Miscellaneous Forest Types-I

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Sundry Bottomland Forest Types
 
   
 

28 .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).

 
   
 

29. 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.  
   

 

30. 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).

 
   
 

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

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

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

 
   
 

32. 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).

 
   
 

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

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

 
   

 

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

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

 
   
 

35. Pecan timber- The forest form of pecan bottoms was represented by this stand of midddle-aged trees on the first terrace of a small creek floodplain in the patchwork of range plant communities where there is an intermixing of Western Cross Timbers and Grand Prairie in northcentral Texas. All trees were pecan whose tall straight trunks supported such woody vines as mustang grape, fiddleleaf or fringed greenbriar, and Alabama supplejack. Dominant understorey shrub was Carolina buckthorn. Most woody ground cover was the invasive, alien Japanese honeysuckle (Lonicera japonica), a treadful, horrid, etc. (adjectives fall short of vile description) woody weed. Major grasses included Canada wildrye, purpletop, Johnsongrass, and Texas wintergrass in that order. Major forbs were Baldwin ironweed (Vernonia baldwinii), and Indian plantain (Cacalia plantaginea).

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

 
   

 

36. Bottomland pecan forest- Higher stocking of pecan trees on bottomland produces a forest in contrast to the savanna oar open woodland form of this range type as shown above. All trees in these two photographs were pecan. Dense stocking resulted in tall, relatively straight boles in marked contradistinction from the widely spreading crowns of pecans with more spacing among trees. Pecans with stocking rates represented by forest vegetation presented in these two photographs produce wood and not fruit. This is the forest community of a lumberman not an orchardman, and the form of vegetation that develops in absence of fire.

There was a well-developed woody vine layer that extended from ground level to tops of forest canopy which was comprised of mustang grape, rattanvine or Alabama supplejack, and fiddleleaf greenbriar. Carolina buckthorn grew as a smaller tree or larger shrub in the lower woody layer. The herbaceous layers included tallgrass species like Canada wildrye, purpletop, and the naturalized Johnsongrass along with composite forbs the two most common of which were Baldwin ironweed and Indian plantain. Much of the lower layer was composed of the invsive (and apparantely naturalized) Japanese honeysuckle.

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

 
   
 

37. Woody understorey of pecan bottom- From ground level and extending upward just shy of tree branching the range vegetation of a bottomland pecan forest was comprised of three woody vine species (mustang grape, fiddleleaf greenbriar, and Alabama supplejack) and the shrub or small tree, Carolina buckthorn. The wsidespread exotic invader, Japanese honeysuckle, was absent from this photoplot. The herbaceous portion of this forest vegetation visible in the background was presented in the next photograph.

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

 
   
 

38. Herbaceous layer (s) of a bottomland pecan forest- In addition to the woody vines of mustang grape, fiddleleaf greenbrier, and Alabama subblejack or rattanvine in right foreground and background of this photoquadrant important forbs of the herbaceous component included Indian plantain (left corner of foreground) and scattered smaller, apparently shade-stunted Baldwin ironweed. Many of the small green shoots with forb-like leaves were new sprouts of greenbrier and supplejack.

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

 
   
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.

 
   
 

39. 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).

 
   
 

40. 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).

 
   

 

41. 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).

 
   

 

42. 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).

 
   

 

43. 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.

 
   

 

44. 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.

 
   
   
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.

 
   
 
45. 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).  
   
 

46. 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).

 
   

 

47. Semiarid gallery forest- Exterior view of a riparian zone forest of Rio Grande cottonwood (Populus deltoides ssp. wislizenii), netleaf hackberry (Celtis reticulata), Mexican walnut (Juglans major), and lanceleaf sumac (Rhus lanceolata) At outer (least mesic) edge of this gallery forest some of the more moisture-loving grasses formed a thick sward. The large cespitose grass was Wright's sacaton (Sporobolus wrightii) the dominant understorey species; Canada wildrye was the major associate species. The most common forb was horehound (Marrubium vulgare), a naturalized Eurasian weed.

This range vegetation was in the Trans-Pecos Basin and Range vegetational area, the west Texas portion of the Chihuhuan Desert Region. It was in Chihuhuan Deserts 24 (ecoregion level III), Chihuhuan Montane Woodlands 24d (ecoregion level IV) based on Griffith et al. (2004). It was an example of mesic forest in a semiarid climate. FRES No 17 (Elm-Ash-Cottonwood Forest Ecosystem). No aqppropriate Kuchler unit for southern and.or southwestern cottonwood gallery forest. SAF 63 (Cottonwood). Populus deltoides Association (if and when such is recognized), Cottonwoow-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Trans-Pecos Basin & Range Vegetational Area: Chihuhuan Deserts- Chihuhuan Montane Woodlands Ecoregion 24d (Griffith et al., 2004).

June, summer aspect. Calamity Creek, Jeff Davis County, Texas.

 
   

 

48. 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.

 
   

 
49. 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.  
   
   

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

 

51. 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.

 
   
 

52. 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.

 
   
 

53. 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.

 
   
 

54. Understorey of a cottonwood-willow bottomland forest- A lower woody layer of regenerating plains cottonwood and willow, an herbaceous layer of grasses and grass-like plants, and a multi-storied liana layer of greenbriar (Smilax sp.) and grape (Vitis sp.) added to the mature tree layer such that this range plant community formed a true forest structure on a wide expanse of river floodplain. Understorey graminoids included bottomland switchgrass, Canada wildrye, Johnsongrass, sedges, spike rushes, rushes, and cattail. This was another of the widespread and naturalized Johnsongrass. The most common forb was wild or American licorice.

Proper range management with emphasis on light and periodic grazing/browsing (vs. the widespread practice of continuous heavy grazing) permitted perpetuation of this forest. This was an outstanding example of good stewardship and sustainability of range resources.

FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). No Kuchler unit for southern and southwestern cottonwood and/or willow gallery forest. SAF 63 (Cottonwood). Populus deltoides association(if and when such is recognized), Cottonwood-Willow Series in Plains and Great Basin Riparian Deciduous Forest biotic community of Brown et al. (1998). Rolling Red Plains Vegetational Area: Central Great Plains- Red Prairie Ecoregion 27h (Griffith et al., 2004).

June, summer aspect. Flood plain, North Fork of Red River. Wheeler County, Texas.

 
   

 

55. 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).

 
   
 

56. 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.

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).

 
   
 

57. 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).

 
   
 

58. 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.

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).

 
   
 

59. Along a dry reach of the interupted stream- Range vegetation of a bottomland forest dominated by plains cottonwood along East Bijou Creek, an interrupted 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).

 
   
 

60. 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.

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).

 
   

 

61. 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.

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).

 
   

 

62. 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.

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).

 
   

 

63. 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).

East Bijou Creek, Elbert County, Colorado. Mid-July; initiation of anthesis.

 
   

 

64. 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.

East Bijou Creek, Elbert County, Colorado. Mid-July; initiation of anthesis.

 
   

 

65. 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".

Larimer County, Colorado. Mid-June (late spring), seed-shatter stage.

 
   

 

66. 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".

Larimer County, Colorado. Mid-June (late spring), seed-shatter stage.

 
   

 

67. Palatable resprouts- Root sprouts (suckers) from a felled, senescing eastern cottonwood on a prairie in the western Ozark Plateau. The three sets of root shoots presented here (first and second photographs) were two-year-old suckers or heterophyllous shoots produced from lateral roots (third photograph) of a relatively old, "half-rotten" eastern cottonwood after it was felled for fuel wood. The shoots, some of which in these photographs were over eight feet in height, had been completely defoliated (100% of leaves removed) by commercial beef cattle (cows and calves of mixed breeding) in both the first and second years of shoot development. The small leaves present on shoots at time of photographs were current-season's regrowth (early part of current--second--warm growing season).

Several (a high proprotion) of cottonwood root suckers were broken off by browsing cattle so that this extreme or severe degree of use included both eating of leaves and upper twigs of current growing season as well as complete breaking off of many root suckers that were the production of two growing seasons. Several of the broken shoots and branches were visible in some of the photographs presented here and immediately below. Cattle defoliation in the first growing season was heavy, but not total as in the second season of growth. In the second growing season many of the first and current year's root shoots were broken off at their point of origin on the lateral roots. This was done by cattle who "rode down" the root suckers in order to reach the uppermost leaves and terminal buds (ie. cattle ate all leaves and much of the upper parts of shoots, terminal twigs, plus broke off entire shoots by the browsing technique of "riding down").

"Riding down" is the feeding behavior whereby browsing animals reach up as high as they can on plants and then commence walking forward so that forage or browse plants bend down beneath the animals' weight and pass progressively beneath the feeding animals' brisket and belly until the animals have fed as far as the plant will bend to within reach of the animals' mouth. In the process, some of the brittle shoots and branches are broken off while the more limber shoots spring back-- minus leaves, fruit, buds, twigs, etc. that were consumed by the animals. In this feeding pattern animals can remove plant material at much greater heights than they could otherwise reach by merely streatching their necks and extending their tongues.

Herbaceous plant species growing adjacent to and beneath eastern cottonwood shoots was mostly the introduced pasture grass, tall fescue (Festuca arundinaceae), which has naturalized across most of the greater Ozark Region and had self-seeded onto this former tallgrass prairie. The second-most common herbaceous species on this local habitat was Japanese chess or Japanese brome (Bromus japonicus), a naturalized Eurasian annual grass. It appeared that none of the herbage of these two non-native grasses had been eaten by cattle although considerable quantities of biomass had been trampled. Cattle seemingly had quite a browsing bout on this spot as quantities of dung were left. There was no evidence of browsing by deer, rodents or rabbits.

Total herbivory (=herbivorous action) that did and would continue to influence regeneration of eastern cottonwood on this pasture included at least four components: 1) complete removal of leaves from cottonwood shoots, 2) total elimination of some cottonwood shoots by breakage at ground level (ie. total loss of some two-year old trees or two growing seasons of plant regeneration), 3) feeding selectivity that severely injured or setback cottonwood while at the same time having much less defoliation impact on annual and perennial grasses (both non-natives), and 4) deposition of dung with importation of more potential plant nutrients than was recycled or otherwise generated on this local environment.

Question to the "village idiot" (so that he can be "above average"): "will grasses or cottonwood derive most benefit for plant growth and/or survival from the combined influences of cattle feeding?"

Eastern cottonwood (both eastern and plains varieties), like most other members of the Salicaceae, readily coppices or resprouts following major injury or defoliation. Fowells (1965, ps. 516, 521) and Burns and Honkala (1990, vol. 2, ps. 533, 539) described vegetative reproduction in eastern cottonwood. Eastern cottonwood sprouts vigerously from both stumps and roots. In the pattern of most sprouting species, vegetative (asexual) reproduction declines with increaseing age of trees while vegetative propagation, both natural and atrtificial, is vigerous in young plants. Fowells (1965, p. 516) and Burns and Honkala (1990, vol. 2, ps. 533) cited research which found that eastern cottonwood (P. deltoides var. deltoides) stump-sprouted up to ages of 25 years.

Heterophyllous shoots (root sprouts or suckers) shown here were from a much older tree. Over half of the trunk (beyond the center growth rings) of this eastern cottonwood had rotted away so that it was impossible to accurately determine tree age. (Rotting appeared to have begun around an old fire scar.) Recollections by the author and his conversations with elderly observers firmly established age of this felled tree in excess of 80 years. Root suckering began in early spring following sawing of the old tree in the preceding winter.

On various occasions this author has observed resprouting by various kinds of cottonwood (Populus spp.) in larger and older tindividual trees following sawing, bulldozing, f lood breakage, etc. Obviously resprouting in cottonwood is not limited to trees that are younger than a quarter century.

It was also obvious that regeneration of eastern cottonwood depends upon protection from overbrowsing. It was possible that some of the herbivory described above benefitted (or had potential to benefit) eastern cottonwood. Dunging provided extra plant nutrients, especially nitrogen, available to surviving cottonwood shoots. Dung might also benefit grass proportionately more than cottonwood so that increased competition from grass resulting from soil fertilization was detrimental to cottonwood. It was conceivable that breakage of some cottonwood root sprouts (suckers) had the effect of beneficial prunning which would enable surviving shoots to grow larger faster and become too large for cattle to "ride down" and thereby escape most further browsing impacts (other than continued dunging if and when cattle rested in shade of growing cottonwoods).

Overall, however, and up to the point of time recorded here, even the village idiot made 100% of this pop quiz by explaining (in simple talk) that the browsing shown here was excessive and could result in death of resprouts and ultimate loss of eastern cottonwood from this pasture.

Ottawa County, Oklahoma. June.

 
   

 

68. Overutilization of palatable eastern cottonwood by cattle- Excessive (extreme or severe) defoliation of two-year old root suckers of eastern cottonwood on a prairie in the Ozark Plateau. Beef cattle (crossbred cows and calves) removed all leaves and many shoot tips (including developing buds) from these heterophyllous shoots by early to mid-spring of the second growing season. This necessitated production and development of a new set of leaves (the leaves present in these and preceding photographs). Numerous branches were broken from shoots that survived "rding down" browsing by cattle, and lots of the two-year old shoots (root suckers) did not survive the aggressive feeding by cattle.

This was an example of overuse which, if continued, might well result in killing and loss of eastern cottonwood from this pasture. These slides demonstrated the importance of protection of young eastern cottonwoods that are extremely palatable (even to animal species that are grazers more than browsers) from excessive browsing. Regeneration of eastern or plains cottonwood such as that shown above in a plains cottonwood-willow bottomland forest would not have been possible with continued overuse at the severity shown here and in the three immediately preceding slides. If degree(s) of use showh here continued for just a few more years (number unknown; would vary depending on many factors) the eastern cottonwood will die out on this pasture. The old felled cottonwood tree described in the preceding caption will be dead afterall.

The main lesson from these photographs was the concept of proper degree of use (the first and most important of the Four Cardinal Principles of Range Management), often expressed as Proper Use Factors. Secondary lessons from these slides involved range animal behavior, including both the phenomenon of "riding down" and feeding selectivity. A degree of defoliation approaching "anililation" of eastern cottonwood coupled with non-eating of tall fescue and Japanese brome (though with some trampling and covering with dung of these grasses) provided a "textbook example" of feeding selectivity, the threeway interaction of animal preferences for certain plant species, relative palatability of different plant species, and availability of herbage and browse of certain plant species.

Ottawa County, Oklahoma. June.

 
   
 

69. It'll do its part and its best, but eastern cottonwood cannot rise from the dead indefinitely- Close-up view of one of the severely browsed shoots that suckered from roots of a felled eastern cottonwood described above. All leaves are regrowth that replaced earlier full-grown leaves that were eaten by browsing beef cattle. These were some shoots that survived extreme degrees of browsing. Many shoots did not survive. (This exterme defoliation might be expected from overpopulations of browsers like deer, but this was from just "plain ole cows".)

If the rangeman takes care of the feed resource it will take care of him. If this cattleman wanted natural replacement of a nice shade tree a drastic change of management was in order. A quicky, makeshift exclosure of wovn wire would protect these young shoots until they could grown beyond reach of the cattle. If this cattleman had the goal of elimination of eastern cottonwood from this pasture he had a good start toward that objective. In other words, if eastern cottonwood was regarded as a brush species (a woody noxious plant) and a single-species stand of tall fescue was the desired result then cattle browsing appeared to be a very effective method of biological control to reduce (perhaps totally eliminate) this woody species from this grazing (or haying) unit of land. As it happened such was not the case. This cattle-owner was simply running cattle for tax purposes and social status (ie."dodging taxes" and "bragging rights").

This photographic case study was used to illustrate how excessive degree of use and/or general imporper animal browsing can prevent or greatly reduce regeneration of eastern cottonwood. Degree of use on eastern cottonwood would almost certainly have been much less if cattle-grazing had been restricted to late autumn through winter-early spring when cottonwood was dormant and nutritive value and palatability of the naturalized cool-season grasses was higher.

Ottawa County, Oklahoma. June.

 
   
 

70. 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
 
There are various forest range types dominated by different Populus species that are important in the Western Range. Included below are some examples of these. They are probably more important for wildlife than livestock range. In fact, many are easily injured by excessive concentrations of animals which is usually more apt to occur with livestock than big game. These cottonwood-dominated forests are primarily floodplain or, at least, bottomland range plant communities such that they are extremely valuable for watershed protection and flood control.  
   
 

71.   Bottomland (floodplain) cottonwood forest- Exterior (physiogonomic) view of a local forest community dominated by narrowleaf and black cottonwood (Populus angustifolia, P. trichocarpa, respectively). A deciduous forest with an herbaceous understory of introduced Eurasian grasses developed along this alluvial bar of the Gallatin River.

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

Gallatin County, Montana. FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem).  K-89 (Northern Flood-Plain Forest). SAF 235 (Cottonwood-Willow). Cottonwood-Willow Series (222.51) in Rocky Mountain Riparian Deciduous Forest biotic community (222.5) of Brown et al. (1998, p. 43). Middle Rockies- Townsend Basin Ecoregion, 17w (Woods et al., 2002).

 
   
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73.   Branches, leaves, and inflorescences of narrowleaf cottonwood- Utah County, Utah. June.  
   
 
74. Black cottonwood (Poplar trichocarpa)- This specimen of nearly giantic size was an associate to western redcedar (Thuja plicata) at edge of a grove of the latter species. Glacier National Park, Montana. July.  
   

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

 
76. 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.  
   
 

77. Cottonwoods and water- Outer edge or perimeter of a black cottonwood with an herbaceous understorey along the Methow River in the Okanogan Valley at base of the Northern Cascades foothills. Basin wildrye dominated the rangeland just outside of the black cottonwood floodplain forest.

Okanogan County, Washington. June (early summer). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem).  K-89 (Northern Flood-Plain Forest). SAF 235 (Cottonwood-Willow). Cascade-Sierran Riparian Deciduous Forest (222.4)- Cottonwood-Willow Series (222.41) of Brown et al. 1998, p. 43). Columbia Plateau- Okanogan Valley Ecoregion, 10m (Environmental Protection Agency, undated).
 
