Miscellaneous

Glades or, as they are more commonly known by the local hillbillies, "balds" or "bald knobs" are one form of "barrens". Glades are classified as to their parent material. Missouri has greater diversity in balds than any state and  has limsteone, dolomite, chert, sandstone, shale and igneous glades. The bald seen here in the Ozark Highlands of southwestern Missouri is a chert glade derived from massive brecciated chert of the Elsey Formation (Nelson, 1985, p. 106). An ephemeral (ie. "wet-weather") stream flows through the center draw in early to mid-spring and there are seeps from under the chert outcrops which support the"runt" or scrub form of blackjack and post oaks (Quercus marilandica and Q. stellata) and common "hackberry (Celtis occidentalis). The domiant grass with the "lacy" or "dainty" appearance in the center and left-center foreground is Festuca paradoxica. The dominant yellow composite is tickseed coreopsis (Coreopsis lanceloata). This is the bottom of the bald which supports the vernal, cool-season, most mesophytic species. The surrounding oak-hickory forest was described immediately below. Wildcat Glade Natural Area, Newton County, Missouri. Vernal aspect, May. FRES of the glade vegetation is 39 (Prairie Ecosystem), K-66 (Bluestem Prairie), FRES of the neighboring forest is No. 91 (Oak-Hickory Forest), and FRES of the combined glade and forest vegetation is No. 73 (Mosaic of Numbers 66 and 91). SRM 803 (Missouri Glades). Range site is Limestone Rock Outcrop. Northwestern Great Plains-Semiarid Pierre Shale Plains Ecoregion, 43g ( Bryce et al., undated).

4. Ground surface of a chert barrens (= chert glade)- Except for the five depauperate big bluestem plants, the plant life on the soil surface of this chert glade in the Ozark Mountains are nonvascular cryptogams. Cryptogams are plants that reproduce sexually by spores (ie. they are not seed plants or spermatophytes). Vascular (having vessels which convey fluids, specifically having the vascular tissues of xylem and phloem that function as conduits through roots, stems, and leaves) plants that reproduce by spores such as ferns, horsetails or scouring rushes, club mosses, and spike mosses are known as pteridophytes (Pteridophyta is the Division of the Plant Kingdom made up of pteridophytes). Pteridophytes are often known generically as “the ferns and fern allies”. Tracheophyta is in some plant classification/taxonomic schemes the Division of the Plant Kingdom encompassing both the former Divisions of Pteridophyta and Spermatophyta. In some other taxonomic arrangements the Tracheophyta is elevated to the Kingdom designated Metaphyta, or sometimes, the Plantae. In the taxonomic schemes that use Tracheophyta (the tracheophytes), the Bryophyta (bryophytes are the mosses) and Thallophyta (thallophytes include the algae and fungi) along with the Pteridophyta are the cryptogams with the pteridophytes being vascular cryptogams and the thallphytes and bryophytes being nonvascular cryptogams. The spermatophytes (the gymnosperms and the angiosperms) are designated phanerogams.  Embryophtes are those plants in the taxon designated Embryophyta (plants distinguished by having a zygote that develops into a multicellular embryo while in the female sex organ or in the embryo sac) which includes the Bryophyta and the Tracheophyta.
 
Many of these terms are of the older, classical, or traditional systems that viewed all organisms as either plant or animal. This view was an oversimplification and less precise than the contemporary five kingdom system, but like most of that “generation” of knowledge it was practical and utilitarian for professions and disciplines in the Agricultural Sciences.

The plant community (“plant” by the classical, traditional view) or layer of plant community seen here and in the preceding layer dominated by a species of fern includes spermatophytes, pteridophytes, and thallophytes. Dominant plants of this soil-rock layer are lichens (foliose, fructicose and crustose species); a species of little spike moss is the associate. Observe that the dominant spermatophyte of the prairie, big bluestem, “held its own” even though dwarfed. On this xeric microsite this is the entire plant community: the scrub tree layer is absent as is most of the tallgrass layer. This xeric community (or soil-rock layer) has already progressed phenologically to the autumnal aspect even though it is only August and the rest of the chert glade vegetation is still at the estival aspect. Wildcat Glade Natural Area, Newton County, Missouri. SRM 803 (Missouri Glades). Range site is Limestone Rock Outcrop.

Lichens were defined by Allaby (1998) as: “ a type of composite organism, which consist of a fungus (the mycobiont) and an alga or cyanobacterium (the phycobiont) living in symbiotic [mutualistic] association. A lichen thallus may be crust-like (crustose), scaly or leafy (foliose), or shrubby (fruticose), according to the species.

--Thallus- “a primitive type of vegetative plant body that is not differentiated into stems, leaves, and roots, although analogous structures may be present [used mostly for nonvascular cryptogams]” (Allaby, 1998).  

Lichens are major range forage plants on certain types of range in the Arctic Region where they are the mainstay for the caribou/reindeer. On this chert glade they are interesting plants contributing more to soil formationH than to animal diets. Lichens visible include Acarospora chlorophana, Cladonia caroliniana, and Xanthoparmelia spp.). Little spike moss is growing (was growing and is now desicated and dormant) in the cracks among the lichens. Autumnal aspect, August (on this xeric microsite plant phenology has passed to dormancy for the little spike moss and near-dormancy for the lichens). Wildcat Glade Natural Area, Newton County, Missouri. SRM 803 (Missouri Glades). Range site is Limestone Rock Outcrop (this is a microsite thereof).

5. Composite shot of the soil-rock community (or, perhaps more specifically, the nonvascular cryptogam vegetation layer) of a chert barrens in the Ozark Plateau- This is a textbook example of a cryptogamic soil surface. The dominant plants are lichens, one of those miraculous kinds of mutualism which like ruminants and their rumen microbes, legumes and their nitrogen-fixing bacteria, and pollinating plants and their pollinators are essential for functioning of range ecosystems and the profession of Range Management. 
 

Lichens were defined by Allaby (1998) as: “ a type of composite organism, which consist of a fungus (the mycobiont) and an alga or cyanobacterium (the phycobiont) living in symbiotic [mutualistic] association. A lichen thallus may be crust-like (crustose), scaly or leafy (foliose), or shrubby (fruticose), according to the species.

--Thallus- “a primitive type of vegetative plant body that is not differentiated into stems, leaves, and roots, although analogous structures may be present [used mostly for nonvascular cryptogams]” (Allaby, 1998).  

Lichens are major range forage plants on certain types of range in the Arctic Region where they are the mainstay for the caribou/reindeer. On this chert glade they are interesting plants contributing more to soil formationH than to animal diets. Lichens visible include Acarospora chlorophana, Cladonia caroliniana, and Xanthoparmelia spp.). Little spike moss is growing (was growing and is now desicated and dormant) in the cracks among the lichens. Autumnal aspect, August (on this xeric microsite plant phenology has passed to dormancy for the little spike moss and near-dormancy for the lichens). Wildcat Glade Natural Area, Newton County, Missouri. SRM 803 (Missouri Glades). Range site is Limestone Rock Outcrop (this is a microsite thereof).

8. Ozark Bald Knob- This is a dolomite glade or dolomite barrens in the Ozark Plateau. Such distinctive habitats and their vegetation are known to the local hillsmen as "balds" or "bald knobs". In virgin condition dolomite balds were almost entirely grasslands devoid of woody growth except for scattered individuals of yellow-wood or smoke tree (Cotinus obovatus), the most common shrub, smooth sumac (Rhus glabra), skunkbush sumac, sassafras (Sassafras albidum) red bud, and persimmon (Diospyros virginiana), all of which are in this scene. Dominant plants are the prairie grasses Indiangrass, prairie dropseed, and big bluestem with sideoats grama, poverty dropseed (Sporobolus  neglectus), and Junegrass (Koleria cristata) as associates. The dominant forb which is co-dominant with the three dominant grasses is prairie dock (Silphium terebinthinaceum). Forb associates include calamint (Satureja arkansana), longleaf bluets (Houstonia longifolia), and spike lobelia (Lobelia spicata). Except for the dominant prairie dock, composites are rare and locally limited to yellow coneflower (Echinacea paradoxa), gayfeather (Liatris spp.), and compassplant (Silphium laciniatum). Lichens were common only on xeric microsites and much less developed than on the chert glade seen immediately above. The tall blue-green grass in the right foreground is Indiangrass, the dominant species of the bald. The tall yellow-flowered composite in center foreground is prairie dock, the dominant forb. 

The Ozarks are ancient mountains worn down to their roots. The dolomite geologic strata are of the Cambrian-Ordovician formations, some of the oldest in these "everlasting hills". Note the concentric ring arrangement of the Cambrian-Ordovician formations.

Note also that this is bald is relatively free of invading trees, which unfortunately surround this oasis of pristine vegetation. The dominant invaders are eastern red cedar (Juniperus virginiana) and Ashe cedar or Ashe juniper (J. ashei). This glade has been kept open by prescribed burning which handily controls the non-sprouting noxious cedars yet allows maintenance of endemic shrubs like yellow-wood. Several ecological reviews of Ozark vegetation prior to European settlement have been made based on surveyers records, travelers journals, Army accounts, etc. (Steyermark, 1940; Beilmann and Brenner, 1951; Howell and Kucera, 1956; Steyermark, 1959). All these ecological surveys proved conclusively that throughout the Ozark Mountain Region oak-hickory-pine forests had expanded and invaded the pre-white man prairies and savannas. Cessation of fire along with farming, overgrazing, and commercial activity weakened the prairie sward and allowed the woody invasion that reached proportions of massive afforestation. The Soil Survey of Taney County Missouri, the adjoining county to the west of this glade, cited a mean fire frequency of 3.2 years on some Ozark glades and credited this as "“the main reason for this open grassy upland landscape" (Natural Resources Conservation Service, 1996, p. 41).

A more colorful testimony of the original openness of these balds is the gripping tale of the "Bald Knobbers", post-Civil War "night riders" (vigillantes) who held clandestined meetings and lite signal fires atop bald knobs so as to be able to spot any unwelcome visitors and send coded messages to citizen peace officers (at least before vigilante justice degenerated into blood fueds). Today, most of the Bald Knobbers grassland meeting places are forests or cedar thickets.

 McClurg Glade, Ava Ranger District, Mark Twain National Forest, Ozark County, Missouri. Estival aspect, July. No specific FRES or Kuchler designations for vegetation at this restricted scale, but it is primarily an "island"of FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie). However, at landscape scale this is K-74 (Cedar Glades, Quercus-Juniperus-Sporobolus) which is contradictory because by definition glades are open areas of herbaceous vegetation so there is no way that oak and juniper could be the dominant genera. SRM 803 (Missouri Glades). Ozark Highlands- White River Hills Ecoregion, 39c (Chapman et al., 2002).

