United States

Department of

Agriculture

Forest Service

Rocky Mountain

Research Station

Fort Collins,

Colorado 80526

General Technical

Report RMRS–GTR–16

Ecology and Management of

Sand Shinnery Communities:

A Literature Review

Roger S. Peterson and Chad S. Boyd

D2 H

D2
Abstract

Peterson, Roger and Chad S. Boyd. Ecology and management of sand shinnery communities: a

literature review. Gen. Tech. Rep. RMRS-GTR-16. Fort Collins, CO: U.S. Department of Agriculture,

Forest Service, Rocky Mountain Research Station. 44 p.

Sand shinnery is codominated by oak shrubs and mid and tallgrasses; the grasses are usually

taller than the oaks. The shrubs are the small, visible shoots of massive underground stem systems,

which are hundreds or thousands of years old. Sand shinnery occupies 5 to 7 M acres in western

Oklahoma, western Texas, and southeastern New Mexico. This area is a decrease from the original

due to land clearing for agriculture and eradication of oak to improve livestock-grazing. Oak control

is controversial because it can open sandy soils to wind erosion and can conflict with wildlife-habitat

quality. Of special concern are the lesser prairie-chicken and the sand dune lizard, which are heavily

dependent on shinnery vegetation. This review of climate, soils, vegetation, ecosystem dynamics

(including responses to drought and fire), wildlife and hunting, livestock grazing, and oak control in

the sand shinnery provides managers with a knowledge-base for decision-making and points to areas

where research is needed.

Keywords: sand shinnery, Quercus havardii, drought, fire, livestock grazing, brush control, wildlife,

lesser prairie-chicken, sand dune lizard

The Authors

Roger Peterson is a retired ecologist who worked for the USDA Forest Service Rocky Mountain

Research Station.

Chad Boyd is a doctoral candidate in range ecology at Oklahoma State University.

Publisher

Rocky Mountain Research Station

Fort Collins, Colorado

August 1998

You may order additional copies of this publication by sending your

mailing information in label form through one of the following media.

Please send the publication title and number.

Telephone (970) 498-1719

E-mail rschneider/rmrs@fs.fed.us

FAX (970) 498-1660

Mailing Address Publications Distribution

Rocky Mountain Research Station

3825 E. Mulberry Street

Fort Collins, CO 80524-8597
Contents

Introduction ..................................................................................................... 1

Ecology ........................................................................................................... 1

Historical Accounts ..................................................................................... 1

Geographical Distribution ........................................................................... 1

Plant Composition....................................................................................... 2

Ecological Classification ............................................................................. 3

Shin-oak and Its Hybrids ............................................................................ 4

Soils ............................................................................................................ 6

Climate ........................................................................................................ 8

Phenology ................................................................................................... 8

Autoecology of Shin-oak............................................................................. 9

Vegetation Dynamics and Water Relations ................................................ 10

Organic Matter and Decomposition ............................................................ 12

Nutrient Cycling .......................................................................................... 13

Drought ....................................................................................................... 14

Fire .............................................................................................................. 14

Wildlife Species .......................................................................................... 15

Mule Deer ............................................................................................... 15

White-tailed Deer.................................................................................... 15

Pronghorn Antelope ............................................................................... 15

Peccary (Javelina) .................................................................................. 15

Barbary Sheep ....................................................................................... 16

Lagomorphs ........................................................................................... 16

Rodents .................................................................................................. 16

Insectivores ............................................................................................ 17

Ecology and Management of Sand

Shinnery Communities: A Literature Review

Roger S. Peterson and Chad S. Boyd
Contents (Cont’d.)

Lesser prairie-chicken ............................................................................ 17

Quail ....................................................................................................... 19

Turkey .................................................................................................... 19

Mourning Doves ..................................................................................... 19

Perching Birds ........................................................................................ 20

Predators (Mammals and Birds) ............................................................ 20

Reptiles .................................................................................................. 20

Invertebrates .......................................................................................... 21

Management ................................................................................................... 22

Hunting ....................................................................................................... 22

Livestock Grazing ....................................................................................... 23

Management of Shin-oak ........................................................................... 26

Mechanical Means ................................................................................. 26

Fire ......................................................................................................... 26

Herbivory ................................................................................................ 27

Herbicides .............................................................................................. 27

Conclusions .................................................................................................... 30

Literature Cited ............................................................................................... 30

Appendix ......................................................................................................... 39

Acknowledgments

Two longtime shinnery researchers helpfully shared their knowledge: E. P. McIlvain,

retired supervisor of research, Southern Great Plains Field Station, Agricultural

Research Service (Woodward, OK), and R. D. Pettit, retired professor of range

management, Texas Tech University (Lubbock). James Zimmerman, research entomologist retired from The Smithsonian Institution provided information on shin-oak

gall insects. W. A. Dick-Peddie, professor of biology at New Mexico State University,

provided information on historical surveys of shinnery distribution. Calvin Smith,

director of the Strecker Museum, Baylor University (Waco, TX) gave information on

the New Mexico shinnery and provided figures 6 and 13. The New Mexico Department of Game and Fish provided figures 11 and 14. D. A. Swepston, district leader of

the Wildlife Division, Texas Parks and Wildlife Department (Canyon, TX), provided

information on mammals and hunting in Texas. Gesina (Niki) Threlkeld kindly

created the drawings. The authors are also grateful to T. G. Bidwell (Oklahoma State

University) and D. G. Brockway (Rocky Mountain Research Station) for review

comments that improved the manuscript, and to Dale Brockway and Deborah Finch

(Rocky Mountain Research Station) for facilitating publication of the review.
USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998 1

Introduction

Sand shinnery communities comprise the nation’s largest stand of oak and occupy 5 to 7 M acres of the southern

Great Plains, extending from northern Texas and western

Oklahoma southward into Chihuahuan Desert scrubland.

The stand is made up of ancient plants, most of them

hundreds or thousands of years old.

This oak forest is only 1 to 4 ft tall. The visible shrubs are

mostly the short-lived twigs of massive underground

stem systems, which slowly spread under areas up to 2

acres or more. The oaks codominate the community with

mid and tallgrasses that are usually taller than themselves.

Sand shinnery is defined by shin-oak (Quercus havardii

Rydb.), also called Havard oak, midget oak, and sand

shinnery oak. Shin and shinnery are Louisiana French

from chêne and chênière, which mean oak and oak woodland. Other shinneries cover millions of acres from Louisiana to central Texas and Oklahoma, but they are dominated by oak species that are small trees, rather than

codominated by oak-shrubs and grasses as is sand shinnery.

Management of this grass-oak mix is difficult and controversial. This review assembles what is known about

sand shinnery communities as part of the search for rational management.

Ecology

Historical Accounts

Historical observation of the sand shinnery comes from

nineteenth-century travellers. Josiah Gregg (1844) described a camp on his 1839 journey from Fort Smith,

Arkansas, to Santa Fe, New Mexico, as follows:

...we encamped in a region covered with sandy

hillocks, where there was not a drop of water to

be found: in fact, an immense sand-plain was

now opening before us,...being entirely barren

of vegetation in some places, while others were

completely covered with an extraordinarily

diminutive growth which has been called shinoak, and a curious plum-bush of equally dwarfish stature. These singular-looking plants

(undistinguishable at a distance from the grass

of the prairies) were heavily laden with acorns

and plums, which, when ripe, are of considerable size, although the trunks of either were

seldom thicker than oat-straws, and frequently

not a foot high. We also met with the same in

many other places on the Prairies.

In 1845, Lt. J. W. Abert (1846) recorded along the

Washita that “...the uplands were thickly spread over by

diminutive species of oak commonly called shin-oak, not

exceeding 2 ft in height. We occasionally saw clumps of

oak of greater size.” In 1854, R. B. Marcy described a

landscape encountered in present-day Beckham County,

Oklahoma:

...the country we traversed was exceedingly

monotonous and uninteresting being a continuous succession of barren sand-hills, producing no other herbage than the artemisia,

and a dense growth of dwarf oak bushes, about

eighteen inches high, which seem to have attained their full maturity, and bear an abundance of small acorns. The same bush is frequently met with upon the Canadian river,

near this longitude, and is always found upon

very sandy soil.

To the south, Valery Havard (1885), an army physician,

described the sand hills country of the Pecos River area of

Texas from visits in the early 1880s. He found only 4

shrubs: mesquite, catclaw acacia, Southwest rabbitbrush,

and shin-oak, which he thought to be close to Quercus

undulata variety Jamesii, “spreading into a low thicket,

with shallow, strongly tuberculated cups and very large,

edible acorns.” Of Quercus undulata, “several of the smaller

forms (Shin Oak) produce edible acorns,...very large in the

Sand Hills, which are eaten by Mexicans, raw or baked.

They afford excellent mast to hogs in the vicinity of settlements” (Havard 1885). Rydberg (1901) described Quercus

havardii from one of Havard’s collections. Long before the

Mexicans, shin-oak acorns were used by the Querecho,

Maljamar, and Ochoa phases of the Jornada Mogollon

culture, and more recently by the Apaches (Beckett 1976).

Older ranchers remember when the shinnery was dominated by tallgrass. Although oaks were inconspicuous, we

deduce from the ages of today’s oaks that they were

present (Osborn 1942). Probably the invasion and increase

reported for shin-oak is the result of the changing appearance of the shinnery due to overgrazing and disappearance of tallgrasses and perhaps to an increase in the height

of shin-oak due to fire suppression.

Geographical Distribution

Sand shinnery occurs on sandy soils in western Oklahoma, northern and western Texas, and southeastern

New Mexico (figure 1). Scattered populations are also in

southern Utah and adjacent Arizona (Welsh et al. 1993).

Disagreements about acreage of the sand shinnery are
2 USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998

CO KS

OK

NM TX

Figure 1. Distribution of sand shinnery in New Mexico, Texas,

and Oklahoma. There are also small outlying stands in these

states and in Arizona and Utah. The total distribution area of

sand shinnery is estimated at 5 to 7 M acres.

striking. Gribble (1981) and McIlvain and Armstrong (1959)

think that there are about 15 M acres; McArthur and Ott

(1996) estimate 6,400,000 acres. Other estimates are made

by Allred (1949), Deering and Pettit (1972), and Scifres

(1972). Duck and Fletcher (1944) estimate 750,000 acres in

Oklahoma. The Texas brush inventory of 1982 included

2.5 M acres of shinnery in that state, 920,000 of them with

an oak canopy less than 11% (USDA SCS 1987). Garrison

and McDaniel (1982) estimate 2,640,000 acres in New

Mexico. Considering all sources we estimate that historically there were about 1 M acres in Oklahoma, 3.5 M in

Texas, and 1.5 M in New Mexico; about one-third of this

acreage in each state had a light or scattered oak canopy.

If the exceptional figures from Duck and Fletcher, the

Texas brush inventory, and Garrison and McDaniel are

correct, our estimates may be high in Oklahoma and Texas

and low for New Mexico. These acreages comprise about

2% of the area of each of the 3 states. Of these areas, more

than 100,000 acres in Oklahoma, almost 1 M in Texas, and

an unknown, smaller acreage in New Mexico have been

converted to cropland or grassland (Deering and Pettit

1972). Also, 100,000 acres in New Mexico have been treated

with herbicide in the past 15 yrs and it is too soon to tell

how many of these acres will come back as shinnery.

Recent advances in the ability to distinguish shinnery by

remote sensing will enable more accurate estimates of its

distribution (Everitt et al. 1993, Middleton et al. 1987).

The shinnery has existed for thousands of years (Beckett

1976, Hafsten 1961). It was believed that the shinnery in

New Mexico had expanded considerably in historical

times (New Mexico Inter-Agency Range Committee 1970,

York and Dick-Peddie 1969). However, mid nineteenthcentury survey records and present-day boundaries of the

shinnery are remarkably similar (Gross and Dick-Peddie

1979, W. A. Dick-Peddie personal communication 1988).

The shinnery range has not increased perceptibly in recent

years (McIlvain 1954); in fact, it has decreased, due to oakcontrol efforts and conversion to cultivation. Dhillion et

al. (1994) consider the community highly threatened in

Texas, as do Bailey and Painter (1994) for shinnery habitat

in New Mexico.

Extension of a mid and tallgrass prairie southwestward

through shortgrass plains into the northern Chihuahuan

Desert has been explained by Weaver and Clements (1938)

as survival of a relict of wetter times. Deep sands developed in certain topographic positions during the postPleistocene drying of the southern Great Plains (Huffington

and Albritton 1941). Weaver and Clements regard the

compensatory influence of sand as responsible for the

survival of the shinnery, which they call a postclimax

community.

Plant Composition

Sand shinnery is always codominated by shrubs and

grasses; usually there is also a fairly rich mix of forbs and

nondominant grasses. See the appendix for species from 3

areas and scientific names.

Shin-oak occurs as the dominant shrub in shinnery or

codominates with sand sagebrush. Broom snakeweed, a

half-shrub, is also widespread in the shinnery, as are

mesquite and the less common lotebush, netleaf hackberry, and soapberry (chinaberry). In the south, additional shrubs of the shinnery are catclaw acacia, catclaw

mimosa, Southwest rabbitbrush, javelina bush, and vine

ephedra (Holland 1994, Martin 1990, Sullivan 1980). In the

Texas Panhandle and Oklahoma sand plum, Oklahoma

plum, prairie baccharis, wolfberry, and skunkbush occur

in shinnery, while post oak and eastern red cedar may

occur on the eastern fringe (Sullivan 1980, Wiedeman

1960, Boyd 1997 unpublished data). On sands with shallow water sources (for instance, the dunes of Chaves
USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998 3

County, New Mexico) tree poplars and shrub willows

occur with shin-oak. On sands near flowing water

Goodding’s willow and saltcedar may be interspersed

with shin-oak. Succulents in sand shinnery include yuccas, pricklypears, and, in the south, barrel cacti (Martin

1990, Smith 1971, Sullivan 1980).

The prominence and composition of grasses in shinnery are affected by grazing (Holland 1994, Sullivan 1980,

Wolfe1978). The appearance of ungrazed shinnery may be

that of mixed tall and midgrass prairie in which grasses

hide the oak shrubs (Brown 1982, Duck and Fletcher 1944),

but some mid nineteenth-century descriptions (previously quoted) differ with this description.

Grasses and sedges of the shinnery have been described by many authors (Deering 1972, Dickerson 1985,

Jones 1963, Martin 1990, Sullivan 1980, Test 1972).

Tallgrasses throughout the shinnery, especially on dunes

and other deep sands, include sand bluestem, big bluestem,

switchgrass, and giant dropseed. Havard panicum is common in the south; giant sandreed and Indiangrass are

common east of New Mexico. Of midgrasses, little bluestem

and sand dropseed are common throughout the shinnery

and, over considerable areas, are the most prominent

grasses. Other widespread midgrasses include spike and

mesa dropseeds, fall witchgrass, sand paspalum, plains

bristlegrass, side-oats grama, cane bluestem, and red and

sand lovegrasses. Short grasses include blue, black, and

hairy gramas, sandbur, windmillgrasses, false

buffalograss, and three-awns. Schweinitz and one-flower

flatsedges dominate the sedge component.

Dominant forbs of the shinnery include annual buckwheat, annual sunflower, and western ragweed (Pettit

1994, Sullivan 1980, Boyd 1997 unpublished data). Other

common, widespread forbs are the composites prairie

coneflower, camphorweed, false dandelion, gaillardias,

gray greenthread, yellow woollywhite, false boneset,

palafoxia, greeneyes, cowpen daisy, fleabanes, and woolly

paperflower; the legumes woolly dalea, sensitive brier,

rushpeas, and prairieclovers; spurge family members

queen’s-delight, Texas croton, sand reverchonia, and several spurges, and members of other families including

erect dayflower, prairie spiderwort, tumble ringwing,

snakecotton, James nailwort, snowballs (sandverbena),

linear-leaved four-oclock, carpetweeds, portulacas,

spectaclepod, flaxes, globemallows, scarlet and woolly

gauras, evening primroses, milkweeds, James cryptantha,

bindweed heliotrope, woolly plantain, blue sage, and mat

bluets (Martin 1990, Sullivan 1980).

Ecological Classification

Sullivan (1980) and Pettit and Sullivan (1981) recognized 3 community types within the shinnery in 6 Texas

counties, from Bailey County in the north through Winkler

County in the south. Type A is on actively moving sand

dunes in Winkler and Andrews counties and comprises 3

mapping units: 1) shin-oak/Havard panicum-giant

sandreed, with canopy cover usually less than 5%; 2) shinoak/Havard panicum-giant dropseed, with canopy cover

from 10% to 50%; and 3) shin-oak/giant dropseed-Havard

panicum on more stable sand, with plant cover from 50%

to 80%. Type B, in all 6 counties, is on sands with slopes up

to 30% and with higher percentages of silt, clay, and

organic matter in the soil surface layer than in type A. It

comprises 2 mapping units: 1) shin-oak/sand dropseedgiant dropseed (figure 2) and 2) shin-oak/giant dropseed.

The oak is most vigorous in the latter unit, sometimes

almost eliminating associated plants due to the effects of

shading and moisture competition. Type C, in all 6 counties, is characterized by honey mesquite. The amount of

honey mesquite varied in the 3 mapping units from an

average of 6 to 65 mesquite plants per acre; the greater

number was on finer-textured soils.

Figure 2. Giant dropseed dominates most of the southernmost

sand shinnery communities.
4 USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998

Working in Cochran and Yoakum counties, Texas, Roebuck (1982) distinguished 2

subtypes within Sullivan’s type B: 1) oak/

grass, with an almost continuous cover of shinoak to 30 inches tall, with threeawns and

sand dropseed and seasonal abundance of

false-buffalograss and purple sandgrass;

2) sage/oak, similar to oak/grass but with more

sandsage and slightly more grass and forbs.

Martin (1990) classified shinnery vegetation of 4 southeastern New Mexico counties.

He recognized 8 distinct communities and 3

outlier communities of limited area. Overall,

plant community composition was relatively

simple. Shin-oak, yucca, purple three-awn,

fall witchgrass, and sand dropseed were in

80% or more of Martin’s 149 10-by-20 m plots.

An additional 15 species were present in 20%

or more of the plots. Major community types

included: 1) active duneland, 2) mesquite/

shin-oak/hairy grama-purple three-awn,

3) mesquite/shin-oak/purple three-awnsand dropseed, 4) shin-oak/purple three-awnfall witchgrass (most common and widespread, in 65 plots), 5) shin-oak/little

bluestem-sand bluestem (figure 3), 6) shinoak/snakeweed/little bluestem-purple threeawn, 7) shin-oak/purple three-awn-sand

dropseed, and 8) shin-oak/purple three-awnsand bluestem.

