows to no cottonwood- Western virgin's bower or western clematis (Clematis ligusticifolia) was infrequent but locally vigerous in the Methow River floodplain black cottonwood forest described herein. Plants of this liana species reached into crowns of mature black cottonwood to form a continuous though highly sporadic layer from understorey to canopy. Okanogan County, Washington. June (early summer). Obvious full-bloom stage.

59. The woods suckle honeysuckle- Bush honeysuckle, twin bearberry, or black twinberry (Lonicera involucrata) is a widespread, often dominant low shrub in forests of the Northern Cascades, including foothills, and forest that develop along streams throughout the Okanogan Upland Region.This stand was growing in the understorey of a Douglas-fir-Engelmann spruce forest at mid-elevation in the Northern Cascades. Okanogan National Forest, Okanogan County, Washington. June (early summer).

60. Bush honeysuckle blooming - Shoot of bush honeysuckle at peak bloom (first photograph) and details of bush honeysuckle infloresecence (second photograph). These examples were in the colony of bush honeysuckle or black twinberry in understorey of a mid-elevation Douglas-fir-Engelmann spruce forest in the Northern Cascades. Okanogan National Forest, Okanogan County, Washington. June (early summer).

61. Bush honeysuckle bearing fruit- Bush honeysuckle or black twinberry in understorey of a black cottonwood floodplain forest along the Methow River. Specimens shown here were photographed the day following photographs of this species in the understorey of a Douglas-fir-Engelmann spruce forest at a mid-elevation in the Northern Cascades range. Onset of spring at lower elevations enabled this species to bloom and bear fruit earlier than members of the same species growing at greater elevation in a conifer forest. This was a good application of Hopkin's Bioclimatic Law. Floodplain of Methow River, Okanogan County, Washington. June (early summer).

62. Seal of the woods- Local colony of star Solomon's eseal (Smilacina stellata) in a better-lite part of the understorey of the black cottonwood floodplain forest along the Methow River. This plant was growing next to a stand of blue wildrye, both of which were presented in preceding photographs. Okanogan County, Washington. June (early summer).

63. Seal of the seal- Shoot apex with fruit in star Solomon's seal. This specimen was one of several shoots in the stand shown in the