9. Bald knob range- Two photo-plots of a dolomite glade in the Missouri Ozarks maintained (more-or-less) by prescribed fire consistent with the natural fire regime. In the first photograph eastern red cedar and Ashe or post cedar were reinvading the bald knob. This is the Kuchler (1964; in map of Garrison, 1977) physiogonomic type designated Cedar Glades which recognized Juniperus virginiana and J. ashei as the dominant woody species of this potential natural vegetation. In absence of fire (and probably aided by improper grazing, especially overgrazing) these two junipers increase to such cover and density to result in a post-climax juniper woodland instead of the climax juniper-tallgrass savanna.

In the second photograph junipers were largely limited to the rock outcrop perimeter of the dolomite glade (where there is insufficient fuel to carry fire that kills these non-sprouting conifers). Instead the invading woody species was smoke tree (smoke bush) or yellow-wood. Also on the perimeter were individuals of smooth sumac, skunkbush sumac, sassafras, and redbud. These shrubs are well-adapted to fire. In fact, smooth sumac typically increases with range burning. Yellow-wood matures quickly and becomes senescent, but fire serves to "rejuvenate" older individuals of smoke tree through resprouting as was case for the young stump sprout in left center foreground of this second photograph.

Major (dominant) grasses were Indiangrass, prairie dropseed, and big bluestem. Numerous individuals of Indiangrass were readily distinguishable in these photographs as photographs as the tallest, bluish-green clumps of grass. Interestingly and for unknown reasons, many of the grass clumps of light-green (a "washed-out" green) color were also Indiangrass. In fact, several of these Indiangrass plants (prominent in center midground of first slide) had shoots of both the regular chalky blue coloration and the uncommon pale green color. Indiangrass is a long-shoot tallgrass that elongates the culm early in the warm growing season whereas big bluestem is a short-shoot tallgrass species that does not elongate the culm until much latter in the growing season (often two months or more after Indiangrass). Large specimens of Indiangrass were visible in front of the yellow-wood sprout in the second slide. Dead shoots topped with prominent panicles were specimens of the cool-season (hence, now dormant) Junegrass.

A nice specimen of prairie dock, the common dominant forb on bald knob glades, was in front of the young cedar in center foreground of the first slide. White-tailed deer (Odcoileus virginianus) bucks had been burnishing their antlers on this sapling juniper resulting in severe damage to the conifer and range imporvement (aid in restoring the range plant community to potential natural vegetation) on this dolomite glade.

FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie) and K-74 (Cedar Glades). Prescribed fire has been used here at intervals of three to eight years. SRM 803 (Missouri Glades). Ozark Highlands- White River Hills Ecoregion, 39c (Chapman et al., 2002).

10. Grass and dock on the glade- Details of species composition of the herbaceous layers of this Ozark glade range. This example of bald knob vegetation was a consociation of prairie dropseed. It was a example of Clements' true prairie (in contrast to tallgrass prairie dominated by bluestems and Indiangrass). Tallgrass species of Indiangrass, big bluestem, and little bluestem (in that order) and prairie dock were associates to prairie dropseed. Other grasses included sideoats grama, poverty dropseed, and Junegrass. Other forbs were gayfeather willow-leaf sunflower (Helianthus salicifolius), compassplant, calamint and longleaf bluets.

The sward of this bald knob range had been maintained by three prescribed fires in recent records: 6, 12, and 22 years before time of this photograph.

FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie) and K-74 (Cedar Glades). Prescribed fire has been used here at intervals of three to eight years. SRM 803 (Missouri Glades). Ozark Highlands- White River Hills Ecoregion, 39c (Chapman et al., 2002).

13. Invasion of dolomite glades or bald knobs by eastern red cedar, primarily, and with some Ashe cedar. At landscape scale this vegetation was mapped by Kuchler (1966) as K-74 (Cedar Glades, Quercus-Juniperus-Sporobolus) which as shown above was contradictory because grassland is climax and oaks and junipers are at best only aspect dominants. True dominants are Indiangrass, prairie dropseed, and big bluestem. Detailed studies clearly proved that woody invasion occurred and caused (or is the result of) range retrogression due mostly as a result of fire suppression. Fortunately that dim-wit bruin Smokey Bear died (cremation would have been more appropriate than his expensive burial). Now the Forest Service is using prescribed fires to clean up the bald knob country and restore the climax prairie vegetation. It was none-too-soon as seen on these cedar-invested glades over which a fire tower stands sentinel. How ironic, prophetic, and symbolic of good intentions that wrought bad! Fortunately man is a thinking animal.

 Three Sisters viewed from Gladetop Trail, Caney Campground, Ava Ranger District, Mark Twain National Forest, Ozark County, Missouri. Estival aspect, July. FRES No. 39 (Prairie Ecosystem), K-74 (Cedar Glades). SRM 803 (Missouri Glades). Ozark Highlands- White River Hills Ecoregion, 39c (Chapman et al., 2002).

14. Cedar encroachment on Ozark Glades- Eastern red cedar and Ashe or post cedar with "tag-along" individuals of yellow-wood or smoke tree, sassafras, persimmon, smooth sumac (which increases rapidly and profusly with fire), and chittam-wood or gum-elastic (Bumelia languinosa var. albicans= B. languinosa var. oblongifolia).

This invading woody component was on the edge of the glade presented in the immediately preceding seven photographs where this glade joined unburnt portions. The woody vegetation at this edge served as a control plot from which response to prescribed fire of range vegetation on the prescription-burnt glade could be compared. Patchworks or mosaics of such burned and unburned range vegetation on Ozark Highlands handily fit the concept in Landscape Ecology and Wildlife Management of edge effect. Small tracts of cedar-dominated vegetation were patches within a matrix of glades just as at larger spatial scale glades were patches in an overall matrix of oak-hickory forest (patches inside patches).

Edges between adjoining dolomite glades and cedar-infested glades were small ecotnes that benefitted wildlife species. However, the ultimate state of this Ozark Plateau range type in absence of natural fire or prescribed fire that simulated natural fire regimes is all too often a cedar (juniper) woodland or, ultimately, a cedar forest of closed canopy and no understorey, often with accelerated erosion of soil surface.

The sermon for this range type is a reversal of words, but the same moral as "repent or burn". On range such as this the prophetic message is "repent and burn or regret and perish".

FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie) and K-74 (Cedar Glades). Prescribed fire has been used here at intervals of three to eight years. SRM 803 (Missouri Glades). Ozark Highlands- White River Hills Ecoregion, 39c (Chapman et al., 2002).

16. Complex of Stony Land- Rock outcrops are some of the most widely distributed forms of barrens. These occur on small areas or in scattered restricted locations within a landscape-scale or regional-sized general plant community (eg. rock outcrops on hill sides or mountain tops within a grassland association or forest formation (eg. granite outcrops within the Sierra Nevada Mixed Conifer Forest  and in the Appalachian Highland or Ozark Plateau of the Eastern Deciduous Forest Formation).

 Closely related to rock outcrops are talus or scree slopes, cliffs and bluffs, the various kinds of rock glades, sand and gravel bars along streams, sand dunes, and beach and dune habitats. These sometimes occur together and, as the case with the continuum of savanna vegetation, intergrade to the point that it becomes arbitrary to distinguish an outcrop from a rock glade from a talus slope. These all share the common feature of having immature, shallow soils, parent material on or close to the ground surface, and rapid rates of water runoff such that the habitats are droughty, generally harsh, and overall hostile to plant life. Nelson (1987, p. 99) designated these as primary natural communities which is certainly consistent with primary succession, primary materials, primary structure, or even primary schools. 

The complex of primary areas and range sites seen here is in the Wichita Mountains of Oklahoma. Coming down from the top of this weathered mountain are the following designated categories of land: 1) Granite Outcrop Escarpment, 2) Granite Rock Outcrop, and 3) Stony Rock Land which in turn is divided into 4) Hilly Stony range site and 5) Hilly Stony Savanna range site. These were given in various geologic references and range site descriptions in Soil Conservation Service Soil Surveys (eg. Commanche County, Oklahoma). The vegetation is extremely variable and different distinct plant communities could be recognized at different spatial scales. Species include eastern red cedar, netleaf hackberry (Celtis reticulata), scrub form of blackjack and post oaks, slippery or red elm (Ulmus rubra), smooth and skunkbush sumac, big bluestem, Indiangrass, little bluestem, sideoats grama. Soils are mostly of the Lithosol great group (the 1938 soil classification system). It is all Land Capability Unit VIII.    

Wichita Mountains National Wildlife Refuge, Commanche County, Oklahoma. Autumnal aspect, October. This is an “island” of rock outcrop or barrens within the “sea” of the mixed prairie range type (FRES No. 38 [Plains Grassland], K-62 [Bluestem-Grama Prairie]) There is no appropriate FRES or Kuchler Unit designation for this rock outcrop. As much as anything else it resembles a scrub form of Cross Timbers and legitimately could be interpreted as such (FRES No. 15 [Oak-Hickory], K- 75 [Cross Timbers]), but the author felt that it was unique from that (and all other vegetation) and should be categorized as a savannah or barrens. Central Great Plains- Wichita Mountains Eoregion, 27k (Woods et al., 2005).

17. Mountaintop savannah-  Here on the top of Mount Scott (elevation 2464 feet) in the Wichita Mountains is a big bluestem-eastern red cedar savanna. Indiangrass and little bluestem are tallgrass associates and sideoats grama is the midgrass associate. The dominant forb is broom snakeweed (Gutierrezia sarothrae). Eastern red cedar is wind-pruned thereby having a semi-krumholtz form and forming a tallgrass-juniper savanna or perhaps a mixed prairie-juniper savanna. There is some Ashe juniper (Juniperus asheii) in the Wichitas, but eastern red cedar (J. virginiana) is overwhelmingly the dominant cedar. On dry, rocky hill tops like this the juniper would be less apt to burn and would logically be a component of the climax community such that the potential natural vegetation has to be interpreted as a savanna. As discussed under the previous slide no FRES or Kuchler Units exist at this restricted scale. Likewise there is no SRM rangeland cover type although some of the Texas Edwards Plateau types have a similar floristics and habitat (eg. SRM 733, Juniper-Oak Woodland). Consistent with the granite parent material of this "boulder field" and the inclusion of glades as a form of barrens and primary natural plant community it seemed that the most likely published classification of this unique range vegetation was Cedar Glades (Kuchler Unit 74) or, alternatively, some designation based on parent material (eg. Granite Glade or Granite Rock Outcrop) or perhaps a combination of geologic material and dominant species (eg. Granite Cedar Glades, Granitic Bluestem-Cedar Savanna). The range site designation is most fitting: Hilly Stony Savanna.

