Forests and Woodlands - Range Types of North America

1. 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).

2. 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.

3. 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).

4. 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).

5. 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).

6. 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 (Eyre, 1980). 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).

7. 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 (Eyre, 1980). 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).

8. 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 (Eyre, 1980). 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).

9. 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.

10. 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.

11. 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 of SAF 94 (Sycamore-Sweetgum-American Elm) that had former SAF designation of Sycamore-Pecan-American Elm (Eyre, 1980). 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).

Tree-lined on a wet prairie- Marmaton River lined with a "pecan bottoms", a climax floodplain forest dominated by pecan and with pin oak (Quercus palustris) and bur oak (Q. macrocarpa) as associated tree species. The understorey, which was obviously not visible in this slide, was nonetheless well-developed. This herbaceous understorey was shown and described in subsequent slides.

Vernon County, Missouri. Late July; mid-estival aspect. There was not an FRES Forest and Woodland Ecosytem for climax vegetation at this small a spatial scale. Closest Kuchler unit would be K-89 (Northern Flood-plain Forest). Variant of SAF 94 (Sycamore-Sweetgum-American Elm) that had former SAF designation of Sycamore-Pecan-American Elm. (Eyre, 1980). Not a good unit of wetland biotic community, but closest in Brown et al. (1998, p. 43) was Mixed Hardwood Series 223.13 of Southeastern Swamp and Riparian Forest 223.1 of Warm Temperate Swamp and Riparian forest 223. This seemed contrary to Kuchler unit. Central Irregular Plains- Cherokee Plains ecoregion 40d (Chapman et al., 2002).

Point of contact- Outer edge of a wet prairie dominated by prairie cordgrass (Spartina pectinata) in foreground and a floodplain forest dominated by pecan with pin oak as the associate tree species while bur oak was the other major tree. Understorey species of this bottomland or floodplain forest were listed in the next two captions.

Relict tract, The Nature Consrvancy, Vernon County, Missouri. Late July; mid-estival aspect. There was not an FRES Forest and Woodland Ecosytem for climax vegetation at this small a spatial scale. Closest Kuchler unit would be K-89 (Northern Flood-plain Forest). Variant of SAF 94 (Sycamore-Sweetgum-American Elm) that had former SAF designation of Sycamore-Pecan-American Elm. (Eyre, 1980). Not a good unit of wetland biotic community, but closest in Brown et al. (1998, p. 43) was Mixed Hardwood Series 223.13 of Southeastern Swamp and Riparian Forest 223.1 of Warm Temperate Swamp and Riparian forest 223. This seemed contrary to Kuchler unit. Central Irregular Plains- Cherokee Plains ecoregion 40d (Chapman et al., 2002).

Directions for continuity: a wet prairie dominated by prairie cordgrass (Spartina pectinata) had developed at the edge of this pecan bottomland forest. The outer primeter of that was shown in the foreground of the immediately preceding slide. That example of prairie cordgrass wet prairie was covered in Range Types of North America in the chapter entitled, Tallgrass Prairie-I.

Sometimes the water stands deep here- Interior of a floodplain (of the Marmaton River) forest dominated by pecn with pin oak as the associate tree species and bur oak as the third important arboreal actor on this sylvan stage. The understorey as seen here was co-dominated by Virginia wildrye (Elymus virginicus) and broadleaf woodoats (Uniola latifolia). The most common or abundant forb was purple or sweet joe-pye weed (Eupatorum purpureum). Two other common forbs (both annuals) were Pennsylvania smartweed (Polygonum pennsylvanicum) and giant ragweed (Ambrosia trifica). All of these species (from tree therough grass and annual forb) were native species.

This climax floodplain forest was at the edge of a prairie cordgrass wet prairie (a consociation of Sprtina pectinata) as shown in the immediately preceding slide. This potential natural vegetation was in the Osage Plains of western Missouri.

When the Marmaton River flooded water in this floodplain forest somtimes reached heights of 20 feet. This author was not present when such flooding took place, but he was told this fact by a resident manager of native pecan forests whose owners contracted with pecan companies who sold and distributed pecans to retailers.

This was an example of multiple use. The main crop of this forest was pecans, but beef cattle as well as wildlife species such as white-tailed deer (Odocoileus virginianus) grazed many such forests so that forage (ie. ultimately, beef) and general habitat (hence, wildlife) as well as nuts were agricultural commodities and, ultimately, food and fiber for human use. Perhaps the most important ecological services provided by this natural vegetation were watershed protection and flood abatement. Or, just maybe, the greatest food from such forest range was its priceless aesthetic value. The beauty of this woods was beyond mere price.

Relict tract, The Nature Conservancy, Vernon County, Missouri. Late July; mid-estival aspect. There was not an FRES Forest and Woodland Ecosytem for climax vegetation at this small a spatial scale. Closest Kuchler unit would be K-89 (Northern Flood-plain Forest). Variant of SAF 94 (Sycamore-Sweetgum-American Elm) that had former SAF designation of Sycamore-Pecan-American Elm. (Eyre, 1980). Not a good unit of wetland biotic community, but closest in Brown et al. (1998, p. 43) was Mixed Hardwood Series 223.13 of Southeastern Swamp and Riparian Forest 223.1 of Warm Temperate Swamp and Riparian forest 223. This seemed contrary to Kuchler unit. Central Irregular Plains- Cherokee Plains ecoregion 40d (Chapman et al., 2002).

When it floods...- At perimeter of a climax bottomland forest that developed on the floodplain of the Marmaton River. This was a "pecan bottoms" forest in which pin oak was the associate species and bur oak was the "first runner-up" for associate status on this pecan-dominated forest that was surrounded by wet prairie which was a consociation of prairie cordgrass. The herbaceous understorey of this floodplain forest range was co-dominated by Virginia wildrye and broadleaf woodoats, but there were local colonies of purple or sweet joe-pye weed, such as the one shown in foreground of these two slides, exclusively made ujp of this single species such that purple joe-pye weed was a local exclusionary dominant. Other Two other important and locally abundant forbs were the native annuals, Pennsylvania smartweed and giant ragweed.

Ultimate expression (except for younger trees )- Another "pecan bottoms" forest on floodplain of Marmaton River in the Osage Plains of western Missouri. In something of a contrast to the mixed tree species forest described immediately above where pin oak and bur oak joined pecan, the tree component of this pecan bottomland forest was comprised exclusively of pecan, and of substantially smaller trees (supposedly younger trees though tree-ring data were lacking). Likewise, the herbaceous understorey was less species-diverse with the herbaceous component being comprised almost entirely of Virginia wildrye except in laarge, local areas where Pennsylvania smartweed accompanied Virginia wildrye.

This climax range vegetation would comport with the official definition of a forest (Helms, 1998), but in some parts of this simple range plant community the designation of woodland was more appropriate because crowns of individual pecans did not come into contact with (did not "touch") each other or did not interlock. Either way this permitted enough light to penetrate through the tree canopy that the understorey was one of the most beautiful stands of Virginia wildrye imaginable.

Forest range like this takes rangemen back to the days of the redman. Did the Osages have campfires in these woods before the whiteman stole this land from them?

A more immediately relevant question would involve the possibility that this forest had been logged at some point in time. All the trees in this bottomland forest were obviously second-growth individuals of roughly "middle age" (trees were approximately at the half-way stage between seedling and senescent or slowly dying mature age (stage of a tree's life cycle). There were no old-growth individuals of pecan present.

This forest (and other forests like the one shown immediately above) on the floodplain of Marmaton River were managed to produce "hardshell" (native or non- domesticated varieties) pecans. The pecan nuts produced on these trees were gathered from the ground and sold to large companies that, in turn, sold them to retail outlets.

Vernon County, Missouri. Mid-July; early estival aspect. There was not an FRES Forest and Woodland Ecosytem for climax vegetation at this small a spatial scale. Closest Kuchler unit would be K-89 (Northern Flood-plain Forest). Variant of SAF 94 (Sycamore-Sweetgum-American Elm) that had former SAF designation of Sycamore-Pecan-American Elm. (Eyre, 1980). Not a good unit of wetland biotic community, but closest in Brown et al. (1998, p. 43) was Mixed Hardwood Series 223.13 of Southeastern Swamp and Riparian Forest 223.1 of Warm Temperate Swamp and Riparian forest 223. This seemed contrary to Kuchler unit. Central Irregular Plains- Cherokee Plains ecoregion 40d (Chapman et al., 2002).

Pecan bottoms inside out- Interior of a pecan bottoms forest. This was a consociation of pecan with an herbaceous understorey comprised almost exclusively of Virginia wildrye except for patches of Pennsylvania smartweed. Trunk of the foremost pecan was being "clumb up" by a still-yound trumpet creeper (Campestrus radicans). The herbaceous was almost entirely Virginia wildrye with local pateches of Pennsylvania smartweed.

This was a second-growth forest, there being no old-growth trees remaining. Even the largest individuals in this forest community, such as the one featured in the second slide, had trunk diameters that were hardly half the size of ancient pecans seen in other forest tracts like the one shown and described in the immediately preceding example (compare size and shape of trunks of pecans in this and that tract).

Although the oldest trees in this forest tract were still decades away from reaching full-adult size and habit (outward appearance of plants or their general or grass morphology) and having the fully developed structure of climax vegetation, species composition of the total forest community was that of climax for this floodplain forest.

Under the pecans- Sward of the herbaceous understorey of a second-growth yet climax floodplain pecan-dominated forest. The single herbaceous layer of this forest range was composed almost exclusively of Virginia wildrye (first slide), but there were local areas where Pennsylvania smartweed (a native annual forb) was co-dominant with Virginia wildrye (second slide). This was the single herbaceous layer of the "pecan bottoms" presented in the two immediately preceding slide/caption sets.

Green Ash (Fraxinus pennsylvanica)

Green ash (Fraxinus pennsylvanica) has the largest biological range of any Fraxinus species in North America extending from northern parts of the Prairie Privinces to southeast Texas. Green ash is one of the domiant species of certain bottomland forests such as those covered under forest cover type, Sugarberry-American Elm-Green Ash (SAF 93), but green ash does not constitute a forest cover type "of its own" so to speak such as those that are named after a single species. In the Northern Great Plains a unit of natural vegetation dominated by green ash was described by Barker and Whitman (1989). This native plant community--though highly modified with understories comprised of naturalized agronomic grasses from Eurasia--exist primarily as groves (rather than as forests or woodlands) much like groves of bur oak (Quercus macrocarpa).

Green ash groves are not a major cover type. This tree-dominated vegetation was not recognized by the Society of American Foresters (Eyre, 1980), but it is an important unit of range vegetation parts of the Great Plains and the Central Lowlands physiographic provinces. For that reason, green ash dominated/defined range was includedin the category of Miscellaneous under Forest and Woodland units of vegetation.

Location note: An example of a gallery forest dominated by green ash (with several accompanying associate and minor tree species) that developed on the floodplain of Wichita River in northcentral Texas ( a variant of SAF 93) was included at end of the chapter, Central and Southern Forests-I.

12. Local forest in the badlands- This and the next three slide-caption sets described the Green Ash type described by Barker and Whitman (1989). These authors remarkded that this unit of vegetation was a domint (= dominance) type on the more more mesic habitats of north- and east-slopes in draws, lower stream terraces, and bottomlands. These occur as small stands, often of only a few acres (Barker and Whitman, 1989).

The Green Ash cover type is likely more readily viewed as a woodland than a forest type, even though trees typically grow close enough together to have interlocking crowns. Grove was this author's choice of terms for these localized groups of trees. There is almost always an herbaceous understorey, usually dominated by grasses, and frequently one to two lower woody layers (mostly of shrubs of various heights). The dominant grass in the herbaceous layer of the green ash grove presented here was smooth brome (Bromus inermis). In fact, this naturalized agronomic--turned weedy--Eurasian grass often excludes other herbaceous species (as in the instance of this green ash grove) . Otherwise, dominant grasses would have been western wheatgrass (Agropyron smithii), needle-and-thread (Stipa comata), little bluestem (Andropogon scoparius), green needlegrass (Stipa viridula), blue grama (Bouteloua gracilis), and sideoats grama (B. curtipendula).

In badlands such as those of the Little Missouri River the Green Ash type is commonly associated with river-bottom--often riparian--forests of eastern cottonwood (Populus deltoides subsp. moniliferaa) and willow species like peachleaf willow (Salix amygdaloides) (SAF 235) and at higher, harsher environments with the Rocky Mountain Juniper type (SAF 220).