   
 

78. Black cottonwood river bottom forest- Physiogonomy, structure, and species composition of a black cottonwood forest that developed on the floodplain of Methow River. Black cottonwood constituted the sole tree species and tree (upper) layer of this forest range community. Larger saplings and pole-size young trees of black cottonwood made up a sporadic or interrupted lower tree or upper shrub layer. The lowest layer of vascular plants was composed of both herbaceous and shrub species with species composition and structure of this layer varying considerably based on forest gaps from fallen or felled trees, the latter by beaver (Castor canadensis). There were colonies of blue wildrye, the dominant native grass, that ranged from "pure" (ie.single-species) stands to local stands mixed with introduced and naturalized Eurasian perennial grasses including smooth brome, Kentucky bluegrass, reed canarygrass (Phlaris arundinaceae) and intermediate wheatgrass (Agropyron intermedium) and star Solomon's seal, the main range forb. Bush honeysuckle, black twinberry, or black bearberry (Lonicera involucrata) was the dominant understorey shrub throughout much of this understorey although common snowberry (Symphoricarpos alba) was a local dominant in better-lite microsites. Western virgin's bower or western clematis (Clematis ligusticifolia) united the lower and uppermost layer of this forest range vegetation by twining its way to the cottonwood canopy.

There were scattered ponderosa pines along outermost edges of the cottonwood forest, but pines did not penetrate into the cottonwood-dominated community where lack of light was a (probably, the) major limiting factor that excluded ponerosa pine. Felling of cottonwood (mostly mid-size trees) by beaver appeared to be the major phenomenon that permitted establishment of ponderosa pine around perimeter of the black cottonwood forest. There was no Douglas-fir, period.

The larger log (far left) was a black cottonwood casualty to wind; the smaller log of black cottonwood (center foreground) fell to hungry, industrous beaver.

Okanogan County, Washington. June (early summer). FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem).  K-89 (Northern Flood-Plain Forest). SAF 235 (Cottonwood-Willow). Cascade-Sierran Riparian Deciduous Forest (222.4)- Cottonwood-Willow Series (222.41) of Brown et al. 1998, p. 43). Columbia Plateau- Okanogan Valley Ecoregion, 10m (Environmental Protection Agency, undated).
 
   
 

79. 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).
 
   
 

80. 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).
 
   

 

81. 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).
 
   
 

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

Okanogan County, Washington. June (early summer).

 
   

 

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

Okanogan County, Washington. June (early summer). Obvious full-bloom stage.

 
   
 

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

Okanogan National Forest, Okanogan County, Washington. June (early summer).

 
   

 

85. Bush honeysuckle blooming - Shoot of bush honeysuckle at peak bloom (first photograph) and details of bush honeysuckle infloresecence (second photograph). These examples were in the colony of bush honeysuckle or black twinberry in understorey of a mid-elevation Douglas-fir-Engelmann spruce forest in the Northern Cascades.

Okanogan National Forest, Okanogan County, Washington. June (early summer).

 
   

 

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

Floodplain of Methow River, Okanogan County, Washington. June (early summer).

 
   
 

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

Okanogan County, Washington. June (early summer).

 
   

 
88. 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).  
   
 

89. 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).

 
   

 

90. 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).

Okanogan County, Washington. June (early summer).

 
   

 

91. 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.

Okanogan County, Washington. June (early summer).

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

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

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

 
   
Niobrara River
 

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

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

 
   
 

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

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

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

 
   

 

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

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

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

 
   
 

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

Cherry County, Nebraska. Late June.

 
   

 

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

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

 
   

 

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

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

 
   
Some Kansas Rivers
 
   

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

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

 
   

 

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

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

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

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

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

 
   

 

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

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

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

 
   

 

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

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

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

 
   

 

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

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

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

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

 
   

 

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

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

 
   
 

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

Ottawa County, Oklahoma. July.

 
   
 

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

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

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

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

 
   
 

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

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

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

 

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

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

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

 
   

 

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

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

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

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

 
   
 

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

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

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

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

 
   
 

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

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

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

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

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

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

 
   
 

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

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

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

 
   

 

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

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

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

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

 
   

 

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

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

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

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

 
   
 

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

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

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

 
   

 

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

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

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

 
   
 

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

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

 
   
 

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

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

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

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

 
   
 

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

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

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

 
   

 

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

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

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

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

 
 
   

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

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

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

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

 
   
 

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

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

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

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

 
   

 

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

Newton County, Missouri.. Late April.

 
   

 

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

Newton County, Missouri.. Late April.

 
   

 

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

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

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

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

 
   

 

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

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

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

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

 
   
 

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

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

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

 
   
 

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

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

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

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

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

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

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

 
   

 

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

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

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

 
   

 

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

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

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

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

 
   

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

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

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

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

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

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

 
   
 

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

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

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

 
   
 

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

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

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

Osage County, Kansas. Mid-June.

 
   

 

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

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

 
   

 

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

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

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

 
   
Bosque River- A Texas Example of a Subhumid Zone River
 
   

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

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

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

 
   

 

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

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

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

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

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

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

Bosqu;e River, Erath County, Texas. Late January.

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

133. Bosque River in spring under the heavy hand of European man- Riparian zone and adjacent mixed hardwood bottomland forest on bank and adjacent floodplain of Bosque River in a typical first third of spring. Bosque River is to some degree a dysfunctional stream due to a large flood control system of earthen dams and "conservation lakes" (large ponds or reserviors) that effectively catch and retain much of the overland flow of water that would otherwise drain into the main stem of this stream. Even in one of the most soil-regolith moisture-rich seasons (spring rains combined with low evapotranspiration) stream flow in Bosque River was reduced to a shallow, slow-moving current (note the numerous thin, flat rocks extending out of the water in the middle of the channel). Woody debris was proof that this river still floods, but flood waters in all but the heaviest, hardest rainstorms are confined to banks of this water-starved stream. What, if any, impact this man-modified (unnatural), restricted stream flow has had on riparian and forest vegetation was unknown.

The riparian vegetation zone was primarily herbaceous and of two major communities: 1) climax Canada wildrye (Elymus canadensis)-broadleaf woodoats (Uniola latifolia) and 2) seral populations of giant ragweed (Ambrosia trifida). The former were larger in area and developed less-disturbed microsites whereas the latter occupied smaller-area microhabitats of freshly disturbed soil (generally spots along the stream bank that were scoured by flood waters).

The bottomland forest was a sugarberry (Celtis laevigata)-cedar elm (Ulmus crassifolia)-pecan (Carya illinoinsis) climax with associate tree species including eastern cottonwood, black walnut (Juglands nigra), bois d'arc (Maclura pomifera), and red mulberry (Morus rubra). Major shrubs in this portion of the floodplain forest included green-briar (Smilax bona-nox), mustang grape (Vitis mustangensis), Virniginia creeper (Parthenocissus quinquefolia), trumpet creeper (Campsis radicans), and rusty black-haw (Viburnum rufidulum). The highly invasive exotic, Japnese honeysuckle (Lonicera japonica) was another important shrub. In addition to co-dominants Canada wildrye and broadleaf woodoats, other major grasses included (Bromus purgans= B pubescens), purpletop (Tridens flavus), sand lovegrass (Eragrostis trichodes), sideoats grama (Bouteloua curtipendula), three-flower melic (Melica nitens) ), Jophnsongrass (Sorghum halepense), common sandbur (Cenchrus incertus), hairy crabgrass (Digitaria sanguinalis), and the two, naturalized, Eurasian annual bromes, Japanese chess (Bromus japonicus) and downy brome or cheatgrass (B. tectorum), and the naturalized, South American, annual brome called rescuegrass (B. unioloides= B. catharticuas). Major forbs in addition to giant ragweed included the naturalized Eurasian umbel, hedge parsley (Torilis nodosa),.dominant cool-season forb, frostweed or white crownbeard (Verbesina virginica), dominant warm-season forb, pokeberry (Phytolacca americana), pigeon-berry (Rivina humilis), and Louisiana sage- or mugwort (Artemisia ludoviciana subsp. mexicana).

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

 
   

 

134. Grassy banks- Along many stretches of Bosque River where there are gently sloping (lower rather than higher) stream banks that permitted development of a relatively wide riparian zone. On this narrow zone (a linear microsite) the range plant community was an overwhelmingly herbaceous one dominated overwhelmingly by Canada wildrye and broadleaf woodoats. There sere short patches of the riparian zone that had been created by rushing flood waters that scoured out bare areas the freshest (most reacent) of which supported colonies of giant ragweed whle "scour scars" made a few seasons previously were taken over by Johnsongrass. Annual ryegrass (Lolium multiflorum), Japanese brome, and hedge parsley (naturalized Eurasian species) were also growing on disturbed patches of the stream bank. Overall, however, Canada wildrye and broadleaf woodoats called all the important shots in the herbaceous riparian zone.

A few seedlings and small saplings of pecan and red mulberry had also established on the stream bank even within the grassy riparian zone. Two larger seedlings of red mulberry were visible in both of these slides, especially in the second one. A large seedling of pecan was visible in the first photograph to the left of the large "lanky" tree at far right. This mid-sized, adult tree was also a pecan as was the short snag in the lower left corner that was accompanied by a common green-briar and a mustang grape. In the second slide the immense, leaning tree on the far bank (center background) was one of several old-growth (and nearly ancient) pecan. Thus in these two slides pecn was represented by age/size classes ranging from seedling to senescing adults. Throughout this climax mixed hardwood forest range pecan was most frequently second to sugrberry and cedar elm as a dominant, but pecan was one of the dominant tree species. Keep that thought as you scroll to the next pair of photographs.

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

 
   
 

135. Vernal vestiture- Dry Branch in spring. In the wet season the seasonally wet Dry Branch, an ephemeral stream into Bosque River, had its bank covered by Canada wildrye and broadleaf woodlats, the dominants of the herbaceous layer, and green briar (Smilax bona-nox) and roughleaf dogwood (Cornus drummondii), dominant shrubs. Associate shrubs were poison oak (Rhus toxicendron = Toxicodendron radicans) and trumpet creeper (Campestris radicans) A locally dominant, but overall minor herbaceous species was downy wood brome (Bromus pubescens = B. nottowayanus). Quite a bit of this native festucoid grass was in foreground.

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

 
   

 

136. Higher on the banks- Farther up on the banks of Bosque River just above the herbaceous riparian zone shown in the last two photgraphs Canada wildrye and broadleaf woodoats remained co-dominants of the herbaceous layer but they were joined by greater cover and density of trees, especially pecan. Other tree species--most numerous of which were sugarberry and cedar elm--became "thicker" (greater canopy cover and trunk density) still farther away from the riparian zone (eg. trees in the backgouund of these two slides).

Pecan commonly formed the leading or outer edge edge of woody range vegetation along Bosque River. The "hemongous" specimen of pecan featured in these two slides (each from a different vantage point) served as a fine example of this phenomenon. Some of the splendid specimens of old-growth pecan like this one have limbs almost as large as their massive trunk. Ain't redwoods, but they're damned impressive. And they certainly serve as a host for more species of vascular plants such as woody climbers like mustang grape, trumpet creeper, green-briar, and coral honeysuckle all of which found this stately pecan a convenient and comfortable source of physical support. (Need it be mentioned that pecan is the State Tree of the Lone Star State? While on the subject of size, viewers should take note of the dimensions of that mustang grape. If Texas ever had a "State Vine" mustang grape more than likely would be it. No, there is not a longhorn grape.)

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

 
   

 

137. Down on the Bosque- Heavy rains and high water blessed a climax elm-sugarberry-pecan forest in the bottomland of the Bosque River in northcentral Texas. Trees in these two "photo-plots" were two pecans at far left, two cedar elm in center, and an American elm at distant right (right background). Much of the river bank vegetation consisted of giant ragweed (Ambrosia trifida) where recurrent flowing water cut or deposited fresh alluvium. Farther back from areas affected by water flow the dominant herbaceous species was made up of Canada wildrye (Elymus canadensis) and broadleaf woodoats (Uniola latifolia), climax decreaser grasses. Major shrubs were common greenbrian (Smilax bona-nox) and mustang grape (Vitis mustangensis). Smooth sumac (Rhus glabra) was common in some habitats.

The short stretch of Bosque River presented in these two "photographic dendrograms" had higher stream banks and a less pronounced riparian zone than in stretches shown above. Here the climax mixed hardwood forest extended into the stream channel so that there was not a treeless, herbaceous (mostly decreaser grasses) developing away from water's edge.

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

 
   
 

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

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

 
   

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

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

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

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

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

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

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

 

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

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

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

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

 
   

 

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

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

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

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

 
   

 

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

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

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

 
   

 

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

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

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

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

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

 
   

 

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

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

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

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

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

 
   

 

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

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

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

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

 
   

 

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

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

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

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

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

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

 
   

 

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

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

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

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

 
   

 

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

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

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

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

 
   
 

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

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

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

 
   

 

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

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

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

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

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

 
   

 

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

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

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

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

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

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

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

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

Erath County, Texas. Late April.

 
   

 

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

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

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

 
   

 

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

Stay tuned.

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

 
   

 

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

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

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

Erath County, Texas. Mid-March.

 
   

 

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

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

Erath County, Texas. Mid-March.

 
   
 

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

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

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

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

 
   

 

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

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

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

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

 
   

 

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

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

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

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

 
   
 

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

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

 
   

 

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

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

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

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

 
   
Concluding note with regards snags, fallen limbs and branches, and downed logs- The Society of American Foresters recognized value of "dead wood" remaining in forests as part of these forests' biological legacy, "an organism, a reproductive portion of an organism, or a biologically derived structure or pattern inherited from a previous ecosystem" including "large trees, snags, and down logs left after harvesting to provide regugia and to structurally enrich the new stand" (Helms, 1998). Biological legacy is a more recent form of Frederic Clements' reaction whereby preceding organisms facilitate (hence, later workers referred to this as facilitation) establishment, survival, reproduction, etc. of later (more successionally advanced) species, stages, communities, etc. Such successional goods and services (eg. addition of soil organic matter, habitat for reducers or decomposers like fungi and microorganisms, nesting cavities for vertebrates, shelter for seedlings of vascular plants) were described in the short section mmediately above in regards to dead pecan wood in form of snags and downed logs.  
   

 

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

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

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

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

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

 

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

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

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

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

 
   
 

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

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

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

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

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

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

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

 
   
 

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

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

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

 
   

 

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

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

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

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

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

 
   
 

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

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

 
   
 

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

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

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

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

 
   
 

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

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

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

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

 
   

 

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

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

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

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

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

 
   

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

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

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

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

 
   
 

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

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

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

 
   

 

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

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

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

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

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

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

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

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

 

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

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

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

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

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

 

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

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

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

 
   

 

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

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

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

 
   

 

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

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

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

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

 
   

 

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

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

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

 
   
 

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

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

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

 
   

 

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

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

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

 
   
 

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

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

 
   

 

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

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

 
   
 

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

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

 
   

 

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

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

 
   

 

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

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

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

 
   
 

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

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

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

 

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

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

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

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

 
   

 

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

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

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

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

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

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

 
   

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

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

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

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

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

 
   

 

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

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

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

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

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

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

 
   
 

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

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

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

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

This one photograph provided pictoral evidence of the same.

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

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

 
   
 

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

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

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

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

 
   

 

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

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

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

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

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

 
   
 

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

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

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

 
   

 

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

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

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

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

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

 
   
 

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

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

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

 
   

 

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

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

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

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

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

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

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

 
   

 

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

Erath County, Texas. Early November.

 
   
 

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

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

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

 
   

 

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

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

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

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

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

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

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

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

 

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

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

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

 
   
 

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

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

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

 
   

 

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

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

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

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

 
   

 

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

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

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

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

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

 
   

 

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

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

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

 
   

 

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

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

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

 
   

 

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

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

 
   

 

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

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

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

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

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

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

 
   
 

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

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

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

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

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

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

 
   
 

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

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

 
   

 

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

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

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

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

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

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

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

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

 
   

 

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

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

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

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

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

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

 
   

 

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

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

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

 
   

 

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

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

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

 
   
 

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

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

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

 
   
 

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

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

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

 
   
 

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

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

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

 
   
 

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

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

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

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

 
   
 

222. Newly arrived pioneer- A seedling of eastern cottonwood becoming established in a higher part of the stream bed of Bosque River. This plant had germinated and emerged sometime during the previous growing season and overwintered in the dormant stage. Leaves for the current (this) year had already emerged from terminal (preformed) buds during late winter as this photograph was taken on the official first day of spring (vernal equinox). This was another example that at this latitude (approximately 32 degrees north) warm-season species start leaf growth before spring. It was specified above that this set of photographs depicted range vegetation in the period that was transition from late winter to early spring.

Eastern cottonwood is an Intolerant tree species (Wenger, 1984; Burns and Honkala, 1990) and a pioneer or initial colonizer on freshly denuded (severely disturbed) land ("new ground"). The few--though quite large--adult trees of eastern cottonwood that were still present in thic climax sugarberry-cedar elm-pecan forest had persisted since they had colonized "new land" made available by some disturbance(s) like flooding, windthrow, wild fire, beaver browsing etc. About the only "new ground" available to eastern cottonwood in this forest stand that was in or, at least, approaching old-growth state was the stream channel, either in the actual stream bed (as shown here) or on stream banks (as in the succeeding set of slides).

Time alone would tell if this tiny tree would be able to reach adulthood before being uprooted or broken off by future raging floods.

Erath County, Texas. Mid-March (late winter; hibernal aspect); seedling at four-leaf (second set of true leaves) stage.

 
   

 

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

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

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

 
   
Plant Succession on Bosque River bottomland forest
 

The city of Stephenville, Texas spent 2.2 million tax dollars to build a concrete street for biking, walking, jogging, etc. that was about one and a half miles long including a bridge over a drainage branch on Bosque River. This streatch of an extra-wide sidewalk was part of Stephenville's city park system. It was a classic pork barrel project; a textbook case of a street and bridge to nowhere. It was not as spectacular as the infamous bridge to nowhere in Alaska, the nation's largest state, but same thing (only smaller, less wasteful). The public works project was made possible by a grant from the Texas Department of Transportation, Statewide Transportation Enhancement Program (National Transportation Equity Act for the 21st Century) with $443,000 spent by city tax revenues ( 20% of total cost).