Wichita Mountains National Wildlife Refuge, Commanche County, Oklahoma. Autumnal aspect, October.Central Great Plains- Wichita Mountains Eoregion, 27k (Woods et al., 2005).

19. Serpentine Barrens Rock outcrop- An extremely harsh edaphic environment created by the presence of the mineral serpentine is one of the more common forms of barrens along the Pacific Coast. When serpentine barrens is combined with rock outcrop, one of the most common forms of barrens continent-wide, the habitat is definitely BAD.

Serpentine forms a unique, if harsh, habitat and, following an initial period of mostly being ignored by ecologists, serpentine sites became a “pet” subject of study in the Pacific Region. Readers are first referred to the relatively recent reviews by Kruckeberg (1984) and Kruckeberg in Anderson et al. (1999, ps. 309-321) as well as Tyndall and Hull in Anderson et al. (1999. Ps. 67-82). The older work, some of it classic, was given in the references of these recent reviews.

Serpentine is a mineral conisisting basically of a variable hydrous magnesium silicate, a complex magnesium-rich silicate- chiefly as Mg₃Si₂O₅(OH)₄- in fine waxy masses (but locally fibrous as asbestos), having colors ranging from dull or light green to dark green to black with intermediate brownish-red or yellow and often with a mottled appearance. Serpentine formed by alteration of peridotite (olivine) in presence of water and thus can occur in or as rock, both altered peridotite and pyroxenites. Serpentine is one of the ultramafic rocks (igneous rocks composed mostly of mafic minerals which are dark-colored iron-magnesian rock-forming silicates with olivine, chromite, and pyroxene as major minerals).        

Serpentine parent material weathers into soils rich in magnesium, iron, nickel, and  silicates. Serpentine soils have montmorillonite as the common clay mineral. Serpentine soils include both alluvial and upland series. The upland serpentine soils are azonal soils that are stony, shallow, and lack regular profile development. In addition, serpentine soils have high levels of exchangeable magnesium and exrtremely low levels of exchangeable calicum. They are typically deficient in nitrogen and phosphorus among macrominerals and molybdenum among trace elements while containing hyper-levels of the heavy metals iron and nickel. This creates a “serpentine syndrome” of infertile soils combined with toxicity of magnesium and heavy metals (Kruckeberg, 1984, ps. xii, 6-8, 18-26). In short, not the place for a plant to plan on having a party. 

Yet, certain plants survive, if not thrive, here. Some of them are serpentine endemics. Others have distribution patterns varying between serpentine and non-serpentine soils while yet others are categorized as indifferent (= bodenvag= ubiquist) species and occur on and off serpentine soils with no apparent taxonomic differences within species on both categories of soil (but ecotypic variation cannot be ruled out definitely). See Kruckeberg (1984, ps. 34-44) for discussion of plant species relations on serpentine soils.

Response of range animals to serpentine sites and vegetation thereon has received much less study than has the more basic plant respones.   

The Serpentine Barrens Rock Outcrop Site seen here is dominated by big squirreltail (Sitanion jubatum) which is one of the indifferent or bodenvag serpentine species. This is a very restricted site (microsite) in the northern Coast Range. The surrounding oak woodland on either side and the California annual grassland behind are not on serpentine soils. Foster Pasture, University of California Hopland Field Station, Mendocino County, California. Late May, nearing end of growing season. 

Serpentine Barrens were too small to be mapped at the scale used by Kuchler (1966, 1977), but this local site is obviously an “island” of the California bunchgrass prairie (K-41, California Steppe). As discussed earlier, SRM failed to designate and describe California steppe or Pacific bunchgrass prairie.

20. Serpentine Barrens range site of California annual grassland- Kruckeberg (1984, p. 41) listed a number of Eurasian annual grasses which he felt must be indifferent (= bodenvaq or ubiquist) species on serpentine and non-serpentine sites. He listed wild oats, soft brome, red brome, nitgrass (Gastridium ventricosum), and foxtail fescue (Festuca megalura). All of these species except wild oats are common on the parcel of naturalized annual grassland in this photograph. There is a relatively higher proportion of lower-seral stage species in this community with soft chess (= soft brome) being the only species that sometimes responds as a naturalized decreaser. Almost all of the grasses seen here are naturalized invaders like red brome, nitgrass, and foxtail fescue. Rosiere et al. (1986) studied the range plant community and domestic sheep diets on this same pasture and compared them to those from non-serpentine range sites on this same ranch. They found that there were few real differences in diets and the main difference in range species compostion was a lower proportion of decreaser and increaser grasses on the Serpentine Barrens.

 Foster Pasture, University of California Hopland Field Station, Mendocino County, California. Late May. FRES No. 42 (Annual Grasslands Ecosystem), Kuchler (1966, 1977) did not map naturalized vegetation. Serpentine sites on California annual grassland were included under SRM 215 (Valley Grassland).

21. Serpentine Barrens soil- The soil series of the Serpentine Barrens range site just seen is Montara, a Vertisol. Cracking of this azonal soil is due to it’s high shrink-swell potential resulting from the high content of montmorillonitic clay (mentioned above) and the shallowness of this upland soil. The physical features of cracking and shallowness together with chemical infertility creates a harsh edaphic habitat for range plants. What is so interesting, however, is that the annual grass species are the same on the Serpentine Barrens as on the non-serpentine site except there are higher proportions of the less palatable species, especially naturalized invaders, on the Barrens. This is because on this range sheep are attracted to and spend more time grazing and, generally, congregating on the Serpentine Barrens sites. The actual reason why sheep preferred grazing and loafing/resting on the barrens was not determined when Rosiere et al. (1986) compared sheep diets from the barrens to diets from non-serpentine sites. Note that even under the relatively heavy degree of use on this site soft chess still set seed for next year.

Foster Pasture, University of California Hopland Field Station, Mendocino County, California. Late May. FRES No. 42 (Annual Grasslands Ecosystem), no Kuchler designation. SRM 215 (Valley Grassland).

22. Comparison of grazed and non-grazed vegetation on a Serpentine Barrens range site-  On the left is the grazed Serpentine Barrens range seen in the previous slides. On the right is a 25 year-old exclosure. Naturalized Mediterranean annual grasses dominate grassland communities on both sides of the fence, but there are more individual plants of the native perennial bunchgrass species in the 25-year exclosure (none could be found on the grazed portion). California melic (Melica californica) and purple needlegrass can be seen in the exclosure.  Rosiere et al. (1986) concluded that the range vegetation on Serpentine Barrens was not strictly determined by edaphic conditions and features but that it was a combination edaphic-zootic climax. All California annual grassland is disclimax but on serpentine sites the role of grazing appeared even greater. In fact, it seemed (based strictly on personal observation  and judgment) that the native bunchgrasses were more common on serpentine than on non-serpentine soils. This is not to say that the native climax grasses would recover and recreate the pre-white man Pacific prairie on serpentine soils if protected from grazing. It does suggest that the native perennial grasses are comparatively better adapted to and more competitive with Mediterranean annual grasses on serpentine sites. Natives like needlegrass, melicgrass, and squirreltail are frequently much more abundant in exclosures on serpentine soils than in exclosures on non-serpentine soils. This is an example of how interactions of management factors (eg. stocking rate, animal distribution) and plant growing conditions influence range vegetation. 
 

Note how perennials extend the green feed season. The annual species have been brown for three weeks and the native bunchgrasses are “still green”. On Hopland Field Station the author observed purple needlegrass that retained green shoots through June even on south slopes and through most of July on north and east slopes.

Foster Pasture, University of California Hopland Field Station, Mendocino County, California. FRES No. 42 (Annual Grasslands Ecosystem), SRM 215 (Valley Grassland).

25. Canebrake above the reaches of the Bosque- The once vast North American canebrakes of giantcane and switchcane apparently extended some distance above their major habitat of river bottoms. The famous Cane Ridge that figured so prominently in development of backwoods camp meetings and onset of the Second Great Awakening gives some suggestion of this. The above two photographs were of relicts of a former canebrake range community that extended upland for considerable distance from the floodplain of the Brazos River in Texas' Western Cross Timbers. In the concept of Landscape Ecology such former extensive canebrakes along creeks and rivers in the Cross Timbers constituted ribbon-shaped patches imbedded within the matrix of the surrounding Cross Timbers. These waterways and their canebrakes also functioned as corridors connecting grasslands of such prairies as Grand Prairie and woodlands/forests of the Cross Timbers.

Large trees in the background of both slides were post oaks. Smaller trees of hackberry or sugarberry (Celtis laevigata) were in background of the first of these two slides. The tree in left foreground of second slide was pecan. For several years prior to taking of these photogrphs this canebrake had been heavily grazed by horses with high-lining" of branches within reach while most main shoots were not consumed. This demonstrated that canebrake range has some resilency.

April. Vernal aspect; shoot emergence/elongation phenological stage. No FRES, Kuchler, or SRM designations. Cross Timbers- Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

26. Rapid growth in a favorable spring- Edge of Arundinaria gigantea canebrake in mid-spring during development of this year's shoots. Some bamboo shoots had already reached their mature height (first photograph) while limbs were still coming into their final position as short, ephemeral, husklike sheaths were shed to that limbs were released from their appressed position on the culms. This positioning of limbs along sides of culms was shown at outer margin of the canebrake in the second photograph.

Erath County, Texas. May: various phenological stages of shoot emergence and elongation

27. Shoot emergence of Arundinaria gigantea- Floor of a canebrake at onset of annual grass growth cycle. Shoot growth (emergence and elongation of main culm and branches) of giant cane typically occurs only once per year. This is at onset of spring green-up when previous year’s leaves are shed and current year’s leaves emerge essentially simultaneously. Young shoots at this stage were used as a pot herb by Indians and backwoodsmen. The shoots had to be boiled “for an eternity” (they ain’t exactly asparagus).

Erath County, Texas. April: early stage of shoot emergence.

29. At 13 days post-emergence the bamboo shoots of the last two slides have reached their adult (maximum) height of six to 10 feet or more. No more aboveground growth will occur this growing season even though there will be over five months of frost-free temperatures and soil moisture adequate for growth (plus leaves remain green until shed the next spring). Instead all growth occurs underground as rhizomes development and extend for distances of up to 15 feet. Tillers will emerge from these rhizomes the next spring to begin the annual growth cycle again.

"I was raised in a canebrake by an ole mamma lion; cain’ t no high-toned woman make me walk the line"

(Sixteen Ton by Merle Travis and made famous by Tennessee Ernie Ford).         