Theodore Rooseveldt National Park (South Unit), Billings County, North Dakota. FRES No. 17 (Elm--Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash Forest). No SAF. Green Ash type of Barker and Whitman, 1989). Brown et al. (1998, 37) did not include any appropriate vegetational unit (neither biotic community nor series) below that of Cold Temperate Forest and Woodland 122. Northwestern Great Plains- Little Missouri Badlands Ecoregion Ecoregion 43b (Byce et al., Undated).

13. Outside of a young grove- A small (about two acres) forest stand or, as preferred by this author, grove of young green ash growing on a disturbance climax of smooth brome. In the context of Landscape Ecology, this ash grove can be visualized as a patch (a small one) within a matrix of Northern Mixed Prairie. The potential natural (climax) plant community of this mixed prairie was western wheatgrass, thickspike wheatgrass (Agropyron dasystachum), little bluestem, sideoats grama, blue grama, green needlegrass, and needle-and-thread. Here in this photograph of range vegetation these native grasses had been largely displace by the aggressive, weedy smooth bormegrass. It appeared that a previous disturbance of earth-moving (to get fill for road construction) had permitted establishment of the smooth brome--and perhaps of green ash.

(Mixed prairie consisting of many of the native grasses just listed,and in Excellent range condition class, was adjacent to this green ash-smooth bromegrass vegetation, but photographs that would have included that climax grassland would have put the grove of green ash too far away for instructional purposes. This photographer did find forests dominated by green ash (groves of green ash) in which native plant species, including numerous kinds of shrubs and grasses, formed layers resulting in a natural understorey. Two examples of such natural plant communities were presented in subsequent sets of slides and caption.)

Burns and Honkala (1990) cited work explaining that green ash had tolerance ratings ranging from Intolerant to Moderately Tolerant in northern environments. These authors concluded that green ash was an early successional species serving as either a pioneer (first seral stage) species or as a colonizer in second or third successional stages after colonization by Populus or Salix species, especially on alluvial soils. It seemed likely that the disturbance due to earth-moving had prepared this site for establishment of green ash (and smooth brome).

This range vegetation had developed in the unglaciated (sedimentary) and more western portion of the Northern Great Plains.

Little Missouri National Grassland, Golden Valley County, North Dakota. FRES No. 17 (Elm--Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash Forest). No SAF. Green Ash type of Barker and Whitman, 1989). Brown et al. (1998, 37) did not include any appropriate vegetational unit (neither biotic community nor series) below that of Cold Temperate Forest and Woodland 122. Northwestern Great Plains- Missouri Plateau Ecoregion 43a (Bryce et al., Undated).

14. Green grew the draw- A woody draw in the Little Missouri River Badlands dominated by green ash (ie. an example of the Green Ash cover type) with two or three woody layeries and one herbaceous layer. The co-dominant shrubs were buffaloberry (Shepherdia argentea) and chokecherry (Prunus virginiana). Western snowberry (Symphoricarpos occidentalis) and silver sagebrush were associate shrub species. The dominant herbaceous species was western wheatgrass. The most abundant forbs was Indian hemp (Apocynum cannabium) and western yarrow (Achillea lanulosa= A. millefolium subsp. lanulosa).

Physiogonomy, structure, and botanical composition of this forest range cover type was visible in these two photographs.This woodland range vegetation was in the more western portion, the sedimentary or unglaciated part, of the Northern Great Plains.

Theodore Rooseveldt National Park (north unit), McKinzie County, North Dakota. FRES No. 17 (Elm--Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash Forest). No SAF. Green Ash type of Barker and Whitman, 1989). Brown et al. (1998, 37) did not include any appropriate vegetational unit (neither biotic community nor series) below that of Cold Temperate Forest and Woodland 122.Northwestern Great Plains- Little Missouri Badlands Ecoregion Ecoregion 43b (Byce et al., Undated).

15. Fire and high water- A grove of green ash with one shrub and one herbaceous layer on the farther portion of the first terrace of the Little Missouri River within the badlands land form. A wild fire (and a hot one at that) had burnt this forest two years earlier so that some adult ash trees had been topkilled (or at least mostly topkilled) and the woody and herbaceous understorey substantially impacted as well. Much of the shrub layer was chokecherry with western snowberry being the weak associate to this dominant. There were a few widely scattered plants of silver sagebrush around the perimeter of this grove, but these had not invaded into the interior of the forest stand. There was no herbaceous layer in this burnt-over forest and, in fact, very few herbaceous plants other than very stunted individuals of Kentucky bluegrass (foreground of second slide) and even fewer plants of smooth brome and needle-and-thread. It was notable that the most common herbaceous species was a naturalized Eurasian grass, and the perennial grass most abundant on overgrazed ranges in the Northern Great Plains Region (see discussion of Kentucky bluegrass disturbance climax in the chapter, Introduced Forages-Grasses, under Grasslands biomes).

This grove of green ash was just beyond a riparian zone of common reed (Phragmites communis= P. australis) and then a floodplain forest of eastern cottonwood (Populus deltoides subsp. monilifera) and peachleaf willow (Salix amygdaloides). (These two units of bottomland vegetation were presented below in the section of eastern cottonwood forest.) These three distinct plant communities along with a savanna of silver sagebrush as associate to the dominants of needle-and-thread and prairie sandreed (Calamolvia longifolia) comprised the four units of range vegetation on this floodplain of the Little Missouri River.

This entire landscape of Little Missouri River Badlands was in the unglaciated Northern Great Plains.

Theodore Rooseveldt National Park (South Unit), Billings County, North Dakota. FRES No. 17 (Elm--Ash-Cottonwood Forest Ecosystem). K-92 (Elm-Ash Forest). No SAF. Green Ash type of Barker and Whitman, 1989). Brown et al. (1998, 37) did not include any appropriate vegetational unit (neither biotic community nor series) below that of Cold Temperate Forest and Woodland 122.Northwestern Great Plains- Little Missouri Badlands Ecoregion Ecoregion 43b (Byce et al., Undated).

16. Green ash (Fraxinus pennsylvanica)- Some examples of seedlings and saplings of green ash on tne perimeter of a greeh ash grove in the Northern Great Plains. The herbaceous layer in these slides was primarily smooth brome, but there were a number of plants (sizable cover) of such native grasses as western wheatgrass, thickspike wheatgrass, green needlegrass, little bluestem, and blue grama.

Little Missouri National Grassland, Golden Valley County, North Dakota. Late June.

17. Green against northern blue- Leaves and clusters of samaras of green ash that was growing in a grove in the Northern Great Plains. Green ash has a species range extending from northern and eastern Canada southward to Florida and father west as far south as southeast Texas. Green ash is a dioecious tree that is most abundant in more moist environments such as riparian habitats (floodplains in general) and coulees or ravines in the Great Plains.

The ash fruit, a one-seeded samara with conspicuous terminal wings, is borne in clusters.

Little Missouri National Grassland, Golden Valley County, North Dakota. Late June; immature fruit phenological stage.

Floodplain Forests

In several sections below various bottomland forest cover types (Eyre, 1954; Eyre, 1980) that function as (are used for) forest range were described.

Although most students of Range Management and Forestry (or, for that matter, general readers) comprehend the idea of floodplain the following official definition was provided to avoid any confusion: "(a) The surface or strip of relatively smooth land adjacent to a river channel, constructed by the present river in its existing regimen and cover with water when the river overflows its banks. It is built of alluvium carried by the river during floods and deposited in the sluggish water beyond the influence of the swiftest current. A river has one flood plain and may have one or more [stream] terraces representing abandoned flood plains. (b) Any flat or nearly flat lowland that borders a stream and that may be covered by its waters at flood satges; the land described by the perimeter of the maxiumm probable flood. Syn. floodland; valley floor" (Wilson and Moore, 1998). As applied herein by the current author, floodplain (can also be spelled as flood plain or flood-plain according to the cited authors) was used for any low-lying land adjacent to a stream or watercourse (brook, creek, or river) so as to be flooded or inundated periodically by water overflowing the stream bank. By extension, floodplain forests are any that develop on floodplains and that are influenced by floodplain phenomena. Bottomland is more or less synonymous with floodplain (Wilson and Moore, 1998) although strictly speaking bottom land (or, often, bottoms) is a broader (no pun intended) term as it can include lowland along the outer margin of a floodplain. Bottomland implies or connotes "[l]ow-lying, level land, usually highly fertile..." (Wilson and Moore, 1998).

It must always be borne in mind that flooding is a natural process. Flooding is a disturbance in the same context and as a factor in dynamics of vegetation just as drought, fire, wind and ice storms, frost/freeze, land slides, etc. (Obviously different disturbances bring about different effects of vegetation as, for contrast, flood versus drought.). In this regard, floodplain or bottomland forest cover types (forest range types) are flood types just as there are fire types, topographic types, and zootic types. These disturbance types can be regarded as climax types (in contrast to climatic climax types) if viewed from the perspective of polyclimax theory tracable to Sir Arthur Tansley or the climax pattern theory of Prof. Robert Whittaker. Certain forest cover types such as those of Intolerant species (eg. Populus, Salix, Alnus, Platanus, Sequoia) require fresh, alluvial deposits, such as bare sand or mineral soil, for seedling establishment. Some of these species are among the largest trees on Earth and, certainly, within any given geographic area or region.

It was, of course, muddy flood water over millenia that carried the rich alluvium which was deposited by water (aided and abetted by gravity) that built the extremely fertile soil on bottomland (floodplain) forests. Students should remember that river valleys (Mississippi, Missouri, Red, Arkansas, Shenandoah, Tennessee, Ohio, Salmon, Eel, Rio Grande, Colorado, the list is almost endless) are as much a part of a river as are its stream bank and stream channel.

Man's interference with flooding-- the natural, soil-forming, community and ecosystem-sustaining process--ultimately can only exacerbate flooding when it inevitably (maybe later than sooner, but surely) will occur again. Man cannot confine forever or indefinitely a river. Slavery is a doomed system, a flawed vision. Eventually watercourses from the big ones like Ole Man River and the Big Muddy to the little ditches like Lost Creek will bust free--channelized, dammed, dredged, or otherwise. Ultimately, the river will win. Meanwhile, enjoy the floodplain forests and their range resources.

Organization note: Forest range types shown below are those of bottomland or floodplain forests and woodlands including the entire land area of these tree-dominated range types with emphasis on the general floodplain or stream bottom. The riparian zone was sometimes included--usually unavoidably--but the emphasis was not on the riparian zone. The riparian zone is that part of the stream bottom associated directly and immediately with the stream channel, especially stream banks. Riparian vegetation is the plant community associated only directly with the stream channel (vegetation that develops along the stream bank) and extending outward only to parts of the stream bottom or floodplain on which soil remains --excepting prolonged outstanding droughts--saturated or waterlogged by lateral water flow. Riparian zone as interpreted herein did not extend outward from streams to subirrigated lands or to range vegetation that developed on subirrigated lands.

Riparian vegetation--including range plant communities that develop along streams, especially stream banks, in grasslands, deserts, tundra--was included in various chapters throughout Range Types of North America. The chapter in Range Types entitled, Willow and Riparian was devoted exclusively to woody riparian vegetation including some forest and shrubland range types dominated by cottonwood (Populus species). Readers were herewith directed there for further coverage woody range vegetation dominated by cottonwood and willow (Salix) species.

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). Other authorities published somewhat different names (taxa). Weber (1990, p.326) referred to eastern cottonwood in Colorado as P. deltoides subsp. monilifera. His treatment was likely based on that in Flora of the Great Plains where McGregor et al. (1986, ps 278-279) used subspecies monilifera as distinct from subspecies deltoides, which they limited to the easterm margin of the Great Plains, and from subspecies wislizenii, which they regarded as to the west. 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.

18. 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).

19. 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).

20. 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).

21. 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 narrowleaf cattail (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.

Members of the Salicaceae, the cottonwood-willow family, are generally dioecious; that is having the characteristic or phenomenon of unisexual genotypes (of separate male and female plants). In the section below examples of male and female catkins on eastern cottonwood were presented. These unisexual (imperfect) flowers were borne on two separate (genetically distinct) trees that were located less than one adult cottonwood crown apart.

Gold and blue- Rich amber-colored staminate catkins of eastern cottonwood against a backdrop of a rare (for Texas) azure blue sky demonstrated colorfully and at progressively closer camera distance the male-flower cluster of this iconic tree of the tallgrass prairie.

Several spring-swollen lateral buds were visible on the leader presented in the first two of these three slides.

This example came from the Grand Prairie-Western Cross Timbers vegetational area of northcentral Texas.

Paler gold with the same blue- Pale amber-colored pistillate catkins of eastern cottonwood against a backdrop of a rare (for Texas) azure blue sky demonstrated colorfully and at progressively closer camera distance the female-flower cluster of this iconic tree of the tallgrass prairie.