Much of this street ran along Bosque River, including portions of the immediate stream bank as well as the river floodplain, hence the designtion Bosque River Trail. Bosque River Trail was designed by the firm of Schrickel, Rollins and Associates, Arlington, Texas. Construction was by local contractor, Jay Mills Contracting, (webmaster@ci.stephenville.tx.us).

Bosque River Trail was not a trail or nature walkway, but just a city street of sidewalk thickness on which motor vehicles were excluded, except of course for city workers. Construction activity for Bosque River Trail was not that of a hiking trail or bridle path, but of road-building complete with bulldozing and earth-moving followed by pouring of ready mix concrete. In other words this project involved use of heavy equipment with destruction of vegetation followed by severe soil erosion resulting in water pollution. Bosque River Trail was not done by manual labor and delicate "environmentally friendly" construction with hand-held implements like axes, shovels, etc.

Following street construction, haul-back soil on the street berm was seeded to something (probably included Lolium multiflorum, annual ryegrass) and fertilized with some commercial fertilizer that contained urea. Two days later a flood came and washed much of this soil and fertilizer into Bosque River. As much as a third of a foot of soil from the upper slope above Bosque River Trail was crried into Bosque River. Portions of the street-like sidewalk of Bosque River Trail on which natural revegetation was occurring were mowed and weed-eaten, practices that further contributed to soil erosion. Clearly there was no post-construction activity that consistemt with best management practices.

This environmental fiasco--local devestation of a floodplain range ecosystem--afforded an opportunity for a case study of secondary plant succession on mixed hardwood bottomland forest. This author had worked on this area and an adjacent location over a period of three years prior to destruction/construction actions of so-called Bosque River Trail. Three days before devegetation of recovering plant life and seeding/fertilizing along the street-construction site this author took photographs of the pioneer plant community. Visual documentation and discussion of secondary plant succession of the devastated portion of the Bosque River floodplain forest ecosystem was presented in the following portion of this publication.

Findings and conclusions (up to time of street completion and end of first warm-growing season) were placed at end of this portion.

 
   
 

224. First stage of recovery- Plant life of the first seral stage of secondary succession on part of the bank of Bosque River subjected to earth-moving activity approximately six months after destruction of all existing climax vegetation of a mixed hardwood bottomland forest. This was the plant community near end of the first warm-growing season following devastation of by bulldozing, grading, and street paving . This scene was on the road (street) berm at outer river bank. The climax understorey that was removed--along with sevral feet of soil (river alluvium)--had been dominated by broadleaf woodoat and Canada or nodding wildrye..Some of this climax herbaceous layer of woodoats remained (upper left corner) as did some of the herbaceous layer made up of frostweed or white crownbeard along with giant ragweed (to immediate right of woodoats).

The seral vegetation (pioneer plant community) in fore- and midground consisted of lambsquarters (Chenopodium album) at right margin, Johnsongrass, hairy crabgrass, red sprangletop, common bermudagrass, Palmer pigweed (Amaranthus palmeri) plus resprouts of bois d'arc and the highly invasive, exotic tree-of-heaven (Ailanthus altissima). Johnsongrass and bermudagrass are naturalized, exotic (introduced) agronomic perennials; hairy crabgrass is a naturalized introduced annual; Palmer pigwood and red sprangletop are native annuals forb; bois d'arc is a native climax tree; and tree-of-heaven is an extremely invasive, exotic, weed tree. Sprout (heterophyllus) shoots of the two trees were from roots remaining after bulldozing and earth-moving activities as where shoots of Johnsongrass and bermudagrass which arose from surviving rhizomes and stolons and rhizomes, respectively. Plants of frostweed were also resprout shoots from underground perenniating parts.

The pioneer plant community was thus a combination of seedlings of typical annual colonizing species (crabgrass, lambsquarters, pigweed, red sprangletop) and vegetative shoots of pre-existing adult plants which had all of their aboveground parts removed (trees and a perennial composite forb). There were no plants of the climax, perennial bunchgrasses.

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

 
   

 

225. First seral stage- Vegetation at end of first warm-growing season of a mixed hardwood bottomland forest that had been destroyed six months earlier for construction of a street (a street to nowhere at cost of $2.2 million of tax-payer money). This pioneer (first seral) stage of secondary plant succession was on the street berm at outer bank of Bosque River. This was the vegetation that had developed at end of the first warm-growing season following destruction of climax vegetation. The first photograph was taken under a full-sun sky and the second photograph was under an overcast sky within one hour of each exposure. These two slides were of the same location, same seral vegetation, only from two slightly different angles or camera focl points. In the background of both slides were remnants of climax vegettion of this bottomland forest range which consisted of an herbaceous layer dminated by broadleaf woodoats and Canada wildrye along with lower limbs of sugarberry and cedar elm, co-dominant trees, and bois d'arc, an associate tree. Also in the remnant of vegettion not subjected to street-construction was the horribly invasive, exotic tree-of-heaven.

The pioneer plant community was a mixture of regrowth (sprouts; heterophyllus shoots) of sprouts of previously existing adult bois d'arc trees (foreground) and typical pioneer annuals, most plants of which were naturalized exotic annual species (primarily hairy crabgrass and lambsquarters). Three native annual species that are also typical pioneers were the eragrostoid-grass, red sprangletop, amaranth forb, Palmer's pigweed, and buffalo-bur, a forb of the nightshade family. Other annual grasses present included goosegrass (Eleusine indica) and barnyardgrass (Echinochloa crusgallii) which are both introduced species that have naturalized and are extremely well-adapted to disturbance.

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

 
   

 

226. Pioneers on a river bank- Two "photoquadrants" of the pioneer plant community on a portion of the bank of Bosque River that had been totally devegetated by bulldozing and blade-grading for construction of a street (that led to nonwhere except pockets of contracting firms). Both of these photographs were of the same local plant community though taken from different camera angles. The first photograph was from greater distance so as to include a slightly larger area and more vegetation. The second photograph was taken at shorter distance for a "closer-in" view and to show more detail of individual plants.

This was the colonizing vegetation at end of first warm-growing season following devegetation and creation of "new land" for secondary plant succession (bare soil surface). This was approximately six months following earth-moving action. It was a textbook example of old field succession or recovery of vegetation on "go-back land".

The pioneer plant community consisted of resprouts from bulldozed, adult bois d'arc trees and shoots of pre-existing Johnsongrass and common bermudagrass plus (and primarily) plants of typical, annual species of colonizing grasses and forbs. These annual species included red sprangletop, native grass; hairy crabgrass, naturalized, alien grass; goosegrass, naturalized exotic grass; barnyardgrass, naturalized alien grass; buffalo-bur, native, nightshade forb; Palmer's pigweed, native, annual amaranth; lambsquarters; naturalized, Eurasian forb; hop-hornbeam copperleaf (Acalypha ostryifolia), native euphorb; giant ragweed; feverfew or false ragweed (Parthenium hysterohorus), naturalized, exotic composite; and (in right background of first slide) cutleaf ground-cherry (Physalis angulata), annual nightshade forb.

The pioneer species of most lasting importance was eastern cottonwood. This tree species was represented by two seedlings (two seedlings in left midground of first slide; only the larger of the two in left midground of second slide). Some very large trees of eastern cottonwood were in this mixed hardwood bottomland forest. These cottonwood trees persisted into the climax forest dominated by sugarberry, cedar elm, and pecan. Eastern cottonwood lives long enough to be part of the climax forest although cottonwood is not a climax species in the same sense as tolerant species like sugarberry and cedar elm. Pecan has a life cycle pattern similar to that of eastern cottonwood, but pecan was more abundant and persisted into the climax forest as a dominant with sugarberry and cedr elm. This successional phenomenon of pecan was shown and discussed below.

Easterrn cottonwood has traditionally be classified as Very Intolerant (Wenger, 198; Burns and Honkala,1990) with cover types dominated by it as seral, in fact as pioneer forest communities (Eyre, 1954; Eyre, 1980). Sexual reproduction (successful recruitment of seedlings) of eastern cottonwood requires availability of bare, moist, mineral soil as a suitable seedbed (Eyre,1954; Eyre, 1980; Burns and Honkala, 1990). In other words, disturbances severe enough to result in bare mineral soil are essential for establishment (= germination, emergence, root/shoot development, and seedling survival) of eastrn cottonwood. This pioneer tree species could not become established in this climax sugarberry-cedar elm-pecan forest without creation of "new land" (soil bare of duff and other organic matter) by disturbances such as tree blowdown (creation of "tree cradles"), flood scouring, or human mechanical scarification.

The Bosque River Trail project that squandered $2.2 million in tax revenue to build a street to nowwhere did create several areas so severely disturbed (completely devegetated by bulldozing and blading) that there were numerous seedlings of the pioneer species, eastern cottonwood. Whether the busybody bosses of the Stephenville City Park permit growth of these seedlings into large, handsome, adult cottonwoods or, altrnatively, these public servants order the seedlings cut off with lawn-mowers or weed-whackers so as to have a cropped turf of alien hairy crabgrass is another matter. Nonetheless, the lesson of vegetation development on this bottomland forest was obvious: eastern cottonwood, a pioneer tree species that persist into climax forest, did regenerate. In fact, regeneration was instaneous in successional time scale. Within six months of a suitable seedbed becoming available, seedlings of eastern cottonwood had become established and were on their lives' journey to becoming mature trees. That is, as long as so-called park managers realize that trees belong in a river bottom forest and that maintenance of such trees requires successful reproduction for replacement of trees that eventually die.

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

 
   
 

227. First colonizers and an increaser- Two seedlings of eastern cottonwood amid a few plants of silver bluestem (Andropogon saccharoides) on a patch of soil on the bank of Bosque River that had been bulldozed and bladed six months prior to time of photograph. Removal of all existing aboveground plant life resulted in land laid bare to mineral soil which permitted germination, emergence, and rapid establishment of the seedlings of the pioneer tree species, eastern cottonwood. The plants of silver bluestem regrew from rootcrowns of this perennial grass that miraculously had survived the earth-moving activity. In the same fashion, surviving rhizomes of Johnsongrass gave rise to shoots of this naturalized, African, perennial grass. Other plants present in this "photoplot" included hairy crabgrasse and Pennsylvania smartweed or Pennsylvania knotwee (Polygonum pensylvanicum).

Forest range vegetation on this microsite did not have the same first seral--the colonizing--stage of secondary or old-field succession as that on almost all of the other such "minisites", but seedlings of the pioneer tree species had established.

Erath County, Texas. Early September (late estival aspect of the pioneer plant community).

 
   
 

228. Pioneers with different approaches- Local spot of bare soil on the bank of Bosque River left after street-construction activity that involved bulldozing, grading, form-building, and concrete delivery and working. Plant species included annual and perennial and native and naturalized exotic monocotyledons. These species were red sprangletop, native, annual grass; hairy crabgrass, exotic, annual grass; common bermudagrass, exotic, perennial grass, Johnsongrass, exotic, perennial grass; and purple nutsedge or purple flatsedge (Cyperus rotundus). Note that four of these five graminoids were introduced species--by white man, purposely or accidentially--that had naturalized. Even more telling was the fact that three of these four species were listed and discussed as some of the worst weeds on Earth: purple nutsedge, world's worst weed; bermudagrass, world's second worst weed; and Johnsongrass, sixth worst weed on Earth (Holm et al., 1991, ps. 8-31, 54-61).

This pioneer plant community was made possible by 2.2 million tax dollars to build a city street known as Bosque River Trail, a part of the city park of Stephenville Texas, which was allegedly (at least partly) justified as a "nature trail". Nice start to educate trail-users to natural plant communities, hugh!

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

 
   

 

229. Woody invaders that are climax species- The berm of a street on the bank of Bosque River supported a number of woody plant species late in the first warm-growing season following devegetation about six months earlier. This portion of the river bank had been bulldozed, balded, and subjected to street construction work that resulted in removal of all of the organic matter layer from the alluvial soil (Bunyon series). All plants in these two photographs were woody species except for one plant of frostweed or white crownbeard, an herbaceous composite that had regrown from its perennial rootcrown, and one plant of sharppod morning-glory (Ipomoea trichocarpa= I. cordatotriloba), a twinning perennial forb. Frostweed was the most abundant forb overall in the herbaceous layer of the climax sugarberry-cedar elm-pecan bottomland forest that was locally removed for street construction.

Woody species included pecan seedlings and resprouts of poison oak/ivy, and trumpet creeper that arose from rootstocks. The second of these two slides was a close-up of two pecan seedlings (a small vine of morning-glory was in background).

Pecan has been interpreted as Intolerant though somewhat less so than cottonwood and willow (Burns and Honkala, 1990). These authors regarded pecan as a subclimax species. On this mixed hardwood bottomland forest pecan was found to be one of three dominant, climax, tree species (sugarberry-cedar elm-pecan climax) because pecan trees had such great longevity that they persisted into climax, and as the largest trees of the climax forest. It was discussed immediately above that eastern cottonwood also persisted into the climax (as a minor species only) even though cottonwood is a pioneer or colonizer (member of the first seral stage). Pecan is also a pioneer tree species, but its larger fruits do not appear to be as senestive to seedbed conditions as the much smaller seeds of cottonwood.

It was obvious from these (and accompanying) photographs that sexual regeneration of pecan was substantial on bare mineral soil of this alluvial deposit. Several slides presented above included established seedlings and saplings in the herbaceous layer of this climax forest. Regeneration (sexual reproduction) of pecan in competition with such herbaceous species as Canada wildrye and broadleaf woodoats, co-dominants of this layer, was "almost nothing" compared to seedling establishment on the bare soil of this street berm. Clearly, pecan reproduction is much greater on exposed soil and with reduced (or eliminated) competition.

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

 
   

 

230. .Invaders of all sorts- An array of plants had established on the bare, exposed soil of a street berm on the bank of Bosque River near end of the first warm-growing season following destruction of the climax vegetation of a mixed hardwood bottomland forest. All aboveground parts of plants in this range plant community had been removed by earth-moving and shallow excavation followed by street construction about six months prior to time of these photographs. Established shoots visible in these first of these two "photoquadrants" included pecan seedlings along with stump/root sprouts of Virginia creeper and trumpet creeper and grass shoots of hairy crabgrass. Pecan and the two species of woody vines are natives whereas crabgrass is a naturalized annual. Plants in the second "photoplot" included five seedlings of pecan, one seedling of cedar elm, plus one shoot of sharppod morning-glory and some hairy crabgrass. Only the crabgrass was a non-native species. One leaf of eastern cottonwood and one leaf of red mulberry in this second photograph attested to presence of nearby trees of these two native tree species.

Seedlings of pecan in these two slides and the two immediately preceding slides were photographic evidence that pecan quickly and readily establishes from seed on "new land" created by disturbance severe enough to result in bare soil. Observations in this climax sugarberry-cedar elm-pecan floodplain forest handily showed that establishment of pecan seedlings is much greater on bare soil than on the dense sward of the herbaceous understorey dominated by Canada wildrye and broadleaf woodoats and under partial shade from crowns of adult trees. Pecan was recorded as Intolerant by Burns and Honkala (1990), but not as much so as was eastern cottonwood that also established seedlings on the bare soil of this street berm (see above).

Lianas (Virginia creeper, trumpet creeper, poison oak/ivy) regrew from established underground organs. These woody climbers were all part of the climax forest vegetation (as was mustang grape). Pecan is such a long-lived tree that it persisted into the climax (and as the largest plants of the forest) even though it is a colonizing species. The same relationship obtained for pioneers of eastern cottonwood, but there were fewer cottonwood trees in the climax whereras pecan persisted in such numbers and of such size and crown coverage as to be one of the dominant, climax tree species.

Invaders in the meaning of plants that established secondary plant succession, especially in early stages thereof, has a slightly different meaning from invaders as used in the more specific application of successional status for determination of range condition classs (ie. decreasers, increasers, invaders). The more general and original meaning (as the term was invented by F.E. Clements) of invaders refers to plants--both individuals and species--that had completed the process of invasion. Weaver and Clements (1938, p. 166) described Clementsian invasion with these words: "Invasion is the movement of one or more plants from one area into another and their establishment in the latter It is thus the complte and complex process of which migration, ecesis, and competition are the essential parts". Later, one of J.E. Weaver's students (Dykersthuis, 1949) used invader to mean a species that continues to increase in abundance under abnormal or extreme disturbances, in particular to overgrazing. Responses to overgrazing were determined (estimated) as to different range sites (ie. they are range site-specific). Responses of specific species to other disturbances (eg. drought, unusual fire, extreme temperature, tillage) were understood to be similar though not exactly the same as responses to overgrazing which is the main abusive practice and extreme disturbance that rangemen can prevent.

Invasion as a central part of plant succession interpreted in the Clementsian model of "dynamic vegetation" (= vegetation development) begins with initial causes, disturbances that "...produce a new or denuded soil upon which invasion is possible" (Weaver and Clements, 1938, p. 106). The initial cause of this invasion on the bank of Bosque River was human earth-moving action in conjuction with street construction (and destruction of existing climax or subclimax vegetation). Continuous or recurrent invasion is the kind or form of new plant establishment tha occurs when pioneer tree species such as willows or cottonwoods "lay claim" to the fresh land or seedbed created by disturbances such as flood, fire, or tillage (Weaver and Clements, 1938, p. 167). The invaders shown in these and preceding photographs were the products of continuous invasion (seeds and fruits are more or less continually produced--dispersed or disseminated [dispersal or dissemination]--on an annual basis whether or not they encounter a suitable medium for establishment or achieve the completed process of migration).

Invasion includes migration (successful arrival of plant progagules or disseminules on a denuded or otherwise suitable area for growth), ecesis (germination, growth and some form of reproduction such as production of more shoots or leaves), aggregation (the process by which migrants form groups of new plants through reproduction), and competition (a main driving force in later stages of plant succession).