Ahlborn (1980), working in southern

Roosevelt and northern Lea counties, New

Mexico, delineated 3 subdivisions of the

shinnery: 1) shin-oak/sand bluestem-little

bluestem, in which bluestems accounted

for nearly 40% of all grasses, which he regarded as in good to excellent range condition; 2) shin-oak/midgrass that largely

lacked bluestems and had a high abundance

of gramas, dropseeds, and three-awns,

which he regarded as in poor to fair range

condition having lost much of its climax

vegetation to heavy grazing by cattle, and 3) sandhills,

dominated by shin-oak with few grasses, with range

condition fair to good (figure 4).

At least 39 county soil surveys by the Soil Conservation

Service (SCS; now the Natural Resources Conservation

Service) treat the sand shinnery. They describe the supposed potential natural vegetation of each range site (soil

types that support similar plant communities). The SCS

descriptions discuss soils and vegetation dynamics, addressed in sections below. However, 9 Oklahoma surveys

(for instance Burgess et al. 1963, Ford et al. 1980, Nance et

al. 1963) provide the only classification of shinnery for that

state. They differentiate 2 range sites, deep sand and deep

sand savanna, both dominated by shin-oak and mid and

tallgrasses including sand and little bluestem, switchgrass, Indiangrass, and sand lovegrass. The main difference between these 2 sites is the higher clay content of the

subsoil on deep sand savanna that increases water-holding capacity and promotes better water relations in dry

years.

Shin-oak and Its Hybrids

Quercus havardii, a member of the white oak group, is

the defining plant of the shinnery. In its genetically pure

Figure 4. On dune sands, shin-oak often grows in almost pure stands.

Figure 3. Little bluestem dominates northern sand shinnery communities,

but in this scene most grasses have been removed by livestock grazing.
USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998 5

form, shin-oak is a low shrub with up to 100 or

more short aerial shoots from a massive underground stem system and a deep root system

(figure 5; Pettit 1986, USDI BLM 1979). The

underground stems are commonly 1 to 4

inches in diameter, some “... are as big in

diameter as a man’s thigh; these may occur on

plants with above-ground portions no more

than 2 ft tall and with individual stems not

thicker than a pencil” (McIlvain 1954). The

underground stems are mostly horizontal (figure 6) and are mostly within 1 or 2 ft from the

surface (Bóo and Pettit 1975, Galbraith 1983,

Wiedeman 1960). Sears (1982, Sears et al.

1986a) found 73% by weight of the “roots” in

the top 30 cm of soil. Where shifting dunes

have built up and caused additional layers of

stems to be added, the stem system may be 30 ft

or more deep (Peterson personal observations).

In the Texas Panhandle, Sears et al. (1986a)

found growing-season above-ground biomass

in shinnery to be 2,004 kg/ha, with oak comprising 1,821 kg/ha and herbaceous plants

183 kg/ha. They reported below-ground

growing-season biomass as 19,841 kg/ha for

oak and 1,394 kg/ha for herbaceous plants.

Pettit (1986) found the ratio of underground

to above-ground tissues in shin-oak to be 10:1

to 16:1, perhaps greater than for any other

American shrub. From level ground the vertical root system penetrates 15 to 20 ft below

the surface (McIlvain 1954). The above-ground

shoots, mostly 1 to 2.5 ft tall (up to twice as tall

in Oklahoma), may grow densely; Jones (1982)

found 30 m-2 in Cochran County, Texas, and

Zhang (1996) measured from 32 to 75 m-2 in

Yoakum County, Texas. Pettit (1986) found

11 to 15 yrs to be a usual lifespan for aboveground shoots though Muller (1951a) found

no shoots older than 11 yrs and Wiedeman

(1960) found that the smaller shrubs attain

ages of only 1 to 5 yrs or 10 to 12 yrs in drier

regions.

The underground stems commonly spread to form

plants 10 to 50 ft or more in diameter (Muller 1951a). Using

electrophoresis of enzymes to distinguish individuals,

Mayes (1994) studied the extent of single plants in a 40,000

m2 plot in Yoakum County, Texas. He found 56 clones in

the plot, but a few large individuals dominated. He estimated the largest clone at 7,000 m2

, equivalent to a square

84 m or 274 ft on a side, about 1.7 acres. “Age has no

significance to an individual of this species except that it

offers greater opportunity to spread and multiply by

fracture. Senescence is limited to individual areal shoots;

the entire clone is characterized by continuous rejuvenation” (Muller 1951a). Muller’s suggestion of multiplication by fracture for these shrubs remains unconfirmed.

The few excavations, for instance by Bóo and Pettit (1975),

suggest that the underground system remains intact.

Mayes’ (1994) data on this point are equivocal; his sampling technique found some evidence for fractured individuals. In species where fracturing occurs so that the

original plant (genet) produces many individuals (ramets)

vegetatively, single clones are reported to cover up to 81

hectares (205 acres) and to achieve ages over 13,000 yrs

(Cook 1985) and maybe 43,000 yrs (Anonymous 1997);

maximal extent and age of shin-oak are unknown.

Figure 5. Shifting sands have exposed a small shin-oak (Quercus havardii).

Figure 6. These exposed horizontal shin-oak stems measure to 8 inches in

diameter.
6 USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998

No measurements are available for normal lateral

growth rates of the oaks. McIlvain (1954) reports, “Old

fields cultivated around 1900 and then abandoned after a

few years of cultivation have often returned to solid

stands of grass even though dense stands of shinnery oak

surround the fields on all sides;” that is, there was no

expansion of oaks adjacent to the fields. Exceptionally,

from the edge of abandoned fields, oaks spread up to 30 ft

in 50 yrs (Sikes and Pettit 1980).

Quercus havardii hybridizes with other oak species,

giving rise to distinct growth forms that are important in

shinnery ecology (Muller 1952, Wiedeman and Penfound

1960). Small populations of Q. havardii var. tuckeri in

southeastern Utah and adjacent Arizona (Tucker 1961,

Welsh et al. 1993) are similar to larger populations of Q.

havardii in morphology and ecology; Q. X eastwoodiae is

thought to be its hybrid with Q. gambelii (Welsh et al. 1993).

On rocky slopes at the breaks of the Texas Plains, Q.

havardii hybridizes with the scrub oak Q. mohriana (Muller

1952). The resulting low shrubs have characteristics intermediate between the parents except that leaves are larger

than those of either parent species. The parent species are

ecologically separated; Q. havardii on deep sands and Q.

mohriana on exposed limestone (Muller 1952). According

to Muller, very dilute hybrids of Q. havardii with a few

morphological characters of Q. mohriana occasionally occur on sand and the reverse on limestone, but the markedly intermediate hybrids are limited to intermediate

habitat and the vicinity of both parents (Muller 1951b).

Very different is the hybrid of Q. havardii with post oak,

Q. stellata, whose distribution overlaps Q. havardii in western Oklahoma and slightly in the adjacent Texas Panhandle (Muller 1951b, 1952, Correll and Johnston 1970).

The hybrids are tall shrubs or small trees up to 20 or more

feet high (figure 7); their fruits vary from post oak’s small

size to the twice-as-large acorns of shin-oak. Post oak, like

shin-oak, can grow on sandy soils. Their hybrids are

abundant in Oklahoma and the eastern Texas Panhandle,

common in northwest Texas, and rare to occasional in

New Mexico and the Texas counties near its southeastern

corner, far from the post oak parent (Bruner 1931, Muller

1952, Pettit and Kauffman 1978). Great distance from one

parental species might indicate that it is the pollen parent,

but Muller (1952) thinks it unlikely that pollen was transported 150 mi against prevailing winds. Instead, he speculates that post oak ranged west to New Mexico in wetter

times and that the hybrids survive from then. The hybrids

share the rhizomatous habit of shin-oak; a “motte” of

these taller plants amid the low-growing shin-oak comprises a single plant commonly with 50 to 150 stems (Pettit

and Kauffman 1978, Wiedeman and Penfound 1960).

Pettit and Kauffman (1978) studied a motte in the Texas

Panhandle of 89 stems of varied ages, the oldest dating

from around 1900. They judged that motte to be decadent,

with ring growth less than .2 mm/yr and a “heart rot like”

condition in stems more than 50 yrs old. They also observed breakup of the horizontal underground system

between upright stems; Wiedeman and Penfound (1960)

made the same observation in tall mottes. As for shin-oak,

no young hybrids have been reported. These taller, hybrid

mottes in areas subjected to periodic fire may be due to

chance protection and reductions in understory fine fuel

as tree height increases. These

reductions in fine fuel would decrease the chance of a surface fire

spreading through the motte

(Wiedeman and Penfound 1960).

Soils

Soils of the shinnery are

Entisols or Alfisols according to

Pettit (1994); Garrison et al. (1977)

add Aridsols and “for most of the

shinnery ecosystem” Mollisols.

Huffington and Albritton (1941),

Lotspeich and Coover (1962), and

Lotspeich and Everhart (1962)

discuss the Pleistocene and postPleistocene formation of these

soils. The SCS county soil surveys list and describe more than

50 soil types that support shinnery (Pettit 1986). Nearly all are

Figure 7. A 14-ft shin-oak hybrid stands among 1- to 2-ft shin-oaks. sandy, often fine sands, some-
USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998 7

times with layers of sandy loams or loamy sands. Typical

are the Brownfield soils, which occur in all 3 sand shinnery

states: the surface horizon of 92% sand, which is yellowish-red to reddish-yellow, is 10 to 40 inches thick and is

highly permeable; under it lies a reddish, noncalcareous

sandy-clay-loam subsoil that is up to 63 inches deep, up to

40% clay, and only moderately permeable (Dittemore and

Hyde 1964, Moldenhauer et al. 1958, Pettit and Deering

1974, Zobeck et al. 1989). Other shinnery soils overlie a

hard caliche layer (calcium carbonate accumulation), and

effects of lime extend through the subsoil. For instance,

the Faskin Series, with a sticky subsoil from 14 to 56 inches

deep, has soft masses of calcium carbonate (Lenfesty

1983). Other soils are sandy throughout, lacking clay

layers. As the soil clay content increases the oak cover

decreases (Pettit 1986). According to Pettit (1978b) soils

under shin-oak have distinct, bleached lower A horizons

due to the acid content of its leaves.

One report from the Llano Estacado of west Texas

(Lotspeich and Everhart 1962) mentions shinnery on a

rocky loam not described as sand. These oaks might be the

Quercus havardii-Q. mohriana hybrids discussed above,

which occur in that area. Wiedeman and Penfound (1968)

report shinnery on a reddish fine sandy loam and on

sandstone in Oklahoma, but regard these as unusual

occurrences. Whitford et al. (1986) report shinnery on

sandy loams in Eddy County, New Mexico.

Calcium carbonate in shinnery soils increases with

depth. Wiedeman and Penfound (1960) concluded that

shin-oak does not occur regularly where there is much

calcium carbonate in the soil. Small (1975) thought that his

study did not support that conclusion, but in fact he did

find that the calcium carbonate equivalents of the surface

horizons were inversely related to the number of oak

stems per unit area. Sullivan (1980) found that when the

caliche layer was within a meter of the soil surface, shinoak was smaller or was replaced by mesquite. Similarly

Pettit (1978b) reported that when subsoils are rich in

carbonate or clay, sand sagebrush or mimosa dominates.

The permeability of shinnery soils is generally high and

water erosion very low. For instance, in Brownfield fine

sand, one of the widespread soil types, the A horizon (40

to 70 cm deep) is 95% sand and overlies a sandy clay loam

B horizon. Infiltration rates are rapid, up to 70 cm/hr, and

percolation is rapid until the B horizon is reached (Sears et

al. 1986a). Small (1975) found maximal densities of shinoak where the soil surface was sand, especially where the

sand layer was relatively thick, permitting water infiltration to greater depths. No oak grew on an adjacent sandy

loam site dominated by blue grama and underlain by

calcium carbonate layers at shallower depths than the

sandy sites. Shin-oak density was most negatively correlated with percentage clay, runoff, and water retention at

-1/10 and -1/3 bar.

Especially under dune sands, water may perch above

less permeable rock or soil layers, affording a source for

riparian-type species, including cottonwoods, willows,

and saltcedar (tamarisk), which can then grow with shinoak. Smith (1985) reports coyotes digging to water 30

inches deep in the Mescalero Sands of Chaves and Eddy

counties, New Mexico. In the Mescalero Sands, water

wells for domestic or livestock use average only 37 ft deep

(Smith 1985) where the depth to groundwater varies from

200 to 400 ft (Geohydrology Associates 1978). It is unknown whether shin-oaks on dunes are larger and more

vigorous than those on stable sites because their roots

reach the perched water or because of the relative lack of

competition from other plants on shifting sands.

Nutrient characteristics of soils supporting shinnery

have been examined by several authors. Sears et al. (1986b)

reported that total nitrogen concentration of the soils

supporting shinnery was relatively low at about .017%.

Highest concentrations were in the upper 15 cm of the

surface and in the B horizon. This bimodal stratification

was attributed to nitrogen leaching through the sandy

surface horizon. Secor et al. (1983) compared soils that

supported shin-oak and sandsage with soils that supported mesquite and javelina bush. They found nitrogen

(especially nitrate), phosphorus, calcium, and magnesium levels higher under mesquite and javelina bush.

Surface soils under mesquite and javelina bush contained

about twice as much organic matter as those under shinoak-sandsage. Small (1975) reported that shin-oak grew

on sandy sites that averaged only .67% organic matter in

the surface horizon; shin-oak grew best on sites with an

almost pure sand cover.

Near-surface soil pH in shinnery was measured by

Kyrouac (1988) at 6.9. In Small’s (1975) shinnery plot, pH

increased in the A horizon from 7 to 8.1 and in the C

horizon from 7.9 to 8.5; pH was only slightly higher on his

adjacent sandy loam plot. Wiedeman and Penfound (1960)

found lower pH values in soils of shinnery mottes than

under shin-oak shrubs, approximating the pH of forest

soil types even though rainfall was less than 25 inches/yr.

Sand shinnery soils are highly susceptible to wind

erosion (Pettit 1978a, SCS soil surveys). Moldenhauer et

al. (1958) reported that newly broken surfaces of Brownfield

soils would lose from 150 to 240 tons/acre/yr under a

sustained 38 mi/hr wind. Spring fine sands would lose

650 tons, and Tivoli fine sands, which form dunes in

shinnery, would lose 1,600 tons. Zobeck et al. (1989)

studied erosion in relation to vegetation cover, soil-particle

size, and soil nutrients. An early SCS treatment (Goodding

1938) says of the shinnery: “What this region would be

without the oak is hard to conjecture. The bushes are seldom

more than 2 ft high but the whole country is completely

covered and the sand stays put.” Similarly Moldenhauer

et al. (1958) regard shinnery vegetation as the most reliable protection for Tivoli fine sands (figure 8). Parks (1937)

calls shin-oak “the most valuable of all sand binders.”
8 USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998

Climate

The sand shinnery extends southwestward from a zone

of 25 inches average annual precipitation in western Oklahoma 400 mi to a zone of 12 inches precipitation in southeastern New Mexico (Canales 1968, McIlvain 1954). The

annual cycle of precipitation also differs between the

Oklahoma and New Mexico extremes. To the north, 3 to 4

inches of rain during each spring and summer month are

fairly reliable but late summer and fall droughts are

common (USDC NOAA 1995). To the south, April through

June are generally dry and July through September relatively wet; many grasses do not begin growth until July

(USDC NOAA 1996). Snowfall in the shinnery of western

Oklahoma averages 12 inches/yr (Burgess et al. 1963)

whereas in the southern shinnery snow averages less than

6 inches/yr and none falls in some winters (Chugg et al.

1971, Orton 1964). Table 1 reflects data from 3 disparate

climatic regimes of the shinnery.

Because water availability is a limiting factor for plant

growth in this community (Pettit 1978a, Sikes and Pettit

1980, Galbraith 1983), climatic differences correlate with

profound differences in shinnery vegetation as previously discussed. For additional discussion of shinnery

climate see Allred and Mitchell (1955), Campbell (1936),

Lotspeich and Everhart (1962), Roebuck (1982), Sullivan

(1980), USDOE (1980), USDI BLM (1977), Wiedeman (1960),

and SCS soil surveys. Drought is discussed in a separate

section. Paleoclimates (under which the shinnery formed)

are discussed by Hafsten (1961), Lotspeich and Everhart

(1962), Reeves (1965), and USDI BLM (1980).

Phenology

Plant activity in the shinnery

occurs from about February

through December; winter annuals can begin growth even earlier

in the south, although there can

be freezing weather into April.

The buds of shin-oak swell in

early to late March (Pettit 1986,

USDI BLM 1979), leaves open

mainly in April and May (Bóo

and Pettit 1975, Pettit 1975), and

flowering occurs in April and

May (Rowell 1967). Early-season

grasses are few in the southern

shinnery, but those few, such as

switchgrass and purple threeawn, begin growth in March or

April. In western Oklahoma and

adjacent Texas, most grass species begin growth in early May

while forb growth begins in early

April.

In the northern shinnery, grass and forb growth continues from spring through summer. Most grasses of the

southern shinnery begin growth following summer rains,

which generally begin in early July. Galbraith (1983) followed perennial grass development in Cochran County

for 2 yrs. On June 1 he found greenup, on July 15 vegetative growth, on September 1 flowering, and on October 15

senescence. Most forb growth in the south also follows

summer rains, especially that of annual forbs, which may

be absent or abundant depending on the timing of rainfall

(Pettit 1979). Growth of grasses and forbs in western

Figure 8. Shin-oak anchors sands.

Table 1. Climatic statistics.

Roger Mills Co. Yoakum Co. Eddy Co.

Variable OK TX NM

Average annual

precipitation (cm) 65.1 40.4 31.6

Average date of

last frost 2 April 13 April 31 March

Average date of

first frost 28 Oct. 1 Nov. 7 Nov.

Average Jan.

 temperature (o

C) 2.7 5.6 6.7

Average July

temperature (oC) 28.2 26.4 27.5

Sources: Burgess et al. 1963, Orton 1964, which uses some

data from adjacent Gaines County and Yoakum Co., and

Chugg et al. 1971.
USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998 9

Oklahoma and adjacent Texas is not as closely tied to

precipitation events due to a more regular cycle of precipitation through the year. Shin-oak acorns mature in July

(personal observations). Ninety-five percent of growth of

fine roots of shin-oak occurs from July through September, with small amounts detected in fall and winter (Zhang

1996).

Plants remain active in the shinnery well into autumn.

Especially in the south, some grasses do not flower until

September or October and remain green (depending on

moisture and freezing weather) into November or December, although an October frost can end growth (personal

observations). Oak leaves normally fall in early November (Bóo and Pettit 1975).

Autoecology of Shin-oak

Pettit (1978a) states that shin-oak does not use more

water than the grasses and forbs in the community; however, it begins spring growth about 2 weeks earlier than

competing vegetation (Jones and Pettit 1980). Shin-oak

rhizomes absorb much water during wet periods, with up

to 50% of their weight being water, This is particularly

important during dry springs because grasses and forbs

cannot mine or store water (Pettit 1986).