The female tree that bore these lady catkins was less than felling distance (less than a felled trees distance) from the gentleman catkins seen immediately above. In addition, the second image shown here sported a large and swollen terminal or apical preformed bud.

Flowers of both sexes- Female (pistillate) catkins were placed above and below a male (staminate) catkin of eastern cottonwood that was growing in the Grand Prairie-Western Cross Timbers vegetational area of northcentral Texas. These catkins were some of those presented in the two immediately preceding slide-caption sets.

Cotton candy on a tree- About one month later (after the bearing of unisexual flowers) some of the pistillate (female) catkins presented above bore ripened ovaries that were now capsules which were dehiscing and releasing seeds equipped with their namessake "cottonwood" parachute-like pilose hairs. This morphological featurewas described by Smith (1977, ps. 128, 292) as comose meaning to have a coma which is a tuft of hairs.

Parts of a foliaceous, voluptous female- Young, full-grown leaves and a pendulous cluster of immature capsules of eastern cottonwood (Populus deltoides sp. deltoides) in the Western Cross Timbers of northcentral Texas. Populus species are dioecious so this was obviously a female tree.

The cobalt-blue sky was result of a late-season norther that took out drab, high-humidity, polluted air and replaced it with fresh, clean Arctic air.

Spring-bedecked female parts- Young, fully developed leaves and a cluster of immature capsules of eastern cottonwood in the West Cross Timbers of northcentral Texas. There are any number of fine references for this State Tree of Kansas including the following which were given in contest of taxonomy and morphology of eastern cottonwood.

Systematics of Populus species (even within P. deltoides) has and continues to be a matter of confusion and conflict with any number of taxonomic treatments ranging from those of Coulter (1891-1894, ps. 419-420) and Sargent (1933, ps. 132-137 passim) through Vines (1960, ps. 89-92), Steyermark (1963, ps. 507-508), Correll and Johnston (1979, p.455), Great Plains Flora Association (1986, ps. 278-279), Diggs et al. (1999, ps. 975-976), and Allred and Ivey (2012, p. 529-530) for the general region herein involved. It seemed to this rather independent author that the treatment by Diggs et al. (1999, ps. 975-976) was most accurate and appropriate, perhaps primarily because it was the closest flora for this immediate area.

Diggs et al. (1999, ps. 975-976) elevated to two subspecies of P. deltoides some taxa (species, varieties, forms) formerly used for eastern cottonwood, plains cottonwood, and perhaps others such as Sargent's cottonwood by previously published work. In their dichotomy of these subspecies, subspecies deltoides had 10 to 20 coarse teeth on each side of the deltoid leaf whereas subspecies monilifera had only five to 10 coarse teeth along each. Viewers can handily see in various of the views presented in this short treatment that this example was clearly P. deltoides subsp. deltoides.

Great Plains Flora Association (1986, p. 279) lumped all Populus deltoides for the Great Plains as being P. deltoides subsp. monilifera with a note that it and P. deltoides subsp. deltoides intergraded with each other along eastern margins of the Great Plains. These authors (Great Plains Flora Association, 1986, p. 2) extended their version of the "Great Plains" (presumedly so that they could include all of Kansas and northward states) into the Central Lowlands physiographic province. This implied that there would be no specimens of P. deltoides subsp. deltoides in Kansas, western Missouri and so on (Great Plains Flora Association, 1986, ps. 2, 279). This would certainly to be erroneous, but again these authorities distinguished between numbers of teeth on leaf margins as between early leaves and late leaves with many more margin teeth on late leaves.

Either way, treatment by Great Plains Flora Association (1985, p. 279) clearly conflicted with treatment by Allred and Ivey (2012, ps. 529-530) who broke P. deltoides into four varieties including P. deltoides var. deltoides which entered ("scarcely") northeastern New Mexco. This is, of course, west of Kansas, Nebraska, Missouri, etc.

Burns and Honkala (1990)--official US Forest Service version--treated eastern cottonwood (P. deltoides) as P. deltoides var. deltoides for "eastern cottonwood (typical)" and P. deltoides var. occidentalis for "plains cottonwood".

How're they hanging?- Immature many-seeded capsules of eastern cottonwood. The pendulous position of the fruit cluster in the first slide is the natural or undisturbed habit whereas the "wrapped" or "coiled" position in the second slide was the result of wind action. The Populus species are not only wind-pollinated and with wind-dispersed seeds, but they also subject to numerous other "avenues", "venues", or "expressions" of wind action

22. Foliage and fruit of prairie silvan queen- First of these two photographs was young shoots (two small seedlings or small saplings) of eastern cottonwood. They had been flattened by running water from recent spring (June) rains. (They straightened up.) Second photograph was the typical deltoid leaf with ripe fruit ( a capsule) of eastern cottonwood. Cottonwood capsules are described as comose, an adjective referring to having a coma (a tuft of hairs). Cottonwood derived its common name from the phenomenon of conspicuous, usually copious (and inconvenient) shedding of the comose ("cottony") fruit. All members of the Salicaceae are dioecious so that a tree is male or female and thus produces only staminate or only pistillate flowers. Of course only female plants produce fruit so that occurrence of the "messy fluff" comes only from the ladies. (Some range men find girls troublesome while others enjoy the mess.) Female catkins from this same plant were shown in the next photograph. Erath County, Texas. May.

23. Wild or American licorice (Glycyrrhiza lepidota)- Wild licorice is one of the more common legumes along watercourses throughout the Great P0lains. 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.

24. Licoriced sticks- Shoots with ripening legumes of wild or American licorice in the floodplain of the Platte River. The two closer-in (more detailed) shots when compared to similar photographs of other range species of Leguminosae served as examples of the great diversity in shapes, sizes, pubescence, etc. of legumes.

Adams County, Colorado. Early July; advanced immature stage of fruit development.

25. 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.

26. 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.

27. Understorey of a cottonwood-willow bottomland forest- A lower woody layer of regenerating plains cottonwood and willow, an herbaceous er 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.

28. Riverbottom vegetation without a place to go- Along a recently flooded Little Missouri River there were two distinct range plant communities: 1) a reedbed of common reed (Phragmites communis= P. australis), the vegetation zone nearest the river comprised solely of short, green grass and 2) a floodplain forest co-dominated by eastern cottonwood (Populus deltoides subsp. monilifera) and peachleaf willow (Salix amygdaloides) adjoining the reedbed.

The stump in foreground of the second slide was that of a cottonwood that had been felled by feeding beaver (Castor canadensis) with the log (trunk) carried downstream by the recent record flood. Details of this coppicing (sprouting) cottonwood were shown in the next subsequent slide.

Obviously this range vegetation was as much--if not more--grassland than forest, but the cottonwood-willow forest cover type clearly certainly had to be included in this section.

Along Little Missouri River, Theodore Roosevelt National Park (North Unit), McKenzie County, North Dakota. Late June. FRES No. 17 (Elm--Ash-Cottonwood Forest Ecosystem). K-89 (Northern Floodplain Forest). SAF 235 (Cottonwood-Willow). Cottonwood-Willow Series 221.21 in Plains and Great Basin Riparian Deciduous Forest biotic community 222.2 and Plains Interior Marshland 242.3, but no Common Reed Series given of Brown et al. (1998, p.43 ). Northwestern Great Plains- Little Missouri Badlands Ecoregion Ecoregion 43b (Byce et al., Undated).

29. Browsing in a reedbank- On the immediate or near (oputermost) channel of the Little Missouri River an eastern cottonwood felled by beaver was resprouting at the edge of a cotton-willow riparian forest and a bed of common reed that had just emerged from dormancy. This entire riverbottom vegetation was recovering from recent flooding. There were two distinct floodplain plant communities: 1) the bed of common reed and 2) botttomland forest of eastern cottonwood and peachleaf willow.

30. Plains cottonwood bottomland forest in the far-western plains- Introduced in this photograph and treated further in subsequent slides was a forest range that developed along an *intermittant stream or, perhaps, interrupted stream (East Bijou Creek) in the Colordo Piedmont on which the dominant, the only, tree species was plains cottonwood (Populus deltoides ssp. monilifera). Plains cottonwood is another subspecies of eastern cottonwood native to the semiarid zone This forest range had been grazed by cattle for decades and was still in Good range condition class so as to approach the state of potential natural (climax) vegetation. There was limited reproduction of cottonwood, but this appeared adequate to maintain the uneven-aged population that dominated and defined this range plant community. Besides the upper woody (crown or canopy) layer there was a lower woody layer--though it was sporadic or "spotty" in its pattern of distribution-- made up of saplings and young poles of plains cottonwood. (This structural feature was shown in sebsequent slides.) A few seedlings of cottonwood were present in the herbaceous understorey. There were three herbaceous layers all of whch were dominated by grass species. These three were tallgrass, midgrass, and shortgrass layers so the understorey was mixed prairie. In fact this range vegetation could be visualized or even described as mixed prairie savanna with plains cottonwood as the woody element. Cottonwoods were dense enough, at least in numerous microhabitats where there were local populations (or, perhaps, individual stands would be a more apt term) that crowns typically interlocked. Thus, forest was the proper designation. Still, the overall or general vegetation could be conceived of as a savanna or, at least, as having a savanna-like physiogonomy, and a wetland savanna (or savanna wetland) at that.

Portions of this range plant community actually comprised a gallery forest along the channel of this seasonally and spatially intermittant stream. Such a dispersion pattern and structure was shown in this photograph and the next photgraph where East Bijou Creek actually held water. Along other short reaches of this stream the creek channel was dry with only moister soil to distinguish the more-or-less interrupted riparian zone from the environment of the farther (outer) channel banks and adjacent floodplain.

The herbaceous vegetation that was actually in the stream channel and extending outward on the nearest (lowr) banks was made up of prairie cordgrass (Spartina pectinata), bottomland ecotype(s) of switchgrass, and Canada wildrye (in that order of cover and biomass or standing crop).

It should be noted that in addition to the light degree of use of tallgrass species there were a lot of cottonwood leaves and lower branches within browsing reach of cattle. This forest range was not being abused by the rancher who used. A well-earned tip of the hat and public praise to the faithful steward of this land. If the author had known he name of this commendable range manager he would have printed it.

A note on stream terminology- The author was "only pasing through" this country and, as he did reside here, could not determine the nature of East Bijou Creek and use the most accurate designation. Specifically, the photographer could not distinguish whether it was an intermittent or intermediate stream.

*Intermittent stream- "A stream or reach of a stream tht flows only at certain times of the year, as when it receives water from springs or from some surface source....A stream that does not flow continuously, as when water losses from evaporation or seepage exceed the available streamflow...Synonyms: temporaray stream, seasonal stream" and

*Interrupeted stream- "A stream that contains perennial reaches with intervening intermittent or emphemeral reaches, or a stream that contains intermittent reaches with intervening ephemeral reaches" (Wilson and Moore, 1998).

East Bijou Creek, Elbert County, Colorado. Mid-July (estival aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). No Kuchler unit for southern and southwestern cottonwood forest. No SRM. 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). Southwestern Tablelands-Piedmont Plains and Tablelands Ecosystem, 26e (Chapman et al., 2006).

31. A little farther downstream- At another short reach with surface water of the channel of an intermediate or interupted stream (East Bijou Creek) along which a forest of plains cottonwood with a n herbaceous understorey had developed. In the reach of the creek viewed here the stream chaannel was smaller (narrower and shallower) so that water persisted for shorter periods than in the reach shown immediately above.

Range vegetation along this segment of East Bijou Creek was more of a mixture of aquatic and also of less mesic grass species than in the reach with a bigger channel and greater riparian zone surface seen above. In this stream segment major range plants were Torrey's rush (Juncus torreyi) and western wheatgrass as well as Canada wildrye and prairie cordgrass which were relatively more abundant around the larger area of water shown in the preceding slide. Cheatgrass or downy bromegrass (Bromus tectorum) was present in minor down to trace amounts.

Both smaller (presumedly younger) and full-sized adult trees of plains cottonwood were present in this portion of the floodplain forest.

Only range vegetation of the general riparizn zone was described in this and the preceding photograph. Composition and structure of the floodplain away from the immediate riparian zone was treated in the slides and their captions that followed immediately below.

32. Along a dry reach of the interupted stream- Range vegetation of a bottomland forest dominated by plains cottonwood along East Bijou Creek, an terrupted or intermittent (whichever it was) stream. This view across across (at a rough right angle to) the stream channel rather than a headon or inline view as in the two preceding photographs. Different age/size classes of trees were present showing this to be an uneven-aged population of plains cottonwood. There were relatively few cottonwood seedlings, but their number appeared adequate (though barely) to maintain the only tree species in this plant community.