Prevalence and importance of canebrakes in the Cumberland country as livestock and wildlife feed and habitat and as an impediment to travel was discussed by Arnow, 1960, ps. 20, 38, 51, 56, 98, 127, 168, 237, 247, 401, 408; Arnow, 1963, ps. 214-215, 239, 241). Sitton (1995, ps. 6, 7,25, 34, 49, 66, 145, 148, 200, 224, 226, 231) documented canebrakes as forage sources in the open range ranching region of the Big Thicket (deep east Texas) which as discussed under the Forest and Woodland Slides (oak-hickory range type) was the last actual open range in the United States. Blevins (2002, ps. 18, 27, 35) explained that in the Arkansas Ozarks switch canebrakes either became some of this states richest farmland or were used as open range by cattlemen during roughly the last two-thirds of the Nineteenth Century. Beilmannn and Brenner (1951, p. 276) reported from a historical review of travlers' accounts, explorers' diaries,  and especially surveyors' notes that the "impenetrable" canebrakes were a "popular retreat of game of all sorts", especially "one of the favored haunts of the black bear". One of the most interesting (and least remembered) "habitat" uses of canebrakes was as a hideaway for runaway slaves, renegade Indians, missionaries, and southern "white trash". Accounts by Peter Cartwright, one of the most influencial and colorful Methodist circuit riders in America, described villages made up of mixed-blood misfits and social outcasts hidden in the immense canebrake that streatched for miles along the floodplain of the Ohio River (Greenbie and Greenbie, 1962, ps. 14-18, 468-469, 478-482).

30. Advance of a canebrake- Invasion of "new land" by Arundinaria gigantea. Shoots (tillers) from remarkably long rhizomes produced over course of the previous growing season formed the vanguard as a canebrake advanced into ground not previously colonized (at least not in recent years) by this native species of bamboo. Newly emerged shoots were in various stages of development. Most shoots had just shed the "husks" from their culms thereby permitting the until-then appressed limbs of the shoots to be released and attain their mature positions along the culms. These shoots ranged in height from less than one foot to over six feet in height.

Erath County, Texas. May: various stages of shoot emergence and early development..

31. Shortleaf pine-oak forest with a canebrake understorey, a pine-oak-giant cane savanna- This was a most unique composite vegetation. It was an oak-pine forest with tree and shrub layers that were diverse in their own rights to which was "added" a canebrake of giant or southern cane, the native bamboo (Arundinaria gigantea subsp. gigantea). There was no layer of vegetation (at least of macroscopic vascular plants) beneath the canebrake which is the typical condition for canebrakes of this species. Transeau et al. (1940, p. 762) specified that canebrakes were "often invaded" by evergreen oaks. As described in the next paragraph evergreen species in this community were pines and the oak species were not "live oaks". This community was similar, however, to that described in a standard Botany text widely used decades ago (ie. old textbook knowledge). Prior to settlement by European man, giant cane formed vast "brakes" or colonies (as in endless miles of bamboo) along water courses ranging in size from major rivers to creeks throughout southeastern North America. These seemingly infinite canebrakes occurred as far north as the Ohio River and westward to central Texas. Giant cane also grew (and still grows) as isolated plants of one or a few shoots up to small colonies of several hundred square feet as part of the understorey vegetation in oak-hickory and pine-hardwood forests. An exclusive (a homogenous or "pure") stand of uninterrupted canebrake understorey beneath an adult tree canopy was-- in this vegetation hunter's experience-- quite rare. The species composition, structure, and physiogonomy of this forest-woody grass vegetation was likely that of the climax. This was relict vegetation. Even though it was not virgin forest this was interpreted as a climax community tht lacked the component of very old trees of the species present. It was one of the most unique assemblages of native plants encountered by your photographer-author.

Shortleaf pine was the dominant tree, but there were a few individuals of loblolly pine (or perhaps of the widespread natural hybrid of these two species). Other trees included white oak, southern red oak, water oak, and sweet gum in about that order. There was a shrub layer composed of small individuals of persimmon and sassafras as well as smooth sumac. These shrubs and/or short trees grew in the canebrake, but were shorter than the taller shoots of the bamboo except at the edge of the canebrake where there was light of greater intensity and frequency. Pensacola bahiagrass (Paspalum notatum var. saurae) and hurrahgrass (P. pubescens= P. muhlenbergii= P. setaceum var. muhlenbergii) also grew on these extreme edges where they got adequate light for survival.

The development of spreading, horizonal branches on all trees, including the dominant pines that are usually strongly apically dominant, indicated that these trees had developed on a savanna which was exactly what this community appeared to be: an oak-pine-giant cane savanna. The soil was moist enough to support water oak and the scarce loblolly pine (or a hybrid thereof), but soil water conditions were not periodically ponded let alone swampish.

The grassland expression of Arundinaria gigantea canebrake (exclusive, homogenous stands or colonies of giant cane with widely scattered or infrequent trees) was covered in the Miscellaneous portion of the Grasslands section.

Miller County, Arkansas. July. West Gulf Coastal Plain portion of the Coastal Plain physiographic province. Pineywoods vegetation, part of the Southern Pine Region. According to U.S. Forest Service Ecological Divisions (Forest Service, Final Environmental Impact Statement, 1999) this was in the Southeastern Mixed Forest Province (231), Mid Coastal Plains, Westen Section (231E), South Central Arkansas Subsection (231Ea). FRES No. 14 (Oak-Pine Forest Ecosystem). K-102 (Southern Mixed Forest). SAF 76 (Shortleaf Pine-Oak, variant of). No Kuchler or SRM units for canebrake. South Central Plains- Tertiary Uplands Ecoregion, 35a (Woods et al., 2004).

32. Shortleaf pine-mixed oak-giant cane savanna or a shortleaf pine-mixed oak forest with a giant cane understorey- A more interior view of the composite pineywoods vegetation displayed in the preceding slide. Interpretation of this vegetation was subject to many subjective factors including disciplinary background or professional allegiance. It could be viewed as a mixed oak-southern pine forest with an understorey comprised exclusively of a "brake" of giant cane, the one species of native North American bamboo. Alternatively this could be viewed as a combination of diverse pineywoods forest and giant cane or bamboo grassland. Beyond any debate these two "elements" (components) were present. Also beyond debate is the ecological fact that both of these vegetational elements do occur (they formerly occurred on massive scales in the virgin vegetation) in their own forms separate and distinct from each other as potential natural vegetation. It was a different form of canebrake; it was a unique developmental expression of oak-pine forest growing as it were "out of" or in the midst of a canebrake.

The characteristic leaves of southern red oak appeared very distinctive along both the left and right borders of this photograph. Smooth sumac and shrublike or scrubby trees of persimmon and sassafras were overwhelmed in a "sea" of native bamboo.

Miller County, Arkansas. July. West Gulf Coastal Plain unit of Coastal Plain physiographic province. Gorest Service ecological divisions given in preceding caption. FRES No. 14 (Oak Pine Forest Ecosystem). K-102 (Southern Mixed Forest). SAF 76 (Shortleaf Pine-Oak, variant of). No Kuchler or SRM unit for canebrake. South Central Plains- Tertiary Uplands Ecoregion, 35a (Woods et al., 2004).

Some of the greatest diversity-- from standpoint of both species and plant community-- in range vegetation occurs in wetlands. Wetlands are also extremely diverse including such important range plant communities as those of riparain, marshes, wet prairies, swamps, estuaries, wet mountain meadows, and playa lakes. Classification and delineation of wetlands has traditionally varied almost as much as wetlands themselves depending on criteria used and objective (eg. legal, historical, ecological, etc.). Even from these more specific perspectives or discipline emphases the bases for classification or recognition frequently are different. For instance ecological distinction and description has been varied depending on criteria such as biotic community, vegetation, body of water features, soil properties, or duration of water coverage on land surface. In this regard the outstanding encyclopedia, Wetlands, by Mitsch and Gosselink (2000) is definitive (for eg. see Chapter 21 for wetland classification).

One of the wetlands featured variously in the current publication was that of streams (and associated landscapes) because the vegetation of such included several range (rangeland and forest) cover types. Vegetation of the the Rio Grande provided examples of willow scrub and mesquite bosque that were treated under Miscellaneous Shrublands. A third major range plant community of the Rio Grande was patches of grassland dominated by or exclusive communities of common reed or, in Spanish, carrizo (Phragamites communis= P. australis). Such "reedbanks" or "reedbeds"(the Spanish of which is "carrizal"), as they are commonly known, were covered here because they are a minor (very small in area) yet unique and very important grassland that can be interpreted as a range cover type. Reedbanks are a kind of grassland wetland comprised primarily of a native North American grass species that forms or is part of natural range vegetation. This range vegetation was viewed herein as climax.

This was consistent with its classification or designation as the Phragmites communis-Carex spp, et al. Series which together with the Distichlis stricta-Allenrolpea sp Series made up the Chihuhuan Interior Marshland (Brown, 1994, Fig 104, p. 177). Unfortunately, Brown et al. (1998, p. 46) erroneously omitted the Phragmites-Carex Series (a biotic community) from Chihuhuan Interior Marshland, 243.6. The Saltgrass Series, 243.61, included in Brown (1994, Fig , p. 177), was given in Brown et al. (1998, p. 46); therefore, the reedbed vegetation should be Common Reed or Carrizo (Brown , 1994, p. 336)-Sedge Series 243.62.

Common reed was long recognized as forming or developing into the reed-swamp association (Warming, 1909, p. 145; Tansley, 1923, ps. 31; Clements, 1936, ps. 264, 278; Polunin, 1960, p. 463). In fact, as was detailed in the Literature Review (Associations and Consociations section of Range Type), Phragmites communis was one of the most commonly used examples of association (or associes, the seral counterpart) in the Anglo-American School of Plant Ecology.Clements (1920, p. 110) reed-swamp was "[t]he most familar example" of an associes that was usually composed of three consocies one of which was P. communis. Consocies was Clements' term for the seral counterpart of consociation.Common reed (as the "reed-swamp") was one of the "kinds" of vegetation used in derivation of "type", including cover or dominance types (thus, rangeland and forest cover types) from the plant association (again, see Literature Review).

Inclusion of common reed (forming reedbeds or reedbanks) as the common reed cover type provided photographs of vegetation that was of historic significance in development of the cover type concept. Reedbanks were also instructive of a very restricted but a highly productive kind of rangeland and critical habitat along an important stream in the Trans-Pecos Basin and Range. The reedbank or reedbed provided an example of range vegetation along a stream inside a general desert and desert climate. Like willow scrub and mesquite bosque, reedbank was not desert vegetation and therefore these was not included with the Chihu;huan Desert but rather were interpreted as miscellaneous forms of shrubland and grassland, respectively.