Strictly speaking, the new plants (seedlings, root sprounts, stc.) in the above "photoplots" where in earlier stages of yet-to-be-completed invasion. The seedlings,especially, were at the stage of ecesis. Whether or not they grow and reproduce adequately to complete the full set of processes of invasion would depend on various factors, the main one of which being if city park workers would permit these plants to grow or cut them off with power implements. In fact that is exactly what happened. Four days after this photographer took the photographs presented here employees of the city of Stephenville, Texas and/or Jay Mills Contracting mowed and weed-whacked off all of these plants. The process of seondary plant succession was set back to the beginning or starting stage on "ground zero" of Bosque River Trail. Apparently "nature trail" meant mowed turf of naturalized alien species, namely hairy crabgrasss and common bermudagrass (neither of which were planted by these workers).

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

 
   
 

231. Rapid return- A three- or four-month old root sprout of cedar elm that grew on the berm of a street named Bosque River Trail built on the bank of Bosque River in the West Cross Timbers of northcentral Texas. This specimen was the result of secondary plant succession on a sere, the area of denuded raw soil on which vegetation development could take place, of a climax sugarberry-cedar elm-pecan bottomland forest. This root sprout was an example that secondary succession in this forest cover type was a combination of both seedlings and resprouts or stump sprouts (mostly heterophyllous shoots), of perennials as well as annual species, and of both native and naturalized exotic species.

There were plants of the typical pioneer annual species (native and naturalized) such as buffalo-bur, red sprangletop, cutleaf ground-cherry, Palmer's pigweed, lambsquarters, and hairy crabgrass along with seedings of pecan (a native, climax, dominant tree species), sprouts from perennial rootcrowns of native lianas (common green-briar, trumpet creeper, Virginia creeper, poison oak/ivy), new shoots from frostweed (native, perennial, composite forb) and of Johnsongrass and bermudagrass (naturalized, alien, perennial grasses), and resprout shoots of cedar elm (one of the native, climax, dominant trees) and of bois d'arc (a native, climax, associate tree species).

The rest of the story (so far) is that this new plant of cedar elm (again, one of three climax dominant tree species of this forest type) was cut off by workers applying seed (apparently of the exotic, annual ryegrass) and a urea-containing commercial fertilizer. Two days following this seeding/fertilizing operation, flood waters and heavy upslope runoff from a heavy rainstorm carried the soil, seed, and urea-fertilizer from this and the rest of the area--all except for the concrete street--into Bosque River. This photographer witnessed and documented the entire process, including the severe sheet and rill erosion as it was taking place.

Erath County, Texas. Early Septermber.

 
   
 

232. Younger elderberries- Resprouts of elderberry (Sambucus canadensis) growing on the bern of a street constructed on the outer bank of Bosque River. These four shoots were resprouts growing about one year following removal of all aboveground parts of plants in the destruction/construction operation. These shoot were approximately five to six months old. In the relatively mild winters of northcentral Texas elderberry with its extremely freeze-hearty foliage, typically overwinters with green leaves.

Ground cover was leaves of various hardwood trees (pecan, sugrberry, eastern cottonwood, cedar and America elms, and live or plateau oak) that escaped the bulldozers of street contractors.

Erath County, Texas. Late February.

 
   

 

233. Can't keep good vegetation down- Range vegetation of a mixed hardwood bottomland forest developing on the berm of a city street in early spring following removal of all aboveground parts of plants (total elimination of all plant life above soil surface) the previous spring. This is the pioneer stage of forest vegetation on land totally denuded one year earlier l(ie. plant life in the first spring following removal of existing vegetation). Plants shown in the first photograph included two seedlings of eastern cottonwood; one seedling of American elm; resprouting greenbriar, chittamwood, and Carolina snailseed (Cocculus carolinus); and broadleaf woodoats, Canada or nodding wildrye (both native perennial grasses) as well as seedlings of such annual herbaceous species as giant ragweed, hedge parsley, and Japanese chess or Japanese brome. Plant species presented in the second photograph included two seedlings of eastern cottonwood along with several seedlings of sugarberry and American elm accompanied by broadleaf woodoats and the white-umbeled hedge parsley.

This assemblage of plant life ranged from climax, shade-tolerant trees to exotic, naturalized annual forbs (hedge parsley) so that the pioneer or colonizing stage of secondry plant succession did not follow the classic revegetation pattern reported (and learned by generations of students) for old fields and cutover forests. Initial stages of old-field succession included major components of climax trees as well as typical colonizing species such as the native giant ragweed, an annual forb, and eastern cottonwood, a pioneering tree that typically requires bare mineral soil for seddling establishment.

This pattern or revegetation was perhaps a reflection of the comparatively small area of denudation: a long, narrow strip of mineral soil that had a rich seedbank and in close proximity to germinule-yielding plants including both climax and seral tree species. Likewise, there were stump sprouts (long shoots) of shrubs and tillers from rootcrowns of native perennial grasses.

Erath County, Texas. Late April.

 
   

 

234. Details of recovering forest vegetation- Two close-in "photoquadrants" of recovering floodplain forest vegetation in early spring one year after street construction denuded a corridor of sugarberry-cedar elm-pecan climax forest on the outer bank of Bosque River. The first slide showed a seedling of eastern cottonwood and one seedling of American elm along with elderberry, rescuegrass, Japanese chess or Japanese brome, and broadleaf woodoats. The second "photoquadrant" contained two seedlings of American elm plus Johnsongrass, Carolina snailseed, Canada wildrye, and hedge-parsley.

This first (pioneer) stage of secondary pland succession was a combination of climax native trees (American elm), colonizing native trees (eastern cottonwood), native shrubs (elderberry), native perennial grass species arising from belowground parts (Canada wildrye, broadleaf woodoats), .introduced perennial grass (Johnsongrass), exotic annual grasses (rescuegrass, Japanese brome), and introduced annual forbs (hedge-parsley). The initial seral stage was tnot comprised totally of herbaceous annuals and therefore did not conform with the classic pattern of old-field succession.

Erath County, Texas. Late April.

 
   
 

235. Some of the beautification received by tax payers - Severe gully erosion on the bank of Bosque River resulting from construction of Bosque River Trail. Inadequate erosion protection by contractor Jay Mills Contracting resulted in formation of these new gullies when flood waters from rainfall of moderate intensity flowed completely unabated from the immediate watershed into the stream. Most of the small, reactangular blaes of straw loosely anchored on stream banks washed into Bosque River. Beaver used one of these bales as the "cornerstone" of a dam they built across the stream (shown and described above). lAccording to the city web page, (webmaster@ci.stephenville.tx.us). Stephenville Texas spent 2.2 million tax dollars (80% grant from Texas Department of Transportation, Statewide Transportation Enhancement Program, 20% city revenue) to construct a walking trail (it was more like a city street that handily accomodated city motor vehicles) on the outer bank of Bosque River. There was obviously not effective enforcement of regulations by the Texas Commission of Environmental Quality to oprevent such damage from runoff.

Most of the green clumps of grass on the immediate river bank were Canada or nodding wildrye and broadleaf woodoats. Note how these two native grasses effectively protected the far bank of Bosque River (background).

Erath County, Texas. Late April.

 
   

 

236. Severe soil erosion in the name of beautification (and paid for by tax dollars)- Pronounced sheet erosion of Purves clay (clayey, montmorillonitic, thermic lithic calcustolls) immediately above outer bank of Bosque River. Removal of all vegetation from a large area of land (having five percent slope) during street construction (so-called Bosque River Trail") left this soil unprotected. The same moderate-intensity rainstorm that resulted in erosion of the river bank shown in preceding slide was the weather event that carried most of the upper A horizon of this soil into the river. Depth of soil removal varied from six to ten inches. Depth of soil loss where stockman's knife was standing (second slide) exceeded six inches, but there had been two to three inches swept off of the "shelf" immediately above the knife. This land had been scraped clear of almost all protective cover by bulldozing and grading. The contractor who was responsible for this soil erosion was Jay Mills Contracting, Stephenville, Texas.

This eroded land was in the city park of Stephenville, Texas. It should be remarked (sadly though instructively) in passing that historically much of the waste and mismanagement of natural resources has been by or with approval of government entities. This can occur at any level of government from federal down through municipal as in the example seen here.

This was on the perimeter of a mixed hardwood bottomland forest immediately above Bosque River. Plateau live oak (Quercus virginiana var. fusiformis) was the local dominant tree with cedar elm and pecan associate species. Leaves on soil surface (shed subsequently to land clearing) were those of plateau live oak.

Erath County, Texas. Late April.

 
   
 

237. By way of contrast- A local undisturbed microsite of the herbaceous layer of climax sugarberry-cedar elm-pecan bottomland forest at the outer margin of the Bosque River floodplain. This less mesic, upper slope of the floodplain forest had an understorey more typical of tallgrass prairie rather nhan the herbaceous layer of Canada wildrye and broadleaf woodoats that was climax for lower, more mesic portions of the river floodplain.

Major species in this drier zone of the forest herbaceous layer were big bluestem, sideoats grama, sand lovegrass, purpletop, pokeberry, pigeonberry, Louisiana sagewort, and frostweed (all native perennials) along with naturalized alien species, namely hairy crabgrass, Johnsongrass, and hedge parsley. Woody species in the background included cear elm, live oak, and bur oak (native, climax trees) and tree-of-heaven (horribly invasive exotic tree).

Range vegetation in this "photoplot" as well as those presented in which Canada wildrye and broadleaf woodoats dominated the understorey can be compared to the pioneer--the first seral stage--of vegetation displayed and discussed in the last slide-caption sets presented immediately above.

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

 
   
Farther along- Treated in the short section that follows was a seral stage of plant succession in a climax sugarberry-cedr elm-pecan bottomland forest that was more advanced than the pioneer or first successional stage presented above. This local forest vegetation was on the upper (more upland; slightly higher elevation) part of this mixed hardwood floodplain forest where it was advancing into an old field (abandoned cropland). This was a "farther along" stage of secondary plant succession of this forest range type.  
   
 

238. Advancing edge- Outermost edge of a climax sugarberry-cedar elm-pecan bottomland forest on the Bosque River floodplain that was invading (encroaching on or extending into) abandoned farmland (ie. an old field). Number of years since the last tillage of this crop field was not known, but it was certainly in excess of a decade and was probably in excess of a quarter-century. The last field cropon this field had been Coastal bermudagrass (Cynodon dactylon cultivar Coastal). Considerable cover of Coastal bermudagrass remained, but this was exceeded and overshaded by Johnsongrass along with much cover of hairy crabgrass in the warm-growing season and such weedy annuals as cheatgrass, Japanese chess, and rescuegrass (annual Eurasian Bromus species) plus hedge parsley, redstem filaree (Erodium cicutarinum), bur-clovers (Medicago species), and other naturaized Eurasian forbs.

This field was mowed for hay at irregular intervals, but it had not been hayed for several years immediately preceding time of this slide and the two slides of the next set. This photograph was an oblique-angle view of the outermost edge of the bottomland forest when it met the old field. The limit of mowing was conspicuous in this and the first of the two slides immediately below. This is a nearly universal pattern of vegetation encroachment into hay fields and mowed meadows. The hay mower carefully keeps the most distant part of the mowing implement at a safe distance from any visible woody plant to avoid damaging the mowing machine. As years pass and woody plant life grows while the mowing implement is kept safely away from such woody growth, the edge of the mowed area slowly and inexorably withdraws inward so that total area (acerage) of mowed field declines year-by-year while area of land dominated by woody plants expands and encroaches into the cropland. Even with mowing operations--other than those specifically for brush control (shredding)--size (acerage) of field gradually becomes less while vegetation dominated by woody species expands commensurably.

Such was the inevitable pattern of forest expansion into this old field as shown in this and the two succeeding photographs. Such is the farming lesson. Laziness, indifference, and stupidity lead to neglect and the farmer's field eventually becomes overrun by brush. The forest reclaims its own. From the forester's perspective this process of plant succession results in reforestation of cutover or cleared land (ie. restoration of the forest crop).

The specific lesson in forest succession shown here was that the principal invading woody plant was pecan (present as many small saplings and--though not visible here--sedlings) which is one of the dominants of this sugarberry-cedar elm-pecan forest. In the immediately preceding section (devoted to the first seral stage or colonizing plant community of this range forest cover type) it was shown and explained that pecan is a pioneer tree species (along with eastern cottonwood). Pecan, an Intolerant tree species, does best--probably requires--bare, mineral soil for emergence and establishment of seedlings. On this forest dominance type, pecan and eastern cottonwood (and sycamore in some areas and on similar bottomland forests) are pioneering trees along with annual herbaceous species. These long-lived tree species persist into the climax or, at least, subclimax forest. On this forest dominance type--the sugarberry-cedar elm-pecan climax--the Intolerant pecan has little (yet, perhaps, adequate) regeneration in the climax forest. Regeneration (primarily sexual reporduction other than some asexual shoots of cedar elm arising from roots) is almost entirely of sugarberry and elm (a very small proportion of American elm along with the dominant cedar elm).

Presence and dominance of pecan in the climax canopy is from crowns of immense pecan trees that became established in the first (at least, earlier) seral stage(s) of plant succesion. That pattern of successional development was shown in this and the next two slides (for the sapling stage of pecan) and in preceding photographs (for the seedling stage of pecan). The limited regeneration of pecan (establishment of pecan seedlings) in more open areas of this climax forest was also presented above. Pecan regeneration in later stages of forest succession (say, subclimax and climax) might be sufficient to maintain a "presence", moderate abundance, or even slight dominance in the climax of this cover type. However, most of the "clear and convincing" dominance cover--crown and basal (stump or diameter breast height)--is that from old, monumental-sized trees that established on "new land" created by disturbances severe enough to result in bare, mineral soil. It was explained above that this is the initial cause of plant succession in the Clementsian model (Weaver and Clements, 1938, p. 106).

The other important species of invading tree was honey locust. Honey locust had obviously become established, and it was also obviously the associate to pecan. Honey locust is also an Intolerant species that can only become established in forest openings (Burns and Honkala, 1990) and, better yet, on bare ground such as that of old fields. Invasion of abandoned cropland by honey locust is a widespread phenomenon. Honey locust is a pioneer tree species and one that is very sensitive to shade so that it can continue to survive in forest only with crowns in the canopy (Burns and Honkala, 1990). Unlike pecan, honey locust is a relatively short-lived tree maturing at a litttle past the century mark (Harlow et al, 1979, p. 387).

Other woody invaders in this old field included bois d'arc (which also persist as a lower-growing tree into climax), American or white elm (in contrast to the more abundant cedar elm that was a climax dominant), eastern redbud (a shrub largely absent from the shrub layer in the climax vegetation), and mustang grape (a liana that clearly is part of the climax forest as shown below).

The white-flowered composite (DWC= Damn White Composite) in the advancing line of forest was frostweed or white crownbeard, the dominant climax forb in the herbaceous layer of the sugarberry-cedar elm-pecan cover type. Note the absence of this climax composite in the mowed area. Mowing had killed it out of (or, perhaps more accurately speaking, prevented its invasion into) the hay field. The main grass species was purpletop although Johnsongrass was very common as it was on the mowed portion (hay field). Another important forb was the native papilionaceous legume, sessileleaf tick-trefoil (Desmodium sessillfolium).

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

 
   

 

239 Forest frontier- Outermost edge of a sugarberry-cedar elm-pecan bottomland forest that was advancing into (invading) an abandoned crop field through secondary forest successsion. The most abundant (greatest cover, highest biomass) was pecan. Pecan is a pioneer species of such long life that it persist into the climax canopy as one of the dominant tree species. The second most abundant tree--though a distant runner-up--was honey locust. The third (and second runner-up) tree was American elm. This latter was a minor member of the adjoining and fully developed climax forest in which cedar elm was second only to sugarberry. Reasons for this difference were not evident. Eastern redbud was another woody species. This surub was not present to any degree in the climax mixed hardwood floodplain forest. Redbud was obviously a seral species in this forest type. Mustang grape was also common. This woody vine is a "proud member" of the climax forest. An eaxmple (and a damn big one) of mustang grape in the climax state of this forest range type was shown below.

The most abundant and important herbaceous species were white crownbeard or frostweed, which is also the dominant forb of the climax forest, and purpletop. Purpletop is almost always classified as an increaser. It was clearly seral to the climax Canada wildrye and broadleaf woodoats in the sugarberry-cedar elm-pecan bottomland forest type, but it and Johnsongrass were the major grasses in this seral range vegetation. Other common herbaceous species included hedge parsley, a widespread Eurasian annual forb, and sessileleaf tick-trefoil or sessileleaf tick-clover, a native papilionaceous legume.There was both common bermudagrass and some remnant plants of Coastal bermudagrass, this latter being the last field crop grown on this now cropland.

Pecan and honey locust are both Intolerant species and require bare (or nearly so) mineral soil for establishment of seedlings. Relevant silvical aspects of these two species as related to succession and forest development were detailed in the immediately preceding caption (as well as several others above) so such discussion was unnecessary in this caption. Other details, (eg. those related to farming practices) that were pertinent to forest invasion (expansion or encroachment) into this old field were also given in the preceding caption and not redundantly included here.

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

 
   
Some of the major range plants on the climax sugarberry-cedar elm-pecan that developed on the Bosque River floodplain
 
   

 

240. Native brome on bottomland land- Hairy or woodland brome or bromegrass (Bromus purgans= B. pubescens) on floor of floodplain hardwood forest on floodplain of Bosque River. The first photograph showed a small, local colony of hairy wood brome growing in shade of (at one time or the other during daylight) pecan, black walnut, cedr elm, sugarberry, and red mulberry. Second photograph was a typical panicle of hairy brome in this restricted population.