Density and spatial arrangement (dispersion pattern) of cottonwood varied from closed canopy forest along the riparian zone to such wide (sparse) spacing of trees as to constitute a savanna on the outskirts of this bottomland range plant community. Cottonwoods grew (occurred) as individual mature adult trees or small groups of two or three trunks up small groves of eight to ten trees. Groves characteristically included pole-sized as well as fully grown trees. These larger groups were closer to the stream (generally in the riparian zone) and tree density declined with distance from the creek until the savanna spacing at outer border of the bottomland community. Dead wood at base of the adult cottonwood across the creek was mostly debris washed in and lodged against the trunk although a minor amount of this wood pile was branches from the crown of the tree that served as a barrier to water-borne material on this floodplain.

Although it was not obvious in this wide view there were three layers comprising the herbaceous understorey: tallgrass, midgrass, and shortgrass that were dominated by prairie sandreed, western wheatgrass, and buffalograss, respectively. Downy brome or cheatgrass was present (as it alway is) but it was minor on this range that was under superb management (see shortly below).

Forbs present included wild licorice, the most common and important; prairie coneflower or Mexican hat (Ratibida columnifera); showy milkweed (Asclepias speciosa); annual sunflower (Helianthus annus); four o'clock (Mirabilis linearis); prairie sagewort (Artemisia frigida); western ragweed (Ambrosia psilostachya) and kochia (Kochia scoparia). There were sparsely scattered plants of plains pricklypear (Opuntia polycantha).

Range condition class was Good. Cattle had most likely grazed this range for decades, certainly over the last several years to the present time as proven by dung piles of varying ages. In all these photographs there were cottonwood leaves and leaders within reach of cattle indicating that there had not been heavy browsing of of this species which has relatively high palatability to cattle. (High preference by cattle for Populus deltoides and the relative ease with which this palatable species is browsed out was described in detail below.) Presence of this palatable browse and fairly high abundance (author's subjective judgement based on cover, density, vigor, etc.) of prairie sandreed and prairie cordgrass along the creek was undeniable evidence that this range had not been abused (overgrazing/overbrowsing had not taken place) in the recent history of use by livestock. Accolades to the range manager.

East Bijou Creek, Elbert County, Colorado. Mid-July (estival.aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). No Kuchler unit for southern and southwestern cottonwood forest. No SRM. 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). Southwestern Tablelands-Piedmont Plains and Tablelands Ecosystem, 26e (Chapman et al., 2006).

33. On the higher and outer bank of the interrupted stream- View looking up the channel and along the bank of East Bijou Creek, an interrupted or intermittent (whichever it was) stream, in the semiarid Colorado Piedmont. From the physiogonomic and structural perspectives, an upper woody layer of plains cottonwood dominated this bottomland range vegetation. Tree density varied from that of a gallery forest in the riparian zones which remained wetter later into the summer to a savanna at outer margins of the floodplain plant community. This range community was, in essence, a mixed prairie with a tree element that varied greatly in tree density and dispersion pattern.

From perspective of annual biomass production (standing crop productivity) the dominant plant species of this range ecosystem was almost assuredly prairie sandreed. Western wheatgrass was the overall community associate. Switchgrass held this honor within the riparain zone in which the dominant herbaceous species was prairie cordgrass. Thus it was that a tallgrass species was the herbaceous dominant of both the wetland (riparian zone) and the greater floodplain.

Dominance of the greater floodplain (beginning at top of stream bank) by prairie sandreed and the close association of western wheatgrass, the major midgrass species, was depicted in this photograph. Buffalograss and blue grama were not conspicuously featured, but they were certainly well-represented. Mesic tallgrass species (notably prairie cordgrass and switchgrass) that dominated the lower stream channel were absent. Forb species included wild licorice, generally the most important; prairie coneflower of Mexican hat, showy milkweed, annual sunflower, kochia, prairie sagewort, and narrowleaf four o'clock. There were sparsely scattered plants of plains pricklypear, but nothing approaching density and cover to form even an interrupted shrub layer.

34. A savanna form on the outer boundary- At outside margins of plains cottonwood-dominated bottomland range vegetation the plant community was actually a savanna or, at best from a crown canopy criterion, a woodland rather than a forest. This lower stand density or wider tree spacing was in contrast to the gallery forest that developed in the riparian zone of this interrupted stream as was shown above.

Downed limbs immediately beneath cottonwood boles were a combination of those that fell from directly overhead off of senescing adult trees as well as those that had been carried by flood water to rest against the trunks. Such debris offered protection for the all-too-scant number of cottonwood seedlings available to replace their aging elders.

The understorey consisted of three distinct herbaceous layers of tallgrass, midgrass, and (to a lesser degree) shortgrass species, the dominants of whch were, respectively, prairie sandreed, western wheatgrass, and buffalograss (blue grama was less abundant having lower relative cover than buffalograss). Forbs included wild licorice, the most common and important; prairie coneflower or Mexican hat; showy milkweed, narrowleaf four o'clock, prairie sagewort, annual sunflower, and kochia.

35. Village meeting- Local assemblage of range plants on plains cottonwood-dominated bottomland range on floodplain of an interrupted or, possibly, an intermittent stream in the semiarid zone. Dominant plant species was prairie sandreed. Other range plant species included prairie sagewort, western ragweed, Mexican hat or prairie coneflower, and wild licorice. The second (vertical) slide was a closer-in, more detailed view of the left foreground of the first slide. This second photograph presented a better view of shoots of praiirie sandreed.

36. Atop the reed of a reed- The panicle--the entire structure--of prairie sandreed (first slide), upper or distal portion of this panicle (second slide), and one branch of this panicle at onset of anthesis (third slide).

37. Unfulfilled florets- Spikelets of prairie sandreed showing florets at pre-anthesis stage. There is only one floret per spikelet in this species and the upper glume is considerably longer than the lower (Shaw, 2008, p.90). Both of these features were presented in these two photographs.

38. Hanging lightly- Drooping leaders (woody shoots) with pendulous clusters of fruit on eastern cottonwood (Populus deltoides subsp. monilifera). The delta-shaped leaves, obvious basis of the specific epithet,.is a key identifying feture of this species. Vines (1960, p. 91) described leaf shape of P. deltoides as "broadly deltoid, margin crenate-serrate, apex abruptly acute or acuminate, base truncate to heart-shaped or abruptly cunete".

39 Lightly but effectively- Fruit (first slide) and individual seeds (second slide) of eastern cottonwood. Eastern cottonwood "wrote the book" on anemochory or aerochory, dispersion of plant disseminules or propagules by wind. Many prairie plants evolved (natural selection, survival of the fittest) anemochory as a means of effectively and efficiently dispersing pollen, but cottonwood went a step farther and has one of the best adaptations to dispersal of its fruit and seed via aerochory of any woody plant in North America. The liguliferous composites like thistles and dandelions have nothing on Populus species.

Eastern cottonwood is a dioecious species. Its fruit is a capsule, numerous of which are borne in racemes (first slide) and each of which consist of 10 to 30 seeds (second slide) with each of these being equipped with pubescence in the form of numerous, long, cottony fibers (Vines, 1960, p. 91). It is the cottony pubescence that permits cottonwood seeds to become air-borne so as to float and finally settle by gravity into every available "nook and cranny".

40. 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.

41. 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.

42. 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.

43. 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.

Northwestern and General Western Cottonwood Forest

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

49. 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.

51. 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 Ecopregion 10m (Environmental Protection Agency, undated).

52. 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).

53. 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).

54. 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).

61. 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 immediately preceding slide. Floodplain of Methow River, Okanogan County, Washington. June (early summer).

62. Cottonwoods nourish grass too- Stand of blue wildrye (Elymus glaucus) in understorey of Methow River bottom black cottonwood forest. In some parts of the herbaceous understorey of this forest range blue wildrye grew as a single-species stand whereas in other localized environments this domiant native grass was joined by Eurasian perennial grasses including smooth brome, Kentucky bluegrass, reed canarygrass, and intermediate wheatgrass, along with star Solomon's seal, the main range forb, and bush honeysuckle, dominant understorey shrub throughout much of the understorey.

Blue wildrye is a major woodland and forest species throughout much of the Sierra-Cascade Range and Pacific Slope, including the Coast Range. Blue wildrye was featured in several chapters of Range Types of North American because it is an important species in various rangeland and forest cover types.

A definitive description and taxonomic treatment of Elymus glaucus was provided in the encyclopedic Flora of North America-...Poaaceae, part 1 (Barkworth et al., 2007, ps. 306-309) to which readers were referred.

Okanogan County, Washington. June (early summer); phenological stage varied considerably across this small local stand. Most of the shoots were asexual (or at least pre-boot stage) whereas others were in the soft to mid-dough stage (see immediately below).

63. Spikes of blue wildrye- Two spikelets with maturing grain (mid-dough phenological stage) of blue wildrye in understorey of the black cottonwood forest community along Methow River described above. Most shoots in the stand of blue wildrye shown above were asexual (strictly vegetative).

64. Cottony seeds on black cottonwood- The fruit of members of the Salicaceae is a capsule which splits open at maturity releasing the comose ( adjective referring to a coma, a tuft or clump of hairs) seeds. The ripe, opened capsule with numerous seeds twas shown in the first of these photographs. Details of the comose seeds was presented in the second photograph. Corresponding details of bark were presented in the respective slides.

65. "For whom the bell tolls"- Physiogonomy of a lanceleaf cottonwood (Populus acuminata)-dominated floodplain forest along the Bell River with an understorey consisting of a shrub/small tree layer composed of mostly of Virginia chokecherry (Prunus virginiana) with some Wood's rose (Rosa woodsii). Limited herbaceous zone, but one layer consisting mostly of naturalized smooth brome (Bromus inermis) with some spreading dogbane (Apocynum androsaemifolium).

Cascade County, Montana. Late June. FRES No. 17 (Elm-Ash-Cottonwood), K-89 (Northern Flood-Plain Forest). SAF 235 (Cottonwood-Willow). Cotttonwood-Willow Series 222.21 of Plains and Great Basin Riparian Deciduous Forest 222.2 of Cold Temperate Swamp and Riparian Forest 222 (Brown et al., 1998, p. 43). Northwestern Glaciated Plains- North Central Brown Glaciated Plains Ecoregion 42o (Woods et al., 2002).

66. Trees along the Bell- Small trees of lanceleaf cottonwood and the much smaller (most plants of shrub category/size) Virginia chokecherry. Some Wood's rose. Dominant herbaceous species was naturalized smooth brome. Locallly spreading dogbane was the dominant herb; it was certainly the dominant forb overall.

Numerous seedlings and saplings of both major woody species were evident at this margin or perimeter of this gallery forest that was slowly advancing into a grassy area dominanted by smooth brome.

67. The shrub or middle layer- The shrub or middle layer of a lanceleaf cottonwood-dominated bottomland forest that developed along Bell River. This lower woody or shrub layer was co-dominated by young plants (seedlings to large saplings) of lanceleaf cottonwood and shrubs plus some small trees of Virginia chokecherry. Some cover of Wood's rose was also evident. This margin or perimeter of the gallery forest had much regeneration of these two co-cominant woody species.

If the habitat type of Daubenmire was applied this would be a lanceleaf cottonwood-Virginia chokecherry (Populus acuminata-Prunus virginiana) habitat type.

68. The herbaceous layer-A local opening (a forest gap created by falling of a lanceleaf cottonwood) supported a local stand of spreading dogbane, the native plant dominant of the herbaceous understorey (herbaceous layer) of a gallery forest along Bell River. This floodplain forest was an example of the lanceleaf cottonwood-Virginia chokecherry habitat type. Young plants of these two co-dominant woody species were growing around the spreading dogbane. A large--and still largely unvegetated--gravel bar of Bell River shone prominently behind this forest gap.

Details of this gravel br and its recent colonization by pioneering by the Very Intolerant cottonwood species was shown in two slides of the next slide-caption set.

69. Closing in on the Bell- A gravel bar or beach constituted the bank of Bell River. The raw parent material of this stream bank was being colonized by the Very Intolerant, pioneer tree species, lanceleaf cottonwood. Colonization by cottonwood was most obvious in the second of these two slides (though small cottonwood plants were visible at greater distance in the first slide). These "baby" lanceleaf cottonwoods were seedlings and not clonal offshoots from older cottonwoods.

The raw gravel of the stream bank appeared to have been the result of recent flooding and stream scouring that washed out and away already established lanceleaf cottonwoods such as those in background and at outer edge of the stream bank and riparian zone. This fresh "new land" or fresh sere for establishment of floodplain forest was created by the defoliating action of flood water. This is one form or kind of denucation the initiating process of plant succession (Weaver and Clements, 1938, chapter V). This stream erosion forming a new area on which vegettion development can proceed was an example of a topographic initial cause.