Following a drought, shin-oak may not leaf out in the

spring, or may lose its leaves. In 1980, shin-oak shed 36% of

its leaves in summer but leafout in 1981 was normal (Jones

and Pettit 1980, 1984). Having failed to leaf out in spring or

having lost its leaves in summer, oak can leaf out later in the

growing season after water becomes available. Leaf structure

varies over the distribution area

of shin-oak, with adaptations to

resist dryness, such as thicker leaf

epidermis bearing more hairs, in

New Mexico and adjacent Texas

that are lacking in Oklahoma

(Canales 1968).

Bóo and Pettit (1975) studied

the annual cycle of nonstructural

carbohydrates in the rhizomes

and roots of shin-oak. Reserves

were gradually depleted through

the dormant season, November

to April, until a low of 6.5% to 7%

was reached in early May. Carbohydrates then began to accumulate when new leaves were 1/

3 to 1/2 full-size and reached concentrations of 17% to 25% from

early August to early November.

Pettit and Hungerford (1973) followed the total available carbohydrate content of shin-oak roots (underground stems)

for 14 mos. Concentrations ranged from a low of 6% to 7%

in early May, when leaves were 1/2 to 2/3 full-size, to a

high of 13% to 15% in January, June, and August.

Plumb (1984) studied variation in shin-oak’s crude

protein and phosphorus through 2 growing seasons. Protein decreased from about 9% to 5% from June to November while phosphorus trends, although less clear, were

generally upward over the same period.

Quercus havardii rarely, if ever, reproduces through the

agency of acorns. “...I saw not a single case of germination

of acorns” (Wiedeman 1960, Dhillion et al. 1994). Pettit

(1977) states that, “It is rare to find a young sand shin-oak

plant in the field which had originated from an acorn”

except seedlings underground in woodrat nests. Pettit

and E. H. McIlvain said that they never found an aboveground seedling in the field (personal communications

1988). Many clonal plants put little energy into seed production and produce seedlings rarely (Cook 1985). Shin-oak’s

failure to produce seedlings may be from unsuitable moisture conditions rather than from a diversion of reproductive energy. Shin-oak stems are often weighed down to the

ground with the large crop of heavy acorns (Palmer 1934).

Under laboratory conditions shin-oak acorns (figure 9)

are viable and quickly produce roots that may extend 12

inches before leaves emerge (Pettit 1977, 1986). Sikes and

Pettit (1980) measured root growth of seedlings at .05 cm/

day in dry soil to .24 cm/day in moist soil. Acorn production may be inhibited by inadequate spring rains or by a

freeze after flowering (USDOE 1980). Pettit (1986) estimates that acorn crops occur in 3 out of 10 yrs on average;

Figure 9. A dime shows the size of shin-oak acorns.
10 USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998

in New Mexico the Bureau of Land Management (USDI

BLM 1979) estimates that crops occur from 1 yr in 10 to 1

yr in 5. In New Mexico, 1977-1997, we found heavy local

crops occurring somewhere every year but in a specific

location crops occurred not more than 2 yrs in 5.

Reproduction appears to be almost completely by rhizomes. If destruction of the above ground parts of shrubs

through the 18- to 20-yr age group occurs, the buds of the

rhizomes sprout and within 1 or 2 mos new growth

appears above the ground. The growth during the first

year proceeds at a rapid pace until stems one to 2 ft all are

produced (Wiedeman 1960, Wiedeman and Penfound

1960). Most new shoots arise from buds covered with no

more than 3 inches of sand. If sand is eroded away from

the stem, new shoots can develop from more deeply

located buds (Pettit 1977).

Zhang (1996) found 2 kinds of ectomycorrhizae on

shin-oak roots, 1) a white type was bifurcate and had a

smooth gray to white surface; 2) a black type was short,

often club-shaped, with radiating black hyphae, formed

by the fungus Cenococcum geophilum, a highly droughtresistant species. Levels of soil organic matter had the

largest direct effect on ectomycorrhizal colonization. The

effect of total nitrogen on ectomycorrhizal colonization

was significant and negative.

Shin-oak may also have direct allelopathic effects on

seedlings of other species. Matizha and Dahl (1991) found

that an extract from oak leaves depressed shoot growth of

weeping lovegrass by 92%. Pettit and Harbert (1974)

found that 5% tannic acid drastically reduced germination of weeping lovegrass and sand dropseed.

Several insects attack shin-oak but reportedly do not

cause mortality. A buckmoth larva, Hemileuca slosseri,

completely defoliates areas of a few acres (Peigler 1989,

Pettit 1986, Pettit and Sullivan 1981, Stone and Smith 1990,

Turrentine 1971, Wangberg 1983). The lined bird grasshopper (Schistocerca alutacea ssp.lineata) partially defoliates oaks over large areas (Dodson 1987, Peterson unpublished data). Cynipid gall-wasps are everywhere abundant on shin-oak stems and leaves (figure 10; Pettit 1977);

Tucker (1970) lists Atrusca bella, Callirhytis frequens, C.

juvenca, Disholcaspis rubens, D. spissa, and Neuroteras

howertoni and Zimmerman (personal comm. 1997) adds

Xystoteras sp. and Xanthoteras eburneum, the latter studied

by Dodson (1987). Large roots are infested by long-horned

beetle larvae (Cerambycidae) that leave only a small part

of a root intact, yet do not limit rhizome and shoot growth

(Pettit 1977). Most acorns become infested by weevils

(Curculio sp.), often 2 or more larvae per fruit, but these do

not always destroy acorn viability (Pettit 1973, 1977,

Wiedeman 1960). Acorns decompose rapidly after falling

(Mayes 1994) and few survive after January 1 (Hanson

1957).

A number of birds and mammals affect shin-oak by

consumption of leaves, roots, acorns, buds, and catkins.

These species are reviewed under the section Wildlife

Species.

Vegetation Dynamics and Water Relations

Sand shinnery is an unusually stable community; individuals of its principal species, shin-oak, grow slowly and

live for hundreds of years, and the species has produced

a continuous pollen profile for at least 3000 yrs (Gross and

Dick Peddie 1979, Hafsten 1961). Drought, fire, livestock

grazing, and brush treatment do modify or transform the

community. Each of these disturbance factors is discussed

below.

Although shin-oak may constitute 80% to 90% of plant

mass or ground cover in shinnery (Biondini et al. 1986,

Dhillion et al. 1994, Pettit 1994, Plumb 1984), in SCS

surveys shin-oak is usually said to make up 5% to 25% of

the plant composition or biomass of the pre-settlement

plant community (Conner et al. 1974, Hodson et al. 1980).

Correspondence with SCS offices in the 3 shinnery states

failed to identify data that support these low percentages.

All of the soil surveys describe shin-oak as an “increaser”

Figure 10. Galls of cyniped wasps are abundant on shin-oak

and provide an important food source for prairie-chickens.
USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998 11

under livestock grazing. Tallgrasses and most midgrasses

decrease under grazing. When these studies were prepared (1963 to1983) “increaser” was a species that increased in percentage composition of the community,

although it could be decreasing in absolute amount. So

defined, shin-oak is an increaser under grazing pressure.

However, some of the studies (e.g. Lenfesty 1983) also

claim that shin-oak increases in absolute amount under

grazing pressure, which is the current definition of “increaser” (Society for Range Management 1995). The SCS

in 3 shinnery states has been unable to support this claim

with data. Although it seems possible that shin-oak would

increase when without grass competition (Muller 1951a),

this assumes that shin-oak is limited by competition with

grasses, which is not supported by data. It is clear that

under some circumstances, perhaps all circumstances, the

abundance of shin-oak under grazing pressure either

remains constant (Dickerson 1985, Holland 1994) or decreases along with grasses. Davis et al. (1979) found that,

unlike instances where shrubs have invaded herbaceous

communities, oak and grass densities are positively correlated; the more oak, the more grass (table 2).

The SCS and others claim that shin-oak invades overgrazed rangeland (Herbel 1979, Herndon 1981, Lenfesty

1983, York and Dick-Peddie 1969). BLM stated that shinoak increases in absolute amount under grazing pressure

and that former grassland was invaded and taken over by

shin-oak (USDI BLM 1979). The idea that shin-oak was not

part of the historic plant community has influenced BLM’s

decision to treat 100,000 acres in New Mexico with herbicides (USDI BLM 1979, BLM Roswell District Manager

letter to the Sierra Club September 24, 1991). However,

stable existence of shinnery for hundreds of years (Beckett

1976, Gross and Dick-Peddie 1979) and lack of spread of

shin-oak by seeds negate claims that shinnery have increased in historical time.

Several SCS writers believe that mottes are the historic

form of shin-oak, and that the low-growing shrubs surrounding them have spread from the mottes as the grass

cover disappears under heavy grazing (e.g. Newman

1964). Apparently this was also the view of Osborn (1942).

This hypothesis seems unreasonable because the taller

oak mottes are single clones (Pettit and Kauffman 1978)

and are hybrids distinguishable by botanical characteristics from the lower-growing shin-oak surrounding them

(Muller 1952, Pettit 1986).

Water is a limiting factor for plant growth across the

range of shinnery (Galbraith 1983, Pettit 1978a, Sikes and

Pettit 1980). Shin-oak absorbs water efficiently when available (Pettit 1986), “even while the plant is still in winter

dormancy” (Sullivan 1980). “Grasses in a vigorous state

extract more deep soil water than sand-shinnery oak”

(Jones and Pettit 1980), but when water is in short supply,

shin-oak biomass may decline less than that of competing

herbaceous vegetation. When water is available tallgrasses

may overtop the oak and perhaps by shading the oak they

achieve a balance. In southern shinnery, tallgrasses may

grow twice as high as the oak. Muller (1951a) observed

that, “The clones of Quercus Havardi are not so luxuriant

and dense where they are subject to heavy competition by

grasses.” Similarly Frary (1957) opined that “shinoak

seems to alter its growth form when under heavy competition from other vegetation. The vigor of the plant seems

to be lower and fruiting is practically nonexistent even

when other plants are producing acorns.” Moldenhauer et

al. (1958) believed that in the original vegetation, “The

grasses were dominant and kept the Shinnery and Yucca

from spreading.”

Dhillion et al. (1994) studied the influence of animal

disturbances (rodent and lagomorph burrows and ant

hills) on the structure of shinnery communities. They

found that density of herbaceous seedlings was not influenced by oak cover. However, the density of herbaceous

seedlings was positively correlated with increasing bare

ground and with the number of animal disturbance types.

The number of seedlings was correlated, though not significantly, with the absolute number of animal disturbances. Undisturbed areas were covered in a mat of oakleaf litter and offered little opportunity for plant establishment. In fact, over 70% of the herbaceous seedling density

was associated with animal-disturbed sites. Shin-oak

ground cover was not correlated with the number of

disturbances.

Holland (1994) working in the same area as Dhillion et

al. found that the density of herbs was much lower where

there was a dense oak cover. Holland explained this by

noting that Dhillion et al. were concerned with herbaceous seedlings whereas she dealt with mature herbaceous plants. Holland hypothesized that oak canopy cover

does not impede colonization by herbaceous seedlings

but has a negative effect on seedling recruitment and

density of adult herbaceous plants.

Pettit and Holder (1973), Pettit and Small (1974), and

Test (1972) manipulated vegetation composition in shinnery communities and examined soil-water use to a depth

of 150 cm by different classes of vegetation. Treatments

included: 1) remove shin-oak and leave herbaceous vegetation, 2) remove herbaceous vegetation and leave oak, 3)

remove all vegetation, and 4) control (leave all vegetaTable 2. Percentage of ground covered by plant type.

Plant type Most grass Medium Least grass

Oak 6 3 4

Grass 11 7 4
12 USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998

tion). From September 1972 through August 1973 results

of the 4 treatments were similar at depths from 15 cm to 45

cm, but from 90 cm to 150 cm the remove-all-vegetation

treatment stored 24.3 cm of water and the other 3 treatments about 17.5 cm of water (Pettit and Holder 1973).

Because the authors did not quantify evaporative water

losses from bare ground, it is difficult to assess water use

among herbs and oaks from these data.

Jones (1982) presents 28-mo records of precipitation

and water in the soil horizons A and B of shinnery in

Cochran County, Texas. He concluded from the position

of oak roots and soil-water responses that shin-oak concentrates its water absorption near the soil surface and at

the interface between A and B horizons. Test (1972) and

Galbraith (1983) found a concentration of fine oak roots

just above that interface, possibly in response to water

perched above the less permeable B horizon. Small (1975)

and Zhang (1996) also report the annual course of percentage soil moisture for shinnery areas in Texas. Biondini and

Pettit (1980) modified a soil water-flow model to predict

changes in soil water content through a growing season at

the 30 cm depth in shinnery from precipitation data. Data

from this study indicate that peak water concentration

occurs in August and a low-point concentration in May.

Shin-oak is an effective water-gatherer. In response to

precipitation, shin-oak produces from 41 to 113 g/m2

/yr

of fine roots (Zhang 1996). Bóo (1974) measured xylem

water potentials as approximately -25 bars with a pressure

bomb. Galbraith (1983) found shin-oak to be a highly

efficient water-user due in part to the oak’s maintaining

more negative leaf water potentials than associated plants.

Oak’s negative leaf water potential was between -25 to -32

bars at midday compared with -12 to -18 bars for grasses

and -12 to -25 bars for forbs. Galbraith found that oak

absorbed large quantities of water in the top 137 cm of soil

when water was available at low potentials, and to 198 cm

depth when water supplies became limiting. Other authors emphasize the shallowness of shin-oak’s underground system and suggest that grasses are more efficient

at harvesting water at greater depths than shin-oak (Jones

and Pettit 1981, Jones 1982). Galbraith (1983) emphasizes

the greater effectiveness of the oak’s absorption down to

2 m depth compared with associated plants.

Given the limited water supply, shinnery is a productive ecosystem. SCS surveys estimate that in untreated

shinnery up to 3,000 lbs of air-dry forage are produced per

acre per year in New Mexico (Lenfesty 1983), up to 3,500

in Texas (Crump and Williams 1975, Richardson et al.

1975), and up to 4,500 in Oklahoma (Nance et al. 1963).

Production is much lower on shifting sands and other

poor soils or where vegetation is in poor condition or

during low-rainfall years. In depauperate stands, up to

90% of production can be oak (Pettit 1994).

Garrison et al. (1977) recognized 4 productivity classes

in the shinnery. Class 1 produces an average of 2,000 lbs of

herbage per acre per year on Mollisols (Calciustolls plus

Haplustolls). Class 2 produces 1,500 lbs of herbage per

acre per year on Alfisols (Haplustolls). Class 3 produces

1000 lbs of herbage per acre per year on Aridisols

(Haplargids). Class 4 produces 500 lbs of herbage per acre

per year on Entisols (Ustipsamments) of stabilized sand

dunes.

Sullivan and Pettit (1977) found the most productive

shinnery in west Texas on younger sands, which are

deeper and lighter in color than older sands. These most

productive stands were also the most easily disturbed and

were often interspersed with blow-out areas. Dark red or

brown soils, characterized by a clay layer close to ground

level, support a stable but marginal shin-oak community.

Sullivan and Pettit (1977) found that in heavier soils shinoak and tallgrasses are replaced by mesquite and shortgrasses and midgrasses including sand dropseed.

Organic Matter and Decomposition

The organic-matter content of shinnery soils is low

even in the A horizon. Secor et al. (1983) found a mean

value of .36% under shin-oak in Eddy County, New Mexico,

less than half of what they measured under adjacent

vegetation dominated by mesquite and javelina bush. In

Roosevelt County, New Mexico, Keesee (1992) measured

organic-matter contents of .40% to .48% in the top 3.75 cm

of soil and .35% to .40% from 3.75 to 7.5 cm. In Yoakum

County, Texas, Small (1975) found a range of .20% to

1.39% (average .67%) in shinnery, which was not significantly less than in adjacent blue grama-buffalo grass

(average .73%). Sears (1982) and Sears et al. (1986a) measured soil organic matter in west Texas shinnery at 3,200

kg/ha.

Elkins and Whitford (1982) and Elkins et al. (1982)

measured annual weight losses of shin-oak leaves in Eddy

County, New Mexico, of 20% for unburied and 35% for

buried leaves. Working in Yoakum County, Texas, Zhang

(1996) reported that over a 3-mo period, fine shin-oak

roots lost 12 to 16% of their weight. Decomposition then

slowed, due to the exhaustion of soluble carbon, and after

15 mos, weight loss totaled 22% of initial weight.

Sears et al. (1986a) found that below-ground biomass

totaled 27,000 kg/ha, of which 22,500 kg/ha were roots,

94% of them oak roots including underground stems.

Comparing an untreated plot with plots in which oak had

been killed with herbicides 3 and 6 yrs previously, they

found that large (> 2 mm) oak-root biomass decreased 8%

in 3 yrs and 41% in 6 yrs. Small oak-root biomass was 19%

lower in the 3-yr plot and 29% in the 6-yr plot. Sears et al.

regarded these as slow rates of decomposition. These

authors measured carbon-nitrogen ratios for large oak

roots from 86:1 to 92:1 and stated that ratios greater than
USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998 13

20 favor immobilization of nitrogen and slow decomposition rates.

Nutrient Cycling

The soils of shinnery communities are critically low in

nitrogen and phosphorus (Pettit 1983b). In Eddy County,

New Mexico, Secor et al. (1983) measured .5 ppm of NO3

+ NH4 nitrogen in shinnery soils, compared with 5.2 ppm

under mesquite, and .1 ppm of phosphorus, compared

with .2 under mesquite. In Garza County, Texas, Matizha

and Dahl (1991) measured 3.4 ppm of available nitrogen,

3.1 ppm available phosphorus, and 93.8 ppm of available

potassium. In the same Brownfield fine sand in Yoakum

County, Texas, Sears et al. (1986b) measured a soil nitrogen concentration of .017% and Dittemore and Hyde

(1964) measured a range of .011% to .019% nearby.

Forage production can be increased with fertilizers.

Pettit and Deering (1974) working in Yoakum County

found that 60 kg/ha of nitrogen in the form of ammonium

sulfate or as ammonium-phosphate-sulfate increased total vegetative production by 51% to 54% including increased grass yields of 84% to 100%. Sulfur as well as

nitrogen was important; in this study they did not find

evidence that phosphorus was beneficial. Deering (1972)

reported that nitrogen applied at 30 lb/ac to a Yoakum

County shinnery community had no effect on shin-oak

stem density or grass canopy cover but did increase forb

cover canopy at the end of the growing season. Holland

(1994) found a strong association between oak height and

soil potassium, which lead her to suggest that oak growth

is limited by potassium.

In Cochran County, Texas, Biondini et al. (1986) measured the crude protein content of little bluestem and

purple three-awn at 6.3%, red lovegrass at 7.1%, and sand

dropseed at 8.6%. The first 3 species had phosphorus

contents of .08% and sand dropseed contents of .11%.