This was a rather rare find because hairy wood brome is not a common range plant in the Cross Timbers and Prairies vegetational area. This is undoubtedly largely due to acreage limitations. Even prior to appearance of European man mixed hardwood bottomland forests comprised a small fraction of natural vegetation in this area. Now almost all of this range type that remains has been severely degraded due to numerous disturbances including farming (more in the past), overgrazing, elimination of fire, and real estate developments. Even with that, it is also a fact that this has been more-or-less overlooked by professional workers in this area. For example, the definitive authority for grasses in the Cross Timbers-Prairie area (Highnight et al., 1988) failed to include Bromus pubescens as also did Gould (1962, 1975) in the checklist of Texas plants. Woodland brome probably still persist in this area only in small parcels of relict vegetation like that treated in this section.

Erath County, Texas. Mid-May, milk to soft-dough phenological stage.

 
   

 

241. Hairy details- Views showing node and internode along with spikelets of hairy wood brome in herbaceous vegetation of a on sugarberry-cedar elm-pecn bottomland forest that developed on the floodplain of Bosque River. Observe the dense though short pubescence on spikelets as well as shoot.

Erath County, Texas. Mid-May, milk to soft-dough phenological stage.

 
   
 

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

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

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

 
   

 

243. Herbaceous dominant- Sexual shoots of a robust plant of Canada or nodding wildrye (first photograph) growing on the floodplain (second terrace) of Bosque River. The was the first dominant (broadleaf woodoats being a close second-place dominant) of the herbaceous layer of a sugarberry-cedar elm-pecan bottomland forest. Warm, wet weather throughout winter to early spring enabled nodding wildrye to produce an unusually high proportion of shoots that progressed to sexual reproduction. Many large, caryopsis-filled spikes (like the two shown in the second photograph) resulted in a abundant grain crop.

Canada wildrye is one of the most imporrtant native cool-season grasses in the immense region of tallgrass prairie and associated oak-hickory-tallgrass savanna, including the Western Cross Timbers-Prairies area where this bottomland forest range had developed. All of the standard flora and grass manuals have good botanical descriptions of Canada or nodding wildrye. Barkworth et al. (2007, ps. 303-305) was the most up-to-date and the North American "grasss bible" for years to come (even though the "King James equivalent" probably will always remain Hitchcock and Chase [1950, p. 260-261]). From a Range Management perspective the classics Range Plant Handbook (Forest Service, 1940, G50) and Pasture & Range Plants (Phillips Petroleum Company, 1963, p. 35) remain two of the best. Also recommended from the range plant view was Bush (2005). Genetic studies of nodding wildrye included those by Sanders and Hamrick (1980) and Saha et al. (2009).

The common name of Canada wildrye was obviously derived from the commerative specific epithet. Equally obvious from these two slides was origin of the common name, nodding wildrye.

Bosque River floodplain, Erath County, Texas. Late May; peak shoot biomass, late stage of anthesis.

 
   

 

244. Sexy details of the dominant- Close-in views of a large spike of Canada or nodding wildrye produced on the second terrace of Bosque River following warm, wet weather from winter through early spring. In nodding wildrye there are typically two to three spikelets per rachis node with two to four florets in each spikelet (Barkworth et al, 2007, ps. 303). Large spikes with several flowers per spikelet and an abundant crop of pollen in this wet year later resulted in a high grain yield from Canada wildrye plants in the understorey of the sugarberry-cedar elm-pecan bottomland forest described herein

Bosque River floodplain, Erath County, Texas. Late May; peak anthesis stage.

 
   

 

245. Associate grass species- Sand lovegrass (Eragrostis trichodes) was a third important native, perennial grass of the mixed-hardwood, bottomland forest along this stretch of Bosque River. Sand lovegrass was clearly an associate grass compared to Canada wildrye and broadleaf woodoats, but this eragrostoid grass was a local dominant. Plus it was a C4 and warm-season species, two features much more common among grass species in this southcentral region of North America.

In the first of these two "photoquadrants" sand lovegrass was joined by the two dominant cool-season grasses. In the second "photoquadrant" the "phyto-escorts" were a seedling of cedar elm as woody cohort along with an herbaceous component of pigeon-berry and horsepen or straggle daisy (Calyptocarpus vialis).

Erath County, Texas. October, peak standing crop and ripening grain stage of sand lovegrass.

 
   
 
246. An example of its kind- Sand lovegrass on the floodplain of Bosque River with some broadleaf woodoats behind. The panicle of this species can sometimes account for two-thirds of shoot length. Erath County, Texas. October, peak standing crop with ripe grain.  
   
 

247. Spikelets on sandstone- Tiny spikelets of sand lovegrass lying against the parent material of Cross Timbers soils. Sand lovegrass usually has four to 18 florets (Gould, 1975, p. 185). These spikelets were gathered from the plants shown in the preceding slide.

Bosque River, Erath County, Texas. October, grain-ripe stage.

 
   

 

248. An uncommon assembly- Stand of Canada or hairy wood brome (Bromus pubescens= B. purgans) on the upper bank of Dry Branch draining into Bosque River. This native perennial bromegrass has a species range extending from south to Florida and westward to Texas. Plants in this stand had attained a height of approximately one yard or slightly more.

Due to the heavy hand of white man weedy annual Bromus species introduced from Eurasian (and now naturalized in much of North American) are much more common than this native perennial species. This robust festucoid grass (one of the author's favorites) grows on more mesic habitats along streams; generally rather above them and typically not in the riparian zone. For example, Canada or hairy wood brome grows on relict sugar maple (Acer saaccharum)-basswood (Tillia americana) forests that develop along bluffs aligned beside streams in the Ozark Plateau (examples of this grass on that forest range cover type were also shown in Range Types with the Oak-Hickory Forest under Wooodlands and Forests).

Hairy wood brome is relatively rare in Texas' Cross Timbers and Prairies vegetational area (this stand was an exhilirating and m ost rewarding occasion), but its presence on (above) Bosque River along with other diagnostic species like bois d'arc, red mulberry, and black walnut showed the ecological affinity of the sugarberry-pecan-cedar bottomland forest type with riverine forests far to the north such as those from northern Kansas show above. This species has historically not been shown to grow in the Cross Timbers and Prairies area. Such highly respected works as Gould (1962), Gould (1975), Hignight et al. (1988) failed to include B. pubsecens (B. purgans) in this vegetational or land resource area.

Erath County, Texas. January.

 
   

 

249. Brome above the Bosque- Plants of the native perennial Bromus pubescens (= B. purgans) on a bank high above an ephemeral stream (Dry Branch) draining into Bosque River in West Cross Timbers of northcentral Texas.

Erath County, Texas. January; early basal leaf stage of current season's growth.

 
   
 

250. Young leaaves- Features of early basal leaves in Canada or hairy wood brome on floor of a sugarberry-pecan-cedar elm bottomland forest. Basis of the common name adjective, hairy, and the specific epithet, pubescens, was evident in this close-up photograph of leaves of the plant shown in the immediately preceding photograph (second slide of the above two-slide set).

Plants were growing during a Severe drought (Palmer scale rating) at time of photographs, but warmer-than-typical temperatures had amellorated some of the reduction and delayed growth of leaves. Native plants simply are not as adversely impacted by stresses due to abnormal growing conditions to the extent that agronomic and horticultural species are.

Erath County, Texas. January; early basal leaf stage (leaves ranged from three to six inches in length).

 
   

 

251. Last year's and this year's- Young basal leaves of the current cool-growing season of hairy wood brome overlaid with some fallen branches of last year's panicle. Details of new leaves and spikelets showed distinctly. The spikelets presented in these two photographs appeared to have their full yield of florets there having been little grain-shattering during a drought of Moderate to Severe rating (Palmer Index). The three to five florets per spikelet were at the lower end of that cited for this species by Barkworth et al. (2003, p. 220). Leaf growth (number of leaves and lengths of them) was somewhat less than that in a year of more typical precipitation, but there was not as much reduction as might be expected due to warmer temperatures.

Erath County, Texas. January; early basal leaf stage (leaves ranged from three to six inches in length).

 
   

 

252. Hanging high above the Bosque- Portion of a panicle of hairy wood brome showing secondary panicle branches (first photograph) followed by detailed view of the two centermost spikelets (second photograph). Canada brome has fewer florets per spikelet--typically four to ten (Barkworth et al., 2007, p. 220)--than many of the Bromus species, especially the more common annuals. Some of the florets had been shed from spikelets presented in these slides whereas other spikelets (such as those shown in the immediately preceding photographs) appeared to have their full complement of florets. All of these slides were taken in early winter following one of the driest dormant periods of this species (Moderate to Severe rating drought from summer through autumn) in recent decades. Thus there was less precipitation, wind, etc. to shatter grains.

Erath County, Texas. January.

 
   
 

253. A species largely grazed out- A healthy population of three-flower melic (Melica nitens) grew in a protected cover just beyond the immediate bank of Bosque River served as a reminder, an indication (= an indicator species), as to what the species-rich, highly productive herbaceous understorey of Bosque River was once like before the whiteman's influence so modified this mixed hardwood bottomland forest. Most of the Melica species, including M. nitens, are unusually sensitive to heavy grazing. Undoubted the Bosque River floodplain had been severely overgrazed by livestock. In fact, overgrazing/overbrowsing would have been essentially unavoidable in this area of bottomland forest range that was adjacent to a county seat town (within the city limits at time of this photograph).

Erath County, Texas. Late April; grain-ripe phenological stage.

 
   

 

254. Other specimens not grazed out- A local population (first slide) and a large individual plant (second slide) of three-flower melic growing on the immediate bank of Bosque River. Three-flower melic has short rhizomes (Barkworth et al., 2007, ps. 100-101) so it was possible that some of the shoots and, even, clumps of shoots in this population were clonal units. Plants (inc,luding clonal units or modules) of three-flower made up almost all of the herbaceous layer of a sugrberry-cedar elm-pecan bottomland forest. These plants grew closest to trees of pecan and eastern cottonwood on what decades earlier had been a small area of disturbance created by forces known but to God.

Three-flower melic is generally regarded as an extremely palatable forage species, but it is not a major forage producer escept at local scale as shown here. There is next to nothing about this species in the scientific literature beyond standard taxonomic treatment.

Bank of Bosque River, Erath County, Texas. Mid-April; peak development and biomass of shoots, peak anthesis.

 
   

 

255. Panicle on the bank- Sexual shoot with large panicle (first slide) and overall view of panicle (second slide) of three-flower melic on a large clonal plant gfrowing on immediate bank of Bosque River. These slides viewed together with the next two slides demonstrated the compound branching pattern of this species.

Erath County, Texas. Mid-April; peak stage of pollination.

 
   

 

256. Spikelets of two more than three- Although "three-flower" melic is appropriately named for most areas where plants of this species have spikelets mostly with three florets (Hitchccock and Chase, 1950,, p. 201) and , rarely, four (Barkworth et al, 2007, p. 100), this is not the case for most plants in Texas where spikelets more commonly have only two florets (so that the adjective "three-flower" is misleading [Gould, 1975, p. 64]). If one knows what to look for at an in-the-field, naked-eye view it can be readily seen in these slides that there are, in fact, only two florets in these spikelets.

Another instructive feature of florets presented in these two photographs was exertion of both stigmas and anthers. The compound branching pattern of three-flower melic was evident when these photographs were viewed in conjuction with the two imediately preceding phtographs of the entire panicle of this species.

Three-flower melic is a particularily showy native grass that makes a fine choice for landscaping with native plants. Three-flower melid is absolutely gorgeous when in flower as was readily seen from examples seen herein. It is also an extremely valuable forage species as noted earlier.

Bank of Bosque River, Erath County, Texas. Mid-April; full-flower stage of phenology..

 
   

 

257. Phytomers portrayed- Phytomers--the fundamental unit of the grass shoot consisting of two parts of consecutive nodes and the internode between them--of three-flower melic on the bank of Bosque River in understorey of a sugarberry-cedar elm-pecan bottomland forest.. Leaf blades, leaf sheaths, and leaf axil featured in slides one, two, and three, respectively.

Band of Bosque River, Erath County, Texas. Late April; peak shoot development and biomass.

 
   
 

258. First responders- Red sprangletop (Leptochloa filiformis), in foreground, and, in right background, cutleaf or lanceleaf groundcherry (Physalis angulata) growing on denuded soil of Bosque River. This river bank had been plowed and scraped by earth-moving equipment bank for street construction approximately three and a half months prior to time of photograph which was near later stage of the first warm-growing following destruction. These two annual species are typical pioneer species in old field (secondary) plant succession.

Other typical annual species that pioneered the denuded creek bank included hairy crabgrass (Digitrtia sanguinalis), buffalo-bur (Solanum rostratum), giant ragweed (Ambrosia trifida), lambsquarters (Chenopodium album), and Palmer's pigweed (Amaranthus palmerii). Secondary sucession at this location was treated above. Most of these early colonizers are common on partially denuded or habitually disturbed land, red sprangletop is even more of a disturbance-obligate species as it is more restricted to more devestated (severely degraded) land. For esample, hairy crabgrass thrives on overmowed lawns and heavily pastured barnyards, but red sprangletop seems to need greater disturbance such as plowed ground, at least in the area being considered here.

Erath County, Texas. Late October; peak standing crop and immature grain stage.

 
   

 

259. An annual for disturbed river bottom- Shoots of red sprangletop that colonized plowed ground along the bank of Bosque River in the West Cross Timbers of northcentral Texas. Red sprangletop is a tall, rank-growing, cespitose, eragrostoid grass. Specimens shown in these photographs exceeded three and a half feet in height (the panicle-bearing shoot in the first slide was bent over for photographic purposes). Both of these plants were growing on plowed and graded land on the bank of Bosque River at end of first warm-growing season following tillage.

Although red sprangletop is a typical weedy annual grass (much like the crabgrasses) it commonly grows to such large size--of whole plant as well as leaves/culms-- that it furnishs an abundance of fairly high-quality feed that it has good forage value for cattle and horses. Its livestock forage value and adaptation to colonize drastically disturbed land renders red sprangletop a desirable grass for flood-prone areas like the mixed hardwood bottomland forest recently devestated by careless construction workers on which these plants were growing.

Erath County, Texas. Late October (first slide) and mid-September (second slide); peak standing crop and immature grain stage.

 
   

 

260. Notable panicle- Overall view of sexual shoots (first slide) and features of panicles (second slide) of red sprangletop that grew on tilled soil along the bank of Bosque River. Red sprangletop has textbook characteristics of an r-selected species with a high allocation of its resources directed to ssexual reproduction. The large panicle of this annual bunchgrass commonly makes up one-third to over half of total shoot length.

In addition to desirable features of this generally large-sized annual grass (described in the preceding caption) the showy "over-sized" panicle makes this native species an attractive plant.

Erath County, Texas. Late October and immature grain stage (first slide); early September and mature (ripe) grain stage (second slide). Note that different plants of this annual species completed their life cycle at different points (dates) of time in the warm-growing season.

 
   
 

261. Another pioneering grass- Common witchgrass or witchgrass (Panicum capillare) is an native annual panicgrass--and not to be confused with fall witchgrass (Leptoloma cognatum)--that is a first invader of disturbed, especially drastically disturbed land such as that devegetated by tillage, extremely intense fire, heavy traffic, or extreme overgrazing. Witchgrass is thus a fellow colonizing species with the various crabgrasses, all of which are naturalized exotics; giant ragweed and horseweed or mare's tail, native annual composites; lambsquarters, a naturalized Eurasian chenopod; buffalobur, a native annual nightshade; and Texas millet, a fellow native panicgrass which unlike the previously listed is typically restricted to the deeper south of North America.(An example of colonization by these annual species of cleared land on bank of Bosque River was presented above.)

Along with the annual herbs just listed, witchgrass is the quitessential pioneering or colonizing species. All these species are textbook examples of r-selected species. Like the other annual pioneer grasses, witchgrass is fair to good in forage value depending on animal species. It is quite palatable to cattle and horses, especially at less mature states of maturity.

Witchgrass commonly breaks off at shoot bases so that the plant is blown to-and-fro across the land as a "tumble-weed" thereby dispersing seeds for the next crop of this annual grass, the opportunities for establishment varying from the next growing season to decades later when conditions are again favorable for this opportunistic, colonizing species.

Erath County, Texas. Mid-September, peak-bloom stage.

 
   

 

262. A pioneer plant up close- Details of leaves, clums, and panicles of the witchgrass plant that was introduced in the immediately preceding slide. The broad leaves of this annual panicoid grass are typically somewhat tattered by late season. The large-diameter culm is relatively weak and readily broken apart by weathering and decomposition so that it breaks off to become a "tumble-weed" with wind serving as a primary grain dispersal agent (anemochory).

Erath County, Texas. Mid-September, peak-bloom stage.

 
   
 

263. A common but atypical one- Plant of false nut-sedge or strawcolor flat (= umbrella) sedge (Cyperus strigosus) on a wet bank of a tributary into Bosque River. This particular umbrella or flat sedge has a large biological range tand it is one of the more striking grasslike plants in the Cross Timbers and Prairies vegetational area of Texas and an atypical pattern of inflorescence. For these reasons (and because the author had plenty of time to photograph plants under ideal conditions) strawcolored umbrella sedge was treated in detail herein.

Erath County, Texas. Late September, full bloom phenological stage.

 
   

 

264. .Showy representative of the grasslike group- Strawcolored flat sedge growing on bank of Bosque River. This species of umbrella sedge is a cespitose perennial with a large species range (Diggs et al., 1999, p. 1136). This individual was growing in the understorey of a sugarberry-cedar elm-pecan floodplain forest on the bank of Bosque River.

Erath County, Texas. Early September; immediate anthesis phase of otherwise full-bloom stage.

 
   

 

265. Beauty of a compound, umbel-resembling panicle- Panicluate units (groups of) spikelets (first photograph) and spikelets (second photograph) of strawcolored flat sedge or false nut-sedge. These floral units were on the same plant as introduced in the immediately preceding set of slides.

Bank of tributary of Bosque River, Erath County, Texas. Early September; immediate anthesis phase of otherwise full-bloom stage.