70. Geology at its root- Montana Department of Transportation hisstorical site marker that explained the geology of an area that strictly speaking was interpreted as part of the Northern Rocky Mountains (Fenneman, 1931, ps. 214-224 passim). This plaque discussed local areas such as Prickly Pear Canyon, Lyons Creek, Confederate Gulch, and Gates of the Rocky Mountains, this latter named by the Lewis and Clark Expedition. Most of the geologic material was sedimentary.

71. Outer appearnces- Three views of the physiogonomy as well as composition and structure of a grove of lanceleaf cottonwoods that developed on the floodplain of Dearborn River in the Northern Rocky Mountains. This bottomland forest had a structure consisting of three woody layers (zones) and, sporadically in local spots beneath openigs in the tree canopy, an herbaceous layer. Crowns of adult trees of lanceleaf cottonwood formed the upper woody or canopy layer. The middle woody layer was composed of younger trees--ranging from saplings to pole-sized shoots--of lanceleaf cottonwood and adult shrubs to small trees of Virginia chokecherry. The lower(est) woody layer consisted of seedlings and vegetative (asexual) shoots of lanceleaf plus smaller plants of Virginia chokecherry. The herbaceous layer consisted almost completely of smooth brome, an introduced, widely seeded, (and now naturalized), Eurasian perennial. There were highway right-of-way and pasture seedings of smooth brome all over this area.

Based on the Daubernmire (1952, 1968, 1984) method of classifying natural plant communities this was a lanceleaf cottonwood-Virginia chokecherry (Populus acuminata-Prunus virginiana) habitat type.

This floodplain forest was not open to livestock. Wildlife species were abundant throughout this general area (it was northern Montana afterall).

Floodplain of Dearborn River, Lewis & Clark County, Montana. Late June. FRES No. 17 (Elm-Ash-Cottonwood), K-89 (Northern Flood-Plain Forest). SAF 235 (Cottonwood-Willow). Cotttonwood-Willow Series 222.21 of Plains and Great Basin Riparian Deciduous Forest 222.2 of Cold Temperate Swamp and Riparian Forest 222 (Brown et al., 1998, p. 43).Middle Rockies- Eastern Divide Mountains Ecoregion 17aj (Woods et al., 2002).

72. Standing at the edge- Perimeter of a grove of lanceleaf cottonwood that developed on the floodplain of Dearborn River. A middle woody layer was present consisting of Virginia chokecherry and younger trees of lanceleaf cottonwood. Debris from past flooding was conspicuous. There was regeneration of cottonwood (probably both sexual and vegetative) as well as chokecherry.

The herbaceous (grassy) growth in foreground of the first slide was naturalized smooth brome, a widely planted, Eurasian perennial that has naturalized across much of central and northrn North America.

Floodplain of Dearborn River, Lewis & Clark County, Montana. Late June. FRES No. 17 (Elm-Ash-Cottonwood), K-89 (Northern Flood-Plain Forest). SAF 235 (Cottonwood-Willow). Cotttonwood-Willow Series 222.21 of Plains and Great Basin Riparian Deciduous Forest 222.2 of Cold Temperate Swamp and Riparian Forest 222 (Brown et al., 1998, p. 43). Middle Rockies- Eastern Divide Mountains Ecoregion 17aj (Woods et al., 2002).

73. Expansion of a grove- Regeneration of lanceleaf cottonwood at the outer edge or perimeter of a cottonwood grove resulting in expansion of the grove. Invasion of a pasture of smooth bromegrass through tree reproduction. Some of the smaller shoots of lanceleaf cottonwood were obviously seedlings, but it could not be determined definitively if all regeneration was sexual or, as appeared to be case, some was also asexual reproduction (cloning or development of offshoots). Offshoots are new modules or clonal units of existing or established trees. This asexual reproduction is proliferation of existing genotypes from, in this case, roots and/or rootstocks (woody rhizomes).

74. Inside a grove- Two views of the interior of a grove of lanceleaf cottonwoods with Virginia chokecherry and smaller trees (saplings to small poles) forming a middle woody layer while seedlings (and, probably, clonal offshoots) of cottonwood along with seedlings of chokecherry comprised a lower woody layer. There was sporadic or spotty cover of naturalized smooth brome that fomred an erratic herbaceous layer. Native herbaceous plants were not seen in the interior of this bottomland forest.

Wildlife of numerous species were present, but there was no livestock in this bottomland forest range.

75. Interior of a grove- Another brace of views of the interior of a grove of lanceleaf cottonwoods, a bottomland forest dominated by cottonwood with Virginia chokecherry as the dominant shrub. This was a Populus acuminata-Prunus virginiana habitat type based on the method of Daubenmire (1952, 1968, 1984).Sapling up to small pole-sized cottonwood and chokecherry comprised a middle woody or shrub layer beneath the canopy or crown layer of this forest range. A lower woody layer consisted of seedlings of chokecherry plus seedlings (and, probably. vegetative offshoots) of lanceleaf cottonwood. There was a discontinuous herbaceous layer consisting solely of smooth brome, an agronomic Eurasian perennial grss that became a naturalized range plant.

There were no livestock in this forest, but wildlife were plentiful in this part of Big Sky Country which was still east of the Continental Divide.

This was at the perimeter of a grove of lanceleaf cottonwood and Virginia chokecherry. The herbaceous in foreground of first slide was smooth brome, an introduced and widely naturalized Eurasian, cool-season perennial. Smooth brome has been widely planted throughout this area (as, for that matter, across huge portions of central and northern North America).

The grassy (herbaceous) understorey was primarily smooth brome, an introduced (and now naturalized) Eurasian perennial.

78. "Lost my top, heck; I'm growing it back!"- One or more of Nature's agents of defoliation (wind, snow and ice, lightening, etc) took off the crown of this lanceleaf cottonwood growing on the floodplain of Dearborn River in the Northern Rocky Mountain Region. With loss of apical meristem of the central shoot and consequent release from apical dominance (largely the influence of the auxin, indole acetic acid) a lower limb or, perhaps, intercalary meristem living lower on the shoot gained dominance and is in process of regrowing the crown on this young, actively growing, adult cottonwood. The new crown is developing from a lower, previously subordinate limb that was spared in the detopping disturbance or, again, perhaps it is of intercalary meristematic origin and developed from that source. At any rate, the replacement upper shoot is conspicuously smaller in diameter than that part of the trunk below point of breakage. The stump of the lost upper trunk and crown was visible to the right opposite the smaller, replacement limb.

Populus species are remarkably adapted to such severe defoliation having the ability to replace lost parts and, especially, to grown entire new shoots clonally (vegetatively, asexually). This phenomenon ws shown in the setion immediately below.

Taxonomic reminder: At various points in this treatment of bottomland cottonwood forest, students were reminded that positive identification of Populus species is problematic without having identifiabily distinctive plant organs and, even better, having these plant parts at various stages of phenological development. This includes plant organs like catkins (of both sexes), fruit, leaves, terminal buds, and bark (young and mature). Rangemen visiting a bottomland forest for a short period like a day or week typically do not have opportunities to examine all these organs (excepting short visits during short-lived, identifiably critical periods like plant flowering or fruit shedding). In absence of these short-time period features the way to obtain positive identification as to Populus species is by asking authorities who already know the species (because they were present during the important species-identifying phenomena like flowering or fruit drop).

Identification of Populus species is further confounded and made more difficult by presence of several Populus species--and hybridization among these species in addition to the ever-present integrading of taxonomic features (eg. as in leaves) between any two species. For example, P. acuminata, P. angustifolia, and P. deltoides have all been collected in the comparatively small Montana county of Wheatland (Lesica, 2013, ps. 175, 177), in which the following slides were taken along Musselshell River. In the adjoining Montana county of Sweetgrass P. angustifolia, P. balsamifera, P. deltoides, and P. tremuloides but not P. acuminata have been documented (Lesica, 2013, ps. 175, 177). Recorded absence of (= no record of presence of ) one Populus species from one relatively small county to another as shown in distribution maps is likely the artifact of limited collection activiety (or, perhaps, lost herbarium mounts or, even, loss of the species from that part of its species range by human activity like logging, damming rivers, or building roads) more than actual absence or absence from the pre-whiteman vegetation. This author is convinced tht he found P. acuminata in Sweetgrass county, Montana within a few miles of the Sweetgrass-Wheatland county line.

Upshot of all this is that your author did his best to get correct identification of Populus species, but there was the possibility that plants he interpreted as P. acuminata where actually variants of P. angustifolia or even P. deltoides. Afterall, P. acuminata is a natural hybrid of these two species (Crawford, 1974; Great Plains Flora Association, 1986, p. 276; Lesica, 2013, p. 175). It was also possible that all three of these Populus species were represented by a a few plants in the floodplain forests shown in this section.

79. From a distance- Esterior view of a wetland cottonwood-willow forest. This floodplain or riverbottom forest consisted of two prominent woody layers, a "sometime" (eradic or sporadic) lower woody layer, and a consistent herbaceous layer. Lanceleaf cottonwood and peachleaf willow (Salix amygdaloides) were co-dominants of the upper or crown ( forest canopy) layer while coyote or sandbar willow (Salix exigua= S. interior= Salix exigua ssp. interior) was the dominant of a middle woody or shrub layer with prickly or Say's rose (Rosa acicularis subsp. sayi= R. sayi) frequented associated with it. The herbaceous layer was composed primarily of co-domiants smooth brome(grass) (Bromus inermis) and quackgrass (Agropyron repens), two naturalized Eurasian grasses, and with the native, perennial American mannagrass (Glyceria grandis) as an associate to local dominant herbaceous species.

This wetland forest range developed on a backwater habitat of Musselshell River in central Montana. The greater part of this bottomland forest range was privately owned land that was grazed/browsed by beef cattle (Bos taurus) and white-tailed deer (Odocoileus virginianus). This area constituted a Range Management treatment of grazing/browsing by all herbivore species with both ruminant species being the major consumers (ie. all species-grazing treatment). Another part of this forest range was a small city park and an area adjacent to it in which there no livestock, but white-tailed deer were abundant. This served as a second de facto Range Management treatment with grazing/browsing limited to one ruminant species, the native cervid). A third area of this floodplain forest was a strip of highway right-of-way from which cattle were excluded and which deer (for whatever reasons) avoided. This third area (the smallest parcel) served as a de facto large animal exclusure (no grazing/browsing by ruminants or, for that matter, any ungulates).

Results of this de facto Range Management experiment (ie. a grazing/browsing trial) were shown and discussed below.

Floodplain of Musselshell River, Wheatland County, Montana. Late June. FRES No. 17 (Elm-Ash-Cottonwood), K-89 (Northern Flood-Plain Forest). SAF 235 (Cottonwood-Willow). Cotttonwood-Willow Series 222.21 of Plains and Great Basin Riparian Deciduous Forest 222.2 of Cold Temperate Swamp and Riparian Forest 222 (Brown et al., 1998, p. 43). Northwestern Great Plains- border of Judith Basin Grasslands and Unglaciated Montana High Plains 43m and 43o Ecoregions, respectively (Woods et al., 2002).

80. Critters in the woods- Two parts of an area of a cottonwood-willow bottomland forest open to grazing/browsing by cattle (cow-calf pairs) and white-tailed deer. All (or almost all) plants of lanceleaf cottonwood, peachleaf willow and coyote or sandbar willow had a high broswe line. Deer obviously browsed the three salicaecous species. Feeding on these species by beef cattle was less obvious given that cattle are, by preference, grass-feeders more than browsers (Holechek et al.2004, ps. 343-349), but this photographer watched (although he could not see close details) brood cows feed on plants of these three species on this part of the floodplain forest. There were hoof prints of cattle as well as those of deer in the mud or damp soil around salicaeous plants.

Note one black cow in left midground of both of these two slides. The cow in the second slide was behind the highly browsed clump of small cottonwood shoots in left foreground. These cows were as about as rank as the deer and did not let the author get close enough to get good slides of them feeding on willow or cottonwood (see the immediately following two slides), but this so-so cowboy and experienced observer of range animal behavior caught them in the act.

Browsing on lanceleaf cottonwood and willow species was without doubt a feeding phenomenon of both ruminant species.

The herbaceous layer of this forest range was made up mostly of the naturalized, Eurasian, perennial grasses of smooth brome and, away from the wetter soil, quackgrass.