Pettit (1983b) reported the crude protein content of both

shin-oak leaves and forbs at 9% to 11% in June, but the

forbs contained only 6% by late summer. Phosphorus

levels in all forages ranged from .04% to .12%. Pettit

concluded that phosphorus must be provided as a supplement for livestock year-round. Villena-Rodriguez (1987)

found crude protein values for shin-oak leaves and current-years twig growth to range from 8.5% in late June to

6.9% in late August.

Zhang (1996) considered the shinnery ecosystem to be

nitrogen-limited. He stated that soil nitrogen dynamics

and microbial biomass are tightly coupled and provide for

efficient nitrogen retention. Microbial biomass increased

from late summer through late fall, decreased during the

winter, and continued at a minimal, steady level from

May through August in the 2 yrs sampled. Nitrogen

mineralization rates changed seasonally, with higher rates

from February through September; the year-long monthly

average was .75 mg N/g soil, of which 85% was nitrate

nitrogen. Nitrification rates were also highest during late

summer and declined through the fall as leaves became

senescent; monthly nitrification averaged .60 mg N/g

soil. Thus, N mineralization and nitrification rates were

inverse to microbial biomass and highest during the growing season. Zhang suggested that microbial biomass responds to carbon inputs as plants shed leaves and fine

roots during late fall and winter. Soil organic matter, total

nitrogen, and above-ground stem density had large, significant, positive, direct effects on microbial biomass...,

soil nitrogen mineralization and nitrification rates. Low

nitrogen availability is related to increased colonization of

oak roots by ectomycorrhizal fungi, which help the oak

obtain nitrogen. The addition of 50 g ammonium nitrate/

m2

 depressed mycorrhizal fungi but sharply increased

microbial biomass and nitrogen mineralization for up to 4

mos.

Kyrouac (1988) examined vesicular-arbuscular mycorrhizae (VAM) of little bluestem and purple three-awn

grasses in shinnery, formed by Scutellospora, Glomus, and

Acaulospora species. From 8% to 25% of grass plants were

infected. VAM root lengths in little bluestem increased

from 200 to 540 cm from April to August, then fell off to

400 cm by October. In purple three-awn, VAM root length

declined from 270 cm in April to 100 cm in August, then

rose to 190 cm in October. It is difficult to see how these

differing responses could relate to nutrient availability in

the same soil. McGinley et al. (1994) found that the importance of mycorrhizae to little bluestem varied in shinnery

depending on differences in nutrient availability, which

are caused by animal (mainly ant) disturbances.

McIlvain and Armstrong (1965) pointed out the importance of deep-rooted oaks in recirculation of deeply leached

soil nutrients. Important to nutrient cycling are the thousand species of insects collected in one shinnery area; the

biomass of termites alone equalled that of cattle grazing

the surface (USDOE 1980). Whitford et al. (1986) calculated that ants bring 84 g/m2

/yr of soil to the surface in

Eddy County shinnery.

In the shinnery ecosystem of Cochran and Yoakum

counties, Texas, Sears et al. (1986a, 1986b) determined that

where oak comprised 92% of the living above-ground

biomass, above-ground nitrogen (kg/ha) was distributed

as follows: live oak stems 4.4, dead oak stems 1.6, live oak

leaves 5.4, herbaceous shoots 2, and above-ground litter

31, for a total of 44.2 kg/ha. Nitrogen (kg/ha) associated

with below-ground biomass averaged: large oak roots

75.8, small oak roots 41.5, herbaceous roots 8.3, and litter

36.5, for a total of 167.1 kg/ha. Soil nitrogen averaged 405

kg/ha. About 5% of soil nitrogen was available for plant

use (Sears et al. 1983). Soil nitrogen was significantly

higher in July (507 kg/ha) than in April or December (337

and 370 kg/ha). Sears et al. also sampled plots where oak
14 USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998

had been killed with herbicide 3 and 6 yrs previously.

Above-ground nitrogen did not differ between control

and herbicide-treated plots; however, below-ground biomass-associated nitrogen decreased in the 6-yr plots. Total soil nitrogen was similar across treatments for the July

and December sampling periods, and was higher for the 6-

yr plots in April.

Working in the shinnery of Yoakum County, McGinley

et al. (1994) found the soils associated with nests of a

harvester ant, Pogonomyrmex barbatus, to be much richer in

NO3 + NH4 nitrogen than soils associated with nests of

Pheidole dentata or Crematogaster punctulata or soils not

associated with ant nests. At the same site Dhillion et al.

(1994) found disturbed soil associated with rabbit burrows to be richer in organic matter, total nitrogen, ammonium, nitrate, iron, and magnesium than off-mound soils.

Drought

Season-long, multiple-year, or droughts for periods as

long as a decade are common in shinnery. In this century,

the droughts of 1906-1911, 1921-1926, 1933-1936, 1946-

1948, and especially 1951-1956 have been the most serious

in the shinnery area; 1994 to mid-1996 was the most recent

drought. In the northern shinnery region, in any 10-yr

period expectation is for 1 or 2 yrs with less than 75% of the

average annual precipitation. In the southern shinnery

region, in any 10-yr period expectation is for 2 or 3 yrs with

less than 75% of the average annual precipitation (Campbell

1936).

The effects of drought are more severe on grasses than

on shrubs in the shinnery, since shrubs can store water and

carbohydrates (USDI BLM 1977). Even a mild drought is

apt to decrease grass production from 1200 to 1400 lbs/

ac/yr down to 400 to 600 lbs/ac/yr. This happened in

Garza County, Texas, shinnery in 1983 and 1984, when

precipitation totaled 8 and 6.9 inches rather than the

historical average of 13.9 inches (Mosley et al. 1985b).

Drought will have stronger negative effects on grass

production than on oak, given the oak’s extensive root

system, its ability to store water, and its strongly negative

leaf water potentials (Galbraith 1983). Observations in

New Mexico (Peterson unpublished data) at the end of the

1994 to 1996 drought support this speculation. Peterson

found that oak began sprouting vigorously after rains in

June 1996, but many grasses had died (or nearly so) and

were slowly replaced, mostly by annual herbs. However,

there are no quantitative studies that show a relative

increase of oak under drought.

 Forage removal by grasshoppers is considerable during drought (Coupland 1958). The 1994 to 1996 drought

was accompanied and followed by a grasshopper outbreak in the sand shinnery of west Texas and New Mexico.

At 17 points sampled in 1996 the predominant grasshopper was Schistocerca alutacea subspecies lineata, which

was eating oak predominantly or exclusively. Grasses

were not noticeably damaged by grasshoppers (Peterson

unpublished data).

Fire

In the twentieth century, wildfire has become less common in the western United States due to human control of

fire and to the loss of fine fuel due to livestock grazing (Box

1967). Shin-oak and associated vegetation have evolved

under the influence of fire. However, the pre-European

settlement fire regime for shin-oak is unknown. Shoop

and McIlvain (1964) speculated that shinnery in western

Oklahoma was historically burned on an annual basis by

Indians. In this same region, historical references to shinoak (Marcy 1854) indicated that it was a low-growing

shrub, generally less than 18 inches high. Possibly these

plants were under a reliable and strong pyritic influence

because shin-oak plants in this region today may approach or exceed 40 inches in height.

Knowledge about the response of shinnery to fire is

lacking, relative to that for other vegetation types of the

Great Plains. However, vegetation trends following singleevent, planned spring burns in Woodward County, Oklahoma, have been described by McIlvain and Armstrong

(1966), McIlvain and Shoop (1965), and Shoop and McIlvain

(1964). These authors reported a variable plant response

to fire, relative to the timing of the burn and amount of

precipitation. In 1964, a dry year, burning on April 8

increased total grass and forb production and shin-oak

density, but total grass production remained relatively

unchanged. In the same year, burning on May 8 increased

shin-oak density and decreased forb and grass production. In 1965, a wet year, grass and forb production showed

dramatic increases in response to burning. Grass production increased 29% after a burn on April 19 and 66% after

a burn on May 11, while forb production increased 140%

after the April 19 burn and decreased 19% after the May

burn. No shin-oak response data were reported for this

year. Spring burning in years of adequate rainfall will

increase production of sand bluestem and switchgrass,

decrease production of little bluestem, and induce prolific

resprouting of top-killed shin-oak (McIlvain and

Armstrong 1966). Shin-oak stems are highly susceptible to

fire, with post-burn top-kill often approaching or equaling 100% (Boyd and Bidwell unpublished data1997).

Slosser et al. (1985) reported that a March 11 burn of

shinnery in Kent County, Texas, had little effect on vegetative composition but decreased leaf litter for 3 yrs following treatment. These authors noted that although top-kill

of shin-oak was near 100%, frequency of oak occurrence

had equalized between burned and unburned treatments

within 8 mos due to vigorous oak resprouting. Frary
USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998 15

(1955) reported shifts in cover of plant species 1 yr following a controlled burn on March 10 in Roosevelt County,

New Mexico. Total plant density was reduced from 37% to

26%, annual forbs increased from 43% to 51% of total

cover, grasses decreased from 49% to 44% (including no

change in little bluestem at 10%), and shin-oak decreased

from 8% to 5%.

Zobeck et al. (1989) studied the effects of fire on wind

erosion and soil nutrients. They measured 330 times more

sediment from burned plots relative to controls. However,

measurement of sediment used a qualitative measure,

making it difficult to assess the biological significance of

increased wind-borne sediments. Pettit (1984) observed

little recovery of grasses from a September wildfire, leaving insufficient cover for soil protection during winter.

Wildlife Species

Mammoth, bison, and elk remains are found in the

shinnery, accompanied by Indian sites dating from 11,500

yrs ago to historic times (Beckett 1976, Smith 1971, Smith

1985). The shinnery remains rich in wildlife.

Mule deer

Mule deer (Odocoileus hemionus) are common in shinnery in Texas and New Mexico (Bryant and Morrison

1985) and occasionally cross from Texas into southwestern Oklahoma (Caire et al. 1989). Oak, including acorns,

buds, and leaves, is the principal food for mule deer that

inhabit shinnery (Ligon 1927). Studies in Palo Duro Canyon in the Texas Panhandle have shown oak to comprise

37% of mule deer diets even where shinnery was only 1 of

several vegetation types comprising the habitat (Krysl et

al. 1980); in autumn the figure was 78% (Gray et al. 1978).

However, although these studies claimed to deal with

Quercus havardii, Q. mohriana seems more likely for Palo

Duro Canyon. “Generally, Texas and Eastern New Mexico

mule deer are on a ‘poor to fair’ nutritional plane” (Bryant

and Morrison 1985), but deer in the shinnery appear

heavy-bodied with broad antlers, probably due to the

nutritional value of acorns (USDI BLM 1979). “The deer

appear to be heavily dependent upon shinnery oak yearlong” (USDI BLM 1980). The tall mottes of hybrid oak

provide important cover for deer (Turrentine 1971). Shinoak is a species of prime importance and should be protected as mule deer habitat. Small patches (less than 5

acres) of shinnery should be burned in February or March

to encourage sprouting of nutritious new oak growth

(Bryant and Morrison 1985).

White-tailed deer

Shinnery is a principal home for white-tailed deer

(Odocoileus virginianus) in the southern Great Plains. For

instance, in Harmon County, Oklahoma, white-tails were

reported only in a riparian area and shinnery (Martin and

Preston 1970). In New Mexico, white-tails on the plains

may be limited to shinnery habitat (Ligon 1927, Raught

1967). Bailey (1905) thought that the sandhills deer

might be a separate subspecies, an idea entertained by

Ligon (1927), pursued by Calvin Smith (Huey 1970, Smith

1985), and accepted by Raught (1967), but Findley et al.

(1975) assign them to subspecies texanus. Ligon (1927)

described acorns as superior food for the sandhills whitetails, and Raught (1967) also referred to the importance of

acorns. Bryant and Demarais (1992) emphasized the importance of forbs and acorns in white-tail diets and the

importance of a wide diversity of plant species. They

recommended the use of prescribed fire to top-kill shrubs

and obtain nutritious new sprouts. Darr and Klebenow

(1975) reported that shin-oak constituted 11% of whitetails’ autumn diet on a ranch that included only 1.6%

shinnery. Oak mottes provided important cover; of 6

vegetative types on the ranch, only the riparian type

supported higher deer densities than shinnery, although

others provided better cover than the 18-inch tall shinoak.

Pronghorn antelope

Now uncommon in Oklahoma and adjacent Texas,

pronghorns (Antilocapra americana) are common in the

southwestern Texas Panhandle and southward in shinnery (Pettit 1978a, USDI BLM 1980; D. A. Swepston personal communiction 1997), including areas far from open

grassland. Pronghorns are not limited to shinnery. They

are opportunistic feeders, with a diet of up to 71% shrubs

(Yoakum 1980), which they may rely on for winter survival (Harmel 1979, Howard et al. 1990). In shinnery of

Cochran and Yoakum counties, however, Roebuck (1982)

and Roebuck and Simpson (1982) found pronghorns eating an average of 68% forbs, 22% shrubs (including 13%

shin-oak), and 4% grass. Roebuck compared the amount

eaten to the relative availability of the various forages and

derived selection indices. A selection index of 1 indicates

a plant eaten in the same proportion as its availability. On

that scale trailing ratany had an index of over 2,000, from

a trace to .2% of available forage but 21% to 34% of diet.

The index for spectaclepod rose to 940 in autumn and that

for mesquite to 510 in winter. The index for sandsage was

2.3 in winter and 3.7 in spring and shin-oak indices, the

most abundant plant in the community, ranged from .1 in

winter to .7 in summer and fall.

Peccary (Javelina)

Collared peccaries (Dicotyles tajacu) occur in the southernmost shinnery in Eddy and Lea counties, New Mexico,

and there are a few peccaries in the adjacent Texas counties (Andrews, Loving, and Winkler). In 1927, Ligon re-
16 USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998

ported that only a few individuals remained in shinnery

country, and Donaldson (1967) reported none, but they

have been sighted often in the 1990s. Although Donaldson

lists peccary foods in southwestern New Mexico, of which

acorns, mesquite beans, prickly pear and cholla, hog potato (rushpea), and filaree would also be available in the

shinnery area, there are no studies of peccaries specifically

in shinnery. In Arizona studies, Knipe (1956) describes

acorns as their most sought-after food, leading to local

migrations in search of good crops.

Barbary sheep

In 1957 and 1958, 44 Barbary sheep (Ammotragus lervia)

were introduced into Palo Duro Canyon near Amarillo.

The population prospered and by 1980 was competing for

food resources with native mule deer. Of the several

habitat types available, the sheep favored shinnery, and

31% of their diets was shin-oak. Other shrubs, mostly not

in shinnery, constituted 19%, forbs 26%, and grasses 24%

of their diet (Krysl et al. 1980).

Lagomorphs

Black-tailed jackrabbits (Lepus californicus) and desert

cottontails (Sylvilagus auduboni) occur throughout the shinnery. Eastern cottontails (S. floridanus) are in shinnery in

Oklahoma (Martin and Preston 1970), but because of its

use of heavily wooded cover, the Eastern cottontail is less

common than its congener in shinnery. A subspecies, S.

floridanus llanensis, was described from the shinnery country of west Texas (Blair 1938).

Little is known about the ecology of hares and rabbits

in this vegetation. Davis et al. (1975a) and Griffing and

Davis (1976b) observed the feeding of jackrabbits on yucca

leaves, mesquite leaves and pods, snakeweed, and forbs

near the southern limit of the shinnery. Jackrabbits preferred more open grass-shrub stands compared with cottontails, which depend upon low-hanging vegetation for

escape cover and concealment (Davis et al. 1975a). The

ecology of rabbit mounds in relation to soil nutrients,

mycorrhizal spores, and other fungi and bacteria, and as

a disturbance factor permitting establishment of seedlings is discussed by Dhillion et al. (1994).

Rodents

About 16 rodent species commonly make their homes

in shinnery; none is limited to the community. Occurrences are given in table 3 for 3 widely separate sampling

points (Martin and Preston 1970, USDOE 1980, Caire et al.

1989, Willig et al. 1993).

The greater number of species at the Department of

Energy’s (DOE) Eddy County site, driest and least productive of the sites recorded in table 3, reflects the intenTable 3. Rodents of sand shinnery communities.

Harmon Co. Yoakum Co. Eddy Co.

OK TX NM

Dipodomys ordii, Ord’s kangaroo rat + + +

Erethizon dorsatum, porcupine + - +

Geomys bursarius, plains pocket gopher + - +

Mus musculus, house mouse - + +

Neotoma micropus, southern plains woodrat + + +

Onychomys leucogaster, northern grasshopper + + +

mouse

Perognathus flavescens, plains pocket mouse + + +

Perognathus flavus, silky pocket mouse - - +

Perognathus hispidus, hispid pocket mouse + - +

Peromyscus leucopus, white-footed mouse + - +

Peromyscus maniculatus, deer mouse + + +

Reithrodontomys megalotis, western harvest - - +

mouse

Sigmodon hispidus, hispid cotton rat + - +

Spermophilus mexicanus, Mexican ground - - +

squirrel

Spermophilus spilosoma, spotted ground + - +

squirrel
USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998 17

sity of DOE’s several-year trapping effort. The desert

pocket mouse, Perognathus penicillatus, and Merriam’s

kangaroo rat, Dipodomys merriami, were also trapped on

dunes at the site; shin-oak was present but their habitat

was not recorded as shinnery. The nonnative house mouse

was not trapped regularly nor year-round at either site

where it was recorded. Colbert (1986) regarded this species

as unexpected because it is usually associated with more

mesic areas. Although not listed by Willig et al., pocket

gophers do occur in west Texas shinnery. In the northern

counties, Geomys bursarius occurs and from Yoakum and

Terry counties southward, G. knoxjonesii, which has been

split from G. bursarius, occurs (Davis and Schmidly 1994).

Although not commonly found in shinnery, prairie

dogs (Cynomys ludovicianus) in Oklahoma converted more

than 100 acres of shinnery to forb-rich, depauperate grassland (Osborn 1942). Osborn stated that human intervention, especially overgrazing by livestock in the tallgrasses

in shinnery, which was previously unsuitable habitat for

prairie dogs, and destruction of prairie dog predators

contributed to this conversion. Prairie dogs are not found

in shinnery when its tallgrasses are abundant (Allan and

Osborn 1954).

Best et al. (1993) studied food habits of the 4 rodents

most common in shinnery at the DOE site. Ord’s kangaroo

rats primarily ate seeds and fruits of purple sandgrass,

mesquite, and euphorbia, plus arthropods. Northern grasshopper mice predominantly ate arthropods. Southern

plains woodrats consumed plant fibers, new forb growth,

and mesquite, mostly green forage. Spotted ground squirrels

ate mostly arthropods, new forb growth, and grasses. All

4, especially the woodrat, ate shin-oak. The ground squirrel

hibernates, and the others are active throughout the year.