 
   

 

266. A confusing arrangement- An entire and atypical inflorescence of strawcolored unbrella sedge with primary and secondary peduncles above long, spreading leaves subtending it (first slide) and a unit of secondary peduncles (second slide) of this umbrella sedge. The key field identification feature of this Cyperus species is the yellow coloration and long, narrow spikelets. However in these specimens there was a division of longer pednucles with groups of spikelets at bottom of the inflorescence that then branched into shorter peduncles which also bore floral (spikelet) units. The only Cyperus species in this area that has this compound branching was C. elegans which has distinctly different spikelets from C. strigosus. C. strigosus does not develop such a secondary branching pattern Diggs et al. (1999, 1136), but these plants possessed this morphological feature. This pattern was similar to that of umbels; in this instance, umbel-resembling units arising from and situated above lower umbel-like arrangements. This author wondered if these plants could be hybrids between C. elegans, which was reported for this locality by Diggs et al. (1999, p. 1136) and C. strigosus which is extremely common throughout the eastern half of Texas (Diggs et al., 1999, p. 1144).

Erath County, Texas. Late September, full bloom phenological stage

 
   

 
267. Not exactly straw-colored- Shoot bases of strawcolored flat or umbrella sedge (known also as false nut-sedge) growing in the main channel of Bosque River. Erath County, Texas. Late September, full-bloom phenological stage.  
   

 

268. Cariced southerner- Southern caric sedge (Carix austrina) was one of several Carex species in the understorey of a sugarberry-cedar elm-pecan bottomland forest range. This caric sedge was presented here with at least (a minimum of) four plants in the first slide . The second slide provided a closer view of a larger specimen of C. austrina.

Bosque River floodplain, Erath County, Texas. Late April, peak shoot development, maximum biomass, and full-bloom phenological stage.

 
   

 

269. Southern spikes- Spike inflorescences of southern caric sedge. There was a total of five spikes in these three slides of C. austrina. This Carex species is in subgenus, Vignea.

Bosque River floodplain, Erath County, Texas. Late April, peak shoot development, full-bloom phenological stage

 
   

 

270. Sedge named after a grass- Local stand of fescue-like caric sedge (Carex festucacea) on the greater floodplain of a small river in northcentral Texas. Whereas C. austrina (presented immediately above) occurred in widely scattered patches of a few plants, C. festucacea grew in large, expansive areas over the same bottomland forest range.

Bosque River floodplain, Erath County, Texas. Late April, peak shoot development, maximum biomass, and full-bloom phenological stage.

 
   

 

271. Crowd of spikes- Numerous spike inflorescences of fescue-like caric sedge on the greater floodplain of Bosque River. This species of caric sedge often covered comparatively extensive areas of the land surface of this forest range. Fescue-like caric sedge was a major rather than an incidental range plant in this bottomland forest vegetation which was contrast to other Carex species in this herbaceous understorey. C. festucacea is in genus, Eu-Carex.

Erath County, Texas. Late April, immature achene stage of phenology..

 
   

 

272. Tickled Tweedy- Tweedy tick-clover or tick-trefoil (Desmodium tweedyi) growing on outer margin of the sugarberry-cedar elm-pecan bottomland forest that developed on the Bosque River floodplain. A number of plants of this species were growing at higher elevations above the river bottom on habitat similar to that of woodland or hairy wood borme. This is a beautiful species of native legume. Diggs et al. (1999, ps.655-656) listed twelve Desmodium species in their Flora of Northcentral Texas. The coloration pattern of the oblong to rhomboid trifoliate leaflets make this one of the most distinctive of these species.

All Desmodium species should be regarded as extremely palatable to almost all ruminants. A local herd of white-tailed deer routinely fed in this bottomland forest range and the author had difficulty finding sexual shoots of this species that had not been grazed off by these deer. (God provides: all shots of this species were "quail and manna from Heaven".)

Erath County, Texas. Mid-May, phenology varied from full bloom to ripening fruit.

 
   
 
273. Tweedly bloom- Inflorescence of Tweedy tick-clover or tick-trefoil. This plant was growing at a high point of the floodplain of Bosque River at outer edge of a sugarberry-cedar elm-pecan forest. Erath County, Texas. Mid-May, full-bloom phenological stage.  
   
 
274. Ticked up- Upper part of shoot of Tweedy's tick-clover bearing legumes. Desmodium species are in the same tribe, Hedysareae, as peanuts (Arachis spp.). Seeds of the tick-trefoil species have a peanut-like flavor. Bosque River floodplain, Erath County, Texas. Mid-May, fruit-ripening phenological stage.  
   
 

275. Tickled twice- Sessileleaf tick-clover or sessileleaf tick-trefoil (Desmodium sessillfolium) growing on the perimeter of a sugarberry-cedar elm-pecan bottomland forest. This plant was growing at the outermost edge of a mixed hardwood forest that was advancing (invading) an old field (abandoned cropland). Its more abundant herbaceous neighbors were Johnsongrass and purpletop.This was location was about 40 steps from the outermost bank of Bosque River in the West Cross Timbers.

Erath County, Texas. Mid-September, peak-bloom stage.

 
   

 

276. Sessiled on forest's edege- Details of inflorescence (flower cluster) and papilionaceous flowers of sessileleaf tick-clover that was growing at the advancing edge of sugarberry-cedar elm-pecan bottomland forest that was invading an old field. This was the same plant that was presented in the preceding slide.

Erath County, Texas. Mid-September, peak-bloom stage.

 
   

 

277. Alien forb in the Bosque Bottoms- Hedge parsley (Torillis arvensis), an annual Eurasian weed of the Umbelliferae, was the locally dominant forb in understorey of an elm (cedar and American)-pecan-sugarberry climax lowland forest. Plants of this species were at this stage of development in mid-spring. In a matter of a few more weeks these plants would be dead. The dead, fruit-bearing stalks remained to shed their tight-sticking schizocarps throughout summer and into autumn to start the life cycle of this naturalized cool-season species all over again.

Above the Bosque River, Erath County, Texas. May; full-bloom stage of phenology.

 
   

 

278.. Texas nightshade (Solanum triquetrum)- This distinctive forb was an occasional member of the herbaceous component of the sugarberry-pecan-cedar elm bottomland forest along Bosque River. Texas nightshade is much less common than other Solanum species of which there are three that commonly grow on Cross Timbers and Grand Prairie ranges in this area.

Bank of Bosque River, Erath County, Texas. November.

 
   

 

279. Inflorescences of Texas nightshade- Details of flowers of a member of the Solonaceae that was frequently associated with such herbaceous species as broadleaf woodoats, Canada wildrye, frostweed, white four o'clock, and giant ragweed.

Bank of Bosque River, Erath County, Texas. Mid-November.

 
   
 

280. Two of a family- Pokeberry or pokeweed (Phytolacca americana), at left, and pigeon-berry or pigeonberry (Rivina humilis), at right, in understorey of a sugarberry-cedar elm-pecan bottomland forest on the floodplain of Bosque River in West Cross Timbers. Phytolacaceae is a small family, mostly of forbs, in North America with only 11 species according to Flora of North America. Two of the major species frequently grew side-by-side on this mixed hardwood forest range. Other plant species in this "photo-plot" included hairy crabgrass, sideoats grama, and Johnsongrass.

Erath County, Texas. Late September.; fruit-ripe stage in pokeberry and full-bloom stage in pigeonberry.

 
   
 

281. Pigeon-berry (Rivina humilis)- Whole plant of pigeon-berry in pre-bloom stage. Along with frostweed (= white crownbeard) and giant ragweed, pigeon-berry was a major warm-season forb on the sugarberry-pecan-cedar elm bottomland forest that developed along Bosque River. This perennial forb was also well-represented in parts of this forest high above the river. It grew in both climax herbaceous communities dominated by Canada wildrye and/or broadleaf woodoats as well as in recently disturbed local sites where it was commonly associated with giant ragweed.

There is considerable variation in size of individuals of this species. This specimen was about two and a half feet tall. Plants of pigeon-berry (a member of the Phytolaccaceae) in this bottomland forest were much smaller than those of the large (and much less abundant) pokeberry or pokeweed (Phytolacca americana) that also grew in this herbaceous undertstorey.

Bank of Bosque River, Erath County, Texas. Early November.

 
   
 
282. Prime age- Adult plant of pigeon-berry at mid-bloom stage on floor of a sugarberry-cedar elm-pecan bottomland forest. Floodplain (bank) of Bosque River, Erath County, Texas. Mid-May.  
   

 
283. Pigeon-berries in the making- Upper shoot (first photograph) and inflorescence at peak bloom (second photograph) of pigeon-berry (Rivina humilis). These slides were of a plant neighboring the plant shown in the preceding slide. Both were growing in the herbaceous layer of a sugarberry-cedr elm-pecan bottomland forest that developed on the floodplain of Bosque River in the West Cross Timbers. Erath County, Texas. Mid-May, peak-bloom stage.  
   

 

284. Sexual shoots of pigeon-berry- Details of leaves and fruit of white four o'clock growing in the understorey of a sugarberry-pecan-cedar elm bottomland forest. The fruit was interpreted as a berry by Diggs et al.(1999, p. 881) with a single seed per fruit (the ovary is unilocular but with only one basal ovule). This appeared as a contradictary description. Smith (1977, p. 290) defined berry as "a multiseeded, inndehiscent fruit in which the pericarp is fleshy [soft] thoughout" . Fernald (1950, p. 606) provided a more coherent description when it was explained that in Phytolaccceae the ovary was multi-locular and consisted of numerous carpels arranged in a ringlike structure all of which formed a berry.

Bank of Bosque River, Erath County, Texas. Early November; fruit-ripe phenological stage.

 
   
 
285. Hay fever source up close- Close-up of staminate flowers of giant ragweed. Bank of Bosque River. Erath County, Texas. Mid-October; anthesis.  
   

 

286. Frostweed or white crownbeard (Verbensina virginica)- This composite is a locally dominant forb in understorey of sugarberry-pecan-cedar elm forests in this West Cross Timbers region. This species occurs over a region extending from Iowa south and west to Texas, including parts of the Cross Timbers, Edwards Plateau, and Coastal Prairies and Marshes regions. In this forest range type frostweed was found on stable microenvironments with such climax perennial grasses as broadleaf woodoats and Canada wildrye, the two dominant herbaceous species in the forest understorey.

Erath County, Texas. September; full-bloom stage of penology.

 
   
 

287.. Inflorescence of frostweed or white crownbeard- Flower cluster of the dominant perennial composite of a sugarberry-pecan-cedar elm forest in the West Cross Timbers. Giant ragweed is another member of the Compositae that is locally dominant (one of three major warm-season dominant forbs) in this forest range vegetation, but this species is an annual and was generally restricted to disturbed microsites especially freshly eroded banks of Bosque River.

Erath County, Texas. September; full-bloom stage of phenology.

 
   

 

288. Prickley poppety, bobbity blue, along with white and yeller too- White or Texas prickly-poppy (Argemone albiflora var. texana) of floodplain of Bosque River. This species and A. polyanthemos are extremely similar with main distinction being descriptive--and highly subjective--leaf features: degree to which leaves are divided to the midrib, the qualitative thiness or thickness of leaves, and degree of glaucousness of leaves (Diggs et al., 1999. p. 873). Correll and Johnston (1979, ps. 666-667) stated that shoots of A. albiflora were usually solitary in contrast to those of A. polyanthemos that ranged from single to five shoots. After viewing many photographs of living and mounted specimens of both these species this author noted that unopened flower buds of A. polyanthemos were vertically oriented while those of A. albiflora were horizontal in orientation. The single-stemed specimen seen here with horizontal floral buds and deeply cut leaves seemed--and quite convincingly so--to be Argemone albiflora.

Texas or white prickly-poppy were not plentiful along Bosque River, but they are a characteristic species of this floodplain vegetation.

Erath County, Texas. Late April; mid-bloom stage.

 
   

 

289. Enough for a poppy high- Views of the inflorescence of white or Texas prickly-poppy. The insect amid stamen was common flower or bumble flower scarab beetles (Euphoria kernii) which is usually more abundant in flowers of pricklypear cactus (Opuntia spp.), but still at home feeding on these anthers which are often eaten off near base of filaments. .

Erath County, Texas. Late April;

 
   

 

290. Plant that ain't no gift- Christmas mistletoe (Phoradendron tomentosum= P. flavescens) growing on sugarberry. Christmas mistletoe is a parasitic plant that is not completely parasitic. It does conduct some photosynthesis (ie. semiautotrophic), but not enough for its survival. Itstead each mistletoe plant gradually weakens the host as both plants grow. Meanwhile, seeds rom mistletoe fruit germinate on other locations of the host so that infestation of the parasite spreads through new (younger, smaller) plants, many of which were dispersed by fruit-eating birds. As parasite load increases the host is weakened until if finally succubs to the parasitism, at least in most cases. In other instances, especially where the environment is less favorable for Christmas mistletoe, the host may reach maturity and senescence before being killed by the parasitic mistletoe. Even under these consitions Christmas mistletoe causes some deformity of the host tree through lost or weakened limbs and branches.

Erath County, Texas. Mid-March; peak bloom phenological stage.

 
   

 
291. Spiked female-Female spikes of the monecious Christmas mistletoe. Erath County, Texas. Mid-March; peak bloom phenological stage.  
   

 
292. Pistillate details-Close-in views of flowers on pistillate spikes of Christmas mistletoe. Erath County, Texas. Mid-March; peak bloom phenological stage.  
   

 

293. Smooth or sugar hackberry, sugarberry, or palo blanco (Celtis laevigata)- This is a widely distributed species (Virginia to Florida westward to New Mexico and northward to Illinois) with many recognized varieties and much ecotypic variation across it's geographic range. It grows on fertile bottomlands, where it frequently reaches large dimensions, and on infertile, generally harsh habitats such as shallow rock outcrops where the species survives at dimutive sizes. There has been considerable author-to-author variation in treatment of taxa below species level.

Sugar hackberry is a common component of various range types and sites. In Texas, for example, it is frequently a dominant woody plant in various range communities in vegetational areas as diverse as the Cross Timbers, the Edwards Plateau, and the Rio Grande Plains. Sugarberry occurs singly and in small thickets through parts of adjoining vegetational areas like that of the Coastal Prairies and Marshes. It is typically the major dominant tree species in river bottom forests throughout much of southern North America.

Celtis species provide fruits that are of value to species of smaller wildlife like birds and rodents. Their twigs, buds, and leaves are taken less frequently by browsing animals. Sugarberry and netleaf hackberry (C. reticulata= C. laevigata var. reticulata) are usually more important as brush species (noxious woody plants) whose cover and density increased under disturbances that induced range deterioration such as overgrazing, underburning, oil and gas development, and abandonment of farm land. In other words hackberry, like most of the other shrubs in this section, are invaders rather than climax species or potential natural dominants of range communities in higher seral states. They are typically indicator plants: indicative of range retrogression.

Mills County, Texas. October.

 
   

 

294. From last year and for this year- Two leaders of sugarberry, smooth hackberry, or palo blanco (Celtis laevigata var. laevigata) with last year's fruit and current year's pistillate and staminate flowers. New leaves are about three/fourths growth. Details of unisexual flowers were shown in the next two sets of photographs.

West Cross Timbers, Erath County, Texas. April.

 
   

 

295. Spring time for sugarberry- Two views of the pistillate and staminate flowers of smooth hackberry, sugar hackberry, or sugarberry. The first slide gave a general view of the inflorescences (along with new spring leaves) of Celtis laevigata var. laevigata. The second slide showed details of the flowers of this monoecious species (all Celtis species are monecious) with pistillate (female) flowers at left and staminate (male) flowers to the right along on portion of a leader.

West Cross Timbers, Erath County, Texas. April.

 
   

 

296. "...male and female created them" (Genesis 1: 27 KJV)- Two slides of sexual reproduction in sugarberry, smooth hackberry, or southern hackberry. In both photographs female (pistillate) flowers at left and male (staminate) flowers at right on leader. A corolla is lacking in pistillate flowers though there is a calyx of five or six sepals. Gynoecium (generic term for female part orf flower; carpels collectively) is unicarpellate (each ovary has one suspended ovule) with two stigmas. The androecium (general term for male part of flower; more specifically, stamens collectively) consist of five or six stamens.

West Cross Timbers, Erath County, Texas. April.

 
   
 

297. Pair of drupes- Two drupes on sugarberry, smooth or southern hackberry (Celtis laevigata var. laevigata). This fruit was on a leader that was bearing just-opened monoecious flowers. Drupes were also in the immediately preceding two photographs where their close proximity to newly opened flowers was shown. Fruit got sole emphasis in this shot. Drupe is a fruit type defined as being fleshy and indehiscent with a single seed inside a stony or boney endocarp, the innermost layer of the fruit wall (Smith, 19789, p. 65, 69).

West Cross Timbers, Erath County, Texas. April.

 
   
 

298. Catchy branching pattern in sugarberry- A combination of alternate branches coming off of opposite sides of a leader is a frequent branching arrangement in Celtis species, especially C. laevigata. Perhaps this pattern permits large numbers of leaves with a minimal degree of shading so as to facilitate photosynthesis in forest canopies. Would make a eye-catching pattern for neck ties, particularily for dressed-up foresters.

Erath County, Texas. September; fruit-ripe stage.

 
   

 

299. Shiny leaves and sweet fruit (well, sort of)- Leaves and fruit of sugarberry shown in photographs that featured leaf position and density along with a better-than-typical crop of fruit. Fruit type in Celtis species is a drupe. Fernald (1950, p. 553) remarked that these sweet-tasting drupes were basis of the name Celtis as used by Pliny for what he termed lotus. Anyway, wildlife of various species consume this fruit and undoubtedly contribute to dispersal of Celtis species.

Erath County, Texas. September; fruit-ripe stage.

 
   
   

 

300. Early sexual stage- Clusters of flowers on leader of American or white elm (Ulmus americana). This species flowers before it puts forth leaves.

Erath County, Texas. Early March; anthesis.

 
   

 

301. Later sexual stage- Clusters of flowers on leader of American or white elm one week after the stage (anthesis) shown in than preceding pair of slides. Samaras were just forming.

Erath County, Texas. Mid-March; early fruit stage.

 
   
 

302. Young leaves and fruit- Newly emerged leaves and growing samaras of American or white elm. Samara is a dry fruit type that is indehiscent an single-seeded with a large wing.