33. Route of the river- Land forms and vegetation defined by two rivers:. the Big Bend stretch of Rio Grande in the Trans-Pecos drainage of the Basin and Range province provides an environment for riverine vegetation in the midst of the Chihuhuan Desert. In the far background the Rio Grande emerges from Santa Elena Canyon. This is a chasm with sheer walls up to 1500 feet deep cut through the limestone of Sierra Ponce (the Chihuhua side in the left two-thirds of the background) and Mesa de Anguila (the Texas side in the right one-third background) by sand-laden, moving water over geologic time. Sierra Ponce- Mesa de Anguila are a mesa, "an uplifted block of Lower Cretaceous limestone", that is the remainder of a former and much higher mesa from which the softer Upper Cretaceous limestones and shales were eroded (Maxwell, 1968, ps. 87-89). Even short of its previous glory the mesa is a backdrop land form to a once-mightier river that now relies on another river, the Rio Conco, for most of its water.Even with most Rio Grande water dammed and diverted for human agricultural and municipal uses, there is adequate aquatic environment for well-defined range cover types.

Immediately along the river banks (or back a short distance where there are recently formed sand bars) there is a zone of riparian vegetation made of two end-to-end disparate plant communities or, perhaps more descriptively, stands: 1) willow and 2) reed. The woody vegetation (a willow shrubland) and the herbaceous vegetation (a grassland of common reed [Phragmites communis]) are each spatially short and narrow "strips" of distinct plant communities which obviously interact yet are physically segregated. There are of course other plant species in the willow and reed communities but the two species so dominate their respective vegetation that most other species seem of miniscule importnce, at least from standpoint of range cover types.

Adjacent to riparian vegetation on the river flood plain and covering a much broader breath of rangeland is vegetation known as bosque or bosques (mostly an Anglo-adopted Spanish word). Bosque vegetation is dominated by mesquite and other woody plants, most of which are of shrub rather than tree form and dimension.

Willows are also usually shrubs or, at most, small trees. In restricted locations along short stretches of the Rio Grande riparian vegetation consist of large cottonwood trees, often with willows as a lower woody layer, that form small gallery forests. These gallery forests were treated separately under Miscellaneous Forest Types and Southern and Central Forests (Woodlands and Forests). Gallery forests were not discernable in these landscape-scale photographs.

Technical Note: The term riparian may have somewhat different meanings or connotations as used Hydrology, Aquatic and Riparian Ecology, Range Management, or Forestry not to mention legal usages in the extremely specialized area of water law and policy. The Glossary of Hydrology (Wilson and Moore, 1998) defined riparian thusly: "Pertaining to or situated on the bank of a body of water, esp. of a watercourse such as a river; e.g. 'riparian land' situated along or abutting upon a stream bank..." (listed as synonyms were riparial and riparious). A Glossary of Terms Used in Range Management (Jacoby, 1989), an official publication of the Society for Range Management, defined riparian as: "Referring to or relating to areas adjacent to water or influencecd by free water associated with streams or rivers on geologic surfaces occupying the lowest position on a waatershed". Jacoby (1989) defined riparian zone: "The banks and adjaacent areas of water bodies, water courses, seeps and springs whose waters provide soil moisture sufficiently in excess of that otherwise available locally so as to provide a more moist habitat that that of contiguous flood plains and uplands".

Clearly, hydrophytic or streamside (= riparian) vegetation such as that dominated by willows and reeds on the edge of the Rio Grande constitutes plant communities not only different from other vegetation, but growing on a different environment from that on and of the flood plain. Alternating patches or strips of willow and reed vegetation were clearly visible in both of these photographs. Also readily discernable was the more extensive (and drier-soil) area of mesquite bosque that had developed adjacent to riparian zone vegetation.

Land in the immediate foreground was on an upland overlooking the Rio Grande and Sierra Ponce-Mesa de Anguila that provided a vista of riparian and flood plain vegetation. Vegetation on the upland foreground (vista point) was Chihuhuan Desert dominated by the climax species, creosotebush (Larrea tridentata) and tasajillo or pencil cholla (Opuntia leptocaulis). Quite a study of and contrast in range vegetation.

Big Bend National Park, Brewster County, Texas. June.

34. Two Rio Grande riparian range types- On the north bank of the Rio Grande (south bank is the Rio Bravo) the two major and segretated riparian plant communites of this strean were holding forth: 1) willow scrub made up primarily of sandbar or coyote willow (mostly Salix interior= S. exigua) on the left and 2) common reed grassland on the right.

The back "curtain" of this study in range vegetation was the immense block of Lower Cretaceous limestone that formed the mesa, Sierra Ponce, in Chihuhua. The formation outcrops from top to bottom were: Santa Elena, Sue Peaks, and Del Carmen (Maxwell, 1968, Fig. 92, p. 89). This stretch of river was just east of Santa Elena Canyon.

Rio Grande, Big Bend National Park, Brewster County, Texas. June, early estival aspect and pre-bloom stage.

35. Santa Elena carrizo- Stand of common reed (carrizo in Spanish) at mouth of Santa Elena Canyon. An example of reedbed or reedbank vegetation. Common reed forms dense colonies that develop as linear strips of single-species vegetation which alternate with willow scrub communities of similar size and shape as the two major types of riparian vegetation growing along the Rio Grande as it flows through the arid climate of the Chihuhuan Region, including the Chihuhuan Desert, and eastward into semiarid climate where the regional vegetation is a mixed grass species-thornscrub savanna.

Common reed (Arundineae tribe of Arundionoideae subfamily) is one of the most widely distributed grass species in the world growing in North America, South America, Africa, and Eurasia. Polunin (1960, p. 98) reported that common reed "is often claimed to be the most widely distributed vascular plant species in the world".

Carrizo was the native grass most commonly used for purposes aside from forage or game habitat by Indians and, later, Mexican peasants living along Chihuhuan Region rivers like the Rio Grande. Maxwell (1968, ps. 105-106) described use of the culms of common reed as building material for huts. Arrow shafts were supposedly fashioned from the smaller distal portions of carrizo culms.

Santa Elena Canyon, Big Bend National Park, Brewster County, Texas. June, late pre-bloom stage. No FRES, Kuchler (1964, in Garrison et al., 1977), or Society for Range Management (Shiflet, 1994) units for common reed or carrizo range cover type. Erroneously omitted by Brown et al. (1998, p. 46) but there should be a Common Reed-Sedge Series, 243.62, of Chihuhuan Interior Marshland, 243.6. Chihuhuan Deserts- Low Mountains and Bajadas Ecoregion, 24c (Griffith et al., 2004).

36. Reeds along the Rio- In this short reach of the Rio Grande (Rio Bravo) common reed or carrizo was growing on both stream banks in direct contact with river water. Reedbeds or reedbanks were the riparian vegetation. This riparian strip of grassland was interrupted at intervals by willow scrub (or, alternatively, riparian willow vegetation was "broken" periodically by stands of carrizo). Though extremely small in size (narrow often short colonies) reedbeds were the major source of forage along the river. Reedbanks appeared to produce appreciably more biomass than willows, but a combination of reed forage and willow browse provided a good combination diet plus cover for game animals like deer. Water, high-quality feed, and shade makes for great wildlife habitat. Reedbeds were ideal range. Maxwell (1968, p. 105) told how Mexican stockmen fired the carrizal (reedbeds) in the winter to insure green feed from tender carrizo shoots in the spring.

Common reed was far more valuable than would be suspected by the extremely small portion it made up of total Chihuhuan Region vegetation. This was yet another example of the incaluable importance of riparian range vegetation. Importance of reedbanks in prevention or reduction of soil erosion was shown in the next two photographs.

Rio Grande (Rio Bravo), Big Bend National Park, Brewster County, Texas. Sierra Ponce in Chihuhua in background. June, early estival aspect and late pre-bloom stage. NO FRES, Kuchler (1964, in Garrison et al., 1977), or Society for Range Management (Shiflet, 1994) units for carrizo or reed rangeland cover type. Should be a Common Reed-Sedge Series, 243.62, of Chihuhuan Interior Marshland, 243.6 (Brown et al., 1998, p. 46). Chihuhuan Deserts- Low Mountains and Bajadas Ecoregion, 24c (Griffith et al., 2004).

37. Reedbed on the river bank- Common reed or carrizo growing on bank of the Rio Grande. Photographs showed structure typical of a stand or colony of carrizo. Specifics of bank-cutting was unknown. Obviously there had been recent flooding without enough subsequent river flow to carry away shoots of reed. Whether this was natural (= geologic) or accelerated soil erosion was not known. River banks are dynamic with some stream banks always being eroded while other banks are being rebuilt (vertical accretion) by sediment deposited from eroded banks. Soil (mostly sand) eroded from this bank was being deposited as a point bar on a stream meander downriver. This location was in close proximity to mouth of Sant Elena Canyon where confined stream flow rushed onto a broad floodplain of that reach of the Rio Grnde. A handy reference or guide for discussion of stream processes and patterns was the small book by Morisawa (1968).

Big Bend National Park, Brewster County, Texas. June, late pre-bloom phenological stage. No FRES, Kuchler (1964, in Garrison et al., 1977), or Society for Range Management (Shiflet, 1994) for reedbed (carrizal) range cover type. If had not been mistakenly omitted from Brown et al. (1998, p. 46) would be Common Reed-Sedge Series, 243.62, of Chihuhuan Interior Marshland, 243.6. Chihuhuan Deserts- Low Mountains and Bajadas Ecoregion, 24c (Griffith et al., 2004).

38. Reeds in the Basin and Range- Common reed or carrizo on a bank of the Rio Grande. This photograph of the north bank of the Rio Grande with Bofecillos Mountains in the background showed physiography of the Basin and Range province from a "close-in" perspective. This slide also presented a good example of the physiogonomy of a reedbed as riparian range vegetation. The perennial nature of common reed was apparent, including some perennality in shoots. More-or-less dead shoots from previous growing seasons (those of last year or several years ago was not determined) were mixed in with current season's shoots. Some of last year's shoots had grown small secondary shoots from intercalary meristem of internodes. Thus plants of common reed sometimes have perennial shoots (ie. in this herbaceous species it is not just the rootcrown or proaxis that lives for more than one growing season; some shoots of this herbacsous species develop lateral shoots proving that there is some aboveground perennating tissue).

Presidio County, Texas. June, early estival aspect and late pre-bloom stage. No FRES, Kuchler (1964, in Garrison et al., 1977), or Society for Range Management (Shiflet, 1994) designations for reedbed (carrizal) rangeland cover type. Common Reed-Sedge Series, 243.61, of Chihuhuan Interior Marshland, 243.6 (Brown et al., 1998, p. 46, but inadvertently left out).Chihuhuan Deserts- Low Mountains and Bajadas Ecoregion, 24c (Griffith et al., 2004).