In general, but not necessarily in shinnery, rodents

disperse seeds, shred vegetation and contribute it to humus, mix and aerate soils, and are major food sources for

predators (Jones and Manning 1996). Rodents, especially

pocket gophers, as disturbance factors and in relation to

plant succession in shinnery are discussed by Dhillion et

al. (1994). Several important midgrass and tallgrass species of the shinnery are highly preferred by plains pocket

gophers. In Texas, but not in shinnery, biomass increased

22% when plains pocket gophers were excluded

(Fagerstone and Ramey 1996).

Insectivores

Desert shrews (Notiosorex crawfordi) may occur throughout the shinnery, though records are few. Eastern moles

(Scalopus aquaticus) occur in shinnery in Oklahoma (Caire

et al. 1989).

Lesser prairie-chicken

Lesser prairie-chickens (Tympanuchus pallidicinctus) are

closely associated with shinnery vegetation through most

of their range. (North of the shinnery they occur in

sandsage-dominated communities.) From extreme abundance in the nineteenth century, when they were used to

supply meat in place of domestic poultry, populations

declined to a few thousand in their 5-state area in this

century. Crawford (1980) estimated a 97% decline in lesser

prairie-chickens, while Taylor and Guthery (1980a) estimated a 92% decrease in occupied range. Hunting was

prohibited in Oklahoma from 1915, in New Mexico from

1934, and in Texas from 1937 (Duck and Fletcher 1944, Lee

1950, Litton et al. 1994).

Populations increased and limited hunts were allowed

in some years in Oklahoma and New Mexico, but not

during the drought of the 1950s. Edminster (1954) thought

that hunting is warranted when populations reach 20/

mi2

, which at the time occurred in only about half the

years. Since 1958 in New Mexico, 1959 in Oklahoma, and

1967 in Texas limited hunts have been allowed almost

every year. Crawford (1980) estimated that the kill averaged 12% of the fall population. In the face of a new

population decline throughout the species’ range, New

Mexico cancelled its 1996 to 1998 hunts, Oklahoma cancelled its 1998 hunt, and the U. S. Fish and Wildlife Service

is considering a 1995 petition to list the species as threatened. Concern for the lesser prairie-chicken strongly affects management of the shinnery on public lands and

some private ranches.

Taylor and Guthery (1980b) reported that lesser prairie-chickens in Yoakum County, Texas, preferred sunflower, shin-oak/bluestem, and shin-oak/sand sagebrush

vegetation, in that order, while shin-oak/mesquite and

shin-oak without other shrubs were used less than in

proportion to their abundance. At least in spring, prairiechickens in shinnery avoid areas of dense oak cover and

favor areas in which grasses are more abundant (figure 11;

Cannon and Knopf 1981, Davis et al. 1979).

Lesser prairie-chickens in shinnery eat mostly plant

material except in summer, when insects, mainly grasshoppers, predominate. Table 4 records foods by season

from Chaves County, New Mexico, in 1976 to 1978 (Davis

et al. 1980, Riley et al. 1993a). Davis et al. (1981) reported

dietary composition of lesser prairie-chickens in Roosevelt

County, New Mexico, in 1979 and 1980 with similar results; major differences were a shortage of acorns (1%, 0%,

14%, and 25% by season) and greater consumption of

insects (1%, 88%, 69%, and 0% by season). In Cochran

County, Texas, Crawford and Bolen (1976a) found autumn diets of prairie-chickens, with access to cultivated

fields, to be 62% grain sorghum, 7% acorns, 8% oak galls,

13% other plants, 6% grasshoppers, and 4% other insects.

Davis et al. (1981) sampled crops of hunter-killed birds

from Cochran County and obtained similar results. Doerr

and Guthery (1983a) studied frequency of foods in prairiechicken droppings in Cochran County. Their figures were

close to those in table 4 except that in spring catkins were
18 USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998

19% of diet while insects were 27%, acorns were lacking

from in summer and fall diets, while percentages of insects were 60% in summer and 65% in fall. respectively.

An absence of acorns in the diet probably relates less to

preference and more to the variability of shin-oak acorn

production.

Shin-oak catkins, and perhaps buds, may be an important food source during midspring, given that they are

high in crude protein content (catkins = 22.3%, buds = 19.1%

crude protein) and are available when other food sources

may be in limited supply (Vermeire and Boyd unpublished manuscript 1997). Consumption of these food items

is of interest given their high phenolic content (catkins = 17.5% and buds = 16.6% gallic acid units) (Vermeire

and Boyd unpublished manuscript 1997). High levels of

phenolic intake in domestic bird species leads to decreased

weight gain and to egg shell thinning (Sell et al. 1983),

although other free-ranging gallinaceous birds have been

reported to consume forages high in phenols (Moss 1972).

Diets of chicks and juveniles are comprised of 85% to

99% insects (Davis et al. 1981, Taylor and Guthery 1980a).

In New Mexico, brood foraging sites are less grassy, more

shrubby, and the vegetation is shorter than in nesting sites

(Riley and Davis 1993). In west Texas and eastern New

Mexico, brooding territory was more dune-like than general habitat and had the greatest coverage and height of

shin-oak (Ahlborn 1980, Sell 1979). In Oklahoma, Copelin

(1963) found broods mostly in oak mottes. Jones (1963)

noted that communities rich in forbs produced more

insects than other habitat types.

The supply of safe nest sites limits prairie-chicken

populations (Davis et al. 1981, Doerr and Guthery 1983b,

Haukos and Smith 1989b, Sell 1979). Most nesting attempts are unsuccessful in Chaves County due to coyotes,

striped skunks, and snakes (Riley et al. 1992) and in

Oklahoma due to bullsnakes, Cooper’s hawks, sharpshinned hawks, crows, and ravens (Duck and Fletcher

1944). In 3 studies, nest failures were 5 of 8, 26 of 36, and

11 of 13 (Haukos and Smith 1989a, Riley et al. 1992, Sell

1979). Successful nests are mostly in dense grass, surrounded by tallgrass (Riley et al. 1992). Overhead cover,

which can be oak, should be at least 50% for successful

nests (Haukos and Smith 1989b).

Management of lesser prairie-chicken habitat involves

maintenance of a mosaic of vegetative conditions. Nesting

cover may be improved with light stocking rates, periodic

deferment, or exclosure of grazing livestock (Ahlborn

1980, Davis et al. 1979, 1981, Donaldson 1969, Frary 1957,

Haukos and Smith 1989b, Jackson and DeArment 1963,

Ligon 1927, Litton et al. 1994, N. M. Dept. Game & Fish

1984, Taylor and Guthery 1980a). Although prairie fires

were the nesting hen’s worst enemy (Bent 1932), prescribed fire may be used to improve courting areas (Cannon 1979), increase forbs (McIlvain and Shoop 1965), and

improve the quality of nesting habitat in years subsequent

to burning (Cannon 1979). Additionally, prairie-chickens

require shady resting cover, escape cover, and roosting

sites as discussed by Copelin (1963), Crawford and Bolen

(1976b), Davis et al. (1981), Donaldson (1969), Jones (1963),

Riley et al. (1993b), Sell (1979), Snyder (1967), Taylor and

Guthery (1980a, 1980b), and Turrentine and Klebenow

(1972).

Figure 11. A male prairie-chicken struts in springtime with an

upright horn of feathers.

Table 4. Adult prairie-chicken foods.

Percent by volume

Food Spring Summer Autumn Winter

Acorns 15 21 39 69

Other seeds 0 0 4 0

Oak insect galls 0 1 10 0

Oak catkins, leaves 34 0 2 0

Other plants 45 22 27 26

Grasshoppers 0 39 15 0

Other insects 6 16 3 5
USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998 19

Quail

Scaled quail (Callipepla squamata) inhabit the southern

and central parts of the shinnery; they become less common northward but do reach western Oklahoma. Bobwhite quail (Colinus virginianus) inhabit the northern and

central parts of the shinnery, becoming less common

southwestward, although they are spreading and increasing in that direction. Neither species is limited to shinnery

but both use the type extensively. Both species nest on the

ground, usually under shrubs (yucca, mesquite, shin-oak)

surrounded by relatively dense cover mostly of mid and

tallgrasses, or in the case of bobwhite, nests may be located

within perennial grass clumps (Bidwell et al. 1991). Both

species use shin-oak and other shrubs as loafing, hiding,

and roosting cover (Davis et al. 1974, Guthery 1982, Sawyer 1973, Tharp 1971, Turrentine 1971, Turrentine and

Klebenow 1971, Webb and Guthery 1982, Wood and Schnell

1984).

Scaled quail eat mainly seeds, mostly of woody plants

and forbs but also of grasses. In winter, mesquite seeds

may supply as much as 37% of their food and snakeweed

seeds up to 35%; shin-oak acorns are a minor item. Green

vegetation can supply up to 20% of scaled quail diet, and, in

summer, insects supply up to 48%. Insects are the primary

food for quail chicks (Best and Smartt 1985, Campbell et al.

1973, Davis et al. 1975b, Dixon and Knight 1993).

Bobwhite foods are similar to those of scaled quail in

shinnery (Guthery 1982, Jackson 1969), but bobwhites

may be more effective at cracking and eating shin-oak

acorns (Bird and Bird 1931, Guthery 1980, Jackson et al.

1962). Food supplies may become critical from the first

frost until spring (Turrentine 1971); competition with

kangaroo rats can be acute (Inglis 1960). Early winter

foods of bobwhites in shinnery in Oklahoma, by percentage volume, were cultivated grains 29, grass seeds 16,

ragweed seeds 22, other forb seeds 12, acorns 11, other

woody-plant seeds 9, green herbage 1, and animal matter

1 (Hanson 1957). Bird and Bird (1931) reported that seeds

of sunflower and ground cherry may also be important

winter food items in the shinnery region of Oklahoma.

Brood habitat is characterized by high insect (Hurst 1972)

and forb availability with sufficient bare ground to allow

movement of chicks through understory vegetation

(Bidwell et al. 1991). Insect and forb availability may be

increased by burning or spring disking (Bidwell et al.

1991, Hurst 1972, Turrentine 1972).

Turkey

Rio Grande turkeys (Melagris gallopavo subsp. intermedia)

occur in shinnery in southwestern Oklahoma and the

adjacent Texas Panhandle. Logan (1973) described seasonal behavior of Rio Grande turkeys in southern Roger

Mills and northern Beckham counties, Oklahoma. Use of

shrub habitat, which included a mix of shin-oak, sand

plum, skunkbrush, and sandsage, was primarily associated with loafing activities during the midday period and

with nesting (figure 12). Twenty percent of located nests

were within shrub habitat, however, most birds selected

nests sites within alfalfa fields (56% of located nests).

Mourning doves

Mourning doves (Zenaidura macroura) occur throughout the shinnery during the warm seasons and sporadically in winter, or regularly in the south. They are not

restricted to the community, but Downing (1957) noted

that ground-nesting mourning doves had higher nesting

success in shinnery than in other vegetation types studied

in northwestern Oklahoma. They nest from May through

early September. In the southern shinnery, their summer

foods (by percentage of volume) were croton seeds 81,

leaf-flower seeds 16, other forb seeds 3; in winter leafflower seeds 48, spurge seeds 23, other forb seeds 10, grass

seeds 18 (Griffing and Davis 1974). Broader studies in

Texas and northwestern Oklahoma, applying partly to

shinnery, found croton, sunflower, amaranth, and sand

paspalum seeds predominant in mourning dove diets,

Figure 12. A turkey nest is under sandsage and shin-oak in

western Oklahoma.
20 USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998

plus sorghum and wheat when available (Carpenter 1971,

Dillon 1961). In the southern shinnery, mourning dove

and scaled quail diets conflicted only with regard to

croton seeds (Griffing and Davis 1976a).

Perching birds

About 20 species of songbird nest in shinnery and

perhaps 80 more visit it regularly, but none is restricted to

the community. In the southernmost shinnery, where 31

species were observed, the most common passerines were

white-necked raven, loggerhead shrike, pyrrhuloxia, blackthroated sparrow, and lark bunting, all but the last nesting

in shinnery (Davis et al. 1974). In Yoakum and Bailey

counties, Texas, 50 passerine species were censused in

shinnery, 17 of them nesting, 21 wintering, and 23 migrant

(Fischer and Bolen 1980). In Oklahoma, county-wide

records for Beckham, Ellis, and Roger Mills counties indicated the occurrence of 142 species of passerine birds. Of

these 94 species are associated with brush habitat—

shin-oak, fence rows, and windbreaks (ODWC 1985).

Predators (mammals and birds)

Several predators have been previously mentioned:

insect-eating rodents, insectivores, gallinaceous birds, and

perching birds. In addition, the cave myotis, pallid bat,

and Brazilian free-tailed bat, and probably several others,

are associated with water (primarily stock tanks) in shinnery (USDOE 1980). Mainly the meat-eating predators are

the Carnivora and raptors discussed here.

Gray wolves and cougars were formerly top carnivores

of the plains; omnivorous black bears and grizzlies also

ranged into shinnery country. Currently the principal

native carnivores of the shinnery are coyote, bobcat, badger (figure 13), striped skunk, and swift fox. Cougars are

occasional in the New Mexico shinnery and are rare

farther east and north. Carnivorous gray fox and red fox

and omnivorous ringtail and raccoon are uncommon to

rare in the shinnery. None of these species is restricted to

shinnery (Caire et al. 1989, Davis and Schmidly 1994,

Findley et al. 1975, Kilgore 1969, Martin and Preston 1970,

USDOE 1980).

Northern harriers, Swainson’s hawks, red-tailed hawks,

kestrels, burrowing owls, and short-eared owls are fairly

regular inhabitants of (or foragers in) all of the shinnery.

Mississippi kites are common in the northern and eastern

parts of the shinnery, Harris’ hawks are common in the

south. Less common are sharp-shinned hawks, roughlegged hawks, ferruginous hawks, golden eagles, prairie

falcons, barn owls, and great-horned owls. Though not

raptors, roadrunners and loggerhead shrikes are also

predatory birds in shinnery. Although none of these species is limited to shinnery, Harris’ and Swainson’s hawks

are nearly so-limited in eastern New Mexico (Bailey 1928,

Bednarz 1988, USDOE 1980, Wood and Schnell 1984).

On a shinnery ranch in Oklahoma, the ranchers “figure

each coyote worth $25/yr increased beef production and

we are of the opinion that the bobcat is even more valuable” for their control of forage-consuming rodents and

lagomorphs (Bunch 1961). Rodents, lagomorphs, and

grasshoppers are the main foods of most of the raptors

previously named; cumulatively, raptors probably equal

the coyote in controlling rodent populations. However,

only Swainson’s and Harris’ hawk diets have been studied in shinnery (Bednarz 1986a, 1986b).

Reptiles

Western box turtles (Terrapene ornata) are the only

terrestrial turtles to inhabit shinnery regularly, although

several aquatic turtles occur in water bodies. The turtles

eat insects, plant material, and carrion (Degenhardt et al.

1996, Sievert and Sievert 1993).

About 25 snake species are recorded in shinnery, of

which the plains hognose snake (Heterodon nasicus), night

snake (Hypsiglena torquata), coachwhip (Masticophis flagellum), bullsnake or gopher snake (Pituophis melanoleucus),

milksnake (Lampropeltis triangulum), massasauga (Sistrurus

catenatus), and prairie rattlesnake (Crotalus viridis) are

Figure 13. A badger amid shin-oaks.
USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998 21

most common (Degenhardt et al. 1996, Sievert and Sievert

1993, Webb 1970, Wolfe 1978). These snakes eat rodents,

lizards, snakes, and eggs. The larger snakes also eat cottontails. Smaller species and individuals eat centipedes,

grasshoppers, and other arthropods (Degenhardt et al.

1996, Stebbins 1985).

Ten lizard species are in shinnery in southeastern New

Mexico: prairie lined racerunner (Cnemidophorus

sexlineatus), western whiptail (C. tigris or C. marmoratus),

Great Plains skink (Eumeces obsoletus), leopard lizard

(Gambelia wislizenii), lesser earless lizard (Holbrookia

maculata), Texas horned lizard (Phrynosoma cornutum),

roundtail horned lizard (P. modestum), prairie lizard

(Sceloporus undulatus), and side-blotched lizard (Uta

stansburiana) (Degenhardt and Jones 1972, Degenhardt et

al. 1996, Gorum et al. 1995, Wolfe 1978). None of these

species is limited to shinnery, but, the shinnery oakmesquite habitat seems to be particularly suitable for sideblotched and western whiptail lizards (Wolfe 1978) and

these 2 species are subjects of ecological studies in shinnery (Best and Gennaro 1984, 1985).

In addition, in 4 New Mexico and 4 Texas counties,

occurs the sand dune lizard (Sceloporus arenicolus), the

only reptile restricted to shinnery (figure 14; Degenhardt

et al. 1996, Degenhardt and Jones 1972). This lizard occurs

only in areas with open sand (especially in large, deep

blow-outs), but it forages and takes refuge under shin-oak

and is seldom more than 4 to 6 ft from the nearest plant. The

sand dune lizard is lacking where blow-outs, topographic

relief, or shin-oak are lacking and where 15% or more of

the sand has a grain size less than .149 mm (Gorum et al.

1995). Because this reptile is listed as endangered by New

Mexico and is under consideration for federal listing, its

presence affects management of public lands. For instance,

herbicidal treatment of shin-oak is forbidden on public

lands in sand dune lizard habitat (USDI BLM 1997). Bailey

and Painter (1994) are concerned about the possible extinction of the sand dune lizard where shinnery is managed

for livestock and oil-and-gas production because of habitat

lost to shin-oak control, well pads, roads, and pipelines.

We found no studies of reptiles in shinnery north of

southeastern New Mexico. Of the species named previously, 6 are throughout the shinnery but the sand dune

lizard, western whiptail, and leopard lizard are only in the

southern part. The side-blotched and roundtail horned

lizards are missing from the central shinnery and probably Oklahoma, though they are found in the Texas Panhandle (Dixon 1987). Regional inventories in Oklahoma

record 9 lizard species from counties containing shinnery,

but it is unknown which species are in shinnery. Species

not listed previously are spotted whiptail (Cnemidophorus

gularis), eastern collared lizard (Crotaphytus collaris),

ground skink (Lygosoma laterale), and western slender

glasslizard (Ophisaurus attenuatus) (Sievert and Sievert

1993, Webb 1970).

These lizards eat snails, insects, and other arthropods

such as millipedes, scorpions, and spider. The larger

species eat small rodents (pocket mice) and other lizards.

Several include plant material in their diets (Best and

Gennaro 1984, 1985, Degenhardt et al. 1996, Stebbins 1985).