81. High-headed (and headed for higher ground) in the woods- A brood cow grazing smooth brome in the herbaceous layer--floor--of a cottonwood-willow forest (first slide) and another cow that had been browsing on a clump of lanceleaf cottonwood and was startled then ran uphill as the author got close enough to get this sho (second slide)t. These slides were taken using a 28mm wide-angle Nikkor lens as the photographer was taking landscape shots. Hence, the cameraman was considerably closer to subjects than would be indicated otherwise in photographs. He was close enough to observe cattle browsing on woody species (eg. lanceleaf cottonwood in this case).

The herbaceous zone of this bottomland forest range consisted mostly of smooth brome and, away from the drier soil, quackgrass. Both of these species are naturalized, Eurasian, agronomic grasses. It was likely that the smooth bromegrass had been seeded on this land (whereas quackgrass grew as a weed species) and that cottonwoods and willows were invading (repossessing the land) on which they were potential natural vegetation. Potential natural vegetation on this forest site might (or might not) have been synonymous with climax vegetation depending on which theoretical perspective (eg. monoclimax, polyclimax, climax pattern theory) the range plant community was viewed from.

Lanceleaf cottonwood, peachleaf willow, and coyote (interior) willow were all overbrowsed as evidenced by highlining of every plant of these species to within feeding reach of deer and/or cattle. This excessive defoliation by cattle and deer had undeniably removed the option of browse from these species in future diets--at least in the near term. This overuse of the browse resource was shown in various slides below.

82. Herbivore overreach- Highlining of young plants (clumps of small saplings) of lanceleaf cottonwood by cattle and/or white-tailed deer on a botoomland cottonwood-willow forest in westcentral Montana. The author did not know the agricultural history of this land, but it supported a pasture of smooth brome that almost assuredly had been seeded (drilled) which would have required some degree of tillage. It seemed likely that there had been some mechanical removal of cottonwood (along with peachleaf and coyote/interior willow) from this land. These species were clearly invading or re-colonizing this forest site.

A floodplain forest dominated by these three woody species was probably the potential natural vegetation, with successional status (climax or seral) subject to interpretation depending on theoretical perspective. Clearly, smooth brome (a Eurasian, agronomic, forage grass) was not the natural vegetation. Smooth brome is, however, a naturalized range grass over a vast areaf of North America (it is on the Society for Range Management list of 200 plant species used for the International Intercollegiate Range Plant Identification Contest) so the brome was considered by the current author to be part of the man-modified vegettion of this floodplain forest range.

Overuse of the native cottonwood and willow species by livestock and/or wildlife was not sound or proper use management for this forest range. Impact of browse overutilization on cottonwood and willow populations was not determined, but it was self-evident that with this degree of overuse on established trees or shrubs there was very little survival of cottonwood or willow seedling or of small, vegetative offshoots. It was also obvious that the highly palatable browse of these species was no longer possible as part of range animal diets. Current management was not sound practice for plants or animals.

If this landowner wished to manage this forage crop as a monoculture of smooth brome for pasture and/or hay production proper tame pasture management would have to include brush control as these native woody species would, under tame pasture criteria, be brush that needed to be controlled (reduced to one degree of the other). As it was, what students saw here was neither sound range or pasture management. Given that the orientation of this publication is to management of ranges not tame pastures, the production of animal feed from native/naturalized vegetation and not domesticated or agronomic forages, the "management sin" shown here was of improper management or misuse of forest range. Specifically it was excessive defoliation of valuable browse plants-- by cattle and deer. Adjustments in management and use of the forest range were needed.

83. Textbook example- This view of the Musselshell River provided a classic example of a floodplain-complete with its floodplain forest dominated by lanceleaf cottonwood. The next two slide-caption sets showed and described the interior of this forest while the third slide-caption set (from here) presented the exterior of this forest.

84. Under a cottonwood canopy- Interior of a bottomland forest that developed on the floodplain of Musselshell River in westcentral Montana. This woody vegetation served as a forest range for beef cattle and white-tailed deer. In this part of the forest lanceleaf cottonwood (and perhaps some trees of eastern cottonwood [Populus deltoides] and/or narrowleaf cottonwood [P. angustifolia]) was the dominant species of both the canopy layer and the broken or sporadic lower woody layer. There were a few widely scattered trees of peachleaf willow present also. A continuous herbaceous layer of naturalizeed smooth bromegrass occurred throughout this forest range.

There was debris that included both 1) in-place fallen limbs and toppled trees and 2) flood-borne woody material brought in and deposited from upstream.

Yes, the first of these two slides, which was a beautiful slide (Provia 100F film), was turned into a piece of junk by an Epson Perfection 700 scanner (never buy Epson equipment), but your author kept it in because it showed that this cottonwood forest had developed next to waters edge of Musselshell River (right midground to right margin of slide). The second slide was of the forest vegetation immediately adjacent to vegetation attempted to be shown in the first slide and farther inside the forest and away from the river.

85. Living and dead in the interior- Inside of a lanceleaf cottonwood-dominated floodplain forest range grazed/browsed by beef cattle and white-tailed deer. This population of lanceleaf cottonwood. Note absence of regeneration of cottonwoods in this overbrowsed forest. Absence of tree regeneration indicated that this was a disfunctional forest ecosystem, at least in regards of ecosystem maintenance and renewal.

86. Unbrowsed edge-The perimeter or outer margin of a bottomland forest on the floodplain of Musselshell River in westcentral Montana. This floodplain forest was co-dominated by lanceleaf cottonwood (and probably with some trees of eastern cottonwood and/or narrowleaf cottonwood) and peachleaf willow (Salix amygdaloides). The forest vegetation presented here was on the right of way of a locally busy highway so that white-tailed deer did not tarry on it long enough to browse or graze any of thie forest vegetation. The highway right of way served as a de facto exclosure for both beef cattle and deer such that it served as a basis of comparison for the heavily browsed (by both cattle and deer) interior of the cottonwood forest that was presented above.

Absence of browsing was evident in the small saplings, most of which were lanceleaf cottonwood. These young trees had no evidence of vertebrate feeding (no removal of leaves or twigs by browsing deer or cattle).

The larger adult trees in background were either at peak adulthood or at the early senescent stage of their life cycles.

87. Basis of comparison- Two closer-in views of lanceleaf cottonwood (with a few peachleaf willow) at the edge of a bottomland forest growing on the right of way of a busy highway from which cattle had been fenced out and white-tailed deer avoided due to noise and frequency of vehicular traffic. Certainly deer undoubtedly crossed over this right of way and the highway, but they did not feed along the road--at least not enough to leave "sign" of browsing (visible evidence of defoliation of trees). Neither lower branches of adult trees nor leaves and twigs of seedlings and saplings were browsed by the resident herd of white-tailed deer that highlined saplings and ate most cottonwood and willow seedlings to the extent that there was very little regeneration of lanceleaf cottonwood or peachleaf willow in the area of forest that was subject to deer feeding.

Foliage (leaves and twigs) of lanceleaf cottonwood and the general abundance or crown cover of cottonwoods of all ages seen in these two "phototransects" taken inside the de facto exclosure served as basis for comparison to the overbrowsing evident in images in parts of the floodplain forest that were subjected to browsing (and overbrowsing at that) by both cattle and deer.

88. Big trees on a backwater- Three views showing physiogonomy, structure, and composition of a bottomland forest dominated by peachleaf willow (Salix that developed on a backwater of the Musselshell River. The lower woody layer of this forest was comprised of coyote or sandbar willow (Salix exigua= S. exigua var. exigua= S. interior) with sizable cover of prickly or Say's rose (Rosa acicularis subsp. sayi= R. sayi). There was an herbaceous understorey dominated by the naturalized Eurasian smooth brome, a widely introduced agronomic perennial. Couchgrass or quackgrass (Agropyron repens), another Eurasian perennial introduced as an agronomic forage crop, was the associate grass species. There was some cover of the native American mannagrass (Glyceria grandis). The locally major forb was the still yet another Eurasian species, but this one was an inadvertently brought-in weed known as houndstongue (Cynoglossum officinale) of the borage family (Boraginaceae).

These three slides presented progressively closer-in views of this forest range vegetation. The first slide gave an indiction as to size of peachleaf willow, adults of which are the size of neighboring adult cottonwoods. In this first slide the two trees in center midground were peachleaf willow while the tree to their right was a lanceleaf cottonwood. Coyote or sandbar willow was to left of the two peachleaf willow. Coyote or sandbar willow was present in the center of the second slide (including considerable dead growth) with Say's or prickly rose. coyote willow was obvious in foreground of the third slide.

Most of the largest trees in background of the second slide were also peachleaf willlow although there were also some lanceleaf cottonwoods. Largest trees in background of third slide were peachleaf willow. Your author never before this saw such large willows!

89. The lower woody layer plus some of the herbaceous zone- Three views at progressively closer camera distance of the shrub layer of a lanceleaf cottonwood-peachleaf willow bottomland forest that developed on the backwater floodplain of the Musselshell River in westcentral Montana. The three slides in the immediately preceding slide/caption unit presented the general view of this wetland range vegetation as to phisiogonomy, structure, and composition. The lower woody or shrub layer of this floodplain forest was featured in this current three-slide/caption set.

The first slide showed a wider-angle photograph of the general shrub layer of this backwater salicaceous forest with a large, adult peachleaf willow in center background and part of the crown of an adult lanceleaf cottonwood wasin right background (right rear margin) while there was a colony of the highly rhizomatous (woody rootstocks) coyote or sandbar willow accompanied by Say's olr prickly rose in foreground. The herbaceous layer (visible in both the first and, especially, the third slide consisted primarily of the agronomic Eurasian forage grasses, smooth bromegrass (the dominant) and quackgrass (the associate herbaceous species). There was also appreciable cover of American mannagrass, a native perennial grass which was probably the natural herbaceous dominant (and now this decreaser had probably overgrazed and largely displaced by the naturalized Eurasian perennials).

, sThe second slide included a young sapling of lanceleaf cottonwood in the middle of thicket of coyote or sandbar willow while adult lanceleaf cottonwoods and peachleaf willows loomed in the distant background.

The third slide showed coyote willow (Salix exigua var. exigua was the taxonomic variety in this wetland forest) with some shoots of Say's or prickly rose in front of an immense peachleaf willow. Most of the herbage in the lower layer (immediate foreground) was American mannagrass, the native (pre-whiteman) perennial, a decreaser species that was the potential natural dominant of the herbaceeous layer of this lanceleaf cottonwood-peachleaf willow riverine forest.

90. Two highlined browse species- Part of the colony of the thicket-forming (highly clonal) coyote willow (left margin of foreground in both slides) and a colony of young saplings of lanceleaf cottonwood (right half and right two-thirds, respectively, of the first and second slide) that developed along the backwater of the Musselshell River in westcentral Montana.

The featured topic of this photographic duo was the overbrowsing of these two salicaceous species by a resident herd of white-tailed deer. This area was part of a small town park from which livestock were excluded. The highlining on these two browse species was exclusively that of the deer. These two at-about-time-of-sundown slides showed the extent to which white-tailed deer can overbrowse their feed resource. The absence of cattle from this area of a lanceleaf cottonwood-peachleaf willow forest in contrast to a contiguous part of this riverine forest grazed/browsed by both ruminant species allowed a photographic comparison of dual- versus single-species browsing on cottonwood and willow species.

Another contiuous (conterminous) area of this floodplain forest was a highway right of way from which both cattle and white-tailed deer were effectively excluded (deer avoided the noisy traffic of the highway). This constituted a de facto exclosure and served as a control area of no browsing by any large vertebrate. This browsing-excluded part of the bottomland was presented above and contrasted with areas of this riverine forest on which there was almost no regeneration of cottonwood or willow species. A combination of cattle and deer browsing had prevented tree reproduction, and by both sexual and asexual means.

These two "phototransects" of highlining (overbrowsing to height that deer could reach) that were highlighted by light of the setting sun showed students that any range animal can overuse its feed source. Just because range animals are wildlife or native species does not mean that they will not exhaust their food supply. Obviously man had a determinative influence on native as well as domestic grazers and browsers in this floodplain forest. The question naturally arose, "What kept white-tailed deer from overbrowsing before arrival of man (whit or red)?" Or, alternatively, "Did deer overbrowse even before humans arrived in North America?" What--if any--was the role of predators? Of course, man is the ultimate predator.

The dioecious Salix species are a "bearcat", a "rabid bitch", of a species when it comes to identification. They are on par with (or worse than) Carex and Crataegus species in this regard. Positive, I-have-no-doubt-whatsoever identifiction requires flowers and fruit, and twigs with terminal buds along with lots of leaves do not hurt a thing. The definitive--though not necessarily correct or even the best--reference for North American Salix species is volume of Flora of North America. Certainly the various flora or manuals are also definitive (as definitive as humanly possibly). For instance, Lesica (2012, ps. 177-185) listed 33 Salix species for Montana The Great Plains Flora Association (1986, ps. 281-291) included 21 species for a region that included most (not all) of the Great PlainsThe old standby Gray's Manual of Botany (Fernald, 1950, ps. 487-519) is always helpful and, more recently, Flora Neomexicana (Allred and Ivey, 2013, ps. 531-535) for the Land of Enchantment should prove useful for wide-ranging species like S. exigua and S. amygdaloides.