39. "Reed swamp" on the Rio Grande- It was explained in the introduction to the reedbed portion that one of the most commonly used examples of a plant association or, the seral equivalent, associes by pioneer British and American plant ecologists (eg. Warming, Clements, Weaver, Tansley) was the so-called "reed-swamp". This included the Phragmites comminus or P. australis consociation or, the seral unit, consocies. Since those fledging years in which successional and vegeetation classification concepts were formulated wetland specialists, including ecologists, have defined swamp as wetland vegetation dominated by woody plants whereas wetland plant communities dominated by herbaceous species are marshes. Common reed-dominated vegetation (which is limited to permanently wet or, at least, very moist soils) is now recognized as a marsh not swamp cover type. Even the term marsh (ie. reed marsh) is not as appropriate or as descriptive of the Rio Grande reedbeds as the title, common reed-dominated riparian vegetation. Plant communities that are restricted to stream banks or adjacent habitats should most properly be entitled and described as having developed in the riparian zone; hence,as riparian vegetation.

Reedbanks are, therefore, riparain range plant communities or, in terminology and context of this publication, the common reed (Spanish, carrizo) or reedbed (Spanish, carrizal) range cover type is riparian range. Plant communities of P. australis in many other areas, including elsewhere in North America, are actual or true marshes. These are properly termed reed marshes. One of the largest and most notable of these reed marshes is the vast network of P. australis and Typha spp.- dominated wetlands known as the Mesopotamian marshlands that developed over the immense lowlands of the Tigris and Euphrates Rivers. Most of this reed-cattail marsh was in Iraq and home to the Ma'dan or Marsh Arabs. This wetland ecosystem and the Ma'dan cultural system is slowly being restored following occupation by American military forces in the current war for Iraqui "liberation". Compared to such vast reed marshes the scattered patches of riparian reedbed range along banks of the Rio Grande were mere parcels of a riparian oasis in the Chihuhuan Desertscrub and Tamaulipan thornscrub savanna. Common reed range communities were nontheless priceless for their forage, cover, soil protection, biodiversity, and so on.

This view of a reedbed was on the outermost edge of the Rio Grande where young shoots of carrizo were in early growth. Older (larger) shoots, including both dead and previous-growing season shoots having intercalary branches as well as current- season shoots, were farther back (deeper) in the colony. Most grazing by ungulates would be on this vegetational "frontier" with consumption of biomass deep inside the reedbed limited to smaller vertebrates like rodents and invertebrates (eg. insects).

Big Bend National Park, Brewster County, Texas. June, early vegetative growth stage (see "very next" photograph). No FRES, Kuchler, Society for Range Management (Shiflet, 1994) units for what was obviously a reed rangeland cover type. Common Reed-Sedge Series, 243.62, of Chihuhuan Interior Marshland, 243.6 (Brown et al., 1998, p. 46 but inadvertently omitted). Chihuhuan Deserts- Low Mountains and Bajadas Ecoregion, 24c (Griffith et al., 2004).

40. Reed reproduction- Asexual reproduction in common reed or carrizo includes all three types of shoots in grasses: 1) tillers which are vertical, intravaginated shoots, 2) rhizomes which are horizonal, belowground, extravaginated shoots, and 3) stolons which are horizonal, aboveground, extravaginated shoots. Seen here were stolons that had produced tillers at the "frontier line" of the advancing colony of this clonal or modular plant.

At edge of water line in bed of Rio Grande, Presidio County, Texas. June, early growth phenological stage.

41. More reed reproduction ("if and when it rains")- Two stolons developed from a rhizome of common reed growing beneath sand along the Rio Grande. This soil was on the backside of a reedbed (the dry side or side of the reed colony away from the water's edge). Green leaves had recently emerged from these shoots which otherwise appeared dead. Will the new shoots (young ramets of the established genet) survive by being able to translocate water, mineral nutrients, and photosynthate from the parent plant or are they dependent on water filling the pores of the dry sand in which they were rooted in?

Presidio County, Texas. June, early summer but early "green-up" for these young 'uns.

42. Reed tiller tops- Distal or apical portions of tillers of common reed or carrizo. The most obvious shoots of common reed or carrizo are tillers, but these tall vertical.shoots typically arise from rhizomes or stolons the "unsung shoots". Polunin (1960, p.98) felt that P. communis (= P. australis) was not only one of the most widely distributed plants on Earth but also one best adapted for dispersal as it has wind-carried, parachute caryopses; water-borne, buoyant rhizomes; and soil surface-covering stolons. Dispersal of propagules or disseminules by wind is anemochory while dispersal of germules by water is hydrochory, and dispersal by gravity is barochory. Dispersal of propagules by more than one agent or mechanism is polychory. Is it any wonder that the polychorous P. communis has one of the largest discontinuous (= disjunct) ranges (geographical distributions) of any extant plant species?

Chapman (1996, p. 145) summarized sexual reproduction in P. australis. He reported that establishment of new reed plants from seed varied tremendously. Although common reed flowers plenteously, grain production is generally low and seedling establishmen is usually rare in native populations. Sexual reproduction is much greater in domestic reed than in natural reedbeds (Chapman, 1996, p. 146). Even phenotypic variation within a clone (genet) is fairly limited.

It would be interesting to know if the isolated stands or colonies of carrizo along the Rio Grande are distinct genotypes (genets, clones, or modules) or if some of these colonies are of the same clone with a "broken" distribution due to disturbances (possibly flood, fire, grazing, human harvest for building materials) that resulted in death of parts of clones (loss of ramets) at various intervals in space and time.

Tillers in the upper (top) photograph were young, current-season's shoots. These tillers had arisen in the current growing season from rhizomes. There were no secondary shoot or "branches" arising from intecalary meristem along the tiller. Tillers in the lower (bottom) photograph were last year's or the previous growing season's shoots with secondary tillers or "branches" that developed from intecalary meristem during the current growing season. The "branches" or secondary (intecalary) shoots (which are also tillers) demonstrated that some tillers are perennial. It is not just the plant (the genetic individual= genotype= genet) that is perennial, but some shoots (some modules= clones) are perennating organs.

Tillers in both photographs were on the river (water) side of reedbeds growing along the Rio Grande. Presidio County, Texas. Advanced pre-bloom phenological stage.

Clarifying Note: Beginning students could easily confuse common reed (P. communis= P. australis) with giant reed (Arundo donax) from both similar physical appearance and morphology of plants as well as from the common name of "reed". (The Spanish names, carrizo and carrizal, would apply only to the native P. communis and not A. donax this latter of which was introduced and naturalized after establishment of Hispanic culture.) Both of these arundinoid grasses occur in some areas and, to add to confusion, in the same general habitat therein. Both species grow along the Rio Grande: in fact, sometimes side-by-side (Powell, 2000, p. 46). A. donax is an exotic species introduced from the Old World (Hitchcock and Chase, 1951) that appears to have naturalized. In North America spread of A. donax appears to be strictly by asexual reproduction and not by sexual means because viable seed production seldom if ever occurs. Instead dispersal is by fragmentation of rhizomes and/or root systems or, less commonly, by secondary shoot development from detached tillers. This fragmentation is often human propagation (culms of tillers are used for reed instruments, the entire shoot has some agronomic potential as a biomass fuel crop).

A. donax has become a very aggressive, exotic weed in riparian and general aquatic ecosystems. A. donax should not be propagated along rivers. Rather it should be controlled or, better yet, eradicated if possible from natural range ecosystems such as those of the various range types along the Rio Grande.

This author/photographer did not find any A. donax on the Rio Grande. All photographs of reedbeds presented above consistedly strictly of P. communis (or associated native willow scrub or mesquite bosque).

54. Range vegetation on stabilized sand dune along Pacific Ocean- The unusual plant community on this high dune was in the Coast Ranges physiographic and geologic province in northern Oregon. This ocean-front and its herbaceous community could be readily visualized and interpreted as a "headland" though certainly not the more characteristic high and rocky headlands found in the same area. The ocean-most extremity of vegetation (foreground) was within the swash zone at high tide such that the beach-bordering vegetation of this range ecosystem could also be viewed as a dune plant community.

Zonation of plant species and groups was remarkably well-defined. The first zone or swash zone (foreground) was composed of salt rush (Juncus lesueurii= J. balticus var. lesueurii). Immediately behind the salt rush zone was another zone dominated by the native legume, giant vetch (Vicia gigantea). In turn the umbels sea-watch (Angelica lucida) and beach lovage (Ligustichum scoticum) formed a small "patch" behind the giant vetch-dominated "strip". Behind the umbel "patch" vegetation was dominated by giant horsetail (Equisetum telmateia) but with the grasses Idaho bentgrass (Agrostis idahoensis), seashore bluegrass (Poa macrantha), and Pacific reedgrass (Calamagrostis nutkaensis) and silverweed (Potentilla anserina).

Yaquina Bay State Park, Lincoln County, Oregon. June. Coast Range- Coastal Lowlands Ecoregion, 1a (Thorson et al., 2003).

55. High sand dune Pacific Ocean range community- This high sand dune immediately beyond the Pacific Ocean beach had been stabilized by a plant community dominated by giant horsetail with several grass species, including seashore bluegrass, Pacific reedgrass, and Idaho bentgrass, serving as associates. Also present, though widely scattered, was Oregon Iris (Iris tenax).

Yaquina Bay State Park, Lincoln County, Oregon. June. Coast Range- Coastal Lowlands Ecoregion, 1a (Thorson et al., 2003).

56. Sand dune range plant community along Pacific Ocean- Adjacent to swash zone of Pacific Ocean in the Oregon Coast Ranges was this natural vegetation of unusual composition. The low shrub in immediate foreground was deer brush (Ceanothus intergerrimus). Behind this shrub was a zone of vegetation consisting of giant horsetail, Pacific reedgrass, seashore bluegrass, Oregon iris, giant vetch, and silverweed.

Yaquina Bay State Park, Lincoln County, Oregon. June. Coast Range- Coastal Lowlands Ecoregion, 1a (Thorson et al., 2003).

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57. Salt rush (Juncus lesueuri= J. balticus var. lesueuri)- A dominant range species at the edge of swash zone that formed the lowermost segment of vegetation at base of sand dunes along the Pacific Ocean in the Oregon Coast Ranges. Inflorescences at anthesis.

Yaquina Bay State Park, Lincoln County, Oregon. June.