Invertebrates

Lists of arthropod groups from

Eddy County, New Mexico, come

from the most complete study of

invertebrates in shinnery, but

identifications are not carried to

the species nor often to the genus

level. From one grass-rich shinnery site, Wolfe (1978) lists 6 families of Orthoptera, 1 of

Psocoptera, 10 of Hemiptera, 8 of

Homoptera, 19 of Coleoptera, 3

of Neuroptera, 4 of Lepidoptera,

17 of Diptera, and 19 of Hymenoptera plus members of Scorpionida, Solpugida, Acarina,

Araneida, and Ostracoda. From

sand sagebrush, Wolfe lists 17

families of insects, and from the

stomachs of 2 lizard species in

shinnery, 24 families. More detailed studies of arthropods, inFigure 14. A sand dune lizard on shin-oak litter. cluding termites, especially as
22 USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998

they are involved in decomposition, were carried out at

the same DOE site (Whitford 1982, Whitford et al. 1980).

Davis et al. (1979) identified insects from prairie-chicken

crops in Chaves County, New Mexico, shinnery: cockroaches, mantids, crickets, walking sticks, treehoppers,

scentless plant bugs, shield-backed bugs, short-horned

and long-horned grasshoppers, robber flies, click beetles,

darkling beetles, ground beetles, leaf beetles, scarab beetles,

silken fungus beetles, snout beetles, weevils, caterpillars

and moths, ants, and plant-gall insects (cynipid wasps,

mainly).

Tucker (1970) lists 6 species of gall wasps (Cynipidae)

from shin-oak and James Zimmerman (personal

communiction 1997) adds Xystoteras sp. and Xanthoteras

eburneum, the latter studied by Dodson (1987). Other

insects that attack this host are discussed under

Autoecology of Shin-oak.

Ants recorded at the Eddy County site are Campanotus

sp., Conomyrma insanum, Crematogaster sp., Iridomyrmex

prunosum, Myrmecocystus dipilis, Pogonomyrmex apache, P.

desertorum, and P. rugosus (Whitford et al. 1980). Dhillion

et al. (1994) recorded Aphaenogaster cockerelli, Campanotus

vicinis, Crematogaster punctulata, Myrmecocystus mimicus,

Pheidole crassiconis, P. dentata, and Pogonomyrmex barbatus

in west Texas shinnery. Ant-plant-microbe interactions

were studied by McGinley et al. (1994).

In 1996, the most frequently encountered short-horned

grasshoppers in shinnery in New Mexico and adjacent

Texas were (in descending order of frequency) Schistocerca

alutacea subsp. lineata (which eats shin-oak), Campylacantha

olivacea, Melanoplus glaucipes, Mermiria bivittata, Dactylotum

bicolor, Phlibostroma quadrimaculatum, Spharagemon

cristatum, Melanoplus aridus, Paropomala wyomingensis, P.

pallida, Melanoplus flavidus, and M. foedus; 18 additional

species were found at only 1 or 2 of 17 collection points

(Peterson unpublished data).

An important and much-studied insect resident of shinnery in west Texas is the boll weevil (Anthonomus grandis),

an introduced pest of cotton that overwinters in litter and

then in spring infests cotton fields up to half a mile away

(Rummel and Adkisson 1970). Shin-oak litter provides

one of the best overwintering habitats for the weevil in the

Rolling Plains and adjacent High Plains (Brown and

Phillips 1989, Carroll et al. 1993). Mottes of taller oaks are

particularly favorable to the weevils. Emergence from

litter in mottes lasts a month longer than from low shinnery, which is long enough to ensure that some weevils

live to infest fruiting cotton in late June and early July

(Slosser et al. 1984).

Soil microarthropods and nematodes are discussed but

not identified beyond family by Elkins and Whitford

(1982). Small fungus-eating mites (Pyemotidae,

Lordalychidae, and Tarsonemidae) dominated soils in

winter, and these were replaced by large predaceous

mites (Rhodacaridae, Laelapidae) in spring and summer.

Collembola and Psocoptera were present in buried litter

but not in soil cores. Nematodes were common in buried

litter. Only 3 families of mites were observed in surface

litter: Nanorchestidae during summer and, sporadically,

Nanorchestidae, Anystidae, and Bdellidae during nonsummer months.

Management

Nearly all shinnery in Texas and Oklahoma is on private lands; the exceptions are: scattered parcels in highway rights-of-way and state parks; 13,000 acres in Black

Kettle National Grassland (USDA Forest Service) in Roger

Mills County, Oklahoma; and 16,000 acres in Packsaddle

Wildlife Management Area (Oklahoma Department of

Wildlife Conservation). In New Mexico, the BLM estimates its shinnery at 480,000 acres (USDI BLM 1979) to

1,200,000 acres (USDI BLM 1986, 1994); state trust lands

include about 500,000 acres, and the New Mexico Department of Game and Fish (1984) manages 21,000 acres.

Nearly all of the shinnery is used for livestock (cattle)

grazing and hunting. Recreational uses other than hunting are limited: birdwatching, especially prairie-chickens

during their spring courting, and dune-hiking and dunebuggy-riding. Small areas are protected for scientific and

wildlife purposes including BLM’s Mescalero Sands Area

of Critical Enviromental Concern and Mathers Research

Natural Area in New Mexico (Peterson and Rasmussen

1986, USDI BLM 1997), and 21,000 acres protected as

prairie-chicken habitat in tracts of 26 to 6,550 acres in 4

New Mexico counties (New Mexico Dept. Game & Fish

1984). In the Permian Basin (southeastern New Mexico

and adjacent Texas), petroleum activities strongly affect

shinnery management.

Hunting

From the abundance of both PaleoIndian and modern

Indian artifacts, thought mainly to represent hunting

camps, we know that game harvesting has occurred in the

shinnery for millennia (Beckett 1976, Smith 1985). In the

nineteeth century, harvests were large and provided food

for explorers and settlers (Gregg 1844, Ligon 1927). Around

1900, railways ran special trains for sportsmen to towns in

the Texas Panhandle and placed iced cars on sidings for

preservation of the lesser prairie-chicken harvest (Jackson

and DeArment 1963). Most game populations are now

much smaller, but hunting is an important activity in

shinnery and increasingly affects management of the rangeland.
USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998 23

Mule deer, pronghorn, and scaled quail are hunted in

shinnery in New Mexico and Texas, white-tail deer and

turkey in Texas and Oklahoma, feral hogs and Barbary

sheep in Texas, and lesser prairie-chicken, bobwhite quail,

mourning dove, and small game (rabbits and jackrabbits)

in all 3 states. Trapping of coyotes, bobcats, foxes, skunks,

and other mammals occurs in shinnery in all 3 states

(Davis and Schmidly 1994, Duck and Fletcher 1944, NM

Dept. Game & Fish 1967, D. A. Swepston personal communication 1997).

Management of shinnery has focused on livestock grazing, but unfavorable economics in that industry plus the

increasing amounts that sportsmen are willing to pay for

hunting privileges make management for wildlife attractive (Hamilton 1979, Holechek 1981, Knight 1985, Teer

1996). This is most true on the private-land ranches of

Texas, Oklahoma, and the easternmost New Mexico shinnery. The BLM, which manages much of the New Mexico

shinnery, is shifting toward active habitat protection for

wildlife including restrictions on oil-and-gas activities

and livestock grazing (USDI BLM 1997). Increased emphasis on wildlife can affect livestock stocking rates and

grazing systems, brush control practices, revegetation,

wildlife transplants, and the design of fences (Bryant et al.

1982, Bryant and Morrison 1985, Hanson 1957, Holechek

1981, Lamb and Pieper 1971, Taylor and Guthery 1980a,

Teer 1996, Tharp 1971, Webb and Guthery 1982, Yoakum

1980).

Managing for livestock and big game can be beneficial

to both, for instance, by preventing unwanted vegetation

trends due to a single species’ use (Bernardo et al. 1994,

Holechek 1980). More frequently there are tradeoffs (discussed under Livestock Grazing below). For instance,

hunting leases in bobwhite and whitetail country in central Oklahoma vary from $1 to $10/acr/yr, depending on

habitat quality. The cost of achieving high habitat ratings

in that non-shinnery area is higher for quail than for deer

due to reductions in cattle numbers required to protect

brood habitat and nesting cover (Bernardo et al. 1994, T.

Bidwell personal communication 1997).

Livestock Grazing

Plants of the shinnery coevolved with grazing mammals. The larger indigenous herbivores included pronghorn antelope, bison, elk, mule deer, and white-tailed deer

(Bailey 1905, Haines 1995, Smith 1971, Tyler and Anderson 1990, Yoakum et al. 1996). Today, cattle are grazed on

nearly all of the shinnery range; sheep, horses, and goats

are minor species. High stocking densities of cattle are

thought to have transformed much of the shinnery from

grass-oak communities to systems dominated by shin-oak

(Dittemore and Blum 1975 and many other SCS publications, Duck and Fletcher 1944, Jackson and DeArment

1963, Litton et al. 1994, Pettit 1994, USDI BLM 1977, 1979),

to have increased annual grasses and forbs at the expense

of perennial grasses (Lenfesty 1983, Richardson et al.

1975), and to have increased bare ground, subjecting soils

to wind erosion (Holland 1994). In 1970, the Forest Service

estimated that 1.5% of the shinnery was in good condition,

92.3% in fair, and 6.2% in poor (USDA Forest Service

1972). These condition ratings were based on resemblance

of current vegetation to the presumed original vegetation,

and on its cover and vigor and on soil factors.

Evaluating the effects of livestock grazing on the structure of shinnery is problematic without at the same time

considering the influence of fire and the interaction between historic fire and grazing in this system. In other

systems, decreased fire frequency allowed woody vegetation to increase, often to the detriment of herbaceous

species. Grazing may act to decrease fire frequency by

reducing herbaceous fine fuels (Box 1967).

The herbaceous plants of ungrazed and heavily grazed

areas differ. In ungrazed, there exist mostly perennial

grasses, while in grazed areas there is a shift toward

annual forbs and annual grasses and changes in perennial

grass species. For example, in the southern shinnery,

species in ungrazed or lightly grazed areas include

bluestems, switchgrass, side-oats grama, plains

bristlegrass, giant dropseed, and big sandreed; in heavily

grazed areas species include mostly annual forbs and

annual grasses (sandbur, purple sandgrass, fringed

signalgrass, false buffalograss), and the main perennial

grasses become three-awns, hooded windmillgrass, and

red lovegrass. Hairy grama, sand muhly, and sand

paspalum also increase in percentage under heavy grazing, while sand dropseed may increase or decrease

(Lenfesty 1983, Pettit 1986, Richardson et al. 1975, USDI

BLM 1977).

Holland (1994) examined the effects of grazing on plant

species richness and distribution in shinnery in Yoakum

County where shin-oak constituted 60% to 80% of plant

cover. Using 2x2 m quadrats, she found that in ungrazed

areas 252 m2

 of quadrats were needed to find 80% of the

total 25 species present while in grazed areas only 115 m2

of quadrats were needed to find 80% of the total 25 species

present. A conclusion is that species are more evenly

spread where grazed. The average number of species per

2x2 m quadrat was twice as high in the grazed treatment,

although there was no difference in species richness between treatments. The density of herbaceous plants was

almost twice as high in grazed (4.7 individuals per m2) as

in ungrazed treatments (2.5 individuals per m2) due to

cattle’s creating open sites in the shin-oak litter for seedling establishment (Holland et al. 1994).

According to McIlvain (1954), “Grass growing in shinnery oak is much less nutritious than grass growing in

open sunlight. The effect of shade is to increase fiber
24 USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998

content and lower sugar content greatly. Consequently,

livestock reluctantly graze these shaded grasses. Instead,

they concentrate in open areas where the grass is weakened and often killed by heavy grazing use.” There are no

data to support these conclusions.

Selection indices (relative amount eaten divided by

relative amount available) were determined for cattle use

of plants in shinnery in Eddy County, New Mexico.

Dropseeds, sandbur, sand paspalum, plains bristlegrass,

gyp phacelia, plains sunflower, evening primrose, and

camphorweed had selection indices above 1; black grama,

lovegrasses, and Schweinitz flatsedge had values about 1;

shin-oak, sand sagebrush, mesquite, three-awns, false

buffalograss, fall witchgrass, hairy grama, fringed

signalgrass, Pectis species, leafflower, and crotons had

values below 1, where summer cattle diets were 81%

grasses, 17% forbs, and 2% shrubs (LaBaume et al. 1980).

It is sometimes claimed that cattle do not eat shin-oak or

eat very little shin-oak when other forage is available

(Villena-Rodriguez 1987), but this is not generally true. In

Yoakum County, the relative frequency of shin-oak in

cattle diets averaged 4.2%, 16.2%, 20%, 4.1%, and 9.5% at

monthly intervals from June to October (Plumb 1984,

Plumb and Pettit 1983). Roebuck (1982) reported that shinoak had a relative frequency of 14% (spring through fall)

in the diets of cattle grazing shinnery in northwestern

Texas. However, in some studies oak was not eaten significantly during the period of study (LaBaume et al. 1980,

Weir 1990). In Dickerson’s (1987) study, cattle ate significant amounts of shin-oak (20% of diet) in only 1 mo of 13.

Shin-oak “provides dependable forage during drought

periods” (Holechek and Herbel 1982) and “is of much

importance as forage” (Dayton 1931). “Shinnery oak provides safe forage during midsummer and fall, and receives heavy grazing pressure during drought years. A

resilient shrub, its vigor and density are rarely harmed by

heavy grazing” (USDI BLM 1979). Holland (1994) found

that oak was significantly shorter in grazed areas compared with ungrazed, though not differing in density.

Cattle are generally removed from shinnery pastures

during the 4-week period in spring following bud-break

when shin-oak buds and immature leaves are poisonous

to them (Allison 1994, Dollahite 1961). Livestock consumption of shin-oak buds and leaves during this time

may lead to malaise, reduced conception rates, lower

weight gains, and death (Jones and Pettit 1984). Several

Texas counties have reported death due to shin-oak poisoning of more than 1,000 cows in a single year. Losses

also occur due to chronic poisoning and increased management costs (moving cattle from shinnery pastures)

(Dollahite 1961). The phenolic compounds responsible for

the toxicity of shin-oak may have evolved as a defense

against grazing animals; buds of grazed oak have been

reported to contain higher phenoic levels that those of

ungrazed (Vermeire and Boyd unpublished manuscript

1997). Ill effects resulting from shinnery consumption

may be minimized by feeding protein-rich supplement

containing 9% hydrated lime (Sperry et al. 1964).

Dowhower (1981) studied effects of trampling by human

walking, vehicles, and (Dowhower et al. 1981) livestock.

Forbs were most affected by trampling and grasses least;

shrubs were intermediate. Livestock affected shin-oak

more than 2 mi from centers of concentrated activity, but

it quickly made up part of its losses by re-sprouting.

In west Texas, shinnery is sometimes converted to

planted grass pastures, especially weeping lovegrass

(Eragrostis curvula) (Goen et al. 1978). Plowing or disking

is generally necessary, with plowing preferred to bury

weed seeds, distribute nitrogen better, and to bury allelopathic residues from shin-oak and sandbur (Matizha and

Dahl 1991). An attempt was made to plant weeping

lovegrass in undisturbed shinnery; up to 17 seedlings/

foot of row survived from May to August, but half were

gone in September (Pettit et al. 1977).

Because cattle-grazing has the potential to decrease the

productivity of shinnery and to decrease production of

perennial grasses, there is considerable research to identify stocking rates and grazing systems that would produce a healthy ecosystem. Results are compared with

traditional stocking of shinnery, which was continuous

grazing averaging 45 acres (Pettit 1986) to 52 acres (USDA

Forest Service 1972) per AU (animal unit, a cow-calf pair).

In the dry years 1983 and 1984, Mosley et al. (1985b) found

that stocking for 100% of proper use fell from an average

of 19 acres/AU to 50 acres/AU in a highly productive area.

The efficacy of rotational grazing, in which certain

pastures are rested during the various seasons has been

evaluated in shinnery. In Yoakum County, Texas, Plumb

(1984) found no difference in weaning weights or average

daily gain of calves using a 3-herd, 4-pasture deferred

rotation system stocked at 74 acres/AU relative to continuous grazing at 62 a/AU. The BLM mandated deferred

grazing on 1,400,000 acres in 90 allotments in shinnery

country of its Roswell District, with rest periods from 4

mos to 2 yrs (USDI BLM 1979), but the BLM has not

enforced these decisions (USDI BLM 1997). Deferment

during spring when shin-oak is poisonous to cattle is

helpful but insufficient for grass recovery, since tillering

and seed production come later in the season (USDI BLM

1977). Dahl (1986) stated that rotation and deferment are

better tools to maintain rather than to improve range

condition, but over several years they provide for about

25% more animals than does continuous year-long stocking.

Extensive tests have been made for short-duration grazing in shinnery in which a single herd is moved through 6 or

more paddocks, with moves every 1 to 7 days. Dickerson

(1985, 1987, summarized by Bryant et al. 1989) found that

an 18-paddock system in Garza County, Texas, did not

differ from continuous grazing with regard to herbage
USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998 25

yield, forage digestibility or protein content, animal performance, animal production per acre, or vegetation composition, but it did permit higher stocking rates. In the

same study, Mosley et al. (1985a) found 25% more crown

death of little bluestem, sand dropseed, hairy grama, and

fall witchgrass under continuous grazing. Shifting from 7

days in each pasture to flexible 0-to-14 day periods depending upon available herbage, did not improve results

(Mosley and Dahl 1990). Plumb (1984, Bryant et al. 1989),

working in Yoakum County, Texas, likewise found no

difference in animal performance or production in a 6-

paddock cell stocked at 35 acres/AU compared with

continuous grazing at 62 acres/AU, though again, higher

stocking was possible.

Weir (1990) evaluated cattle performance for 161 days

in Roosevelt County, New Mexico, under continuous and

short-duration grazing at the same stocking rates (19 to 26

acres/head) and found that daily weight gains were higher

under continuous grazing than short-duration grazing;

no differences appeared in quality or composition of diets.

Working on the same ranch, Keesee (1992) measured

physical and hydrologic properties of soils under continuous and short-duration grazing at the same stocking rate;

he was unable to confirm the improvements claimed as a

result of hoof action under short-duration grazing. Dahl

(1986) discussed Texas Tech’s experiments with shortduration grazing and concluded that it reduces sacrifice

areas around range developments, provides opportunity

for plants to recover from grazing, and, in large pastures,

provides opportunity to increase stocking rates. However, short-duration grazing does not improve diet qualities,

animal performance (at stocking rates equal to continuous

grazing), infiltration of water into the soil, or revegetation

of eroded gullies. Ethridge et al. (1987b) discussed the

economics of an 8-paddock system in shinnery.

The only grazing system claimed to improve grass

production in shinnery was the 1-herd, 4-pasture system

described for Eddy County, New Mexico, by Moore (1954).