Erath County, Texas.Late March; immature fruit stage of phenology.

 
   
 

303. Fruit of a real American- Ripening fruit of American elm (Ulmus americana) on twig. The fruit elms is a samara or a winged achene. More specifically the Ulmus fruit type is a dry, indehiscent, single-seeded fruit fitted with a prominent wing (Smith, 1977, ps. 66, 90, 307). Cedar elm produces its samaras in later summer or early autumn, but American (white) elm and slippery (red) elm bear their fruit first thing in late winter to early spring befgore emergence of leaves.

Erath County, Texas. Mid-March (late winter), fruit-ripening phenological stage.

 
   

 

304. Enter, cedar elm (Ulmus crassifolia)- Leaves and samaras (or winged achenes) of cedar elm. In contrast to American or white and slippery or red elm (U. rubra) cedar elm bears it fruit at end rather beginning of it's growing season so that leaves and fruit are present together. On mixed hardwood bottomland forests in much of Texas, especially central and south Texas, it is cedar elm and not American or red (slippery) elm that is a dominant. On such forests cedar elm is a "three-way" dominant (would that be a "tri-dominant'?) with sugarberry and pecan.

Floodplain of Bosque River, Erath county, Texas. Mid-September; fruit ripe stage (immediately preceding shedding stage).

 
   

 

305. Green ash (Fraxinus pennsuylvanica var. integerrima)- Green ash is a species (or, sometimes, subspecies or varierty) treated differently by various authors such that confusion has arisen frequently as to whether or not this is the plant found by different vegetation scientists. Nonetheless, it is a taxon that defines certain forest vegetation, especially bottomland forest types such as the sugarberry-American elm-green ash (forest cover type 93) of the Society of
American Foresters ( Eyre, 1980, ps. 65). In bottomland forests of this type (and also SAF 94, sycamore-sweetgum-American elm, where green ash is the first-listed associate species [Eyre, 1980, ps. 65-66]) in the south, such as much of Texas, green ash is an infrequent yet a defining and diagnostic species. Such was the situation for the bottomland forest range in this example of the Bosque River.

The small and showy specimen presented here stuck out like the proverbial sore thumb in this forest dominated by sugarberry, pecan, and cedar elm-- even during the autumnal aspect when leaf color of dominants is at its "prime" (scroll back up to above photographs of the north bank of Bosque River).

North bank of Bosque River, Erath County, Texas. Early November; peak autumn leaf coloration.

 
   

 

306. Winter twigs of pecan (Carya illinoensis)- Examples of winter twigs of pecan growing on a climax sugarberry-pecan-cedar elm bottomland forest in West Cross Timbers of northcentral Texas. Winter twigs, especially features of lenticels, leaf scars, and terminal buds provide some of the most definitive means of distinguishing among the various Carya species growing in a given locality. The narrow (thin) terminal bud often with "peeled-up" or peeling bud scales is a common feature of pecan twigs in their winter phase.

Top of upper bank of Bosque River, Erath County, Texas. January; winter dormancy.

 
   
The presence of members of Moraceae, bois d'arc (Maclura pomifera) and red mulberry (Morus rubra), clearly showed the affinity/continuity of this forest vegetation with riverine forest flora northward to southern Nebraska. Likewise, abundance of cedar elm (Ulmus crassifilia) as co-dominant to associate
showed ecological connectedness of this forest community with river forests to the north in which white or American elm (U. Americana) and/or red or slipper elm (U. rubra) had role as dominants (ie ecological equivalents).
 
   

 

307. Bois d'arc, Osage orange, or hedge apple (Macula pomifera)- Bark and fruiting organ of bois d'arc, a defining associate woody species, in centrtal bottomland forests from the south as far north as southern South Dakota (McGregor et al, 1977, p. 34) and extending eastward into the deciduous forests from Texas and Arkansas to New Enmgland (Fernald, 1950, p.555). Strictly speaking the fruit of Osage orange is an achene, numerous of which are "buried in the greatly enlarged fleshy calyx (Fernald, 1955; p. 555).

The name Osage orange traces to the traditional use of wood of this species for making long bows by American Indians, presumedly especially the Osages. Hedge apple traces to the planting (and extended introduction) of this species as a living (and suposedly impenetrable) hedge to fence in (or out) livestock, especially cattle and horses. Such usage was of "mixed success", but after adoption of barbed wire as developed and sold by Joseph Glidden, the unparalled durability (rot-resistance) of this species for fence posts enshrined its designation as hedge apple or simply hedge (hence, hedge posts). Horse apple, another common name, indicates the frequent tendency of horses to eat the fleshy calyx, often with dire to deadly consequences in cases where this structure lodges in the esophagus and results in closure of the trachea and suffication of the careless equine.

Bank of Bosque River, Erath County Texas. Mid-Novermber, fruit-ripe stage.

 
   

 

308. Male attire- Leader of male boisd'arc with clusters of staminate inflorescences. Osage orange is a dioecious species. A female and her fruit was shown immediately above. Here the male of this species was presented for "closer inspection". Although they were still young and tender the leaves were typical of Macula pomifera.

Bosque River, Erath County, Texas. Mid-April, early bloom phenological stage..

 
   
 
309 Opening flowers against bark- Young (just opened and opening) staminate infloresences of bois d'arc laid onto characteristic bark of this species. Bosque River, Erath County, Texas. Mid-April.  
   

 

310. After Injuns, hosses, and hedges- Osage orange, horse apple, or hedge apple are commonly used descriptive names for the fruit of bois d'arc, Osage orange, hedge apple, etc. This is an example of multiple fruit, the fruit type formed by fusion or coalescence of the entire flower cluster (inflorescence). It is one form of a so-called false fruit (Smith, 1977, ps.69, 301). Sycamore is another tree species of bottomland forests that produces a multiple fruit though that false fruit is dry in contrast to the sap-filled (and extremely sticky and persistent) fleshy multiple fruit of bois d'arc. By some interpretations the multiple fruits of Moraceae (mulberry family) are composed of minute drupes called druplets with the whole multiple fleshy fruit designated a syncarpet.

This hardwood species has been associated more with members of the Osage Nation than of other North American Indians because it was native to land controlled by the Osages. Bois d'arc is an Anglicization of the French "wood of the bow" or "bow wood". Numerous tribes in addition to the Osage used the wood of bois d'arc for long bows and war clubs. (It is unlikely the Osages got hungry enough to try to eat this fruit.) Osage orange was indigenous only to what is today parts of Texas, Arkansas, and Oklahoma. It was spread by white settlers seeking to use it as a living fence against otherwise free-ranging livestock, especially cattle.

Livestock, including horses, rarely attempt to eat this fruit which has resulted in choking (obstruction of the esophagus), hence the common name of horse apple.

Bosque River, Erath County, Texas. Mid-September, fruit-ripe stage.

 
   

 

311. Osage orange halved- Cross-section of the multiple fruit of bois d'arc or Osage orange. Seeds are in individual carpels that fused to form this fleshy multiple fruit sometimes known as a syncarpet (from the adjective, syncarpous meaning a gynoecium comprised of fused carpels (Smith, 1977, p. 310). Usually this fruit is used as feed by very few animals with notable exceptions like squirrels. Attempts by cattle and horses to eat this large fruit have not uncommonly resulted in choking from esophageal obstruction.

Horse apples, hedge apples, Osage oranges (or by whatever other common names are applied to this fruit) were commonly placed in houses as insect and spider repellents based on folk treatment or home remedy recommendations. At least they had decorative value to some (at least until they started to rot down).

Bosque River, Erath County, Texas. Mid-September, fruit-ripe stage.

 
   

 

312. Smaller trunk on bottomland- Nice, straight trunk (bole) of a young adult red mulberry (Morus rubra) on the outer bank of a small river in the West Cross Timbers of northcentral Texas. General niche of this native, small tree was discussed above. Species range of red mulberry extends from Massachusetts across to South Dakota then southward to Trans Pecos Texas and eastward to south Florida (Sargent, 1933, p. 329-330). Red mulberry along with bois d'arc (both are members of Moraceae) comprised a small tree zone below the crown canopy of the three forest dominants, sugarberry, cedar elm, and pecan. Large trunks in background were those of large, senescing pecan; individual trees of which grew to the largest size among three dominant tree species in this forest cover type .

Bosque River, Erath County, Texas. Mid-May.

 
   

 

313. Big leaves along the Bosque- Outermost (distal) portion of a lower limb of red mulberry growing on the bank of Bosque River. The characteristic large leaves of this species are generally ovate with a somewhat rounded base and pointed apex. Leaves of red mulberry can be be lobed (all of these were unlobed). Margins are serrate.

Tree trunk at left in second photograph was cedar elm supporting the woody shoot of a large truppet creeper.

Erath County Texas. Late October.

 
   
 

314. Male doing his thing- A large, male red mulberry with young leaves and catkins at peak anthesis with backdrop of a rare, cobalt-blue, spring sky. Like its moraceous "cousin", bois d'arc, red mulberry is a dioecious species. Likewise, both of these members of the Moradceae were associate tree species in a sugarberry-cedar elm-pecan bottomland forest that developed along a river in the West Cross Timbers.

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

 
   

 

315. New garb and exposed male parts- Young leaves and just-past-peak pollen production staminate catkins of red mulberry that was an associate tree species of a sugarberry-cedar elm-pecan bottonland forest.

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

 
   

 
316. Male detail- Staminate catkins of red mulberry with baby leaves (first photograph) and laid against red mulberry (second photograph). The latter had been shed from the tree. Floodplain of Bosque River, Erath County, Texas. Mid-April.  
   

 

317. Satisfied with leaves and fruit- Part of crown (first slide) followed by terminal portion of twig both topside--abaxial--or above (second slide) and bottom side (adaxial) or below (third slide) of red mulberry growing in a mixed hardwood bottomland forest in northcentral Texas.

Floodplain of Bosque River, Erath County, Texas. Mid-May; immature fruit to fruit-ripening phenological stages.

 
   

 

318. Ripening underneath- Underside (adaxial surface) of leaves with both immature (white) fruit and ripening (wine-colored) fruit of red mulberry growing on outer bank of a small river in West Cross Timbers of northcentral Texas. Fruit type of mulberry is multiple fruit consisting of "... achenes covered by sweet, juicy, white to dark purple calyces" where calyx refers collectively to sepals, the outer bracts of the flower structure or the modified leaves that enclose the other floral parts, (Diggs et al, 1999, ps.831, 1426).

Floodplain of Bosque River, Erath County, Texas. Mid-May; immature fruit to fruit-ripening phenological stages.

 
   

.

 

319. Fruit and feeding friends- Hackberry emperor or hackberry butterfly (Asterocampa celtis; family of brushfooted butterflies, Nymphalidae) along with some species of beetle and, of course, housefly (Musca domestica) feeding on the juicy calyces surrounding achenes in the multiple fruit of red mulberry growing on the outer bank of a small river in a mixed hardwood bottomland forest in the West Cross Timbers of northcentral Texas. At time of this photograph there were numerous hackberry emperor butterflies throughout this floodplain forest. They appeared to be most common around trees of red mulberry such as the tree featured in this section. Adults of Asterocampa celtis and the closely related A. leilia feed on juicy fruits (both fresh and rotten) as well such as high-moisture feeds as dung and rotten flesh of vertebrates, but seldom visit flowers (Neck, 1983). Langlois and Lanflolis (1964) reported that A. celtis feed on mulberries.

It is generally recognized that Celtis species (apparently mostly leaves) are the host feed plant for caterpillars of A. celtis and A. leilia. This was even indicated by the common name of hackberry butterfly or hackberry emperor. (It is standard convention to include the common name of larval host food plant as part of the common name of butterflies.) Sugarberry, the most common species of hackberry in southcentral North America, was the first among three dominant tree species in this floodplain forest. Not only are Celtis species the main food source for larvae of Asterocampa species, especially hackberry emperor, but, as described in detail by Rutowski et al. (1991), males of A leilia perch and defend mating territories in desert hackberry (C. pallida).

Many animal species including insects, birds, reptiles such as turtles, and mammals (even humans) feed on the multiple fruit of red mulberry. Over many years of observation the current author concluded that mulberry fruit is one of the most attractive feeds to song birds (and, once fallen on the ground, to poultry) and rodents such as squirrels. He also observed white-tail deer and, with somewhat less dexterity, cattle consuming mulberries. Sargent (1933,.p. 330) claimed that red mulberry fruit was "valued for fattening hogs", and even "occasionally planted" for this purpose. The current author questioned the fattening part, but assumed that swine also found mulberry fruit quite palatable (and much less likely to be met up with again than drupe fruits like peaches).

Mini lesson: in a nontraditional sense the hackberry emperor butterfly could be viewed as a range animal on this forest range. Given that Asterocampa species have been found to feed on animal materials, included carcases, as well as vegetable matter (both browse as larvae and mast as adults) they are, by definition, omnivores.When it is remembered that there are more species of insects (Order, Insecta) than of all other living things combined, the rationale for presentation and discussion of insects in this slide-caption set (note that in this example three orders of inscets were feeding on one multiple fruit) becomes obvious. Range ecosystems are complex.

Floodplain of Bosque River, Erath County, Texas. Mid-May; mature fruit (fruit-ripe) stage of phenology.

 
   
 

320. Blooming above the branch- Rusty or southern black-haw (Virburnum rufidulum) on the west bank of Dry Branch that drains into Bosque River (shown in detail above). Virburnum includes some of the largest members of the Caprifoliaceae in North America. Several of the Virburnum species are understorey shrubs in deciduous forest communities. V. rufidulum is one of the most common of these in southern forests as indicated by the common name of southern black-haw. It is also one of the earlier blooming woody plants in the range vegetation of which it is a component.

Bosque River, Erath County, Texas.Late March (early spring).

 
   
 
321. Spring shoot- Blooming vernal leader of rusty or southern black-haw. This branch was on the plant introduced in the preceding slide. Bosque River, Erath County, Texas.Late March (early spring).  
   

 
322. White flowers on a black-haw- Inflorescence of rusty black-haw. Flower cluster on the same plant shown above Dry Branch into Bosque River. Erath County, Texas.Late March (early spring).  
   

 

323. Blue fruit of black haw- Clusters of fruit (one-seeded drupe) of rusty or southern black haw. The umbel arrangement of flowers is carried through to the fruit-bearing stage resulting in colorful "bunches" of pleasant-tasting drupes. Wildlife--from deer and squirrel to passerines--are extremely fond of this fruit.

Ottawa County, Oklahoma. November; ripe fruit phenological stage.

 
   

 

324.. Leaves of a southern arborescent shrub- Southern or rusty black-haw or southern blackhaw or, sometimes written as black-haw, (Viburnum rufidulum) was one of the few upright shrub (non-climbing shrub, non- woody vine) species in this sugarberry-pecan-cedar elm forest. Rusty black-haw was a distant "back-aways" runner-up behind such shrubs as common greenbriar (Smilax bona-nox), overall dominant, poison oak or ivy (Rhus radicans= Toxicodendron radicans), and mustang grape (Vitis mustangensis). In contrast, rusty black-haw was more common than eastern redbud (Cercis canadensis), smooth sumac (Rhus glabra), Virginia creeper (Parthenocissus quinquefolia), and trumpet creeper (Campsis radicans) in cover, density, etc. (general relative abundance) in this range vegetation. While this shrub was not common in this plant community its presence both here and in bottomland forests through central North America (southern Ohio, Indiana, and Illinois) and extending from the Ozark Plateau through the Applachian Mountains almost to the Atlantic Coast in Virginia and North Carolina stood as testament to the affinity of this species for bottomland forests over a vast territory. In more humid areas such as the Ozarks and Applachians rusty or southern black-haw grows under oaks and hickory on uplands and with sugar maple and beech on north slopes. In short, rusty black-haw is adapted to diverse habitats and forest communities.

The leaves on this particular specimen were less lanceolate (generally more ) than is typical of this species. Still, these leaves were typical enough (including their glossy surface) to facilitate ready field identification-- especially when augmented by the unique texture of bark on the trunk as shown in the next slide.

Above Bosque River, Erath County, Texas. Early Noverber.

 
   
 

325. Large trunk of a southern shrub- Bole of southern or rusty black-haw with the characteristic and very distinctive bark pattern of this species. This was another (a larger, so preseumedly, older) plant than the one introduced and detailed above. This one was growing on the north bank of Bosque River.

Erath County, Texas.Late March (early spring).

 
   
 

326. Trunk of rusty or southern black-haw (blackhaw)- The knobby bark of this species is very distinctive and a foolproof, identifying characteristic of this species. This feature and its glossy leaves along with general morphology makes this species obvious in the shrub layer of forests.

Above Bosque River, Erath County, Texas. Early November.

 
   

 

327. A deceivingly attractive alien- Inflorescence of Japnese honeysuckle (Lonicera japanica) blooming on the floodplain of Bosque River. Japanese honeysuckle is one of the widespread and insideous, introduced woody invaders throughout much of the Eastern Deciduous Forest Region of North America. Rosiere et al. (2013) described the threat this naturalized woody vine poses for forest of the southcentral portion of this forest super-region

Japanese honeysuckle was introduced and widely distributed by nurserymen to homeowners who quicly fell in love with the fragerant scent and hardiness of this easily established exotic. Japanese honeysuckle has not yet reached the horror movie-like invasiveness of some other intentionally introduced exotics, but it is widespread and one of the most more aggressive of woody aliens in southeastern North America. For example, Japanese honeysuckle was the most abundant (cover and density) non-native woody species in the sugarberry-cedar elm-pecan bottomland forest described in this section (Rosiere et al., 2013).

Upper bank of Bosque River, Erath County, Texas. Peak-bloom phenological stage; mid-April..

 
   
 

328. Basis of it all- Profile of Bunyan soil series that was the predominant soil of the sugarberry-pecan-cedar elm bottomland treated in this section. Bunyan series consist of fine-loamy, mixed, nonaacid, thermic typic ustifluvents (Soil Conservation Service, 1990). Range site was Bottomland (Soil Conservation Service, 1973).