59. "Shades of the Past": consociation of Great Basin, basin, or giant wildrye (Elymus cinereus). This relict stand of basin wildrye was but a remnant example (relatively large though it was) of what was once a major climax plant community in much of the Intermountain Range Region. Grassland comprised of natural single-species stands of basin wildrye or of basin wildrye and inland saltgrass (Distichlis stricta) was often the most productive natural (pre-Columbian) vegetation of the Great Basin and parts of adjoining areas. This remarkable high yield was in terms of both biomass (= herbage) and useful forage. In frontier times basin wildrye was cut for hay and was the largest source of harvested forage for U.S. Army cavalry horses (A.L. Lesperance, personal communication). Young and Sparks (2002, p. 51) mentioned "extensive areas" of Great Basin wildrye dispersed with wetlands, salt desert scrub, and other Basin and Range vegetation.

In Classification of Native Vegetation of Oregon by the Oregon Natural Heritage group (Kagan et al., 2004) two Great Basin wildrye plant associations were recognized: 1) basin wildrye bottomlands (Leymus [= Elymus] cinereus) and 2) giant wildrye alkali marsh-saltgrass (Leymus [= Elymus] cinereus- Distichlis spicata). The example of giant wildrye shown here was the basin wildrye bottomlands association.

Undoubtedly much of the original climax basin wildrye-dominated vegetation was destroyed to make way of farming and other land uses. Much of the remaining giant wildrye range was overgrazed and/or overmowed for hay. As a result relatively little of the original basin wildrye range vegetation remains.

The relict vegetation shown above was in the Harney Basin of the High Lava Plains province (Franklin and Dyrness, ps. 6, 32-34). This extensive stand was one of the few tracts of its species that remained in the former grassland-marsh complex that developed in the Silvies River valley. Inland saltgrass was absent from this stand which again was an example of the basin wildrye bottomlands form of this type.

Efforts continue to restore this basin wildrye to some of the land once dominated by it. Magnar and Trailhead are two accessions of Great Basin wildrye that were selected and released by Soil Conservation Service Plant Materials Centers. Seed is available from several seedsmen.

Harney County, Oregon. June. Mid-bloom phenological stage. Basin wildrye bottomlands association of Kagan et al. (2004). Northern Basin and Range- High Desert Wetlands Ecoregion, 80e (Thorson et al., 2003).

60. Basin wildrye range- Interior of the relict Great Basin wildrye grassland presented in the immediately preceding photograph showing species composition, plant morphology and dispersion, and community structure. Botanical composition and vegetation stucture were simple: one species and one layer of vegetation. Plant growth form was largely cespitose (=bunched or clumped). Basin wildrye does have short rhizomes, but it retains the basic morphological form of a bunchgrass.

In the virgin landscape much of the graminoid vegetation here in the Silvies River valley was wetland (bulrush-dominated marsh) and associated mesic grassland consisting largely of basin wildrye. Much of the marsh was converted to hay fields of introduced Eurasian perennial grasses while the climax virgin grassland was farmed, overgrazed, and otherwise largely destroyed or highly modified so that most of the basin wildrye was replaced by other plants.

Unfortunately the value of basin wildrye communities was not appreciated until most of it was gone. Even plant ecologists seem not to have paid much attention to this robust, highly productive species and the range plant communities dominated by or made up of it (see for eg. brief coverage given by Franklin and Dyrness, 1973, p. ). Even the Society for Range Management (Shiflet, 1994) did not designate a basin wild rye rangeland cover type. Perhaps so much of the original basin wildrye vegetation was gone that it had little practical use.

Some taxonomic authorities recognized two varieties of Elymus cinereus. Hitchcock and Cronquist (1973) distinguished between E. cinereus var. cinereus having glabrous sheaths and E. cinereus var. pubescens having pubescent sheaths.

Harney County, Oregon. June. Mid-bloom phenological stage. Basin wildrye bottomlands association of Kagan et al. (2004). Northern Basin and Range- High Desert Wetlands Ecoregion, 80e (Thorson et al., 2003).

61. Giant wildrye in the woods- Basin wildrye is widely distributed in northwestern North America. In its geographic range basin wildrye grows on diverse habitats varying from Northern Great Plains grasslands southward into wet and fairly saline sites in Great Basin vegetation. Between these latitudinal extremes giant wildrye grows in mesic environments in mountain forests. Shown here was an example of a consociation of basin wildrye had developed on a moist yet well-drained opening within a ponderosa pine (Pinus ponderosa) forest in the Blue Mountains province (Franklin and Dyrness, 1973, ps.6, 27-29). Based on the Orgeon Natural Heritage Classification of Native vegetation of Oregon (Kagan et al., 2004) this was a basin wildrye bottomlands plant association, there being no inland saltgrass in this specific community and no vegetation unit for this basin wildrye in ponderosa pine forest.

Grant County, Oregon. June. Soft-dough phenological stage. Variant of basin wildrye bottomlands association of Kagan et al. (2004). Blue Mountains- John Day/Clarno Uplands Ecoregion, 11a (Thorson et al., 2003).

Throughout much of eastern North America the original forest vegetation was highly modified by European (white) man. Over much of this vast domain forest were cleared and converted into cropland, including introduced pasture. Some land was cutover forests that through secondary plant succession partially recovered as forest vegetation, often with appreciable propotions of naturalized introduced plant species. Over the course of time (decades in many instances) much of this land that had been farmed was either abandoned (in most cases of which it reverted to second-growth forests) or used as grazing land managed extensively and, usually, improperly (ie. mostly just grazed and fertilized, reseeded, renovated, etc. only periodically, if that, and generally used without regard to proper husbandry-- of pasture or livestock). Some of the second-growth forest on cutover land was also grazed/browsed by livestock and/or wildlife. Aside from stocking with livestock and supporting free-ranging wildlife that might be hunted, about the only human input to this mostly mismanaged, more-or-less defacto pasture was mowing or shredding (when the weeds and brush had gotten "too thick" and/ or grass shoots had grown so tall as to appear unsightly and prompt an oral response--which might or might not be followed by action--such as "It looks sorta rough; guess I ought to knock her down some").

This category of grazing land is probably the major form of naturalized pasture (naturalized range) in North America. (Naturalized annual grassland in California is the most recognized type of naturalized range, and it is generally much better managed than naturalized pasture to the east.) In northen and central regions of North America naturalized pasture (grazing land and range are appropriate synonyms for pasture when the adjective naturalized is applied) is typically dominated by Eurasian domesticated perennial grass species and/or weedy (ruderal) annual grasses of cool-season tribes especially Festuceae (Poaeae) and Aveneae. In more southern regions of North America naturalized, introduced perennial grasses tend to be of African or South American origin and in warm-season tribes, notably Andropogoneae, Paniceae, and Eragrosteae. Annual grasses on these southern anthropogenic pastures--which can be grasslands, savannahs, woodlands, or forests-- include many of the same genera and species (in particular Bromus, Avena, Hordeum) as those on naturalized grazing lands to the north and east and on the Pacific Slope. Some of the annual cool-season grasses in the south are native. Little barley (Hordeum pusillum) is the most common (widespread) example. In addition to these annual festucoid grasses, annual panicoid species are frequently abundant on naturalized southern range. Some of these are also native. Texas millet or Texas panicgrass (Panicum texanum) is perhaps the most common of these species. Most warm-season annual grasses are naturalized (primarily weeds imported from Eurasia). Hairy crabgrass (Digitaria sanguinalis), southern crabgrass (D. ciliaris), and stinkgrass (Eragrostis cilianensis ) are examples of this group that are common in the Texas Pineywoods.

An example of naturalized pasture typical of southern North America was provided by a Texas Pineywoods woodland (or savannah, depending on interpretation) dominated by native oaks, water oak (Quercus nigra) and southern red oak (Q. falcata), in the tree layer and agronomic (introduced) perennial panicoid grasses, bahiagrass (Paspalum notatum) and Vaseygrass (P. urvillei) in the herbaceous layer. Both of these domestic Paspalum species were introduced from South America. Interestingly enough, Dallisgrass (P. dilatum) which is a more widely naturalized Paspalum species in Texas was not observed on this example pasture by the author. Apparently Dallisgrass is not competitive with the rhizomatous bahiagrass and taller, ranker Vaseygrass in the more mesic habitat of the Pineywoods, at least on sites represented by the example shown here. There were some native Paspalum species, including thinseed paspalum (P. setaceum) and brownseed paspalum (P. plictum), but these were "totally overwhelmed" by the two exotic domestic paspalums. Other grasses present included the natives, purpletop (Tridens flavus), broomsedge (Andropogon virginicus), beaked panicgrass (Panicum anceps), and Texas millet. With the exception of the latter, these native grasses were most common under oak canopy. It seemed that they were more competitive with the two dominant Paspalum species mostly under shade. Common bermudagrass (Cynodon dactylodon) was present, but not abundant. Like naturalized Dallisgrass, bermudagrass appeared to be not competitive with bahiagrass and Vaseygrass. The most common forb in this naturalized pasture was wooly croton or hogwort (Croton capitatus), a native annual.

The herbaceous layer(s) of vegetation of this grass-oak woodland was, once again, largely dominated by introduced species of Paspalum (P. notatum and P. urvillei) as well as such other naturalized exotic grasses as Cyndon dactylon and even less locally common exotics as P. dilatum and Bothriochloa ischaemum along natives like Tridens flavus, Andropogon virginicus, and and scattered individuals of various Paspalum species like P. plicatulum, P. setaceum, P. pubiflorum, and P. bifidum. Herbage was largely or overwhelmingly of naturalized alien panicoid grasses. This range vegetation is not widely distributed like California annual grassland so there was no SRM rangeland cover type for it. Nor was there an SAF forest cover type for woodland dominated by water oak and southern red oak.

62. Naturalized woodland (or savannah) pasture in Texas Pineywoods- An herbaceous understorey dominated by bahiagrass with Vaseygrass as the associate species that developed beneath scattered young trees of water and southern red oak was used as naturalized range for beef cattle and white-tail deer at western edge of Texas Pineywoods. The largely anthropogenic (man-made) vegetation of this woodland range included some native grasses of which the major perennials were (in approximate order of abundance) purpletop, broomsedge bluestem, beaked panicgrass, thinseed paspalum, and brownseed paspalum. These species were most abundant (greatest density with highest cover) in shade of oaks. The native Texas millet was the most common and productive annual grass. The introduced( and widely naturalized) common bermudagrass and Johnsongrass were also locally abundant species, but Dallisgrass was not present at other than trace amounts (ie. it was not found at cover or density that was floristically meaningful). There were some Carex and Cyperus species, but these were infrequent and could not be identified without inflorescences which had been removed by recent mowing. The most common forb was the native annual euphorb, hogwort or wooly croton.