Bunching cattle in 1 pasture had the advantage of forcing

them to eat the less palatable grasses, which cost something in weight gains but, “we feel that the benefit to the

range is greater than the loss to the stock” (Moore 1954).

Moore offered no data to support his claim and there has

been no controlled experiment to test it. However, this

grazing system is probably closest to that used for thousands of years by bison, which moved through shinnery in

close-knit groups in search of good forage (Shaw 1996),

although they probably moved more than did Moore’s

cattle.

Rotational grazing is recommended for improvement

of lesser prairie-chicken, bobwhite quail, scaled quail, and

turkey habitat (Bryant et al. 1982, Holechek 1981, Taylor

and Guthery 1980a). Ahlborn (1980) suggested a flexible

strategy such as a best-pasture system (choice of pasture

based on vegetative condition). Litton et al. (1994) recommended short-duration grazing for its hoof-action effect,

believing that ground disturbance favors forbs that provide prairie-chicken food.

More important than the grazing system for groundnesting birds are stocking rates. Light to moderate grazing

secures nesting cover for bobwhite quail (Guthery 1982),

while serious reductions in prairie-chicken and scaled

quail populations may follow overgrazing (Dixon and

Knight 1993, Duck and Fletcher 1944, Litton et al. 1994).

Haukos and Smith (1989) noted that prairie-chicken hens

selected nest sites within lightly grazed shinnery with

>75% vertical screening in the first 13 inches and 50%

overhead cover. Riley et al. (1992) found, on average, 26-

inch bluestem grasses at successful prairie-chicken nests

and 14-inch grasses at unsuccessful ones. One important

way to achieve this degree of cover is with livestock

exclosures, which are discussed by Davis et al. (1979),

Frary (1957), Sell (1979), Snyder (1967), and USDI BLM

(1979) for prairie-chickens; by Dixon and Knight (1993) for

scaled quail; by Holechek (1981) and Webb and Guthery

(1982) for bobwhite habitat; by Bryant et al. (1982) and

Holechek et al. (1982) for turkeys; and for all these species

by Brown (1978).

Effects of grazing on other wildlife vary. Dietary overlap between deer and cattle in shinnery is minimal (Bryant

and Demarais 1992, Bryant and Morrison 1985). Deer may

avoid ranching operations and often select for deferred

pastures over actively grazed areas (Holechek et al. 1982,

Hood and Inglis 1974). The significance of this behavior to

the well-being of deer populations is unclear. Shin-oak

was the main item of dietary overlap between pronghorns

and cattle in the Texas Panhandle but because there was

plenty, the overlap posed no problem (Roebuck 1982).

Livestock grazing alters food and cover for non-game

birds, particularly by altering the vertical structure of

habitat. A few species benefit from grazing, while many

species react unfavorably. However, species response varies

drastically between sites and between years within sites

so that lists of species are highly conditional (Knopf 1996).

Effects of livestock grazing on small mammal populations are variable and species-dependent. Plains pocket

gophers are most abundant on moderately grazed range,

preferring the varied vegetation of disturbed sites, whereas

woodrats, cotton rats, and harvest mice prefer ungrazed

range (Fagerstone and Ramey 1996). These generalizations

are not based on studies in shinnery. Black-tailed jackrabbits have been variously reported to do best on ungrazed,

moderately grazed, or overgrazed rangeland; dietary overlap is moderate as jackrabbits consume less grass and

more shrubs than cattle (Fagerstone and Ramey 1996).

Effects of cattle-grazing on the sand dune lizard are unclear.

In sites where most shin-oak had been killed, there were

fewer lizards under heavy grazing than in lightly grazed

areas, but heavily grazed untreated areas had slightly

more lizards than those lightly grazed (Snell et al. 1993).
26 USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998

kinds of mowers, beaters, cutters, choppers, and other

mechanical devices have been unsuccessfully used to

control shinnery oak” (McIlvain 1954). Removal of aboveground shoots of the oak increases grass production primarily in the eastern shinnery, “where rainfall is high and

potential production is good. However, the economics of

the practice are questionable” (McIlvain 1954). Shredding

above-ground parts of oaks weakens the plants, but they

resprout and after a few months their levels of available

carbohydrate have recovered (Bóo and Pettit 1975).

Fire

Control of shin-oak using prescribed fire is hampered

because the oak typically resprouts vigorously in the

growing season following fire, with regrowth reaching 3

to 4 ft within 2 or 3 growing seasons (McIlvain 1954).

Although more research is needed, the use of annual or

biennial prescribed fire may decrease the canopy height of

shin-oak and increase production of perennial grasses.

McIlvain (1954) speculated that annual fires were used to

control shin-oak during early settlement in Oklahoma.

Management of Shin-oak

Shin-oak is often regarded as a noxious and undesirable weed for agriculture and livestock grazing: Shin-oak

competes with better livestock forage and its buds and

leaves are toxic to cattle for several weeks in spring

(Dollahite 1961, Marsh 1919). In the last 25 yrs, an additional reason for control of shin-oak has developed. In 4

Texas counties where cotton is grown, boll weevils overwinter in shin-oak litter and infest nearby cotton fields in

spring (Slosser et al. 1984, 1985). Most research and other

human effort concerning shin-oak have been devoted to

its eradication.

Mechanical Means

Plowing shinnery to create fields for crops is expensive

but successful; almost 1 M acres have been cleared in

Texas (Deering and Pettit 1972). The practice is ecologically questionable given the high regional potential for

wind erosion (Lotspeich and Everhart 1962). Where improved rangeland for cattle is the goal, “Many different

Figures 16. Plant growth during a wet year, 4 months after an

April 29, 1997, prescribed fire in Ellis County, Oklahoma.

Grasses dominate the post-fire community. The measuring

board is marked in 10-cm increments to a height of 130 cm.

Figure 15. Prescribed fire moves through sand shinnery

during the growing season in Ellis County, Oklahoma. The

pole is 2 m high.
USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998 27

and forbs without changing total production, and also

greatly increased nitrogen, phophorus, and potassium in

soil. Goats kept continuously on shin-oak lost weight. The

authors recommended rotating goats out of target pastures when defoliation of shin-oak reaches 80% and returning them when regrowth leaves have attained one

half of their mature size (Escobar et al. 1995). Control by

goats is promising.

Herbicides

Chemical-control trials by the Agricultural Research

Service began at Woodward, Oklahoma, in 1947 (McIlvain

1954). In 50 yrs, a great deal of effort at Woodward, Texas

Tech, and Texas A & M has gone into herbicidal control of

shin-oak. This effort has resulted in about 75 articles and

theses on how and when to control, another 40 on the

effects of control on vegetation and soils, and others on the

effects on mammals, birds, and reptiles.

Most early trials involved phenoxy herbicides including 2, 4-D and 2, 4, 5-T and Silvex (2-(2,4,5-T) propionic

acid), benzoic acids, including dicamba, and a picolenic

acid, picloram (Tordon), which are applied as liquids and

absorbed through foliage. At a pound or more per acre of

active ingredient, these herbicides top-killed 85% to 95%

of the oak and root-killed a small percentage (Greer et al.

1968, NM Inter-Agency Comm. 1970, Pettit 1977). Lighter

application of herbicide was usually recommended because of adverse effects on forbs. Control was expensive

but effective. Spray applications that killed 10% of the oak

doubled grass production, which was measured 3 yrs

later. At 20% kill, production was tripled 2 yrs after the

second annual spraying. A 90% kill, resulting from 3

annual sprayings, quadrupled production a year after the

third spraying, although grass production declined to the

level of twice the original production within a few years

(Greer et al. 1968, McIlvain and Armstrong 1959). Much

higher application rates, 2.7 to 7 lb/acre of picloram,

killed oak with 1 spraying but desirable forbs and seedlings of perennial grasses had scarcely reentered the

sprayed plots 6 yrs later (Pettit 1976, 1977).

Beginning in 1974, a pellet form of tebuthiuron (Spike

or Graslan, a thiadiazol combined with dimethlyurea

and dimethylethylene), which acts through the soil and is

absorbed by roots, was tested on shin-oak. Tebuthiuron

defoliates the oak, which then leafs out repeatedly with

decreasing vigor, followed by death in the second, third,

or later years after treatment (Jones and Pettit 1984). At 2.7

lb active ingredient/acre, the oak was killed and many

non-target forbs and grasses were set back (Pettit 1979). At

.36 lb active incredient/acre, oak canopy was reduced

95% and grass yield was 2.5 times the control after 3 yrs.

At .54 to .9 lb/acre all oak was killed (figure 18). At .72 lb/

acre, the maximal grass yield of 4 times the control was

obtained (Jones 1982, Jones and Pettit 1984).

Given the resilience of shin-oak to fire, it is unlikely that

annual fires would eliminate shin-oak from the community. Current research by Oklahoma State University (figures 15 to17) may help determine the use of annual and

timed spring burns for managing shin-oak.

Herbivory

Goats have long been used in Texas for control of

woody plants, including oaks, and there were early trials

in southeastern New Mexico to control shin-oak with

goats. In shinnery in west Texas where oak provided 79%

of available forage, oak increased in the diets of Angora

and Spanish goats from 31% in June to 51% in August, of

which 85% was oak leaves and 15% stems. The increase

was likely due to the goats’ previous inexperience with

oak and decreasing availability of forbs (Villena et al.

1987, Villena-Rodriguez 1987). Trials over 3 yrs in Black

Kettle National Grassland (Roger Mills County, Oklahoma) used both meat-type and mohair-producing goats.

The goats succeeded in shifting forage production from

95% oak and 5% grass and forbs to 50% oak and 50% grass

Figure 17. Plant growth during a wet year, 4 months after an

April 29, 1997, prescribed fire in Ellis County, Oklahoma.

Shin-oak resprouts form an understory 30-40 cm tall.
28 USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998

Total production was generally constant. The herbicide

shifted its composition from oak toward grass, which

increased from 26% to 700% following control depending

on the degree of oak cover, percentage kill, initial grass

cover, soil type, and precipitation (Gribble 1981, Jacoby et

al. 1983, Jones and Pettit 1982, 1984, Pettit 1979, 1986,

Plumb 1984). Much of the increase was annual grasses and

undesirable sandbur (Jacoby et al. 1983, Plumb 1984).

Grasses begin growth from 3 to 6 weeks earlier in treated

pastures than in untreated (Biondini et al. 1986, Plumb

1984). In the 4 grass species sampled, crude protein was

28% higher in treated than in untreated shinnery in the

treatment year, but the effect was gone a year later (Biondini

et al. 1986). Plumb (1984) found that cows on treated

pasture were above protein maintenance levels in all

seasons except winter. On untreated pastures, cows required protein supplement in all seasons except summer.

Forb response to tebuthiuron is uneven but generally

after 1 or 2 yrs of depleted production, the variety and

yield of forbs become greater than in untreated pastures

(Doerr and Guthery 1983b, Jacoby et al. 1983, Jones and

Pettit 1982, Olawsky and Smith 1991, Pettit 1986, Willig et

al. 1993). Bare soil plus litter decreased from 35% in

untreated plots to 20% in treated, while the percentage of

living-plant cover increased (Jacoby et al. 1983). By 1995,

320,000 acres of shinnery had been treated with tebuthiuron

in Texas (Johnson and Ethridge 1996). From 1981 to 1993,

the BLM sprayed 100,000 acres in New Mexico (USDI

BLM 1997).

In the eastern two-thirds of the shinnery region, the

economics of shinnery conversion can be favorable on

livestock ranches, though very sensitive to rainfall, depth

of fine sand, herbicide price, and beef prices (Etheridge et

al. 1987a, 1987b, Jones 1982, Neal 1983, Pettit 1986,

Richardson and Badger 1974). By extension of the calculations of Etheridge et al. (1987a) it appears that shin-oak

control loses money farther west, including New Mexico,

where precipitation is less and forage increases are smaller,

or where wildlife values should be included in the calculations.

Pettit (1983a) reported that brownspine prickly pear

was indirectly controlled by tebuthiuron: after shin-oak is

defoliated the cactus gains vigor or at least remains static,

but as grasses begin to dominate the cactus shows signs of

stress. Within 2 yrs, many cactus pads are infested with

borers and scale insects, and Pettit estimated that the cacti

would die about 10 yrs after herbicidal treatment.

Herbicidal treatment alters the distribution of biomass

in shinnery. Sears et al. (1986a) reported that 3 yrs after

tebuthiuron treatment, total above-ground biomass decreased by 2% relative to control plots, which is the result

of a 17% decrease in oak and litter and a 266% increase in

herbaceous vegetation. Six yrs following treatment, aboveground biomass had decreased by 29% (41% decrease in

oak, 32% decrease in litter, and 161% increase in herbaceous vegetation). Surprisingly, the ratio of above-ground

to below-ground biomass was the same in controls and in

6-yr treated shinnery, 1: 5.7 (Sears et al. 1986a). Litter can

cover 100% of the soil under oak.

This decreases after control but is

partly compensated by an increase in herbs (Sears 1982). Soil

organic matter increases with the

decay of oak (Sears et al. 1986b).

Soil nutrient and water relations are changed by herbicidal

control. Nitrogen (nitrate + ammonium) concentrations were

higher in soils of treated shinnery than in untreated and increased for years after control

(Kyrouac 1988, Sears et al. 1986b).

Phosphorus may also have increased in soils after control but

the effect was not as long-lasting

(Kyrouac 1988). Removal of oak

adds little to soil moisture if herbs

are allowed to flourish, nor does

removal of herbs add much to

soil moisture since the oaks will

use more (Pettit and Holder 1973,

Pettit and Small 1974). Galbraith

(1983) did find some increase in

soil water after treatment but

Figure 18. This treated sand shinnery is a rich mix of grasses and forbs; little bluestem,

sand bluestem, annual wild buckwheat, and fall witchgrass are visible.
USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998 29

mainly found water differently distributed; the increase

was from 35 to 100 inches depth due to death of the deep

oak roots. Test (1972) also found more soil water in treated

plots than untreated (37% more in July, 9% in wetter

September soils) but most of the increase was at depths

less than 30 inches.

Small differences between treated and untreated areas

in development of vesicular-arbuscular mycorrhizae on

little bluestem and purple three-awn grasses were likely

due to increases in soil nutrients and moisture due to

treatment (Kyrouac 1988).

Experimenters with tebuthiuron treatment have issued

several cautions: shin-oak should not be treated in drought

years, in areas of large dunes, or where perennial grasses

are few, and treated pastures should not be burned or

summer-grazed for at least 2 yrs after treatment (Doerr

and Guthery 1980b, McIlvain and Armstrong 1959, N. M.

Inter-Agency Comm. 1970). “These herbicides are not

recommended for application to an oak community in

poor range condition. These lands are perhaps the most

fragile of all ecosystems on the southern High Plains of

Texas and the landowner cannot afford to abuse them”

(Pettit 1979). A year after shin-oak treatment, a pasture

“supporting a midgrass prairie” was many times more

susceptible to wind erosion than untreated shinnery

(Thurmond et al. 1986). Even where surface cover increased from 60% to 65% due to herbicidal treatment,

there was an 80% increase in wind-blown sediment (Zobeck

et al. 1989). Because of the fine fuel load, treated land also

becomes more susceptible to wildfire. In sand dune habitat, Petit (1985) reported that 50% of grasses were killed by

fire or by blowing sand and over 3 inches of sand were

removed. The biggest danger from herbicide-based control is erosion following drought. The grasses and forbs

that replace oak are not as resistant to drought as is shinoak, and their death would leave the soil exposed.

The usefulness of shin-oak as livestock feed during

drought, as well as erosion control, indicate that preserving some oak is beneficial. “Research results in Oklahoma

show greater forage production when shinnery makes up

part of the vegetation than when grasses are alone on these

soil types” (N. M. Inter-Agency Comm. 1970). According

to McIlvain and Armstrong (1963), on steep dunes the goal

for brush control should be 25% kill, on low dunes without

thick grass stands kill should be less than 50%, on gently

rolling or nearly flat areas that support dense grasses kill

should be 70% to 80%.

On multiple use land, the question of treatment becomes more complex, especially regarding the lesser prariechicken. Jackson and DeArment (1963) claimed that largescale brush control produces communities that the prairie-chicken does not readily adjust to. Even a 25% kill

prevented acorn production for 2 yrs and the effects were

apparent in lower prairie-chicken populations. Copelin

(1963) discussed the difficulties of determining whether

brush control helped or hurt prairie-chicken populations.

But Donaldson (1966, 1969) claimed benefits from shinoak control. He counted birds for 1 season where 75% of

shin-oaks had been killed, forb production had decreased,

and grass production had doubled. He found 10 times as

many prairie-chickens in treated areas as in untreated

areas. However, shin-oak was the plant that the birds used

most, especially in the treated areas, and he proposed

supplemental winter feeding to make up for loss of acorns.

In the 1960s, the New Mexico Department of Game and

Fish negotiated an understanding with the BLM prevening

shin-oak control in federal prairie-chicken areas (Sands

1968). The BLM (USDI BLM 1984) later abrogated the

memorandum and from 1981 through 1993 treated 100,000

acres. There is no evidence from Texas or New Mexico

indicating that shin-oak treatment with tebuthiuron benefits prairie-chickens. Two studies used paired plots to

compare prairie-chickens in treated and untreated shinnery: Olawsky (1987, Olawsky et al. 1988) found little

change in populations but in treated areas there was a

decrease in lipid levels that indicated worse overall condition, which he tentatively blamed on lack of acorns. Martin (1990) found 86% fewer prairie-chickens in treated

areas than in untreated areas but seeing the birds in grass

was a problem. A third study used radiotelemetry to track

females and found that they preferred nesting in untreated rather than treated shinnery (Haukos and Smith

1989b). If shin-oak control benefited prairie-chickens it

would do so by creating a mosaic of small blocks of

vegetation (Davis et al. 1979, Sell 1979, Taylor and Guthery

1980a). “Because of the importance of shinnery oak grassland to prairie-chickens for both food and cover, broadscale eradication of this community should be avoided”

(Riley et al. 1993a, Olawsky and Smith 1991). Use of

picloram to treat 150 to 300 ha blocks was recommended

by Doerr and Guthery (1980b) to improve nesting cover,

providing that control was partial and admitting that

these large blocks were a compromise between the habitat

requirements of the birds and the economics of treatment.

Control of shin-oak affects other wildlife species variously. For mule deer habitat, “Any mechanical or chemical treatments should be avoided where damage to

skunkbush sumac, wild plum, or sand-shinnery oak is a

possibility” (Bryant and Morrison 1985). Javelina presumably do better in untreated shinnery because of the importance to them of shin-oak and prickly pear (Bryant et al.

1982). Bryant and Demarais (1992) emphasized whitetailed deer use of acorns and browse. They thiought that

opening up dense browse can be beneficial if openings are

less than 300 yards wide, ideally 50 to 100 yards wide.

Darr and Klebenow (1975) discouraged treatment of shinnery, especially oak mottes.