91. A willow the size of a cottonwood- Textbook example (even if an Epson scanner did over-expose and "crystalize" it) of an immense peachleaf willow (Salix amygdaloides) growing on a subirrigated, backwater wetland of the Mussellshell River in in the Judith Basin of westcentral Montana. The smaller, highly rhizomatious sandbar or coyote willow (S. exigua) and colony forming lanceleaf cottonwood (Populus acuminata) grew in association with peachleaf willow to form the native woody vegetation of this riverine wetland. The comparatively great size of peachleaf willow was a major part what awed this rangeman about this range type.

Peachleaf willow has received good treatment in the native plant literature. Some recommended sources were

92. Peachy branches- Drooping outer leaders (woody branches) of peachleaf willow at early bloom stage thriving in/on a backwater wetland (on the greater floodplain) of the Mussellshell River in southcentral Montana.

93. Peachy trunks- Two huge peachleaf willow trees with the multi-trunked habit characteristc of this species. These thing were not just impressive; they were damn impressive. Two key morphological features that distinguish trees from shrubs are: 1) height (shrubs are generally defined as being less than 15 feet tall) and 2) multiple trunks or boles in shrubs. Adult peachleaf willow in this neck of the woods are too damn big for shrubs, but multiple shoots were typical. These rascals are just plain doggone big for "willers".

Backwater wetland on floodplain of Mussellshell River in in the Judith Basin of westcentral Montana. (Wheatland County). Mid-June; mid-bloom phenological stage.

94. Peachy leaves and blooms- Leaders (woody branches) of a female peachleaf willow that grew on the low bank of a backwater slough of Mussellshell River in Species in Salicaceae, the willow or cottonwood family, are dioecious with separate male and female plants. These pistillate (female) catkins, with the characteristic namesake peach leaf-like leaves, were produced by a large multi-shoot tree that was quite obviously was a "girl tree" (and a big girl at that).

A lengthy section devoted to sandbar or coyote willow (Salix exigua= S. interior): S exigua is one of the more widespread, densely populated, and, thus, more important of the Salix species in the Western Range Region. The biological range of S. exigua is vast extending from Baja California to British Columbia and Alaska eastward to Quebec and New Brunswick south to Maryland and westward to Oklahoma. Given this huge species (biological) range it is not surpising that this species carries numerous common names including streambank willow, basket willow, narrowleaf willow, slender willow, and gray willow. Of course, some of these common names are applied to other Salix species as well.

A most important morphological feature of S. esigua from a Range Management, Wildlife Management-Fisheries, Watershed Manageament, and Forestry standpoint is its highly clonal (due to stolons and rootstocks), colony forming nature. Overbrowsing of willow species by these animals is a major, a serious problem from thewse professional standpoints and relative to conservation of renewable natural resources. That subject was dealt with throughout this section. Certainly the colony forming sandbar, coyote, or narrowleaf willow is no exception. Dayton (1931, p. 16) described S. exigua as being relished by livestock. Your author has seen vast populations of this species browsed out by everything from horses to elk to beaver--and on private and public lands (National Wildlife Refuges and National Parks are some of the worse cases as matter of fact).

Good sources for sandbar or coyote willow included the "usual suspects" such as Forest Service (1940, B141), Vines (1963, ps. 101-102), Stephens (1973, ps. 38-41), Francis (2004, ps. 664-667) and Many, if not most (almost all) of the wilows are palatable to all kinds of livestock and ungulate and rodent species. Overbrowsing of these valuabale browse plants was a major source of concern expressed throughout this chapter.

95. Sunset on sandbar (willow, that is)- Staminate leaders (= male branches) of sandbar or coyote willow on a clonal tree on a subirrigated backwater slough of the Mussellshell River in the Judith Basin of westcentral Montana. These shoots were lite aglow by the setting sun. (Another godsend: your phytophographer barely got there in time for these rather neat shots. "Guide Me O' Thou Great Jehova".)

96. Bears some resemblance to coyote's tail- Staminate (=male) catkins of coyote or sandbar willow along a slough of Mussellshell River in the Judith Basin (Northern Great Plains) in westcentral Montana.

97. Smooth 'nough for gals legs, but these were boy shanks- Lower woody shoots (trunks, boles) of a male clonal plant along a slough of the Mussellshell River in the Judith Basin (Northern Great Plains) in westcentral Montana.

98. Time for the girls- Clonal unit or module (first slide) and pistillate leaders (= female branches) (second slide) of female coyote or sandbar willow. These shoots were produced along a small stream in the Judith Basin of the Northern Great Plains in westcentral Montana.

99. Bearing much fruit on a sandbar- Female leaders (first or upper slide) and pistillate catkins with immature capsules (second or lower slide) of sandbar or coyote wilow in the Judith Basin of the Northern Great Plains in westcentral Montana.

100. Ladies, right; gents, left- Male and female plants--and therefore male and female shoots with their catkins/fruit (left and right, respectively)--grew far enough apart that a close side-by-side comparison or flowers and/or fruit (capsules) was not possible in this setting. Details of male and female flowers and clusters of capsules were shown immediately above. If viewers know how to get these two images through a server such as Google, differences can be seen readily (the same as when projected in a slide show which is still the "gold standard" for audience viewing of photographic images).

The following short piece was devoted to lanceleaf cottonwood (Populus acuminata), the third member of the Salicaceae that dominated and defined a primarily woody plant wetland in the Judith Basin in the Northern Great Plains of westcentral Montana.

The Populus species present an identification and taxonomic challenge to the plant systematist, rangeman, wildlifer, and causal observer. No, Populus is not as difficult as Salix because the former has fewer species than the latter. Nonetheless, positive identifiction as to Populus species is frequently difficult (and sometimes impossible when flowers and fruits are lacking). Hybridization between Populus species is a major confounding problem. In this instance, P. acuminata is thought to be a natural hybrid between P. deltoides and P. angustifolia (Stephens, 1973, p. 60; Great Plains Flora Association, 1986, p. 276) shown in taxonomic treatments (eg. Lesica, 2013, p. 175) as Populus x acuminata Rydb. Crawford (1974) donfirmed this hybrid based on morphological and phytochemical criteria.

For treatment of the cottonwoods this author still prefers Sargent (1933, ps. 128-129 for P. acuminata). P. acuminata was not included in Silvics of North America (Burns and Hankala, 1990).

101. A highlined clone- A clonal unit of young (immature) shoots of lanceleaf cottonwood (Populus acuminata) growing along a slough of the Mussellshell River in the Judith Basin of westcentral Montana. The high browse line was the result of feeding by a resident herd of white-tailed deer (Odocoileus virginianus). It was an example of the phenomenon known as highlining, overbrowsing (excessive defoliation of browse material) by browsing animals. This area was either in or immediately adjacent to a small city park and no livestock were in this area. It served as a de facto Range Management treatment (browsing by ruminants was limited to a single, native cervid species). There was a conterminous range that was privately owned land in which both this herd of white-tailed deer and beef cattle (Bos taurus) browsed willow and cottonwood species. This constituted a second Range Manageament treatment. Finally there was a third area, a streatch of highway right-of-way, from which cattle were excluded and which the deer (for whatever reasons) avoided.

Height of highlining (overbrowsing) exceeded five feet. The herbaceous understorey was primarily American mannagrass (Glyceria grandis).

102. A lanced grove- Grove of lanceleaf cottonwood with an obvious clonal unit or module of five shoots (first slide) and a view of upper trunks and branches of the clonal unit (second slide) tht grew on the floodplain of the Mussellshell River in the Judith Basin of westcentral Montana.

103. Lanced leader- A leader (branch) of lanceleaf cottonwood on a tree that grew along a slough in the floodplain of Musssellshell River in the Judith Basin of westecntral Montana.

104. Roses amid the willows- Plant of prickly or Say's rose (Rosa acicularis subsp. sayi= R. sayi) growing with sandbar or coyote willow on a backwater of the Musselshell River. There are a number of wild rose species across the Western Range Region. Lesica (2012, ps. 284-285) listed only five Rosa species for Montana, but Dayton (1931, ps. 46-49) discussed 12 Rosa species as important browse plants across the western range. He noted that Say's rose has thicker leaves and is less palatable but more abundant and, thus, an important browse plant (Dayton, 1931, p. 49).

Floodplain of Musselshell River, Wheatland County, Montana. Mid-June; peak-bloom stage of phenology.

105. Prickly parts in the willows- Blooming shoots (first slide) and a sinlge flower of prickly or Say's rose growing with coyote or sandbar willow on a backwater wetland of the Musselshell River in central Montana.

Floodplain of Musselshell River, Wheatland County, Montana. Mid-June; peak-bloom stage of phenology.

106. Manna in Montana- American mannagrass (Glyceria grandis) growing in a community of willow and along a backwater drainage of the Musselshell River in southcentral Montana. American mannagrass was not common in this wwetland range community, but it certainly was an indicator plant, given that American mannagrass is a climax and decreaser grass species on this range type/site.

Wheatland County, Montana. Mid-June (late spring); milk- to soft-dough grain stage.

107. Essence of manna- Sexual shoots and panicles of American mannagrass growing in a backwater slough in floodplain of Musselshell River. The obvious resenblance of these spikelets and the panicle overall to that of domestic rice (Oryza sativa) tells the astute rangeman or agronomist that this species is a member of the rice subfamily, Oryzoideae.

Wheatland County, Montana. Mid-June (late spring); milk- to soft-dough grain stage.

108. Beauty ruined by Epson- All the king's horses men, material, etc. plus the best of Adobe Photoshop could not restorea nearly flawless slide of this local colony of dame's rocket (Hesperis matronalis ) on a floodplain of the Musselshell River southcentral Montana. This member of the mustard family (Cruciferae or Brassicaceae) is one of the largest and most showy of all forbs in this "neck of the river", but as might be expected for anything this large it was introduced from Eurasia by the white man for an ornamental and then proceeded to naturalize (Great Plains Flora Association, 1986, p. 315) so as to produce robust specimens like the local colony seen here.

Wheatland County, Montana. Mid-June; peak-bloom stage (but you'd never know it following Epson scanning).

109. Some for the dame of the river- Sexual shoots (first or upper dslide) and close-in view of flower clusters and flowers (second or lower slide) of dame's rocket in a sprawling colony that grew on floodplain of the Musselshell River in southcentral Monatana. This cricifer (member of the Crucifereae) is an introduced ornamental that has naturalized (Great Plains Flora Association, 1986, p. 315). As yet it is not a troublesome weed, but do'nt hold your breath.

Another form of the general cottonwood-willow range type in the Northern Great Plains was a wetland range plant community that was more of a shrubland than forest because it consisted primarily of moisture-loving shrubs including coyote or sandbar willow, red-ozier dogwood or, sometimes, red-osier cornel (Cornus stolonifer= C. sericea ssp. sericea), goldern current (Ribes aureum), western snowberry (Symphoricarpos occidentalis), and silverberry or wolf willow (Elaegnus communtata) yet with scattered large trees of narrowleaf cottonwood (Popoulus angustifolia). Kentucky bluegrass (Poa pratensis) and smooth brome (two, naturalized, Eurasian, perennial grass species) formed a scattered or sporadic herbaceous layer.

This natural range vegetation had large cottonwood trees as a major component of the wetland range community and, therefore, something of an arborescent physiogonomy. Given this similarity of range vegetation, plus the desirability of keeping range plant communities of similar composition and structure together, this shrubland with a tree component was retained in this chapter along with the lanceleaf cottonwood-peachleaf willow-coyote willow-grass community covered immediately above.

This natural wetland community was also included in the Shrubland chapter entitled Willow & Riparian Types. This was redundant coverage, but 1) it was more consistent in treatment of very similar range types (and range types that developed in close proximity to each other) and 2) was make more convenient for viewers to find similar (especially from standpoint of species composition even if physiogonomy and community structure differed).

110. Willows below the cottonwoods- Riparian vegetation along a small, slow-moving stream in the Northern Great Plains that consisted of the overwhelming dominant of coyote or sandbar willow (Salxi exigua ssp. interior) along with red-osier cornel (Cornus stolonifer= C. sericea ssp. sericea), silverberry or wolf willow (Elaegnus communtata), western snowberry (Symphoricarpos occidentalis), and goldern current (Ribes aureum) with a widely scattered tree layer of narrowleaf cottonwood (Popoulus angustifolia). There was a "hit-or-miss", "here-and-there" (sporadic) herbaceous layer comprised of Kentucky bluegrass and smooth bromegrass (two introduced and now naturalized Eurasian perennial species).