This pasture had been recently destocked of cattle and shredded. Prior to destocking it had been grazed to light degree of use, but subjected to frequent mowing at comparatively low stubbgle height. Aggressiveness and highly competitive features of bahiagrass and Vaseygrass effectively excluded most other grasses. These two agronomic paspalums apparently were less competitive under conditions of shade. It was likely that frequent, short mowing (shredding) had limited cover and density of Johnsongrass, a common and aggressive weedy perennial through much of southern and central North America. The conventional wisdom that Johnsongrass can tolerate almost any abuse except repeated close grazing and/or mowing was probably demonstrated on this naturalized pasture.

Domination of herbaceous cover by bahiagrass became a widespread phenomenon over much of the North American southland within a few decades of its introduction. Bahiagrass became an aggressive weed much like Johnsongrass though it has a considerably smaller area of distribution. Since introduction of bahiagrass numerous cultivars of bermudagrass were developed that are much more productive (higher-yielding) than bahiagrass or common bermudagrass, but bahiagrass and, to a lesser extent, common bermudagrass outcompete hybrid bermudagrasses and take over fields planted to the superior forages (and inferior competitors). The ranker-growing Vaseygrass maintains cover and spreads less effectively than bahiagrass under heavier defoliation, but it typically fares better under closer mowing and grazing than the tall-growing Johnsongrass and the shorter, stoloniferous bermudagrass (at least under the warm, mesic.habitats of the Pineywoods). Although Vaseygrass is semirhizomatous it usually grows more as a bunchgrass (much like the smaller Dallisgrass), which combined with its elevated apical meristems, puts it at competitive disadvantage to bahiagrass.

Freestone County, Texas. October; grain-ripe phenological stage in the two major paspalums. No units of native vegetation were applicable for this naturalized woodland pasture. East Central Texas Plains- Floodplains and Low Terraces Ecoregion, 33f (Griffith et al., 2004).

63. Naturalized grass under native trees- Physiography and structure of a naturalized herbaceous layer beneath scattered young to mid-age water and southern red oaks. Bahiagrass and Vaseygrass were the principal herbaceous species with Vaseygrass the associate to co-dominant species on local spots (microsites) having less cover of bahiagrass which was the overall dominant. Common bermudagrass and Johnsongrass were other locally abundant introduced perennial grasses. Native perennial grasses included (in this approximate order of abundance) purpletop, broomsedge bluestem, beaked panicgrass, thinseed paspalum, and brownseed paspalum. Texas millet was a common and even locally dominant native annual grass. There were some species of caric sedge (Carex spp.) and flat or umbrella sedge (Cyperus spp.) which could not be identified in their existing vegetative state. Hogwort or wooly croton, a native monecious euphorb, was the major forb.

The oak species were present as individuals as well as clumps (assemblages) of trees. Bahiagrass and Vaseygrass were comparatively less abundant under tree shade, sometimes even being associate to mere incidental species under dominance of native grasses. Tree regeneration was limited to seedlings so short that the shredder blades passed over them. Browsing was not evident, probably because mowing had twisted off taller seedlings and saplings. There were some larger (taller) young trees close enough to existing trees that the rotary shredder could not get to them. The history of mowing and grazing management was unknown though this man-modified naturalized pasturage was representative of others in this western edge of Texas Pineywoods.

The spatial distribution of trees on this naturalized grazing land was subject to interpretation as either woodland or savanna. There were stands of water and southern red oak in which tree density and proximity were such that crowns interlocked to form a forest canopy (though a relatively open canopy layer). There were other locations on this naturalized range where there were no trees and enough distance among clumps or stands of trees that such treeless vegetation was grassland. These treeless (grassland) areas made up more of the total land area of the pasture than did that supporting trees. From that perspective the total or general vegetation of this pasture could be interpreted as that of a savanna. Conversely, it could be viewed as an overall woodland (as a combination of closed-canopy stands of oak and open grassland).

Freestone County, Texas. October; grain-ripe phenological stage in bahiagrass and Vaseygrass. No units of native vegetation were applicable for this naturalized woodland pasture. East Central Texas Plains- Floodplains and Low Terraces Ecoregion, 33f (Griffith et al., 2004).

64. Sward of Pineywoods naturalized pasture- Herbaceous layer of a naturalized woodland or savanna pasture on the western edge of Texas Pineywoods. Bahiagrass and Vaseygrass were the two major herbaceous species with bahiagrass almost always dominant or co-dominant with Vaseygrass which was, communitywide, the associate herb. Other grasses included the introduced species of common bermudagrass and Johnsongrass along with native grasses. These latter included perennial species of purpletop, broomsedge bluestem, beaked panicgrass, thinseed paspalum, and brownseed paspalum (in that approximate order) and the annual, Texas millet. There was some cover of caric sedges and umbrella or flat sedges. Forbs in this naturalized vegetation were limited to hogwort or wooly croton which grew in patches, none of which were in the sward presented here.

Freestone County, Texas. October; grain-ripe phenological stage in bahiagrass and Vaseygrass. No units of native vegetation were applicable for this naturalized woodland pasture. East Central Texas Plains- Floodplains and Low Terraces Ecoregion, 33f (Griffith et al., 2004).

65. Bahiagrass (Paspalum notatum) on naturalized Pineywoods range- Turf and sexual shoots of bahiagrass (first slide) and bahiagrass panicle (second slide), the dominant herbaceous species on naturalized pasture on water oak-southern red oak woodland or savanna at western perimeter of Texas Pineywoods. The "dead giveaway" characteristic of the bahiagrass panicle is the physical arrangement in which the two spicate branches usually arise at a common point on the central axis of this inflorescence and then frequently curving symmetrically downward (as in the second of these slides). Sometimes one panicle branch arises slightly higher on the floral axis (ie. one branch is a "little higher up on the seed stalk" than the other branch). Bahiagrass is the only Psapalum species in the Texas Pineywoods that regularly has this point of common origin for main branches in its panicle (Gould, 1975, p. 501).

Freestone County, Texas. October; grain-ripe phenological stage.

66. Spikelets of the naturalized dominant- Ripened caryopses on branches of a panicle of bahiagrass, the dominant herbaceous species on naturalized water oak-southern red oak woodland or savanna range at western extremity of Pineywoods in east Texas.

Freestone County, Texas. October; grain-ripe to grain-shedding phenological stages.

67. Spikelets of the natralized associate- Inflorescence of Vaseygrass, an agronomic perennial grass that was the associate herbaceous species on water oak-southern red oak woodland or savanna having an understorey dominated by naturalized domestic grasses at western perimeter of Pineywoods in east Texas. Panicle and flag leaf were shown in first photograph while the second slide featured ripening spikelets.

The Paspalum inflorescence type has traditionally been regarded as a panicle. Contemporary terminology described the major units of this panicle as "unilateral spicate branches" (Gould, 1975, p. 500). These branches in P. urvillei are ascending or erect and closely spaced among themselves and arising off both sides of the central axis (or rachis) of the panicle.

.Freestone County, Texas. October; hard-dough to grain-shatter stages of phenology (and all on the same branch of a panicle).

68. Texas millet or Texas panicgrass (Panicum texanum), a native, warm-season annual grass on naturalized (and, mostly, perennial) pasture- Local stand of Texas millet growing in the herbaceous understorey of water oak-southern red oak woodland or savannal range. The plants representing this species were growing on the example of naturalized Pineywoods grazing land shown and described above. Texas millet was not a major species in this pasture vegetation, but it was important and even dominant on some local sites (microhabitats).

.Freestone County, Texas. October; soft-dough stage of phenology.

69. A native, warm-season annual and its sexual shoots- A single plant of Texas millet (first photograph) and two sexual shoots of this species (second photograph). This small to mid-size panicgrass is widespread in North America ranging from the Atlantic Coast (Massachusetts) to Pacific Slope (California). Texas millet is often a weedy or ruderal species, even in areas where it is a native. More specifically, it tends to be a pioneer (at least early seral) species on disturbed sites. The specimen presented here was growing in the West Cross Timbers on land subjected to repeated heavy (close) mowing.

Erath County, Texas. October; anthesis.

70. Spikelets of a native warm-season annual grass- Spikelets on contracted panicle of Texas millet. Different spikelets were in various stages of flowering and fruit formation ranging from anthesis to grain in milk-stage. Examples were from plants growing in the West Cross Timbers on overmowed land. This annual species was well-represented in a Texas Pineywoods woodland dominated by water and southern red oak in the tree layer and bahiagrass and Vaseygrass in the herbaceous layer. Texas millet added two floristic components to this understorey: 1) annual life cycle and 2) native species. Other native grasses and grasslike plants in this herbaceous zone of vegetation were perennials.

The inflorescence of Texas millet is made up of erect or ascending branches that lie closely to and spread slightly away from the central axis (or rachis) resulting in a "tight" panicle yet with branching conspicuous enough that it is not a contracted panicle.

Erath County, Texas. October.

71. A native annual forb on naturalized (perennial) Pineywoods range- Hogwort or wooly croton (Croton capitatus) on water oak and southern red oak-dominated woodland with an understorey composed largely of naturalized bahiagrass and Vaseygrass. Hogwort was the most common forb in the herbaceous layer(s) of this highly man-modified vegetation.

.Freestone County, Texas. October.

72. Wooly croton or hogwort- Shoot apex of Croton capitatus showing fruit which is interpreted as a capsule composed of two to four carpels each of which contains one seed (Fernald, 1950, p. 959). All Croton species are monecious. The apt adjective "wooly" was clearly evident in this photograph.

.Freestone County, Texas. October.

73. Water oak (Quercus nigra)- Leader (first photograph) and detail of leaves (second photograph) of water oak growing on a naturalized woodland or savanna pasture on the western margin of the Texas Pineywoods. Southern red oak was the co-cominant woody species on this highly man-modified grazing land. Bahiagrass and Vaseygrass dominated most of the herbaceous layer of this naturalized range. There was limited regeneration of oaks on this frequently close mowed (overmowed) pasture with tree reproduction limited to short seedlings or saplings growing next to trunks of established trees where they were protected from a tractor-mounted shredder.

Freestone County, Texas. October.

74. Southern red oak (Quercus falcata)- Leaves of the co-dominant tree on a naturalized (anthropogenic) woodland or savannah pasture on western edge of Texas Pineywoods. Bahiagrass and Vaseygrass were understorey dominants over most of this naturalized range. Water oak was co-dominant with southern red oak in the tree layer(s). Regeneration of Quercus species was limited to seedlings that had not grown to mowing height or where growing close enough to trunks of larger trees that they could not be reached by the tractor-mounted rotary mower.

Freestone County, Texas. October