In Eddy County, New Mexico, Bednarz (1987) found

lagomorph populations depressed the year following complete elimination of shin-oak, with some recovery in the
30 USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998

second and third years following spraying. In Cochran

County, Texas, rodent populations were 41% higher on

untreated plots than where the shin-oak had been 75%

removed by tebuthiuron (Doerr and Guthery 1980a). In

Yoakum County, Texas, where treatment removed 90% of

the oak, populations of Ord’s kangaroo rat more than

doubled on treated plots; northern grasshopper mouse

populations were lower in winter and spring on treated

plots but higher in fall, plains pocket mice prospered on

treated plots, and deer mice were fewer on treated plots

(Colbert 1986, Willig et al. 1993). In Eddy County, Fischer

(1985) also found larger Ord’s kangaroo rat populations

on treated areas; other species seemed unaffected.

Davis et al. (1974) reviewed the effects of brush control

on bird species in Eddy County. They thought that any

control should be in swaths. When control was in large

blocks in Eddy County, Bednarz (1987) found that the

success of raven nesting “crashed” following treatment

although populations did not change. He hypothesized

that lack of suitable foods was the cause. On plots in

Chaves County, New Mexico, where shin-oak was reduced 90% by treatment and grasses doubled, numbers of

birds and bird species were not significantly different

from untreated plots. Cassin’s sparrows may have increased slightly on treated plots, only prairie-chickens

sharply decreased (Martin 1990). Bobwhite quail use of

treated areas was almost double that of untreated areas

where control was in 20-ft wide strips (Webb and Guthery

1982). In Eddy County, scaled quail populations fell following treatment, while bobwhite quail and most other

bird species were unaffected and loggerhead shrike populations increased (Bednarz 1987). Meadowlark populations may double in treated shinnery (Olawsky et al.

1987).

In Chaves County, 6 of 7 species of lizard censused

were more common in untreated than in treated shinnery,

the endangered sand-dune lizard was 5 to 6 times more

common. The fence lizard, lesser earless lizard, sideblotched lizard, western whiptail, and six-lined racerunner

were significantly more common in untreated areas. The

Great Plains skink was significantly more common in

treated areas (Gorum et al. 1995, Snell et al. 1997).

A possible compromise between tebuthiuron treatments that destroy shin-oak and no control is to use the

foliar herbicides of the 1960s that merely defoliate the oak,

giving its herbaceous competitors a period of time in

which to regain dominance. The use of goats and of fire are

also promising control methods that may not affect wildlife unduly. Most studies ( e.g. Bednarz 1987, Harmel

1979) suggest interspersing untreated areas with treated

so that shin-oak can provide food and cover.

Conclusions

Shaped by a balance between an extremely long-lived

shrub and grasses and forbs that come and go with climatic and human-caused changes, the shinnery is an

unusual biological community with unusual problems.

Problems concern sand shin-oak, whose virtues are tempered by its negative effects on livestock grazing. Shinoak is targeted because it is seasonally poisonous to cattle,

because grass production can be increased by herbicidal

treatment, and because of the erroneous beliefs that it has

invaded and increased. Conversion to cultivated land or

to seeded pasture destroys the shinnery ecosystem. Loss

of tallgrasses to livestock or of shin-oak to herbicides

destroys the integrity of the ecosystem. Only small samples

resembling undisturbed shinnery remain.

The shinnery is rich habitat for wildlife compared with

the surrounding short grass plains. Protecting sandy soils

from wind erosion, maintaining wildlife habitat, and improving vegetation for livestock grazing are often in conflict and should be carefully considered before undertaking control of shin-oak.

More research is needed especially to: 1) identify grazing management to help restore and maintain grasses of

the shinnery, 2) explore the role of fire in restoring and

maintaining the shinnery ecosystem, and 3) examine the

effects of different methods and configurations of shinoak control on wildlife species.

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38 USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998

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New Mexico during the past hundred years. pp. 157-166. In: W.G.

McGinnies & B. J. Goldman, editors, Arid Lands in Perspective.

Tucson: University of Arizona Press. 421 pp.

Zhang, Q. 1996. Fungal community structure and microbial biomass in

a semi-arid environment: roles in root decomposition, root growth,

and soil nitrogen dynamics. Ph.D. thesis, Texas Tech University.

234 pp.

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wind-eroded sediment and plant nutrients. Journal of Soil and Water

Conservation 44: 160-163.
USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998 39

Appendix

Plants of Shinnery Communities

Plants are listed that were recorded in shinnery by Sullivan (1980) for 6 counties of western Texas adjacent to New

Mexico, by R. Peterson (unpublished) for the Mescalero Sands of Chaves County, NM, and by C. Boyd (unpublished) for

Black Kettle National Grassland (Roger Mills County) and southern Ellis County, OK. Plant lists for shinnery of the Texas

Panhandle such as that of Sears (1982) resemble Boyd’s list. Nomenclature largely accords with the Great Plains Flora

(1986) except in Bothriochloa, Digitaria, Elymus, Schizachyrium, Vulpia, Brickellia, Dimorphocarpa, Heterotheca, and Mimosa.

TX NM OK

Woody Plants

Amorpha canescens leadplant +

Artemisia filifolia sandsage + + +

Baccharis texana prairie baccharis +

Celtis reticulata netleaf hackberry + + +

Chrysothamnus pulchellus Southwest rabbitbrush + +

Coryphantha (Mammillaria) vivipara pincushion, ball cactus +

Ephedra antisyphilitica vine ephedra +

Ephedra sp. ephedra +

Ferocactus wislizenii barrel cactus +

Gutierrezia sarothrae broom snakeweed + + +

Lycium berlandieri wolfberry +

Mimosa biuncifera catclaw mimosa +

Opuntia imbricata cholla +

Opuntia leptocaulis tasajillo +

Opuntia macrorhiza plains pricklypear +

Opuntia phaeacantha brown-spine pricklypear +

Opuntia polyacantha plains pricklypear + +

Populus fremontii subsp. wislizenii Rio Grande cottonwood +

Prosopis glandulosa honey mesquite + + +

Prunus angustifolia Chickasaw plum +

Prunus gracilis Oklahoma plum + +

Quercus havardii sand shin-oak + + +

Rhus aromatica skunkbush, squawbush + + +

Salix exigua coyote willow +

Salix nigra (S. gooddingii) Goodding’s willow +

Sapindus saponaria soapberry + +

Tamarix spp. tamarisk, saltcedar + +

Yucca campestris plains yucca +

Yucca glauca soapweed yucca + + +

Ziziphus obtusifolia lotebush +

Graminoids

Andropogon gerardii big bluestem +

Andropogon hallii sand bluestem + + +

Aristida divaricata var. havardii Havard’s three-awn + +

Aristida purpurea (4 varieties) purple three-awn + + +

Bothriochloa barbinodis cane bluestem + +

Bothriochloa laguroides silver bluestem + + +

Bothriochloa springfieldii bluestem +

Bouteloua curtipendula side-oats grama + + +

Bouteloua eriopoda black grama + +

Bouteloua gracilis blue grama + + +
40 USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998

Graminoids (Cont’d.)

Bouteloua hirsuta hairy grama + + +

Bouteloua rigidiseta Texas grama +

Brachiaria ciliatissima fringed signalgrass + +

Bromus tectorum cheatgrass +

Buchloe dactyloides buffalograss +

Calamovilfa gigantea giant sandreed + +

Cenchrus incertus sandbur + + +

Chloris cucullata hooded windmillgrass + +

Chloris verticillata tumble windmillgrass + + +

Cyperus onerosus flatsedge +

Cyperus schweinitzii Schweinitz flatsedge + + +

Cyperus uniflorus one-flower flatsedge +

Dichanthelium oligosanthes rosette-grass +

Digitaria (Leptoloma) cognata fall witchgrass + + +

Elymus canadensis Canada wild rye +

Elymus longifolius squirreltail grass +

Enneapogon desvauxii spike pappus-grass +

Eragrostis cilianensis stinkgrass +

Eragrostis curtipedicellata gummy lovegrass + + +

Eragrostis curvula weeping lovegrass +

Eragrostis lehmanniana Lehmann lovegrass + +

Eragrostis secundiflora red lovegrass + + +

Eragrostis spectabilis purple lovegrass +

Eragrostis sessilispica tumble lovegrass + +

Eragrostis trichodes sand lovegrass + +

Erioneuron (Dasychloa) pulchellum fluffgrass +

Juncus scirpoides needlepod rush +

Lycurus phleoides (L. setifolia) wolftail +

Muhlenbergia arenacea ear muhly +

Muhlenbergia porteri bush muhly +

Munroa squarrosa false buffalograss + +

Panicum capillare witchgrass +

Panicum hallii Hall’s panicum +

Panicum havardii Havard panicum + +

Panicum obtusum vine-mesquite +

Panicum virgatum switchgrass +

Paspalum setaceum sand paspalum + + +

Schedonnardus paniculatus tumble-will +

Schizachyrium scoparium little bluestem + + +

Scirpus acutus hardstem bulrush +

Scleropogon brevifolius burrograss +

Setaria leucopila (S. macrostachya) plains bristlegrass + +

Setaria (Panicum) ramisetum bristle panicum +

Setaria viridis green foxtail +

Sorghastrum nutans Indiangrass + +

Sporobolus contractus spike dropseed +

Sporobolus cryptandrus sand dropseed + + +

Sporobolus flexuosus mesa dropseed + +

Sporobolus giganteus giant dropseed + +

Sporobolus vaginiflorus poverty grass +

Stipa comata needle-and-thread + +

Triplasis purpurea purple sandgrass +

Vulpia octoflora six-weeks fescue + + +

Forbs

Abronia fragrans snowballs, sandverbena + +

Achillea millefolium yarrow +

TX NM OK
USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998 41

Forbs (Cont’d.)

Allionia incarnata trailing four-o’clock, windmills +

Allium drummondii wild onion +

Amaranthus retroflexus redroot pigweed +

Ambrosia psilostachya western ragweed + + +

Aphanostephus skirrhobasis sleepydaisy +

Apocynum cannabinum dogbane + +

Argemone polyanthemos prickly poppy +

Argemone sp. prickly poppy +

Artemisia caudata threadleaf sagewort +

Artemisia frigida estafiata, prairie sagewort +

Artemisia ludoviciana white sage +

Asclepias arenaria sand milkweed + +

Asclepias asperula antelopehorns +

Asclepias engelmanniana Engelmann milkweed +

Asclepias pumila plains (or dwarf) milkweed + +

Asclepias subverticillata poison milkweed +

Asclepias tuberosa butterfly milkweed +

Asclepias verticillata whorled milkweed +

Asclepias viridiflora green antelopehorns + +

Aster ericoides white aster +

Aster patens aster +

Astragalus mollissimus woolly loco + + +

Baptisia australis blue false indigo +

Berlandiera lyrata greeneyes, chocolate flower +

Brickellia (Kuhnia) eupatorioides false boneset + +

Caesalpinia jamesii James’ rushpea + +

Callirhoe involucrata poppy mallow +

Calylophus berlandieri sundrops +

Calylophus hartwegii sundrops +

Calylophus serrulatus halfshrub sundrops + + +

Cassia chamaecrista partridgepea +

Chenopodium album lamb’s quarters +

Chenopodium leptophyllum slimleaf goosefoot +

Cirsium ochrocentrum yellow-spine thistle + + +

Cirsium undulatum wavy-leaf thistle +

Cnidoscolus texanus Texas bull-nettle +

Comandra umbellata subsp. pallida bastard toadflax +

Commelina erecta dayflower + + +

Conyza canadensis horseweed +

Corispermum hyssopifolium tickseed, bugseed +

Corispermum nitidum tickseed, bugseed + +

Croton dioicus hierbo del gato +

Croton pottsii leatherweed croton + +

Croton texensis Texas croton + + +

Cryptantha cinerea hiddenflower + +

Cryptantha minima small hiddenflower +

Cuscuta glabrior dodder +

Cycloloma atriplicifolium tumble ringwing + +

Dalea aurea golden prairie-clover +

Dalea candida white prairie-clover +

Dalea lanata woolly dalea + +

Dalea nana dwarf dalea + +

Dalea purpurea purple prairie-clover + +

Dalea tenuifolia slimleaf prairie-clover +

Dalea villosa silky prairie-clover + +

Datura quercifolia oak-leaf thornapple +

Datura stramonium jimsonweed +

TX NM OK
42 USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998

Forbs (Cont’d.)

Delphinium virescens plains larkspur + +

Desmodium sessilifolium sessile-leaved tickclover +

Dimorphocarpa (Dithyrea) candicans spectaclepod +

Dimorphocarpa (Dithyrea) wislizenii spectaclepod + +

Erigeron bellidiastrum fleabane +

Erigeron modestus (broad sense) plains fleabane + + +

Erigeron spp. fleabane +

Eriogonum annuum annual wild buckwheat + + +

Erodium texanum Texas stork’s-bill +

Erysimum capitatum wallflower +

Euphorbia carunculata spurge +

Euphorbia fendleri spurge + +

Euphorbia geyeri spurge +

Euphorbia micromera spurge +

Euphorbia missurica spurge + +

Euphorbia parryi spurge +

Euphorbia prostrata spurge +

Euthamia gymnospermoides viscid euthamia +

Evolvulus alsinoides ojo de víbora +

Evolvulus nuttallianus hairy evolvulus + +

Evolvulus sericeus silky evolvulus +

Froelichia floridana snakecotton + + +

Froelichia gracilis snakecotton +

Gaillardia pinnatifida yellow blanketflower +

Gaillardia pulchella firewheel, Indianblanket + + +

Gaillardia suavia rayless gaillardia +

Gaura coccinea scarlet gaura + + +

Gaura suffulta gaura +

Gaura villosa woolly gaura + +

Gnaphalium sp. cudweed +

Guilleminea lanuginosa woolly cottonflower +

Gutierrezia dracunculoides annual broomweed +

Haplopappus spinulosus goldenweed, ironplant + + +

Hedyotis humifusa mat bluets + +

Helianthus annuus annual sunflower + + +

Helianthus petiolaris plains sunflower + +

Heliotropium convolvulaceum bindweed heliotrope + +

Heterotheca canescens golden aster +

Heterotheca latifolia camphorweed + + +

Heterotheca villosa golden aster +

Hoffmanseggia drepanocarpa sicklepod rushpea + +

Hymenopappus flavescens yellow woollywhite + +

Hymenopappus tenuifolius woollywhite +

Hymenoxys acaulis bitterweed +

Hymenoxys odorata bitterweed +

Hymenoxys scaposa bitterweed +

Ipomoea leptophylla bush morning-glory +

Ipomopsis (Gilia) longiflora blue trumpets + +

Krameria lanceolata trailing ratany + + +

Lactuca ludoviciana western wild lettuce +

Lechea mucronata pinweed +

Lepidium virginicum peppergrass +

Lespedeza stuevei tall-bush lespedeza +

Lesquerella fendleri bladderpod +

Leucelene ericoides baby aster, sand aster +

Liatris punctata gayfeather + +

Linaria canadensis toadflax +

TX NM OK
USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998 43

Forbs (Cont’d.)

Linum aristatum yellow flax +

Linum pratense Norton’s flax +

Linum rigidum stiffstem flax + +

Linum sulcatum grooved flax +

Lithospermum arvense puccoon +

Lithospermum carolinense puccoon +

Lithospermum incisum gromwell, puccoon + + +

Lygodesmia juncea rush skeletonplant +

Machaeranthera canescens daisy (lavender, white forms) +

Machaeranthera tanacetifolia Tahoka daisy + +

Maurandya antirrhiniflora snapdragon vine +

Melampodium leucanthum blackfoot daisy + +

Mentzelia nuda blazingstar + +

Mentzelia perennis blazingstar +

Mentzelia strictissima blazingstar +

Mimosa (Schrankia) quadrivalvis sensitive brier + + +

Mirabilis (Oxybaphus) carletonii four-o’clock +

Mirabilis (Oxybaphus) glabra four-o’clock +

Mirabilis (Oxybaphus) linearis four-o’clock + + +

Mollugo cerviana carpetweed +

Mollugo verticillata Indian chickweed, carpetweed +

Monarda citriodora lemon beebalm +

Monarda pectinata plains beebalm +

Monarda punctata dotted beebalm +

Oenothera albicaulis evening primrose +

Oenothera engelmannii evening primrose +

Oenothera rhombipetala evening primrose + +

Orobanche ludoviciana broomrape +

Palafoxia sphacelata palafoxia + +

Paronychia jamesii nailwort + +

Pectis angustifolia limoncillo +

Pectis papposa chinchweed +

Penstemon albidus white beardtongue +

Penstemon ambiguus plains penstemon + +

Penstemon buckleyi Buckley penstemon + + +

Phacelia integrifolia scorpionweed +

Phyllanthus abnormis leaf-flower + +

Physalis viscosa groundcherry + +

Physalis sp. groundcherry +

Plantago patagonica Indianwheat + + +

Polanisia dodecahedra clammyweed +

Polanisia jamesii clammyweed +

Polygala alba white milkwort +

Portulaca mundula chisme, wild purslane + +

Portulaca oleracea purslane + +

Psilostrophe villosa hairy paperflower +

Psoralea tenuiflora scurfpea + +

Pyrrhopappus multicaulis false dandelion +

Ratibida columnifera prairie coneflower + + +

Reverchonia arenaria sand reverchonia + +

Ruellia humulis fringe-leaf ruellia +

Salsola iberica Russian thistle + +

Salvia azurea blue sage +

Senecio flaccidus (2 varieties) threadleaf groundsel + +

Senecio riddellii Riddell ragwort +

Senecio spartioides broom groundsel + +

Solanum elaeagnifolium horsenettle, nightshade + + +

TX NM OK
44 USDA Forest Service Gen. Tech. Rep. RMRS–GTR–16. 1998

Forbs (Cont’d.)

Sphaeralcea coccinea scarlet globemallow + +

Stillingia sylvatica queen’s-delight + + +

Streptanthus hyacinthoides twistflower +

Talinum calycinum rockpink +

Talinum parviflorum fame-flower +

Tephrosia virginiana goat’s rue +

Teucrium laciniatum germander +

Thelesperma filifolium gray greenthread +

Thelesperma megapotamicum greenthread, Navajo tea + +

Tidestromia lanuginosa tidestromia + +

Townsendia exscapa Easter daisy + +

Tradescantia occidentalis prairie spiderwort + + +

Tragia ramosa noseburn +

Tribulus terrestris puncturevine +

Triodanis perfoliata Venus looking-glass +

Verbena bracteata prostrate vervain +

Verbesina encelioides cowpen daisy; crownbeard +

Vernonia baldwinii western ironweed +

Xanthisma texanum sleepydaisy + +

Xanthium strumarium cocklebur + +

Zinnia grandiflora wild zinnia +

TX NM OK

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and technology to improve management, protection, and use of forests

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Forest managers, federal and state agencies, public and private

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