These two photographic views were used to try to impart that structure and composition of this wetland range plant community that was extremely valuable for watershed protetion and wildlife habitat, particularly as browse for cervid species and beaver.

Small, slow stream, Sweetgrass County, Montana. Late June. FRES No. 17 (Elm-Ash-Cottonwood), K-89 (Northern Flood-Plain Forest). SAF 235 (Cottonwood-Willow). Cotttonwood-Willow Series 222.21 of Plains and Great Basin Riparian Deciduous Forest 222.2 of Cold Temperate Swamp and Riparian Forest 222 (Brown et al., 1998, p. 43). Northwestern Great Plains- Unglaciated Montana High Plains 43o Ecoregion. (Woods et al., 2002).

111. Thicket in middle of nowhere- A thicket of coyote or sandbar willow, red-osier dogwood, silverberry or wolf willow, and golden current plus scattered old and senescing trees of narrowleaf cottonwood made up this riparian vegetation that developed along a small stream in the Northerrn Great Plains. An herbaceous layer comprised of Kentucky bluegrass and smooth brome occurred intermittantly in both the shrub thicket and beneath the aging/dying cottonwoods.

This wetland range vegetation was beside a recently abandoned, unpaved road. A newly built, high, six-wire fence excluded (at least up to time of these photographs) deer and cattle so that plants of all riparian species had produced profuse growth along the creek bed. Overutilized shoots of willow and dogwood along with old cow chips (cattle dung) indicated overbrowsing (assurredly by white-tailed deer, that were abundant in the immediate area, as well as by cattle). Recovery of plants protected (by the new barbed wire fence) from overbrowsing had been remarkable.

Rapid recovery of riparian vegetation from overbrowsing/overgrazing after initiation of proper grazing management has been a commonly observed (and welcomed) response. Typical abundance of soil moisture in riparian zones permits recovery (including essential, increased reproduction) of plant life and restored protection of straams and their watersheds.

112. New lease on life- A "dirt" (ungraveled, unpaved) road ran along what remained of a grove of ancient narrowleaf cottonwood trees. The road (apparently previously maintained by the county) had recently been abandoned (decommissioned is the proper description or official designation) and fenced off. Immediately (as measured on Nature's temporal scale) root sprouts or clonal offshoots from the obviously senescing (naturally dying of "old age") narrowleaf cottonwoods sprang up in the abandoned roadbed. This phenomenon was shown in the foreground of these two slidces. visible. Of course what actually took place is that these root suckers, which are new shoots of the aging cottonwoods, had been trying to grow for decades but grading of the county "dirt" road kept cutting them off. With decommissioning and ceased maintenance of the county road the new shoots were able to continue to grow (and, if allowed to continue their growth, hopefully establish "new trees"). This continued growth, including reproduction, would be recovery of plant life and, ultimately, restoration of the natural vegetation. This is the same "healing process" as recovery of coyote willows from overbrowsing by ruminants.

A more physiologically explicit description of cottonwood recovery is that the "old, dying trees" are only mature or adult shoots that are senescing because they have completed their life cycle. The natural life span of these shoots of a genetic tree (a genotype or genetically unique individual) has been reached and now those shoots are nearing the end of their life cycle. In other words, what are seen as cottonwood "trees" are actually just a one generation, the current generation, of shoots that have now matured, reproduced (both sexually by flowering/fruit production and asexually by cloning offshoots), and are dying. The next generation of shoots (the new asexual or vegetative crop of shoots), as seen in the offshoots in the old road bed, has grown from roots and/or rootstocks (woody rhizomes) and is replacing the senescing (dying of "old age") shoots of the last generation. Both of these generations (and perhaps one or several more before this one) are "trees" (shoots) of the original genetic individuals (genotypes) of narrowleaf cottonwood. This is asexual reproduction or vegetative regeneration.

Note that there are shoots of various ages in this local grove of cottonwood shoots. There was a pole-sized young shoot seen as the right-most "tree" in the first of these two slides. In the second slide there are several shoots or, perhaps, genetically unique trees (genotypes) that are smaller (presumedly, younger) than the largest tree which had a largely dead crown (this senesced crown was visible in the first slide).

Actual age of trees (genotypic individuals) has to be measured as, and understood to be, generations of grown-reproduced-and-died shoots es . True age of any given tree (a genotype of narrowleaf cottonwood) is known only by God. Life span of an actual tree might extend back for centuries or even millenia. Again, "God only knows". What we do know is that it is up to man "to dress the garden". Adam can browse it out, cut it off, pave it over, or let it grow and wisely harvest the crop. "One generation passeth and another generation cometh, but the Earth abidith forever" (Ecclesiastes 1:4).

113. Whole new life- Offshoots of narrowleaf cottonwood emerging from a recently abandoned roadbed. These clumps of clonal shoots or modules that grew (sprouted) from roots and/or rhizomes are vegetative reproduction of senescing adult shoots seen as "dying old trees". These sprouts are the next (the most recent) genertion of a single or of several genetic narrowleaf cottonwoods (genotypes of narrowleaf cottonwood).

Leaves and twigs of Populus species are palatable to most vertebrate herbivores, including cattle. If this area, from which cattle and deer were currently fenced off (excluded) from, was overbroswed there might not be enough offshoots remaining to insure survival of these narrowleaf cottonwoods. Some cottonwood shoots could be browsed and used as a feed source(they are a renewable resource), but the proper stocking rate (manifest as proper degree of use) and proper season of use would have to be assured to achieve both proper use of browse and management of the watershed. Proper management of the watershed begins with protection of soil againsts accelerated erosion. Start with stream bank stabilization through maintenance of riparian vegetation.

The bare soil and absence of shade made this abandoned roadbed a very likely spot for sexual reproduction in addition to the asexual reproduction seen here. All Populus species are rated as Very Intolerant, meaning that they are least likely to regenerate or establish beneath older or adult tree cover. Instead, cottonwoods are pioneer species or "first colonizers" on "new land" (recently bared soil that is free of plants that could outcompete the Very Intolerant cottonwoods). Abandoned fields and roads, cutover forests, old school yards, and so on are the "prime habitat" of cottonwoods like those featured here. This is even more the case for sexual than asexual reproduction: seedlings are at even more of a competitive disadvantage than offshoots that receive some "support" from existing tree shoots. Had this rangeman returned the next year it was likely that he could have photographed seedlings of narrowleaf cottonwoods. Unfortunately that was not to be the case so viewers will have to use their imagination for that slide.

Phreatophyte- " a plant that derives it water supply from groundwater and is more or less independent of precipitation" (Helms, 1998). More speifically phreatophytes (literally "wel plants" from the Greek) have enough of their roots in the capillary fringe (the continually wet zone immediately above an aquifer) that can survive where they otherwise could not. This is one form of natural subirrigation.

Narrowleaf cottonwoods presented in this section were examples of phretophytes. In this case it was most likely that the phreatophytic source was lateral or outward flow of water from the stream. Even though these narrowleaf cottonwoods were not growing on the stream bank they were, in actuality, part of the riparian vegetation.

The first of these three slides showed new shoots of coyote or sandbar willow growing from the abandoned roadbed (foreground) and a thicket of older willlow shoots on the other side of (behind) the recently decommissioned county road (background). The second or vertical slide showed this same comparison though at closer distance with new or current season's shoots at edge of roadbed (foreground) and older, larger willow shoots immediately adjacent to the roadbed (background). The third slide presented (or attempted to present after scanning--and marring--by an Epson Perfection 700 scanner) this comparison (at an oblique angle) with current season's willow shoots in the abandoned road (foreground) and taller, older shoots of several year's growth to the immediate side of the road (background).

This example of prolific production of new willow shoots by asexual or vegetative reproduction showed why (and how) abused riparian vegetation is capable of phenomenally rapid restoration under desirable or, even more so as was the case here, ideal conditions. Sandbar or coyote willow is an extremely important species for streambank bank stabilization (Francis, 2004, p. 665).

Older shoots of sandbar or coyote willow in this same dense population were flowering; some shoots even had immature capsules (the willow fruit type). Staminate catkins and capsules on these shoots were presented above. They are designated in the captions as being in Sweetgrass County, Montana (Late June).

The main ecological role of silveberry would appear to be erosion control, especially in the riparian zone of streams, and cover for certain wildlife species (the thicket-forming feature of silverberry certainly provides an invincible covert for animals that can get into it).

Location Note: Another example of coyote, sandbar, or basket willow-dominated wetlands--a riparian community that was both cattail marsh and willow carr along the Shields River in southcentral Montana--was presented and described in the Shrublands chapter entitled, Willow & Riparian Types. There is a continuum of range plant communities comprised of or dominated by willow and cottonwood species that varies from riparian, wetland scrub to floodplain forests. Given this natural variation (quite an extreme range) in range vegetation it was unavoidable that there some confusion regrading location or placement of range types and thus difficulty for viewers in "navigating" such wetland range types. Northwestern Great Plains- Unglaciated Montana High Plains 43o Ecoregion. (Woods et al., 2002).

120. Another variation- Riparian vegetation formed a narrow gallery foresst along the Gila River Fremont cottonwood (Populus fremontii) and coyote or sandbar willow (Salix exigua) which together constitute a habitat type by the method of Daubenmire, ). Also present as an associate tree species was the beautiful Arizona sycamore (Platanus wrightii). Box elder (Accer negundo) was present at small cover so as to be a minor tree species. There was limited understorey, but Chihuahuan brickellbush (Brickellia floribunda) was the principal shrub besides sandbar or coyote willow. Blue grama (Bouteloua gracilis) was the major herbaceous species.

The Gila River is one of the Southwest's most beautiful rivers (even when dry as seen here). As of this writing the Gila is still free-flowing throughout its entire length in New Mexico. On-going attempts to build diversion(s) on this stream have met with stiff and well-organized political opposition reaching to all levels of government from local, state, and federal. Such a river war is understandable and justifiable and, in this author's judgment, must be won in favor of keeping the Gila wild and free-flowing. If in any way these images help save Gila River from short-sighted, pork barrel destruction it will have been well worth this small effort.

Long live Gila River. Personal: the Land of Enchantment is one of your author's favorite states. This fondness is because of New Mexico's various natural treasures like Gila River. Keep New Mexico New Mexico; preserve Gila River.

Grant County, New Mexico. Late June; estival aspect. FRES No. 17 (Elm-Ash-Cottonwood), K-None. SAF 235 (Cottonwood-Willow). Mixed Broadleaf Series 223.22 of Southwestern Riparian Deciduous Forest and Woodland 223.2 of Warm Temperate Swamp and Riparian Forest 223 (Brown et al., 1998, p. 43).Chihuhuan Deserts- Chihuhuan Desert Grasslands Ecoregion 24b (Griffith et al., 2006).

121. Still yet another variation on the same theme- A gallery forest of Fremont cottonwood and Arizona walnut, Arizona black walnut, river walnut, or mountain walnut, (Juglans major) with a sporadic understorey dominataed by rubber rabbitbrush (Chrysothamnus nauseosus) that developed along a stream in the vegetational mosaic of semidesertgrasslands and the Chihuhuan Desert in southwestern New Mexico.

This form of a more seric or less mesic form of cottonwood bottomland is found as forest patches "here and there" throughout the greater Chihuhuan Desert Region. It was included here to show the diversity of cottonwood forests in westrn North America.

Hildago County, New Mexico. Late June; estival aspect. FRES No. 17 (Elm-Ash-Cottonwood), K-None. SAF 235 (Cottonwood-Willow). Mixed Broadleaf Series 223.22 of Southwestern Riparian Deciduous Forest and Woodland 223.2 of Warm Temperate Swamp and Riparian Forest 223 (Brown et al., 1998, p. 43). Madrean Archipelago- Apachian Valleys and Low Hills Ecoregion 79a (Griffith et al., 2006).

Almost in Arizona- Leaves and fruit (nuts) of Arizona walnut Arizona black walnut, or mountain walnut on one of the trees shown in the preceding image of a gallery forest in the Chihuhuan Desert. Good sources for basic discussion of this valuable, native nut species included (Sargent, 1933, ps. 172-173; Little, 1950, p. 42; Vines, 1960, ps. 125-126).

Miscellaneous Forest Types- IA

Sundry Bottomland Forest Types

Pecan (Carya illinoinenisis)

Green Ash (Fraxinus pennsylvanica)

Floodplain Forests

Eastern Cottonwood (Populus deltoides)

Northwestern and General Western Cottonwood Forest

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