Erath County, Texas. July.

 
   
Example of an Ecotonal Hardwood Forest in the Humid Zone
 
In the transition belt between the Texas Pineywoods and Gulf Coast Prairies-Marshes vegetational areas there are several forms of hardwood forests, some with and others without a coniferous element. One example of these ecotonal or transition forest was presented in the synopsis showing that follows. Average annual precipitation is 58 inches and with average temperature of 74 degrees Fahrenheit some authories interprete this as a form of subtropical climate. While climate is maritime (or, at least, maritime as to prevailing atmospheric influence), the forest vegetation was clearly a hardwood-dominated community having a prominent woody vine component with plant species which were essentially the same as those of interior forests in continental climate.  
   
 

329. Outlier of Texas Big Thicket- Outer edge of a sugarberry-sweetgum-green ash forest in a transition zone between pine-mixed hardwoods forest and a cordgrass (Spartina spp.)-dominated marsh-prairie. This hardwood forest qualifies as a margin of the Big Thicket only as to location and proximity to the general East Texas Timberlands. Forest range vegetation represented by this second-growth forest of climax species composition was an ecotone of the Texas-Louisiana Pineywoods and the western Gulf Coast Prairies and Marshes vegetational areas (= zonal vegetation).

Tthe two foremost trees were green ash, which in this forest, was a major and defining climax species along with sugarberry and sweetgum which were the most common (abundant) trees in this forest which was a vriant of the general sugarberry-American elm-green ash. forest cover type described by the Sopciety of American Foresters (Eyre, 1980, p. 65). The shrub that was most distinctive, especially in combination with the three dominant tree species, (and most conspicuous) was dwarf palmetto (Sabal minor). This monocotyledonous (and relatively) small shrub demonstrated the affilitation of this forest range type with the general Pineywoods zonal vegetation. Palmetto is not generally a component of the coastal prairies and marshes. Other prominent shrubs in this forest were mostly lianas (woody vines) including rattanvine or Alabama supplejack, trumpet creeper, greenbriar (Smilax sp[p].), peppervine (Ampelopsis arborea), yaupon (Ilex vomitoria), and poison oak/ivy.

There was a sporadic though well-developed herbaceous component (more of an irregular rather than continuous layer) made up mostly of broadleaf woodoats, one or more unidentified Paspalum species, and southern waxy cric sedge (Carex glaucescens). The most abundant forbs were seacoast goldenrod (Solidago sempervirens) and giant ragweed. Commonness of this annual composite and broadleaf woodoats along with the dominant sugarberry clearly showed the floristic and structural affinity of this forest with the sugarberry-cedar elm-pecan forest discussed above.

Jefferson County, Texas. Early October, autumnal aspect. FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash); sweetgum as one of the dominants shared some features of SAF 94 (Sycamore-Sweetgum-American Elm). Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Western Gulf Coastal Plain-Northern Humid Gulf Coastal Prairies Ecoregion, 29c (Griffith et al., 2004).

 
   
 

330. Defining members of a unique variant form- Green ash (the large featured tree) and dwarf palmetto (conspicuous at base of ash) in combination with sugarberry and sweetgum (the other dominant tree species) defined this forest vegetation as a variant of the general sugarberry-dominated forest cover type that is widespread throughout much of Texas and much of southeastern North America. This was a second-growth forest of small to mid-size trees but it obviously had the species composition and structure of the climax stage of this forest range type.

In addition to dwarf palmetto other shrub species included trumpet creeper, rattanvine or Alabama supplejack, poison oak/ivy, greenbriar, peppervine, and yaupon. The predominance of woody vines was striking. This was a discontinuous herbaceous layer consisting primarily of broadleaf woodoats, southern waxy caric sedge, and paspalums with two composites as the major forbs (one a perennial, seacoast goldenrod, and the other the common annual, giant ragweed).

Jefferson County, Texas. Early October, autumnal aspect. FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash); sweetgum as one of the dominants shared some features of SAF 94 (Sycamore-Sweetgum-American Elm). Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Western Gulf Coastal Plain-Northern Humid Gulf Coastal Prairies Ecoregion, 29c (Griffith et al., 2004).

 
   
 

331. Composite "scenery"- General view of a sugarberry-sweetgum-green ash forest range in a transition zone between Pineywoods and Coastal Prairies and Marshes vegetation. As to relative cover and density green ash was more of an associate species in this particular tract of forest. This was a second-growth forest of more-or-less climax composition and it seemed plausible that green ash would become generally and relatively more important than sweetgum, a pioneering species that typically persist into the climax forest.

In addition to the three dominant trees, dwarf palmetto was a conspicuous shrub that was joined by yaupon and liana species including rattanvine or Alabama supplejack, poison oak/ivy, trumpet creeper, greenbriar, and peppervine. The discontinous herbaceous layer included broadleaf woodoats (generally the most abundant herb), southern waxy cric sedge, paspalums, giant ragweed, and seacoast goldenrod. Age and size of trees at stage of young maturity combined with a relatively low density of these trees resulted in a fairly open forest having well-developed shrub and herbaceous strata. A natural regime of periodic surface fire could be expected to maintain this forest structure. Conversely, fire suppression would likely lead to greater canopy closure and less-developed lower layers of range vegetation. This would probably accompanied by reduced cover and density of sweetgum, a pioneer species, and (as mentioned above) greater relative cover of green ash, a less Intolerant tree species.

Jefferson County, Texas. Early October, autumnal aspect. FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash); sweetgum as one of the dominants shared some features of SAF 94 (Sycamore-Sweetgum-American Elm). Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Western Gulf Coastal Plain-Northern Humid Gulf Coastal Prairies Ecoregion, 29c (Griffith et al., 2004).

 
   
 

332. Unmistakable trunks- The blotched gray bark of sugarberry (foremost bole) and and finely furrowed, brown bark of sweetgum (trunks in background) bespoke the unmistakable dominance of these species in a climax composition, second-growth forest. Sweetgum and dwarf palmetto were some of the more obvious indicators that this range plant community was an ecotone between Big Thicket Pineywoods and Gulf Coast Prairies-Marsh vegetation. Numerous plants of giant ragweed indicated the widespread biological range and adaptation of this annual, weedy composite. Abundance of a pioneer or colonizing forb (though as smaller, stunted-appearing plants) was likely still possible because the canopy of this second-growth forest had not closed sufficiently to exclude critical quantities of light from lower levels of this forest range.

Jefferson County, Texas. Early October, autumnal aspect. FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash); sweetgum as one of the dominants shared some features of SAF 94 (Sycamore-Sweetgum-American Elm). Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Western Gulf Coastal Plain-Northern Humid Gulf Coastal Prairies Ecoregion, 29c (Griffith et al., 2004).

 
   
 

333. Featuring the lower levels- Floor of the sugarberry-sweetgum-green ash forest described above. Species makeup and structure of the lower layer of shrubs (or lower part a shrub layer) and discontinuous herbaceous layer of this second-growth (and climax composition) forest range. Poison ivy/oak (a colony forming ecotype or genotype) was locally dominant, but dwarf palmetto, peppervine, rattanvine, greenbriar, and trumpet creeper were also visible in this "photoplot" of the forest floor. Also conspicuous was southern waxy caric sedge, broadleaf woodoats, and giant ragweed (although this light-limited specimen did not live up to the adjective of its common name). Trunk in background was sweetgum.

Jefferson County, Texas. Early October, autumnal aspect. FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash); sweetgum as one of the dominants shared some features of SAF 94 (Sycamore-Sweetgum-American Elm). Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Western Gulf Coastal Plain-Northern Humid Gulf Coastal Prairies Ecoregion, 29c (Griffith et al., 2004).

 
   
 

334. Light made the difference- Man-made clearing in the above second-growth forest was a patch of Gulf Coast prairie dominated by sugarcane plumegrass (Erianthus giganteus) and with two composites, seacoast goldenrod (Solidago sempervirens) and late [flowering] thoroughwort or late boneset (Eupatorium serotinum), as associates. Trumpet creeper, which was content to climb the ground, was the major woody species in this clearing. Along margins of this man-made patch of grassland palmetto was dominant. Tree species in the badkground were primarily sweetgum and sugarberry with less green ash.

Jefferson County, Texas. Early October; early bloom stage of plumegrass, peak-bloom stage of both composites..

 
   
Arid Zone Riparian or Gallary Forest
 

Woody (tree and/or shrub dominated) riparian vegetation along streams in deserts constitutes a unique category or form of range plant community. In context of Landscape Ecology such gallary forests and woodlands comprise corridors in a matrix of arid scrubland. Sometimes such corridor vegetation differs only slightly from adjoining desert scrub in having narrow strips of potential natural vegetation that is slightly less xeric (or more mesic) than adjacent desert plant communities. In such range vegetation the riparian community (ies) amount to little more than bigger, taller "bushes" growing at greater densities. In other instances, however, an actual forest or woodland develops beside the stream. Although such forests or woodlands can be viewed or interpreted as part of the larger desert ecosystem, landscape, or general community this strikingly distinct range vegetation can be regarded as forests in much the same context as montane and subalpine forests on mountain ranges within deserts.

Such narrow, winding, small-spatial scale forests or woodlands would be interpreted as post-climax in the Clementsian monoclimax theory and as climaxes in their own right when interpreted in the polyclimax theory of Tansley or the climax pattern model of Whittaker. Regardless of theoretical ecological perspective these linear or "ribbon-like" forests were deemed herein as singularily unique and worthy of a special category in this chapter of miscellaneous tree-dominated range (= forest) types.

Woody riparain vegetation dominated by shrubs rather than trees was regarded by this author as a form of scrub (=scrubland or shrubland) and not forest or woodland. As such, shrub-dominated riparian range vegetation in arid regions was treated in the chapters devoted to the deserts in which such wetland range vegetation developed (ie. Chihuhuan, Sonoran, Great Basin).

In tree-and/or shrub-dominated riparian range vegetation there are typically herbaceous species which often form layers or strata that are distinctive--often diagnostic--of such wetland plant communities. Some of the characteristic herbaceous plants were included in the following treatment for which McKittrick Creek in McKittrick Canyon (Guadalupe National Park, Culbertson County, Texas) served as an example of a stream in arid regions. For historic purposes it was noted that this creek and canyon were originally spelled McKitterick, that is with an "r" (Tarr, 1892).

 
   
 

335. Rock bottom- Near the mouth of McKittrick Creek at base of Capitan Reef the bed of this stream flows over plates of limestone, at least during wet periods. McKittrick Creek has a intermittant flow pattern with some short and interrupted stream stretches having yearlong flow (more-or-less) aboveground while other stretches frequently have subsurface (ie. stream flow is belowground). Overall, McKittrick Creek has permanent surface water only at sporatic or infrequent intervals along the stream's reach.

The geologic material of McKittrick Creek and Canyon is interpreted as travertine, "a hard, dense, finely crystalline, compact or massive but often concretionary limestone, of white, tan, or cream color, often having a fibrous or concentric structure and splintery fracture, formed by rapid chemical precipitation of calcium carbonate from solution in surface and ground waters, as by agitation of stream water or by evaporation around the mouth or in the conduit of a spring ..." (Gary et al, 1972).

Upslope range plants vary from various species of scrub oaks such as Emory oak (Quercus emoryi), wabyleaf oak (Q. undulata), gray oak (Q. grisea), and sandpaper oak (Q. pungens) to smooth soto (Dasylirion leiophyllum) and lechuguilla (Agave lechuguilla) to Apache plume (Fallugia paradoxa) and mountain mahogany (Cercocarpus montanus) among shrubs to small trees to grasses like Chino grama (Bouteloua ramosa), sideoats grama (B. curtipendula), New Mexico feathergrass (Stipa neomexicana), sundry species of muhly (Muhlenbergia spp.), and various threeawns (Aristida spp.).

In this short presentation of riparian range vegetation the plant communities of upslope woodland and forest types was ignored because that was covered separately as desert scrub in the Chihuhuan Desert chapter and as an outlier or southern extension of Southern and Central Rocky Mountain Forests under that chapter in the Forest and Woodlands section herein.

Major trees along the riparian zone of McKittrick Creek included the dominant little walnut, river walnut, or Texas black walnut (Juglans microcarpa) and such local associate species as velvet ash (Fraxinus velutina), Texas madrone or lady's leg (Arbutus texana), bigtooth maple (Acer grandidentatum), and the composite shrub, seepwillow baccharis (Baccharis salicifolia).

The riparian vegetation along McKittrick Creek has often been descripted as travertine vegetation with travertine the adjective describing the parent geologic material of stream bed as it was defined above.

Guadalupe National Park, Culbertson County, Texas. June.

 
   
 

336. Segment of travertine vegetation- The specific riparian range vegetation along McKittrick Creek has traditionally been described as travertine in reference to the specific form of limestone as defined/described in the preceding photo caption. Also as explained in the previous caption of this short section, upslope nonriparian vegetation of ponderosa pine-dominated forest and scrub oak woodland was treated elsewhere and, hence, ignored within this portion that was devoted strictly to riparian range vegetation.

The foreground section of riparian vegetation along McKittrick Creek presented here was little walnut (represented by two widely spreading scrub trees). The background section of streamside vegetation was dominated by Jamacian sawgrass (Cladium jamaicense) of which more was covered below.

Guadalupe National Park, Culbertson County, Texas. June.

 
   

 

337. Little walnut along a mostly dry desert stream- Little walnut or river walnut was the dominant woody species along much of the streatch of McKittrick Creek. Taller shrubs or small trees in bckground of the first of these two slides was velvet ash which was featured below. Farther upstream a few ponderosa pine (Pinus ponderosa) grew in close proximity to the riparian plant community, but these conifers were not actually in the streamside habitat (ecosystem) and therefore were not part of the riparian vegetation. There were some individuals of seepwillow baccharis in this riparian range community but this species was not a locally dominant nor associate member of the travertine vegetation.

Guadalupe National Park, Culbertson County, Texas. June.

 
   
 

338. Dry creek by an ancient reef- The limestone of Cpitan Reef extended to the bed of McKittrick Creek where little or river walnut, velvet ash, Texas madrone, and an occasional bigtooth maple comprised most of the woody component of a riparian range plant community.

Guadalupe National Park, Culbertson County, Texas. June.

 
   

 

339. Little, river, or Texas black walnut (Juglans microcarpa)- A few representative specimens of the dominant woody plant of the riparian vegetation of McKittrick Creek.

Guadalupe National Park, Culbertson County, Texas. June.

 
   
 

340. Foliage of little, river, or Texas black walnut- Example of shoots and leaves of Juglans microcarpa along McKittrick Creek.

Guadalupe National Park, Culbertson County, Texas. June.

 
   
 

341. Water-dependent walnuts- A relatively large population of little or river walnut dominated the riparian vegetation so as to form a spatially broad community (including velvet ash, bigtooth walnut, and Texas madrone) extending considerable distance from the streamside zone of McKittrick Creek. Farther up McKittrick Canyon and Creek (base of tallest peak in center m;idground) a few ponderosa pine and alligator juniper (Juniperous deppeana) extended down the hillsides to grow in a zone adjacent to the actual riparian range vegetation.

Guadalupe National Park, Culbertson County, Texas. June.

 
   

 

342. Ash edging the creek- Velvet ash was locally a dominant and frequently an associate species in the riparian vegetation along mcKittrick Creek, essentially a desert stream flowing out of the Guadalupe Mountains. A few representative individuals of this species of small tree were introduced here. Tip your leaves to the folks ashes.

In between the two larger ashes in the second photograph a bushy (small, rounded) bigtooth maple represented its species while a saw-toothed smooth sotol stood at base of foremost ash. Featured stars shared spotlight with their shorter sidekicks.

Guadalupe National Park, Culbertson County, Texas. June.

 
   
 

343. A strange (at least, a unique) trio- Jamaca sawgrass, velvet ash, and little walnut "held down the fort" at this short streatch of McKittrick Creek. The travertine stone that comprised most of the bed of this stream was conspicouts. Tree species around the "next bend" (left mid- to background) included more velvet ash, Texas madrone, and bigtooth maple. Sawgrass grew at base and comprised an herbaceous layer in this local riparian plant community.

Guadalupe National Park, Culbertson County, Texas. June.

 
   

 

344. A little bit of the Florida Everglades- Unbelieveable as it seems, the dominant herbaceous species along much of McKittrick Creek was the same plant that dominates the Everglades of south Florida: Jamacian sawgrass (Cladium jamacense). At very edge of this desert stream a major grasslike plant more common to the Gulf Coast had found a faraway home much to its liking.

Guadalupe National Park, Culbertson County, Texas. June.

 
   

 

345. Shoots of sawgrass- Several of the individual shoots of Jamaca sawgrass along McKittrick Creek had unusually large inflorescences with several whorls of fruit-bearing branches. Some range plants occupy unique arrangements of environments. The species range or area of Cladium jamacense is an example of a diffuse type (= form) of discontiuity or discontinuous range (Polunin, 1960, p. 188).

Guadalupe National Park, Culbertson County, Texas. June.

 
   

 

346. Texas madrone along McKittrick Creek- Two picturesque specimens of Arbutus texana adorned this mountain stream within the greater region of the Chihuhuan Desert. In the first of these two slides a little walnut grew at immediate left of the featured Texas madrone. while in the second of these slides a bigtooth maple, Guadalupe sacahuiste (Nolina micrantha), and smooth sotol flanked the Texas madrone (known also as lady's leg and naked Indian).

Guadalupe National Park, Culbertson County, Texas. June.

 
   
 
347. Blooming madrone- Shoots of Texas madrone or naked Indian at full-bloom stage. Williamson County, Texas. March.  
   

 
348. Sexual shoots- Two shoots of Texas madrone at peak bloom. Also examples of leaf characteristics of this ericaceous shrub. Williamson County, Texas. March.  
   

 
349.. Flowers of Texas madrone- Details of flower cluster and individual flowers of Texas madrone or lady's leg. This was a good example of the basic floral features of the Ericaceae as viewed on the range. Williamson County, Texas. March.  
   
   

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